Abstract
A new fern species, Dryopteris calciphila (Dryopteridaceae), is described from the karst limestone regions of Tuyen Quang Province in northern Vietnam. Morphologically, the new species is placed in Dryopteris subg. Erythrovariae and most closely resembles D. tahmingensis in having dense black scales and pinnules or lobes with obtuse apices. However, it differs from D. tahmingensis in having thickly papery lamina and nearly entire pinnule or lobe margins, whereas D. tahmingensis has herbaceous lamina and toothed margins. Phylogenetic analyses based on five plastid markers (rbcL, trnL, trnL–F, rps4, and rps4–trnS) confirm its placement within Dryopteris sect. Erythrovariae and recover it as a distinct lineage within the section. A detailed taxonomic description, color plate, and information on its morphology, habitat, and distribution are provided. Given its narrow distribution and small observed population size, the species is provisionally assessed as Endangered (EN) according to the IUCN Red List Categories and Criteria. This discovery highlights the high level of endemism in the limestone ecosystems of northern Vietnam and further enriches the known diversity of Dryopteris in the region.
References
- Adanson, M. (1763) Familles des Plantes, vol. 2. Chez Vincent, Paris, 640 pp.
- Averyanov, L., Nguyen, K.S., Tran, T., Averyanova, A., Maisak, T. & Nguyen, H. (2020) Plant diversity, flora and vegetation of Bat Dai Son Mountain area, northern Vietnam. Strata, Saint Petersburg, 561 pp.
- Blume, C.L. (1828) Enumeratio Plantarum Javae et insularum adjacentium: minus cognitarum vel novarum ex herbariis Reinwardtii, Kuhlii, Hasseltii et Blumii. Fasc. 2. Apud J.W. van Leeuwen, Lugduni Batavorum, pp. 99–274.
- Chen, C.-W., Hsu, T.-C., Doan, H.S., Do, V.H., Luu, H.T., Le, V.S., Liu, Y.-C., Li, C.-W., Huang, Y.-M. & Chung, K.-F. (2023) Studies of Vietnamese pteridophyte flora 2. Systematic Botany 48: 159–172. https://doi.org/10.1600/036364423x16847773873161
- Ching, R.-C. (1938) A revision of the Chinese and Sikkim-Himalayan Dryopteris with reference to some species from neighbouring regions (Part 10). Bulletin of the Fan Memorial Institute of Biology, Botany 8: 363–507.
- Christensen, C. (1906) Index Filicum: sive enumeratio omnium generum specierumque Filicum et Hydropteridum. H. Hagerup, Copenhagen, 744 pp. https://doi.org/10.5962/bhl.title.402
- Christensen, C. (1917) Index Filicum, Supplément préliminaire, pour les années, 1913, 1914, 1915, 1916. Typis Triers Bogtrykkeri, Copenhagen, 60 pp.
- Christensen, C. (1937) Taxonomic fern-studies. Dansk Botanisk Arkiv 9: 1–67.
- Ebihara, A. (2016) The Standard of Ferns and Lycophytes in Japan Ⅰ. Gakken Plus, Tokyo, 475 pp.
- Fraser-Jenkins, C.R. (1986) A classification of the genus Dryopteris (Pteridophyta: Dryopteridaceae). Bulletin of the British Museum (Natural History), Botany 14: 183–218.
- Hassler, M. (1994–2026) World Ferns. Synonymic checklist and distribution of ferns and lycophytes of the World. Version 26.02. Available from: https://www.worldplants.de/world-ferns/ferns-and-lycophytes-list/ (accessed 27 February 2026)
- Hooker, W.J. (1853) Notice of two new ferns from China. Hooker’s Journal of Botany and Kew Garden Miscellany 5: 236–238.
- Huang, J.-K., Li, X.-J., Zhou, L.-X., Zuo, Z.-Y. & Dong, S.-Y. (2025) Dryopteris pycnolepis, a new fern species from northern Vietnam, with miscellaneous notes on Dryopteris sect. Splendentes (Dryopteridaceae). PhytoKeys 263: 1–15. https://doi.org/10.3897/phytokeys.263.163351
- Itô, H. (1936) Filices Japonenses. III. Shokubutsugaku Zasshi 50: 32–39.
