Open Access via the Springer Compact Agreement. This study was funded by IBioIC PhD studentship (LW), Leverhulme Trust Research Project (HD and MHT, project No. RPG-2014-418), The Elphinstone Scholarship of University of Aberdeen (QF), Leverhulme Trust-Royal Society Africa award (KK and HD, AA090088) and the jointly funded UK Medical Research Council – UK Department for International Development (MRC/DFID) Concordat agreement African Research Leaders Award (KK and HD, MR/S00520X/1), Biotechnology and Biological Sciences Research Council UK (HD and SW, BB/P00380X/1) and National Natural Science Foundation of China (31,570,033, 31,811,530,299, and 31,870,035 to YY), and the Royal Society-NSFC Newton Mobility Grant Award (IEC\NSFC\170,617 to HD and YY).Peer reviewedPublisher PD
