Asplenium wrightii Eaton ex Hooker, Sp. Fil. 3: 113. pl. 182. 1860.

Xu, Ke-Wang, Wang, Lu-Lu & Zhang, Li-Bing, 2021, Taxonomic revision of the Asplenium wrightii complex (Aspleniaceae) with reinstatement of A. alatulum and A. subcrenatum, PhytoKeys 172, pp. 75-91 : 75

Asplenium wrightii Eaton ex Hooker, Sp. Fil. 3: 113. pl. 182. 1860.

Asplenium duplicatoserratum Ching ex S.H. Wu, Bull. Bot. Res., Harbin 9(2): 86, f. 4. 1989.

Asplenium fujianense Ching ex S.H. Wu, Bull. Bot. Res., Harbin 9(2): 21, 88-89, f. 7. 1989.

Asplenium laui Ching, Acta Phytotax. Sin. 9(4): 360-361. 1964.

Asplenium neomultijugum Ching ex S. H. Wu, Bull. Bot. Res. 9 (2): 21, f. 6. 1989.

Asplenium pseudowrightii Ching, Acta Phytotax. Sin. 9(4): 360. 1964.

Asplenium serratissimum Ching, Bull. Bot. Res., Harbin 9(2): 87. 1989.

Asplenium taiwanense Ching ex S.H. Wu, Bull. Bot. Res., Harbin 9(2): 85-86, f. 3. 1989.

Asplenium wrightioides Christ, Bull. Acad. Int. Géogr. Bot. 11(153-154): 238-239. 1902.

Type.

Japan. Kagoshima: Ryukyu Islands, 1853-1856, C. Wright, #s.n. (holotype: NY (NY-00128031 [image!], isotype: GH (GH00020612 [image!])). Fig. 3C, D .

Description.

Plants 35-70(-100) cm tall. Rhizome erect, short, scaly; scales brown to dark brown, lanceolate-triangular, 5-12 × 0.6-1.2 mm, denticulate glandular margin or long fibrillose. Fronds tufted; stipe greyish-green to brown, (18-)23-29(-31) cm, base densely scaly, scales brown, lanceolate to linear-lanceolate, (4.6-)6.1-7.8(-8.8) mm long, with multicellular hairs, subglabrous towards apex; lamina ovate-lanceolate to lanceolate, (19-)39-70(-88) × (9-)17-27(-35) cm, base truncate, apex acute, 1-pinnate; pinnae (12-)17-25(-34) pairs, basal pinnae subopposite, others alternate, at an angle of ca. 50°-60° to rachis, with stalks (2-)4-8 mm, lower pinnae slightly reduced, suprabasal pinnae narrowly oval-lanceolate and often falcate, (6-)9.1-13(-20) × (0.9-)1.2-1.8(-2.5) cm, base asymmetrical, acroscopic side truncate at an angle of (40°-)55°-75°(-85°) to costa and often auriculate, basiscopic side cuneate, at an angle of (20°-)30°-40°(-60°), becoming decurrent on rachis in central part of lamina, margin serrate to coarsely dentate, apex acuminate. Veins (1 or)2-forked, with terminal hydathode (Fig. 6B, C, D ). Fronds papery, green to brownish-green when dry, subglabrous; rachis dull green to reddish-brown, terete abaxially, winged towards apex. Sori linear, (3-)6-10(-12) mm, on acroscopic veinlets, medial (Fig. 6B, C ); indusia brown, linear, papery, opening towards costa, persistent. Spores with average exospore length 32-45 μm, perispore cristato-alate.

Distribution and habitat.

Asplenium wrightii is commonly distributed in China and Japan and it might occur in Korea. This species is found in damp valleys under evergreen broad-leaved forests where it grows as a lithophyte by streams (Fig. 6A ).

Additional specimens examined.

