The disappearance of muntjac (Muntiacus reevesi) and roe deer (Capreolus capreolus) pellet groups in a pine forest of lowland England

Reliable knowledge of the disappearance rate of faecal pellets is essential for converting pellet density to deer density when using standing-crop pellet-group counts. Disappearance of muntjac and roe deer pellet groups was monitored in four growth stages of a pine forest of lowland England over a 15-month period. Time to disappearance of the pellet groups (days) of both species significantly differed between habitats and months; it was shorter in late summer to early autumn and in habitats with more ground vegetation. Muntjac pellet groups disappeared more quickly than roe deer pellet groups. Time to disappearance of roe deer pellet groups was negatively correlated with air and grass temperature in pre-thicket and pre-fell habitats, while time to disappearance of muntjac pellet groups was negatively correlated with frequency of rainfall and positively correlated with the run of wind (air passage over a site within a 24-h period measured in km) in pre-thicket habitats. It is the time of the standing-crop pellet-group counts and the disappearance rate of pellet groups deposited in different months and habitats that determine the appropriate method for conversion of pellet-group density to deer density.     

Local and seasonal variations of roe deer diet in relation to food resource availability in a Mediterranean environment

Feeding behaviour patterns of European roe deer are well known but not in their southern range, where in contrast to Central and northern Europe the climate is characterised by mild winters, but hot and dry summer months. Spatial and temporal variations of the roe deer diet were investigated by analyses of 597 faecal samples in a coastal area in Italy. The aim was to evaluate the adaptive strategies of dietary intake to seasonal availability of different forage types in areas of fairly natural Mediterranean scrubwood and different areas dominated by agricultural habitats. A wide range of plant taxa (151 species/genera) was used, belonging to eight categories. A gradient was found from a diet dominated by woody plants and fruits in natural habitats to one characterised by cultivated plants in the agricultural areas. Usage of forage categories in areas dominated by woodland was proportional to their availability, whereas diet selection was found in the agricultural habitats, particularly marked in summer, the time at which food quantity and quality decreased due to harvest and drought. As to seasonal variation, the summer and autumn diets included more deciduous browse, whereas the winter diet was characterised by an increasing use of cereals and grass. In most areas, half-woody browse and fruit were preferred; grass and cereals were used less than expected by their availability. Most probably, selective behaviour patterns are linked to the difference of quality between plants together with their variable abundance, which is more evident in agricultural areas than in the woodland.     

Estimation of roe deerCapreolus capreolus and mouflonOvis aries densities, abundance and habitat use in a mountainous Mediterranean area

The abundance, density, and habitat use of roe deerCapreolus capreolus (Linnaeus, 1758) and mouflonOvis aries Linnaeus, 1758 were studied in a confined Mediterranean area in Greece with a dung survey based on the faecal accumulation rate (FAR) technique. Estimated density was modelled with generalized additive models using altitude, habitat type, and slope as potential covariates. Model selection among the set of candidate models was conducted based on their generalized cross-validation score. Roe deer had an estimated mean density of 13.9 ind./km² and the best model included slope and habitat type as covariates. The mean density of mouflon in the study area was 22.1 ind./km² and the best model used altitude and habitat type as covariates. For both species, the highest densities were encountered in abandoned cultivations and glades, followed by conifer forests, while the lowest densities were observed in maquis. However, use of open habitats by mouflon was much greater than it was for roe deer. The strong preference of mouflon (a grazer species) for open habitats that were abundant with grasses probably reflected food availability and contrasted with the more diverse habitat use by roe deer (a selective browser).     

