The genus Horikawaella S. Hatt. & Amakawa in China

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1 Nova Hedwigia 2013 Vol. 96 issue 3 4, published online December 07, 2012 Article The genus Horikawaella S. Hatt. & Amakawa in China Wei Li 1, Jiří Váňa 2 and Chien Gao 1 1 Institute of Applied Ecology, Chinese Academy of Sciences, Shenyang, , China; liw@iae.ac.cn 2 Department of Botany, Charles University, Benátská 2, Praha 2, Czech Republic; jiri.vana@natur.cuni.cz With 3 figures Abstract: The genus Horikawaella is rather common in the Himalayan region. Of its three described species, H. grosse-verrucosa is reported here for the first time from China. H. rotundifolia is found to be a new synonym of Mylia taylorii. Key words: Horikawaella rotundifolia; Horikawaella grosse-verrucosa; Mylia taylorii; Marchantiophyta, China. Introduction The genus Horikawaella S.Hatt. & Amakawa was described by Hattori & Amakawa (1971) for the species Anastrophyllum subacutum Herzog (Herzog 1939) [= Horikawaella subacuta (Herzog) S.Hatt. et Amakawa]. This species is rather common in the Himalayan region. The second species, H. grosse-verrucosa Amakawa & S.Hatt., distributed in Eastern Nepal (Eastern Himalaya), was published by Amakawa (1975). Recently we found it also in Yunnan (China). Yi et al. (1998) added the third species Horikawaella rotundifolia C.Gao et Y.-J.Yi on the basis of a sterile collection from Yunnan (China). While, examining the type specimen of H. rotundifolia and some other specimens from adjacent regions, we found that the species should be reduced to a new synonymy of Mylia taylorii (Hook.) Gray J. Cramer in Gebr. Borntraeger Verlagsbuchhandlung, Stuttgart, Germany. DOI: / /2012/ /2012/0083 $

2 Results and discussion Mylia taylorii (Hook.) Gray, Nat. Arr. Brit. Pl. 1: Schuster, Hep. Anth. North Amer. 2: 1042, fig. 300(1 7), 301, 1969; Gao et Chang, Flora Hepaticarum Chinae Boreali-Orientalis. 84, 1981; Piippo, Journ. Hattori Bot. Lab. 68: Synonymies: Jungermannia taylorii Hook., Brit. Jungerm. Tab Aplozia taylorii (Hook.) Dumort., Rec. d Observ Leptoscyphus taylorii (Hook.) Mitt., Journ. Bot. Kew Garden Miscel. 3: Coleochila taylori (Hook.) Dumort., Hep. Eur Type: Ireland, County Wicklow, Toulagee, leg. T. Taylor (lectotype: BM!). Distr.: Europe, North America, northern Asia to Himalaya. Jungermannia reticulato-papillata Steph., Mem. Soc. Nat. & Math. Cherbourg. 29: Type: China, Yunnan, Tsung-Yang-Tschang, May 24, 1899, leg. Delavay (lectotype: G!) Horikawaella rotundifolia C.Gao et Y.J.Yi, Acta Phytotax. Sinica 36(3): 284, 1998, syn. nov. Type: China, Yunnan, Dulongjiang, leg. Zang Mu 611 (holotype: KUN!; isotype: IFP!). Distribution in China: yunnan, XiZanG (tibet), GuanGXi, LiaoninG, JiLin, heilongjiang. CharaCters of M. taylorii are: (1) PLants green or reddish purple, coarse plates covered the cell surface of the whole plant. (2) underleaves lanceolate. (3) Perianth weakly compressed below but strongly compressed laterally and flattened above, slightly narrowed to the relatively wide mouth, in shape similar to Plagiochilaceae, and perianth surface with cell surface like that of leaves, with or without