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Prey species may display anti-predatory behavior, i.e., flight, increased vigilance, and decreased feeding, in response to the true presence of a predator or to the implied presence of a predator through, e.g., acoustic cues. In this study, we investigated the anti-predatory reactions of moose (

In the last few decades wild boar populations have expanded northwards, colonizing boreal forests. The soil disturbances caused by wild boar rooting may have an impact on soil organisms that play a key role in organic matter turnover. However, the impact of wild boar colonization on boreal forest ecosystems and soil organisms remains largely unknown. We investigated the effect of natural and simulated rooting on decomposer and predatory soil mites (total, adult and juvenile abundances; and adult–juvenile proportion). Our simulated rooting experiment aimed to disentangle the effects of (i) bioturbation due to soil mixing and (ii) removing organic material (wild boar food resources) on soil mites. Our results showed a decline in the abundance of adult soil mites in response to both natural and artificial rooting, while juvenile abundance and the relative proportion of adults and juveniles were not affected. The expansion of wild boar northwards and into new habitats has negative effects on soil decomposer abundances in boreal forests which may cascade through the soil food web ultimately affecting ecosystem processes. Our study also suggests that a combined use of natural and controlled experimental approaches is the way forward to reveal any subtle interaction between aboveground and belowground organisms and the ecosystem functions they drive.

Little is known about whether female mating tactics vary with age based on their preference for mates. To fill this knowledge gap, we examined how maternal age is related to the age of their mates using detailed individual long-term monitoring of a genotyped and pedigreed European roe deer (Capreolus capreolus Linnaeus, 1758) population. We found that mating between old females and prime-aged males was more frequent than mating between prime-aged females and prime-aged males. This suggests that old females avoid old mates. Old females might be more selective in their mate choice than prime-aged females owing to increased mate-sampling effort. Our finding is in line with the terminal investment/allocation hypothesis. The study of age-related variation in female mating behaviour is particularly important because this behaviour can influence the intensity and direction of sexual selection and the maintenance of variation in male sexually selected traits. Further studies are needed to quantify the exact fitness benefits of age-specific mating tactics in females.

The taiga tick, Ixodes persulcatus, has previously been limited to eastern Europe and northern Asia, but recently its range has expanded to Finland and northern Sweden. The species is of medical importance, as it, along with a string of other pathogens, may carry the Siberian and Far Eastern subtypes of tick-borne encephalitis virus. These subtypes appear to cause more severe disease, with higher fatality rates than the central European subtype. Until recently, the meadow tick, Dermacentor reticulatus, has been absent from Scandinavia, but has now been detected in Denmark, Norway and Sweden. Dermacentor reticulatus carries, along with other pathogens, Babesia canis and Rickettsia raoultii. Babesia canis causes severe and often fatal canine babesiosis, and R. raoultii may cause disease in humans. We collected 600 tick nymphs from each of 50 randomly selected sites in Denmark, southern Norway and south-eastern Sweden in August–September 2016. We tested pools of 10 nymphs in a Fluidigm real time PCR chip to screen for I. persulcatus and D. reticulatus, as well as tick-borne pathogens. Of all the 30,000 nymphs tested, none were I. persulcatus or D. reticulatus. Our results suggest that I. persulcatus is still limited to the northern parts of Sweden, and have not expanded into southern parts of Scandinavia. According to literature reports and supported by our screening results, D. reticulatus may yet only be an occasional guest in Scandinavia without established populations.

Deepened knowledge on response of biota and ecological processes following fire is essential for a future with warmer climate and more disturbances. In 2014 the first mega-fire (13,100 ha) for at least a century in Scandinavia hit south-central Sweden, in a production forest landscape shaped by clearcutting forestry. Ecological dynamics is followed in >20 projects from universities, authorities and citizen science initiatives, rapidly accumulating substantial amounts of data. We outline projects and summarize their results during the first four years, demonstrating a rapid succession of fungi, lichens, vascular plants, birds, mammals, ticks, butterflies, beetles, and drastically altered carbon dynamics. We characterize forest operations including regeneration measures and point to patterns in pest and pathogen infestations. 8,000 ha is set aside for natural succession, with the rest harvested and managed for forest production, offering excellent opportunities for studies on salvage logging effects, already evident for birds. We demonstrate a strong regrowth of deciduous trees, and the protected part will in some decades likely develop into the largest deciduous-dominated area in boreal north Europe outside Russia. Continued studies of biodiversity and ecological processes are urgent for this unique area.

Browsing can reduce forest productivity, particularly when the apical shoots of trees are damaged. Repellents are used widely to reduce browsing, but application is costly. To improve efficiency, it may be possible to take advantage of associational plant refuge effects, requiring repellents to be applied only to some trees or parts of trees, or reapplied less frequently. Using captive moose (Alces alces) and constructed stands of Scots pine (Pinus sylvestris), we tested for potential refuges by applying a commercial repellent (HaTe2) to all, alternate or none of the apical shoots, or all of the previous-year apical shoots. We also tested for potential refuges under field conditions, applying the repellent to all, alternate or none of the apical pine shoots in forest stands. Captive moose (two individuals in a 2.1-ha enclosure, ~ 95 individuals km−2) browsed 100% of trees, but were significantly less likely to browse apical shoots treated with repellent. Associational refuge was ineffective both within and between trees. In the field (0.84 moose km−2), only 1.3% of trees sustained browsing damage. Applying the repellent to the apical shoots of pines had no direct repellent effect nor any within-plant associational effects. Trees with treated apical shoots provided some protection for untreated neighbouring trees, but this was not biologically meaningful given the low percentage of trees browsed overall. Here, a simple captive experiment was not predictive of the browsing response observed in the field, demonstrating the need to test repellent application strategies in situ.

