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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.

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.

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.

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.

For social foragers, movement as a group could increase foraging efficiency through collective discovery of high-quality food sources. This would require an efficient mechanism for transferring information about food quality between individuals. Conversely, the constraints of foraging as a cohesive group could decrease efficiency; grouping may persist to serve other functions such as protection from predators. To test what drives cohesion in herbivores, we manipulated patch shape and within-patch pattern of food quality and quantified the effects on group level diet selection by a social herbivore, the fallow deer, Dama dama. We arranged feeders containing fodder in lines or blocks, and manipulated the pattern of food quality within patches by adding tannin, a plant secondary compound that decreases palatability. We quantified the relative consumption of low- and high-tannin food to compare diet selectivity at the group level between patch treatments. If group foraging evolved to increase foraging efficiency, altering the spatial arrangement of food should not affect diet selectivity because information about food location and quality is shared. We found, however, that the herd expressed different levels of selectivity between both patch shapes and food quality patterns. Deer selected better diets in blocks than lines. In lines, the herd selected better diets when quality varied between alternate feeders rather than between the two halves of the patch, suggesting a reliance on personal rather than group information. Deer consumed the most at patch centres in all treatments except in blocks with high-tannin centres, but diet selection was poorer in the latter compared to blocks with low-tannin centres. Aggregation at the centre of patches appears to have restricted exploitation of the best food. Predation pressure and/or resource variability may have favoured the evolution of a foraging strategy that prioritizes social cohesion over effective diet selection.

Mammalian browsers can cause ecologically and economically significant damage to important tree species, particularly at the vulnerable seedling stage. Some tree species or particular parts of trees are less preferred as a result of chemical defences – harnessing such defences could reduce the appeal of more preferred trees and thus their loss to browsing. We tested the potential of birch bark extract, a forestry by-product, to protect seedlings of Scots pine Pinus sylvestris from two globally significant browsers: moose Alces alces and red deer Cervus elaphus . We quantified the responses by captive animals to extract application as a function of both the distance between seedlings and whether the extract was applied to all or alternate seedlings. Both moose and red deer consumed pine seedlings less frequently when seedlings were treated with birch bark extract, but their responses differed when only alternate seedlings were treated. At inter-seedling distances equivalent to those used in forestry, red deer browsed untreated seedlings more frequently when alternate seedlings were treated than when none were treated (associational susceptibility), whereas moose browsed untreated seedlings less frequently with treated than untreated neighbours (associational refuge). These neighbourhood effects were not evident at inter-seedling distances three times larger or smaller than those used in forestry. There was also no significant difference in the frequency of browsing on treated seedlings in alternate compared to completely treated arrangements at any inter-seedling distance. Red deer removed significantly less foliage from treated than untreated seedlings once browsed. Browsing by moose followed a similar but non-significant trend. The relative number of untreated to treated seedlings browsed was 2:1 for red deer and 4:1 for moose. Red deer used a finer scale of selection than moose for choosing between seedlings, consistent with their smaller body mass. However, the coarser-scale of selection used by moose resulted in consumption of more untreated relative to treated seedlings. Our study highlights the potential of existing plant defensive chemistry, harvested from forestry by-products, to reduce herbivore damage to tree seedlings. We also demonstrate the need to consider both plant associational effects and the foraging strategies of the dominant herbivores in applying repellents.

Background Browsing repellents are widely used to deter large herbivores from consuming plants of ecological, economic and aesthetic importance. Understanding how these repellents function on a behavioural mechanistic level is critical to predicting effectiveness. Here, we illustrate how these mechanisms can be tested, by exposing a model mammalian herbivore, the fallow deer, to different concentrations of a commercial chemical repellent (HaTe2) in two-choice feeding trials. Results The repellent acted as a defensive chemical for the food by both reducing visitation and the amount consumed. Deer favoured the less defended feeders before ingesting any food, suggesting that the repellent altered olfactory and/or visual cues. Deer also consumed less of the more defended food when choosing between low and high repellent feeders than no and low repellent feeders, indicating that the repellent modified flavour and/or sensation. Repellent effectiveness declined with increased exposure, suggesting that consumption had no negative post-ingestive effects, and thus, deterrence was not caused by a conditioned aversion or irritation. Instead, this pattern suggests that deer learned, through repeated sampling of repellent-treated food, that there was no adverse physiological effect of ingesting it. Conclusions These results imply that HaTe2 repellent will not be effective over prolonged periods or in the absence of alternative untreated food. Understanding the mechanisms driving repellent function using two-choice trials could help practitioners decide whether a particular repellent is likely to be effective against mammalian herbivory in their management scenario.

