Hedwigia filiformis (Michx.) P.Beauv., Prodr. Aethéogam.

Huang, Wen-Zhuan, Ma, Wen-Zhang, Ignatov, Michael S., Ignatova, Elena A., Sulayman, Mamtimin, Wu, Yu-Huan & Zhu, Rui-Liang, 2024, A taxonomic revision of Hedwigia (Hedwigiaceae, Hedwigiales) in China, Phytotaxa 634 (3), pp. 215-234 : 224-225

Hedwigia filiformis (Michx.) P.Beauv., Prodr. Aethéogam. : 60. 1805.

Type:— UNITES STATES OF AMERICA. États-Unis, A. M. F. J. Palisot de Beauvois s.n. (Lectotype: P, selected and commented by De

Luna 2022). = Hedwigia nemoralis Ignatova, Ignatov & Fedosov, Arctoa 25 (2): 252. 2016. Syn. nov. Type:— RUSSIA. Primorsky Territory, Olga District, waterfall on Milogradovka Creek, 43°27.5′ N, 134°19.3′ E, 370 m alt., dry cliffs near waterfall, 21 Aug. 2007. Ignatov 07-324 (holotype: MHA).

Note: — Hedwigia filiformis was described based on a specimen originally determined as Anictangium filiforme Michaux (1983: 287) from Eastern North America by Michaux (1983), and a lectotype specimen was selected and commented by De Luna (2022). Hedwigia nemoralis was described based on a specimen from Primorsky Territory, eastern Asian Russia by Ignatova et al. (2016), and reported also from Europe and North America. Both H. nemoralis and H. filiformis are characterized by (1) small to medium-sized plants, (2) ovate leaves, (3) stem leaves with short hyaline apices, (4) narrowly recurved leaf margins, and (5) median leaf cells with 2–3(–4) simple or branched papillae on abaxial surfaces ( Ignatova et al. 2016; De Luna 2022). Though De Luna (2022) has emphasized that they are indistinguishable morphologically, and suggested that H. nemoralis might be a synonym of H. filiformis , he did not synonymize them.

We also could not find distinct differences to separate Eurasian H. nemoralis from North American H. filiformis morphologically. In addition, the phylogenetic topology showed that the obtained additional sequences of H. filiformis from U.S.A. (including Connecticut, Maryland, and Virginia) and accessions from Eurasian H. nemoralis formed a highly supported clade ( Fig. 1 ), which confirms they are the same species. Hedwigia nemoralis is thus proposed as a new synonym of H. filiformis .

This species is distributed in Asian Russia ( Ignatova et al. 2016), Japan and South Korea ( De Luna 2022), Europe ( Ignatova et al. 2016; De Luna 2022; Pihlaja et al. 2023), and North America ( Ignatova et al. 2016; De Luna 2022). In China, H. filiformis was previously reported in Anhui (as H. nemoralis by Ignatova et al. 2017) and Heibei province (as Hedwigidium integrifolium by Chen 1978), and the present study expands its range to Fujian, Guangdong, Guangxi, Henan, Hunan, Jiangxi, Liaoning, Shaanxi, Taiwan, Xinjiang, and Zhejiang.

