Exotic asphyxiation: interactions between invasive species and hypoxia.

Non-indigenous species (NIS) and hypoxia (<2 mg O 2 l -1 ) can disturb and restructure aquatic communities. Both are heavily influenced by human activities and are intensifying with global change. As these disturbances increase, understanding how they interact to affect native species and systems is essential. To expose patterns, outcomes, and generalizations, we thoroughly reviewed the biological invasion literature and compiled 100 studies that examine the interaction of hypoxia and NIS. We found that 64% of studies showed that NIS are tolerant of hypoxia, and 62% showed that NIS perform better than native species under hypoxia. Only one-quarter of studies examined NIS as creators of hypoxia; thus, NIS are more often considered passengers associated with hypoxia, rather than drivers of it. Paradoxically, the NIS that most commonly create hypoxia are primary producers. Taxa like molluscs are typically more hypoxia tolerant than mobile taxa like fish and crustaceans. Most studies examine individual-level or localized responses to hypoxia; however, the most extensive impacts occur when hypoxia associated with NIS affects communities and ecosystems. We discuss how these influences of hypoxia at higher levels of organization better inform net outcomes of the biological invasion process, i.e. establishment, spread, and impact, and are thus most useful to management. Our review identifies wide variation in the way in which the interaction between hypoxia and NIS is studied in the literature, and suggests ways to address the number of variables that affect their interaction and refine insight gleaned from future studies. We also identify a clear need for resource management to consider the interactive effects of these two global stressors which are almost exclusively managed independently.