Personality exists in non-human animals and can impact fitness. There is, however, a shortage of empirical studies in certain areas within the field, and fundamental evolutionary theory on personality remains largely untested. For example, little is known on how variation in personality is maintained over evolutionary time. Theory suggests that fluctuating selection pressures due to spatiotemporal variation in conditions, e.g. food availability, is a possible mechanism and a few studies have shown that the success of different personality types varies with spatiotemporal conditions. However, it remains unknown whether different mechanisms can maintain personality within a species. Here we use a reintroduction programme for the critically endangered European mink (Mustela lutreola) to test whether multiple personality trait domains (boldness, exploration and sociability) affected survival in two different years and islands. This was done through pre-release personality tests and post-release radio-tracking monitoring. Survival was positively correlated with boldness, whereas the relationship with exploration was either negative or positive depending on year/island. The results show a complex relationship between personality and survival and suggest that exploration can be maintained over evolutionary time via spatiotemporal variation in conditions. However, in contrast to exploration, boldness did not vary spatiotemporally and sociability had no impact on survival. This indicates that different personality trait domains might be maintained by different mechanisms. To date, personality has been studied primarily within behavioural sciences, but through empirical findings we highlight the importance of personality also in ecology and conservation biology. Electronic supplementary material The online version of this article (doi:10.1007/s00442-016-3740-0) 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.

It is increasingly common to quantify and describe behavioral variation in domestic and wild animals in terms of personality. Correlating behavioral traits are referred to as personality dimensions ...

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. Electronic supplementary material The online version of this article (doi:10.1186/s12917-017-1045-0) contains supplementary material, which is available to authorized users.

Non-human animal personality is defined as consistent behavioural differences across time and situations/contexts. Behaviours are, however, often plastic and to explain how plasticity and personality may coexist an adaptive framework has been developed. Still, there is little information on how personality is impacted by situations and contexts on an individual level. We investigated this in the European mink (Mustela lutreola) by performing a set of five experiments in two situations consisting of non-breeding and breeding season, and by using different test contexts. Three personality trait domains were identified; boldness, exploration and sociability. The levels of boldness and exploration changed between seasons but remained repeatable, which implies behavioural reaction norms and supports that the concept of personality remained applicable despite plasticity. Whilst males became bolder and more explorative in the breeding season females became shyer, which reflects European mink breeding behaviour. Furthermore, behaviours performed in mirror stimulus tests fell into different domains depending on whether, the test was conducted in the own territory or not, suggesting plasticity in the response towards conspecifics. To conclude, our results highlight the importance of situation and context for the expression of personality, and the significance of measuring multiple personality trait domains with several methods.

A major problem within forest industry is unwanted browsing on seedlings from mammalian herbivores. The aim of this study was to evaluate the effects of birch bark extracts as repellents towards fallow deer. Birch bark was extracted in a conventional way with ethanol as solvent at ambient temperature and with a new method, liquid CO2 extraction. An analysis of the ethanol-extracted birch bark showed that it contained large amounts of terpenoids, of which the most abundant was betulin. In seven different treatment trials, we used 15 individually handled fallow deer. To investigate the binary taste preferences, birch bark extract was added to food and presented in two bowls in typical two-choice tests. We found that the amount of a food type consumed during a trial and the number of shifts between food bowls were dependent on the amount of the birch extract the food contained. Concentrations of above 1 % by dry weight of birch extract acted as a repellent. In addition, such concentrations produced shorter feeding bouts by a greater willingness to change bowls. Therefore, our conclusion is that birch bark extract acts as a repellent towards fallow deer and is therefore likely to act as a repellent against other deer species. In addition, we show that birch bark extract produced by the new and more environmentally sustainable method employing liquid CO2 mixed with ethanol has the same repellent effect as the traditional ethanol extraction.

