Resource partitioning between large herbivores in Hustai National Park, Mongolia

Re-introduced Przewalski horses in Hustai National Park, Mongolia could suffer from food competition with other herbivore species through food resource depletion. Diet composition of the Przewalski horse (Equus ferus przewalskii), red deer (Cervus elaphus) and four livestock species (sheep, goat, cattle and horse) were studied, using micro histological analysis of faecal samples in the summer of 2005 and winter of 2006 – 2007. We expected that herbivores become less selective in food choice in winter regarding to summer, resulting in a larger diet breadth, a larger similarity in diet and a larger dietary overlap in winter, potentially triggering exploitative competition by depletion of shared resources. Vegetation biomass decreased during winter, and the different herbivores species in HNP changed their diet from summer to winter. As expected diet breadth, diet similarity and dietary overlap were significantly larger in winter in comparison to summer. The existence of competition by resource depletion between the different species cannot be ruled out. Vegetation biomass was probably not a limiting factor according to the correlation between annual rainfall and herbivore species biomass, however the forage quality may be limiting, triggering competition.     

Reflections of ecological differences? Stress responses of sympatric Alpine chamois and red deer to weather,forage quality,and human disturbance

Depending on the habitats they live in, temperate ungulates have adapted to different degrees to seasonally changing forage and weather conditions, and to specific escape strategies from predators. Alpine chamois, a mountain ungulate, and red deer, originally adapted to open plains, would therefore be expected to differ in their physiological responses to potential stressors. Based on 742 chamois and 1557 red deer fecal samples collected year‐round every 2 weeks for 4 years at the same locations within a strictly protected area in the Swiss Alps, we analyzed glucocorticoid metabolite (FGM) concentrations for both species. Results from linear mixed effects models revealed no physiological stress response to changing visitor numbers, but instead to drought conditions for both species during summer. In winter, FGM concentrations increased with increasing snow height in both species, but this response was modulated by temperature in red deer. Chamois showed a stronger stress response to increasing snow height during November and December than between January and March, while FGM concentrations increased with decreasing temperature throughout winter. An increase in FGM concentrations with decreasing forage digestibility during winter was found only for red deer. The results are thus partly in contradiction to expectations based on feeding type and adaptations to different habitats between the two species. The lack of a response to forage digestibility in chamois may reflect either better adaptation to difficult feeding conditions in subalpine forests, or, by contrast, strong constraints imposed by forage quality. The similar responses of both species to weather conditions in winter suggest that climatic factors at the elevations examined here are sufficiently harsh to be limiting to temperate ungulates regardless of their specific adaptations to this environment.     

Spatial variability in wolf diet and prey selection in Galicia (NW Spain)

We studied wolf (Canis lupus) diet for three different landscapes in the north-western Iberian Peninsula, differing in land uses and availability of food for wolves. We examined 2740 scats, collected over a period of 4 years, in order to describe wolf diet, its geographic variation, and trophic preferences. The most consumed species were wild pony, roe deer and cattle. We observed differences in wolf diet among the three study sites, related to the availability and accessibility of food resources in each habitat. For the two study sites in northern and central Galicia, wolves showed similar diet, with high occurrence of wild pony (37 vs. 34%) and cattle (20 vs. 23%), but differing in the consumption of wild ungulates (16 vs. 8%) and carrion (7 vs. 14%). Wolf diet in eastern Galicia’s mountain ranges was entirely different, due to the higher consumption of wild ungulates (70%). Wolves showed clear prey selection patterns. Between wild ponies and livestock, wolves positively selected ponies. Among wild ungulates, wolves positively selected roe deer. Wild pony and roe deer are key species for wolf feeding in Galicia. In the Galician wild pony range, ponies are the main food for wolves. Given that the availability of wild ponies may contribute to the decrease in wolf predation on cattle, it is essential to develop innovative administrative decisions in such areas to preserve this traditional equid population. In the same way, the population of roe deer should be strengthened in the livestock areas outside the range of wild pony.     

Partial migration in expanding red deer populations at northern latitudes – a role for density dependence?

