[1] Harvard Univ, Program Evolutionary Dynam, Cambridge, MA 02138 USA
[2] Harvard Univ, Dept Math, Cambridge, MA 02138 USA
[3] Harvard Univ, Dept Organism & Evolutionary Biol, Cambridge, MA 02138 USA
[4] Emmanuel Coll, Dept Math, Boston, MA 02115 USA
[5] Harvard Univ, Museum Comparat Zool, Cambridge, MA 02138 USA