This work was supported by the National Science Foundation (Dr. Arnosti ,Dr. Jeffrey), EPA Graduate Fellowship (Dr. Steen), and the Deutsche Forschungsgemeinschaft Postdoctoral Fellowship (Dr. Ziervogel). The authors are also grateful to the Max-Planck Institute for Marine Microbiology (Bremen) for supporting the work on Svalbard. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.