Permissivity of DPP4 Orthologs to MERS-Coronavirus is Governed by Glycosylation and 1 Other Complex Determinants 2 3 Kayla M. Pecka, Trevor Scobeyb, Jesica Swanstromb, Kara L. Jensenb, Christina L. Burcha, Ralph 4 S. Baricb,d,*, and Mark T. Heisec,d,* 5 6 Departments of Biologya, Epidemiologyb, Geneticsc, and Microbiology and Immunologyd, 7 University of North Carolina-Chapel Hill, Chapel Hill, North Carolina. 8 9 Running title: DPP4 orthologs have complex interactions with MERS-CoV 10 11 Keywords: MERS-coronavirus, DPP4, orthologs, host range expansion, animal models 12 13 14 15 *Corresponding Authors: Ralph S. Baric (