Program for Evolutionary Dynamics, Department of Mathematics and Department of Organismic & Evolutionary Biology, Harvard University, Cambridge, Massachusetts, United States of America.|Program for Evolutionary Dynamics, Department of Mathematics and Department of Organismic & Evolutionary Biology, Harvard University, Cambridge, Massachusetts, United States of America.|Program for Evolutionary Dynamics, Department of Mathematics and Department of Organismic & Evolutionary Biology, Harvard University, Cambridge, Massachusetts, United States of America.|Department of Biomedical Informatics, Columbia University Medical Center, New York, New York, United States of America.|Program for Evolutionary Dynamics, Department of Mathematics and Department of Organismic & Evolutionary Biology, Harvard University, Cambridge, Massachusetts, United States of America.|Program for Evolutionary Dynamics, Department of Mathematics and Department of Organismic & Evolutionary Biology, Harvard University, Cambridge, Massachusetts, United States of America.