[1] Univ Otago, Dept Microbiol & Immunol, Dunedin, New Zealand
[2] Malaghan Inst Med Res, Wellington, New Zealand
[3] Walter & Eliza Hall Inst Med Res, 1G RoyalParade, Parkville, Vic 3052, Australia
[4] Univ Melbourne, Dept Med Biol, Parkville, Australia