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Background: In Europe, Ixodes ricinus (Acari: Ixodidae) is the most widespread and abundant tick species, acting as a vector for several microorganisms of medical and veterinary importance. In Northern and Central Europe, the tick has a bimodal activity pattern consisting of a peak in spring to the beginning of summer and a second peak at the end of summer. However, several findings of ticks on animals during winter have been reported, which raises the question of whether this is an overwintering strategy or whether ticks are active during winter in Scandinavia. The objectives of our study were to determine (i) whether ticks were active and finding hosts during winter, (ii) whether they parasitize their hosts, and (iii) what climatic factors-i.e., temperature, snow depth and precipitation-govern tick winter activity., Methods: Throughout three winter seasons, we examined wild-living and free-ranging roe deer (Capreolus capreolus) for ticks on 332 occasions. In total, 140 individual roe deer were captured in two climatically contrasting sites in south-central Sweden, Grimsö and the Bogesund research area, respectively. We re-examined individual roe deer up to 10 times within the same winter or approximately once a week (mean 10 days, median 7 days between re-examinations) and recorded the absence or presence of ticks on the animals, and tested to what extent meteorological factors affected tick activity. To determine the attachment day, we used the coxal/scutal index of 18 nymphs and 47 female ticks., Results: In total, 243 I. ricinus were collected from 301 roe deer captures between 14 December and 28 February at the Bogesund study site during three subsequent years (2013/2014-2015/2016). We found attached ticks every third to every second examination (32%, 48% and 32% of the examinations, respectively). However, we collected only three I. ricinus females from 31 roe deer captures at the Grimsö study site between 17 December 2015 and 26 February 2016. At the Bogesund study site, based on 192 captures of previously examined deer, we collected 121 ticks, and ticks were found at 33%, 48% and 26% of the examinations during the respective winters. The probability of finding an attached tick on a roe deer at a temperature of -5 °C was > 8% ± 5 (SE), and that probability increased to almost 20% ± 7 (SE) if the air temperature increased to 5 °C., Conclusions: To the best of our knowledge, this is the first time that winter-active nymphs and female ticks have been documented to attach and feed on roe deer during winter (December to February) in Scandinavia. The main weather conditions regulating winter activity for females were temperature and precipitation, and the lowest estimated air temperature for finding an active tick was well below 5 °C. The behaviour of winter-active and blood-feeding ticks was documented over several winters and in two contrasting areas, implying that it is a common phenomenon that should be investigated more thoroughly, since it may have important consequences for the epidemiology of tick-borne pathogens., (© 2023. The Author(s).)

Background: Wildlife hosts may serve as reservoirs for strongyles, which can be transmitted to domestic livestock. Therefore, studies evaluating nemabiome compositions in wildlife ruminants are of great use in assessing the possibility of transmission of important nematode pathogens to domestic sheep in Sweden., Methods: First, fecal samples were collected from roe deer (n = 125), fallow deer (n = 106), red deer (n = 18) and mouflon (n = 13) in south central Sweden during the hunting season in 2019. Second, after fecal examination samples were cultured and the larvae were harvested, followed by DNA extractions. Third, all samples were barcoded and processed for sequence analysis on the PacBio platform. Finally, bioinformatic sequence analysis was conducted with DADA2, while species diversity and richness, as well as interactions between the different hosts, were calculated and analyzed in R., Results: Nematode ITS2 sequences were found in 225 of 262 (86%) samples. In total, 31 taxa were identified, among which 26 (86%) to the species level. These were found in different combinations, among which 24 (77%) occurred in roe deer, 19 (61%) in fallow deer, 20 (65%) in red deer and 10 (32%) in mouflon. Five of the species found are known to be associated with livestock (Chabertia ovina, Haemonchus contortus, Oesophagostomum venulosum, Teladorsagia circumcincta and Trichostrongylus axei). However, in the present study the relative abundance and prevalence of most of these species were low. The most striking exception was T. axei, which was relatively abundant in all wildlife hosts. Mostly a wide range of wildlife specific nematodes such as Ostertagia leptospicularis and Spiculopteragia spp. were identified including the invasive nematode Spiculopteragia houdemeri, which was found for the first time in red deer, fallow deer, and mouflon in Sweden. The difference in the number of shared species between mouflon and all cervids (n = 6) was less than among all three cervids (n = 8)., Conclusion: In this study, we investigated the community structure of parasitic intestinal nematodes in four wildlife hosts, and we found that the majority of the parasite species identified were wildlife specific. We also found a new, potentially invasive species not reported before. After comparing the nemabiome of the wildlife hosts in this study with a previous study in sheep from the same geographical region, we conclude that the horizontal transmission potential appears to be relatively low. Still, cross-infections of nematodes between game and sheep cannot be completely ignored., (© 2022. The Author(s).)

