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The snowshoe hare (Lepus americanus) possesses a broad suite of adaptations to winter, including a seasonal coat color molt. Recently, climate change has been implicated in the range contraction of snowshoe hares along the southern range boundary. With shortening snow season duration, snowshoe hares are experiencing increased camouflage mismatch with their environment reducing survival. Phenological variation of hare molt at regional scales could facilitate local adaptation in the face of climate change, but the level of variation, especially along the southern range boundary, is unknown. Using a network of trail cameras and historical museum specimens, we (1) developed contemporary and historical molt phenology curves in the Upper Great Lakes region, USA, (2) calculated molt rate and variability in and among populations, and (3) quantified the relationship of molt characteristics to environmental conditions for snowshoe hares across North America. We found that snowshoe hares across the region exhibited similar fall and spring molt phenologies, rates and variation. Yet, an insular island population of hares on Isle Royale National Park, MI, completed their molt a week earlier in the fall and initiated molt almost 2 weeks later in the spring as well as exhibited slower rates of molting in the fall season compared to the mainland. Over the last 100 years, snowshoe hares across the region have not shifted in fall molt timing; though contemporary spring molt appears to have advanced by 17 days (~ 4 days per decade) compared to historical molt phenology. Our research indicates that some variation in molt phenology exists for snowshoe hares in the Upper Great Lakes region, but whether this variation is enough to offset the consequences of climate change remains to be seen. [ABSTRACT FROM AUTHOR]

Intraspecific feeding choices comprise a large portion of herbivore foraging decisions. Plant resource quality is heterogeneously distributed, affected by nutrient availability and growing conditions. Herbivores navigate landscapes, foraging not only according to food qualities, but also energetic and nutritional demands. We test three non-exclusive foraging hypotheses using the snowshoe hare (Lepus americanus): (1) herbivore feeding choices and body conditions respond to intraspecific plant quality variation; (2) high energetic demands mitigate feeding responses; and (3) feeding responses are inflated when nutritional demands are high. We measured black spruce (Picea mariana) nitrogen, phosphorus and terpene compositions, as indicators of quality, within a snowshoe hare trapping grid and found plant growing conditions to explain spruce quality variation (R2 < 0.36). We then offered two qualities of spruce (H1) from the trapping grid to hares in cafeteria-style experiments and measured their feeding and body condition responses (n = 75). We proxied energetic demands (H2) with ambient temperature and coat insulation (% white coat) and nutritional demands (H3) with the spruce quality (nitrogen and phosphorus content) in home ranges. Hares with the strongest preference for high-quality spruce lost on average 2.2% less weight than hares who ate the least high-quality spruce relative to low-quality spruce. The results supported our energetic predictions as follows: hares in colder temperatures and with less-insulative coats (lower % white) consumed more spruce and were less selective towards high-quality spruce. Collectively, we found variation in plant growing conditions within herbivore home ranges substantial enough to affect herbivore body conditions, but energetic stats mediate plant–herbivore interactions. [ABSTRACT FROM AUTHOR]

Home range size of consumers varies with food quality, but the many ways of defining food quality hamper comparisons across studies. Ecological stoichiometry studies the elemental balance of ecological processes and offers a uniquely quantitative, transferrable way to assess food quality using elemental ratios, e.g., carbon (C):nitrogen (N). Here, we test whether snowshoe hares (Lepus americanus) vary their home range size in response to spatial patterns of C:N, C:phosphorus (P), and N:P ratios of two preferred boreal forage species, lowbush blueberry (Vaccinium angustifolium) and red maple (Acer rubrum), in summer months. Boreal forests are N- and P-limited ecosystems and access to N- and P-rich forage is paramount to snowshoe hares' survival. Accordingly, we consider forage with higher C content relative to N and P to be lower quality than forage with lower relative C content. We combine elemental distribution models with summer home range size estimates to test the hypothesis that home range size will be smaller in areas with access to high, homogeneous food quality compared to areas of low, heterogeneous food quality. Our results show snowshoe hares had smaller home ranges in areas where lowbush blueberry foliage quality was higher or more spatially homogenous than in areas of lower, more heterogeneous food quality. By responding to spatial patterns of food quality, consumers may influence community and ecosystem processes by, for example, varying nutrient recycling rates. Our reductionist biogeochemical approach to viewing resources leads us to holistic insights into consumer spatial ecology. [ABSTRACT FROM AUTHOR]

Determining the factors driving cyclic dynamics in species has been a primary focus of ecology. For snowshoe hares (Lepus americanus), explanations of their 10-year population cycles most commonly feature direct predation during the peak and decline, in combination with their curtailment in reproduction. Hares are thought to stop producing third and fourth litters during the cyclic decline and do not recover reproductive output for several years. The demographic effects of these reproductive changes depend on the consistency of this pattern across cycles, and the relative contribution to population change of late-litter versus early litter juveniles. We used monitoring data on snowshoe hares in Yukon, Canada, to examine the contribution of late-litter juveniles to the demography of their cycles, by assigning litter group for individuals caught in autumn based on body size and capture date. We found that fourth-litter juveniles occur consistently during the increase phase of each cycle, but are rare and have low over-winter survival (0.05) suggesting that population increase is unlikely to be caused by their occurrence. The proportion of third-litter juveniles captured in the autumn remains relatively constant across cycle phases, while over-winter survival rates varies particularly for earlier-litter juveniles (0.14–0.39). Juvenile survival from all litters is higher during the population increase and peak, relative to the low and decline. Overall, these results suggest that the transition from low phase to population growth may stem in large part from changes in juvenile survival as opposed to increased reproductive output through the presence of a 4th litter. [ABSTRACT FROM AUTHOR]

