In defense of elemental currencies: can ecological stoichiometry stand as a framework for terrestrial herbivore nutritional ecology?

Nutritional ecologists aim to predict population or landscape-level effects of food availability, but the tools to extrapolate nutrition from small to large extents are often lacking. The appropriate nutritional ecology currencies should be able to represent consumer responses to food while simultaneously be simple enough to expand such responses to large spatial extents and link them to ecosystem functioning. Ecological stoichiometry (ES), a framework of nutritional ecology, can meet these demands, but it is typically associated with ecosystem ecology and nutrient cycling, and less often used to study wildlife nutrition. Despite the emerging zoogeochemical evidence that animals, and thus their diets, play critical roles in nutrient movement, wildlife nutritional ecology has not fully embraced ES, and ES has not incorporated nutrition in many wildlife studies. Here, we discuss how elemental currencies are "nutritionally, organismally, and ecologically explicit" in the context of terrestrial herbivore nutritional ecology. We add that ES and elemental currencies offer a means to measure resource quality across landscapes and compare nutrient availability among regions. Further, we discuss ES shortcomings and solutions, and list future directions to advance the field. As ecological studies increasingly grow in spatial extent, and attempt to link multiple levels of biological organization, integrating more simple and unifying currencies into nutritional studies, like elements, is necessary for nutritional ecology to predict herbivore occurrences and abundances across regions.