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

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 phylo...

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Main Authors: Zheng-Yu Zuo, Germinal Rouhan, Shi-Yong Dong, Hong-Mei Liu, Xin-Yu Du, Li-Bing Zhang, Jin-Mei Lu
Format: Article
Language:English
Published: KeAi Communications Co., Ltd. 2025-01-01
Series:Plant Diversity
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Online Access:http://www.sciencedirect.com/science/article/pii/S2468265924001276
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author Zheng-Yu Zuo
Germinal Rouhan
Shi-Yong Dong
Hong-Mei Liu
Xin-Yu Du
Li-Bing Zhang
Jin-Mei Lu
author_facet Zheng-Yu Zuo
Germinal Rouhan
Shi-Yong Dong
Hong-Mei Liu
Xin-Yu Du
Li-Bing Zhang
Jin-Mei Lu
author_sort Zheng-Yu Zuo
collection DOAJ
description 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.
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spelling doaj-art-528568caf522403184ab148a62bc68952025-02-12T05:31:31ZengKeAi Communications Co., Ltd.Plant Diversity2468-26592025-01-014713452A revised classification of Dryopteridaceae based on plastome phylogenomics and morphological evidence, with the description of a new genus, PseudarachniodesZheng-Yu Zuo0Germinal Rouhan1Shi-Yong Dong2Hong-Mei Liu3Xin-Yu Du4Li-Bing Zhang5Jin-Mei Lu6Germplasm Bank of Wild Species, and Yunnan Key Laboratory of Crop Wild Relatives Omics, Kunming Institute of Botany, Chinese Academy of Sciences, Kunming 650201, Yunnan, ChinaInstitut 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, FranceKey Laboratory of Plant Resources Conservation and Sustainable Utilization, South China Botanical Garden, Chinese Academy of Sciences, Guangzhou 510650, Guangdong, ChinaCenter 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, ChinaGermplasm Bank of Wild Species, and Yunnan Key Laboratory of Crop Wild Relatives Omics, Kunming Institute of Botany, Chinese Academy of Sciences, Kunming 650201, Yunnan, ChinaMissouri Botanical Garden, 4344 Shaw Blvd, St Louis 63110, MO, USA; Chengdu Institute of Biology, Chinese Academy of Sciences, Chengdu 610041, Sichuan, China; Corresponding author. Missouri Botanical Garden, 4344 Shaw Blvd, St Louis, MO, 63110, USA.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; State Key Laboratory of Plant Diversity and Specialty Crops, Kunming Institute of Botany, Chinese Academy of Sciences, Kunming 650201, Yunnan, China; Corresponding author. Germplasm Bank of Wild Species, and Yunnan Key Laboratory of Crop Wild Relatives Omics, Kunming Institute of Botany, Chinese Academy of Sciences, Kunming, Yunnan 650201, China.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.http://www.sciencedirect.com/science/article/pii/S2468265924001276CtenitidoideaeInfra-familialMorphological diagnostic charactersPhylogenomicsRachis-costae architectureStigmatopetris
spellingShingle Zheng-Yu Zuo
Germinal Rouhan
Shi-Yong Dong
Hong-Mei Liu
Xin-Yu Du
Li-Bing Zhang
Jin-Mei Lu
A revised classification of Dryopteridaceae based on plastome phylogenomics and morphological evidence, with the description of a new genus, Pseudarachniodes
Plant Diversity
Ctenitidoideae
Infra-familial
Morphological diagnostic characters
Phylogenomics
Rachis-costae architecture
Stigmatopetris
title A revised classification of Dryopteridaceae based on plastome phylogenomics and morphological evidence, with the description of a new genus, Pseudarachniodes
title_full A revised classification of Dryopteridaceae based on plastome phylogenomics and morphological evidence, with the description of a new genus, Pseudarachniodes
title_fullStr A revised classification of Dryopteridaceae based on plastome phylogenomics and morphological evidence, with the description of a new genus, Pseudarachniodes
title_full_unstemmed A revised classification of Dryopteridaceae based on plastome phylogenomics and morphological evidence, with the description of a new genus, Pseudarachniodes
title_short A revised classification of Dryopteridaceae based on plastome phylogenomics and morphological evidence, with the description of a new genus, Pseudarachniodes
title_sort revised classification of dryopteridaceae based on plastome phylogenomics and morphological evidence with the description of a new genus pseudarachniodes
topic Ctenitidoideae
Infra-familial
Morphological diagnostic characters
Phylogenomics
Rachis-costae architecture
Stigmatopetris
url http://www.sciencedirect.com/science/article/pii/S2468265924001276
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