On an evolutionary time scale, competition for food drives species formation by genetic adaptations to the environment and subsequent niche separation. On a short-term scale, animals use different strategies to meet their nutritional requirements, which ultimately influence their fitness. Understanding these adaptations in herbivores is especially important in temperate climates where animals have adapted both physiologically and behaviorally to seasonal variations in order to meet their nutritional requirements. The aim of this project was to investigate temporal variation in chemical composition of rumen content between two coexisting species of large herbivores, the native roe deer (Capreolus capreolus L.) and the introduced fallow deer (Dama dama L.), as well as a potential effect of competition on niche separation (interspecific differences in rumen nutrient composition). We analyzed 345 rumen samples collected from animals at one 95 km2 estate, Koberg, in southwestern Sweden. Based on samples from all seasons, temporal variation in nutrient composition and interspecific differences between the two deer species were investigated under two contrasting fallow deer population densities. Results revealed that nutrient composition varied between species and across seasons. Roe deer had a higher proportion of rumen protein compared to fallow deer, with the highest proportions in spring. In contrast, fallow deer had a higher proportion of rumen hemicellulose compared to roe deer in spring, while no differences in nutrient composition between species could be found in fall. Overall, there were greater differences between the two species when fallow deer density was high and competition likely more pronounced than when fallow deer density was low. The results from this study can be used to understand interspecific competition and how it fosters niche separation between coexisting large herbivores.

International audience; Evolution should favour plasticity in dispersal decisions in response to spatial heterogeneity in social and environmental contexts. Sex differences in individual optimization of dispersal decisions are poorly documented in mammals, because species where both sexes commonly disperse are rare. To elucidate the sex-specific drivers governing dispersal, we investigated sex differences in condition dependence in the propensity and distance of natal dispersal in one such species, the roe deer, using fine-scale monitoring of 146 GPS-collared juveniles in an intensively monitored population in southwest France. Dispersal propensity increased with body mass in males such that 36% of light individuals dispersed, whereas 62% of heavy individuals did so, but there was no evidence for condition dependence in dispersal propensity among females. By contrast, dispersal distance increased with body mass at a similar rate in both sexes such that heavy dispersers travelled around twice as far as light dispersers. Sex differences in the strength of condition-dependent dispersal may result from different selection pressures acting on the behaviour of males and females. We suggest that females disperse prior to habitat saturation being reached, likely in relation to the risk of inbreeding. By contrast, natal dispersal in males is likely governed by competitive exclusion through male–male competition for breeding opportunities in this strongly territorial mammal. Our study is, to our knowledge, a first demonstration that condition dependence in dispersal propensity and dispersal distance may be decoupled, indicating contrasting selection pressures drive the behavioural decisions of whether or not to leave the natal range, and where to settle.

ObjectiveOur aim was to analyse the chromogranin A-derived peptides vasostatin and catestatin in serum from wild boar (Sus scrofa) captured in a corral trap. Acute capture-related stress quickly leads to a release of adrenalin and noradrenalin, but these hormones have a short half-life in blood and are difficult to measure. Chromogranin A (CgA), a glycoprotein which is co-released with noradrenalin and adrenalin, is relatively stable in circulation and the CgA-derived peptides catestatin and vasostatin have been measured in domestic species, but not yet in wildlife.ResultsVasostatin and catestatin could be measured and the median (range) serum concentrations were 0.91 (0.54–2.86) and 0.65 (0.35–2.62) nmol/L, respectively. We conclude that the CgA-derived peptides vasostatin and catestatin can be measured in wild boar serum and may thus be useful as biomarkers of psychophysical stress.

Ticks carry pathogens that can cause disease in both animals and humans, and there is a need to monitor the distribution and abundance of ticks and the pathogens they carry to pinpoint potential high risk areas for tick-borne disease transmission. In a joint Scandinavian study, we measured Ixodes ricinus instar abundance at 159 sites in southern Scandinavia in August-September, 2016, and collected 29,440 tick nymphs at 50 of these sites. We additionally measured abundance at 30 sites in August-September, 2017. We tested the 29,440 tick nymphs in pools of 10 in a Fluidigm real-time PCR chip to screen for 17 different tick-associated pathogens, 2 pathogen groups and 3 tick species. We present data on the geolocation, habitat type and instar abundance of the surveyed sites, as well as presence/absence of each pathogen in all analysed pools from the 50 collection sites and individual prevalence for each site. These data can be used alone or in combination with other data for predictive modelling and mapping of high-risk areas., Measurement(s) pathogen • Ixodida • Abundance • tick instar abundance Technology Type(s) real-time PCR • Manual Count Factor Type(s) geographic location • time of data collection Sample Characteristic - Organism Ixodes ricinus Sample Characteristic - Environment forested area • meadow ecosystem Kingdom of Denmark • Kingdom of Norway • Sweden Machine-accessible metadata file describing the reported data: 10.6084/m9.figshare.12523133

