Changes in abiotic drivers of green sea urchin demographics following the loss of a keystone predator

Sea urchin population demographics can respond to changes in keystone species abundances, with the magnitude of these responses varying depending on environmental influences. In this study, sea urchin populations were surveyed across 15 Aleutian archipelago islands over a 30-year period to understand how patterns of sea urchin demography (density, biomass, and size structure) varied through different ecological regimes that were caused by changes in the abundance of sea otters, a keystone species in this system. To examine long-term changes in sea urchin demographics, four time periods across the recent decline of sea otters were examined: during sea otter presence (1987-1994), nearing absence at the end of the decline (1997-2000), 10 years postdecline (2008-2010), and 15-20 years following the loss of sea otters from the ecosystem (2014-2017). Our results show that when sea otters were broadly present, sea urchin demographics were generally similar across the archipelago, with few urchins that had large-sized bodies. During this time, bottom-up environmental controls were muted relative to top-down forces from keystone predation. However, as sea otters declined and remained absent from the system, abiotic factors became more influential on sea urchin biomass, density, and size structure. In particular, differences among island groups during these periods were correlated with variation in ocean temperature, bathymetric complexity, and habitat availability. Sea urchin recruitment also varied among island groups, corresponding to ecoregions delineated by oceanic passes across the archipelago. The functional extinction of sea otters revealed an increasing influence of abiotic forcing in the absence of top-down control. This study further highlights the importance of understanding how keystone predators regulate herbivore demographics.