“…A mixed linear model (MLM) method (Yu et al, 2006) was proposed to control for population structure and the imbalanced kinships among various individuals (Pritchard et al, 2000). Until recently, determining the molecular basis of heritable trait variation has been challenging in conifer species, and genome-wide association studies (GWAS) have been limited to a few species and traits (Baison et al, 2020;Chen et al, 2021;De La Torre et al, 2021a;Elfstrand et al, 2020;Lu et al, 2017;Weiss et al, 2020). For example, association studies of drought tolerance have only been performed with pre-selected candidate genes (Cumbie et al, 2011;Depardieu et al, 2021;Eckert et al, 2010;Gonzalez-Martinez et al, 2008;Trujillo-Moya et al, 2018) and no large-scale, genome-wide studies have been reported to date.…”