A natural feeding experiment on a declining snowshoe hare population

Summary A snowshoe hare (Lepus americanus) population on a 9-ha area was supplied with extra natural food by chopping down large white spruce (Picea glauca) and aspen (Populus tremuloides) trees throughout 3 winters from 1981 to 1984. Hares fed vigorously on the downed trees, but the phase of decline of the ten-year cycle occurred equally on control and experimental grids from 1981–1983, and we could detect no improvement in survival or reproduction on the food area. Growth rates were improved on the food grid during the first winter of the decline (1981–82). We concluded that food shortage is not necessary for the cyclic decline of snowshoe hares in the southern Yukon.     

Defense of winter-dormant Alaska paper birch against snowshoe hares

Summary Mature growth-phase internodes of Alaska paper birch (Betula resinifera) are preferred by the snowshoe hare (Lepus americanus) over juvenile growth-phase internodes due to the low food value of the latter. While the mature over juvenile preferencec cannot be explained by the levels of inorganic nutrients or gross chemical fractions (resins or phenols), it can be explained by the striking differences in secondary metabolites of the two growth phases. The principle compound which renders the juvenile phase internodes unpalatable is papyriferic acid, a triterpene which is a demonstrated feeding deterrent to snowshoe hares and which is present in juvenile internodes at concentrations 25 times greater than those in mature internodes.     

Powassan virus infection in snowshoe hares (Lepus americanus)

Sera from snowshoe hares (Lepus americanus) trapped near Rochester, Alberta, Canada were tested for Powassan virus antibody by the constant virus/serum dilution neutralization test. Of 1264 serum samples tested, 137 had an antibody titer of at least 1:4 for Powassan virus. Ten hares were inoculated with Powassan virus in the laboratory. Viremia lasted 4-5 days and ceased with the appearance of Powassan antibody in the serum. Neutralizing antibody reached a peak titer of 1:119 on day 15 post-inoculation and was still detectable 13 months post-inoculation.     

Summer diet selection of snowshoe hares: a test of nutritional hypotheses

We examined summer diet selection by snowshoe hares in the southwestern Yukon, Canada, and attempted to explain how plant qualities such as nutritional content (protein, energy, fiber, water) and defensive compounds affect hare food choice. Male hares were placed in enclosures in natural vegetation areas dominated by shrubs and in both open and closed spruce forests. Each site contained a variety of herbaceous plant species. After 24 h, hares were sacrificed, their stomachs collected, and the contents removed and analyzed. Of the 30 plant species recorded in the enclosures only 10 were identified in the stomach contents of the hares. The summer diet is composed predominantly of five plant species; Lupinus arcticus, Salix spp. Shepherdia canadensis, Betula glandulosa and Festuca altaica. The selection of these species changes between sampling times. Protein, energy and water contents were highest in leaf tissues of most species early in the season, and most species had an increase in fiber through the summer. Protein content consistently explains the largest amount of variation in diet selection, although protein selection is modified by extremely high concentrations of defence compounds. It is likely that hares are not protein limited and are selecting for other plant characteristics correlated with protein content such as energy content. It is significant that hares continue to ingest heavily defended species (e.g. Shepherdia canadensis, B. glandulosa and L. arcticus) when many less defended plants are available to be eaten (e.g. F. altaica, Epilobium latifolium and Anemone parviflora). Our results support the idea that hares are selecting dietary items on the basis of energy content. No single plant species can satisfy all of the hares’ nutritional requirements and the selection patterns may reflect the need to balance many conflicting plant qualities.     

An antifeedant in balsam poplar inhibits browsing by snowshoe hares

Summary The plant defense guild hypothesis for the evolution of plant secondary chemicals predicts that plant species defend themselves against generalist herbivores such as the snowshoe hare (Lepus americanus) in the Canadian boreal forest by evolving unique antifeedant chemicals. Plant species may coevolve in an ecosystem by presenting an array of chemicals to herbivores. We report further evidence for this idea from the presence of 2,4,6-trihydroxydihydrochalcone in the CH₂Cl₂ extracts of Populus balsamifera juvenile twigs. These extracts, added to rabbit chow, were offered to hares in choice tests. The bioassay established that the chemical acted as an antifeedant for hares.     

Owl predation on snowshoe hares: consequences of antipredator behaviour

We show evidence of differential predation on snowshoe hares (Lepus americanus) by great horned owls (Bubo virginianus) and ask whether predation mortality is related to antipredator behaviour in prey. We predicted higher predation on (1) young and inexperienced hares, (2) hares in open habitats lacking cover to protect from owl predation, and (3) hares in above average condition assuming that rich food patches are under highest risk of predation. Information on killed hares was obtained at nest sites of owls and by monitoring hares using radio-telemetry. The availability of age classes within the hare population was established from live-trapping and field data on reproduction and survival. Great horned owls preferred juvenile over adult hares. Juveniles were more vulnerable to owl predation before rather than after dispersal, suggesting that displacement or increased mobility were not causes for this increased mortality. Owls killed ratio-collared hares more often in open than in closed forest types, and they avoided or had less hunting success in habitats with dense shrub cover. Also, owls took hares in above average condition, although it is unclear whether samples from early spring are representative for other seasons. In conclusion, these results are consistent with the hypothesis that variation in antipredator behaviours of snowshoe hares leads to differential predation by great horned owls.     