China. Anhui: Qimen County, Xifengsi, Nov. 1957, M.B.Deng 5308 (HZ-028581); Jing County, Dingxi Village, Suhong, 22 Sep 1959, 0431 (NA-S00153661); Tongling County, Xiaokeng, Jun 1985, X.L.Liu 85053 (HUST-00008926); Chongqing: Pengshui County, Hanjia Village, elev. 370 m, 29°25'137.98"N, 108°18'19.1"E, 16 Oct 2012, Pengshui expedition 500243-003-167 (IMC-0003680); Yangshan County, Nanling Nature Reserve, Xinerkeng, 29 Sep 2007, X.L.Zhou & H.F.Chen 785 (HUST-00008767); Fujian: 22 Jul 2015, ZXL05538 (CSH-0101629); 10 Apr 2014, H.Shang & Y.F.Gu SG134 (CSH-0034578); Wuyishan, elev. 750 m, 15 Aug 2011, X.F.Zeng 11215 (CZH-0006554); Zhaoan County, Wushan, 12 Apr 2015, X.F.Zeng ZXF20041 (CZH-0012412); Nanping City, Mangdangshan, 17 Feb 1999, G.S.He 9643 (PE-00913318); Jianning County, 20 Nov 1977, Z.Y.Li 10619 (PE-0913321); Jiangle County, Longxishan, 27 May 1991, Longxishan expedition 0359 (PE-01555190); Guangxi: Lingyun County, Yuhong Village, Donglan, elev. 1303 m, 24°24'13.19"N, 106°29'03.17"E, 16 Aug 2013, Lingyun expedition 451027130816103 (GXMG-0117634); Longshen County, Heping Village, elev. 615 m, 25°41'12.10"N, 110°03'22.40"E, 6 Mar 2013, Longshen expedition 450328130306038LY (IBK-00362003); Xingan County, Maoershan, 5 Oct 2007, L.Wu & X.X.Xu 1042 (HUST-00019436); Sanjiangdongzu County, Sanxingpo, Dudong Village, elev. 700 m, 18 Dec 2007, X.X.Xu & L.C.Qin 255 (HUST-00011550); Guangdong: Heping County, Daba Village, Dafukeng, 2 Jan 2007, C.M.Tan et al. Y06630 (HUST-00004651); Lechang County, 23 Nov 1931, Z.Huang 31490 (IBK-00035278); the same locality, 21 Jun 1942, S.Q.Chen1611 (IBK-00035299); Boluo County, Luofushan, 18 Aug 1930, N.Q.Chen 41631 (IBK-00035279); Xinyi, 13 Aug 1931, Z.Huang 31164 (IBK-00035285); Dawuling, 4 Aug 2003, Y.H.Yue et al. 1568 (PE-01784537); 21 Mar 1931, Z.Huang 31756 (PE-00913341); Ruyuan, Daxiagu, 2 Aug 2005, B.R.Liu 05100 (PE-01785865); Yingde, Shimentai Nature Reserve, Oct 2001, Y.H.Yue & F.W.Xing 13265 (PE-01784538); Mei County, Jiaying, 4 Aug 1932, Tsang, W.T. 21466 (PE-00913338); Guizhou: Yongshun County, Xiaoxi Village, elev. 621 m, 28°82'21.10"N, 110°25'11.00"E, 14 Jan 2014, D.G.Zhang zdg9949 (JIU-04159); Liping County, Pingjia, elev. 670 m, 16 Dec 2007, X.X.Xu & L.C.Qin 349 (HUST-00011558); Libo County, Maolan Nature Reserve, 25 Apr 2015; X.C.Zhang 7259 (PE-02051530); Shiqian County, Qiangyang Village, Longdong, 30 Jul 1988, Wulingshan expedition 2355 (PE-01557785); Jiangkou County, 30 Aug 1986, B.Bartholomew et al. 595 (PE-00913309); Jianhe County, Nanjiaqu, 30 Apr 1992, F.Wang 651 (HGAS-055247); Jinping County, Gaodengpo, 27 Nov 1991, fern expedition 91474 (HGAS-055248); Rongjiang County, Shuiwei Shuizu Village, 15 Oct 2014, Wei et al. WYG036 (CSH-0043860); Shibing County, Maohao Village, 30 May 2016, D.Y.Zhou 522623160530476LY (GZTM-0066044); Danzhai County, 18 Oct 2012, Hou GZDZ201210180003 (GYBG-0009834); Hunan: Sangzhi County, Bamaoxi Village, 21 Oct 2014, X.L.Zhou et al. ZXL09673 (CSH-0045103); Baojing County, Baiyunshan Nature Reserve, elev. 397 m, 28°37'51.90"N, 109°17'11.34"E, 11 Aug 2012, X.J.Su. & H.B.Liu 433125D00030810086 (JIU-06613); Yongshun County, Xiaoxi, Daping, 12 Sep. 2009, L.Xu 090912005 (JIU-02238); Dongzu County, Pingyang Village, Yangdongtan, elev. 300 m, 10-15 Jan 2008, L.C.Qing & H.B.Ouyang 1080 (HUST-00012147); Shangyan, Sanxingpo, elev. 750 m, 14 Dec 2004, L.C.Qing. & X.X.Xu 691 (HUST-00011548); Suining County, Huangsang Nature Reserve, elev. 450 m, 10 Jan 2008, J.M.Xi & Y.B.Qin 07919 (HUST-00012150); Yanling County, Taoyuandong Nature Reserve, Jul 2008, X.L.Zhou &Z.L.Zhu 2154 (PE-01964123); Rucheng County, Donggangling, Shuinishan, elev. 110 m, 31 Jul 2007, L.Wu & H.B.Ouyang 88 (HUST-00008785); Pingjiang County, Sicun Village, Wudeng, elev. 500 m, 15 Dec 2007, L.Wu & S.X.Qi W467 (HUST-00011555); Huaihua County, Bamianshan, 15 Dec 2007, X.L.Zhou & Y.Q.Yao 1744 (HUST-00011562); Hubei: Laifeng County, Dahe Village, Lianghekou, elev. 590 m, 25 Jul 2013, W.Z.Zhu BLF077 (CCAU-0002976); Jiangxi: Xingzi County, Wenquan Village, Xiufeng, 26 Jul 2013, A.M.Dong 2082 (JJF00010847); Wanzai County, Jiulong Forest Park, elev. 190 m, 28°21'38"N, 114°30'47"E, 15 Aug 2013, H.G.Ye & F.Y.Zeng LXP10-2161 (IBSC-0773610); Wuning County, Yuangkou Village, Dongkeng, elev. 400 m, 16 Oct 2012, J.H.Zhang 2045 (JJF-00010846); Chongyi County, Fengzhou Village, Changlongao, elev. 1000 m, 8 Jul 2007, L.Y.Wang W.181 (HUST-00008778); Jianggangshan, elev. 800 m, 10 Apr 2004, Y.H.Yan & J.S.Zhou 3415 (HUST-00001166); Jiangsu: Yixing, Longchi, Shuizhushan, 1956, Liu & Huang 2874 (NAS-00091223); Sichuan: Emeishan, 15 Sep 1963, G.X.Xing & K.Y.Lang 1788 (PE-13307); the same locality, 21 Sep 1979, Z.R.Wang C122 (PE-96543); Zhulian County, 1 Jun 1978, X.R.Kong 5204 (CDBI-003584); Taiwan: Wulai, Fushan-Hapen, 13 Oct 1984, Y.Tateishi et al. 20492 (PE-01451621); Takao, 27 Jan 1939, M.Tagawa 2049 (PE-00913326); W.Hancock 141 (PE-00913329); Taito, 4 Mar 1940, M.Tagawa 3168 (PE-00913330); Huanlian County, Changliang, 8 Jun 2001, T.C.Chen 11303 (PE-01451613); Taibei County, Lujiaokengxi, 28 Jan 2000, T.C.Chen 10393 (PE-01451614); Zhejiang: Tonglu County, Luci, Changzhou, 8 Nov 1989, L.Hong s.n. (HZ-028576); Shouchang Zhen, 19 Jun 1959, Zhejiang expedition 27803 (HZ028578); Taishun County, Liguang, Huangshikeng, 27 Nov 1958, Anonymous 23818 (HZ-028579); Lishui, Suichang Nature Reserve, elev. 780 m, 25°59'11"N, 116°25'11"E, 5 Jun 2012, Q. Tian TQ01773 (CSH-0002430); 25 Nov 2015, H.J.Wei JSL3466 (CSH-0110219); 29 Jul 1958, Shan 5578 (PE-00913289);

