Macronutrient balancing in free‐ranging populations of moose

At northern latitudes, large spatial and temporal variation in the nutritional composition of available foods poses challenges to wild herbivores trying to satisfy their nutrient requirements. Studies conducted in mostly captive settings have shown that animals from a variety of taxonomic groups deal with this challenge by adjusting the amounts and proportions of available food combinations to achieve a target nutrient balance. In this study, we used proportions‐based nutritional geometry to analyze the nutritional composition of rumen samples collected in winter from 481 moose (Alces alces) in southern Sweden and examine whether free‐ranging moose show comparable patterns of nutrient balancing. Our main hypothesis was that wild moose actively regulate their rumen nutrient composition to offset ecologically imposed variation in the nutritional composition of available foods. To test this, we assessed the macronutritional composition (protein, carbohydrates, and lipids) of rumen contents and commonly eaten foods, including supplementary feed, across populations with contrasting winter diets, spanning an area of approximately 10,000 km². Our results suggest that moose balanced the macronutrient composition of their rumen, with the rumen contents having consistently similar proportional relationship between protein and nonstructural carbohydrates, despite differences in available (and eaten) foods. Furthermore, we found that rumen macronutrient balance was tightly related to ingested levels of dietary fiber (cellulose and hemicellulose), such that the greater the fiber content, the less protein was present in the rumen compared with nonstructural carbohydrates. Our results also suggest that moose benefit from access to a greater variety of trees, shrubs, herbs, and grasses, which provides them with a larger nutritional space to maneuver within. Our findings provide novel theoretical insights into a model species for ungulate nutritional ecology, while also generating data of direct relevance to wildlife and forest management, such as silvicultural or supplementary feeding practices.     

Supplementary winter feeding of wild red deer Cervus elaphus in Europe and North America: justifications, feeding practice and effectiveness

1. Supplementary winter feeding of game animals, and particularly deer, is a common practice throughout northern (continental) Europe and parts of North America. Feeding is normally associated with maintaining high densities of animals for hunting, in terms of: (i) maintaining or increasing body weights and condition overwinter; (ii) improving reproductive performance and fertility; (iii) increasing overwinter survival; and (iv) reducing levels of damage caused to agriculture and forestry or the natural heritage. We consider the balance of evidence on the effectiveness of winter feeding of red deer Cervus elaphus in achieving these objectives. Where that evidence is equivocal, we attempt to reconcile apparent contradictions to evaluate the circumstances under which winter feeding may or may not be effective. 2. In general, feeding of red deer on open range appears to have relatively little effect on body weights or fecundity. Effects on increasing antler size and quality are variable and seem to depend on the degree to which animals may be mineral limited on native range. Effects on survival are similarly ambiguous. It is apparent, however, that to be effective in reducing mortality, any supplementation is required early in the season and not simply when heavy mortalities are already being experienced. If provision of supplementary foods is delayed until animals are perceived already to be in poor condition, such feeding may have little effect. 3. One of the primary goals of winter feeding in both Europe and the USA has become the prevention of environmental damage, particularly damage to commercial and native forests, while maintaining deer populations at densities suitable for hunting. Again, empirical evidence for effectiveness in this regard is inconclusive, with some studies showing a decrease in damage caused, some showing no effect and others showing a significant increase in local impact. 4. There are equally a number of problems associated with the provision of supplementary feeds overwinter. Those animals which come to the feeding stations may develop a reliance on the food supplement provided, reducing intake of natural forages to near zero; where feed provided is less than 100% of daily requirement, such animals may regularly lose, rather than gain condition. Feed provision is also extremely uneven at such feeding stations; dominant stags displace younger stags and hinds from the feed provided until they have themselves finished feeding. Concentrations of high densities of animals around small feed‐areas may also increase the risk of infection and lead to development of high parasite burdens. 5. In an attempt to assess the current status and distribution of supplementary winter feeding in Scotland, a questionnaire was circulated to a number of individual across the country. Results of this survey are summarized and conclusions presented on the likely effectiveness of current feeding practices in achieving their aims.     