Tick-borne pathogens cause diseases in animals and humans, and tick-borne disease incidence is increasing in many parts of the world. There is a need to assess the distribution of tick-borne pathogens and identify potential risk areas. We collected 29,440 tick nymphs from 50 sites in Scandinavia from August to September, 2016. We tested ticks in a real-time PCR chip, screening for 19 vector-associated pathogens. We analysed spatial patterns, mapped the prevalence of each pathogen and used machine learning algorithms and environmental variables to develop predictive prevalence models. All 50 sites had a pool prevalence of at least 33% for one or more pathogens, the most prevalent being Borrelia afzelii, B. garinii, Rickettsia helvetica, Anaplasma phagocytophilum, and Neoehrlichia mikurensis. There were large differences in pathogen prevalence between sites, but we identified only limited geographical clustering. The prevalence models performed poorly, with only models for R. helvetica and N. mikurensis having moderate predictive power (normalized RMSE from 0.74–0.75, R2 from 0.43–0.48). The poor performance of the majority of our prevalence models suggest that the used environmental and climatic variables alone do not explain pathogen prevalence patterns in Scandinavia, although previously the same variables successfully predicted spatial patterns of ticks in the same area. Tick-borne pathogens cause diseases in animals and humans, and tick-borne disease incidence is increasing in many parts of the world. There is a need to assess the distribution of tick-borne pathogens and identify potential risk areas. We collected 29,440 tick nymphs from 50 sites in Scandinavia from August to September, 2016. We tested ticks in a real-time PCR chip, screening for 19 vector-associated pathogens. We analysed spatial patterns, mapped the prevalence of each pathogen and used machine learning algorithms and environmental variables to develop predictive prevalence models. All 50 sites had a pool prevalence of at least 33% for one or more pathogens, the most prevalent being Borrelia afzelii, B. garinii, Rickettsia helvetica, Anaplasma phagocytophilum, and Neoehrlichia mikurensis. There were large differences in pathogen prevalence between sites, but we identified only limited geographical clustering. The prevalence models performed poorly, with only models for R. helvetica and N. mikurensis having moderate predictive power (normalized RMSE from 0.74–0.75, R2 from 0.43–0.48). The poor performance of the majority of our prevalence models suggest that the used environmental and climatic variables alone do not explain pathogen prevalence patterns in Scandinavia, although previously the same variables successfully predicted spatial patterns of ticks in the same area.

ticks carry pathogens that can cause disease in both animals and humans, and there is a need to monitor the distribution and abundance of ticks and the pathogens they carry to pinpoint potential high risk areas for tick-borne disease transmission. In a joint Scandinavian study, we measured Ixodes ricinus instar abundance at 159 sites in southern Scandinavia in August-September, 2016, and collected 29,440 tick nymphs at 50 of these sites. We additionally measured abundance at 30 sites in August-September, 2017. We tested the 29,440 tick nymphs in pools of 10 in a Fluidigm real-time PCR chip to screen for 17 different tick-associated pathogens, 2 pathogen groups and 3 tick species. We present data on the geolocation, habitat type and instar abundance of the surveyed sites, as well as presence/absence of each pathogen in all analysed pools from the 50 collection sites and individual prevalence for each site. these data can be used alone or in combination with other data for predictive modelling and mapping of high-risk areas. Ticks carry pathogens that can cause disease in both animals and humans, and there is a need to monitor the distribution and abundance of ticks and the pathogens they carry to pinpoint potential high risk areas for tick-borne disease transmission. In a joint Scandinavian study, we measured Ixodes ricinus instar abundance at 159 sites in southern Scandinavia in August-September, 2016, and collected 29,440 tick nymphs at 50 of these sites. We additionally measured abundance at 30 sites in August-September, 2017. We tested the 29,440 tick nymphs in pools of 10 in a Fluidigm real-time PCR chip to screen for 17 different tick-associated pathogens, 2 pathogen groups and 3 tick species. We present data on the geolocation, habitat type and instar abundance of the surveyed sites, as well as presence/absence of each pathogen in all analysed pools from the 50 collection sites and individual prevalence for each site. These data can be used alone or in combination with other data for predictive modelling and mapping of high-risk areas.

Cloth-dragging is the most widely-used method for collecting and counting ticks, but there are few studies of its reliability. By using cloth-dragging, we applied a replicated line transects survey method, in two areas in Sweden with different Ixodes ricinus tick-densities (low at Grimsö and high at Bogesund) to evaluate developmental stage specific repeatability, agreement and precision in estimates of tick abundance. ‘Repeatability’ was expressed as the Intraclass Correlation Coefficient (ICC), ‘agreement’ with the Total Deviation Index (TDI) and ‘precision’ by the coefficient of variation (CV) for a given dragging distance. Repeatability (ICC) and agreement (TDI) were higher for the most abundant instar (nymphs) and in the area of higher abundance. At Bogesund tick counts were higher than at Grimsö and so also repeatability, with fair to substantial ICC estimates between 0.22 and 0.75, and TDI ranged between 1 and 44.5 counts of difference (thus high to moderate agreement). At Grimsö, ICC was poor to moderate and ranged between 0 and 0.59, whereas TDI remained low with estimates lower or equal to 1 count (thus high agreement). Despite a 100-fold lower abundance at Grimsö, the same level of precision for nymphs could be achieved with a 70% increase of dragging effort. We conclude that the cloth-dragging technique is useful for surveying ticks’ and primarily to estimate abundance of the nymphal stage, whereas it rarely will be recommended for larvae and adults.

