Consumers make space use decisions based on resource quality. Most studies that investigate the influence of resource quality on the spatial ecology of consumers use diverse proxies for quality including measures based on habitat classification, forage species diversity and abundance, and nutritional indicators, e.g., protein. Ecological stoichiometry measures resource quality in terms of elemental ratios, e.g., carbon (C):nitrogen (N) ratio, but rarely have these currencies been used to study consumer space use decisions. Yet, elemental ratios provide a uniquely quantitative way to assess resource quality. Consequently, ecological stoichiometry allows for investigation of how consumers respond to spatial heterogeneity in resource quality by changing their space use, e.g. their home range size, and how this may influence ecosystem dynamics and trophic interactions. Here, we test whether the home range size of a keystone boreal herbivore, the snowshoe hare (Lepus americanus), varies with differences in the C:N, C:phosphorus (P), and N:P ratios of two preferred forage species, lowland blueberry (Vaccinium angustifolium) and red maple (Acer rubrum). We consider forage resources with higher C content relative to N and P to be lower quality than resources with lower relative C content. We use a novel approach, combining elemental distribution models with herbivore home range size estimates to test our hypothesis that hare home range size will be smaller in areas with access to high, homogeneous resource quality compared to areas with access to low, heterogeneous resource quality during summer months. Our results support our prediction for lowland blueberry, but not for red maple. Herbivore home range size decreased with increasing blueberry foliage quality, but also with decreasing spatial heterogeneity in blueberry foliage quality, i.e. N or P content. Herbivores in the boreal forest face strong nutritional constraints due to the paucity of N and P. Access to areas of high, homogeneous resource quality is paramount to meeting their dietary requirements with low effort. In turn, this may influence community (e.g., trophic interactions) and ecosystem (e.g., nutrient cycling) processes. Paradoxically, our study shows that taking a reductionist approach of viewing resources through a biochemical lens can lead to holistic insights of consumer spatial ecology.

Intraspecific variability in ecological traits is widespread in nature. Recent evidence, mostly from aquatic ecosystems, shows individuals differing at the most fundamental level, that of their chemical composition. Age, sex, or body size and condition may be key drivers of intraspecific variability in the body concentrations of carbon (C), nitrogen (N), and phosphorus (P). However, we still have a rudimentary understanding of the patterns and drivers of intraspecific variability in chemical composition of terrestrial consumers, particularly vertebrates. Here, we investigate the elemental composition of the snowshoe hare Lepus americanus. Based on snowshoe hare ecology, we predicted older, larger individuals to have higher concentration of N or P and lower C content compared with younger, smaller individuals. We also predicted females to have higher concentrations of N, P, and lower C than males due to the higher reproductive costs they incur. Finally, we predicted that individuals in better body condition would have higher N and P than those in worse condition, irrespective of age. We obtained C, N, and P concentrations and ratios from a sample of 50 snowshoe hares. We then used general linear models to test our predictions on the relationship between age, sex, body size or condition and stoichiometric variability in hares. We found considerable variation in C, N, and P stoichiometry within our sample. Contrary to our predictions, we found weak evidence of N content decreasing with age. As well, sex appeared to have no relationship with hare body elemental composition. Conversely, as expected, P content increased with body size and condition. Finally, we found no relationship between variability in C content and any of our predictor variables. Snowshoe hare stoichiometry does not appear to vary with individual age, sex, body size, or condition. However, the weak relationship between body N concentration and age may suggest varying nutritional requirements of individuals at different ages. Conversely, body P's weak relationship to body size and condition appears in line with this limiting element's importance in terrestrial ecosystems. Snowshoe hares are keystone herbivores in the boreal forest of North America, and the substantial stoichiometric variability we find in our sample could have important implications for nutrient dynamics, in both boreal and adjacent ecosystems.

Intraspecific feeding choices account for a large portion of herbivore foraging in many ecosystems. Plant resource quality is heterogeneously distributed, affected by nutrient availability and growing conditions. Herbivores navigate landscapes, making feeding decisions according to food qualities, but also energetic and nutritional demands. We test three nonexclusive foraging hypotheses using the snowshoe hare (Lepus americanus): 1) herbivores feeding choices and body conditions respond to intraspecific plant quality variation, 2) feeding responses are mitigated when energetic demands are high, 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 (R 2 < 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 that preferred higher-quality spruce lost less weight during experiments (p = 0.018). The results supported our energetic predictions: 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 any plant-herbivore interactions are also mediated by animal energetic states.

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