Can Wild Ungulate Carcasses Provide Enough Biomass to Maintain Avian Scavenger Populations? An Empirical Assessment Using a Bio-Inspired Computational Model

Background

The reduction in the amount of food available for European avian scavengers as a consequence of restrictive public health policies is a concern for managers and conservationists. Since 2002, the application of several sanitary regulations has limited the availability of feeding resources provided by domestic carcasses, but theoretical studies assessing whether the availability of food resources provided by wild ungulates are enough to cover energetic requirements are lacking.

Methodology/Findings

We assessed food provided by a wild ungulate population in two areas of NE Spain inhabited by three vulture species and developed a P System computational model to assess the effects of the carrion resources provided on their population dynamics. We compared the real population trend with to a hypothetical scenario in which only food provided by wild ungulates was available. Simulation testing of the model suggests that wild ungulates constitute an important food resource in the Pyrenees and the vulture population inhabiting this area could grow if only the food provided by wild ungulates would be available. On the contrary, in the Pre-Pyrenees there is insufficient food to cover the energy requirements of avian scavenger guilds, declining sharply if biomass from domestic animals would not be available.

Conclusions/Significance

Our results suggest that public health legislation can modify scavenger population trends if a large number of domestic ungulate carcasses disappear from the mountains. In this case, food provided by wild ungulates could be not enough and supplementary feeding could be necessary if other alternative food resources are not available (i.e. the reintroduction of wild ungulates), preferably in European Mediterranean scenarios sharing similar and socio-economic conditions where there are low densities of wild ungulates. Managers should anticipate the conservation actions required by assessing food availability and the possible scenarios in order to make the most suitable decisions.     

Ungulates,rodents, shrubs: interactions in a diverse Mediterranean ecosystem

Ungulate abundance has increased dramatically worldwide, having strong impacts on ecosystem functioning. High ungulate densities can reduce the abundance, diversity and/or body condition of small mammals, which has been attributed to reductions in cover shelter and food availability by ungulates. The densities of wild ungulates have increased recently in high-diversity Mediterranean oak ecosystems, where acorn-dispersing small rodents are keystone species. We analysed experimentally ungulate effects on seed-dispersing rodents in two types of oak woodland: a forest with dense shrub layer and in dehesas lacking shrubs. Ungulates had no significant effects on vegetation structure or rodent body mass, but they reduced dramatically rodent abundance in the lacking-shrub dehesas. In the forest, ungulates modified the spatial distribution and space use of rodents, which were more concentrated under shrubs in the presence than in the absence of ungulates. Our results point to the importance of shrubs in mediating ungulate–rodent interactions in Mediterranean areas, suggesting that shrubs serve as shelter for rodents against ungulate physical disturbances such as soil compaction, trampling or rooting. Holm oak seedling density was reduced by ungulates in dehesa plots, but not in forests. Acorn consumption by ungulates may reduce oak recruitment to a great extent. Additionally, we suggest that ungulates may have a negative effect on oak regeneration processes by reducing the abundance of acorn-dispersing rodents. Given that shrubs seem to mediate ungulate effects on acorn dispersers, controlled shrub encroachment could be an effective alternative to ungulate population control or ungulate exclusion for the sustainability of the high-diversity Mediterranean oak ecosystems.     

Rewilding traditional grazing areas affects scavenger assemblages and carcass consumption patterns

The abandonment of traditional livestock farming systems in Mediterranean countries is triggering a large-scale habitat transformation, which, in general, consists of the replacement of open grazing areas by woodlands through non-managed regeneration. As a consequence, wild ungulates are occupying rapidly the empty niche left by domestic ungulates. Both types of ungulates represent the main trophic resource for large vertebrate scavengers. However, a comparison of how vertebrate scavengers consume ungulate carcasses in different habitats with different ungulate species composition is lacking. This knowledge is essential to forecast the possible consequences of the current farmland abandonment on scavenger species. Here, we compared the scavenging patterns of 24 wild and 24 domestic ungulate carcasses in a mountainous region of southern Spain monitored through camera trapping. Our results show that carcasses of domestic ungulates, which concentrate in large numbers in open pasturelands, were detected and consumed earlier than those of wild ungulate carcasses, which frequently occur in much lower densities at more heterogenous habitats such as shrublands and forest. Richness and abundance of scavengers were also higher at domestic ungulate carcasses in open habitats. Vultures, mainly griffons (Gyps fulvus), consumed most of the carcasses, although mammalian facultative scavengers, mainly wild boar (Sus scrofa) and red fox (Vulpes vulpes), also contributed importantly to the consumption of wild ungulate carcasses in areas with higher vegetation cover. Our findings evidence that the abandonment of traditional grazing may entail consequences for the scavenger community, which should be considered by ecologists and wildlife managers.     