Background Wildlife traps are used in many countries without evaluation of their effect on animal welfare. Trap-capture of wild animals should minimise negative effects on animal welfare, irrespective of whether the animals are trapped for hunting, research, or management purposes. Live-trap capture of wild boar (Sus scrofa) followed by killing inside the trap by gunshot is a recently introduced but disputed hunting method in Sweden. Approval of trap constructions is based on gross necropsy findings of 20 trapped and shot wild boars. For improved animal welfare evaluation, our aim was to study wild boar behaviour during live-trapping in a 16 m2 square corral-style trap. Behavioural assessments were conducted after filming 12 capture events of in total 38 wild boars (five adults, 20 subadults, 13 piglets). Selected behavioural traits were compared with pathological changes (trap-related lesions) found at necropsy of the 20 subadults, to determine if these variables were useful proxies of capture-induced stress in wild boar. Results The wild boars spent less time resting in the evening than in the night and morning. Using Friedman’s ANOVA, there was an overall difference in the time spent foraging. However, we only found a difference between the evening and morning in the Wilcoxon matched pairs test after the Sequential Bonferroni correction, where the wild boars spent more time foraging in the evening than in the morning. Single captured individuals showed more escape behaviours and reacted more strongly to external stimuli than individuals captured in a group. It was more common for animals to charge against the mesh walls of the trap upon human approach compared to upon initial capture when the trap door closed. Trap-related pathological findings due to trauma were documented in 13 of the 20 subadults that were necropsied. Behavioural alterations indicative of capture-induced stress (e.g. charging into the trap walls) were documented in trapped wild boars with no or minor physical injuries (e.g. skin abrasions, subcutaneous haemorrhage). Conclusions Behavioural assessment provided valuable information for determination of capture-induced stress in wild boar when evaluating live-trapping in a corral-style trap, whereas pathological evaluation through necropsy did not fully reflect the animal welfare aspects of live-trapping. We emphasize the inclusion of species-specific behavioural data assessment for evaluation of capture-related stress during live-trapping and for testing of new trap constructions before approval.

Summary Animals exhibit a diversity of movement tactics [ 1 ]. Tracking resources that change across space and time is predicted to be a fundamental driver of animal movement [ 2 ]. For example, some migratory ungulates (i.e., hooved mammals) closely track the progression of highly nutritious plant green-up, a phenomenon called “green-wave surfing” [ 3 , 4 , 5 ]. Yet general principles describing how the dynamic nature of resources determine movement tactics are lacking [ 6 ]. We tested an emerging theory that predicts surfing and the existence of migratory behavior will be favored in environments where green-up is fleeting and moves sequentially across large landscapes (i.e., wave-like green-up) [ 7 ]. Landscapes exhibiting wave-like patterns of green-up facilitated surfing and explained the existence of migratory behavior across 61 populations of four ungulate species on two continents (n = 1,696 individuals). At the species level, foraging benefits were equivalent between tactics, suggesting that each movement tactic is fine-tuned to local patterns of plant phenology. For decades, ecologists have sought to understand how animals move to select habitat, commonly defining habitat as a set of static patches [ 8 , 9 ]. Our findings indicate that animal movement tactics emerge as a function of the flux of resources across space and time, underscoring the need to redefine habitat to include its dynamic attributes. As global habitats continue to be modified by anthropogenic disturbance and climate change [ 10 ], our synthesis provides a generalizable framework to understand how animal movement will be influenced by altered patterns of resource phenology.

Faced with rapid environmental changes, individuals may express different magnitude and plasticity in their response to a given stressor. However, little is known about the causes of variation in phenotypic plasticity of the stress response in wild populations. In the present study, we repeatedly captured individual roe deer (Capreolus capreolus) from two wild populations in Sweden exposed to differing levels of predation pressure and measured plasma concentrations of stress-induced cortisol and behavioral docility. While controlling for the marked effects of habituation, we found clear between-population differences in the stress-induced cortisol response. Roe deer living in the area that was recently recolonized by lynx (Lynx lynx) and wolves (Canis lupus) expressed cortisol levels that were around 30% higher than roe deer in the human-dominated landscape free of large carnivores. In addition, for the first time to our knowledge, we investigated the stress-induced cortisol response in free-ranging newborn fawns and found no evidence for hypo-responsiveness during early life in this species. Indeed, stress-induced cortisol levels were of similar magnitude and differed between populations to a similar extent in both neonates and adults. Finally, at an individual level, we found that both cortisol and docility levels were strongly repeatable, and weakly negatively inter-correlated, suggesting that individuals differed consistently in how they respond to a stressor, and supporting the existence of a stress-management syndrome in roe deer. Electronic supplementary material The online version of this article (10.1007/s00442-018-4174-7) contains supplementary material, which is available to authorized users.

