Abstract
Dryopteris sukungiana, a new tetraploid fern species of Dryopteridaceae in Yunnan, Southwest China, is described and illustrated. Morphologically, D. sukungiana belongs to the D. sparsa complex but is unique in spreading blackish brown scales on the stipe, glandular hairs and small fibrillose scales on the abaxial surface of the lamina. Phylogenetic analyses show that D. sukungiana is of hybrid origin; its paternal parent is the Sino-Himalayan D. sparsa, and its maternal parent might be an undetected diploid species closely related to the Sino-Japanese D. sparsa.
References
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<p>Sessa, E.B., Zhang, L.B., Väre, H. & Juslén, A. (2015) What we do (and don’t) know about ferns: <em>Dryopteris</em> (Dryopteridaceae) as a case study. <em>Systematic Biology</em> 40 (2): 387–399. https://doi.org/10.1600/036364415X688844</p>
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<p>Zhang, L.B. (2012) Reducing the fern genus <em>Dryopsis</em> to <em>Dryopteris </em>and the systematics and nomenclature of <em>Dryopteris</em> subgenus <em>Erythrovariae</em> section <em>Dryopsis </em>(Dryopteridaceae). <em>Phytotaxa</em> 71 (1): 17–27. https://doi.org/10.11646/PHYTOTAXA.71.1.4</p>
<p>Zhang, L.B. & Zhang, L. (2012) The inclusion of <em>Acrophorus</em>, <em>Diacalpe</em>, <em>Nothoperanema</em>, and <em>Peranema</em> in <em>Dryopteris</em>: The molecular phylogeny, systematics, and nomenclature of <em>Dryopteris</em> subg. <em>Nothoperanema</em> (Dryopteridaceae). <em>Taxon</em> 61 (6): 1199–1216. https://doi.org/10.1002/tax.616003</p>
<p>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 <em>Dryopteris</em> (Dryopteridaceae). <em>BMC Evolutionary Biology</em> 12: 180. https://doi.org/10.1186/1471-2148-12-180</p>
<p>Zuo, Z.Y., Zhao, T., Du, X.Y., Xiong, Y., Lu, J.M. & Li, D.Z. (2021) A revision of <em>Dryopteris</em> sect. <em>Diclisodon</em> (Dryopteridaceae) based on morphological and molecular evidence with description of a new species. <em>Plant diversity</em>. [in press] https://doi.org/10.1016/j.pld.2021.09.005</p>
<p>Boissier, H. (1904) <em>Bulletin de l’Herbier Boissier Ser. 2. v.4</em>, Genève [https://www.biodiversitylibrary.org/bibliography/49730]</p>
<p>Christensen, C. (1905) <em>Index filicum, sive, Enumeratio omnium generum specierumque filicum et Hydropteridum ab anno 1753 ad finem anni 1905 descriptorium: adjectis synonymis principalibus, area geographica, etc.</em> Copenhagen: Hafniae. https://doi.org/10.5962/bhl.title.402</p>
<p>Darnaedi, D. & Iwatsuki, K. (1987) On the structure and systematic position of the fern <em>Dryopteris yakusilvicola</em> Kurata. <em>Journal of the faculty of science, University of Tokyo: Section III: botany </em>14: 121–136.</p>
<p>Darnaedi, D., Kato, M. & Iwatsuki, K. (1989a) Five new or ill-defined species related to <em>Dryopteris sparsa </em>(Dryopteridaceae). <em>The Journal of Japanese Botany</em> 64 (10): 299–310. [http://www.jjbotany.com/pdf/JJB_064_299_310.pdf]</p>
<p>Darnaedi, D., Kato, M. & Iwatsuki, K. (1989b) A cytotaxonomic study of <em>Dryopteris sparsa</em> and closely related species (Dryopteridaceae). <em>The Journal of Japanese Botany </em>64 (11): 330–340. [http://www.jjbotany.com/pdf/JJB_064_330_340.pdf]</p>