Macronutrient Optimization and Seasonal Diet Mixing in a Large Omnivore,the Grizzly Bear: A Geometric Analysis

Nutrient balance is a strong determinant of animal fitness and demography. It is therefore important to understand how the compositions of available foods relate to required balance of nutrients and habitat suitability for animals in the wild. These relationships are, however, complex, particularly for omnivores that often need to compose balanced diets by combining their intake from diverse nutritionally complementary foods. Here we apply geometric models to understand how the nutritional compositions of foods available to an omnivorous member of the order Carnivora, the grizzly bear (Ursus arctos L.), relate to optimal macronutrient intake, and assess the seasonal nutritional constraints on the study population in west-central Alberta, Canada. The models examined the proportion of macronutrients that bears could consume by mixing their diet from food available in each season, and assessed the extent to which bears could consume the ratio of protein to non-protein energy previously demonstrated using captive bears to optimize mass gain. We found that non-selective feeding on ungulate carcasses provided a non-optimal macronutrient balance with surplus protein relative to fat and carbohydrate, reflecting adaptation to an omnivorous lifestyle, and that optimization through feeding selectively on different tissues of ungulate carcasses is unlikely. Bears were, however, able to dilute protein intake to an optimal ratio by mixing their otherwise high-protein diet with carbohydrate-rich fruit. Some individual food items were close to optimally balanced in protein to non-protein energy (e.g. Hedysarum alpinum roots), which may help explain their dietary prevalence. Ants may be consumed particularly as a source of lipids. Overall, our analysis showed that most food available to bears in the study area were high in protein relative to lipid or carbohydrate, suggesting the lack of non-protein energy limits the fitness (e.g. body size and reproduction) and population density of grizzly bears in this ecosystem.     

MUTUALISM OR PARASITISM? USING A PHYLOGENETIC APPROACH TO CHARACTERIZE THE OXPECKER‐UNGULATE RELATIONSHIP

With their striking predilection for perching on African ungulates and eating their ticks, yellow-billed (Buphagus africanus) and red-billed oxpeckers (B. erythrorhynchus) represent one of the few potentially mutualistic relationships among vertebrates. The nature of the oxpecker-ungulate relationship remains uncertain, however, because oxpeckers are known to consume ungulate tissues, suggesting that the relationship between oxpeckers and ungulates may also be parasitic. To examine this issue further, we obtained data on oxpecker preferences for different ungulate species, the abundance of ticks on these ungulates, and ungulate hide thickness. In support of the mutualism hypothesis, we found that both species of oxpeckers prefer ungulate hosts that harbor a higher abundance of ticks. We found no evidence that hide thickness-a measure of the potential for parasitism by oxpeckers-predicts oxpecker preferences for different ungulate species. Oxpeckers also prefer larger-bodied ungulates, possibly because larger animals have more ticks, provide a more stable platform upon which to forage, or support more oxpeckers feeding simultaneously. However, the preference for ungulates with greater tick abundance was independent of host body mass. These results support the hypothesis that the relationship between oxpeckers and ungulates is primarily mutualistic.     

