Hedwigia filiformis (Michx.) P.Beauv., Prodr. Aethéogam. : 60. 1805 Hedwigia nemoralis Ignatova, Ignatov & Fedosov, Arctoa 25 (2): 252. 2016 A taxonomic revision of Hedwigia (Hedwigiaceae, Hedwigiales) in China Huang, Wen-Zhuan Ma, Wen-Zhang Ignatov, Michael S. Ignatova, Elena A. Sulayman, Mamtimin Wu, Yu-Huan Zhu, Rui-Liang Phytotaxa 2024 2024-01-18 634 3 215 234 3JZKD (Michx.) P. Beauv., Prodr. Aetheogam. P. Beauv., Prodr. Aetheogam. Michx. [136,798,1346,1373] Bryopsida Hedwigiaceae Hedwigia Plantae Leucodontales 9 224 Bryophyta species filiformis   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 nemoralisIgnatova, Ignatov & Fedosov, Arctoa25 (2): 252. 2016. Syn. nov. Type:— RUSSIA. PrimorskyTerritory, Olga District, waterfall on Milogradovka Creek, 43°27.5′ N, 134°19.3′ E, 370 malt., dry cliffs near waterfall, 21 Aug. 2007. Ignatov 07-324( holotype: MHA).  Note:—  Hedwigia filiformiswas described based on a specimen originally determined as  Anictangium filiformeMichaux (1983: 287)from Eastern North America by Michaux (1983), and a lectotypespecimen was selected and commented by De Luna (2022).  Hedwigia nemoraliswas described based on a specimen from PrimorskyTerritory, eastern Asian Russiaby Ignatova et al.(2016), and reported also from Europe and North America. Both  H. nemoralisand  H. filiformisare 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. nemoralismight be a synonym of  H. filiformis, he did not synonymize them. We also could not find distinct differences to separate Eurasian  H. nemoralisfrom North American  H. filiformismorphologically. In addition, the phylogenetic topology showed that the obtained additional sequences of  H. filiformisfrom U.S.A.(including Connecticut, Maryland, and Virginia) and accessions from Eurasian  H. nemoralisformed a highly supported clade (  Fig. 1), which confirms they are the same species.  Hedwigia nemoralisis thus proposed as a new synonym of  H. filiformis. This species is distributed in Asian Russia( Ignatova et al. 2016), Japanand 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. filiformiswas previously reported in Anhui(as  H. nemoralisby Ignatova et al. 2017) and Heibei province (as  Hedwigidium integrifoliumby 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. Wanget al. 20091028-199( HSNU); ibid.,  Y. F. Wanget al. 20091028-274( HSNU); ibid.,  Y. F. Wanget 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. Wu20091102-1188( 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. Wu20100405-299( 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. Wanget al. 201508032( HSNU); ibid., 27°43′33.99″ N, 117°42′50.77″ E,  572 m, on rock,  1 Sept. 2015,  Y. F. Wanget al. 201509428( HSNU).   Guangdong: Lechang City, Baishi Town,  1000 m, on rock,  15 Oct. 1990,  G. Q. Zeng357a( 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. Tang20190519-350( IBK); ibid.,  Q. M. Tang20190519-351( IBK); ibid., Dapeng Tun, 24°45′24.67″ N, 106°20′27.54″ E,  1643 m, on rock,  20 May 2019,  Q. M. Tang20190520-389( 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. Tang4503282023 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. Tang4503282023 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. Liu22301( JZNC); ibid.,  Y. Y. Liu22308( JZNC);  Xinyang City, Shangcheng County, Donghe Village, Jingangtai Nature Reserve, on rock,  9 Apr. 2016,  Y. Y. Liu16065a( JZNC); ibid.,  18 Jun. 2018,  Y. Y. Liu18163( JZNC).   