Keystone predation can be a determinant of community structure, including species diversity, but factors underlying “keystoneness” have been minimally explored. Using the system in which the original keystone, the sea star Pisaster ochraceus, was discovered, we focused on two potential (but overlapping) determinants of keystoneness: intrinsic traits or state variables of the species (e.g., size, density), and extrinsic environmental parameters (e.g., prey productivity) that may provide conditions favorable for keystone predator evolution. Using a comparative-experimental...