1. Department of Ecology and Evolutionary Biology, University of Arizona, Biosciences West 310, Tucson, AZ 85721, USA
2. Department of Biology, University of North Carolina, CB no. 3280 Coker Hall, Chapel Hill, NC 27599 USA
3. Laboratoire Ecologie and Evolution, CNRS UMR 7625, Ecole Normale Superieure, 46 rue d'Ulm, 75005 Paris, France
4. Santa Fe Institute, 1399 Hyde Park Road, Santa Fe, NM 87501, USA