Foraging dynamics of muskoxen in Peary Land, northern Greenland

Muskoxen Ovibos moschatus in northern Greenland (79-83°N) are at the northern limit of their distribution and exist under seasonal extremes dominated by nearly 10 months of winter, much of which is without sunlight. The period of summer vegetative growth is less than two months. In the Kap København area (82°30'N), diversity of plant species is low (76 species of vascular plants) and forage biomass in major vegetation types in summer varies from over 40 g m^-2 in sedge-dominated fens to ≤5 g m^-2 in polar barrens. Nonetheless, 90-95% of the ice-free area consists of barren ground or sparcely vegetated polar desert. During summer, muskoxen apparently foraged opportunistically to maximize intake, with sedges the major food item in fens while willows were the major dietary component when on willow-dominated slopes. Quality of summer forage was high during its early phenological stages, with 21-28% crude protein and 60-75% in vitro digestibility. Microhistological analysis of winter feces indicated dominance by graminoids. Muskoxen spent > 50% of their daily activity feeding, which fits a cline of increasing feeding time with increasing latitude in summer. Increased feeding times at high latitudes appears to be a function of both reduced forage biomass and need to maximize forage intake during the brief summer period when forage quality is high. Movement rates in summer while foraging were inversely related to available forage biomass. Seasonal activity of muskoxen peaks during the rutting period (July-September) and then declines gradually through early winter to a low in late winter (March-April).     

Nutritional Value of Moss for Arctic Ruminants: a Test with Muskoxen

Abstract: Although moss is commonly found in the feces of arctic herbivores, we do not know the digestible value of this forage for ruminants. We compared grass hay (Bromus sp.) with moss (Hylocomium splendens, Tomenthypnum nitens) from 2 locations in Alaska, USA: Cape Krusenstern National Monument and Fairbanks. We evaluated forages by digestion in ruminally fistulated muskoxen (Ovibos moschatus) by suspending forages in polyester bags before and after the rumen was acclimated with moss for 15 consecutive days. Ruminal degradation was not affected by acclimation to moss. Hay lost dry matter during 48 hours of ruminal incubation (-49%), whereas moss gained dry matter (+44-57%). Incubated moss gained nitrogen (+435-680%), as well as fiber (+18%), and one moss gained ash (+121%). Mass gained by moss in the rumen was probably due to the combined effect of microbial colonization and adsorption of fibrous particles onto the sponge-like matrix. We evaluated postruminal degradation of forages by incubation in acid-pepsin. Ruminally incubated mosses lost little nitrogen in acid-pepsin even though ruminally incubated hay lost 23% nitrogen on acid digestion. Consumption of moss during winter may be a net cost of selecting plants within moss communities when lichens and graminoids are scarce. Moss in feces may, therefore, indicate low availability of favored foods for muskoxen and other arctic ruminants that are confined to small winter ranges. Increasing concentrations of moss in the feces and, thus, the diet of muskoxen may alert wildlife managers to shifts in winter range quality or forage access due to changing snow conditions.     