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" [3-5]. 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., Competing Interests: Declaration of Interests The authors declare no competing interests., (Copyright © 2020 Elsevier Inc. All rights reserved.)

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., (© 2019 British Ecological Society.)

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., Competing Interests: The authors have declared that no competing interests exist.

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.

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 multi-range 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.

Background: To understand and reduce the concomitant effects of trapping and handling procedures in wildlife species, it is essential to measure their physiological impact. Here, we examined individual variation in stress levels in non-anesthetized European roe deer (Capreolus capreolus), which were captured in box traps and physically restrained for tagging, biometrics and bio-sampling. In winter 2013, we collected venous blood samples from 28 individuals during 28 capture events and evaluated standard measurements for stress (heart rate, body temperature, neutrophil to lymphocyte ratio, lactate and total cortisol). Additionally, we assessed stress using the immunological tool, Leukocyte Coping Capacity (LCC), a real-time proxy for stress measuring oxygen radical production by leukocytes. Finally, the behavioral response to handling was recorded using a scoring system., Results: LCC and therefore stress levels were negatively influenced by the time animals spent in the box trap with human presence at the capture site prior to handling. In contrast, none of the classical stress measures, including total cortisol, nor the behavioral assessment, were correlated with the stressor tested (time of human presence prior to handling) and thus did not provide a clear depiction regarding the extent of the animals short-term stress response., Conclusions: Overall our study verifies the LCC as a strong method to quantify short-term stress reactions in wildlife. Moreover, our results clearly show that human presence at the trapping site prior to handling should be kept to an absolute minimum in order to reduce stress levels.

The risk allocation hypothesis predicts that vigilance should be adjusted to the temporal variation in risk. We test this hypothesis in wild fallow deer exposed to short term (disturbance) and long term (presence of a fawn after parturition) changes in risk. We recorded the proportion, frequency and type of vigilance and size of used area before and after parturition, in GPS-collared wild female fallow deer. Vigilance was divided in two main groups: "non-grazing vigilance" and "grazing vigilance". The latter group was divided into "grazing vigilance while chewing" and a "grazing vigilance when chewing was interrupted". By recording external disturbance in form of passing cars, we were able to investigate if this altered the amount, and type of vigilance. We found that females increased the proportion and frequency of "grazing vigilance stop chewing" after parturition. The "grazing vigilance chewing" was unaffected, but "non-grazing vigilance" decreased. Disturbance increased the proportion "grazing vigilance stop chewing" to the same extent before and after parturition. We found a clear decrease in female home range size after parturition as a possible behavioural adjustment. The increase in "grazing vigilance stop chewing" after parturition is a rarely described but expected cost of reproduction., (Copyright © 2016 Elsevier B.V. All rights reserved.)

Background: The epidemiology of the zoonotic tick-transmitted parasite Babesia spp. and its occurrence in wild reservoir hosts in Sweden is unclear. In European deer, several parasite species, including Babesia capreoli and the zoonotic B. venatorum and B. divergens has been reported previously. The European roe deer, Capreolus capreolus, is an important and common part of the indigenous fauna in Europe, as well as an important host for Ixodes ricinus ticks, the vector of several Babesia spp. in Europe. Here, we aimed to investigate the occurrence of Babesia spp. in roe deer in Sweden., Findings: Roe deer (n = 77) were caught and sampled for blood. Babesia spp. was detected with a PCR assay targeting the 18S rRNA gene. The prevalence of Babesia spp. was 52%, and two species were detected; B. capreoli and B. venatorum in 44 and 7.8% of the individuals, respectively. Infection occurred both in summer and winter., Conclusions: We showed that roe deer in Sweden, close to the edge of their northern inland distributional range, are infected with Babesia spp. The occurrence of B. venatorum in roe deer imply that it is established in Sweden and the zoonotic implication of this finding should be regarded to a greater extent in future.

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.