Partial migration is common in ungulates living in highly seasonal environments. Typically, at higher latitudes, this involves movement between high elevation summer areas used during breeding and lowland areas with less snow used during winter. Snow depth is regarded the main cause of migration to low elevation, but it is less clear why deer migrate to high elevation in spring. The forage maturation hypothesis explains the upward migration due to plant phenology. We here present also an alternative and non‐exclusive hypothesis, that deer migrate uphill in summer to escape competition due to the high density in winter areas (the competition avoidance hypothesis). We also suggest that social fences may play a role at high population density. Based on a unique study of 141 GPS‐marked red deer from seven regions covering the main distribution in Norway, we found that the proportion of migrants in the populations varied from 38% to 100%. Migration was more common in areas with a diverse topography, i.e. for areas with access to high elevation. Further, we found evidence that migration was negatively density dependent, and that fall migration was delayed at high density. We suggest that a combination of avoidance of competition in high density winter ranges, social fencing during summer in addition to the forage maturation and predation risk avoidance hypotheses, is needed to explain migration patterns of northern ungulates.     

Diet overlap among ruminants in Fennoscandia

Information on overlap in resource use is central to understanding of interspecific exploitation competition and resource partitioning. Despite this, measures of diet overlap among northern ruminants in Fennoscandia is limited to one earlier study (reindeer and sheep). Diet overlap between sympatric moose and roe deer calculated with Schoener’s index was 20.7% and 33.6% during summer (data from one area) and winter (data from two areas), respectively, whereas average diet overlap between moose and red deer was 32.0% during winter (data from four areas). Diet overlap between a coastal island population of red deer and sheep was 59.3% during summer and 63.9% during winter. Summer diet overlap between a sheep and a goat population and a sheep and a reindeer population calculated with data on main types of forage plants was 77.0% and 55.1%, respectively. However, overlap calculated with main plant groups was sometimes considerably higher than when calculated for individual forage species. Neither difference in feeding type nor body mass successfully predicted diet overlap between species pairs (n=9), although there tended to be negative correlation (r _p =–0.586, P=0.098) between diet overlap of main plant groups (calculated across studies) and difference in feeding type. Received: 19 October 1999 / Accepted: 31 January 2000     

Seasonal differences between moose and roe deer in ability to digest browse

The digestive capacity of free-living moose Alces alces (L.) and roe deer Capreolus capreolus (L.) for four plant species was studied with in vitro technique. Moose were found to digest all species better than roe deer. Individual variations were also smaller for moose. In moose the level of digestibility was the same both summer and winter, while roe deer shot in winter digested winter samples of bilberry Vaccinium myrtillus L. and to some extent Scots pine Pinus silvestris L. better than did summer roe deer. This was taken as evidence for a seasonal adaptation in the roe deer rumen. Digestibility of the plant material studied showed a strong negative correlation to the fiber content. The results are discussed in relation to winter problems for roe deer and possible food competition between moose and roe deer.     

Food–niche relationships within a guild of alpine ungulates including an introduced species

To assess competition for trophic resources within a guild of ungulates, we studied the food niches of sympatric ungulates in an alpine area of the Italian Western Alps and evaluated the extent of their dietary overlap, especially with regard to the presence of the introduced mouflon Ovis musimon . The roe deer Capreolus capreolus had the most diverse diet including dicotyledons, young sprouts of trees and an unusually high percentage of graminoids. The red deer Cervus elaphus and chamois Rupicapra rupricapra mainly consumed graminoids, supplemented with dicotyledonous herbs and woody plants. The mouflon was a grass feeder, with graminoids representing the majority of items ingested in all seasons. Among the native species, we recorded a high overlap of feeding choices between red deer and chamois, which frequently used the same areas and are both intermediate feeders. The mouflon showed considerable dietary overlap with the chamois and red deer in all seasons, while the similarity with roe deer was low. High food–niche overlap can imply competition but only if resources are limited and the species use the same habitats.     