<p>Don, D. (1825) <em>Prodromus florae Nepalensis</em>. London. https://doi.org/10.5962/bhl.title.86</p>
<p>Doyle, J.J. & Doyle, J.L. (1987) A rapid DNA isolation procedure for small quantities of fresh leaf tissue. <em>Phytochemical Bulletin</em> 19: 11–15.</p>
<p>Flora Fujianica Compilation Group. (1982) <em>Flora Fujianica</em> <em>Vol. 1: Pteridophyta-Spermatophyta</em>. Fujian Science and Technology Press, Fuzhou.</p>
<p>Fraser-Jenkins, C.R. (1986) A classification of the genus <em>Dryopteris </em>(Pteridophyta: Dryopteridaceae). <em>Bulletin of the British Museum </em>(<em>Natural History</em>)<em> Botany</em> 14: 183–218. [https://www.biodiversitylibrary.org/bibliography/2198]</p>
<p>Fraser-Jenkins, C.R. (2008) <em>Taxonomic revision of three hundred Indian subcontinental pteridophytes with a revised census-list - a new picture of fern-taxonomy and nomenclature in the Indian subcontinent</em>. Dehra Dun: Bishen Singh Mahendra Pal Singh.</p>
<p>Fraser-Jenkins, C.R., Gandhi, K.N. & Kholia, B.S. (2018) <em>An annotated checklist of Indian Pteridophytes II</em>. Dehra Dun: Bishen Singh Mahendra Pal Singh.</p>
<p>Gibby, M. (1985) Cytological observations on Indian subcontinental and Chinese <em>Dryopteris</em> and <em>Polystichum</em> (Pteridophyta: Dryopteridaceae). <em>Bulletin of the British Museum (Natural History) Botany</em> 14: 1–42. [https://www.biodiversitylibrary.org/bibliography/2198]</p>
<p>Hayata, M. (1914) <em>Icones plantarum formosanarum nec non et contributiones ad floram formosanam</em>, Volume: 4. Taipei. https://doi.org/10.5962/bhl.title.1316</p>
<p>Hooker, W.J. & Baker, J.G. (1867) <em>Synopsis filicum</em>. London: R. Hardwicke. https://doi.org/10.5962/bhl.title.30057</p>
<p>Hori, K., Ebihara, A. & Murakami, N. (2021) Origin of the apogamous Japanese fern <em>Dryopteris yakusilvicola</em> (Dryopteridaceae). <em>Taxon</em> 70 (1): 16–26. https://doi.org/10.1002/tax.12416</p>
<p>Itô, H. (1935) Filices Japonenses II. <em>The Botanical Magazine, Tokyo </em>49 (583): 432–437. https://doi.org/10.15281/jplantres1887.49.432</p>
<p>Katoh, K., Misawa, K., Kuma, K. & Miyata, T. (2002) MAFFT: a novel method for rapid multiple sequence alignment based on fast Fourier transform. <em>Nucleic Acids Research</em> 30 (14): 3059–3066. https://doi.org/10.1093/nar/gkf436</p>
<p>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. <em>Bioinformatics</em> 28 (12): 1647–1649. https://doi.org/10.1093/bioinformatics/bts199</p>
<p>Kuntze, O. (1891) <em>Revisio Generum Plantarum: vascularium omnium atque cellularium multarum secundum leges nomeclaturae internationales cum enumeratione plantarum exoticarum in itinere mundi collectarum</em>, v.2. Leipzig. [https://www.biodiversitylibrary.org/bibliography/327]</p>
<p>Kuo, L.Y., Chang, Y.H., Glowienka, J.M., Amoroso, V.B., Dong, S.Y., Kao, T.T., Wang, C.N. & Chiou, W.L. (2016) A revised framework of <em>Dryopteris</em> subg. <em>Nothoperanema</em> (Dryopteridaceae) inferred from phylogenetic evidence, with descriptions of two new sections. <em>Systematic Botany</em> 41(3): 596–605. https://doi.org/10.1600/036364416X692334</p>
<p>Liou, W.T. (2010) <em>A taxonomy study of Dryopteris subgenus Nephrocystis (Dryopteridaceae).</em> Taipei: National Taiwan Normal University.</p>