Foraging strategies and seasonal diet optimization of muskoxen in West Greenland

Various aspects of optimal foraging and seasonal diet composition of bulls (bachelor and dominant), cows, subadults, and yearlings of muskoxen Ovibos moschatus were investigated in West Greenland during the following seasons: calving, post-calving, summer, rut and mid-winter. The following hypotheses were tested: (1) muskoxen maximize daily energy intake during spring and summer, (2) dominant bulls monopolizing cows during the rutting season shift from an energy maximizing to a time minimizing foraging strategy in order to maximize the time available for reproductive activities, and (3) muskoxen employ a time minimizing foraging strategy during winter to conserve energy. As forage quality changed throughout the short Arctic growing season, muskoxen responded by changing the proportions of daily time spent feeding on graminoids (Cyperaceae, Poaceae) and dicots (Salix, Betula), respectively. This seasonal variation in the relative proportion of daily feeding time spent ingesting graminoids followed approximately the energy maximization prediction over the periods calving to rut. Neither time minimizing nor random foraging could explain the observed diets in this period, thus confirming hypothesis 1. Dominant bulls did not shift to the time minimizing strategy as predicted by hypothesis 2. However, during the pre-rutting and rutting seasons bulls deviated from the other sex/age classes by failing to obtain the daily maximum energy predicted by the model, as a result of a higher proportion of time allocated to agonistic and sexual behaviour. During winter, none of the sex/age classes employed a time minimizing strategy, so rejecting hypothesis 3. Instead, muskoxen were found to maximize Na intake, indicating that Na is of major importance for winter survival. The results emerging from a linear programming model with constraint settings varying over seasons confirm that the constraint parameters applied are indeed important limiting factors for muskoxen in natural populations.     

Moose summer and winter diets along a large scale gradient of forage availability in southern Norway

Studies on dietary functional responses in large herbivores are traditionally conducted by following individual animals. The method is very time-consuming, and hence, typically provides only a narrow array of forage species compositions. Here we use a range level approach to look at moose (Alces alces) selectivity for and utilization of forage species in relation to availability in both summer and winter. We compare 12 Norwegian ranges representing a large scale gradient in plant communities. The most important forage species in the diet were birches (Betula spp., comprising 43% of all trees browsed in summer and 27% in winter), rowan (Sorbus aucuparia, 25% of trees browsed in summer, 37% in winter), and bilberry (Vaccinum myrtillus, 42% of herbaceous epidermal fragments in summer feces). Selectivity for birches was positively related to its availability and negatively related to availability of rowan, Salix spp., and aspen (Populus tremula) together (all more selected for than birches). Multiple regression models including availability of several forage species were thus superior to single-species models in explaining the diet content of main forage plants. Selectivity for birches was also stronger in summer than in winter, while the opposite pattern was found for rowan. The finding is relevant for our evaluation of the quality of summer and winter ranges, and hence, their relative influence on population productivity. Our study underlines the need to incorporate species composition of available forage when quantifying dietary functional responses in selective herbivores such as moose. Furthermore, care should be taken when extrapolating data on moose diet across ranges or seasons.     

Beetle community responses to grey willow (Salix cinerea) invasion within three New Zealand wetlands

Abstract

We investigated the effects of invasion by introduced grey willow (Salix cinerea) on beetle communities within four wetland vegetation types: native vegetation, native vegetation following grey willow removal, native vegetation undergoing grey willow invasion and dense grey willow-dominated vegetation. In total, 1505 beetles from 90 species were collected using modified Malaise traps. Native wetland vegetation had significantly lower beetle species richness than willow-dominated vegetation and was dominated by herbivores, whereas detritivores characterised willow-dominated vegetation. Beetle abundance was highest in the willow-dominated vegetation and mostly comprised detritivores. In contrast, beetle abundance was lowest in native wetland vegetation, but had even proportions of herbivores and detritivores. Native wetland vegetation had a high proportion of native beetles present. As grey willows invaded, introduced beetles became more common. The beetle community composition differed significantly between grey willow-dominated vegetation and native wetland vegetation. These compositional differences were mainly due to the increasing complexity of vegetation structure following grey willow invasion. The beetle communities within restored native wetland vegetation were most similar to those within the native wetland vegetation. From a conservation perspective, these results are encouraging and suggest that, although grey willows dramatically alter the composition of beetle communities present, these communities can be restored to a beetle fauna that is similar to those found within native wetland through the removal of the willows.     