Estimating roe deer abundance from pellet group counts in Spain: An assessment of methods suitable for Mediterranean woodlands

Despite of the generalized expansion of wild ungulates in Europe, roe deer (Capreolus capreolus) is experiencing contrasted population trends; it is expanding in some regions while declining in others likely due to the expansion of other deer species. In both extremes, reliable methods to estimate population abundance are required. We evaluated different methods of estimating deer abundance in Mediterranean woodlands based on pellet group counts. Distance Sampling applied to pellet counts and a new easier and cost-effective method based on strip-variable transect counts (FST) were assessed comparing their estimates (pellet group density) with the abundance indices obtained from traditionally used reference methods (faecal standing crop) in 61 localities (n = 183 surveys). The average roe deer density estimated from faecal standing crop was 5.56 ± 0.75 (range 0.01–20.74) deer per 100 ha. Distance Sampling and FST estimates correlated with reference methods. As a first conclusion it may be noted that all indirect methods used here can be used to estimate roe deer abundance. The selection of a given method based on pellet counts to estimate roe deer population abundance should take into account the specific objectives of the research, resources available, and the timescale in which the information is required. Among them, Distance Sampling may be used when human resources and skills are enough but FST is a rapid and efficient alternative to estimate pellet group density when they are not.     

Non-invasive monitoring of adrenocortical activity in free-ranging fallow deer (<Emphasis Type="Italic">Dama dama</Emphasis> L.)

Measurement of faecal glucocorticoid metabolites is increasingly used as a non-invasive tool to examine disturbances in various domestic and wild animals. Because measurements of faecal glucocorticoid metabolites has previously never been reported in fallow deer, we determined 11,17-dioxoandrostanes (11,17-DOA), a group of cortisol metabolites, in the faeces of four fallow deer yearlings after an adrenocorticotropic hormone (ACTH) challenge or control saline injection by an 11-oxoaetiocholanolone enzyme immunoassay (EIA), to validate a method. A 2.9- to 4.3-fold increase in measured cortisol metabolites in challenged animals after approximately 22 h demonstrated the suitability of this group-specific EIA to monitor adrenocortical activity in respective deer species. To determine faecal cortisol metabolites in fallow deer from a Mediterranean habitat, we collected samples during a 1-year study at Veliki Brijuni Island. The study confirmed seasonal pattern of cortisol release in fallow deer. Higher 11,17-DOA concentrations (median; min–max) were determined for November (99; 50–2,035), March (112; 25–315) and May (92; 40–196 ng/g faeces). Significantly lower concentrations were measured during July (30; 10–195 ng/g faeces). This study indicates that the analysis of faecal glucocorticoid metabolites is a valuable non-invasive technique for monitoring adrenocortical activity in fallow deer. This, together with information about the seasonal pattern of glucocorticoid excretion, could help to improve fallow deer management and welfare, especially in the case of farmed and park animals.     

Black bears Ursus americanus are effective seed dispersers,with a little help from their friends

Black bears Ursus americanus are generally considered effective seed dispersal agents for fleshy‐fruited plants because they can consume hundreds of fruits at once and have large home ranges. Although seedlings can emerge from faecal piles, establishment of such seedlings seems to be infrequent. Removal of seeds from faeces by rodents is often considered seed predation. We show that removal of seeds from bear faeces by seed‐caching rodents in the Sierra Nevada, USA, represents a second phase of seed dispersal that benefits some fleshy‐fruited plants. Using Trail Master infrared cameras to photograph animals and scandium‐46, a gamma‐emitting radionuclide, to track seeds, we determined that deer mice Peromyscus maniculatus removed seeds from bear faeces and cached them in soil. Caches typically contained 1–3 seeds buried 5–10 mm deep. These seeds escaped several sources of mortality by being moved to relatively safe locations, but deer mice also eventually eat many of the cached seeds. A field germination study confirmed that seed burial increased seedling emergence. Rodents removed seeds in bear faeces more quickly than those in bird faeces in one year, but seeds in bird faeces were removed faster in another year. Results varied across two years, probably because of availability of alternative food sources or changes in deer mice population sizes. The two‐phase seed dispersal syndrome described here may be important in understanding seed dispersal by carnivores and large ungulates that produce large faecal deposits containing many relatively large seeds.     

