Plant Diversity ›› 2025, Vol. 47 ›› Issue (01): 34-52.DOI: 10.1016/j.pld.2024.07.010

• Articles • Previous Articles    

A revised classification of Dryopteridaceae based on plastome phylogenomics and morphological evidence, with the description of a new genus, Pseudarachniodes

Zheng-Yu Zuoa, Germinal Rouhanb, Shi-Yong Dongc, Hong-Mei Liud, Xin-Yu Dua, Li-Bing Zhange,f, Jin-Mei Lua,g   

  1. a. Germplasm Bank of Wild Species, and Yunnan Key Laboratory of Crop Wild Relatives Omics, Kunming Institute of Botany, Chinese Academy of Sciences, Kunming 650201, Yunnan, China;
    b. Institut de Systématique, Evolution, Biodiversité (ISYEB), Muséum National d'Histoire Naturelle, CNRS, Sorbonne Université, Ecole Pratique des Hautes Etudes, Université des Antilles, CP39, 57 rue Cuvier 75005, Paris, France;
    c. Key Laboratory of Plant Resources Conservation and Sustainable Utilization, South China Botanical Garden, Chinese Academy of Sciences, Guangzhou 510650, Guangdong, China;
    d. Center for Integrative Conservation & Yunnan Key Laboratory for Conservation of Tropical Rainforests and Asian Elephants, Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences, Menglun 666303, Yunnan, China;
    e. Missouri Botanical Garden, 4344 Shaw Blvd, St Louis 63110, MO, USA;
    f. Chengdu Institute of Biology, Chinese Academy of Sciences, Chengdu 610041, Sichuan, China;
    g. State Key Laboratory of Plant Diversity and Specialty Crops, Kunming Institute of Botany, Chinese Academy of Sciences, Kunming 650201, Yunnan, China
  • Received:2023-09-17 Revised:2024-07-24 Published:2025-02-15
  • Contact: Li-Bing Zhang,E-mail:libing.zhang@mobot.org;Jin-Mei Lu,E-mail:lujinmei@mail.kib.ac.cn
  • Supported by:
    The study was funded by the National Natural Science Foundation of China (Grant No. 31970232) and the National Key Basic Research Program of China (Grant No. 2014CB954100).

Abstract: Dryopteridaceae are the largest fern family and include nearly 20% of extant fern diversity, with 24 currently recognized genera. Recognition and delineation of genera within this family have varied greatly. The three-subfamily classification of Dryopteridaceae was based primarily on molecular phylogenetic relationships but lacked morphological evidence, and the phylogenetic relationships of the subfamilies and genera of Dryopteridaceae are only partially resolved. A comprehensive and robust phylogeny is urgently needed. The heterogeneous morphology of the current members of Dryopteridaceae makes the family and its subfamilies difficult to define by single morphological characteristics or even character combinations. We carried out phylogenetic analyses to reconstruct a highly supported phylogeny of Dryopteridaceae. Our analyses recovered 24 strongly supported clades grouped into seven major clades of Dryopteridaceae. Seven morphological characters including habit, rhizome shape, frond morphology, rachis-costae architecture, appendages on stipe base and lamina, and soral arrangement were found to be informative for identifying different major clades and clades in Dryopteridaceae. Based on phylogenetic reconstruction and morphological analysis, we presented an updated infra-familial classification of Dryopteridaceae with seven subfamilies and 24 genera including four newly proposed subfamilies (Ctenitidoideae, Lastreopsidoideae, Pleocnemioideae, and Polystichopsidoideae). Morphological character combinations of each subfamily are summarized, and a key is provided. Most genera sensu PPG I are recognized, with Stigmatopetris reclassified into Dryopteridoideae and Arthrobotrya considered a synonym of Teratophyllum. A new genus Pseudarachniodes is introduced. This revised classification will serve as a foundational framework for future investigations on taxonomy, biogeography, and diversification of the most species-rich Dryopteridaceae in ferns.

Key words: Ctenitidoideae, Infra-familial, Morphological diagnostic characters, Phylogenomics, Rachis-costae architecture, Stigmatopetris