穆棱林区野生东北马鹿种群冬季食物组成模式与植物代谢产物的关系

高纬度地带,在冬季食物资源有限的环境中,野生大型有蹄类动物满足营养需求的同时,需要对植物中次生代谢产物进行平衡,回避有害物质并选择对机体有益的成分,从而形成特定的食物组成模式。以东北马鹿（Cervus elaphus xanthopygus）为研究对象,于2020年11月,在黑龙江穆棱东北红豆杉国家级自然保护区境内,采集东北马鹿粪便和植物样本。通过粪便显微分析法确定保护区内马鹿冬季食物组成,采用k-means聚类分析揭示马鹿冬季食物组成模式。应用广泛靶向代谢组技术对部分植物中次生代谢产物的含量进行全覆盖定性和相对定量检测,将食物组成与次生代谢产物数据整合,进行曼特尔检验（Mantel test）分析,以探究植物次生代谢产物对马鹿种群内食物组成模式的影响。结果表明,林区内马鹿种群冬季共采食30种植物,其中木本植物占92.48%;且种群内分别呈现出以东北红豆杉（Taxus cuspidata）,簇毛槭（Acer barbinerve）,毛榛子（Corylus mandshurica）为主要食物的不同食物组成模式。广泛靶向代谢组技术共检测出638种次生代谢产物,有25种代谢物与马鹿采食频率显著相关,其中多数萜类物质抑制马鹿采食,而鞣质类物质对马鹿的采食选择有一定的正向作用;Mantel test结果显示,上述25种物质中黄酮、鞣质、萜类化合物相对含量与不同马鹿个体食物组成显著相关,说明这些代谢物相对含量和性质的差异会造成种群内不同个体食物组成的差异,是种群内形成不同食物组成模式的原因之一。从植物次生代谢产物角度揭示了该地区东北马鹿种群冬季食物组成模式呈现差异的可能因素,为进一步研究大型有蹄类营养策略和植物化学防御关系提供基础依据。     

Heterogeneity in Primary Productivity Influences Competitive Interactions between Red Deer and Alpine Chamois

Habitat heterogeneity can promote coexistence between herbivores of different body size limited to different extents by resource quantity and quality. Red deer (Cervus elaphus) are known as superior competitors to smaller species with similar diets. We compared competitive interactions and habitat use between red deer and Alpine chamois (Rupicapra rupicapra) in two adjacent valleys in a strictly protected area in the Central Alps. Red deer density was higher in the valley with higher primary productivity. Only here was horn growth in kid and yearling chamois (as a measure for body condition) negatively correlated with red deer population size, suggesting interspecific competition, and chamois selected meadows with steeper slopes and lower productivity than available on average. Conversely, red deer selected meadows of high productivity, particularly in the poorer area. As these were located mainly at lower elevations, this led to strong altitudinal segregation between the two species here. Local differences in interspecific competition thus coincided with differences in habitat preference and–segregation between areas. This suggests that spatial habitat and resource heterogeneity at the scale of adjacent valleys can provide competition refuges for competitively inferior mountain ungulates which differ from their superior competitor in their metabolic requirements.     

Habitat selection by an Alpine ungulate: the significance of forage characteristics varies with scale and season

Habitat selection in ungulates should ensure access to abundant forage of sufficiently high quality. Species living in rugged mountain areas have to face nutritional bottlenecks regularly and should show particularly sophisticated habitat selection behaviour. However, patterns and mechanisms of such adaptations remain little studied. We analysed habitat selection and its seasonal variability of 10 GPS‐collared red deer Cervus elaphus living in a topographically challenging landscape of the Swiss Alps. We hypothesised that resource selection by red deer was scale‐dependent and predicted that scale‐dependence would vary among seasons in relation to seasonal changes of available forage biomass and quality, which we sampled across the entire study area of 250 km². The studied population of Alpine red deer undertook altitudinal migrations and showed scale‐dependent habitat selection that was strongest in winter and declined through spring and summer. Selection occurred mostly at the larger (landscape/home‐range location) scale and less so at the smaller (within home‐range) scale. Topographic parameters were selected mainly at the landscape scale and mostly in winter. About 70% of all instances of preference for habitat parameters were associated with above‐average forage characteristics, represented mostly by higher crude protein content, in a few cases also by higher biomass or both. The overall pattern of space use by red deer characterised by migration and seasonal habitat selection was therefore closely linked to the quality of food resources, although some trade‐offs with avoiding human disturbance may also have been involved.     

