[1] Smithsonian Conservat Biol Inst, Conservat Ecol Ctr, Front Royal, VA 22630 USA
[2] Smithsonian Conservat Biol Inst, Ctr Conservat Genom, Washington, DC USA
[3] Natl Ecol Observ Network, Front Royal, VA USA
[4] Univ Illinois, Dept Nat Resources & Environm Sci, Urbana, IL 61801 USA