Spatial and Temporal Analysis of Contact Rates in Female White-Tailed Deer

Abstract: White-tailed deer (Odocoileus virginianus) are important game mammals and potential reservoirs of diseases of domestic livestock; thus, diseases of deer are of great concern to wildlife managers. Contact, either direct or indirect, is necessary for disease transmission, but we know little about the ecological contexts that promote intrasexual contact among deer. Using pair-wise direct contacts estimated from Global Positioning System collar locations and joint utilization distributions (JUDs), we assessed habitats in which contacts occur to test whether direct contact rates among female white-tailed deer in different social groups differs among land-cover types. We also tested whether contact rates differed among seasons, lunar phases, and times of day. We obtained locations from 27 female deer for periods of 0.5–17 months during 2002–2006. We designated any simultaneous pair of locations for 2 deer <25 m apart as a direct contact. For each season, we used compositional analysis to compare land-cover types where 2 deer had contact to available land-cover weighted by their JUD. We used mixed-model logistic regression to test for effects of season, lunar phase, and time of day on contact rates. Contact rates during the gestation season were greater than expected from random use in forest and grassland cover, whereas contact rates during the fawning period were greater in agricultural fields than in other land-cover types. Contact rates were greatest during the rut and lowest in summer. Diel patterns of contact rates varied with season, and contact rates were elevated during full moon compared to other lunar periods. Both spatial and temporal analyses suggest that contact between female deer in different social groups occurs mainly during feeding, which highlights the potential impact of food distribution and habitat on contact rates among deer. By using methods to associate contacts and land-cover, we have created beneficial tools for more elaborate and detailed studies of disease transmission. Our methods can offer information necessary to develop spatially realistic models of disease transmission in deer.     

老爷岭南部狍冬季移动、卧息生境选择及其适宜性评价

在动物生境研究中,移动生境和卧息生境是生境研究的焦点。开展移动生境和卧息生境选择,并在此基础上进行生境评价,有利于深入了解动物对移动和卧息生境条件的需求,制定科学合理的栖息地保护计划。以东北虎（Panthera tigris altaica）的主要猎物物种之一-狍（Capreolus pygargus）为研究对象,于2017-2019年冬季积雪覆盖期在老爷岭南部通过随机布设28个大样方和84条用于足迹链跟踪的样线收集狍的移动点和卧息点信息,再结合近年来收集的东北虎出现点,利用广义可加模型（GAM）和最大熵模型（MaxEnt）进行狍移动、卧息生境选择及评价研究。移动生境选择研究表明,狍在移动的过程中偏好选择坡度小、距农田距离>500 m、远离道路、居民点和低海拔或较高海拔的区域;移动生境评价分析表明,移动适宜和次适宜生境面积之和为1318.16 km²,占研究区域面积的51.28%,当加入虎活动点影响因子后,狍移动适宜和次适宜生境面积之和为901.52 km²,适宜和次适宜生境面积之和减少了31.61%。狍卧息生境选择研究表明,水源、农田、道路和雪深是影响狍卧息的关键因素,其中雪深对狍卧息生境选择的贡献率达到70.13%;卧息生境评价表明,卧息适宜和次适宜生境面积之和为1243.77 km²,占研究区域面积的48.39%,当加入虎出现点因子后,适宜生境和次适宜生境面积之和减少了61.00%,仅为485.02 km²。研究认为,虎的出现对狍移动和卧息生境选择均产生影响,虎的活动及捕食行为可能会减少狍的活动范围和频次,狍远离虎活动区域卧息休息,压缩了狍适宜卧息的空间。     

Estimating dung decay rates of roe deer (<Emphasis Type="Italic">Capreolus capreolus</Emphasis>) in different habitat types of a Mediterranean ecosystem: an information theory approach

For elusive species living in concealing habitats (e.g. deer in a forest habitat), indirect methods such as faecal pellet counts are considered more practical means of estimating population density and abundance. Accurate estimation of deer density using the faecal standing crop (FSC) method necessitates the reliable estimation of the mean time to decay of pellet groups present during the survey. Mean time to decay is generally habitat specific, and separate estimations should be made for each habitat type in the study area. In a confined mountainous area of Greece, the habitat-specific mean time to decay of roe deer pellet groups was estimated by locating and marking fresh pellet groups on several dates in the lead up to an FSC survey and returning to the marked signs at the time of the survey to record whether or not each pellet group had survived. Several logistic models were fitted to the data, and estimations were based on a multi-model inference (MMI) approach according to information theory. The highest mean time to decay was estimated in coniferous forests, while mid-ranged values were found in maquis shrubs, and the lowest mean time to decay was observed in open areas. MMI by model averaging, based on Akaike weights, is recommended for making robust parameter estimations and for dealing with uncertainty in model selection.     