Herbivores Influence the Growth,Reproduction, and Morphology of a Widespread Arctic Willow

Shrubs have expanded in Arctic ecosystems over the past century, resulting in significant changes to albedo, ecosystem function, and plant community composition. Willow and rock ptarmigan (Lagopus lagopus, L. muta) and moose (Alces alces) extensively browse Arctic shrubs, and may influence their architecture, growth, and reproduction. Furthermore, these herbivores may alter forage plants in such a way as to increase the quantity and accessibility of their own food source. We estimated the effect of winter browsing by ptarmigan and moose on an abundant, early-successional willow (Salix alaxensis) in northern Alaska by comparing browsed to unbrowsed branches. Ptarmigan browsed 82–89% of willows and removed 30–39% of buds, depending on study area and year. Moose browsed 17–44% of willows and browsed 39–55% of shoots. Browsing inhibited apical dominance and activated axillary and adventitious buds to produce new vegetative shoots. Ptarmigan- and moose-browsed willow branches produced twice the volume of shoot growth but significantly fewer catkins the following summer compared with unbrowsed willow branches. Shoots on browsed willows were larger and produced 40–60% more buds compared to unbrowsed shoots. This process of shoot production at basal parts of the branch is the mechanism by which willows develop a highly complex “broomed” architecture after several years of browsing. Broomed willows were shorter and more likely to be re-browsed by ptarmigan, but not moose. Ptarmigan likely benefit from the greater quantity and accessibility of buds on previously browsed willows and may increase the carrying capacity of their own habitat. Despite the observed tolerance of willows to browsing, their vertical growth and reproduction were strongly inhibited by moose and ptarmigan. Browsing by these herbivores therefore needs to be considered in future models of shrub expansion in the Arctic.     

Food habits of sika deer and nutritional value of sika deer diets in eastern Hokkaido, Japan

Rumen content analysis and field observations were used to investigate the food habits and diet quality of sika deer (Cervus nippon yesoensis Heude) from 1991 to 1993 in eastern Hokkaido, Japan. Diets varied seasonally, with deer consuming graminoids and browse in winter, forbs and agricultural crops in spring and summer and all of these plant foods in autumn. Eighty-four plant species with sika deer bite marks were identified and their use also varied seasonally. The diversity of food resources available provided both critical protein and digestible energy, allowing for physiological maintenance and seasonal growth. With these high-quality diets, deer maintained good body condition in eastern Hokkaido, where the population density was relatively low.     

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.     

Characteristics of foraging sites and protein status in wintering muskoxen: insights from isotopes of nitrogen

Identifying links between nutritional condition of individuals and population trajectories greatly enhances our understanding of the ecology, conservation, and management of wildlife. For northern ungulates, the potential impacts of a changing climate to populations are predicted to be nutritionally mediated through an increase in the severity and variance in winter conditions. Foraging conditions and the availability of body protein as a store for reproduction in late winter may constrain productivity in northern ungulates, yet the link between characteristics of wintering habitats and protein status has not been established for a wild ungulate. We used a non‐invasive proxy of protein status derived from isotopes of N in excreta to evaluate the influence of winter habitats on the protein status of muskoxen in three populations in Alaska (2005–2008). Multiple regression and an information‐theoretic approach were used to compare models that evaluated the influence of population, year, and characteristics of foraging sites (components of diet and physiography) on protein status for groups of muskoxen. The observed variance in protein status among groups of muskoxen across populations and years was partially explained (45%) by local foraging conditions that affected forage availability. Protein status improved for groups of muskoxen as the amount of graminoids in the diet increased (?0.430 ± 0.31, β± 95% CI) and elevation of foraging sites decreased (0.824 ± 0.67). Resources available for reproduction in muskoxen are highly dependent upon demographic, environmental, and physiographic constraints that affect forage availability in winter. Due to their very sedentary nature in winter, muskoxen are highly susceptible to localized foraging conditions; therefore, the spatial variance in resource availability may exert a strong effect on productivity. Consequently, there is a clear need to account for climate–topography effects in winter at multiple scales when predicting the potential impacts of climatic shifts on population trajectories of muskoxen.     

