Study system
Our study included 4 sites along the Oregon coast: Strawberry Hill (44.250°N-124.115°W) and Yachats Beach (44.319°N-124.109°W), located on Cape Perpetua, and Fogarty Creek (44.837°N-124.0587°W) and Boiler Bay (44.832°N-124.061°W) located on Cape Foulweather (Fig.1). Cape Perpetua is a rocky headland adjacent to a wide continental shelf offshore that promotes retention of propagules such as larvae and phytoplankton (Menge et al. 2015). As a result, Cape Perpetua intertidal sites are characterized by high phytoplankton productivity and high recruitment of invertebrates (Menge et al. 1997, 2004, 2015). Cape Foulweather, in contrast, is characterized by a narrower offshore continental shelf, which leads to reduced retentiveness, lower invertebrate abundance, and high macrophyte abundance. Prior to the onset of SSWD, densities of P. ochraceus could be as high as 8 individuals m-2 at Cape Perpetua and 4 individuals m-2 at Cape Foulweather sites (Menge et al. 2016).

Subordinate predator removal experiment
To assess the effects of subordinate predators in the absence of the keystone, we conducted a factorial removal experiment at two intertidal sites located on Cape Perpetua, Oregon (Strawberry Hill and Yachats Beach, see Fig. 1). We predicted that whelk predators would affect establishment of the dominant mussel, Mytilus californianus, by consuming the mid-successional prey species that facilitate its recruitment. As such, we chose to follow prey dynamics from a mid-successional stage by placing plots where there was abundant cover of the mussel Mytilus trossulus and several barnacle species. This mid-successional community is where we expected to see the greatest effects of subordinate predators and the largest changes in community structure following SSWD. We originally examined the effects of two groups of subordinate predators, gastropod whelks Nucella canaliculata and N. ostrina (W) and the smaller sea star Leptasterias spp. (L), in a factorial design including four treatments: +W +L, -W +L, +W -L, and -W -L. However, Leptasterias spp. were rare in our plots, and treatments were combined to include control (+W) and whelk removal (-W) treatments only (see data analysis section below). It is important to note that our experiment tested the effects of subordinate predators at reduced P. ochraceus densities, rather than comparing their effects in the presence or absence of P. ochraceus.

The main treatment plots were 0.25 m2 in size and corners were marked with stainless steel lag screws. Each main plot was surrounded by four additional subplots adjacent to each plot side that were meant to act as a buffer for the main plot. We monitored plots either bi-weekly or monthly as tides permitted from experimental initiation in June 2014 through May 2015. At each monitoring, we counted the total number of each subordinate predator species (N. canaliculata, N. ostrina, and Leptasterias spp.), as well as any less common predators, such as the whelk Nucella lamellosa, in all plots. When possible, we conducted a full monitoring with counts and removals of predators in the main plot, the four adjacent subplots, and in the corners between subplots. On some occasions, particularly during winter months with limited site access, we only monitored the main plots. All removal of whelks and Leptasterias spp. was conducted using forceps, and removed predators were relocated away from the plot area. In plots without removals, we mimicked the use of forceps in the plot while counting subordinate predators to limit the possibility that the physical action of predator removal would influence our results. Although SSWD caused declines in Pisaster ochraceus densities, it did not extirpate the species entirely from our sites. We recorded and relocated any P. ochraceus within each plot and any adult and juvenile P. ochraceus from a 3m radius around each plot.

pred_removal.csv