Influence of interspecific competition on mule deer birthing and rearing site selection

Ungulates often alter behavior and space use in response to interspecific competition. Despite observable changes in behavior caused by competitive interactions, research describing the effects of competition on survival or growth is lacking. We used spatial modeling to determine if habitat use by female mule deer (Odocoileus hemionus) was affected by other ungulate species prior to, during, and after parturition. We conducted our study in the Book Cliffs region of eastern Utah, USA, during 2019 and 2020. We used resource selection function (RSF) analysis to model space use of 4 ungulate species that potentially competed with mule deer: bison (Bos bison), cattle, elk (Cervus canadensis), and feral horses. We incorporated RSF models for competing species into a random forest analysis to determine if space use by mule deer was influenced by these other ungulate species. We used survival and growth data from neonate mule deer to directly assess potential negative effects of other ungulates. Habitat use by elk was an important variable in predicting use locations of mule deer during birthing and rearing. The relationship was positive, suggesting interference competition was not occurring. Survival of neonate mule deer increased as the probability of use by elk increased (hazard ratio = 0.185 ± 0.497 [SE]). Further, probability of use by elk in rearing habitat had no influence on growth of neonate mule deer from birth to 6 months of age, suggesting that exploitative competition was not occurring.     

Linking alternative food sources to winter habitat selection of herbivores in overbrowsed landscapes

During winter, ungulates in boreal forests must cope with high energetic costs related to locomotion in deep snow and reduced forage abundance and quality. At high density, ungulates face additional constraints, because heavy browsing reduces availability of woody browse, the main source of forage during winter. Under these severe conditions, large herbivores might forage on alternative food sources likely independent of browsing pressure, such as litterfall or windblown trees. We investigated the influence of alternative food sources on winter habitat selection, by studying female white-tailed deer (Odocoileus virginianus) living in 2 landscapes with contrasted browse abundance, recently logged and regenerated landscapes, in a population at high density and on a large island free of predators. We fitted 21 female white-tailed deer with Global Positioning System (GPS) collars and delineated winter home ranges and core areas. We measured snow conditions in different habitat categories and sampled vegetation in the core areas and in the rest of the home ranges to determine how forage abundance, protective cover, and snow conditions influenced habitat selection within the home range. In both landscapes, deer were less likely to use open habitat categories as snow accumulated on the ground. At a finer scale, deer inhabiting the regenerated landscape intensively used areas where balsam fir cover was intermediate with greater balsam fir browse density than in the rest of the home range. In the recently logged landscape, deer were more likely to be found near edges between clear-cuts and balsam fir stands and in areas where windblown balsam fir trees were present; the latter being the most influential variable. Although balsam fir browse was sparse and mainly out of reach in this landscape, deer increased the use of areas where it was present. Our results offer novel insights into the resource selection processes of northern ungulates, as we showed that access to winter forage, such as woody browse and alternative food sources, depends on climatic conditions and stochastic events, such as abundant compacted snow or windthrows. To compensate for these scarce and unpredictable food supplies, deer selected habitat categories, but mostly areas within those habitat categories, where the likelihood of finding browse, litterfall, and windblown trees was greatest. © 2011 The Wildlife Society.     

Factors affecting the seasonal distribution of red deer (Cervus elaphus) at Glen Dye, north-east Scotland

Part of a red deer herd found on an estate in north-east Scotland in summer spent more time on neighbouring ground during the winter where they damaged farms and woods. Factors associated with the deer's seasonal distribution were studied to help understand how to keep more deer on the estate in winter. There were no indications that food species or food quality were responsible for the observed changes in deer distribution, or that other animals or disturbance influenced distribution over a long period. Deer sought shelter from chilling winds, and areas occupied by deer in winter had more sheltered ground from wind, and the quality of this shelter was higher than on the areas little used. Home-range behaviour and tradition appeared to determine which areas particular deer occupied in winter. The main recommendation for management was to re-open mature woodland to provide more and better shelter on the estate in order to get more control over the deer stock and to reduce marauding.     