obvious protuberances formed by irregular and coarsely polyhedral plates. The occurrence of mentioned protuberances on the perianth cell surface is not constant (also in one specimen) and it was never recognized and described in the populations from areas outside of Himalayan region, such as in the description of Müller ( ) and Schuster (1969). M. taylorii may be confused with M. verrucosa Lindb. by protuberance occurring, but if samples are examined under microscope, protuberance of M. taylorii is the protuberant coarse plates on cell walls, while in the latter one, protuberance is obviously a line containing two or three cells. other specimens examined: CHINA: YUNNAN, Co. Lijiang, 3700 m, S.G.Wu 167B (KUN, IFP), Co. Gongshan, Dulongjiang River, m, A.L.Zhang 31(b), M.Zang 611, 969 (KUN, IFP), Dulong Xiang, W slope of Gaoligong Shan, Irrawadi catchment, Qiqi trail between San Dui and Xishaofang, open hillside with old dead Abies trees and young Abies/Rhododendron scrub, in damp rocky hollow amongst boulders, c m, 1 Nov 2004, D.G.Long (KUN), Bingzhongluo Xiang, E slope of Gaoligong Shan, Nu Jiang (Salween) catchment, ridge above SE end of lake at head of Nianwaluo He valley below Gawagapu Mountain, alpine hillside with scattered Abies trees, shrubs, herb meadows and metamorphic rock outcrops, on side of mossy boulder amongst shrubs, c m, 20 Aug 2006, D.G.Long (KUN), Bingzhongluo Xiang, E slope of Gaoligong Shan, Nu Jiang (Salween) catchment, valley on SW slope of Gawagapu Mountain, U-shaped alpine valley snow patches, scree, scrub slopes, flushes and rock outcrops, on wall of boulder on stony slope, c m, 20 Aug 2006, D.G.Long (KUN), Bingzhongluo Xiang, E slope of Gaoligong Shan, Nu Jiang (Salween) catchment, ridge above SE end of lake at head of Nianwaluo He valley below Gawagapu Mountain, slopes with degraded Abies/ Rhododendron forest, on rotten log, c m, 22 Aug 2006, D.G.Long (KUN), Cikai Xiang, E slope of ridge crest Gaoligong Shan (Nu Jiang catchment), Qiqi trail at pass to Dulong, exposed alpine ridge crest with rock outcrops, hollows and dense Juniperus/ Rhododendron scrub, on branch of dwarf Rhododendron, c m, 29 Sep 2007, D.G.Long & J.Shevock (KUN), Lumadeng Xiang, Yaping Cun, E slope of Gaoligong Shan (Nu Jiang catchment), E side of Burma/ Yunnan border ridge just south of Yaping Pass, exposed alpine 526

3 Fig 1. Mylia taylorii. A, B. Dorsal and ventral view of plant. C, D. Leaves. E, F. Underleaves. G. Leaf cells and coarse plate on leaf. H, I, L P: Female plants with perianth. J. Female plant. K. Ventral view of plant, showing the underleaves. [A D drawn from M.Zang 611, E K from A.L.Zhang 31(b), I from M. Zang 10156, L, M from D.G.Long & J.Shevock 37234, N from D.G.Long 35895, O from G. & S. Miehe , P, Q from G. & S.Miehe a: A, B; b: C, D; c: E, F; d: G; e: H; f: J, g: K; h: I, L, M, N; i: O, P, Q] (Drawn by W.Li). ridge with rock outcrops, dwarf Rhododendron and bamboo, in liverwort turf amongst rocks under dwarf Rhododendrons, c m, 5 Oct 2007, D.G.Long & J.Shevock (KUN), Chang-shan submerged on granite, Y.G.Su 1388 (KUN, IFP), Co. Weixi, E slope of Mt. Biluo, on rock, 2150 m, M.Z.Wang 5410A-C (PE), Co. Zhongdian, Bigu, on trunk of Abies, 3500 m, 28 Jul 1986, M.Zang (KUN); XIZANG (TIBET), Co. Motuo, 4450 m, on rock, Y.G.Su 1644 (IFP), on S slope of 527

