Precipitation and local environment shape the geographic variation of seed size across natural populations of sand rice (Agriophyllum squarrosum)

Sand rice (Agriophyllum squarrosum) is widely distributed on dunes in the Asian interior arid zone, and its large intraspecific trait variation makes it a very good model for investigating the ecological processes underlying its adaptation to the desert environment. In this study, seed size variation across 68 natural populations was used to establish geographic patterns and to quantify the effects of the climate, soil, and collection-year weather variables. The length of the seed major axis and thousand seed weight (TSW) both showed significant longitudinal patterns. Long-term climate variables accounted for most of the explained variances for seed major axis (57.20%) and TSW (91.54%). Specifically, annual precipitation and minimum monthly precipitation had the most significantly positive and negative effects, indicating that longitudinal clines are driven by a precipitation gradient across the species' distribution range. A substantial unique effect of soil variables (27.27%) was found for seed major axis variation, but only 3.64% of TSW variation was explained by soil variables. Two extreme groups were selected to evaluate the genetic and plastic effects on seed size in a common garden experiment. Large-seeded individuals were more competitive in semi-arid regions, and had stronger adaptive plasticity as well as better performance in early seedling establishment, and hence they have potential for use in future domestication projects. Precipitation underlies the longitudinal cline of seed size across the distribution range of sand rice, and large-seeded individuals have strong adaptive plasticity that has potential for use in future domestication.