黑麂粪便DNA提取及其PCR检测

采集了黑麂(Muntiacus crinifrons)的新鲜粪便以及在野外自然条件下保存较长时间的粪便样品,晾干后带回实验室,提取其DNA;同时提取黑麂肌肉、皮张样品的DNA,用以对比粪便样品的提取效果。电泳检测结果显示,此方法使用实验室中常用的分子生物学试剂,可以从黑麂粪便样品中抽提到高质量的粪便DNA并克服分子粪便学研究中常见的PCR反应抑制物的影响。为其它濒危鹿科动物的非损伤性取样提供了的新途径,为其遗传结构、遗传多样性现状等研究提供了更加广阔的取材空间。     

The impacts of landscape structure on the winter movements and habitat selection of female red deer

An area of research that has recently gained more attention is to understand how species respond to environmental change such as the landscape structure and fragmentation. Movement is crucial to select habitats but the landscape structure influences the movement patterns of animals. Characterising the movement characteristics, utilisation distribution (UD) and habitat selection of a single species in different landscapes can provide important insights into species response to changes in the landscape. We investigate these three fields in female red deer (Cervus elaphus) in southern Sweden, in order to understand how landscape structure influences their movement and feeding patterns. Movements are compared between two regions, one dominated by a fragmented agriculture–forest mosaic and the other by managed homogenous forest. Red deer in the agriculture-dominated landscape had larger UDs compared to those in the forest-dominated area, moved larger distances between feeding and resting and left cover later in the day but used a similar duration for their movements, suggesting faster travelling speeds between resting and feeding locations. The habitat selection patterns of red deer indicate a trade-off between forage and cover, selecting for habitats that provide shelter during the day and forage by night. However, the level of trade-off, mediated through movement and space use patterns, is influenced by the landscape structure. Our approach provides further understanding of the link between individual animal space use and changing landscapes and can be applied to many species able to carry tracking devices.     

The relative densities of red (Cervus elaphus) and roe (Capreolus capreolus) deer and their relationship in Scottish plantation forests

Population densities of red and roe deer were estimated in 20 conifer plantations throughout central and northern Scotland. Ten forests were selected on the basis of culling records that were considered likely to cover the range of possible densities. Ten further forests were selected at random to investigate the relationship between red and roe deer densities. Density assessment was by standing-crop dung-group counts, on 1.5 x 100 m transects. Dung decay rates were estimated in four areas and used to convert the values to deer densities. Red deer densities were estimated to range from 0.3-35 km⁻²; roe deer densities ranged from 0.5-25 km⁻². Preferences for different structural habitats within individual forests (i.e. blocks with different age classes of tree) were inferred from dung densities. Red deer preferred unplanted ground, open-thicket, and prethicket; roe deer likewise preferred prethicket (highly) and open-thicket. Both species avoided closed-canopy habitats. Red and roe deer densities were negatively correlated and the relationship was significant when the data were log transformed ( P < 0.05).     

So close and yet so different: The importance of considering temporal dynamics to understand habitat selection

