Comparing patterns of human harvest and predation by Eurasian lynx Lynx lynx on European roe deer Capreolus capreolus in a temperate forest

Human harvest is the most important mortality factor for wild ungulates in Europe and can affect several aspects of ungulate biology. There is a growing concern about possible negative side effects of human harvest. To better understand the differences between human and natural mortality, we compared the extent, age and sex structure, nutritional condition, spatial and temporal distribution of human harvest, and natural predation by the Eurasian lynx Lynx lynx on the European roe deer Capreolus capreolus, the most abundant wild ungulate in Europe. Compared to the human harvest, lynx were less likely to kill fawns and yearlings than adults, and among adult deer, lynx were more likely to kill females. The proportion of roe deer with fat-depleted bone marrow was higher among lynx prey than among harvested animals. Average lynx kill rate was estimated to 47.8 roe deer per year, and lynx predation was considerably lower than the human harvest in the same area. While human harvest increased with higher roe deer density, lynx predation was similar across the gradient of roe deer densities. Comparison with other countries indicated that differences between human harvest and natural mortality of ungulates vary considerably in different parts of Europe. Variation in hunting practices and, even more importantly, carnivore predation may have an important role in buffering unwanted side effects of harvest of wild ungulates.     

Use of winter habitat by roe deer at a northern latitude where Eurasian lynx are present

Winter climate at northern latitudes is a challenge to small-bodied ungulates, and they modify behaviour to save energy and to increase the likelihood of survival. Also, the ongoing expansion of large carnivores in several European countries can lead to the recovery of (potentially energetically costly) anti-predator behaviours. In an area recently recolonized by Eurasian lynx Lynx lynx , we snow-tracked radio-collared roe deer Capreolus capreolus in order to investigate their bedding and feeding behaviour during winter, and assess how environmental factors affect their habitat use. We also tested the prediction that roe deer use more open sites than locally available in areas with a stalking predator such as the lynx. Our results showed that both bed sites and foraging sites had more cover, compared with random sites. Most of the variation in canopy cover and in the distance and foraging sites between bed sites and foraging sites was explained by prevailing weather. As the winter progressed, the presumed depletion of fat reserves promoted the use of more canopy cover at foraging sites by night, less by day and a decrease in the distance between beds, foraging sites and human activities. Males used artificial feeding sites less often and bedded further from humans than females. The data fit the hypothesis of tighter energy budgets for family groups (females with fawns) or that males are more cautious towards humans. There was no support for the hypothesis that roe deer used more open habitat than locally available in order to reduce their vulnerability to lynx predation. Owing to severe winter conditions and the danger of starvation, roe deer seem to be forced to accept a high risk when predators are present, not changing their main pattern of habitat use from comparative areas where predators are absent.     

Habitat use under predation risk: hunting, roads and human dwellings influence the spatial behaviour of roe deer

Wildlife populations are subjected to increasing pressure linked to human activities, which introduce multiple stressors. Recently, in addition to direct effects, it has been shown that indirect (non-lethal) effects of predation risk are predominant in many populations. Predation risk is often structured in space and time, generating a heterogeneous “landscape of fear” within which animals can minimize risks by modifying their habitat use. Furthermore, for ungulates, resource quality seems to be positively correlated with human-related sources of risk. We studied the trade-off between access to resources of high-quality and risk-taking by contrasting habitat use of roe deer during daytime with that during nighttime for 94 roe deer in a hunted population. Our first hypothesis was that roe deer should avoid human disturbance by modifying their habitat use during daytime compared to nighttime. Our results supported this, as roe deer mainly used open fields during nighttime, but used more forested habitats during daytime, when human disturbance is higher. Moreover, we found that diel patterns in habitat use were influenced by hunting disturbance. Indeed, the roe deer decreased their use of high-crops during daytime, an important source of cover and food, during the hunting season. The proximity of roads and dwellings also affected habitat use, since roe deer used open fields during daytime to a greater extent when the distance to these sources of disturbance was higher. Hence, our results suggest that roe deer resolve the trade-off between the acquisition of high-quality resources and risk avoidance by modifying their habitat use between day and night.     

