Division of Biological Sciences, California Institute of Technology, Pasadena, CA 91125, USA. titus@caltech.edu|Division of Biological Sciences, California Institute of Technology, Pasadena, CA 91125, USA. titus@caltech.edu|Division of Biological Sciences, California Institute of Technology, Pasadena, CA 91125, USA. titus@caltech.edu|Division of Biological Sciences, California Institute of Technology, Pasadena, CA 91125, USA. titus@caltech.edu