Habitat selection of animals is influenced by spatial heterogeneity as well as temporal environmental dynamics. In addition, human activities potentially have severe influences on the habitat selection of animals, often resulting in more nocturnal behavior. We investigated seasonal and circadian habitat selection patterns of red deer (Cervus elaphus) on a military training area in Bavaria (Germany). Individual deer ranged on two neighboring, but non-overlapping sites differing in landscape composition and human activity. Using GPS telemetry data, we visually investigated selection patterns and then fitted step-selection functions to assess multiple approaches to account for temporal (i.e., diel and seasonal) effects and for the impact of habitat conditions on selection. We first showed that the way in which time of day is considered in step-selection functions is essential for obtaining ecologically meaningful results, and that the usual ways of including time (i.e., either based on clock-time or based on night vs. day categories) can lead to incomplete or misleading conclusions. Furthermore, we found that individuals followed either circadian or seasonal habitat selection patterns, depending on the site they inhabited. This can be explained by differences in disturbance between the two sites. Except in winter, individuals selected for open habitats during the night irrespective of human activity level, but only individuals from the more disturbed site selected for covered habitats during the day. Our results highlight the importance of carefully considering the appropriate temporal scale for habitat selection analyses. Our findings also indicate that red deer are not a crepuscular or nocturnal species per se, as is often observed in human-dominated landscapes. Instead, our results imply that nocturnality in red deer is an effect of adaption to human activities.     

Seasonal changes in the dietary compositions of rusa deer in Pangandaran Nature Reserve,West Java,Indonesia

Information regarding the food habits of tropical cervids is limited. We studied the food habits of rusa deer (Rusa timorensis), a tropical cervid in the Pangandaran Nature Reserve, West Java, Indonesia. The faecal composition of the rusa deer collected over a year was analysed using microscopy of plant fragments. The results showed marked seasonal changes: in the rainy season, the rusa deer mainly fed on Cynodon dactylon, a lawn-like grass, which accounted for approximately 50% of the faecal composition. During the dry season, C. dactylon in the faeces decreased to approximately 20%, while the composition of fruits increased, suggesting that food conditions for the deer were better in this season. This pattern differs from that of temperate cervids which can feed on abundant plant leaves during summer but are forced to consume low-quality foods such as bark and dead leaves during winter.     

Establishing baseline values of parameters potentially indicative of chronic stress in red deer (<Emphasis Type="Italic">Cervus elaphus</Emphasis>) from different habitats in western Germany

The use of habitats by wild animals is commonly assumed to be decreasing due to human activities, such as tourism or the installation and use of wind-energy plants. These anthropogenic interferences may subject animals to chronic stress. To be able to objectively characterise the effects on animal populations or on individual animals, the collection of data that might be suitable to monitor such chronic stress is required. In this study of hunted red deer, we report data that are related to adrenal activity and are not affected by the acute stress induced by hunting. Adrenal glands and samples from ileal digesta were collected from 75 hunted deer from seven different habitats in the German Rhineland. The adrenal glands were evaluated histomorphometrically; in the digesta, the concentration of cortisol metabolites, i.e. of 11,17-dioxoandrostanes (11,17-DOA), was measured. Digesta were also examined for parasites. Animals were grouped according to age, sex, habitat, and hunting method. Animals were infected with gastrointestinal helminths and lungworms; examination for liver flukes was negative. Significant differences were not established among the different groups for any of the recorded parameters. For sex, a tendency (P=0.11) towards higher DOA levels was observed in female deer when compared to male deer. The variability of the parameters together with the lack of identifiable influences of hunting indicates that chronic stress might indeed have been a relevant factor. However, none of the parameters analysed can presently be validly used to evaluate habitat quality for red deer since physiological stressors cannot be differentiated.     

Temporal variation in habitat selection breaks the catch‐22 of spatially contrasting predation risk from multiple predators

Predator avoidance depends on prey being able to discern how risk varies in space and time, but this is made considerably more complicated if risk is simultaneously present from multiple predators. This is the situation for an increasing number of mammalian prey species, as large carnivores recover or are reintroduced in ecosystems on several continents. Roe deer Capreolus capreolus in southern Norway illustrate a case in which prey face two predators with contrasting patterns of predation risk. They face a catch‐22 situation: spatially avoiding the risk from one predator (lynx Lynx lynx in dense habitat) implies exposure to the other (hunters in open habitat). Using GPS‐data from 29 roe deer, we tested for daily and seasonal variation in roe deer selection for habitat with respect to the habitats’ year‐round average risk level. Generally, roe deer altered their habitat selection between night and day in a pattern consistent with being able to avoid predicted risk from the nocturnal lynx during night and predicted risk from human hunters during day. However, seasonal variation in habitat selection only partially corresponded with the predicted seasonal variation in risk. Whereas roe deer avoided areas with high risk from hunters more strongly during the hunting season than in other seasons, there was a lack of selection towards areas and time periods lowering the risk of lynx predation during winter. It seems likely that the risk of starvation and thermal stress constrain roe deer habitat selection during this energetically challenging season with cold temperatures, snow cover and limited natural forage. The habitat selection pattern of roe deer fits thus only partly with the two contrasting risk gradients they face. Adjusting risk‐avoidance behavior temporally can be an adaptive response in the case of several predators whose predation patterns differ in space and time.     

