Research Article |
Corresponding author: Yue-Hong Yan ( yhyan@sibs.ac.cn ) Corresponding author: Rui-Jiang Wang ( wangrj@scbg.ac.cn ) Academic editor: Blanca León
© 2021 Jiang-Ping Shu, Zi-Yue Liu, Zhi-Rong Gu, Li-Jun Chen, Hong-Jin Wei, Xi-Le Zhou, Yue-Hong Yan, Rui-Jiang Wang.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Shu J-P, Liu Z-Y, Gu Z-R, Chen L-J, Wei H-J, Zhou X-L, Yan Y-H, Wang R-J (2021) Dryopteris wulingshanensis (Dryopteridaceae), a new species from Hunan, China. PhytoKeys 185: 17-26. https://doi.org/10.3897/phytokeys.185.72019
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Dryopteris wulingshanensis, a new species growing on limestone in the Wulingshan Mountains, Hunan, China, is described and illustrated. This species is most similar to D. jishouensis and D. gymnophylla on general morphological traits, such as the form of scales, rhizome and sori, but differs by the number of vascular bundles at the base of the petiole, length to width ratio of lamina, stalk length of basal pinnae, division of the lamina, apex form of the pinnule and habitat. Moreover, molecular phylogenetic analysis using the chloroplast rbcL gene suggested that D. wulingshanensis, as the sister group of D. jishouensis, is a monophyletic clade. According to its restricted geographic range, small populations and few individuals, D. wulingshanensis should be considered endangered, according to the IUCN Red List criteria.
New taxon, rbcL, subg. Dryopteris, phylogeny, fern
Dryopteris Adans. (1763: 20, 551) is one of the largest fern genera with about 400 species, widely distributed all over the world (
During 2016–2021, we surveyed ferns in the Wulingshan Mountains, which occupy the border zone of four Provinces in China (Hubei, Chongqing, Guizhou and Hunan). This region, as one of the biodiversity hotspots, nurtures a large number of endemic plants and preserves many relict plants (
A total of 32 samples, representing 11 species of the genus Dryopteris, were analyzed to infer the phylogenetic relationships amongst the unknown species and its closest relatives. Dryopteris aemula (Aiton) Kuntze was sampled as an outgroup based on the previous phylogenetic studies of the genus Dryopteris (
Taxon | Voucher specimen | Locality | rbcL |
---|---|---|---|
Dryopteris jishouensis | JSL3612 | Guangxi, China | MZ444597 |
JSL3610 | Guangxi, China | MZ444596 | |
JSL3607 | Guangxi, China | MZ444595 | |
ZXL6320 | Hunan, China | MZ444598 | |
ZXL6317 | Hunan, China | MZ444593 | |
YYH7842 | Guizhou, China | MZ444594 | |
Dryopteris wulingshanensis | ZXL6320-3 | Hunan, China | MZ444607 |
ZXL6320-1 | Hunan, China | MZ444606 | |
JSL3935 | Hunan, China | MZ444605 | |
JSL3926 | Hunan, China | MZ444604 | |
Dryopteris gymnophylla | ZXL6626 | Jiangxi, China | MZ444602 |
JSL4354 | Zhejiang, China | MZ444599 | |
JSL3320 | Anhui, China | MZ444600 | |
JSL3393 | Anhui, China | MZ444601 | |
JSL3949 | Anhui, China | MZ444603 | |
Dryopteris chinensis | JSL3359 | Anhui, China | MZ444610 |
JSL3329 | Anhui, China | MZ444609 | |
JSL2983 | Anhui, China | MZ444608 |
The consensus sequences were generated using SeqMan v7.1.0 (DNASTAR, USA) and then 32 sequences used for phylogenetic analysis were aligned with BioEdit v7.2.0 (
A total of 32 samples were used for phylogenetic analysis, based on chloroplast rbcL gene and the length of sequence alignments was 1,204 bp after removing the missing or gap sites. The phylogenetic relationships amongst D. wulingshanensis and its relatives based on ML and BI algorithms, showed a tree topology to be nearly the same as previous studies (
Morphological comparison amongst Dryopteris wulingshanensis, D. jishouensis and D. gymnophylla A–C D. jishouensis (type locality, Hunan A plants and habitats B stalks of basal pinnae C scales at base of stipe) D–F D. wulingshanensis (Zhangjiajie, Hunan D plants and habitats E stalks of basal pinnae F scales at base of stipe) G–I D. gymnophylla (Zhejiang G plants and habitats H stalks of basal pinnae I scales at base of stipe).
The morphological comparison of Dryopteris wulingshanensis, D. jishouensis and D. gymnophylla.
Species characters | D. jishouensis | D. wulingshanensis | D. gymnophylla |
---|---|---|---|
Vascular bundles at the base of petiole | 2–3 | About 5 | 7–8 |
Length/width of the fronds | 1.7–2.0 times | 1.3–1.6 times | 1–1.2 times |
Stalk of basal pinnae | Usually shorter than 1.5 cm | With a long stalk, usually up to 3cm or more | With a long stalk, usually up to 3cm or more |
Division of the lamina | 3× pinnate | 4× pinnate- pinnatipartite | 3× pinnate-pinnatipartite |
Pinnules | Obtuse | Acuminate | Acuminate |
Habitat | Epipetric | Epipetric | Terrestrial |
China. Hunan: Wulingshan Mountains Zhongli Grand Canyon, Sangzhi County, Zhangjiajie City. 29°39'10.08"N, 110°37'04.29"E, 900 m alt., 26 June 2021, Y.-H. Yan & Z.-R. Gu, YYH24468 (holotype, IBSC; isotypes, NOCC!, CSH!,PE!).
