Understanding dietary specialization in herbivores has theoretical and practical implications in ecology, yet defining niche breadth consistently has been problematic. To increase clarity and communication among ecologists and among disciplines (i.e., chemists, pharmacologists), we propose a specialization key for mammalian herbivores that assigns “obligatory” and “facultative” modifiers to the terms “specialist” and “generalist”. These modifiers are assigned based on (1) relative breadth of the animal's realized niche and diet (what it eats), (2) relative breadth of the fundamental niche and available diet (what it could eat), (3) the extent of chemical or physical characteristics, termed “difficulty”, that make food items either low in value or unpalatable to most herbivores, and (4) relevant temporal and spatial scales at which diets and niche breadth were measured. Obligatory specialists always have a narrow realized niche consisting of difficult food items, and morphological adaptations and/or the loss of redundant behavioral flexibility that effectively limit their fundamental niches, precluding them from expanding their diet under changed environmental conditions. Facultative specialists have a consistently narrow realized niche for difficult foods during at least one spatial or temporal scale, but have a broad enough fundamental niche to allow them to expand their diet to include less difficult foods when environmental conditions allow. Facultative generalists have the broadest fundamental niche, allowing them to consume a wide variety of foods. However, they may occasionally demonstrate a narrow realized niche, focused on less difficult plants than is the case with specialists. Finally, the obligatory generalists always have a wide realized niche because of a relatively narrow fundamental niche, precluding them from eating much of any difficult plant. We summarize hypothesized characteristics of mammalian herbivores in each of the four categories of specialization. We demonstrate the need for further work on defining the realized and fundamental niches, comparing among herbivores across categories conducted under similar conditions, and understanding the nature of trade-offs required for specialization and generalization for both community and phylogenetically based analyses.
Mule (Odocoileus hemionus) and white‐tailed deer (Odocoileus virginianus) are congeneric and share similar life histories, yet their distribution is segregated across much of North America. Extensive research on both species within and outside their zone of co‐occurrence has not fully explained these distribution patterns, especially the potential role of diet and foraging behavior. Therefore, we used a common garden experiment to compare diet composition, diet quality, foraging behavior, and intake of tractable mule and white‐tailed deer foraging together within the dry Douglas‐fir (Pseudotsuga menziesii)/ponderosa pine (Pinus ponderosa) forests of northeastern Washington. We sampled at 21 0.5‐ha sites from June to August 2016. We used standard bite count techniques coupled with forage biomass sampling, behavioral observations, and nutritional analyses to compare the foraging ecology of the two species. Mule and white‐tailed deer had similar activity patterns. However, mule deer took larger bites and harvested food faster than white‐tailed deer, and white‐tailed deer consumed more diverse but higher‐quality diets than mule deer. These differences resulted in mule deer acquiring ~25% more dry matter and digestible energy per day. About 90% of the diets consumed by both deer species consisted of deciduous shrubs and forbs, and they selected many of the same plant species. However, overall diet composition was 38% dissimilar, with mule deer consuming diets that were more likely to contain shrubs with higher levels of tannins and lower levels of dry matter digestibility than diets eaten by white‐tailed deer. Dietary overlap was greatest at both very low and very high forage biomass, indicating potential for modest resource competition or partitioning. Our research provides evidence that differences in diet composition of mule and white‐tailed deer do not merely reflect differences in habitat selection, but also suggest the species differ in their fundamental nutritional niches.