Evolution in the Consumer Age: Predators and the History of Life

Three properties of predation make this form of consumption an important agency of evolution: universality (all species have predators), high frequency (encounters of prey with predators test both parties often), and imperfection (many predatory attacks fail, enabling antipredatory selection to take place). On long time scales, predators have two principal effects: they influence their victims' phenotypes, and prey species that are highly vulnerable to all phases of predatory attacks are evolutionarily restricted to environments where predators are rarely encountered. Although predator and prey can affect each other's behavior and morphology on timescales commensurate with individual lifespans, predators have the evolutionary upper hand over the long run, especially in the expression of sensory capacities, locomotor performance, and the application of force. Only in passive defenses (armor, toxicity, large body size) does escalation favor the prey. In a review of methods for inferring predation in the geological past, I argue against the use of whole assemblages, which combine species of contrasting adaptive type, Instead, I strongly favor species-level and clade-level approaches (including examples of clade replacement) in which comparisons among places and among time intervals are made within the same adaptive types and the same physical environments. The available evidence, much of which comes from studies of shell drilling and shell breakage, points to temporal increases in both predator power and prey defenses. Escalation between species and their enemies, including predators, has proceeded episodically against a backdrop of generally increasing productivity and increasing top-down evolutionary control by high-energy predators during the Phanerozoic, the consumer age.