Genetic studies of Domesticated-Weed-Wild Complexes (DWWC) have typically focused on one-way introgression of crop alleles into wild or weedy populations, with little consideration of the entire natural ecosystem. In Sri Lanka, DWWC is diverse, comprising six evolutionarily discrete groups in the genus Oryza. Using 33 neutral simple sequence repeat (SSR) loci, we characterized six Oryza groups to understand the genetic background and evolution of DWWC components. Our analysis found that Oryza groups have large population sizes and high inter-group long-term gene flows. Asymmetric gene flows were found between wild and weedy rice groups, but the rare alleles shared among DWWC components provide additional evidence for extensive and enduring exchange, highlighting the dynamic nature of this complex genetic admixture among different Oryza lineages. We found high genetic diversity at the population and species levels due to mixed DWWC components over the generations. Weedy rice types exhibit genetic incorporation through admixture from both crop and wild species, highlighting the multi-way genetic transfer in the evolution of weedy rice types. Our findings support the idea that the DWWC is an integrated complex in the Sri Lankan rice ecosystem and that its weedy rice has multiple origins, including de-domestication via feralization of cultivated rice, inter-varietal hybridization among distinct cultivated rice types, adaptation, and invasion of rice cultivation areas by wild Oryza species, and hybridization events between crop and wild rice populations. Abandoned rice domesticates can also evolve into weedy forms with less intimate human relationships and contaminate the rice ecosystem.

Ecological divergence due to habitat difference plays a prominent role in the formation of new species but the genetic architecture during ecological speciation and the mechanism underlying phenotypic divergence remain less understood. Two wild rice species (O. rufipogon and O. nivara) are a progenitor-derivative species pair with ecological divergence and provide a unique system for studying ecological adaptation/speciation. Here, we constructed a high-resolved linkage map and conducted a quantitative trait locus (QTL) analysis of 19 phenotypic traits using an F2 population generated from a cross between the two wild rice species. We identified 113 QTLs associated with interspecific divergence of 16 quantitative traits, with effect sizes ranging from 1.61% to 34.1% in terms of the percentage of variation explained (PVE). The distribution of effect sizes of QTLs followed a negative exponential, suggesting that a few genes of large effect and many genes of small effect were responsible for the phenotypic divergence. We observed 18 clusters of QTLs (QTL hotspots) on 11 chromosomes, significantly more than that expected by chance, demonstrating the importance of coinheritance of loci/genes in ecological adaptation/speciation. Analysis of effect direction and v-test statistics revealed that interspecific differentiation of most traits was driven by divergent natural selection, supporting the argument that ecological adaptation/speciation would proceed rapidly under coordinated selection on multiple traits. Our findings provide new insights into the understanding of genetic architecture of ecological adaptation and speciation in plants and helps effective manipulation of specific genes or gene cluster in rice breeding.