Much research on large herbivore movement has focused on the annual scale to distinguish between resident and migratory tactics, commonly assuming that individuals are sedentary at the within-season scale. However, apparently sedentary animals may occupy a number of sub-seasonal functional home ranges (sfHR), particularly when the environment is spatially heterogeneous and/or temporally unpredictable. The roe deer (Capreolus capreolus) experiences sharply contrasting environmental conditions due to its widespread distribution, but appears markedly sedentary over much of its range. Using GPS monitoring from 15 populations across Europe, we evaluated the propensity of this large herbivore to be truly sedentary at the seasonal scale in relation to variation in environmental conditions. We studied movement using net square displacement to identify the possible use of sfHR. We expected that roe deer should be less sedentary within seasons in heterogeneous and unpredictable environments, while migratory individuals should be seasonally more sedentary than residents. Our analyses revealed that, across the 15 populations, all individuals adopted a multi-range tactic, occupying between two and nine sfHR during a given season. In addition, we showed that (i) the number of sfHR was only marginally influenced by variation in resource distribution, but decreased with increasing sfHR size; and (ii) the distance between sfHR increased with increasing heterogeneity and predictability in resource distribution, as well as with increasing sfHR size. We suggest that the multirange tactic is likely widespread among large herbivores, allowing animals to track spatio-temporal variation in resource distribution and, thereby, to cope with changes in their local environment. Migration · Residency · Sub-seasonal functional home range · Plasticity · Roe deer

Herbivory pressure on a forest stand and each plant individual may be affected by the spatial distribution of conspecific and allospecific plants on the site; whether the plants are standing in solitary or groupwise settings; or by the differences in the preference of plants in relation to each other. The study was conducted in southwest Sweden, where 120 plots (1508 m2) were visited and 1280 individual woody plants evaluated and measured. We tested the hypothesis that preferred woody plants can protect unpreferred conspecific and allospecific woody species against herbivory in a system with one dominant, generalist herbivore, the fallow deer (Dama L.), and that the effectiveness of this protection varies depending on the relative preference of the neighboring conspecific and allospecific woody species placed in conspecific groups or standing solitary. Our results support the hypothesis that preferred woody plants can protect unpreferred conspecific and allospecific woody species against herbivory...

Humans, as super predators, can have strong effects on wildlife behaviour, including profound modifications of diel activity patterns. Subsequent to the return of large carnivores to human-modified ecosystems, many prey species have adjusted their spatial behaviour to the contrasting landscapes of fear generated by both their natural predators and anthropogenic pressures. The effects of predation risk on temporal shifts in diel activity of prey, however, remain largely unexplored in human-dominated landscapes. We investigated the influence of the density of lynx Lynx lynx, a nocturnal predator, on the diel activity patterns of their main prey, the roe deer Capreolus capreolus, across a gradient of human disturbance and hunting at the European scale. Based on 11 million activity records from 431 individually GPS-monitored roe deer in 12 populations within the EURODEER network (http://eurodeer.org), we investigated how lynx predation risk in combination with both lethal and non-lethal human activities affected the diurnality of deer. We demonstrated marked plasticity in roe deer diel activity patterns in response to spatio-temporal variations in risk, mostly due to human activities. In particular, roe deer decreased their level of diurnality by a factor of 1.37 when the background level of general human disturbance was high. Hunting exacerbated this effect, as during the hunting season deer switched most of their activity to night-time and, to a lesser extent, to dawn, although this pattern varied noticeably in relation to lynx density. Indeed, in the presence of lynx, their main natural predator, roe deer were relatively more diurnal. Overall, our results revealed a strong influence of human activities and the presence of lynx on diel shifts in roe deer activity. In the context of the recovery of large carnivores across Europe, we provide important insights about the effects of predators on the behavioural responses of their prey in human-dominated ecosystems. Modifications in the temporal partitioning of ungulate activity as a response to human activities may facilitate human-wildlife coexistence, but likely also have knock-on effects for predator-prey interactions, with cascading effects on ecosystem functioning.Résumé Les humains, en tant que ‘super-prédateurs’, peuvent avoir des effets importants sur le comportement de la faune sauvage, y compris des modifications profondes de leurs rythmes circadiens d'activité. A la suite du retour des grands carnivores dans les écosystèmes anthropisés, de nombreuses espèces proies ont ajusté leur comportement spatial à ces paysages de la peur contrastés, générés à la fois par les pressions liées aux risques anthropiques et à la présence de leurs prédateurs naturels. Les effets du risque de prédation sur les modifications temporelles des rythmes circadiens d'activité des proies restent cependant largement inconnus dans les écosystèmes dominés par l'homme. Ici, nous avons étudié l'influence de la densité de lynx Lynx lynx, un prédateur nocturne, sur les rythmes circadiens d'activité de leur proie principale, le chevreuil Capreolus capreolus, à travers un gradient de pressions anthropiques à l’échelle Européenne. Sur la base de plus de 11 million de données d'activité issues de 431 suivis individuels de chevreuils équipés de colliers GPS provenant de 12 populations au sein du réseau EURODEER (http://eurodeer.org), nous avons analysé comment le risque de prédation par le lynx, associé aux risques létaux et non-létaux des activités humaines, influence la diurnalité des chevreuils. Nous avons démontré une forte plasticité des rythmes circadiens d'activité des chevreuils en réponse aux variations spatio-temporelles du risque, et notamment face aux activités humaines. Plus particulièrement, les chevreuils diminuent leur degré de diurnalité d'un facteur de 1.37 lorsque le dérangement humain est important. La chasse accentue cet effet, puisque durant la saison de chasse les chevreuils basculent la plupart de leur activité de nuit, et dans une moindre mesure, durant l'aube également, bien que ce patron soit essentiellement variable en fonction de la densité de lynx. En effet, en présence de lynx, leur principal prédateur, les chevreuils sont relativement plus diurnes. Globalement, nos résultats révèlent une forte influence des activités humaines et de la présence de lynx sur l'ajustement des rythmes circadiens d'activité des chevreuils. Dans le contexte du retour des grands carnivores en Europe, notre étude apporte de nouvelles connaissances sur les effets des prédateurs sur la réponse comportementale de leur proie dans des écosystèmes anthropisés. La modification de la répartition temporelle de l'activité des ongulés en réponse aux activités humaines pourrait être un facteur facilitant la coexistence homme-faune sauvage, avec toutefois des conséquences autres sur les interactions prédateurs-proies et leurs effets en cascade sur le fonctionnement des écosystèmes.