Comparative use of forest habitats by roe deer and moose in a human-modified landscape in southeastern Norway during winter

The negative impact of anthropogenic disturbance and land-use changes on large mammals is generally recognized within conservation biology. In southeastern Norway, both moose (Alces alces) and roe deer (Capreolus capreolus) occur throughout human-modified landscapes, facilitating an interesting comparative study of their habitat use. By using pellet group counts, we looked at the importance of forest structure, vegetation characteristics and human disturbance (e.g., distance to the nearest house, nearest paved road, and nearest edge between field and forest) in shaping the winter distribution of both species at multiple spatial scales, in non-agricultural habitats. Moose occurred more often in areas with higher densities of heather and Vaccinium sp. in the ground layer, and used areas with more open forest structure. The proportion of built-up areas, within a 1,000-m buffer, negatively influenced moose occurrence. Roe deer occurred more often in areas with deciduous trees and patches with juniper and Vaccinium sp. in the ground layer, used areas near roads less, but were significantly associated with areas near the field–forest ecotone. The proportion of built-up areas positively influenced roe deer distribution within a 2,500-m buffer. Roe deer seem to be able to persist in more human-dominated landscapes, possibly due to the availability of field–forest edges providing both high-quality fodder and cover in close proximity. Moose, on the contrary, did not show any preference for areas associated with human disturbance, and their distribution was only associated with patches providing food.     

Plankton predation rates in turbulence: a study of the limitations imposed on a predator with a non-spherical field of sensory perception

This paper presents an extension to previously published work which studied encounter rates of planktonic predators with restricted perception fields, to examine the related problems of prey capture and predation rates. Small-scale turbulence influences planktonic predation in two ways: the extra energy of the flow enhances the number of encounter events between individual predator and prey meso/micro-zooplankton, but it lowers the capture probability (because the time spent by the predator and prey in close proximity is reduced). Typically, an 'encounter' has usually been defined as an event when a potential prey swims (or is advected) to within a distance R of the predator in any direction. However, there is a considerable body of experimental evidence showing that predators perception fields are far from spherical; often they are wedge shaped (e.g. fish larvae), or strongly aligned with the directions of sensory antennae (e.g. copepods); and this is certain to influence optimal predation strategies. This paper presents a theoretical model which for the first time examines the combined problems of both encounter and capture for a predator with a restricted perception field swimming in a turbulent flow. If such a predator adopts a cruising strategy (continuous swimming, possibly with direction changes) the model predictions suggest that predation rates actually vary little with swimming speed, in contrast to predictions made for spherical perception fields. Consequently, cruising predators are predicted to swim at relatively low speeds whilst foraging. However, application of the model to examine the net energy gain of a typical pause-travel predator (the Atlantic cod larva), does predict the existence of an optimal ratio of the length of pauses to time spent swimming (specifically one pause phase to every two travel phases), in line with experimental observations. Kinematic simulations are presented which support these findings.     

Diet of Eurasian lynx, Lynx lynx, in the boreal forest of southeastern Norway: the relative importance of livestock and hares at low roe deer density

The year-round food habits of lynx were studied using radio-telemetry and snow-tracking in the boreal forest of southeastern Norway. The main objectives of the study were to clarify the importance of domestic sheep and small prey species in the diet of lynx in an area with a very low density of roe deer. During the period 1995–1999, we found 193 scats and 358 kills made by lynx. Our results indicate that roe deer were the most common prey species (contributing to 83 and 34% of the biomass consumed in winter and summer, respectively), although a wide range of other species were also found, including mountain hares, tetranoids, red foxes, domestic sheep, wild reindeer, and even moose. Most of the diet was obtained by predation, although we did document several cases of scavenging. Roe deer were more important in the diet in winter than in summer, perhaps because they were easier to locate in winter as they clustered around feeding sites. In summer, domestic sheep and small prey increased in importance. Despite the very low density of roe deer in this study area, lynx seemed to still specialise on them, although domestic sheep did constitute a significant amount to their diet, especially for males and yearlings. However, the contribution of sheep to summer diet was far from that expected if their relative density was considered.     