Hubei: Shennongjia Forestry District, Pingqian, Linghekou,  1500 m, on rock,  18 Jul. 1997,  S. X. Liu510( CCNU);  Xianning City, Tongshan County, Jiugongshan,  450 m, on rock,  15 Jul. 1999,  G. Y. Chen2826( CCNU);  Huanggang City, Luotian County, Jiuzihe Town,  300 m, on rock, 1999,  S. X. Liu3652( 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. Sun117( 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. Sun406( 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. Jia468( IFP);   Shaanxi: Shangluo City, Zhenan County, Maoergou, Naoshikan, on rock,  12 Feb. 2013,  L. Y. Yao13-618( KUN).   Taiwan: Taizhong City, Wuling National Forest Recreation Area, at end of Wuling Roadat Recreation Area Visitor Stationthen on the trail toward Taoshan Waterfall, Taoshan Streamnear 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 Sulayman28180( XJU).  Zhejiang: Quzhoucity, Kaihua county, Gutianshan National Nature Reserve, on rock,  May 1993,  C. Y. Tian0846( HSNU); ibid.,  C. Y. Tian0848( HSNU); ibid.,  C. Y. Tian0653( HSNU); ibid.,  C. Y. Tian0551( HSNU);  Quzhoucity, 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. Zhuet al. 20190414-3( HSNU);  Hangzhoucity, Chunan County, Thousand Island Lake, Mushan Island, 29°31′ N, 119°4′ E,  242 m, on soil,  29 Oct. 2006,  Y. F. Wang245( HSNU); ibid.,  21 Apr. 2007,  Q. Liu& Y. Q. Zhu494( HSNU); ibid.,  Q. Liu& Y. Q. Zhu387 ( HSNU);  Hangzhoucity, 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. Wanget 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. Wanget al. 73( HSNU); ibid., 30°19′12.79″ N, 119°26′38.38″ E,  329 m, on rock,  27 Apr. 2011, D. H. Wanget 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. Wanget al. 1209( HSNU); ibid., 30°22′11.89″ N, 119°25′10.56″ E,  856 m, on rock wall,  28 Apr. 2011, D. H. Wanget al. 1211( HSNU); ibid., 30°20′43.18″ N, 119°27′16.75″ E,  416 m,  24 Apr. 2023, on rock,  J. Wang& Q. Wu20230427-5( HSNU);  Jinhua City, Beishan, near the reservoir, 29°12′8″ N, 119°37′51″ E,  480 m, on rock,  7 Nov. 2020,  R. L. Zhu20201107-5 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. Zhu20190206-6( HSNU);  Shengzhou City, Guimen County, Jiexi Village, 29°21′18″ N, 120°36′26″ E,  371 m, on rock,  5 Apr. 2023,  R. L. Zhu20230405-10( HSNU); ibid., 29°21′13″ N, 120°36′49″ E,  400 m, on rock,  27 Apr. 2009,  R. L. Zhu20090427-1( 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. Maet al. 20220821-32( HTC);  Lishui City, Qingyuan County, Xiyan Village, 27°39′08.48″ N, 119°12′32.35″ E,  694 m, Onsoil,  20 Aug. 2022, X. Y. Maet al. 20220820-05( HTC);  Lishui City, Suichang County, Baimashan Forest Park, 28°38′20.68″ N, 119°10′00.63″ E,  947 m, Onstone,  4 Aug. 2016,  W. Sheng20180804-6( HTC); ibid.,  W. Sheng 2016080410-1( HTC); ibid., Baimashan Forest Park, 28°38′22.30″ N, 119°10′21.65″ E,  840 m, Onstone,  4 Aug. 2016,  W. Sheng2016080429-3( HTC);  Lishui City, Suichang County, Jiulongshan National Forest Park, on stone,  21 Aug. 2016,  W. Sheng2016083144-1( HTC);  Lishui City, Songyang County, Ruoliao Primeval Forest Scenic Area, 28°18′30.32″ N, 119°17′30.38″ E,  672 m, Onstone,  4 Aug. 2017.  W. Sheng20170804-19( HTC);  Longquan City, Madi Village, 28°07′44.5″ N, 119°1′17.99″ E,  410 m, Onsoil,  19 Dec. 2009,  R. L. Zhu20091219-16( HSNU);  Taizhou City, Tiantai County, Baxian Lake, 29°12′58″ N, 120°59′51″ E,  164 m, on stone,  25. Jun. 2023,  R. L. Zhu202300605-28( 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. Tang20210719-29( HTC);  Wenzhou City, Taishun County, Juxi Town, on the way from Zhikeng Villageto Hengkengyuan, 27°31′36.83″N, 120°06′49.06″ E,  744 m, on rock,  13 Apr. 2018,  Y. H. Wuet al. 20180413-138( HTC). 