Background Tick-borne diseases have become increasingly common in recent decades and present a health problem in many parts of Europe. Control and prevention of these diseases require a better understanding of vector distribution. Aim Our aim was to create a model able to predict the distribution of Ixodes ricinus nymphs in southern Scandinavia and to assess how this relates to risk of human exposure. Methods We measured the presence of I. ricinus tick nymphs at 159 stratified random lowland forest and meadow sites in Denmark, Norway and Sweden by dragging 400 m transects from August to September 2016, representing a total distance of 63.6 km. Using climate and remote sensing environmental data and boosted regression tree modelling, we predicted the overall spatial distribution of I. ricinus nymphs in Scandinavia. To assess the potential public health impact, we combined the predicted tick distribution with human density maps to determine the proportion of people at risk. Results Our model predicted the spatial distribution of I. ricinus nymphs with a sensitivity of 91% and a specificity of 60%. Temperature was one of the main drivers in the model followed by vegetation cover. Nymphs were restricted to only 17.5% of the modelled area but, respectively, 73.5%, 67.1% and 78.8% of the human populations lived within 5 km of these areas in Denmark, Norway and Sweden. Conclusion The model suggests that increasing temperatures in the future may expand tick distribution geographically in northern Europe, but this may only affect a small additional proportion of the human population.

The most common framework under which ungulate migration is studied predicts that it is driven by spatio–temporal variation in plant phenology, yet other hypotheses may explain differences within and between species. To disentangle more complex patterns than those based on single species/ single populations, we quantified migration vari-ability using two sympatric ungulate species differing in their foraging strategy, mating system and physiological constraints due to body size. We related observed variation to a set of hypotheses. We used GPS-collar data from 537 individuals in 10 roe Capreolus capreolus and 12 red deer Cervus elaphus populations spanning environmental gra-dients across Europe to assess variation in migration propensity, distance and tim-ing. Using time-to-event models, we explored how the probability of migration varied in relation to sex, landscape (e.g. topography, forest cover) and temporally-varying environmental factors (e.g. plant green-up, snow cover). Migration propensity varied across study areas. Red deer were, on average, three times more migratory than roe deer (56% versus 18%). This relationship was mainly driven by red deer males which were twice as migratory as females (82% versus 38%). The probability of roe deer migration was similar between sexes. Roe deer (both sexes) migrated earliest in spring. While territorial male roe deer migrated last in autumn, male and female red deer migrated around the same time in autumn, likely due to their polygynous mating system. Plant productivity determined the onset of spring migration in both species, but if plant productivity on winter ranges was sufficiently high, roe deer were less likely to leave. In autumn, migration coincided with reduced plant productivity for both species. This relationship was stronger for red deer. Our results confirm that ungulate migration is influenced by plant phenology, but in a novel way, that these effects appear to be modulated by species-specific traits, especially mating strategies.

Background Tick-borne diseases have become increasingly common in recent decades and present a health problem in many parts of Europe. Control and prevention of these diseases require a better understanding of vector distribution. Aim Our aim was to create a model able to predict the distribution of Ixodes ricinus nymphs in southern Scandinavia and to assess how this relates to risk of human exposure. Methods We measured the presence of I. ricinus tick nymphs at 159 stratified random lowland forest and meadow sites in Denmark, Norway and Sweden by dragging 400 m transects from August to September 2016, representing a total distance of 63.6 km. Using climate and remote sensing environmental data and boosted regression tree modelling, we predicted the overall spatial distribution of I. ricinus nymphs in Scandinavia. To assess the potential public health impact, we combined the predicted tick distribution with human density maps to determine the proportion of people at risk. Results Our model predicted the spatial distribution of I. ricinus nymphs with a sensitivity of 91% and a specificity of 60%. Temperature was one of the main drivers in the model followed by vegetation cover. Nymphs were restricted to only 17.5% of the modelled area but, respectively, 73.5%, 67.1% and 78.8% of the human populations lived within 5 km of these areas in Denmark, Norway and Sweden. Conclusion The model suggests that increasing temperatures in the future may expand tick distribution geographically in northern Europe, but this may only affect a small additional proportion of the human population.

Recently, focus on tick-borne diseases has increased as ticks and their pathogens have become widespread and represent a health problem in Europe. Understanding the epidemiology of tick-borne infections requires the ability to predict and map tick abundance. We measured Ixodes ricinus abundance at 159 sites in southern Scandinavia from August-September, 2016. We used field data and environmental variables to develop predictive abundance models using machine learning algorithms, and also tested these models on 2017 data. Larva and nymph abundance models had relatively high predictive power (normalized RMSE from 0.65–0.69, R2 from 0.52–0.58) whereas adult tick models performed poorly (normalized RMSE from 0.94–0.96, R2 from 0.04–0.10). Testing the models on 2017 data produced good results with normalized RMSE values from 0.59–1.13 and R2 from 0.18–0.69. The resulting 2016 maps corresponded well with known tick abundance and distribution in Scandinavia. The models were highly influenced by temperature and vegetation, indicating that climate may be an important driver of I. ricinus distribution and abundance in Scandinavia. Despite varying results, the models predicted abundance in 2017 with high accuracy. The models are a first step towards environmentally driven tick abundance models that can assist in determining risk areas and interpreting human incidence data.

