Persistent kelp forests during a massive decline reveal the importance of land-sea connectivity.

A fundamental goal in ecology is to understand the drivers of stability in natural ecosystems in the face of disturbances. However, this is challenging when biotic and abiotic stressors operate simultaneously across multiple spatial scales. Such is the case for bull kelp forests (Nereocystis luetkeana) in northern California, where losses of predators combined with marine heatwaves have led to shifts from kelp forest to sea urchin barren states. However, despite the >90% loss of bull kelp forests since 2014, some patches remain. Here, we investigate the bull kelp community assemblage in these remnant patches as well as the drivers of bull kelp forest resistance. We used a combination of in situ field surveys (years 2020-2022), remote sensing data (years 2016-2022), and a laboratory grazing experiment with urchins (Strongylocentrotus purpuratus). We found that, in addition to the two dominant states (kelp forest vs. urchin barren), there is a third community state dominated by understory canopy-forming macroalgae that stays subsurface. Moreover, bull kelp abundance and cover were positively associated with freshwater flow and proximity to freshwater sources, and bull kelp persistence was positively associated with sand cover, all of which seem to diminish sea urchin abundance and the negative effects of sea urchin herbivory on bull kelp. This was also shown in the laboratory experiment where sea urchin herbivory rates on bull kelp decreased with decreasing salinity. Overall, these results suggest that freshwater influence in shallow coastal environments could prevent loss of bull kelp and show that land-sea connections should be considered for species-specific management and conservation actions.