Individual and population level effects of ocean acidification on a predator-prey system with inducible defenses: bryozoan-nudibranch interactions in the Salish Sea

Ocean acidification (OA) from increased oceanic CO2 concentrations imposes significant physiological stresses on many calcifying organisms. OA effects on individual organisms may be synergistically amplified or reduced by inter- and intraspecies interactions as they propagate up to population and community levels, altering predictions by studies of calcifier responses in isolation. The calcifying colonial bryozoan Membranipora membranacea and the predatory nudibranch Corambe steinbergae comprise a trophic system strongly regulated by predator-induced defensive responses and space limitation, presenting a unique system to investigate OA effects on these regulatory mechanisms at individual and population levels. We experimentally quantified OA effects across a range of pH from 7.0 to 7.9 on growth, calcification, senescence and predator-induced spine formation in Membranipora, with or without waterborne predator cue, and on zooid consumption rates in Corambe at Friday Harbor Laboratories, San Juan Island, WA. Membranipora exhibited maximum growth and calcification at moderately low pH (7.6), and continued spine formation in all pH treatments. Spines reduced Corambe zooid consumption rates, with lower pH weakening this effect. Using a spatially explicit model of colony growth, where colony area serves as a proxy for colony fitness, we assessed the population-level impacts of these experimentally determined individual-level effects in the context of space limitation. The area-based fitness costs associated with defense measured at the individual level led to amplified effects predicted for the population level due to competition. Our coupled experimental and modeling results demonstrate the need to consider population-level processes when assessing ecological responses to stresses from changing environments.