<p>Nguyen, L.T., Schmidt, H.A., Von Haeseler, A. & Minh, B.Q. (2015) IQ-TREE: a fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. <em>Molecular Biology and Evolution</em> 32 (1): 268–274. https://doi.org/10.1093/molbev/msu300</p>
<p>Ronquist, F., Teslenko, M., van der Mark, P., Ayres, D.L., Darling, A., Höhna, 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. <em>Systematic Biology</em> 61 (3): 539–542. https://doi.org/10.1093/sysbio/sys029</p>
<p>Serizawa, S. (1971) Taxonomical notes on Asian ferns (4). <em>Journal of Japanese Botany</em> 46 (9): 278–282. [http://www.jjbotany.com/pdf/JJB_046_278_282.pdf]</p>
<p>Sessa, E.B., Zhang, L.B., Väre, H. & Juslén, A. (2015) What we do (and don’t) know about ferns: <em>Dryopteris</em> (Dryopteridaceae) as a case study. <em>Systematic Biology</em> 40 (2): 387–399. https://doi.org/10.1600/036364415X688844</p>
<p>Tindale, M.D. & Roy, S.K. (2002) A cytotaxonomic survey of the Pteridophyta of Australia.<em> Australian Systematic Botany</em> 15 (6): 839–937. https://doi.org/10.1071/SB00034</p>
<p>Widén, C.J., Fraser-Jenkins, C.R., Reichstein, T. & Sarvela, J. (2001) A survey of phenolic compounds in <em>Dryopteris</em> and related fern genera. Part III. Phloroglucinol derivatives in subgenera <em>Erythrovaria</em> and <em>Nephrocystis</em> and related genera (Pteridophyta, Dryopteridaceae). <em>Annales Botanici Fennici</em> 38 (2): 99–138. [http://www.sekj.org/PDF/anbf38/anbf38-099p.pdf]</p>
<p>Weng, R.F. (1989) Cytological observation on <em>Dryopteris</em> from East China. <em>In:</em> Shing, K.H. & Kramer, K.U. (Eds.) <em>Proceeds of international symposium on systematic Pteridology</em>. Beijing: China Science and Technology Press, pp. 141–146.</p>
<p>Wu, S.G. & Lu, S.G. (2000) <em>Dryopteris</em>. <em>In: </em>Wu, C.Y. (Ed.) <em>Flora Reipublicae Popularis Sinicae</em>. Vol. 5 (1) (Dryopteridaceae). Beijing: Science Press, pp. 102–220.</p>
<p>Wu, S.G., Xiang, J.Y., Lu, S.G., Wang, F.G., Xing, F.W., Dong, S.Y., He, H., Zhang, L.B., Barrington, D.S. & Christenhusz, M.J.M. (2013) <em>Dryopteris</em>. <em>In: </em>Wu, Z.Y., Raven, P.H., Hong, D.Y. (Eds.) <em>Flora of China</em>. Vol. 2–3 (Pteridophytes). Beijing: Science Press; St. Louis: Missouri Botanical Garden Press, pp. 571–628.</p>
<p>Zhang, L.B. (2012) Reducing the fern genus <em>Dryopsis</em> to <em>Dryopteris </em>and the systematics and nomenclature of <em>Dryopteris</em> subgenus <em>Erythrovariae</em> section <em>Dryopsis </em>(Dryopteridaceae). <em>Phytotaxa</em> 71 (1): 17–27. https://doi.org/10.11646/PHYTOTAXA.71.1.4</p>
<p>Zhang, L.B. & Zhang, L. (2012) The inclusion of <em>Acrophorus</em>, <em>Diacalpe</em>, <em>Nothoperanema</em>, and <em>Peranema</em> in <em>Dryopteris</em>: The molecular phylogeny, systematics, and nomenclature of <em>Dryopteris</em> subg. <em>Nothoperanema</em> (Dryopteridaceae). <em>Taxon</em> 61 (6): 1199–1216. https://doi.org/10.1002/tax.616003</p>
<p>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 <em>Dryopteris</em> (Dryopteridaceae). <em>BMC Evolutionary Biology</em> 12: 180. https://doi.org/10.1186/1471-2148-12-180</p>
<p>Zuo, Z.Y., Zhao, T., Du, X.Y., Xiong, Y., Lu, J.M. & Li, D.Z. (2021) A revision of <em>Dryopteris</em> sect. <em>Diclisodon</em> (Dryopteridaceae) based on morphological and molecular evidence with description of a new species. <em>Plant diversity</em>. [in press] https://doi.org/10.1016/j.pld.2021.09.005</p>