Landscapes composed of agricultural land mixed with forest are desirable since they provide a wide range of diversified ecosystem services, unlike specialized agricultural landscapes, but that creates a trade-off between these land uses since wildlife usually feed on crops and reduce yields. In Nordic countries, where human population density is low and game hunting can be a viable economic alternative, mixed landscape systems are particularly interesting. To evaluate the economic sustainability of such systems we need to quantify wildlife damage to crops. One important species, being popular among Swedish hunters and therefore economically valuable, is fallow deer (Dama dama). Our objective was to evaluate the economic sustainability of mixed landscape systems including cultivated fields and commercial hunting of fallow deer. We studied the effects of excluding fallow deer by using 86 exclosures and adjacent plots in winter wheat and oat fields in south-west Sweden. We analyzed yield losses and interactions between spatial and temporal grazing patterns, anthropogenic landscape features, and topological characteristics of the landscape. We found that animals avoided exposed spots, irrespective of distance from human activity. We also found a seasonal grazing pattern related to the different growing periods of winter wheat (more grazed, emerging in autumn) and spring oat (less grazed, emerging in spring). We then compared the costs of crop damage against the commercial value of fallow deer hunting. The damage amounted to 375 ±196 € ha-1 for wheat and 152 ±138 € ha-1 for oat, corresponding to a total cost per animal of 82.7 ±81.0 €, while each animal had an estimated market value of approximately 100 €. Therefore the value of fallow deer presence compensated for the associated cost of crop damage. Profit could be further improved in this case by adopting additional management strategies. In general our study confirmed the economic feasibility of this particular mixed land management.

Animal welfare concerns are becoming a central issue in wildlife management and conservation. Thus, we investigated stress response of wild ungulates to potentially traumatic situations (shooting injuries, vehicle collisions, entanglement, injuries or diseases) and hunting methods (stalking, battues and hunts with dogs) by means of serum cortisol concentrations from blood collected from killed animals. Cortisol levels in roe deer ranged below and in wild boar above levels for moose, red deer and fallow deer (hence, pooled as a group “deer”). Apart from species, cortisol concentration in trauma situations was mainly explained by trauma type and presence of disturbance after the trauma event. Effect of trauma type differed significantly for “deer”, with animals caught in fences and suffering vehicle collisions experiencing higher cortisol levels than animals injured by shooting. Differences between hunting methods were observed in the cervids (“deer” and roe deer), with stalking leading to lower cortisol levels than hunts with dogs (both groups) and battues (roe deer). Events both before and after the shot, such as duration of pursuit prior to shooting, location of injury, trauma length and presence of disturbance after the shot were relevant for cortisol levels in hunted cervids. Our results indicate that search teams tracking and euthanising wounded animals should behave in a calm way to minimise disturbance. Still, it is important to acknowledge that many situations described in the literature, i.e. reindeer handling, roe deer captures and red deer yarding, seem even more stressful, beside vehicle collisions, than most hunting methods.

Background Understanding the genetic and environmental mechanisms governing variation in morphology or phenology in wild populations is currently an important challenge. While there is a general consensus that selection is stronger under stressful conditions, it remains unclear whether the evolutionary potential of traits should increase or decrease with increasingly stressful conditions. Here, we investigate how contrasting environmental conditions during growth may affect the maternal and genetic components of body mass in roe deer, the most abundant and widespread wild ungulate in Western Europe. Body mass is a key life history trait that strongly influences both survival and reproductive performance in large herbivores. We used pedigrees and animal models to determine the variance components of juvenile and adult winter body mass in two populations experiencing contrasting early-life conditions. Results Our analyses showed that roe deer at Chizé, where habitat was poor and unpredictable, exhibited very low genetic variance in juvenile body mass. Instead, variance in mass was mainly driven by among-cohort differences in early-life conditions and maternal environment. In contrast, roe deer at Bogesund, where resource availability during the critical period of fawn rearing was higher, displayed a substantial level of genetic variance in body mass. We discuss the potential role of past demography and viability selection on fawn body mass on the erosion of genetic variance in the poor habitat. Conclusions Our study highlights the importance of accounting for both spatial (i.e. between-population variation) and temporal (i.e. cohort variation) heterogeneity in environmental conditions, especially in early life, to understand the potential for adaptive responses of wild populations to selection. Electronic supplementary material The online version of this article (10.1186/s12862-018-1154-9) contains supplementary material, which is available to authorized users.

We evaluated impact of the needle length, sex, and body condition on chemical immobilization induction time in 50 (29 males and 21 females) free-ranging fallow deer (Dama dama) in Sweden, 2006-11. Induction time is probably the single most important factor when immobilizing free-ranging wildlife with the use of a remote drug-delivery system. Induction times should be short to minimize stress and risk of injury, and to ensure that immobilized animals can be found and clinically monitored as soon as possible. We measured the distance between the darting location and where we recovered the immobilized animal and also the time occurring between the two events. We used two types of needles: 2.0 × 30- or 2.0 × 40-mm barbed needles with side ports. The most important result is that a 10-mm-longer dart needle can reduce the retrieval time substantially (20 min) until an animal is under monitoring. On average after the darting, the retrieval time decreased from 51 to 29 min and the distance decreased from 519 m from the darting location to 294 m. We suggest that a needle length of 40 mm is preferable for immobilization of wild fallow deer, especially for animals in over-average-to-fat body condition.