An understanding of mortality patterns, and especially the variation in juvenile mortality, is an important component in vertebrate population dynamics. Our study investigates, for the first time, neonate mortality and two levels of spatial behaviour, in a free-ranging fallow deer Dama dama population in southwestern Sweden. In the summers of 2008 and 2009, 36 fawns were marked with radio-collars. Neonate mortality calculated by the Kaplan-Meier method was 23.6%. Mortality caused by predation was low, since only one of eight non-surviving fawns died from predation, probably by red fox Vulpes vulpes. The spatial behaviour of the neonates was examined by habitat selection at home-range level, which in fact is a selection made by the mother, and at bed-site level within that habitat. Compositional analysis revealed a significant preference for arable land, pasture and coniferous forest between 5-15 m high, compared to young forest. Selected bed sites showed significantly lower visibility and higher amount of canopy cover than random sites. Surprisingly, we did not find any relationship between canopy cover and visibility in selected bed sites while it showed a significant and negative relationship at random bed sites. We interpret this finding as while high canopy cover and low visibility covary at the habitat level, fawns seem to select these two bed-site variables independently, perhaps for thermoregulatory reasons. Since there are few predators in our study area and predation pressure is low, this behaviour is not connected to actual survival rates in this area, but would rather be in support of the hypothesis of ‘pleiotropy’ as thermoregulatory reasons for bed-site selection in neonate fawns might be the most important contemporary selection force in the absence of large predators.

A number of studies have reported how neighbouring plants may influence herbivory on palatable or unpalatable plants. Such neighbourhood effects can have important evolutionary consequences as they ...

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.

Recent studies have examined the ecological and evolutionary bases for variation in animal personality. However, only a few such studies have examined how foraging parameters are influenced by different personality domains. In wild ungulates, the trade-off between the time spent on food intake and antipredator behaviour differs between individuals, but the underlying reason for this is not yet well understood. One possibility is that this trade-off reflects personality dimensions such as boldness. To relate foraging decisions to personality we measured personality and performed feeding experiments with familiar and novel food in familiar and novel situations. We measured personality traits in 15 tame fallow deer, using novel object tests (NO), behavioural observations (BO) and personality ratings (PR). Boldness dimensions were found using PR and NO, dominance dimensions were found using BO and PR, and a flexibility dimension was found using BO. Multitrait–multimethod analysis showed that similar dimensions were significantly correlated across different methods and that different dimensions were not significantly correlated, even if measured using the same method. We also found that novel food eaten in familiar situations and familiar food eaten in novel situations were strongly related to boldness but not dominance, flexibility or age. Thus the trade-off between the benefits of gaining more food and the costs of reduced vigilance or increased toxin ingestion reflect boldness. These findings highlight the nature of personality dimensions in ungulates and how boldness impacts foraging behaviour.

Consummatory simultaneous positive and negative contrast in fallow deer: implications for selectivity

