1 National Engineering Laboratory for Tree Breeding; Key Laboratory of Genetics and Breeding in Forest Trees and Ornamental Plants, Ministry
of Education; The Tree and Ornamental Plant Breeding and Biotechnology Laboratory of National Forestry and Grassland Administration,
College of Biological Sciences and Technology, Beijing Forestry University, Beijing 100083, China 2 Key Laboratory of Plant Resources Conservation and Sustainable Utilization, South China Botanical Garden, Chinese Academy of Sciences,
Guangzhou 510650, China 3 Key Laboratory of Resource Biology and Biotechnology in Western China (Ministry of Education), College of Life Science, Northwest
University, Xi'an 710069, China 4 Department of Ecology and Environmental Science, Umeå Plant Science Centre, Umeå University, Umeå 901 87, Sweden
†These authors contributed equally to this work. *Author for correspondence. E‐mail: zhao.wei@umu.se