a, Principal coordinate analysis of 58 citrus accessions based on pairwise nuclear genome distances and metric multidimensional scaling. The first two axes separate the three main citrus groups (citrons, pummelos and mandarins) with interspecific hybrids (oranges, grapefruit, lemon and limes) situated at intermediate positions relative to their parental genotypes. b, Violin plots of the heterozygosity distribution in 58 citrus accessions, representing 10 taxonomic groups as well as 2 related genera, Poncirus (Poncirus trifoliata, also known as Citrus trifoliata) and Chinese box orange (Severinia). White dot, median; bar limits, upper and lower quartiles; whiskers, 1.5× interquartile range. The bimodal separation of intraspecies (light blue) and interspecies (light pink) genetic diversity is manifested among the admixed mandarins and across different genotypes including interspecific hybrids. Three-letter codes are listed in parenthesis with additional descriptions in Supplementary Table 2. c, Chronogram of citrus speciation. Two distinct and temporally well-separated phases of species radiation are apparent, with the southeast Asian citrus radiation followed by the Australian citrus diversification. Age calibration is based on the citrus fossil C. linczangensis16 from the Late Miocene (denoted by a filled red circle). The 95% confidence intervals are derived from 200 bootstraps. Bayesian posterior probability is 1.0 for all nodes. d, Proposed origin of citrus and ancient dispersal routes. Arrows suggest plausible migration directions of the ancestral citrus species from the centre of origin—the triangle formed by northeastern India, northern Myanmar and northwestern Yunnan. The proposal is compatible with citrus biogeography, phylogenetic relationships, the inferred timing of diversification and the paleogeography of the region, especially the geological history of Wallacea and Japan. The red star marks the fossil location of C. linczangensis. Citrus fruit images in c and d are not drawn to scale.