Ungulate herbivory reduces abundance and fluctuations of herbivorous insects in a boreal old-growth forest

Ecological theory predicts the strongest ecosystem effects of herbivory when dominant and ecologically important species are consumed. Bilberry, Vaccinium myrtillus, is such a key plant species, attractive to many other species in the boreal forests, for example ungulate and invertebrate herbivores. Large herbivores may remove substantial biomass and alter plant quality and therefore affect abundance and populations of invertebrate animals sharing the same food plant. We combined experimental exclusion of ungulates with a browsing intensity gradient to investigate the 15-year effect of ungulate (Cervus elaphus and Ovis aries) browsing on bilberry plant size and on bilberry-feeding herbivorous larvae (Lepidoptera and Symphyta), in a Norwegian old growth boreal forest ecosystem. Bilberry ramets in exclosure plots had nearly nine times higher dry mass and three times higher abundance of invertebrates feeding on them than in ungulate-access plots. Sweep-netting data verified these findings as larval numbers were twice as high in exclosure plots. The pattern in the large herbivore effects on bilberry size and abundance of herbivorous larvae were identical along the browsing gradient. Differences in larval abundance between treatments, as indicated by leaf-chewing, increased during the 15-year study period, and the community fluctuations were larger when ungulate herbivores were excluded. The browsing effect was moderated by plant quality as larval densities were lowest on both heavily-browsed and non-browsed plants, and highest on ramets that had 50–74% of annual shoots browsed. Our study supports previous findings in that bilberry is relatively disturbance tolerant and may recover quickly, but that ungulates may compete with herbivorous larvae for food biomass. Additionally, our results strongly indicates that population insect community peaks and fluctuations are dampened by ungulate consumption. Our findings add to the understanding on how ungulates may structure forest ecosystems directly and indirectly.     

Browse supply in lowland forests of eastern Poland

Browse supplies in different growth stages of 4 forest associations and forest site types were estimated. Two ecological situations were compared: exclusion from browsing and unrestrained access of ungulate animals. Harvest technique was applied. Three big forest tracts in eastern Poland provided study locations. Results were tested with the use of analysis of variance. The fresh mixed coniferous forest revealed the highest species diversity in the understorey and regeneration zone, while fresh coniferous forest had the lowest.
No obvious influence of animal exclusion upon the number of plant species was found. Altogether, 41 plant species were found in the understorey and regeneration zones. In young plantations shoot weight was higher than in thickets, pole-sized and timber stands. Shoots clipped in autumn were heavier than those clipped in spring. The browse biomass varied from 400-600 kg ha^-1 in poor communities to 2–3 tonnes ha^-1 in the most productive ones. Generally, low (5%) utilization of browse by ungulates was found.     

The feeding habits of wolves in relation to large prey availability in northern Italy

We investigated wolf feeding habits in relation to the abundance of wild and domestic ungulates to test the hypothesis that large prey are preferred and that their abundance affects the use of other food categories and diet breadth. We determined diet composition by scat analysis from December 1987 to December 1992. The research was carried out in three study areas located in northern Italy and characterised by marked differences in wild and domestic ungulate abundance. In study area A (low wild and domestic ungulate availability) fruits, livestock, other vertebrates and wild ungulates made up the bulk of the diet (71% in volume). In area B (high availability of livestock) wolf diet was mainly based on sheep and wild boars (80% in volume). In study area C (high availability of wild ungulates) wild ungulates were the main food of wolves (90% in volume). Significant differences were found among study areas in the mean percentage volume of all food categories and in particular for wild ungulates, livestock, other vertebrates and fruits (p < 0.0001 in all cases). Diet breadth decreased in areas with high availability of large wild and domestic herbivores. The use of livestock species was lower where there was high abundance, richness and diversity of the wild ungulate guild. Selection for wild ungulate species was partially affected by their abundance: however other factors as prey social behaviour, adaptability to the habitat (for introduced species), and body size could have an important role in species selection by wolves. In particular in area C wild boars were selected for, roe and red deers avoided, and fallow deers and mouflons used as available. Livestock species were used in relation to their abundance and accessibility, in particular sheep were selected for and cattle avoided; but if calves bom in the pastures were considered as the only available cattle, they were selected for and sheep were used as available. Large and in particular wild herbivores were found to be of great importance for the wolf population maintenance in northern Italy, one of the most important recovery areas of Mediterranean wolves.     

