Department of Biology, University of Pennsylvania, Philadelphia, Pennsylvania, United States of America. alstew@sas.upenn.edu|Department of Biology, University of Pennsylvania, Philadelphia, Pennsylvania, United States of America. alstew@sas.upenn.edu|Department of Biology, University of Pennsylvania, Philadelphia, Pennsylvania, United States of America. alstew@sas.upenn.edu|Department of Biology, University of Pennsylvania, Philadelphia, Pennsylvania, United States of America. alstew@sas.upenn.edu