We present the first reference ranges for hematology (n = 35 animals), serum biochemistry (n = 62), and serum protein electrophoresis (n = 32) in physically restrained free-ranging roe deer (Capreolus capreolus). Animals were captured in box traps and physically restrained for blood sampling during the winter in Sweden, 2011-13. No clinically significant sex or age differences were found.

Tracking devices are commonly used to locate and monitor wild animals for studying spatial ecology and survival rates. There is growing interest in capture effects, partially to minimize the impact on the study species, but also for animal welfare reasons. This study aims to examine roe deer behaviour in box-traps, when restrained, when released and during recaptures to quantify injuries and deaths over a period of 41 years. We use data from 2911 captures from 926 individuals between 1973 and 2014. We recorded behaviour inside the box-traps over two seasons. We also recorded behavioural data from 671 catches of 346 individuals during six seasons to study habituation. Additionally, we discuss box-traps in relation to ethological theory and animal welfare. Over a 41-year period, one roe deer suffering from starvation was found dead in a trap (0.035%), which cannot be solely related to capture (N = 926). About 58% of all roe deer were recaptured at least once during their life time. There was a low prevalence of injuries (0.5% of the captures, N = 2911), and they occurred predominately to the nose or antlers in velvet (in males). During the first hour after capture, animals typically stand very tense between eating bouts. Thereafter, the deer tended to move more softly and exhibited resting behaviours (e.g. lying down). Overall, we conclude that this method of capture and handling had very low impact on the welfare or survival of roe deer, which also habituated to recapture over successive events.

SUMMARYHantaviruses are globally distributed and cause severe human disease.Puumalahantavirus (PUUV) is the most common species in Northern Europe, and the only hantavirus confirmed to circulate in Sweden, restricted to the northern regions of the country. In this study, we aimed to further add to the natural ecology of PUUV in Sweden by investigating prevalence, and spatial and host species infection patterns. Specifically, we wanted to ascertain whether PUUV was present in the natural reservoir, the bank vole (Myodes glareolus) further south than Dalälven river, in south-central Sweden, and whether PUUV can be detected in other rodent species in addition to the natural reservoir. In total, 559 animals were collected at Grimsö (59°43′N; 15°28′E), Sala (59°55′N; 16°36′E) and Bogesund (59°24′N; 18°14′E) in south-central Sweden between May 2013 and November 2014. PUUV ELISA-reactive antibodies were found both in 2013 (22/295) and in 2014 (18/264), and nine samples were confirmed as PUUV-specific by focus reduction neutralization test. Most of the PUUV-specific samples were from the natural host, the bank vole, but also from other rodent hosts, indicating viral spill-over. Finally, we showed that PUUV is present in more highly populated central Sweden.

Wildlife management systems face growing challenges to cope with increasingly complex interactions between wildlife populations, the environment and human activities. In this position statement, we address the most important issues characterising current ungulate conservation and management in Europe. We present some key points arising from ecological research that may be critical for a reassessment of ungulate management in the future. Ecosystem . Population sustainability . Science-basedmanagement .Wildlifemanagement .Adaptive management

Partial migration, when only part of the population migrates seasonally while the other part remains resident on the shared range, is the most common form of migration in ungulates. Migration is often defined by spatial separation of seasonal ranges and consequently, classification of individuals as migrants or residents is usually only based on geographic criteria. However, the underlying mechanism for migration is hypothesized to be movement in response to spatiotemporal resource variability and thus, migrants are assumed to travel an “ecological distance” (ED) or shift their realized ecological niches. While ecological and geographic distances should be related, their relationship may depend on landscape heterogeneity. Here, we tested the utility of ecological niche theory to both classify migratory individuals and to understand the underlying ecological factors for migratory behavior. We developed an integrative approach combining measures in geographic and ecological niche space and used this to classify and explain migratory behavior of 71 annual roe deer (Capreolus capreolus) movement trajectories in five European study areas. First, to assess the utility of the ED concept for classifying migratory behavior, we tested whether roe deer sought the same ecological conditions year-round or moved to different ecological conditions by measuring the annual ED travelled and the seasonal niche overlap using multivariate statistics. Comparing methods to classify migrants and residents based on geographic and ecological niche space, we found that migratory roe deer switched between seasons both in geographic and in ecological dimensions. Second, we tested which seasonal ecological factors separated resident from migrant niches using discriminant analysis and which broad-scale determinants (e.g., spatiotemporal forage variation and population density) predicted migration probability using generalized linear models. Our results indicated that factors describing forage and topographic variability discriminated seasonal migrant from resident niches. Determinants for predicting migration probability included the temporal variation (seasonality) and also the spatial variability of forage patches. Last, we also found suggestive evidence for a positive relationship between population density and migration probability. By applying the ecological niche concept to the study of partial migration in ungulates, our work underlines that partial migration is a form of behavioral plasticity. Key words: behavioral plasticity; Capreolus capreolus; ecological distance; large herbivores; niche switching; Normalized Difference Vegetation Index; partial migration; realized niche; spatiotemporal variation; ungulates