Why do migrants move downhill? The effects of increasing predation and density on red deer altitudinal migration in temperate Carpathian forests

Resource selection by ungulates is driven by trade-offs between foraging and predation avoidance or by intraspecific competition. Ungulates use migratory flexibility to optimize access to spatially and temporally variable resources across seasons, sometimes even adaptively switching between migrant and resident strategy as conditions change. After an 80% increase in red deer (Cervus elaphus) population and simultaneous recovery of wolves (Canis lupus) in the Kremnica Mountains, Slovakia, a significant portion of the deer population started to migrate downhill (<?700 m) to marginal habitats during winter. Building on available spatial data on forage availability, predation risk, and deer abundance, we tested for differences in habitat selection of migrant and resident male red deer to assess possible reasons for this change. On high-altitude (700–1100 m) summer ranges, deer were not forced to trade-off forage to avoid predation within their home ranges. However, during winter, residents remaining on high-altitude ranges selected for areas with highly abundant forage only under low predation risk or at high deer abundance. Downhill migration exposed migrants to 15% lower forage availability but simultaneously reduced wolf predation risk by 39% relative to residents. Consequently, the limited access to forage resources at low-altitude ranges have reduced antler growth, especially in young males. Our study represents one of few that address the role of predation risk in driving seasonal migrations in temperate systems where snow is not likely to be the major driver of migration to low-altitude winter ranges.     

Influence of digestive morphology on resource partitioning in Amazonian ungulates

Summary Resource partitioning of diet and habitat use was studied in the entire Amazonian ungulate community of Northeastern Peru, which comprises the red brocket deer (Mazama americana), grey brocket deer (M. gouazoubira), collared peccary (Tayassu tajacu), white-lipped peccary (T. pecari), and lowland tapir (Tapirus terrestris). Each ungulate species partitioned at least one type of resource from every other species. Digestive morphology had a greater influence on resource partitioning of diet than body size. Neither digestive morphology nor body size were related to segregation of habitats. However, species with similar diets partitioned habitats, whereas species with different diets often used the same type of forest. Increases in habitat breadth of ungulates were positively correlated with increases in dietary breadth.     

Facilitative and competitive interactions between sympatric cattle, red deer and wild boar in Dutch woodland pastures

Use of cattle-grazed and ungrazed woodland pastures by red deerCervus elaphus Linnaeus, 1758 and wild boarSus scrofa Linnaeus, 1758 was investigated monthly by measuring dung-deposition rates. CattleBos taurus grazed pastures year-round, with peak intensities during the growing season (May–September). Red deer and wild boar grazed pastures primarily during autumn and winter (October–April) when cattle occupancy was at a minimum. The lower occupancy of cattle in pastures from November to April was interpreted as the result of competition with red deer. Mean sward height in this period fell below 6.5 cm. In autumn and winter a negative relationship was found for red deer and wild boar occupancy with sward height, which indicated that red deer and wild boar preferred swards previously grazed by cattle. At the start of the growing season, when cattle occupancy in the pastures increased, red deer switched their habitat preference and almost totally disappeared from pastures to use alternative feeding grounds. Interpretation of the results lead to the conclusion that facilitative and competitive interactions occurred between sympatric cattle and red deer in woodland pastures, and to some extent also between cattle and wild boar.     

Spatio-temporal segregation of facultative avian scavengers at ungulate carcasses

Spatial and/or temporal segregation of resource use are mechanisms that may allow coexistence between potential competitors. Spatial and temporal patterns of carrion use were studied in the main avian scavengers of Sierra Espuña Natural Park (SE, Spain). We monitored the use of ungulate carcasses provided by hunting during winter and summer of 2005–2006. Non-breeding ravens exploited aggregated carcasses and depleted the resource in a few days while golden eagles used scattered carcasses over a longer consumption period. Moreover, non-breeding ravens exploited carcasses when golden eagles were less active. Almost all available hunter kills were exploited by golden eagles and/or ravens during winter whereas a high number of carcasses were not used during the breeding season, suggesting strong competition in a period of low food availability. Thus carrion arrangement, seasonal behaviour and competitive interactions could be affecting the foraging patterns of these two species. Interestingly, intraspecific aggregation of the smaller species and dominance of the larger one may act on the use of shared resources and trigger segregation mechanisms. Our results support that differences in the spatial and temporal patterns of resource use may allow resource partitioning between two species, thus facilitating their coexistence in sympatric areas.     

