Abstract
Vandenboschia lofoushanensis has been a dubious species since it was first described from Guangdong, southern China, in 1959, due to the lack of additional material beyond its type specimens. This species was recently rediscovered at its type locality and a nearby site. Our field observations, morphological comparisons, and phylogenetic analyses of plastid sequences demonstrate that V. lofoushanensis is conspecific with V. kalamocarpa, a species widely distributed in East Asia. Accordingly, V. lofoushanensis is formally treated as a synonym of V. kalamocarpa in this study. This case study indicates that the flatness or crispation of the laminae should be used cautiously as a character for species delimitation within Vandenboschia and other filmy ferns.
References
- Allan, H.H.B. (1961) Hymenophyllaceae. In: Allan, H.H.B. (Ed.) Flora of New Zealand Vol I Indigenous Tracheophyta: Psilopsida, Lycopsida, Filicopsida, Gymnospermae, Dicotyledones. Government Printer, Wellington, pp. 22–37.
- Beddome, R.H. (1876) Supplement to the Ferns of Southern India and British India. Gantz Brothers, Madras, 28 pp.
- Blume, C.L. (1828) Enumeratio Plantarum Javae et Insularum Adjacentium 2. J.W. van Leeuwen, Leiden, pp. 99–274.
- Bosch, R.B. van den (1859) Synopsis Hymenophyllacearum. Nederlandsch Kruidkundig Archief 4: 341–419.
- Ching, R.C. & Wang, C.H. (1959) Materials for the Pteridophytic Flora of Hainan. Acta Phytotaxonomica Sinica 8: 125–171.
- Ching, R.C. (1959) Hymenophyllaceae. In: Ching, R.C. (Ed.) Flora Republica Popularis Sinicae, vol 2. Science Press, Beijing, pp. 132–196, 350–357.
- Christenhusz, M.J.M., Fay, M.F. & Byng, H.W. (2018) The Global Flora: A practical flora to vascular plant species of the world, vol. 4: GLOVAP nomenclature part 1. Plant Gateway Ltd., Bradford, 155 pp.
- Copeland, E.B. (1933) Trichomanes. Philippine Journal of Science 51: 119–280.
- Copeland, E.B. (1938) Genera Hymenophyllacearum. Philippine Journal of Science 67: 1–110.
- Don, D. (1825) Prodromus Florae Nepalensis. J. Gale, London, 256 pp.
- Dubuisson, J.Y., Boucheron-Dubuisson, E., Le Péchon, T., Bausero, P., Droissart, V., Deblauwe, V., Ebihara, A., Hennequin, S. & Rouhan, G. (2022) Diversity and taxonomy of the fern genus Vandenboschia Copel. (Hymenophyllaceae, Polypodiidae) in the Afro-Malagasy region and description of a new species. Botany Letters 169: 268–283. https://doi.org/10.1080/23818107.2022.2062445
- Ebihara, A., Dubuisson, J.Y., Iwatsuki, K., Hennequin, S. & Ito, M. (2006) A taxonomic revision of Hymenophyllaceae. Blumea 51: 221–280. https://doi.org/10.3767/000651906X622210
- Ebihara, A., Ishikawa, H., Matsumoto, S., Lin, S.J., Iwatsuki, K., Takamiya, N., Watano, Y. & Ito, M. (2005) Nuclear DNA, chloroplast DNA, and ploidy analysis clarified biological complexity of the Vandenboschia radicans complex (Hymenophyllaceae) in Japan and adjacent areas. American Journal of Botany 92: 1535–1547. https://doi.org/10.3732/ajb.92.9.1535
- Ebihara, A., Matsumoto, S. & Ito, M. (2009) Taxonomy of the reticulate Vandenboschia radicans Complex (Hymenophyllaceae) in Japan. Acta Phytotaxonomica et Geobotanica 60: 26–40. https://doi.org/10.18942/apg.kj00005576220
- Fraser-Jenkins, C.R., Gandhi, K.N., Kholia, B.S. & Benniami, A. (2017) An Annotated Checklist of Indian Pteridophytes, part 1. Bishen Singh Mahendra Pal Singh Dehra Dun, Dehra Dun, 562 pp.
- Hassler, M. (1994–2026) World Plants: Synonymic Checklist and Distribution of the World Flora, version 23.02. Available from: http://www.worldplants.de (accessed 13 April 2026)
- Hayata, B. (1915) Icones Plantarum Formosanarum nec non et Contributiones ad Floram Formosanam 5. Taihoku Bureau of Productive Industry, Taihoku, 358 pp.
- Huang, J.-K., Li, X.-J. & Dong, S.-Y. (2025) A checklist survey on the species of lycophytes and ferns in Huidong County of Guangdong, China. Subtropical Plant Science 54: 186–194. https://doi.org/10.3969/j.issn.1009-7791.2025.02.008
- Iwatsuki, K. (1958) Taxonomic studies of Pteridophyta I. Acta Phytotaxonomica et Geobotanica 17: 65–72. https://doi.org/10.18942/bunruichiri.KJ00001077782
- Iwatsuki, K. (1985) The Hymenophyllaceae of Asia, excluding Malesia. Journal of the Faculty of Science, University of Tokyo, Section 3, Botany 13: 501–551.