2019-10-16 Z, HSNU Dai China Qingyang County 116 30.5403 Zhubei Town 1 117.8567 Chizhou City 10 225 R0010-20191016, 09 1 Anhui 2020-10-13 S, W, A, HSNU Tu China Shitai County 394 30.0741 Dayan Town 1 117.5006 Chizhou City 10 225 R002-032 1 2009-10-28 Y, F, HSNU Wang China Dehua County 870 25.685555 Mt. Daiyun 21 118.217224 Quanzhou City 10 225 1 Fujian 2009-10-28 Y, F, HSNU Wang China Dehua County 870 25.685555 Mt. Daiyun 21 118.217224 Quanzhou City 10 225 1 Fujian [450,961,518,544] 2009-10-28 Y, F, HSNU Wang China Dehua County 870 25.685555 Mt. Daiyun 21 118.217224 Quanzhou City 10 225 1 Fujian 2009-11-02 Y, F, W, HSNU Wu China Dehua County 830 25.683332 Chishui Town 21 118.14139 Mt. Daiyun 10 225 1 2010-04-05 Y, F, W, HSNU Wu China Dehua County 561 25.665596 Leifeng Town 1 118.289055 Mt. Daiyun 10 225 1 2015-08-30 Y, F, HSNU Wang China Xingcun County 1222 27.735243 Tongmu Village 1 117.63823 Mt. Wuyi National Nature Reserve 10 225 1 [246,1448,698,727] 2015-09-01 Y, F, HSNU Wang China Xingcun County 572 27.726109 Tongmu Village 1 117.714096 Mt. Wuyi National Nature Reserve 10 225 1 [136,1257,734,761] 1990-10-15 G, Q, IBSC Zeng China 1000 Baishi Town Lechang City 10 225 1 Guangdong 2019-05-19 Q, M, IBK Tang China Leye County 1755 24.7475 Yachang Wild Orchids National Nature Reserve 1 106.33908 Baise City 10 225 1 Guangxi [863,1290,806,833] 2019-05-19 Q, M, IBK Tang China Leye County 1755 24.7475 Yachang Wild Orchids National Nature Reserve 1 106.33908 Baise City 10 225 1 Guangxi 2019-05-20 Q, M, IBK Tang China Leye County 1643 24.756853 Dapeng Tun 1 106.34099 Baise City 10 225 1 Guangxi 2023-03-01 Y, M, Q, IBK Tang China Longsheng Co. 208 25.809616 Longsheng town 1 109.97452 Guilin City 10 225 BRY0936 1 2023-03-01 Y, M, Q, IBK Tang China Jiangdi Township 358 25.878012 Tangjia Tun 1 110.215775 Jianxin Villiage 10 225 BRY1181 1 2022-06-25 Y, JZNC Liu China Xinyang City 90 31.945833 Lingshan Town 20 114.27361 Luoshan county 10 225 1 Henan [711,1039,1022,1048] 2022-06-25 Y, JZNC Liu China Xinyang City 90 31.945833 Lingshan Town 20 114.27361 Luoshan county 10 225 1 Henan 2016-04-09 Y, JZNC Liu China Shangcheng County Donghe Village Xinyang City 10 225 1 2018-06-18 Y, JZNC Liu China Shangcheng County Donghe Village Xinyang City 10 225 1 1997-07-18 S, CCNU X. Liu China Shennongjia Forestry District 1500 Linghekou Pingqian 10 225 1 Hubei [296,1447,1130,1157] 1999-07-15 G, Y, CCNU Chen China Tongshan County 450 Jiugongshan Xianning City 10 225 1 1974-08-27 S, CCNU, IBSC X. Liu & Liuyang City China Luotian County 300 Jiuzihe Town Huanggang City 10 225 1 Hunan 2020-10-30 Y, LBG Sun China Jingan County 1038 28.824202 Baofeng Town 1 115.06144 Yichun City 10 225 1 Jiangxi 2020-11-18 C, F, Y, LBG Chen China Jingan County 966 29.002771 Qingshan Village 1 115.273186 Shuikou Town 10 225 1 Jiangxi 2022-11-22 Y, LBG Sun China Jingan County 321 28.985195 Qingshan Village 1 115.27492 Shuikou Town 10 225 1 Jiangxi 2023-03-29 C, F, T, J, LBG China Pengze County 471 29.82765 Chen 1 116.63023 Jiujiang City 10 225 1 2023-06-05 C, F, Z, Y, LBG China Chongyi County 981 25.83006 Qiyunshan National Nature Reserve 1 113.976006 Ganzhou City 10 225 1 1988-07-13 Y, IFP Jia China Zhuanghe County Xianrendong National Nature Reserve Dalian City 10 225 1 Liaoning 2013-02-12 L, Y, KUN Yao China Zhenan County Maoergou Shangluo City 10 225 1 Shaanxi 1993-05 2016-07-12 1993-05 R, KUN, XJU, C, Y, HSNU Mamtimin Sulayman & Zhejiang & Quzhou & Tian Taiwan Haba County 1925 48.298218 Wuling