Spatial determinants of infection risk in a multi‐species avian malaria system

Spatially‐variable processes can be an important element of host–parasite interactions, but their longer term demographic and evolutionary effects depend on the magnitude of variation in space, the scale at which variation occurs and the degree to which such processes are temporally stable. Here, we use multiple years of data from a study of two closely related tit species (Paridae), infected with two congeneric species of avian malaria (Plasmodium), to evaluate the roles of extrinsic and intrinsic factors in driving spatial heterogeneity in infection risk, and to address questions of scale and temporal stability in these vector‐driven host–parasite interactions. We show that the two malaria parasite species exhibit markedly different spatial epidemiology: P. relictum infections are effectively randomly distributed in space, with no temporal consistency, whereas P. circumflexum infections exhibit pronounced spatial structuring that is stable over the six years of this study and similar in both host species. We show that both conspecific and heterospecific host density contribute to elevated infection risk, but that the main determinants of elevated risk of P. circumflexum infection risk are habitat features probably associated with vector distribution and abundance. We discuss the implications of these findings, both for our understanding of the epidemiology of malaria in the wild, but also in terms of the longer‐term evolutionary and demographic consequences that spatially variable parasite‐mediated selection may have on host populations.     

Risk taking by Eurasian lynx (Lynx lynx) in a human-dominated landscape: effects of sex and reproductive status

This study aimed to test how the sex and reproductive status of Eurasian lynx influenced their use of 'attractive sinks'– habitats with high prey density and high mortality risks. Locations of 24 Eurasian lynx Lynx lynx were obtained by radio-telemetry in a mixed forest and agricultural habitat in south-eastern Norway. Roe deer, the major food source of lynx in the study area, occurred at higher densities closer to areas of human activity and infrastructure. Proximity of lynx locations to human activity and infrastructure was used as a risk index because the most common causes of death among Scandinavian lynx were of anthropogenic origin. This study shows that distances from lynx locations to human activity were significantly greater for females with newborn kittens than for males, but this decreased with kitten age. The data suggest that this response to human activity is influenced by the reproductive strategies of males and females, and might explain male-biased human-induced mortality in this study and in carnivores more generally.     

Partial migration in roe deer: migratory and resident tactics are end points of a behavioural gradient determined by ecological factors

Ungulate populations exhibiting partial migration present a unique opportunity to explore the causes of the general phenomenon of migration. The European roe deer Capreolus capreolus is particularly suited for such studies due to a wide distribution range and a high level of ecological plasticity. In this study we undertook a comparative analysis of roe deer GPS location data from a representative set of European ecosystems available within the EURODEER collaborative project. We aimed at evaluating the ecological factors affecting migration tactic (i.e. occurrence) and pattern (i.e. timing, residence time, number of migratory trips). Migration occurrence varied between and within populations and depended on winter severity and topographic variability. Spring migrations were highly synchronous, while the timing of autumn migrations varied widely between regions, individuals and sexes. Overall, roe deer were faithful to their summer ranges, especially males. In the absence of extreme and predictable winter conditions, roe deer seemed to migrate opportunistically, in response to a tradeoff between the costs of residence in spatially separated ranges and the costs of migratory movements. Animals performed numerous trips between winter and summer ranges which depended on factors influencing the costs of movement such as between‐range distance, slope and habitat openness. Our results support the idea that migration encompasses a behavioural continuum, with one‐trip migration and residence as its end points, while commuting and multi‐trip migration with short residence times in seasonal ranges are intermediate tactics. We believe that a full understanding of the variation in tactics of temporal separation in habitat use will provide important insights on migration and the factors that influence its prevalence.     

Fear factor: prey habitat selection and its consequences in a predation risk landscape

Predation risk influences prey use of space. However, little is known about how predation risk influences breeding habitat selection and the fitness consequences of these decisions. The nest sites of central-place foraging predators may spatially anchor predation risk in the landscape. We explored how the spatial dispersion of avian predator nests influenced prey territory location and fitness related measures. We placed 249 nest boxes for migrant pied flycatchers Ficedula hypoleuca , at distances between 10 and 630 m, around seven different sparrowhawk nests Accipiter nisus . After closely monitoring flycatcher nests we found that flycatcher arrival dates, nest box occupation rates and clutch size showed a unimodal relationship with distance from sparrowhawk nests. This relationship suggested an optimal territory location at intermediate distances between 330 and 430 m from sparrowhawk nests. Furthermore, pied flycatcher nestling quantity and quality increased linearly with distance from sparrowhawk nests. These fitness related measures were between 4 and 26% larger in flycatcher nestlings raised far from, relative to those raised nearby, sparrowhawk nests. Our results suggest that breeding sparrowhawk affected both flycatcher habitat selection and reproductive success. We propose that nesting predators create predictable spatial variation in predation risk for both adult prey and possibly their nests, to which prey individuals are able to adaptively respond. Recognising predictable spatial variation in perceived predation risk may be fundamental for a proper understanding of predator-prey interactions and indeed prey species interactions.     