Japan. Kyushu: Chikushi-yabakei, Nakagawa-cho, Chikushi-gun, Fukuoka Pref., 27 Mar 1975, T.Nakaike et al. 105 (PE-01708808); Is. Yakushima (Kagoshima Pref.), from Kurio to Segiri through Ookonotaki, elev. 30-100 m, 16 Jul 1979, T.Yamazaki et al. 2327 (PE-01708809); Kagoshima Pref., Okuchi-shi, Jusso, 6 Sep 1959, Tetsuji Yamanaka s.n. (PE-01964369); Prefecture Kagoshina, Yakushina Island, Yaku-cho, Tainoko-gawa River, 10-11 Oct 1977, Boufford, David Edward 20118 (MO-3136075); Along trail in gorge, foot of Mt. Awa. Motobu Peninsula, Okinawa Island, 10 Dec 1953, Walker, EH 7614 (L-3508507); Honshu: Owase City, Kuki, Namera Valley, 12 Nov 1991, K.H.Shing 31 (PE-01708810); Pref. Mie,Kuki, Omse-shi, 5 Oct 1971, Kato, Masahiro-259 (MO-3136077); Pref. Schizouka: Aono, Minamiizu-cho, Kamo-gun, 30 Dec 1972, T.Nakaike 5 (L-3508506); Shikoku: Nakaohsaka Ohsakadani, Nakatosa Town, Takaoka Distr. Kouchi Pref., elev. 60-80 m, 33°18'48"N, 133°23'48"E, 6 Jan 2012, Taku Miyazaki 1201164 (PE-02002704); Yatabe, Nakatosa Town, Takaoka District, Kouchi Pref., elev. 40-60 m, 33°15'70"N, 133°15'37"E, 3 Jan 2007, T.Miyazaki 0701137 (PE-01963957); Takanoyama-rindou Yamauchi, Nakatosa Town, Takaoka Distr. Kochi Pref., elev. 60-90 m, 33°17'04"N, 133°12'18"E, 27 Mar 2009, Taku Miyazaki 0903138 (PE-01963829); Ohkawauchi Kaminokae, Nakatosa Town, Takaoka District, Kouchi Pref., elev. 20 m, 33°16'67"N, 133°14'15"E, 29 Dec 2006, Taku Miyazaki 0701025 (PE-01962310).