Identification of the endangered small red brocket deer (Mazama bororo) using noninvasive genetic techniques (Mammalia; Cervidae)

The small red brocket deer Mazama bororo is one of the most endangered deer in the Neotropics. The great morphological similarities with three other sympatric brocket deer species, coupled with the fact that they inhabit densely forested habitats complicate detection and prevent the use of traditional methodologies for accurate identification of species. The ability to determine the presence of this endangered species in an area is crucial for estimating its distribution range, and is critical for establishing conservation management strategies. Here we describe a fast and reliable noninvasive genetic method for species identification of Mazama species from faeces. We designed a primer set that amplifies a short 224-bp fragment of the cytochrome b and demonstrate its effectiveness in successful amplification of DNA isolated from both tissue and faecal samples. This fragment contains a BSTNI/ECORII digestion site that is unique to the endangered M. bororo. The digested polymerase chain reaction products yielded a 160-bp fragment that is clearly visible in a 2% agarose gel. Two other diagnostic sites were identified to differentiate the other three sympatric species, SspI (M. gouazoubira) and AflIII (M. americana, and M. nana).     

The role of deer as vehicles to move ticks, Ixodes ricinus, between contrasting habitats

In Europe the most important hosts maintaining Ixodes ricinus tick populations are deer. Therefore, excluding deer by fencing or culling are potential tick management tools. Here we test the hypothesis that deer act as vehicles for moving ticks between two distinct habitats: forest and open heather moorland. We utilised an ideal “natural experiment” whereby forests were either fenced or unfenced to prevent or allow deer to move between habitats. We aimed to test the hypothesis that deer cause a net movement of ticks from high tick density areas, i.e. forests, to low tick density areas, i.e. open moorland. We recorded I. ricinus and host abundance in 10 unfenced and seven fenced forests and their respective surrounding heather moorland. We found that fenced forests had fewer deer and fewer I. ricinus nymphs than unfenced forests. However, we found no evidence that fencing forests reduced I. ricinus abundance on adjacent heather moorland. Thus there was insufficient evidence for our hypothesis that deer cause a net movement of ticks from forest onto adjacent moorland. However, we found that deer abundance generally correlates with I. ricinus abundance. We conclude that fencing can be used as a tool to reduce ticks and disease risk in forests, but that fencing forests is unlikely to reduce ticks or disease risk on adjacent moorland. Instead, reducing deer numbers could be a potential tool to reduce tick abundance with implications for disease mitigation.     

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

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

Thermal constraints and optimization of winter feeding and habitat choice in white-tailed deer

I examined the activity and habitat choice of white-tailed deer in winter to determine how deer balance feeding in habitats which may expose them to cold with resting in sheltered habitats. Average heat gain or loss of deer for feeding, resting and walking in four habitats in each of 24 h were estimated for two winter periods (early-mid and late winter). These values were used in a dynamic optimization model to predict the hourly behavior-habitat choices which enable deer to maintain thermal homeostasis, minimize the risk of exceeding lethal body temperatures and satisfy daily energy requirements. The behavior and habitat choices observed under natural conditions were consistent with the model predictions. Deer foraged for the maximum time that was thermally possible in both sampling periods and they used exposed habitats without incurring high heat losses. Deer appeared to avoid overheating in sheltered habitats during daylight and avoid hypothermia in exposed habitats at night.