Selected microbial agents in snowshoe hares and other vertebrates of Alberta

Serologic surveillance of populations of snowshoe hares and other vertebrate species of north-central Alberta from 1961 to 1969, revealed activity of one bacterial and eight viral agents. The most prevalent agents infecting the snowshoe hare were California encephalitis and Silverwater viruses, while in other vertebrates California encephalitis and Western equine encephalomyelitis viruses were the most common. The role of the snowshoe hare in the natural history of the agents is considered as is the effect of the agent on the hare ten-year cycle of abundance.     

Habitat-dependent foraging in a classic predator–prey system: a fable from snowshoe hares

Current research contrasting prey habitat use has documented, with virtual unanimity, habitat differences in predation risk. Relatively few studies have considered, either in theory or in practice, simultaneous patterns in prey density. Linear predator–prey models predict that prey habitat preferences should switch toward the safer habitat with increasing prey and predator densities. The density‐dependent preference can be revealed by regression of prey density in safe habitat versus that in the riskier one (the isodar). But at this scale, the predation risk can be revealed only with simultaneous estimates of the number of predators, or with their experimental removal. Theories of optimal foraging demonstrate that we can measure predation risk by giving‐up densities of resource in foraging patches. The foraging theory cannot yet predict the expected pattern as predator and prey populations covary. Both problems are solved by measuring isodars and giving‐up densities in the same predator–prey system. I applied the two approaches to the classic predator–prey dynamics of snowshoe hares in northwestern Ontario, Canada. Hares occupied regenerating cutovers and adjacent mature‐forest habitat equally, and in a manner consistent with density‐dependent habitat selection. Independent measures of predation risk based on experimental, as well as natural, giving‐up densities agreed generally with the equal preference between habitats revealed by the isodar. There was no apparent difference in predation risk between habitats despite obvious differences in physical structure. Complementary studies contrasting a pair of habitats with more extreme differences confirmed that hares do alter their giving‐up densities when one habitat is clearly superior to another. The results are thereby consistent with theories of adaptive behaviour. But the results also demonstrate, when evaluating differences in habitat, that it is crucial to let the organisms we study define their own habitat preference.     

Actinobacillosis in free-ranging snowshoe hares (Lepus americanus) from Alaska

Actinobacillus capsulatus was isolated from lung, liver, and/or spleen tissue of three snowshoe hares (Lepus americanus) in Alaska. This is the first report of the isolation of this bacterium from free-ranging hares. Actinobacillus capsulatus may have a negative impact on the population density of hares.     

Of lemmings and snowshoe hares: the ecology of northern Canada

Two population oscillations dominate terrestrial community dynamics in northern Canada. In the boreal forest, the snowshoe hare (Lepus americanus) fluctuates in cycles with an 8-10 year periodicity and in tundra regions lemmings typically fluctuate in cycles with a 3-4 year periodicity. I review 60 years of research that has uncovered many of the causes of these population cycles, outline areas of controversy that remain and suggest key questions to address. Lemmings are keystone herbivores in tundra ecosystems because they are a key food resource for many avian and mammalian predators and are a major consumer of plant production. There remains much controversy over the role of predation, food shortage and social interactions in causing lemming cycles. Predation is well documented as a significant mortality factor limiting numbers. Food shortage is less likely to be a major limiting factor on population growth in lemmings. Social interactions might play a critical role in reducing the rate of population growth as lemming density rises. Snowshoe hares across the boreal forest are a key food for many predators and their cycles have been the subject of large-scale field experiments that have pinpointed predation as the key limiting factor causing these fluctuations. Predators kill hares directly and indirectly stress them by unsuccessful pursuits. Stress reduces the reproductive rate of female hares and is transmitted to their offspring who also suffer reduced reproductive rates. The maternal effects produced by predation risk induce a time lag in the response of hare reproductive rate to density, aiding the cyclic dynamics.     

