Evolutionary patterns of nucleotide substitution rates in plastid genomes of .

Molecular evolution, including nucleotide substitutions, plays an important role in understanding the dynamics and mechanisms of species evolution. Here, we sequenced whole plastid genomes (plastomes) of , , , and and compared them with 14 other plastomes to explore their evolutionary relationships using 67 shared protein-coding sequences. While many previously identified evolutionary relationships were found, our findings do not support previous research which retrieve subg. sect. as a monophyletic group, with sect. found to be polyphyletic and composed of three strongly supported lineages inserted between sections and . Compared with gymnosperms, plastomes showed higher evolutionary rates (/ = 0.3793). Most protein-coding genes experienced relaxed purifying selection, and the high value (0.1927) indicated that gene functions adjusted to environmental changes effectively. Our findings suggest that gene interval regions play an important role in evolution. We detected greater variation in the intergenic regions (, , , , and ), intron losses ( and ), and pseudogene loss and degradation (). Additionally, the loss of some genes suggested the existence of gene exchanges between plastid and nuclear genomes, which affects the evolutionary rate of the former. However, the connective mechanism between these two genomes is still unclear.