Abstract

In heterogeneous landscapes, large herbivores employ plastic behavioral strategies to buffer themselves against negative effects of environmental variation on fitness. Yet, the mechanisms by which individual responses to such variation scale up to influence population performance remain uncertain. Analyses of space-use behaviors exemplify this knowledge gap, because such behaviors are often assumed, but rarely demonstrated, to have direct fitness consequences. We combined fine-scale data on forage biomass and quality with movement data and measures of somatic energy reserves to determine whether variation in use (the quantity of resource units, e.g., pixels on a landscape, that receive some level of investment by an animal during a specific sampling period) or selection (use of a resource unit relative to its availability to the animal during the same sampling period) of the nutritional landscape predicted early winter body condition of mule deer (Odocoileus hemionus). At the population level, mule deer exhibited stronger selection for high forage biomass at the landscape scale than at the home-range scale, and during summer than during spring. Use of the nutritional landscape varied among individual deer and had important consequences for early winter condition (an important determinant of survival and reproduction in capital-breeding ungulates). Females that consistently used vegetation communities that provided high biomass of preferred forage plants throughout spring and summer entered winter in better condition than females that used those vegetation communities less frequently. In contrast, selection (i.e., use relative to availability) of the nutritional landscape by individual deer was not significantly related to early winter condition at either the landscape or home-range scales. Our results highlight the value of using mechanistic, nutritional approaches to understand the potential fitness consequences of individual variation in behavior. In addition, our study suggests that patterns of forage use by ungulates may sometimes correlate more strongly with fitness than patterns of forage selection, which are scale-dependent and more vulnerable to biases stemming from the need to accurately quantify availability.

Highlights

  • IntroductionFree-ranging animals often are forced to make complex tradeoffs among factors that influence fitness (e.g., forage versus predation risk; Berger, 1991; Barten et al, 2001)

  • In heterogeneous landscapes, free-ranging animals often are forced to make complex tradeoffs among factors that influence fitness

  • Three deer were monitored during both years of the study, and we considered condition measurements from those deer to be independent based on longitudinal condition data obtained from mule deer in Wyoming

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Summary

Introduction

Free-ranging animals often are forced to make complex tradeoffs among factors that influence fitness (e.g., forage versus predation risk; Berger, 1991; Barten et al, 2001). In temperate and polar environments, large terrestrial herbivores exhibit life-history strategies that revolve around seasonal changes in resource availability and the energetic demands imposed by key life-history events (Monteith et al, 2013) These long-lived mammals must accrue sufficient energy and protein reserves during summer to replenish reserves lost over winter, while meeting the demands of lactation without endangering future survival and reproduction (Therrien et al, 2008; Bårdsen et al, 2010; Tollefson et al, 2010; Bårdsen and Tveraa, 2012). A variety of factors (e.g., competition and risk of predation) may limit the ability of herbivores to optimize their use of the nutritional landscape (i.e., variation in the density of energy and protein across space and time; Pretorius et al, 2011) Under these conditions, differences in use of the nutritional landscape among individuals can have important fitness consequences (van Beest and Milner, 2013; Long et al, 2016)

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