Impact of Red Deer Cervus elaphus Grazing on Bilberry Vaccinium myrtillus and Composition of Ground Beetle (Coleoptera, Carabidae) Assemblage

We studied the role of red deer Cervus elaphus L. as ecosystem modifier in boreal forest (Tingvoll municipality, 62°52′ N, 8°20′ E, Norway), during early summer of 2001. The effect of grazing by red deer on ground beetles (Carabidae) abundance and diversity was investigated across a gradient of grazing pressures. We trapped ground beetles by pit-fall traps from three homogeneous winter grazing areas (ungrazed, medium grazed, heavily grazed). Bilberry Vaccinium myrtillus (the main winter food for red deer) was sampled and its dry weight was measured for the three locations. Gradient analyses showed that grazing by red deer affects carabid species composition. Grazing significantly affected the amount of bilberry, which correlated with species variation. According to our predictions, we found a higher abundance of carabids in the heavily grazed location, but the species richness and the diversity indices were similar for the three areas. This study shows that overall species composition is altered along a gradient as consequence of red deer winter grazing and that red deer act as ecosystem engineer, by reducing the bilberry heather which dominates the field layer in early summer.     

Red deer habitat selection and movements in relation to roads

Roads affect wildlife in many direct and indirect ways. For ungulates, roads may inhibit seasonal migration and may cause an effective loss of habitat due to avoidance. On the other hand, roadsides and associated agricultural lands offer high quality forage that may attract ungulates and increase the frequency of car accidents. Mitigating actions require detailed knowledge on space use in relation to roads. Using data from 67 global positioning system (GPS)-marked red deer in Norway, we quantified 1) scale of avoidance of roads, 2) crossing frequency, and 3) selection of crossing sites. Red deer avoided roads only on a very local scale and only during daytime, with minor influence of variation in road size (traffic burden). Marked red deer crossed roads, on average, 2 times per day. Females crossed more frequently than males and crossings were most frequent during autumn and winter and during night. Deer selected forested crossing sites close to agricultural pastures, reflecting that roads are crossed most often on nightly feeding excursions. Our findings imply that red deer in our study area have adjusted to exploit feeding habitat close to roads at times of low traffic burden. The high frequency of crossings suggests a limited influence on seasonal migration patterns. The frequency at which red deer cross highways suggests that mitigation measures to reduce road mortality may be effective if targeted in the right areas. © 2012 The Wildlife Society.     

Solar Radiation Determines Site Occupancy of Coexisting Tropical and Temperate Deer Species Introduced to New Zealand Forests

Assemblages of introduced taxa provide an opportunity to understand how abiotic and biotic factors shape habitat use by coexisting species. We tested hypotheses about habitat selection by two deer species recently introduced to New Zealand’s temperate rainforests. We hypothesised that, due to different thermoregulatory abilities, rusa deer (Cervus timorensis; a tropical species) would prefer warmer locations in winter than red deer (Cervus elaphus scoticus; a temperate species). Since adult male rusa deer are aggressive in winter (the rut), we also hypothesised that rusa deer and red deer would not use the same winter locations. Finally, we hypothesised that in summer both species would prefer locations with fertile soils that supported more plant species preferred as food. We used a 250 × 250 m grid of 25 remote cameras to collect images in a 100-ha montane study area over two winters and summers. Plant composition, solar radiation, and soil fertility were also determined for each camera location. Multiseason occupancy models revealed that direct solar radiation was the best predictor of occupancy and detection probabilities for rusa deer in winter. Multistate, multiseason occupancy models provided strong evidence that the detection probability of adult male rusa deer was greater in winter and when other rusa deer were present at a location. Red deer mostly vacated the study area in winter. For the one season that had sufficient camera images of both species (summer 2011) to allow two-species occupancy models to be fitted, the detection probability of rusa deer also increased with solar radiation. Detection probability also varied with plant composition for both deer species. We conclude that habitat use by coexisting tropical and temperate deer species in New Zealand likely depends on the interplay between the thermoregulatory and behavioural traits of the deer and the abiotic and biotic features of the habitat.