Specimens examined:— CHINA. Anhui: Chizhou City, Qingyang County, Zhubei Town , Dongqiao Village , 30°32′25.08″ N, 117°51′24.12″ E, 116 m, on rock, 16 Oct. 2019, Z. Dai R0010-20191016 - 09 ( HSNU) ; Chizhou City, Shitai County, Dayan Town , Xinnong Village , 30°4′26.76″ N, 117°30′2.16″ E, 394 m, on soil, 13 Oct. 2020, S. W. Tu R002-032 A ( HSNU) . Fujian: Quanzhou City, Dehua County, Mt. Daiyun , Shanyong Town , Houzhai Village , 25°41′8″ N, 118°13′2″ E, 870 m, on rock, 28 Oct. 2009, Y. F. Wang et al. 20091028-199 ( HSNU) ; ibid., Y. F. Wang et al. 20091028-274 ( HSNU) ; ibid., Y. F. Wang et al. 20091028-325 ( HSNU) ; Dehua County, Mt. Daiyun, Chishui Town , Xixi Forest , 25°41′00″ N, 118°8′29″ E, 830 m, on rock, 2 Nov. 2009, Y. F. Wang & W. Y. Wu 20091102-1188 ( HSNU) ; Dehua County, Mt. Daiyun, Leifeng Town , Handou Stream , 25°39′56.15″ N, 118°17′20.61″ E, 561 m, on rock, 5 Apr. 2010, Y. F. Wang & W. Y. Wu 20100405-299 ( HSNU) ; Mt. Wuyi National Nature Reserve, Xingcun County, Tongmu Village , Guadun , 27°44′06.87″ N, 117°38′17.62″ E, 1222 m, on rock, 30 Aug. 2015, Y. F. Wang et al. 201508032 ( HSNU) ; ibid., 27°43′33.99″ N, 117°42′50.77″ E, 572 m, on rock, 1 Sept. 2015, Y. F. Wang et al. 201509428 ( HSNU) . Guangdong: Lechang City, Baishi Town , 1000 m, on rock, 15 Oct. 1990, G. Q. Zeng 357a ( IBSC) . Guangxi: Baise City, Leye County, Yachang Wild Orchids National Nature Reserve , Panguwang , 24°44′51.00″ N, 106°20′20.69″ E, 1755 m, on rock, 19 May 2019, Q. M. Tang 20190519-350 ( IBK) ; ibid., Q. M. Tang 20190519-351 ( IBK) ; ibid., Dapeng Tun , 24°45′24.67″ N, 106°20′27.54″ E, 1643 m, on rock, 20 May 2019, Q. M. Tang 20190520-389 ( IBK) ; Guilin City, Longsheng Co., Longsheng town , Duping Villiage , yinshuitongzhai, 25°48′34.62” N, 109°58′28.25” E, 207.8 m, on rock, 1 Mar. 2023, Y. M. Wei & Q. M. Tang 4503282023 BRY0936 ( IBK) ; ibid., Jiangdi Township, Jianxin Villiage, Tangjia Tun , 25°52′40.84” N, 110°12′56.80” E, 358.2 m, on rock, 1 Mar. 2023, Y. M. Wei & Q. M. Tang 4503282023 BRY1181 ( IBK) . Henan: Xinyang City, Luoshan county, Lingshan Town , Dongqiao Village , 31°56′45″ N, 114°16′25″ E, 90 m, on rock, 25 Jun. 2022, Y. Y. Liu 22301 ( JZNC) ; ibid., Y. Y. Liu 22308 ( JZNC) ; Xinyang City, Shangcheng County, Donghe Village , Jingangtai Nature Reserve , on rock, 9 Apr. 2016, Y. Y. Liu 16065a ( JZNC) ; ibid., 18 Jun. 2018, Y. Y. Liu 18163 ( JZNC) . Hubei: Shennongjia Forestry District, Pingqian, Linghekou , 1500 m, on rock, 18 Jul. 1997, S. X. Liu 510 ( CCNU) ; Xianning City, Tongshan County, Jiugongshan , 450 m, on rock, 15 Jul. 1999, G. Y. Chen 2826 ( CCNU) ; Huanggang City, Luotian County, Jiuzihe Town , 300 m, on rock, 1999, S. X. Liu 3652 ( CCNU). Hunan: Liuyang City, Daweishan Town , Daweishan National Nature Reserve , 1300 m, on rock, 27 Aug. 1974, 1974 collection team 289 ( IBSC) . Jiangxi: Yichun City, Jingan County, Baofeng Town , 28°49′27.12″ N, 115°3′41.18″ E, 1038 m, on stone, 30 Oct. 2020, Y. Sun 117 ( LBG) ; ibid., Shuikou Town, Qingshan Village , 29°0′9.98″ N, 115°16′23.46″ E, 966 m, on stone, 18 Nov. 2020, C. F. Chen & Y. Sun-C148 ( LBG) ; ibid., 28°59′06.70″ N, 115°16′29.70″ E, 321 m, on stone, 22 Nov. 2022, Y. Sun 406 ( LBG) ; Jiujiang City, Pengze County, Taohongling Sika Deer National Nature Reserve, 29°49′39.54″ N, 116°37′48.84″ E, 471 m, on stone, 29 Mar. 2023, C. F. Chen & T. J. Liang-C957 ( LBG) ; Ganzhou City, Chongyi County, Qiyunshan National Nature Reserve , 25°49′48.21″ N, 113°58′33.63″ E, 981 m, on stone, 5 Jun. 2023, C. F. Chen & Z. Y. Li-C1143 ( LBG) . Liaoning: Dalian City, Zhuanghe County, Xianrendong National Nature Reserve , Daguanmen , on rock, 13 Jul. 1988, X. Y. Jia 468 ( IFP) ; Shaanxi: Shangluo City, Zhenan County, Maoergou , Naoshikan , on rock, 12 Feb. 2013, L. Y. Yao 13-618 ( KUN) . Taiwan: Taizhong City, Wuling National Forest Recreation Area , at end of Wuling Road at Recreation Area Visitor Station then on the trail toward Taoshan Waterfall , Taoshan Stream near confluence with Cijiawan River , 24°23′53.0″ N, 121°18′28.7″ E, 1925 m, on metamorphic boulders in filtered light, 13 Oct. 2015, James R. Shevock 47987 ( KUN). Xinjiang: Altai mountains , Haba County, Hutelin Village , 48°17ʹ53.58ʺ N, 86°36ʹ29.93ʺ E, 1140m, on rock, 12 Jul. 2016, Mamtimin Sulayman 28180 ( XJU). Zhejiang: Quzhou city, Kaihua county , Gutianshan National Nature Reserve , on rock, May 1993, C. Y. Tian 0846 ( HSNU) ; ibid., C. Y. Tian 0848 ( HSNU) ; ibid., C. Y. Tian 0653 ( HSNU) ; ibid., C. Y. Tian 0551 ( HSNU) ; Quzhou city, Changhong county, Qiangjiangyuan National Park , Kukeng Reservoir , near the gallery, 29°18′51.48″ N, 118°11′37.94″ E, 408 m, on soil, 14 Apr. 2019, R. L. Zhu et al. 20190414-3 ( HSNU) ; Hangzhou city, Chunan County, Thousand Island Lake , Mushan Island , 29°31′ N, 119°4′ E, 242 m, on soil, 29 Oct. 2006, Y. F. Wang 245 ( HSNU) ; ibid., 21 Apr. 2007, Q. Liu & Y. Q. Zhu 494 ( HSNU) ; ibid., Q. Liu & Y. Q. Zhu 387 ( HSNU) ; Hangzhou city, Linan District, Tianmushan National Nature Reserve , 30°19′19.30″ N, 119°26′26.34″ E, 349 m, on rock, 19 Apr. 2011, D. H. Wang et al. 64 ( HSNU) ; ibid., 30°19′19.30″ N, 119°26′34.80″ E, 349 m, on rock near river, 19 Apr. 2011, D. H. Wang et al. 73 ( HSNU) ; ibid., 30°19′12.79″ N, 119°26′38.38″ E, 329 m, on rock, 27 Apr. 2011, D. H. Wang et al. 1107 ( HSNU) ; ibid., D. H. Wang et al. 1109 ( HSNU) ; ibid., 30°22′11.66″ N, 119°24′53.90″ E, 