Marine Protected Areas That Preserve Trophic Cascades Promote Resilience of Kelp Forests to Marine Heatwaves.

Under accelerating threats from climate-change impacts, marine protected areas (MPAs) have been proposed as climate-adaptation tools to enhance the resilience of marine ecosystems. Yet, debate persists as to whether and how MPAs may promote resilience to climate shocks. Here, we use 38 years of satellite-derived kelp cover to empirically test whether a network of 58 temperate coastal MPAs in Central and Southern California enhances the resistance of kelp forest ecosystems to, and their recovery from, the unprecedented 2014-2016 marine heatwave regime that occurred in the region. We also leverage a 22-year time series of subtidal community surveys to mechanistically understand whether trophic cascades explain emergent patterns in kelp forest resilience within MPAs. We find that fully protected MPAs significantly enhance kelp forests' resistance to and recovery from marine heatwaves in Southern California, but not in Central California. Differences in regional responses to the heatwaves are partly explained by three-level trophic interactions comprising kelp, urchins, and predators of urchins. Urchin densities in Southern California MPAs are lower within fully protected MPAs during and after the heatwave, while the abundances of their main predators-lobster and sheephead-are higher. In Central California, a region without lobster or sheephead, there is no significant difference in urchin or kelp densities within MPAs as the current urchin predator, the sea otter, is protected statewide. Our analyses show that fully protected MPAs can be effective climate-adaptation tools, but their ability to enhance resilience to extreme climate events depends upon region-specific environmental and trophic interactions. As nations progress to protect 30% of the oceans by 2030, scientists and managers should consider whether protection will increase resilience to climate-change impacts given their local ecological contexts, and what additional measures may be needed.