Competition hierarchies among ants and predation by birds jointly determine the strength of multi‐species ant–aphid mutualisms

Protection mutualisms often involve multiple species of protector that vary in quality as mutualists. Because protectors may compete for access to mutualists, concordance between competitive ability and degree of benefit will determine the overall strength of multi‐species mutualisms. We compared the abilities of two similarly sized congener ants as competitors for, and mutualists of pine‐feeding aphids, and how insectivorous birds affected each ant–aphid mutualism. Formica planipilis and F. podzolica were indistinguishable in forager abundance, but the former was 13‐fold more abundant at competition baits and provided 11‐fold more benefits to aphids. These results highlight how, in a single environment, a great ecological distance can exist between two congener ants of similar size. Insectivorous birds disrupted the two mutualisms to a similar extent, reducing aphid and ant abundance by 91% and 39% respectively. Nevertheless, birds had an important influence on the relative benefits of the two ants to aphids: where F. planipilis consistently benefited aphids, F. podzolica only did so in the absence of birds. Consequently, the presence of insectivorous birds and ant species identity jointly determined whether ant–aphid mutualisms occurred in pine canopies and the strength of such interactions. Our study highlights the inter‐relatedness of competition, predation and mutualism, and how competition can serve to strengthen mutualism by filtering inferior mutualists.     

Assessing landscape resistance to roe deer dispersal using fuzzy set theory and multicriteria analysis: a case study in Central Spain

The central Iberian Peninsula has one of the highest densities of roe deer populations in Spain. A new motorway is planned to pass through the middle of the distribution of roe deer, thus making it necessary to conduct a connectivity analysis. A map of resistance to roe deer dispersal movements was obtained based on the literature and expert judgment. Three factors were selected: land use (defined by the ability to hide movements, food source, and degree of naturalness), landforms, and influence due to proximity to elements that increase (such as roads and urban areas) or decrease (water resources and proximity to optimal habitat patches) resistance at the local level. Different combinations of factors derived using the analytical hierarchy and fuzzy logic processes were analysed and compared with the real distribution of the species. More realistic resistance (cost) values were obtained for gamma values close to 0.9. This highlights the greater predominance of the fuzzy sum over the fuzzy product in modelling the cost surface. Better results were obtained in scenarios where the predominant factors were either land use and landforms or land use and proximity to human-modified areas. This indicates that roe deer will readily range far from their optimal patches if the land use provides partial cover. These movements appear to be conditioned by steep terrain. Our case study offers a good example of building a cost resistance matrix to help locate areas where the expansion of the species may be curbed or encouraged.     

Group dynamics and landscape features constrain the exploration of herds in fusion-fission societies: the case of European roe deer

Despite the large number of movement studies, the constraints that grouping imposes on movement decisions remain essentially unexplored, even for highly social species. Such constraints could be key, however, to understanding the dynamics and spatial organisation of species living in group fusion-fission systems. We investigated the winter movements (speed and diffusion coefficient) of groups of free-ranging roe deer (Capreolus capreolus), in an agricultural landscape characterised by a mosaic of food and foodless patches. Most groups were short-lived units that merged and split up frequently during the course of a day. Deer groups decreased their speed and diffusion rate in areas where food patches were abundant, as well as when travelling close to main roads and crest lines and far from forests. While accounting for these behavioural adjustments to habitat features, our study revealed some constraints imposed by group foraging: large groups reached the limit of their diffusion rate faster than small groups. The ability of individuals to move rapidly to new foraging locations following patch depression thus decreases with group size. Our results highlight the importance of considering both habitat heterogeneity and group dynamics when predicting the movements of individuals in group fusion-fission societies. Further, we provide empirical evidence that group cohesion can restrain movement and, therefore, the speed at which group members can explore their environment. When maintaining cohesion reduces foraging gains because of movement constraints, leaving the group may become a fitness-rewarding decision, especially when individuals can join other groups located nearby, which would tend to maintain highly dynamical group fusion-fission systems. Our findings also provide the basis for new hypotheses explaining a broad range of ecological patterns, such as the broader diet and longer residency time reported for larger herbivore groups.     