4 Mt. Duoxiongla, 3600 m, M.Z. Wang d (PE), Co. Chayu, Ridong, Mt. Molongla, m, M.Z. Wang a, a (PE), on the N slope of Mt. Songtashan, m, M.Z.Wang 8504, 8665, 8671 (PE), on the E slope of Mt. Qimala, in the forest of Abies and Picea, m, M.Z. Wang a (PE). BHUTAN: Bumthang: E of Nada La, Rhododendron thicket on NE-facing slope, 4320 m, 15 Oct 1998, G. & S.Miehe (PRC), E of Nada La, Rhododendron thicket on N-facing slope, 4300 m, 15 Oct 1998, G. & S.Miehe (PRC). The genus Horikawaella is characterized by: (1) PLants erect or suberect, 1 3 cm long, robust, usually purplish. (2) Leaves cordate to ovate, subtransversely inserted, usually conduplicate-concave, covered by conspicuous verrucae, apex narrowly obtuse to subacute, ventral margin somewhat recurved, trigones usually nodulose. (3) rhizoids sparse, long, hyaline or reddish. (4) DioiCous; androecia terminal, bracts in 3 5 pairs, ventricose; perianth semi-exserted, fusiform, usually 4-plicate, bracts in 2 pairs. Key to Horikawaella species 1 Plants large, robust, yellowish-green or reddish-purple, in dense mats, up to 2 3 cm long, cm wide with leaves. Branches intercalary, cortex covered by distinct verrucae. Rhizoids rare, sometimes reddish. Leaves imbricate, incubous, nearly transverse, conduplicate concave, leaf base slightly ventral decurrent, cordate with abrupt subacute apices when flattened, mm long, mm wide. Underleaves absent. Cells along leaf margin μm, median leaf cells μm μm, basal leaf cells rectangular, μm μm, trigones large, usually nodulose. Dioicous. Male inflorescence not seen. Gynoecium terminal, perianth fusiform, 1.5 mm long, 0.6 mm wide, bracts 2 pairs, similar to stem leaves. Spores not seen...1. H. subacuta 1' Plants small to robust, reddish-purple, in dense mats, ca. 1.5 cm long, ca. 0.5 mm wide with leaves. Stems ascending, cortex covered by distinct verrucae. Rhizoids rare, long, colorless. Leaves slightly incubously inserted, nearly transverse, conduplicate concave, ovate when flattened, apex obtuse, leaf base somewhat ventrally decurrent mm long, mm wide. Underleaves absent. Cells along leaf margin μm μm, median leaf cells μm μm, basal leaf cells 9 15 μm μm, trigones large, sometimes nodulose. Dioicous. Male inflorescence terminal or intercalary, bracts about 3-5 pairs. Gynoecium terminal, perianth fusiform, about 1.1 mm long, 0.5 mm wide, bracts 2 pairs, similar to stem leaves. Spores not seen...2. H. grosse-verrucosa 1. Horikawaella subacuta (Herzog) S.Hatt. & Amakawa, Miscel. Bryol. Lichenol. 5: ; Flora Eastern Himalaya ; Acta Bot. Yunnan. 24(3): (Fig. 1) basionym: Anastrophyllum subacutum Herzog, Ann. Bryol. 12: type: Sikkim Himalaya, Tsomgo Lake, zwischen Gangtok und Natu La, m, Aug. 1937, leg. C.Troll (holotype: JE!). Distribution: NEPAL (Grolle 1966, Hattori & Amakawa 1971, Hattori 1975, Long & Grolle 1990, Miehe 1990, Váňa & Long 2009). INDIA, SIKKIM (Herzog 1939, Grolle 1964, 1966, Hattori & Amakawa 1971, Hattori 1975, Long & Grolle 1990, Váňa & Long 2009), Darjeeling (Long & Grolle 1990, Váňa & Long 2009). BHUTAN (Long & Grolle 1990, Váňa & Long 2009). CHINA (Sun et al. 2002). other specimens examined: CHINA: XIZANG (TIBET), Co. Milin, W slope of Duoxiongla, Y.G.Su 5038(KUN, IFP), Co. Ridong, Bulaolong, 3780 m, M.Zang 5206 (KUN, IFP); YUNNAN, Co. Gongshan, Dulongjiang River, 3200 m, M.Zang 946(KUN, IFP), Co. Gongshan, Dulongjiang River, 3500 m, M.Zang 969 (KUN, IFP), Co. Gongshan, E slope of Gaoligong Shan, Nu Jiang catchment, 528