Intraspecific variation in plant toxins at different spatial scales can influence foraging decisions by wild herbivores. In order to investigate plant associational defences in relation to spatial scale, we performed an experiment with fallow deer encountering 2 patches of low- and high-tannin hazel branches. One patch was good, consisting of 7 low- and 1 high-tannin branch, and the other bad, with 1 low- and 7 high-tannin branches. We kept the between-patch spatial scale constant and varied the within-patch spatial scale: the branches in a patch were either spread out or close together in a bundle. When the branches were spread out, the deer showed a clear preference for low-tannin branches in both patches and consumed similar amounts from low-tannin branches in the good and the bad patch, which means that there was no associational defence. In contrast, when the branches instead were together in a bundle, within-patch selectivity decreased and between-patch selectivity increased, and the low-tannin branches in the bad patch were less eaten than the low-tannin branches in the good patch, which corresponds to associational defence. We conclude that small inter-plant distances can be crucial for the operation of plant associational defences. Keywords: associational defence, associational susceptibility, fallow deer, selectivity, spatial scale. Resume : La variation intraspecifique des toxines des plantes a differentes echelles spatiales peut influencer les decisions de quete alimentaire des herbivores sauvages. Pour examiner les defenses associatives des plantes en fonction de l'echelle spatiale, nous avons effectue une experience dans laquelle des daims avaient l'option de s'alimenter dans 2 types de parcelles contenant des branches de noisetier, un dont le contenu en tanins des branches etait eleve et l'autre bas. Une des parcelles etait bonne, contenant sept branches a faible contenu en tanins et une seule a fort contenu en tanins. L'autre parcelle etait mauvaise, avec une seule branche a faible contenu en tanins et sept autres a fort contenu en tanins. Nous avons maintenu l'echelle spatiale constante entre les parcelles et avons fait varier l'echelle spatiale a l'interieur des parcelles ou les branches etaient soit dispersees, soit rassemblees dans un paquet. Lorsque les branches etaient dispersees, les daims ont demontre une nette preference pour les branches a faible contenu en tanins dans les 2 types de parcelles et ont consomme des quantites semblables de branches a faible contenu en tanins dans les parcelles bonne et mauvaise, ce qui signifie qu'il n'y avait aucune defense associative des plantes. Au contraire, lorsque les branches etaient rassemblees dans un paquet, les daims ont ete moins selectifs a l'interieur des parcelles et plus selectifs entre les parcelles. Les branches a faible contenu en tanins ont ete moins mangees dans la mauvaise parcelle que dans la bonne, ce qui correspond a une defense associative des plantes. Nous concluons que de faibles distances entre les plantes peuvent etre cruciales pour que les defenses associatives de celles-ci soient fonctionnelles. Mots-cles : daim, defense associative, echelle spatiale, selectivite, sensibilite associative.

Large herbivores select food at several spatial scales: plant communities are chosen at a landscape scale, plant patches are chosen within a plant community, and individual plants within a patch. Foraging decision at the patch level can result in associational effects in plant communities and populations. Several studies have shown that herbivore attack and consumption rates may not only depend on a plant's own defence traits, but also on the defence traits of its neighbours. In the present experiment we investigated whether the spatial scale of the food distribution affects food selection by fallow deer and whether the foraging behaviour gives rise to associational effects in plant defences. In a population of captured wild fallow deer we simulated a natural situation where two separate plant patches are exposed to intense herbivory pressure. We presented different spatial arrangements of low- and high-tannin food to the deer, varying the frequency of the feeder types within and between patches. We found that the deer consumed palatable food among the unpalatable food on average as much as they consumed palatable food among other palatable feeders. However, when unpalatable food occurred among the palatable food it was more consumed than among other unpalatable feeders. Hence, we did not find support for associational defence, but our results supported associational susceptibility. At the between patch level a patch of mainly high-tannin feeders was consumed less when presented near to a patch of mainly low-tannin feeders, suggesting that for well-defended plants having palatable neighbours in a nearby patch might accentuate the effectiveness of their defence.

The study of contrast investigates how rewards influence behaviour when animals are exposed to two or more levels of rewards compared to when they experience only a single level. The appearance of an exaggerated response to a shift in reward is referred to as a contrast effect and is an empirically well-established phenomenon. Although contrast effects could be important in foraging behaviour, no direct experimental tests of contrast effects in foraging by mammalian herbivores exist. During foraging, mammalian herbivores can encounter a range of plants that vary in the amount of nutrients and toxins. They may thus compare food items by taste, which in turn can give rise to contrast effects. In feeding experiments with fallow deer, Dama dama, we investigated the presence of simultaneous negative contrast. We found that the deer consumed less from a bowl of pellets containing 1% tannin when they shifted to it from a bowl with pellets containing only 0.25% tannin than when they shifted from another bowl with pellets containing 1% tannin. We estimated a fourfold difference between treatments in test food consumption at the highest levels of preloading, but none at the lowest levels. We found no support for successive negative contrast in experiments where the deer approached food in a runway, comparing a current reward with the memory of a previous reward. We suggest that simultaneous negative contrast can influence foraging decisions in mammalian herbivores.