Breeding dispersal, defined as the net movement between successive breeding sites, remains a poorly understood and seldom reported phenomenon in mammals, despite its importance for population dynamics and genetics. In large herbivores, females may be more mobile during the breeding season, undertaking short-term trips (excursions) outside their normal home range. If fertilisation occurs, leading to gene flow of the male genome, this behaviour could be considered a form of breeding dispersal from a genetic point of view. Here, we investigated ranging behaviour of 235 adult roe deer using intensive GPS monitoring in six populations across Europe within the EURODEER initiative. We show that excursions are common from June to August among females, with 41.8% (vs. 18.1% of males) making at least one excursion. Most individuals performed only one excursion per season and departure dates for females were concentrated in time, centred on the rutting period, suggesting a link with reproduction. The distance females travelled during excursions was significantly greater than the site-specific average diameter of a male home range, while travel speed decreased once they progressed beyond this diameter, indicating search behaviour or interaction with other male(s) outside the resident male's territory. Because adults are normally highly sedentary, the potential for mating with relatives is substantial; hence, we conclude that rut excursions could be an alternative tactic enabling females to avoid mating with a closely related male. To understand better the ultimate drivers at play, it will be crucial to explore the genetic causes and consequences of this behaviour.

1. Because many large mammal species have wide geographical ranges, spatially distant populations may be confronted with different sets of environmental conditions. Investigating how home range (HR) size varies across environmental gradients should yield a better understanding of the factors affecting large mammal ecology. 2. We evaluated how HR size of a large herbivore, the roe deer (Capreolus capreolus), varies in relation to seasonality, latitude (climate), weather, plant productivity and landscape features across its geographical range in Western Europe. As roe deer are income breeders, expected to adjust HR size continuously to temporal variation in food resources and energetic requirements, our baseline prediction was for HR size to decrease with proxies of resource availability. 3. We used GPS locations of roe deer collected from seven study sites (EURODEER collaborative project) to estimate fixed-kernel HR size at weekly and monthly temporal scales. We performed an unusually comprehensive analysis of variation in HR size among and within populations over time across the geographical range of a single species using generalized additive mixed models and linear mixed models, respectively. 4. Among populations, HR size decreased with increasing values for proxies of forage abundance, but increased with increases in seasonality, stochastic variation of temperature, latitude and snow cover. Within populations, roe deer HR size varied over time in relation to seasonality and proxies of forage abundance in a consistent way across the seven populations. Thus, our findings were broadly consistent across the distributional range of this species, demonstrating a strong and ubiquitous link between the amplitude and timing of environmental seasonality and HR size at the continental scale. 5. Overall, the variability in average HR size of roe deer across Europe reflects the interaction among local weather, climate and seasonality, providing valuable insight into the limiting factors affecting this large herbivore under contrasting conditions. The complexity of the relationships suggests that predicting ranging behaviour of large herbivores in relation to current and future climate change will require detailed knowledge not only about predicted increases in temperature, but also how this interacts with factors such as day length and climate predictability., (© 2013 The Authors. Journal of Animal Ecology © 2013 British Ecological Society.)

Risk of predation is an evolutionary force that affects behaviors of virtually all animals. In this study, we examined how habitat selection by roe deer was affected by risk of predation by Eurasian lynx - the main predator of roe deer in Scandinavia. Specifically, we compared how habitat selection by roe deer varied (1) before and after lynx re-established in the study area and (2) in relation to habitat-specific risk of predation by lynx. All analyses were conducted at the spatial and temporal scales of home ranges and seasons. We did not find any evidence that roe deer avoided habitats in which the risk of predation by lynx was greatest and information-theoretic model selection showed that re-colonization by lynx had limited impact on habitat selection by roe deer despite lynx predation causing 65% of known mortalities after lynx re-colonized the area. Instead we found that habitat selection decreased when habitat availability increased for 2 of 5 habitat types (a pattern referred to as functional response in habitat selection). Limited impact of re-colonization by lynx on habitat selection by roe deer in this study differs from elk in North America altering both daily and seasonal patterns in habitat selection at the spatial scales of habitat patches and home ranges when wolves were reintroduced to Yellowstone National Park. Our study thus provides further evidence of the complexity by which animals respond to risk of predation and suggest that it may vary between ecosystems and predator-prey constellations.

Births are highly synchronized among females in many mammal populations in temperate areas. Although laying date for a given female is also repeatable within populations of birds, limited evidence suggests low repeatability of parturition date for individual females in mammals, and between-population variability in repeatability has never, to our knowledge, been assessed. We quantified the repeatability of parturition date for individual females in five populations of roe deer, which we found to vary between 0.54 and 0.93. Each year, some females gave birth consistently earlier in the year, whereas others gave birth consistently later. In addition, all females followed the same lifetime trajectory for parturition date, giving birth progressively earlier as they aged. Giving birth early should allow mothers to increase offspring survival, although few females managed to do so. The marked repeatability of parturition date in roe deer females is the highest ever reported for a mammal, suggesting low phenotypic plasticity in this trait.