Precommercial forest thinning alters abundance but not survival of snowshoe hares

Anthropogenic landscape modification is a major threat to global biodiversity and ecosystem integrity. Thus, land use practices that reconcile human needs with protection of species and ecological processes are of interest. Precommercial forest thinning (PCT) is a land use practice that is believed to be less ecologically disruptive than other silvicultural approaches (e.g., clear-cutting). The impacts of PCT on wildlife populations are not well understood, however, and the effects of this practice on individual animals have not been rigorously examined. Accordingly, we investigated short-term population and individual responses of snowshoe hares (Lepus americanus) to PCT in western Oregon during 2001–2002. We live-trapped hares (n = 143) seasonally and equipped them with mortality-sensitive radio-collars on 3 manipulated sites that were precommercially thinned and on 3 undisturbed controls. We also quantified the activity (movement rates) of all collared hares using an automated data-logger. As a result, we were able to document the effects of PCT on hare abundance, individual survival, and daily movement. Thinning significantly reduced hare abundance relative to that on control sites, but had no effect on individual mortality or activity. We infer, therefore, that a portion of the hares on thinned sites dispersed to adjacent habitat, where they survived as well as conspecifics on both control and manipulated sites but were unavailable for subsequent capture. We conclude that PCT affects hare density in the short-term (i.e., at least in the first few years post-treatment) largely via altered behavior. Thus, if the immediate impacts of this practice on hare populations are to be minimized, thinned stands should be imbedded in a habitat matrix that facilitates the occupancy of individuals dispersing from disturbed patches. © 2012 The Wildlife Society.     

Prevalence of helminths in a cyclic snowshoe hare population

Five species of helminths were monitored in a population of snowshoe hares (Lepus americanus) near Rochester, Alberta, during 1961-1977. Prevalence of both Obeliscoides cuniculi and Protostrongylus boughtoni among young hares averaged about 50% by age 2 mo, then tended to level off. Prevalence of Taenia pisiformis (cysticerci) and Dirofilaria scapiceps rose more slowly, but continued to increase steadily beyond their mean levels of 8% and 1% at age 2 mo. There were well defined seasonal (within-year) cycles in prevalence of O. cuniculi and P. boughtoni that were generated evidently to a major degree by arrested development of larvae in fall and renewed development in late winter. It was hypothesized that renewed larval development was triggered (in February) in O. cuniculi by the seasonal rise of circulating pituitary gonadotrophins, and (in April) in P. boughtoni by the seasonal rise of gonadal androgens and estrogens. Indices to gonadal hormone levels in hares indicated that these increased most rapidly among males, and may have accounted for the higher prevalences of P. boughtoni in males during April-May. Neither T. pisiformis nor D. scapiceps exhibited conspicuous seasonal changes in prevalence. Maximum prevalence of T. pisiformis was attained at about 1 yr of age, whereas D. scapiceps increased among adult snowshoes through age 2 yr before stabilizing. Long-term (between-year) changes in prevalence of O. cuniculi, T. pisiformis, and D. scapiceps were correlated significantly with the cyclic hare population which declined from a peak in fall 1961 to a low in 1965-1966, rose to another peak by fall 1970, and declined again to a low in 1975. There was no detectable time lage between this "10-yr" cycle in hare density and the cycles of parasite prevalence among juveniles (less than 1 yr of age). Among adult hares, the cycle of O. cuniculi prevalence was likewise synchronous with that of the hare population, but the cycles of D. scapiceps and T. pisiformis lagged by approximately 1 and 2 yr, respectively. This lag in T. pisiformis prevalence was largely inexplicable to us. Our data on P. boughtoni were not suitable for analyses of between-year trends; nor were those for the fifth helminth, Taenia serialis (coenuri), because mean prevalence was less than 1% among both juveniles and adults. An apparent decline in T. serialis after the early 1950's, and its continued scarcity thereafter, paralleled a major change in numbers of one important definitive host--the red fox (Vulpes vulpes).(ABSTRACT TRUNCATED AT 400 WORDS)     

Influence of forest structure on the abundance of snowshoe hares in western Wyoming

Snowshoe hares (Lepus americanus) are a primary prey species for Canada lynx (Lynx canadensis) in western North America. Lynx management plans require knowledge of potential prey distribution and abundance in the western United States. Whether even-aged regenerating forests or multi-storied forests contain more snowshoe hares is currently unknown. During 2006–2008, we estimated snowshoe hare density in 3 classes of 30–70-year-old lodgepole pine (Pinus contorta) and 4 classes of late seral multi-storied forest with a spruce (Picea engelmannii)-fir (Abies lasiocarpa) component in the Bridger-Teton National Forest, Wyoming. We recorded physiographic variables and forest structure characteristics to understand how these factors influence abundance of snowshoe hares. In many instances, snowshoe hares were more abundant in late seral multi-storied forests than regenerating even-aged forests. Forest attributes predicting hare abundance were often more prevalent in multi-storied forests. Late seral multi-storied forests with a spruce–fir component and dense horizontal cover, as well as 30–70-year-old lodgepole pine with high stem density, were disproportionately influential in explaining snowshoe hare densities in western Wyoming. In order to promote improved habitat conditions for snowshoe hares in this region, management agencies should consider shifting their focus towards maintaining, enhancing, and promoting multi-storied forests with dense horizontal cover, as well as developing 30–70-year-old lodgepole pine stands with high stem density that structurally mimic multi-storied forests. © 2012 The Wildlife Society.