Why should a grazer browse? Livestock impact on winter resource use by bharal Pseudois nayaur

Many mammalian herbivores show a temporal diet variation between graminoid-dominated and browse-dominated diets. We determined the causes of such a diet shift and its implications for conservation of a medium-sized ungulate—the bharal Pseudois nayaur. Past studies show that the bharal diet is dominated by graminoids (>80%) during summer, but the contribution of graminoids declines to about 50% in winter. We tested the predictions generated by two alternative hypotheses explaining the decline: low graminoid availability during winter causes bharal to include browse in their diet; bharal include browse, with relatively higher nutritional quality, in their diet to compensate for the poor quality of graminoids during winter. We measured winter graminoid availability in areas with no livestock grazing, areas with relatively moderate livestock grazing, and those with intense livestock grazing pressures. The chemical composition of plants contributing to the bharal diet was analysed. The bharal diet was quantified through signs of feeding on vegetation at feeding locations. Population structures of bharal populations were recorded using a total count method. Graminoid availability was highest in areas without livestock grazing, followed by areas with moderate and intense livestock grazing. The bharal diet was dominated by graminoids (73%) in areas with highest graminoid availability. Graminoid contribution to the bharal diet declined monotonically (50, 36%) with a decline in graminoid availability. Bharal young to female ratio was 3 times higher in areas with high graminoid availability than areas with low graminoid availability. The composition of the bharal winter diet was governed predominantly by the availability of graminoids in the rangelands. Our results suggest that bharal include more browse in their diet during winter due to competition from livestock for graminoids. Since livestock grazing reduces graminoid availability, creation of livestock-free areas is necessary for the conservation of grazing species such as the bharal and its predators including the endangered snow leopard in the Trans-Himalaya.     

Variation at high latitudes: the geography of body size and cranial morphology of the muskox, Ovibos moschatus

Abstract Aim Bergmann's rule, one of the most studied and controversial ecogeographical generalizations, has rarely been tested with observations from high latitudes. We tested the rule using cranial measurements of the muskox [Ovibos moschatus (Zimmerman)], a homeotherm with an extremely northern distribution. We also used these data to describe geographical patterns in the species' dental architecture, an extension of the framework developed from interspecific comparisons. Location Specimens were compiled from arctic Canada, Alaska and Greenland, a latitudinal range of 60° N?83° N. Methods Body size was estimated from principal components analysis (PCA) of five cranial characters from 128 specimens. Mean scores on the first principal component from each locality were regressed against latitude and mean temperature to identify geographical variation in body size; scores on the second principal component were regressed against latitude to assess patterns in dental architecture. Regression analyses of the individual characters were performed as a complement to PCA. Results No latitudinal or climatic trend in body size was observed in either sex. On the other hand, for males, significant latitudinal variation was found for the second PCA axis (r = ?0.434), and the feature which loaded most heavily on it, maxillary tooth row length (r = 0.429). For females, this dental structure also tended to increase with latitude (r = 0.423), but the trend was only marginally significant (P=0.12), perhaps owing to a smaller sample size. Main conclusions The geographically invariant body size of muskoxen failed to support current hypotheses of size variation. Behavioural and physiological adaptations may exempt the muskox from selective pressures underlying these hypotheses. We interpret latitudinal variation in dental architecture as a reflection of a cline in diet, dominated by graminoids at the expense of willows at higher latitudes. This intraspecific geographical trend is a recapitulation of the interspecific framework for large mammalian herbivores.     