1. Territoriality is commonly associated with resource defence polygyny, where males are expected to gain access to females by anticipating how resources will influence female distribution and competing for resource-rich sites to establish their zone of dominance. 2. We tested this hypothesis in European roe deer (Capreolus capreolus) by simultaneously assessing the influence of resources on female distribution and the influence of female distribution on male distribution and breeding success using paternity analyses. 3. Females did not fully distribute themselves among male territories in relation to resources. As a result, relative female abundance in a male's territory depended on territory size, but not on its habitat quality. In turn, relative female abundance in a male's territory determined, at least partially, his breeding success. 4. Interestingly, male territory size, and hence access to females, was partly determined by male body mass (all males) and by residual antler size (subadults only). The latter result suggests that large antlers may be important to young males for establishing their first territory, which is then usually retained for all subsequent reproductive seasons. 5. To conclude, although territoriality of male roe deer has certainly evolved as a tactic for ensuring access to mates, our results suggest that it does not really conform to a conventional resource defence polygyny strategy, as males seem to gain no obvious benefit from defending a territory in an area of high habitat quality in terms of enhanced access to mates. 6. This may explain the stability of male territories between years, suggesting that male territoriality conforms to an 'always stay' and 'low risk-low gain' mating strategy in roe deer.

Identifying factors shaping secondary sexual traits is essential in understanding how their variation may influence male fitness. Little information is available on the allocation of resources to antler growth in territorial ungulates with low sexual size dimorphism. We investigated phenotypic and environmental factors affecting both absolute and relative antler size of male roe deer in three contrasting populations in France and Sweden. In the three populations, we found marked age-specific variation in antler size, with an increase in both absolute and relative antler size between yearling and prime-age stages, followed by a decrease (senescence) for males older than 7 years. Antler size increased allometrically with body mass. This increase was particularly strong for senescent males, suggesting the evolution of two reproductive tactics: heavy old males invested particularly heavily in antler growth (potentially remaining competitive for territories), whereas light old males grew small antlers (potentially abandoning territory defense). Finally, environmental conditions had little effect on antler size: only population density negatively affected absolute antler size in one of the three populations. Antler size may therefore provide an honest signal of male phenotypic quality in roe deer. We discuss the implications of these results in terms of territory tenure and mating competition.

We investigated the influence of population density on juvenile body mass in two contrasting roe deer populations, in Sweden (Bogesund) and France (Chizé), in which density was monitored for >or=15 years. We investigated the effect of population density and climatic conditions on cohort performance. We predicted that: (1) body mass of growing fawns should be sensitive to environmental changes, showing marked between-year variation (i.e., cohort effects), (2) fawns in the less productive (weakly seasonal, weakly predictable summer weather) habitat of Chizé should show stronger density-dependent responses due to more severe food competition during summer than fawns in the more productive (markedly seasonal, moderately predictable summer weather) habitat of Bogesund, and (3) fawns at Bogesund should be heavier both in absolute terms and relative to their size than their conspecifics in Chizé due to a higher degree of fat accumulation in northern environments. In both study sites we found marked cohort variation and clear effects of density, with body mass varying by as much as 29% over years. While neither summer nor winter climate influenced fawn body mass at Bogesund, fawns tended to be lighter after summers with high temperatures at Chizé. In addition, fawns were heavier after acorn mast years experienced in utero at Bogesund. As expected, the strength of the density-dependent response of fawn body mass was greater at Chizé than at Bogesund. For a given density, male fawns were consistently heavier than females in both sites. Lastly, both sexes at Bogesund had higher absolute body mass and were larger for a given body size than in Chizé. Our results clearly demonstrate that absolute density is a poor predictor of roe deer performance and supports the view that habitat quality has an overwhelming importance for determining fawn body mass in roe deer populations.

