1.CSIRO Division of Plant Industry, Canberra, ACT 2601, Australia；Research School of Biology, Australian National University, Canberra, ACT 2601, Australia; 2.CSIRO Division of Plant Industry, Canberra, ACT 2601, Australia；Current address: School of Molecular and Microbial Bioscience, University of Sydney, Sydney NSW 2006, Australia; 3.Research School of Biology, Australian National University, Canberra, ACT 2601, Australia; 4.CSIRO Division of Plant Industry, Canberra, ACT 2601, Australia;