J Syst Evol

• Research Article •    

Phylogeny and biogeography of the fern genus Hymenasplenium (Aspleniaceae), with special reference to island speciation

Ke-Wang Xu1, Meng-Dan Deng1, Lin Zhou2, Hui Shang3, Liang Zhang4, Chun-Xiang Li5, Carl J. Rothfels6, David Lorence7, Kenneth R. Wood7, Tom A. Ranker8, Ngan Thi Lu9, 10, Xin-Mao Zhou11, Ralf Knapp12, Zhao-Rong He13, Yue-Hong Yan3, Xin-Fen Gao2 and Li-Bing Zhang2, 14*   

  1. 1Co‑Innovation Center for Sustainable Forestry in Southern China, College of Life Science, Nanjing Forestry University, Nanjing 210037, China
    2CAS Key Laboratory of Mountain Ecological Restoration and Bioresource Utilization, Chengdu Institute of Biology, Chinese Academy of Sciences, P.O. Box 416, Chengdu 610041, China
    3Shanghai Chenshan Plant Science Research Center, Chinese Academy of Sciences; Shanghai Chenshan Botanical Garden, Shanghai 201602, China
    4Key Laboratory for Plant Diversity and Biogeography of East Asia, Kunming Institute of Botany, Chinese Academy of Sciences, Kunming 100049, China
    5State Key Laboratory of Palaeobiology and Stratigraphy of the Nanjing Institute of Geology and Palaeontology, Chinese Academy of Science, Nanjing 210008, China
    6Ecology Center and Department of Biology, Intermountain Herbarium, Utah State University, Logan, UT 84322, USA
    7National Tropical Botanical Garden, 3530 Papalina Road, Kalaheo, HI 96741, USA
    8School of Life Sciences, University of Hawaii at Mānoa, 3190 Maile Way, Room 101, Honolulu, HI 96822, USA
    9University of Chinese Academy of Sciences, Beijing 100049, China
    10Department of Biology, Vietnam National Museum of Nature, Vietnam Academy of Science and Technology, 18th Hoang Quoc Viet Road, Ha Noi 11307, Vietnam
    11Laboratory of Ecology and Evolutionary Biology, State Key Laboratory for Conservation and Utilization of Bio-Resources in Yunnan, Yunnan University, Kunming 650091, China
    12Correspondent of the Muséum national d'Histoire naturelle (MNHN, Paris, France), Steigestrasse 78, Eberbach 69412, Germany
    13Herbarium, Laboratory of Pteridophyta, Institute of Ecology & Geobotany, Yunnan University, Kunming
    14Missouri Botanical Garden, 4344 Shaw Blvd, St. Louis, MO 63110, USA

    *Author for correspondence: Li-Bing Zhang, Libing.Zhang@mobot.org
  • Received:2025-01-08 Accepted:2025-05-08
  • Supported by:
    This research was partially supported from the National Natural Science Foundation of China (32100167) to K.-W.X.

Abstract: Although considerable progress has recently been made in the phylogeny of Hymenasplenium, the genus remains poorly investigated; specifically, the diversification and historical biogeography of the genus have been little studied. Here, we infer an updated plastid DNA phylogeny and the first large-scale nuclear DNA phylogeny to understand the biogeography of the genus. The plastid phylogeny includes 312 accessions from across the genus’ distribution range (ca. 121% increase of the latest sampling), with special attention paid to island accessions from 14 Indian Ocean and Pacific islands, whereas the nuclear phylogeny includes 161 accessions of the Afro-Eurasian species. We identify one new major clade and two new subclades. Reticulate evolution was revealed both among subclades and among species in the Afro-Eurasian. Our divergence-time analyses show that most of the extant species diversity has arisen from diversification after the Oligocene despite a Cretaceous origin of the genus. Ancestral area reconstruction revealed that vicariance likely played a major role in building biogeographic patterns at deep evolutionary scales (the Afro-Eurasian clade and the American clade) in Hymenasplenium, while the intercontinental disjunctions within the Afro-Eurasian clade among Asia, Africa, and Oceania might have resulted from frequent long-distance dispersal events from Asia to Oceania and Africa.

Key words: Asplenium, boreal-tropics, dispersals, fern evolution, LEAFY gene, island biogeography