Inter-specific synchrony of two contrasting ungulates: wild boar (Sus scrofa) and roe deer (Capreolus capreolus)

Very few studies on ungulates address issues of inter-specific synchrony in population responses to environmental variation such as climate. Depending on whether annual variation in performance of ungulate populations is driven by direct or indirect (trophic) interactions, very different predictions regarding the pattern of inter-specific synchrony can be derived. We compared annual autumn body mass variation in roe deer (Capreolus capreolus) and wild boar (Sus scrofa) from Poland over the period 1982–2002, and related this to variation in winter and summer climate and plant phenological development [the Normalized Difference Vegetation Index (NDVI), derived from satellites]. Roe deer fawns (∼1.3 kg increase from year 1982 to 2002) and yearlings both increased markedly in mass over years. There was also an increase for wild boar mass over years (∼4.2 kg increase for piglets from 1982 to 2002). Despite our failure to link annual body mass to spring or winter conditions or the NDVI, the body mass of roe deer and wild boar fluctuated in synchrony. As this was a field roe deer population, and since wild boar is an omnivore, we suggest this may be linked to annual variation and trends in crop structure (mainly rye). We urge future studies to take advantage of studying multiple species in order to gain further insight into processes of how climate affect ungulate populations. Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

Comparing patterns of human harvest and predation by Eurasian lynx Lynx lynx on European roe deer Capreolus capreolus in a temperate forest

Human harvest is the most important mortality factor for wild ungulates in Europe and can affect several aspects of ungulate biology. There is a growing concern about possible negative side effects of human harvest. To better understand the differences between human and natural mortality, we compared the extent, age and sex structure, nutritional condition, spatial and temporal distribution of human harvest, and natural predation by the Eurasian lynx Lynx lynx on the European roe deer Capreolus capreolus, the most abundant wild ungulate in Europe. Compared to the human harvest, lynx were less likely to kill fawns and yearlings than adults, and among adult deer, lynx were more likely to kill females. The proportion of roe deer with fat-depleted bone marrow was higher among lynx prey than among harvested animals. Average lynx kill rate was estimated to 47.8 roe deer per year, and lynx predation was considerably lower than the human harvest in the same area. While human harvest increased with higher roe deer density, lynx predation was similar across the gradient of roe deer densities. Comparison with other countries indicated that differences between human harvest and natural mortality of ungulates vary considerably in different parts of Europe. Variation in hunting practices and, even more importantly, carnivore predation may have an important role in buffering unwanted side effects of harvest of wild ungulates.     

Short Communication

Abalone farmed in Australia are predominantly fed formulated feeds, while in the wild, their diet consists of a mix of macroalgae. Here we investigated the feeding behaviour of juvenile greenlip abalone, Haliotis laevigata; fed live macroalgae and formulated diets at different rations; and observed their feeding behaviour using a novel ventral video monitoring technique. Four different diets (commercial chip, experimental flake, Ulva sp. and Gracilaria cliftonii) at two feeding rations (excess vs. restricted) were tested. Diet type had no effect on abalone movement, but macroalgal diets resulted in higher feed intakes. Restricting feed rations induced greater movement. Abalone moved little during the light period and moved mostly during darkness, except for animals on the restricted feed ration where feeding commenced during the light period. On farms, this phenomenon may be a useful behavioural indicator for identifying underfed abalone. Despite their lower intake, formulated diets promoted higher energy and nutritional intake, indicating that quantity of feed consumed is not solely indicative of nutritional gain. From a research perspective, the novel ventral monitoring method has created opportunities for further behavioural studies in molluscs.     

Density dependence of daily activity in three ungulate species

Daily activity in herbivores reflects a balance between finding food and safety. The safety‐in‐numbers theory predicts that living in higher population densities increases safety, which should affect this balance. High‐density populations are thus expected to show a more even distribution of activity—that is, spread—and higher activity levels across the day. We tested these predictions for three ungulate species; red deer (Cervus elaphus), roe deer (Capreolus capreolus), and wild boar (Sus scrofa). We used camera traps to measure the level and spread of activity across ten forest sites at the Veluwe, the Netherlands, that widely range in ungulate density. Food availability and hunting levels were included as covariates. Daily activity was more evenly distributed when population density was higher for all three species. Both deer species showed relatively more feeding activity in broad daylight and wild boar during dusk. Activity level increased with population density only for wild boar. Food availability and hunting showed no correlation with activity patterns. These findings indicate that ungulate activity is to some degree density dependent. However, while these patterns might result from larger populations feeling safer as the safety‐in‐numbers theory states, we cannot rule out that they are the outcome of greater intraspecific competition for food, forcing animals to forage during suboptimal times of the day. Overall, this study demonstrates that wild ungulates adjust their activity spread and level based on their population size.     