A Review of Environmental Factors Affecting Elk Winter Diets

Abstract: Decades of research have produced substantial data on elk (Cervus elaphus) diets in winter, when foraging conditions are most likely to affect population dynamics. Using data from 72 studies conducted in western North America between 1938 and 2002, we collated data on elk diets and environmental variables. We used these data to quantify diet selection by elk and to test whether variation in elk diets is associated with habitat type, winter severity, period of winter, human hunting, and study method. Graminoids (grasses and grass-like plants such as sedges) dominated elk diets and consistently occurred at a higher proportion in the diet than in elk foraging habitats, indicating preference. Forbs commonly made up ≤5% of the diet, with no evidence for preference; we conclude that forb use is largely incidental to grazing for graminoids. Browse was consumed in proportion to its availability, implying that the amount of browse in the diet was primarily determined by habitat use rather than selection. Comparing the diets of elk and sympatric ruminants, elk consistently selected graminoids more strongly than sympatric ruminants with the exception of bison (Bison bison), suggesting that elk are not environmentally forced to adopt the graminoid-biased diet that they normally select. The proportion of open meadows and grasslands on winter ranges was strongly and positively associated with graminoid consumption by elk. The proportion of graminoids in the diet was significantly lower in elk experiencing severe winter conditions or predation risk from human hunting. The period of winter (early, middle, and late) had only small effects on elk diets, as did the method by which the diet was determined. Overall, variation in elk diets is well-explained by a consistent tendency to select graminoids if available, modified by winter habitat type, predation risk, and winter severity, which can constrain habitat selection and access to grazing opportunities. To fully understand variation in foraging behavior, biologists should recognize these broad patterns when interpreting resource selection data. Managers should recognize that inconspicuous behavioral responses to environmental stimuli can alter the diet in ways that probably carry nutritional consequences.     

Critical summer foraging tradeoffs in a subarctic ungulate

Summer diets are crucial for large herbivores in the subarctic and are affected by weather, harassment from insects and a variety of environmental changes linked to climate. Yet, understanding foraging behavior and diet of large herbivores is challenging in the subarctic because of their remote ranges. We used GPS video‐camera collars to observe behaviors and summer diets of the migratory Fortymile Caribou Herd (Rangifer tarandus granti) across Alaska, USA and the Yukon, Canada. First, we characterized caribou behavior. Second, we tested if videos could be used to quantify changes in the probability of eating events. Third, we estimated summer diets at the finest taxonomic resolution possible through videos. Finally, we compared summer diet estimates from video collars to microhistological analysis of fecal pellets. We classified 18,134 videos from 30 female caribou over two summers (2018 and 2019). Caribou behaviors included eating (mean = 43.5%), ruminating (25.6%), travelling (14.0%), stationary awake (11.3%) and napping (5.1%). Eating was restricted by insect harassment. We classified forage(s) consumed in 5,549 videos where diet composition (monthly) highlighted a strong tradeoff between lichens and shrubs; shrubs dominated diets in June and July when lichen use declined. We identified 63 species, 70 genus and 33 family groups of summer forages from videos. After adjusting for digestibility, monthly estimates of diet composition were strongly correlated at the scale of the forage functional type (i.e., forage groups composed of forbs, graminoids, mosses, shrubs and lichens; r = 0.79, p < .01). Using video collars, we identified (1) a pronounced tradeoff in summer foraging between lichens and shrubs and (2) the costs of insect harassment on eating. Understanding caribou foraging ecology is needed to plan for their long‐term conservation across the circumpolar north, and video collars can provide a powerful approach across remote regions.     

Ecology of muskoxen in Jameson Land, northeast Greenland

Muskoxen Ovibos moschatus in Jameson Land exist at a density of somewhat more than 1 km⁻² of useable habitat and select moist meadows and snow bed vegetation for summer grazing and wind-exposed, dry dwarf shrub heath vegetation in winter. Graminoids dominate the winter diet and willows are the main component of the summer diet. Quality of the winter diet, as measured by the protein to fiber ratio is about one fourth that of the summer diet. During summer muskoxen supplement dietary sodium by using mineral licks. Muskoxen, especially females, retain considerable unused fat reserves through the winter and these are drawn upon during the post-calving period of lactation. Alternate year breeding is a common occurrence. Calves are frequently not weaned before the end of their first winter. Mean calf mortality is relatively low in the absence of significant predation and annual removal by hunting Inuits approaches the annual increment.     