Survival rates and causes of mortality of roe deer <Emphasis Type="Italic">Capreolus capreolus</Emphasis> in a rural landscape,eastern Poland

Game species in the cultural landscapes of Europe are exposed to several sources of anthropogenic pressure. Whilst hunting is part of coordinated wildlife management, poaching is difficult to quantify, and hence, relatively little is known about its impact on survival patterns of game species. We estimated survival and cause-specific mortality rates of 31 roe deer Capreolus capreolus in the surroundings of the Bia?owie?a Primeval Forest, eastern Poland, from 2010 to 2013. The study area consisted of large, extensively used agricultural fields and a mosaic of pine plantations, which were interspersed with natural forest stands and small agricultural fields and meadows. Roe deer were regularly hunted and the forested part of the study area was traversed by large carnivores. Annual survival rate of all deer was 0.69 and did not differ between seasons. Annual survival rates of females and males in each age class (young and adult) were similar. Poaching was a major source of mortality (6 out of 13 roe deer deaths), and poachers did not prefer any sex or age. Adult survival (0.77) was higher than survival of young individuals (0.53); this trend was stronger when poaching was excluded from the analysis. All cases of poaching were recorded between mid-September and February. Although the two parts of the study area were exposed to a similar degree of anthropogenic pressure, all poached individuals occupied home ranges in the mosaic, where the forest cover in home ranges of roe deer was significantly higher than in the field.     

Diversity and functions of the sheep faecal microbiota: a multi‐omic characterization

Little is currently known on the microbial populations colonizing the sheep large intestine, despite their expected key role in host metabolism, physiology and immunity. This study reports the first characterization of the sheep faecal microbiota composition and functions, obtained through the application of a multi‐omic strategy. An optimized protocol was first devised for DNA extraction and amplification from sheep stool samples. Then, 16S rDNA sequencing, shotgun metagenomics and shotgun metaproteomics were applied to unravel taxonomy, genetic potential and actively expressed functions and pathways respectively. Under a taxonomic perspective, the sheep faecal microbiota appeared globally comparable to that of other ruminants, with Firmicutes being the main phylum. In functional terms, we detected 2097 gene and 441 protein families, finding that the sheep faecal microbiota was primarily involved in catabolism. We investigated carbohydrate transport and degradation activities and identified phylum‐specific pathways, such as methanogenesis for Euryarchaeota and acetogenesis for Firmicutes. Furthermore, our approach enabled the identification of proteins expressed by the eukaryotic component of the microbiota. Taken together, these findings unveil structure and role of the distal gut microbiota in sheep, and open the way to further studies aimed at elucidating its connections with management and dietary variables in sheep farming.     

Living at the edge of the front; reduced predation risk to invasive round goby in a Great Lakes tributary

The objective of this research was to determine if the highly invasive round goby (Neogobius melanostomus) experiences lower predation risk during early stages of invasion. We compared round goby predation rates between a recently invaded area (occupied for ≈1 year) and a longer established area (≈7 years) of the Trent River, Ontario, Canada. Tethering trials were conducted in three habitat types, and comparable habitats in the two areas were similar in water temperature, velocity and depth. Predation rates of tethered round gobies were on average 27% lower in the recently invaded area. Reduced predation in the recently invaded area may be due to the short duration of round goby occupancy and/or differences in predator communities between the two study areas. Data before the round goby invasion suggest that predator communities were similar between the two range areas, but differences in predator abundance cannot be ruled out as a potential mechanism. Other possible mechanisms include a numerical or learned response by predators over time to a novel prey item. Reduced predation rate during the initial stages of invasion may contribute to the fitness of individuals that migrate into areas not previously occupied, and thus facilitate successful range expansion.