1. Department of Biological Sciences National University of Singapore 14 Science Drive 4 Singapore 117558 Singapore
2. UMR5174 EDB (Laboratoire Evolution et Diversité Biologique) CNRS, IRD, UPS, Université Paul Sabatier 118 route de Narbonne Toulouse F‐31062 France
3. Department of Ecology and Evolutionary Biology The University of Tennessee Knoxville TN 37996 U.S.A.