Population dynamics of the collared lemming and the tundra vole at Pearce Point,Northwest Territories,Canada

From 1987 to 1989 we monitored population changes during summer of the collared lemming (Dicrostonyx groenlandicus) and the tundra vole (Microtus oeconomus) at Pearce Point, Northwest Territories, Canada (69° 48 N, 122° 40 W). Populations on four study areas did not cycle but remained at low density (<3/ha) each year and continued at low numbers for the following 3 years (Reid et al. 1995). Lemming numbers often declined throghout the summer in spite of continous reproduction, and population recovery occurred overwinter. Heavy predation losses of radio-collared lemmings occurred each summer and this lemming population may be trapped in a predator-pit. Collared lemmings breed in winter and only because of winter population growth do these populations persist. Tundra vole numbers increased rapidly in most summers but usually declined overwinter. Tundra voles do not seem able to sustain winter reproduction in this extreme environment and this prevents them from reaching high density because of the short summer. Population growth in both these rodents could be prevented by poor food or by predation losses, and landscape patchiness may also help to prevent population growth. For lemmings we do not think that a shortage of shelter or intrinsic limitations could be restricting population increase at Pearce Point. This is the first detailed study of a non-cyclic collared lemming population.     

Does Supplemental Feed Increase Selective Foraging in a Browsing Ungulate?

ABSTRACT Provision of supplemental feed to large herbivores is a common management practice that may motivate selective foraging, thereby influencing plant community composition. Our objective was to assess the effect of a high-quality supplement on diet composition and nutritional quality for white-tailed deer (Odocoileus virginianus). We permanently released hand-reared deer into 4 81-ha enclosures; in 2 enclosures we provided a pelleted supplement. We conducted bite-count studies seasonally to assess diet composition and quality. Supplemented deer reduced mast (fruits and pods of woody plants and cacti) in their diets (P < 0.019) during spring and autumn compared to unsupplemented deer. Diets of deer in supplemented enclosures had 2 times greater proportion of browse during spring (P = 0.065) and 5 times greater proportion of forbs during autumn (P = 0.007). Quality of the forage portion of the diet did not vary by treatment during winter or summer. Metabolizable energy concentration was 13% greater (P = 0.054) in spring and digestible protein content was 3 times greater (P = 0.006) during autumn in diets of supplemented compared to unsupplemented deer. Our results support the selective foraging hypothesis during autumn but not during winter, spring, or summer. Furthermore, white-tailed deer did not reduce the proportion of their diet composed of browse, but did reduce consumption of mast. Supplemented deer continued to eat poor-quality, chemically defended forage, perhaps to alleviate ruminal acidosis induced by the supplement or because nutrients in the supplement increased the deer's ability to detoxify chemically defended browses. A decline in mast consumption by supplemented deer could influence plant communities, depending on the role of deer in seed dispersal and seed predation. Impacts of supplemental feed on selective foraging of white-tailed deer in shrub-dominated rangelands are more complex than suggested by previous research. Long-term studies of vegetation communities are needed before wildlife managers will be able to fully incorporate effects of supplemental feed into management decisions.     

Percentage of Fecal Moss in Arctic Ungulates as an Indicator of Wintering Area Quality

Abstract: I investigated local and regional differences in percent moss in the feces of muskoxen (Ovibos moschatus) and reindeer or caribou (Rangifer tarandus) in northwestern Alaska, USA, and related fecal moss to forage availability, snow conditions, animal density, and terrain ruggedness on wintering areas. Reindeer are a partially domesticated form of Rangifer tarandus originating from Europe and Asia and differ physiologically and ecologically from caribou. Percent moss in feces of muskoxen differed locally among individual wintering areas. Because of the large local variation in moss content of muskox feces, regional differences between the 2 study areas were difficult to resolve. Percent of moss in the feces of reindeer–caribou did not differ between wintering areas within the same study area but did differ between study areas. On muskox wintering areas, fecal moss correlated negatively with graminoid cover and snow hardness and positively with moss cover and muskox density, but fecal moss did not correlate with snow depth or terrain ruggedness. On reindeer–caribou wintering areas, fecal moss correlated positively with moss availability but not with lichen cover or snow depth or hardness. Because muskox groups in Alaska are isolated from each other in winter, even groups wintering on neighboring hills may face different foraging availability and might, therefore, exhibit differences in growth or productivity. Reindeer–caribou are more mobile than muskoxen in winter, and fecal samples may not be representative of vegetative and snow conditions at the wintering area where I collected them. I conclude that managers can use fecal moss as an indicator of overgrazed ranges, severe snow conditions, or crowded conditions on muskox wintering areas, but that the association between fecal moss and range conditions has to be drawn with caution for the more mobile reindeer–caribou.     