911 m, on rock, 28 Apr. 2011, D. H. Wang et al. 1209 ( HSNU) ; ibid., 30°22′11.89″ N, 119°25′10.56″ E, 856 m, on rock wall, 28 Apr. 2011, D. H. Wang et al. 1211 ( HSNU) ; ibid., 30°20′43.18″ N, 119°27′16.75″ E, 416 m, 24 Apr. 2023, on rock, J. Wang & Q. Wu 20230427-5 ( HSNU) ; Jinhua City, Beishan , near the reservoir, 29°12′8″ N, 119°37′51″ E, 480 m, on rock, 7 Nov. 2020, R. L. Zhu 20201107-5 B ( HSNU) ; Shengzhou City , Guimen County, 29°21′25.45″ N, 120°36′43.06″ E, 308 m, on rock, 6 Feb. 2019, R. L. Zhu 20190206-6 ( HSNU) ; Shengzhou City, Guimen County, Jiexi Village , 29°21′18″ N, 120°36′26″ E, 371 m, on rock, 5 Apr. 2023, R. L. Zhu 20230405-10 ( HSNU) ; ibid., 29°21′13″ N, 120°36′49″ E, 400 m, on rock, 27 Apr. 2009, R. L. Zhu 20090427-1 ( HSNU) ; Lishui City, Qingyuan County, Xiyan Village , 27°39′47.35″ N, 119°12′18.81″ E, 795 m, on stone, 21 Aug. 2022, X. Y. Ma et al. 20220821-32 ( HTC) ; Lishui City, Qingyuan County, Xiyan Village , 27°39′08.48″ N, 119°12′32.35″ E, 694 m, On soil, 20 Aug. 2022, X. Y. Ma et al. 20220820-05 ( HTC) ; Lishui City, Suichang County, Baimashan Forest Park , 28°38′20.68″ N, 119°10′00.63″ E, 947 m, On stone, 4 Aug. 2016, W. Sheng 20180804-6 ( HTC) ; ibid., W. Sheng 2016080410-1 ( HTC) ; ibid., Baimashan Forest Park, 28°38′22.30″ N, 119°10′21.65″ E, 840 m, On stone, 4 Aug. 2016, W. Sheng 2016080429-3 ( HTC) ; Lishui City, Suichang County, Jiulongshan National Forest Park , on stone, 21 Aug. 2016, W. Sheng 2016083144-1 ( HTC) ; Lishui City, Songyang County, Ruoliao Primeval Forest Scenic Area , 28°18′30.32″ N, 119°17′30.38″ E, 672 m, On stone, 4 Aug. 2017. W. Sheng 20170804-19 ( HTC) ; Longquan City, Madi Village, 28°07′44.5″ N, 119°1′17.99″ E, 410 m, On soil, 19 Dec. 2009, R. L. Zhu 20091219-16 ( HSNU) ; Taizhou City, Tiantai County, Baxian Lake , 29°12′58″ N, 120°59′51″ E, 164 m, on stone, 25. Jun. 2023, R. L. Zhu 202300605-28 ( HSNU) ; Taizhou City, Tiantai County, Canghua Village , 29°11′54.27″N, 121°09′29.32″ E, 526 m, on Rock , 19 Jul. 2021, X. Y. Ma & S. F. Tang 20210719-29 ( HTC) ; Wenzhou City, Taishun County, Juxi Town , on the way from Zhikeng Village to Hengkengyuan , 27°31′36.83″N, 120°06′49.06″ E, 744 m, on rock, 13 Apr. 2018, Y. H. Wu et al. 20180413-138 ( HTC) .

Figures

FIGURE 1. Phylogeny of Hedwigia species inferred from the combined dataset (nad5, trnL-trnF and ITS). The topology derived from the BI tree is shown. The symbol ‘‘//’’ means the branch length shown in the representative picture is only 1/2 of the actual length. ML bootstrap values BS ≥ 75 are shown on left and Bayesian posterior probabilities values PP ≥ 0.90 on right. The five Chinese accessions of the new species are marked in red and arrowed.