- 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
- Kuntze, O. (1891) Revisio generum plantarum vascularium omnium atque cellularium multarum secundum Leges nomenclaturae internationales cum enumeratione plantarum exoticarum in itinere mundi collectarum, Pars II. A. Felix, Leipzig, pp. 375–1011. https://doi.org/10.5962/bhl.title.124064
- Li, C.-X. & Lu, S.-G. (2006) Phylogenetics of Chinese Dryopteris (Dryopteridaceae) based on the chloroplast rps4-trnS sequence data. Journal of Plant Research 119: 589–598. https://doi.org/10.1007/s10265-006-0003-x
- Lu, N.T., Zhang, L. & Zhang, L.-B. (2014) Polystichum hagiangense (subg. Haplopolystichum; Dryopteridaceae), a new fern species from Ha Giang Province, northern Vietnam. Phytotaxa 175: 293–297. https://doi.org/10.11646/phytotaxa.175.5.9
- Lu, N.T., Zhang, L., Nguyen, D.T. & Zhang, L.-B. (2016) Polystichum quangbinhense sp. nov. (subg. Haplopolystichum; Dryopteridaceae), the southernmost cave species of Polystichum from central Vietnam. Phytotaxa 283: 295–299. https://doi.org/10.11646/phytotaxa.283.3.6
- Lu, N.T., Nguyen, L.P., Le, C.T., Nguyen, T.T. & Zhang, L. (2021) Arachniodes libingii sp. nov. and A. vietnamensis sp. nov. (Dryopteridaceae), two new ferns from southern China and central Vietnam. Phytotaxa 522: 256–262. https://doi.org/10.11646/phytotaxa.522.3.9
- Lu, N.T., Wei, H.-J., Vuong, L.D., Toan, L.C., Zhang, L.-B. & Zhang, L. (2023a) Cyrtomium calcis sp. nov. and six new records of the shield fern family (Dryopteridaceae) from Vietnam. Phytotaxa 583: 15–26. https://doi.org/10.11646/phytotaxa.583.1.2
- Lu, N.T., Zhang, L., Thanh, N.T., Toan, L.C. & Zhang, L.-B. (2023b) Three new species and one new record of the fern genus Polystichum (Dryopteridaceae) from Vietnam. Phytotaxa 607: 85–94. https://doi.org/10.11646/phytotaxa.607.1.7
- Phan, K. (2010) The updated checklist of the fern flora of Vietnam following the classification scheme of A. Smith et al. (2006). Journal of Fairylake Botanical Garden 9: 1–13.
- Presl, C. (1836) Tentamen Pteridographiae. Typis Filiorum Theophili Haase, Pragae, 290 pp.
- Roth, A.W. (1800) Tentamen Florae Germanicae, vol. 3. Gleditsch, Leipzig, 578 pp.
- Stamatakis, A., Hoover, P. & Rougemont, J. (2008) A rapid bootstrap algorithm for the RAxML Web servers. Systematic Biology 57: 758–771. https://doi.org/10.1080/10635150802429642
- Wei, H.-J., Chen, B. & Zuo, Z.-Y. (2024) Two new Dryopteris taxa originated from distantly related congeners in China. Taxon 73: 124–141. https://doi.org/10.1002/tax.13116
- Wu, S.-G., Phan, K.L. & Xiang, J.-Y. (2005) A new genus and two new species of ferns from Vietnam. Novon 15: 245–249.
- Zhang, L.-B., Zhang, L., Dong, S.-Y., Sessa, E.B., Gao, X.-F. & Ebihara, A. (2012) Molecular circumscription and major evolutionary lineages of the fern genus Dryopteris (Dryopteridaceae). BMC Evolutionary Biology 12: 180. https://doi.org/10.1186/1471-2148-12-180
- Zhang, L.-B., Wu, S.-G., Xiang, J.-Y., Xing, F.-W., He, H., Wang, F.-G., Lu, S.-G., Dong, S.-Y., Barrington, D.S., Iwatsuki, K., Christenhusz, M.J.M., Mickel, J.T., Kato, M. & Gilbert, M.G. (2013) Dryopteridaceae. In: Wu, Z.-Y., Raven, P.H. & Hong, D.-Y. (Eds.) Flora of China, vols. 2–3. Science Press, Beijing & Missouri Botanical Garden Press, Saint Louis, pp. 541–724.