Diet overlap between small ruminants and the European hare in a Mediterranean shrubland

Seasonal diets of goats, sheep and European hares (Lepus europaeus) were examined using microhistological analysis of feces collected when these herbivores grazed together in a typical Mediterranean shrubland. Approximately half of the total diet content of goats was shrubs (mainly kermes oak, Quercus coccifera), while that of hares was grasses (mostly brush grass, Chrysopogon gryllus). Sheep had a more balanced diet consisting mainly of grasses, forbs, and shrubs. Dietary overlap between goats and sheep was high throughout the year. In contrast, there was very low dietary overlap between small ruminants and hares. Dietary diversity was high in spring and low in winter across all species, with sheep in general displaying higher dietary diversity across all seasons than goats and hares. Goats had intermediate and hares had low dietary diversity across all seasons. Communal grazing by small ruminants and hares ensures that there is a more uniform use of the available forage resources than if a single herbivore is left to graze an area.     

Seasonal food habits of a sika deer population in the warm temperate forest of the westernmost part of Honshu, Japan

The rumen contents of 63 sika deer (Cervus nippon Temminck) shot in the westernmost part of Honshu were analyzed by the point-frame method. The rumen contents were dominated by forbs and the majority of these were evergreen monocotyledonous plants, probably Ophiopogon spp. Evergreen browse leaves (woody leaves), such as Eurya japonica Thunb., Ilex crenata Thunb. and Trachelospermum asiaticum Nakai, were also important, constituting 20–40% of the rumen contents. Other categories accounted for only small proportions, although in summer, graminoids increased but twigs and bark decreased in importance and in fall and winter, fruits and seeds increased in importance. The diet of the deer was characterized by high-quality foods even in winter and by a stability or less-pronounced seasonality throughout the year than found in the northern deer. These results were compared with those from other populations, including a northern sika deer population in Japan.     

Summer feeding strategy of Spanish ibexCapra pyrenaica and domestic sheepOvis aries in south-eastern Spain

The feeding strategies of the Spanish ibexCapra pyrenaica Schinz, 1838 and sheepOvis aries Linnaeus, 1758 during summer in the alpine area of Sierra Nevada in south-eastern Spain were compared. The degree of vegetation use by the two ungulates, food availability, diet composition, species selection indices and the effect of plant availability and chemical composition on diet selection were analysed. The available biomass of herbaceous and woody resources was 87 and 13%, respectively. The degree of use was 15% for herbaceous vegetation and 10.3% for woody vegetation. The most used species were those with moderate amounts in diet composition of both ruminants. The main plant groups consumed by Spanish ibex were graminoids (57.5%), shrubs (20.5%) and chamaephytes (14.3%). Sheep consumed graminoids (41.6%), forbs (32.3%) and chamaephytes (20.9%). There was a significant correlation between the two diets, with an overlap of 56.3%. The highest indices of selection were found in items with moderate-low availability and which comprised moderate levels in the diet composition of the two species. Spanish ibex mainly selected diet according to availability, while sheep selection depended on protein content. The ibex consumed cell wall-rich resources such as grasses and shrubs, while the sheep consumed higher quality feed and includes more forbs in its diet.