[1] Univ Washington, Dept Biol, Seattle, WA 98195 USA
[2] Univ Washington, BEACON Ctr Study Evolut Act, Seattle, WA 98195 USA
[3] Eawag, Dept Environm Microbiol, CH-8600 Dubendorf, Switzerland
[4] ETH, Dept Environm Syst Sci, CH-8092 Zurich, Switzerland