Vertebrate populations throughout the circumpolar north often exhibit cyclic dynamics, and predation is generally considered to be a primary driver of these cycles in a variety of herbivore species. However, weather and climate play a role in entraining cycles over broad landscapes and may alter cyclic dynamics, although the mechanism by which these processes operate is uncertain. Experimental and observational work has suggested that weather influences primary productivity over multi-year time periods, suggesting a pathway through which weather and climate may influence cyclic herbivore dynamics. Using long-term monitoring data, we investigated the relationships among multi-year weather conditions, measures of primary productivity, and the abundance of two cyclic herbivore species: snowshoe hare and northern red-backed vole. We found that precipitation (rain and snow) and growing season temperatures were strongly associated with variation in primary productivity over multi-year time horizons. In turn, fourfold variation in the amplitude of both the hare and vole cycles observed in our study area corresponded to long-term changes in primary productivity. The congruence of our results for these two species suggests a general mechanism by which weather and climate might influence cyclic herbivore population dynamics. Our findings also suggested that the association between climate warming and the disappearance of cycles might be initiated by changes in primary productivity. This work provides an explanation for observed influences of weather and climate on primary productivity and population cycles and will help our collective understanding of how future climate warming may influence these ecological phenomena in the future. [ABSTRACT FROM AUTHOR]

Isodar theory can help to unveil the fitness consequences of habitat disturbance for wildlife through an evaluation of adaptive habitat selection using patterns of animal abundance in adjacent habitats. By incorporating measures of disturbance intensity or variations in resource availability into fitness-density functions, we can evaluate the functional form of isodars expected under different disturbance-fitness relationships. Using this framework, we investigated how a gradient of forest harvesting disturbance and differences in resource availability influenced habitat quality for snowshoe hares ( Lepus americanus) and red-backed voles ( Myodes gapperi) using pairs of logged and uncut boreal forest. Isodars for both species had positive intercepts, indicating reductions to maximum potential fitness in logged stands. Habitat selection by hares depended on both conspecific density and differences in canopy cover between harvested and uncut stands. Fitness-density curves for hares in logged stands were predicted to shift from diverging to converging with those in uncut forest across a gradient of high to low disturbance intensity. Selection for uncut forests thus became less pronounced with increasing population size at low levels of logging disturbance. Voles responded to differences in moss cover between habitats which reflected moisture availability. Lower moss cover in harvested stands either reduced maximum potential fitness or increased the relative rate of decline in fitness with density. Differences in vole densities between harvested and uncut stands were predicted, however, to diminish as populations increased. Our findings underscore the importance of accounting for density-dependent behaviors when evaluating how changing habitat conditions influence animal distribution. [ABSTRACT FROM AUTHOR]

The carbon/nutrient balance hypothesis fails to correctly predict effects of fertilization and shading on concentrations of defensive metabolites in Alaskan balsam poplar ( Populus balsamifera). Of six metabolites analyzed, only one responded in the predicted fashion to fertilization and one to shading. These results and those of other similar studies suggest that while the carbon/nutrient balance hypothesis may correctly predict the effects of fertilization and shading on the concentrations of metabolic 'end products', it fails for many metabolites because of the dynamics associated with their production and turnover. In metabolites that turn over, static concentration is a poor predictor of defensive investment. [ABSTRACT FROM AUTHOR]

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 CHCl 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. [ABSTRACT FROM AUTHOR]

We relate causes of mortality of snowshoe hares to density of hares over an 8-year period that included a peak in numbers. We then use simulation modeling to examine whether these density-dependent relationships could produce changes in hare density similar to those observed in our study are in Yukon, Canada. Predation during winter was the largest source of mortality for snowshoe hares at Kluane, Yukon during 1978-84. There was a one-year lag in the response of winter predation mortality rate to hare density. There was a two-year lag in the response of winter mortality not caused by predators to hare density. A simple simulation model with density-dependent predation produced 8-11 year cycles only within a narrow range of parameters that are inconsistent with data from the Kluane region. However, a simulation model that predicted winter mortality rates using a delayed density-dependent numerical response and a Type II functional response by predators, produced 8-11 year cycles within the range of parameter values measured in our study. Yet another simulation model that predicted both summer and winter mortality rates using a delayed density-dependent numerical response and a Type II functional response by predators, did not produce 8-11 year cycles within the range of parameter values measured in our study. Lack of data on juvenile mortality may be one reason for this result. [ABSTRACT FROM AUTHOR]

Plant carbon/nutrient balance has been implicated as an important factor in plant defensive chemistry and palatability to herbivores. We tested this hypothesis by fertilizing juvenile growth form Alaska paper birch and green alder with N, P and N-plus-P in a balanced 2x2 factorial experiment. Additionally, we shaded unfertilized plants of both species. Fertilization with N and N-plus-P increased growth of Alaska paper birch, reduced the concentration of papyriferic acid in internodes and increased the palatability of birch twigs to snowshoe hares. Shading decreased birch growth, decreased the concentration of papyriferic acid in internodes and increased twig palatability. These results indicate that the defensive chemistry and palatability of winter-dormant juvenile Alaska paper birch are sensitive to soil fertility and shade. Conversely the defensive chemistry and palatability of green alder twigs to snowshoe hares were not significantly affected by soil fertility or shade. The greater sensitivity of Alaska paper birch defensive chemistry and palatability to snowshoe hares in comparison to green alder is in agreement with the hypothesis that early successional woody plants that are adapted to high resource availability are more plastic in their chemical responses to the physical environment than are species from less favorable environments. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]