Predators indirectly induce stronger prey through a trophic cascade

Many prey species induce defences in direct response to predation cues. However, prey defences could also be enhanced by predators indirectly via mechanisms that increase resource availability to prey, e.g. trophic cascades. We evaluated the relative impacts of these direct and indirect effects on the mechanical strength of the New Zealand sea urchin Evechinus chloroticus. We measured crush-resistance of sea urchin tests (skeletons) in (i) two marine reserves, where predators of sea urchins are relatively common and have initiated a trophic cascade resulting in abundant food for surviving urchins in the form of kelp, and (ii) two adjacent fished areas where predators and kelps are rare. Sea urchins inhabiting protected rocky reefs with abundant predators and food had more crush-resistant tests than individuals on nearby fished reefs where predators and food were relatively rare. A six-month long mesocosm experiment showed that while both food supply and predator cues increased crush-res...