- Iwatsuki, K. (1995) Hymenophyllaceae. In: Iwatsuki, K., Kato, M. & Yamazaki, T. (Eds.) Flora of Japan, vol 1. Kodansha Ltd., Tokyo, pp. 41–53.
- Iwatsuki, K. & Ebihara, A. (2023) Flora Malesiana, Ser. II, vol 5. Hymenophyllaceae. Singapore Botanic Gardens, Singapore, 161 pp.
- Jin, J.J., Yu, W.B., Yang, J.B., Song, Y., dePamphilis, C.W., Yi, T.S. & Li, D.Z. (2020) GetOrganelle: a fast and versatile toolkit for accurate de novo assembly of organelle genomes. Genome Biology 21: 241. https://doi.org/10.1186/s13059-020-02154-5
- Katoh, K. & Standley, D.M. (2013) MAFFT Multiple Sequence Alignment Software, version 7: Improvements in Performance and Usability. Molecular Biology and Evolution 30: 772–780. https://doi.org/10.1093/molbev/mst010
- Kearse, M., Moir, R., Wilson, A., Stones-Havas, S., Cheung, M., Sturrock, S., Buxton, S., Cooper, A., Markowitz, S., Duran, C., Thierer, T., Ashton, B., Meintjes, P. & Drummond, A. (2012) Geneious Basic: An integrated and extendable desktop software platform for the organization and analysis of sequence data. Bioinformatics 28: 1647–1649. https://doi.org/10.1093/bioinformatics/bts199
- Lefort, V., Longueville, J.E. & Gascuel, O. (2017) SMS: Smart Model Selection in PhyML. Molecular Biology and Evolution 34: 2422–2424. https://doi.org/10.1093/molbev/msx149
- Liu, J.X., Zhang, Q.Y., Atsushi, E. & Kunio, I. (2013) Hymenophyllaceae. In: Wu, Z.Y., Raven, P.H. & Hong, D.Y. (Eds.) Flora of China, vol 2–3. Science Press, Beijing; Missouri Botanical Garden Press, St. Louis, pp. 93–109.
- Miller, M.A., Pfeiffer, W. & Schwartz, T. (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic trees. Proceedings of the Gateway Computing Environments Workshop (GCE). Gateway Computing, New Orleans, pp. 1–8. https://doi.org/10.1109/GCE.2010.5676129
- Morton, C.V. (1968) The genera, subgenera, and sections of the Hymenophyllaceae. Contributions from the United States National Herbarium 38: 153–214.
- Nakai, T. (1926) Notes on Japanese Ferns III. Botanical Magazine, Tokyo 40: 239–275. https://doi.org/10.15281/jplantres1887.40.239
- Park, S.H., Kim, J.S. & Kim, H.T. (2025) Hybridization and diversity of the genus Vandenboschia in Korea insights from morphological, cytological, and genotype analyses. Scientific Reports 15: 1619. https://doi.org/10.1038/s41598-025-86000-3
- Ronquist, F., Teslenko, M., van der Mark, P., Ayres, D.L., Darling, A., Hohna, S., Larget, B., Liu, L., Suchard, M.A. & Huelsenbeck, J.P. (2012) MrBayes 3.2: Efficient Bayesian Phylogenetic Inference and Model Choice Across a Large Model Space. Systematic Biology 61: 539–542. https://doi.org/10.1093/sysbio/sys029
- Stamatakis, A. (2014) RAxML version 8: a tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics 30: 1312–1313. https://doi.org/10.1093/bioinformatics/btu033
- Swartz, O. (1801) Genera et Species Filicum. Journal für die Botanik 1800: 1–120.
- Swofford, D.L. (2002) PAUP*: Phylogenetic Analysis Using Parsimony (*and other methods), version 4.0b10. Sinauer Associates, Sunderland, Massachusetts.
- Tagawa, M. (1951) Fern Miscellany (5). Journal of Japanese Botany 26: 185–188.
- Wu, S.H. (2006) Hymenophyllaceae. In: Wu, T.L. (Ed.) Flora of Guangdong, vol 7. Guangdong Science and Technology Press, Guangzhou, pp. 55–71.
- Yabe, Y. (1905) Trichmanes Formosense et Loochooense. Botanical Magazine 19: 31–38. https://doi.org/10.15281/jplantres1887.19.31
- Zhang, D., Gao, F., Jakovlic, I., Zou, H., Zhang, J., Li, W.X. & Wang, G.T. (2020) PhyloSuite: An integrated and scalable desktop platform for streamlined molecular sequence data management and evolutionary phylogenetics studies. Molecular Ecology Resources 20: 348–355. https://doi.org/10.1111/1755-0998.13096
