24Both direct repeats (DR) and inverted repeats (IR) are documented in the published 25 plastomes of four Selaginella species indicating the unusual and diverse plastome 26 structure in the family Selaginellaceae. In this study, we newly sequenced complete 27 plastomes of seven species from five main lineages of Selaginellaceae and also re-28 sequenced three species (S. tamariscina, S. uncinata and S. moellendorffii) to explore 29 the evolutionary trajectory of Selaginellaceae plastomes. Our results showed that the 30 plastomes of Selaginellaceae vary remarkably in size, gene contents, gene order and 31 GC contents. Notably, both DR and IR structure existed in the plastomes of 32 Selaginellaceae with DR structure being an early diverged character. The occurrence 33 of DR structure was right after the Permian-Triassic (P-T) extinction (ca. 246 Ma) and 34 remained in most subgenera of Selaginellaceae, whereas IR structure only reoccurred 35 in the most derived subg. Heterostachys (ca. 23 Ma). The presence of a pair of large 36 repeats psbK-trnQ, together with DR/IR region in S. bisulcata, S. pennata, S. uncinata, 37 and S. hainanensis, could frequently mediate diverse homologous recombination and 38 create approximately equal stoichiometric isomers (IR/DR-coexisting) and 39 subgenomes. High proportion of repeats is presumably responsible for the dynamic 40 IR/DR-coexisting plastomes, which possess a lower synonymous substitution rate (dS) 41 compared with DR-possessing plastomes. We propose that the occurrence of DR 42 structure, together with few repeats, is possibly selected to adapt to the 43 environmental upheaval during the P-T crisis and the IR/DR-coexisting plastomes also 44 reached an equilibrium in plastome organization through highly efficient homologous 45 recombination to maintain stability. 46 47 65 conservation in structures and gene order (Mower and Vickrey 2018). A 30 kb inversion 66 (ycf2-psbM) was detected in the large single copy (LSC) of ferns and seed plants 67 plastomes relative to bryophytes and lycophytes, which is a strong evidence showing 68 lycophytes are located at the basal position of vascular plants (Raubeson and Jansen 69 1992). The plastomes of ferns underwent two hypothetical inversions (CE inversion 70 (trnC to trnE) and DE inversion (trnD-trnY)) within rpoB-psbZ (BZ) region from the 71 ancestral gene order in eusporangiates to the derived gene order in core 72