Benthic invertebrates in a high CO2 world

Ocean acidification (OA), whereby increases in atmospheric carbon dioxide (CO2) over the past 200 years have led to a decline in the pH and carbonate ion availability of the oceans, has emerged as one of the major drivers of twenty- first century marine scientific research. Here we describe the current understanding of OA effects on benthic marine invertebrates, in particular the calcifiers thought to be most sensitive to altered carbonate chemistry. We describe the responses of benthic invertebrates to OA conditions predicted up to the end of the century, examining individual organism response through to ecosystem- level impacts. Research over the past decade has found great variability in the physiological and functional response of different species and communities to OA, with further variability evident between life stages. Over both geological and recent timescales, the presence and calcification rates of marine calcifiers have been inextricably linked to the carbon chemistry of the oceans. Under short-term experimentally enhanced CO2 conditions, many organisms have shown trade-offs in their physiological responses, such as reductions in calcification rate and reproductive output. In addition, carry-over effects from fertilization, larval and juvenile stages, such as enhanced development time and morphological changes, highlight the need for broad- scale studies over multiple life stages. These organism- level responses may propagate through to altered benthic communities under naturally enhanced CO2 conditions, evident in studies of upwelling regions and at shallow- water volcanic CO2 vents. Only by establishing which benthic invertebrates have the ability to acclimate or adapt, via natural selection, to changes from OA, in combination with other environmental stressors, can we begin to predict the consequences of future climate change for these communities.