Selection for nutrients by red deer hinds feeding on a mixed forest edge

We studied the nutritional behaviour of hinds foraging on a mixed-forest edge by direct observation of their choices at each season and by measuring nutrient concentration in the plants. We compared nutrient concentrations in the observed diets with those in the total available vegetation, and with those of 1,000 randomly simulated diets in which we included only those plants that were actually eaten by the animal. Whether the available or the consumed feeds were used as the basis for comparisons had important consequences due to the presence of conifers and ferns, which were high in soluble sugars but were never eaten by the animals (potentially due to their toxicity). The selected diets were lower in sugars than the total available vegetation in summer, but were actually higher in sugars than the random diets generated from consumed forage species only. Hind diets contained more soluble sugars but not more protein than simulated diets in all seasons. Contrary to our prediction, anti-nutritional compounds (ADL and tannins) were avoided only in winter. Compared to simulated diets, hinds consumed more tannins in spring and summer and more ADL in summer and autumn. We suggest that this was a consequence of selection for soluble sugars, because the preferred plant species, which had high soluble sugar concentrations, also contained a large proportion of the anti-nutritional compounds eaten. In winter, the grass-dominated diets contained more fibre (NDF) and less ADL than the simulated diets, indicating that hinds orient their feeding towards digestible fibres. The switch from a browser to a grazer diet was related to a change in the availability of the nutrients, mainly soluble sugars. In our study, grasses contained more soluble sugars and proteins than deciduous browse during winter. This calls into question the dichotomy usually assumed in the literature between grass and browse in terms of nutrient content. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Seed dispersal by ungulates in the point calimere wildlife sanctuary: A scientific and perspective analysis

Exotic woody weed plants are a very serious threat to seed dispersed by ungulate in the tropical forest of Asia. The ungulates in Point Calimere Wildlife Sanctuary (PCWS) are a significant role in native indigenous seed dispersal. The exotic woody weed tree Prosopis juliflora prevalence distributed in the PCWS and they might potentially alter the native medicinal plant species. In the present investigation, we have assessed the seed dispersal by ungulates in PCWS from January to March 2017. Four different ungulate species were selected to understand their seed dispersal rate of different plant species in selected sanctuary. This investigation was planned to confirm the seed dispersal by ungulates of blackbuck, spotted deer, wild boar and feral horse. Among the four different ungulates tested, the maximum numbers of pellets collected from blackbuck and no seed found in their pellets. The low quantities of pellets were collected from wild boar and this study has recorded medium-sized ungulates which dispersed variety of plant. However, the dispersal of the seed of medicinal plants were not considerably high and relatively moderate percentage of seeds dispersal occurred in medium-sized ungulates like wild boar and spotted deer. P. juliflora had 100% seed germination rate were observed from the faecal samples of wild boar and feral horse. The control seed achieved maximum seedling rate than the ungulates seeds.     

Effects of Snow on Sitka Black-Tailed Deer Browse Availability and Nutritional Carrying Capacity in Southeastern Alaska

ABSTRACT Snow affects the nutritional ecology of northern ungulates during winter through burial of important winter forages. We used nonlinear regression analyses to model snow-burial dynamics of blueberry (Vaccinium spp.) browse biomass, a key winter food item of Sitka black-tailed deer (Odocoileus hemionus sitchensis) in southeastern Alaska, USA. During November 2003—March 2004 we collected data from 546 individually marked twigs located on 100 plants of differing sizes and architectures across a range of snow depths. In general, browse biomass became buried and unavailable to deer at snow depths substantially lower than prewinter twig heights. Plant architecture and plant height were related to the probability of a twig being buried. Probability of twig burial was higher on plants with lateral than on those with erect architectures. Twig height also affected the probability of burial by snow but the relationship was complex. For twigs located on erect plants, probability of burial was greatest for twigs near the bottom and top of the plant due to ground-up burial and bending of flexible apex stems, respectively. We used estimated nonlinear equations to model blueberry browse availability in a simulated upland old-growth habitat patch subject to a range of snow depths. We then compared subsequent estimates of deer winter nutritional carrying capacity for this habitat patch to findings derived using an alternative, simple linear (ground-up) model of winter-browse burial by snow. Comparisons indicated that ground-up models of browse burial overestimated browse availability and nutritional carrying capacity for most snow depths. Our findings demonstrate the importance of applying detailed snow-burial models when characterizing nutritional landscape of northern ungulates during winter.     

