The adaptive evolution of herbivory in freshwater systems

Herbivory is thought to be nutritionally inefficient relative to carnivory and omnivory, but herbivory evolved from carnivory in many terrestrial and aquatic lineages, suggesting that there are advantages of eating plants. Herbivory has been well-studied in both terrestrial and aquatic systems, and there is abundant information on feedbacks between herbivores and plants, coevolution of plant and herbivore defenses, mechanisms for mediating nutrient limitation, effects of nutrient limitation on herbivore life history, and, more recently, the origins of the herbivorous diet. Researchers have sufficiently defined the ecological context and evolutionary origins of the herbivorous diet, and these main areas of research have laid the groundwork for studying herbivory as an adaptation. However, we have yet to synthesize this information in a way that allows us to establish a framework of testable adaptive hypotheses. To understand the adaptive significance of this diet transition, we review the current literature and use evidence from these works as support for five hypotheses on the evolution of herbivory from carnivory: (1) intake efficiency—herbivores use part of their food source as habitat, thus minimizing the energy/time spent searching for food and avoiding predators; (2) suboptimal habitat—herbivory allows organisms to invade and establish populations in habitats that have high primary production but low abundance of animal prey; (3) heterotroph facilitation—herbivory is adaptive because herbivores consume microbes associated with producers; (4) lipid allocation—herbivory is adaptive because producers are rich in fatty acids, which fuel reproduction and storage; and (5) disease avoidance—herbivory minimizes animal-facilitated disease transmission. Due to the extensive literature, we have limited this review to discussing herbivory in freshwater systems. To our knowledge, no prior work has compiled a comprehensive list of conditions that favor an herbivorous diet in nature. With backgrounds in both theoretical and experimental ecology, the incorporation of these hypotheses to the current literature will provide information about diet evolution, where it is currently lacking