National Forest Recreation Area 1 86.608315 City 10 225 1 Taichung 1993-05 2016-07-12 1993-05 C, Y, HSNU Tian Taiwan Haba County 1925 48.298218 Wuling National Forest Recreation Area 1 86.608315 City 10 225 1 Taichung [433,784,1742,1769] 1993-05 2016-07-12 1993-05 C, Y, HSNU Tian Taiwan Haba County 1925 48.298218 Wuling National Forest Recreation Area 1 86.608315 City 10 225 1 Taichung [799,1150,1742,1769] 1993-05 2016-07-12 1993-05 C, Y, HSNU Tian Taiwan Haba County 1925 48.298218 Wuling National Forest Recreation Area 1 86.608315 City 10 225 1 Taichung 2019-04-14 R, L, HSNU Zhu Taiwan Quzhou 408 29.314299 Qiangjiangyuan National Park 1 118.19387 Changhong county 10 225 1 2006-10-29 Y, F, HSNU Wang Taiwan Chunan County 242 29.516666 Thousand Island Lake 1226 119.066666 Hangzhou 10 225 1 2007-04-21 Q, Y, HSNU Zhu Taiwan Chunan County 242 29.516666 Liu 1226 119.066666 Hangzhou 10 225 1 [429,853,1886,1913] 2007-04-21 Q, Y, HSNU Zhu Taiwan Chunan County 242 29.516666 Liu 1226 119.066666 Hangzhou 10 225 1 2011-04-19 H, HSNU Wang Taiwan Hangzhou 349 30.32203 Tianmushan National Nature Reserve 1 119.44065 Linan District 10 225 1 [136,1382,1958,1987] 2011-04-19 H, HSNU Wang Taiwan Hangzhou 349 30.32203 Tianmushan National Nature Reserve 1 119.443 Linan District 10 225 1 2011-04-27 H, HSNU Wang Taiwan Hangzhou 329 30.32022 Tianmushan National Nature Reserve 1 119.44399 Linan District 10 225 1 2011-04-27 H, HSNU Wang Taiwan Hangzhou 329 30.32022 Tianmushan National Nature Reserve 1 119.44399 Linan District 10 225 1 2011-04-28 H, HSNU Wang Taiwan Hangzhou 911 30.369905 Tianmushan National Nature Reserve 1 119.41497 Linan District 10 225 1 [246,1447,2066,2095] 2011-04-28 H, HSNU Wang Taiwan Hangzhou 856 30.36997 Tianmushan National Nature Reserve 1 119.41959 Linan District 10 225 1 [136,1367,158,187] 2023-04-24 J, Q, HSNU Wu Taiwan Hangzhou 416 30.34533 Wang 1 119.45465 Linan District 11 226 1 2020-11-07 R, L, B, HSNU Zhu Taiwan 480 29.202223 Beishan 20 119.63084 Jinhua City 11 226 1 2019-02-06 R, L, HSNU Zhu Taiwan Guimen County 308 29.357069 Shengzhou City 1 120.61196 11 226 1 2023-04-05 R, L, HSNU Zhu Taiwan Guimen County 371 29.355 Jiexi Village 20 120.60722 Shengzhou City 11 226 1 2009-04-27 R, L, HSNU Zhu Taiwan Guimen County 400 29.353611 Jiexi Village 20 120.61361 Shengzhou City 11 226 1 2022-08-21 Y, HTC Ma Taiwan Qingyuan County 795 27.663153 Xiyan Village 1 119.20522 Lishui City 11 226 1 2022-08-20 Y, HTC Ma Taiwan Qingyuan County 694 27.652355 Xiyan Village 1 119.208984 Lishui City 11 226 1 2016-08-04 W, HTC Sheng Taiwan Suichang County 947 28.639078 Baimashan Forest Park 1 119.16684 Lishui City 11 226 1 [225,644,482,508] 2016-08-04 W, HTC Sheng Taiwan Suichang County 947 28.639078 Baimashan Forest Park 1 119.16684 Lishui City 11 226 1 2016-08-04 W, HTC Sheng Taiwan Suichang County 840 28.639528 On 1 119.172676 Baimashan Forest Park 11 226 1 2016-08-21 W, HTC Sheng Taiwan Suichang County Jiulongshan National Forest Park Lishui City 11 226 1 2017-08-04 W, HTC Sheng Taiwan Songyang County 672 28.308422 Ruoliao Primeval Forest Scenic Area 1 119.29177 Lishui City 11 226 1 2009-12-19 R, L, HSNU Zhu Taiwan Longquan City 410 28.129028 On 1 119.02167 Madi Village 11 226 1 2023-06-25 R, L, HSNU Zhu Taiwan Tiantai County 164 29.216112 Baxian Lake 20 120.9975 Taizhou City 11 226 1 2021-07-19 Y, S, F, HTC Tang Taiwan Tiantai County 526 29.198408 Canghua Village 1 121.15815 Taizhou City 11 226 1 2018-04-13 Y, H, HTC Wu Taiwan Taishun County 744 27.526897 Juxi Town 1 120.113625 Wenzhou City 11 226 1