Moving ocean acidification research beyond a simple science: Investigating ecological change and their stabilizers

The response of complex ecological communities to ocean acidification reflects interactions among species that propagate or dampen ecological change. Yet, most studies have been based on short-term experiments with limited numbers of interacting species. Both limitations tend to exaggerate measured effects and when combined with our predisposition for investigating change, we reduce insight into pathways of stability, acclimation and adaptation. Here, we review accepted and emerging insights into processes that drive ecological change (top-down and bottom-up) and the stabilizing processes by which ecological complexity may dampen change. With an emphasis on kelp forest examples, we show that boosted primary productivity from enriched CO2 creates competitive imbalances that drive habitat change, but we also recognise intensifying herbivory on these habitats dampens this change. Foraging herbivores thrive on CO2 enriched plants and over successive generations their populations expand. When we consider such population level responses, we open new questions regarding density-effects (e.g. competition, susceptibility to predation and disease), as well as the bottom-up benefits to predators. Nevertheless, research on predators has lagged behind because their wide-ranging behaviour typically imposes logistical difficulties for observational and experimental research. We know that ocean warming imposes elevated metabolic costs on their foraging whilst acidification hampers navigation of their larvae towards suitable habitat and impairs their hunting and avoidance of predators as adults. Connecting such top-down with bottom-up responses is fundamental for progress, and is also contingent on understanding the mechanisms that dampen change. These stabilizers have the potential to keep pace with abiotic change and thereby influence the drivers of acclimation and adaption. Certainly, we acknowledge that investigating change is often simpler and the associated bold messages appeal to citation impact. Yet, if we are to anticipate the ability of complex ecological communities to persist in changing environments, then understanding the shifting balance between the propagation of resource enrichment and its consumption across trophic levels is central to this challenge.