In the field of plant-herbivore interactions research methods manipulating plant secondary metabolites are becoming more and more popular. In most cases food offered to herbivores is manipulated wi ...

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.

Behavioural consistency is a key assumption when evaluating how between-individual differences in behaviour influence life history tactics. Hence, understanding how and why variation in behavioural repeatability occurs is crucial. While analyses of behavioural repeatability are common, few studies of wild populations have investigated variation in repeatability in relation to individual status (e.g. sex, age, condition) and over different timescales. Here, we aimed to fill this gap by assessing within-population variation in the repeatability of docility, as assessed by the individual's response to human handling, in a free-ranging population of European roe deer, Capreolus capreolus. Docility was an equally repeatable behaviour at both short- and long-term timescales, suggesting that this behavioural trait is stable across time. Repeatability did not differ markedly between age and sex categories but tended to be higher in juvenile males than in juvenile females. Finally, contrary to expectation, individual variation in the repeatability of docility was not correlated with individual body mass. Further studies are required to assess the life history consequences of the individual variation in docility we report here. (C) 2015 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

We have investigated food choice in individual fallow deer (Dama dama) encountering different relative frequencies of food types in the form of bowls containing pellets with either high or low conc ...

In order to better understand the development and maintenance of feeding selectivity, several feeding experiments were performed with fallow deer (Dama dama L.). In experiments performed when the fawns were between ten and 27 days old, it was found that all fawns showed preferences for sucrose but aversions towards tannic acid and ascorbic acid. However, differences in selectivity towards tannic acid were present already before the fawns became functional ruminants and these individual differences lasted 5 years. Moreover, individuals that ingested overall less tannic acid, searched more thoroughly between food sources. When the foraging behaviour was compared with age (11-41 days old and 65-97 days old), it was found that the time a fawn spent eating, increased with age, and the time spent on exploration, smelling and tasting plants decreased with age. Furthermore, the fawns increased their intake of grass and herbs, while the intake of soil and dead plant material decreased with age.

A basic idea of plant defences is that a plant should gain protection from its own defence. In addition, there is evidence that defence traits of the neighbouring plants can influence the degree of protection of an individual plant. These associational effects depend in part on the spatial scale of herbivore selectivity. A strong between-patch selectivity together with a weak within-patch selectivity leads to a situation where a palatable plant could avoid being grazed by growing in a patch with unpalatable plants, which is referred to as associational defence. Quite different associational effects will come about if the herbivore instead is unselective between patches and selective within a patch. We studied these effects in a manipulative experiment where we followed the food choice of fallow deer when they encountered two patches of overall different quality. One of the two patches consisted of pellets with low-tannin concentration in seven out of eight buckets and with high concentration in the remaining bucket. The other patch instead had seven high- and one low-tannin bucket. We performed the experiment both with individuals one at a time and with a group of 16–17 deer. We found that the deer were unselective between patches, but selective within a patch, and that the single low-tannin bucket among seven high-tannin buckets was used more than a low-tannin bucket among other low-tannin buckets. This corresponds to a situation where a palatable plant that grows among unpalatable plants is attacked more than if it was growing among its own kind, and for this effect we suggest the term neighbour contrast susceptibility, which is the opposite of associational defence. We also found that the high-tannin bucket in the less defended patch was less used than the high-tannin buckets in the other patch, which corresponds to neighbour contrast defence. The neighbour contrast susceptibility was present both for individual and group foraging, but the strength of the effect was somewhat weaker for groups due to weaker within-patch selectivity.