The influence of snow on the functional response of grazing ungulates

The forage intake rate of grazing ungulates is limited either by the rate at which they encounter food items, or the rate at which food items are handled. Whether an ungulate is encounter‐ or handling‐limited influences spatial and temporal depletion of forage, daily time budgets, and ultimately animal condition. Previously, vegetation abundance has been used as a surrogate for an ungulate's encounter rate with food items and related to observed bite rate to determine whether intake rate is encounter‐ or handling‐limited. In temperate climates snow accumulation during winter limits access to vegetation by forcing animals to wade and paw through snow to consume underlying vegetation, increasing the amount of time required to encounter a food item. As a result, an ungulate may be handling‐limited when foraging in a high biomass system under snow‐free conditions, but becomes encounter‐limited when snow accumulates. We derived a model that provides a frame work for estimating the rate at which a grazing ungulate encounters vegetation by considering foraging velocity, vegetation biomass and the time required to paw away snow when present. We then used data from focal observations of 36 wild elk Cervus canadensis wintering on a montane grassland in the Canadian Rockies of Alberta, Canada, to apply our model and estimate encounter rate over a range of vegetation abundance and snow conditions. Using AIC_c in a model selection approach we found that an asymptotic regression model of observed bite rate as a function of estimated encounter rate provided a better fit than similar models using only vegetation abundance as the explanatory variable. An asymptotic model suggests elk were handling‐limited in the absence of snow, but became encounter‐limited when snow accumulated. Our results demonstrate the importance of considering the influence of factors other than vegetation abundance on the intake rate of grazing ungulates.     

Spatiotemporal Effects of Supplementary Feeding of Wild Boar (Sus scrofa) on Artificial Ground Nest Depredation

Supplementary feeding of ungulates, being widely used in game management, may have unwanted consequences. Its role in agricultural damage is well-studied, but few studies have considered the potential for the practice to attract ground nest predators. Our goal was to identify the factors influencing ground nest predation in the vicinity of year-round supplementary feeding sites for wild boar and to characterise their spatiotemporal scope. We conducted two separate artificial ground nest experiments in five different hunting districts in south-eastern Estonia. The quantity of food provided and distance of a nest from the feeding site were the most important factors determining predation risk. Larger quantities of food resulted in higher predation risk, while predation risk responded in a non-linear fashion to distance from the feeding site. Although predation risk eventually decreases if supplementary feeding is ceased for at least four years, recently abandoned feeding sites still pose a high predation risk.     

Habitat niche relationships within an assemblage of ungulates in Bardia National Park,Nepal

Species co-exist in their individual niches by partitioning resources. The main prey of the big predators – wild boar, chital, and sambar are sympatric in tropical south-Asia. In order to obtain a better understanding of their habitat use and habitat niche relationships, we used an indirect sign (faecal pellets and rooting by wild boar) survey along line transects in Bardia National Park. Ground cover, in particular grass cover for wild boar and chital, and shrub cover for chital and sambar positively affected the occurrence of ungulate signs. We found a niche differentiation between wild boar, which preferred hill sal forest and grassland, and chital and sambar, which avoided hill sal forest and grassland. Habitat niche overlap between chital and sambar in open habitat was clearly evident during the dry season. It is suggested that the fine grained habitat mosaics in space are the underlying causes of the coexistence of the ungulates studied. Therefore, focus on maintaining the habitat mosaic throughout the regions of the species’ distribution may prove successful for effective conservation management.