Competition in benthic marine invertebrates: the unrecognized role of exploitative competition for oxygen.

Competition is a ubiquitous structuring force across systems, but different fields emphasize the role of different types of competition. In benthic marine environments, where some of the classic examples of competition were described, there is a strong emphasis on interference competition: marine invertebrates are assumed to compete fiercely for the limiting resource of space. Much of our understanding of the dynamics of this system is based on this assumption, yet empirical studies often find that increases in density can reduce performance despite free space being available. Furthermore, the assumption that space is the exclusively limiting resource raises paradoxes regarding species coexistence in this system. Here, we measure the availability of oxygen in the field and in the laboratory, as well as the tolerance of resident species to low-oxygen conditions. We show that oxygen can be the primary limiting resource in some instances, and that exploitative competition for this resource is very likely among benthic marine invertebrates. Furthermore, growth form (and the associated risk of oxygen limitation) covaries with the ability to withstand oxygen-poor conditions across a wide range of taxa. Oxygen availability at very small scales may influence the distribution and abundance of sessile marine invertebrates more than is currently appreciated. Furthermore, competition for multiple resources (space and oxygen) and trade-offs in competitive ability for each may promote coexistence in this system.