The morphology of D. wulingshanensis was intermediated between D. jishouensis and D. gymnophylla, but more similar to the former. Dryopteris wulingshanensis and D. jishouensis both grow in alkaline soil, but the former’s fronds are ovate, length/width 1.3–1.6 times, tripinnate to quadripartite; the latter’s fronds are ovate-lanceolate to triangular-lanceolate, length/width 1.7–2.0 times or more, bipinnate to tripinnatisect. Dryopteris gymnophylla grows in acidic soil, the fronds are pentagonal, usually length/width 1–1.2 times, tripinnate to quadripinnnatisect (Fig.
Rhizome short-creeping, apex scaly; scales dark brown, lanceolate, margin entire or 1–2 dentate at base. Frond approximate, (40–)65–70 cm; stipe (23–)31–36 cm, medial diameter 1.5–2 mm, basal scales similar to rhizome scales, antrorse, glabrous, stramineous to brown-stramineous, ventrally grooved; lamina ovate, (19–)32–36 × (12–)19–28 cm, about 1.3–1.6 times as long as wide, base round or cordate, apex acuminate, tripinnate to quadripartite (premature lamina only bipinnate to tripinnatisect); pinnae 6–8 pairs, oblique, distant, falcate, stipitate, basal pair longest and largest, opposite to nearly opposite, significantly falcate, stipe (1.2–)3–4.5(–7) cm, deltate-lanceolate, (8–)12–23 × (3.6–)7–11 cm, apex long-acuminate, base broad-cuneate to round, asymmetric, basiscopic pinnules longer than acroscopic pinnules, suprabasal pinnae with pinnules similar; pinnules 7–8 pairs, discrete, oblique to explanate, falcate, base broadly cuneate, pedicellate, basiscopic ones largest, (3–)5–10 × (1.5–)2.2–4 cm, stipe (1.5–)4–9 mm, trigonal oblong, apex obtuse, ultimate pinnules (lobe) 7–10 pairs, oblique to explanate, trigonal oblong, basiscopic ones longer than acroscopic ones, exstipitate, round-obtuse, base broad-cuneate, connate, pinnatifid to partite, (9–)14–20 (4)8–10 mm; lobes 2–7 pairs, 1.5–2 mm wide, oblong, entire, round, others ultimate pinnules lobate to pinnatifid or crenate, base broad-cuneate to decurrent; other pinnae decrescent, opposite to alternate, trigonal lanceolate to oblong lanceolate; rachis and costa glabrous, stramineous, adaxially sulcate, lamina papyraceous, glabrous; veins pinnate, single or dichotomous; sori round, on apices of veinlets, nearly incision; indusia orbicular-reniform, margin premorse, brown, persistent.
China. Hunan: Pangu Peak, Dehang Scenic Area, Jishou City, Xiangxi Tujia and Miao Autonomous Prefecture. 28°20'50.046"N, 109°34'34.522"E, 914 m alt., 26 Jun 2016, X.-L. Zhou et al. ZXL6320-1, ZXL6320-3 (CSH!); Tianmenshan Scenic Area, Zhangjiajie City. 5 April 2016, H.-J. Wei et al. JSL3926, JSL3935 (CSH!).
Dryopteris wulingshanensis is endemic to the Wulingshan mountains in Jishou and Zhangjiajie Cities, Hunan, China. It is epipetric in limestone crevices at an elevation of 700–1000 m in evergreen and deciduous broad-leaved mixed forest. The associated ferns include: Cheilanthes patula Baker, Cyrtomium fortunei J. Sm., Cyrtomium nephrolepioides (Christ) Copel., Lemmaphyllum drymoglossoides (Baker) Ching, Polystichum tsus-simense (Hook.) J. Sm., Pronephrium penangianum (Hook.) Holttum, Pyrrosia petiolosa (Christ) Ching and Woodwardia unigemmata (Makino) Nakai and associated seed plants include: Celtis sinensis Pers., Choerospondias axillaris (Roxb.) B.L. Burtt & A.W. Hill, Davidia involucrata Baill., Ficus sarmentosavar. henryi (King ex Oliv.) Corner, Hydrangea strigosa Rehder, Loropetalum chinense (R. Br.) Oliv., Mallotus philippinensis (Lam.) Muell. Arg., Miscanthus sinensis Andersson, Platycarya strobilacea Siebold & Zucc., Rhus chinensis Mill. and Viburnum cylindricum Buch.–Ham. ex D. Don.
The specific epithet “wulingshanensis” is derived from the name of type locality Wulingshan Mountains, where the new species is found.
EN(B1ab(iii)). Dryopteris wulingshanensis is only known from three locations of Wulingshan Mountains in Jishou and Zhangjiajie Cities, Hunan, China. Based on its restricted geographic range, small populations and few individuals, Dryopteris wulingshanensis should be considered endangered under the IUCN Red List criteria (
This research was funded by the Strategic Priority Research Program of the Chinese Academy of Sciences (XDA19050404). We are very grateful to the subject editor Blanca León and anonymous reviewers for their helpful comments and modification assistance. We thank Prof. Gong-Xi Chen from Jishou University and Mr. Yang Xiang from Badagongshan National Nature Reserve, Hunan Province, China, for assisting in the field surveys.