We studied the effects of population density, red fox predation risk, individual body mass and longevity on female fitness in a free-ranging roe deer population. During the study, population density varied from 9.3 to 36.1 deer km(-2), and red fox abundance varied strongly over years owing to a sarcoptic mange outbreak. In support of our predictions, long-lived females had higher fitness than short-lived ones. Further, fortunate female roe deer that gave birth in years of low red fox abundance attained much higher fitness than those that gave birth in years of high red fox abundance. Longevity and predation risk explained more than half the variation in fitness observed among roe deer females. As a possible effect of small sample size, we found no effect of female body mass or population density at birth. Our study demonstrates that predation risk, a component of environmental stochasticity, may prevent directional selection even when phenotypic quality influences individual fitness.

Body mass is a key determinant of fitness components in many organisms, and adult mass varies considerably among individuals within populations. These variations have several causes, involve temporal and spatial factors, and are not yet well understood. We use long-term data from 20 roe deer cohorts (1977-96) in a 2600 ha study area (Chizé, western France) with two habitats contrasting in quality (rich oak forest in the North versus poor beech forest in the South) to analyse the effects of both cohort and habitat quality on adult mass (i.e. median body mass between 4 and 10 years of age) of roe deer (Capreolus capreolus). Cohort strongly influenced the adult body mass of roe deer in both sexes: males born in 1994 were 5.2 kg heavier when aged between 4 and 10 years old than males born in 1986, while females born in 1995 were 4.7 kg heavier between 4 and 10 years old than females born in 1982. For a given cohort, adult males were, on average, 0.9 kg heavier in the rich oak forest than in the poor beech forest. A similar trend occurred for adult females (0.5 kg heavier in the oak forest). The effects of cohort and habitat were additive and accounted for ca. 40% of the variation observed in the adult mass of roe deer at Chizé (males: 41.2%; females: 40.2%). Population density during the spring of the birth accounted for about 35% of cohort variation, whereas rainfall in May-June had no effect. Such delayed effects of density at birth on adult body mass probably affect population dynamics, and might constitute a mechanism by which delayed density-dependence occurs in ungulate populations.

ABSTRACT Research on climate change impacts has focused on projecting changes in the geographic ranges of species, with less emphasis on the vital rates giving rise to species distributions. Managing ungulate populations under future climate change will require an understanding of how their vital rates are affected by direct climatic effects and the indirect climatic and non-climatic effects that are often overlooked by climate impact studies. We used generalized linear models and capture-mark-recapture models to assess the influence of a variety of direct climatic, indirect climatic, and non-climatic predictors on the survival of roe deer ( Capreolus capreolus) at 2 sites in Sweden. The models indicated that although direct climatic effects (e.g., precipitation) explained some variation in survival, indirect climatic effects (e.g., an index of vegetation production), and non-climatic effects (hunting by lynx [ Lynx lynx] and humans) had greater explanatory power. Climate change is likely to increase vegetation productivity in northern Europe, and, coupled with the positive effects of vegetation productivity on roe deer survival, might lead to population increases in the future. Survival was negatively affected by lynx presence where these predators occur and by human harvest in the site that lacked predators. In the future, managers might find that a combination of increased harvest and predation by recovering carnivore populations may be necessary to mitigate climate-induced increases in roe deer survival. Considering vegetation availability and predation effects is likely to improve predictions of ungulate population responses to variation in climate and, therefore, inform management under future climate change. © 2016 The Wildlife Society. [ABSTRACT FROM AUTHOR]

Because avoiding predation is crucial for fitness, foraging animals must trade acquisition of high-quality resources against risk avoidance when the best resources occur in locations with high predation risk. Although optimality models predict the distance at which an animal should initiate vigilance and flight, many studies have shown that animals generally flee soon after detecting an approaching threat, supporting the ‘flush early and avoid the risk’ (FEAR) hypothesis. Despite this, flight behaviour varies markedly depending on context, suggesting some behavioural plasticity in the response of prey to a given threat. We evaluated the degree of plasticity in the flight responses of roe deer, Capreolus capreolus , a highly flexible species which thrives in human-dominated landscapes. Based on individually identifiable animals and a standardized flight initiation protocol, we measured the distance at which a deer detected an approaching threat, and the distance at which it subsequently initiated flight. Our results provide strong support for the FEAR hypothesis, suggesting that alert and flight responses are strongly coupled in roe deer. However, the perceived level of both risk (in terms of landscape openness and proximity to human infrastructure) and reward (in terms of habitat quality) influenced the time it took for a deer to detect an approaching threat, and the subsequent time for which the threat was tolerated prior to flight. Overall, our findings indicate that although roe deer minimize monitoring costs when assessing risk by fleeing early, they also adjust their monitoring and flight responses to the local risk–resource trade-off. [ABSTRACT FROM AUTHOR]