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Univ Tennessee, Natl Inst Math & Biol Synth, Ctr Dynam Social Complex, Dept Ecol & Evolutionary Biol,Dept Math, Knoxville, TN 37996 USA Univ Tennessee, Natl Inst Math & Biol Synth, Ctr Dynam Social Complex, Dept Ecol & Evolutionary Biol,Dept Math, Knoxville, TN 37996 USA