Note.

Asplenium wrightii is described based on the type material from Japan and it is widely distributed in China. The morphology of this species is variable, which resulted in the taxonomic chaos amongst A. wrightii and its closely-related species. Several ploidy levels (e.g. tetraploid, octoploid, decaploid and dodecaploid) have been reported for the A. wrightii complex, but no correlation was found between ploidy levels and morphological characters within the complex ( Mitui 1967; Wang 1988; Lin and Viane 2013). Asplenium x shikokianum is a natural hexaploid hybrid between octoploid A. wrightii and tetraploid A. ritoense and Asplenium x wangii is another hybrid between A. wrightii and A. bullatum . These hybrids are usually uncommon in terms of their distributions where their parents grow together ( Kuo 1988; Lin and Viane 2013). Due to hybridisation and polyploidisation, there are a large number of recognised species complexes with ambiguous boundaries between species in Aspleniaceae ( Reichstein 1981; Dyer et al. 2012). More comprehensive taxonomic studies are still needed to elucidate the species delimitation in this species complex.

Asplenium wrightii was also documented in Vietnam ( Pham 1999; Lin and Viane 2013). However, we checked specimens identified as A. wrightii from Vietnam and found none of these specimens is true A. wrightii . It is possible that the name A. wrightii has been used erroneously for species A. subcrenatum in Vietnam.

Figures

Figure 3. Type of Asplenium alatulum Ching (A barcode PE 00059412), A. subcrenatum Ching ex S. H. Wu (B barcode PE 00059494), A. wrightii Eaton ex Hooker (C barcode NY 00128031) and isotype of A. wrightii GH (D barcode GH 00020612).

Figure 6. Asplenium wrightii A Habitat and habit B Abaxial view of upper portion of lamina C Abaxial view of pinna showing the almost entire to crenate-sinuate margin D Adaxial view of upper portion of lamina E Upper portion of rachis showing the lateral wings.