5 Fig. 2. Horikawaella subacuta. A. Female plant, ventral view. B. Perianth. C. Stem leaf, ventral view. D. Stem leaf, dorsal view. E. Basal leaf cells. F. Apical leaf cells. [A, C F drawn from M.Z.Wang 9451C, B from T.Herzog (1939). a: A, B; b: C, D; c: E; d: F.] (Drawn by W.Li). Yipisaka Lake at head of Pula He Valley 2.2 km SSE of tunnel, granitic cliff on alpine lake shore; on rock wall of deep shady cleft on cliff, 3455 m, D.G.Long 35736( KUN), Co. Bijiang, Mt. Gaoligongshan, M.Zang 5906 (KUN, IFP), Co. Gongshan, Qiujiang, on rock, 3300 m, Q.W.Wang 6926 (PE,IFP), Site 45, Co. Gongshan, Dulong Xiang, W slope of Gaoligong Shan, Irrawadi catchment, Qiqi trail between San Dui and Xishaofang, degraded Rhododendron/ Bamboo scrub with old dead Abies trunks and young regenerating Abies, on mossy shady rock outcrop, c m, 1 Nov 2004, D.G.Long (KUN), Site 06-11, Co. Gongshan, Bingzhongluo Xiang, E slope of Gaoligong Shan, Nu Jiang (Salween) catchment, W end of lake at head of Nianwaluo He valley below Gawagapu Mountain, alpine lake shore with metamorphic rock outcrops, peaty flats and small inflow stream; on shady rock outcrop by stream, c m, 18 Aug 2006, D.G.Long (KUN), Site 38, Co. Gongshan, Dulong Xiang, W slope of Gaoligong Shan, Irrawadi catchment, Qiqitrail above San Dui between Shu Gung Qiao (Stone Arch Bridge) and Xishaofang, large ravine at margin of Tsuga/Rhododendron forest with mossy rock outcrops and scrub on banks, on wet mossy rock 529

6 Fig. 3. Horikawaella grosse-verrucosa. A. Male plant, dorsal view. B. Male plant, ventral view. C. Female plant. D. Basal leaf cells. E. Apical leaf cells. F. Stem leaf, dorsal view. G. Stem leaf, ventral view. [A, B, D G drawn from M.Zang 969, C from Hattori (1975). a: C; b: D, E; c: A, B; d: F, G] (Drawn by W.Li). ledges of cliff, c m, 30 Oct 2004, D.G.Long (KUN), Site-07-14, Co. Gongshan, Cikai Xiang, E slope of Gaoligong Shan (Nu Jiang catchment), Qiqi trail above Dengxiaofang. Valley slope with cleared Abies/Rhododendron forest, bamboo and scrub, on small N-facing rock outcrop above stream, c m, 27 Sep 2007, D.G.Long & J.Shevock (KUN), Co. Gongshan, Mt. Gaoligongshan, W slope, m, M.Z.Wang 9451C (PE), Co. Gangshan, Qiannatong, on soil, 3000 m, Q.W.Wang 6383 (PE), Co. Gongshan, Qiujiang, on rock, 3300 m, Q.W.Wang 6908 (PE), Co. Gongshan, Qiujiang, on rock, 3300 m, Q.W.Wang 6926 (PE). 2. Horikawaella grosse-verrucosa Amakawa & S.Hatt., Flora Eastern Himalaya type: Nepal, around Kipuphu, 4550 m, Jun 26, 1972, coll. Z.Iwatsuki, 1699 (holotype: NICH !). Distribution: NEPAL (Hattori 1975, Long 2005, Váňa & Long 2009), new to CHINA. other specimens examined: CHINA: YUNNAN, Co. Changshan, Y.G.Su 1388 (KUN, IFP), Co. Gongshan, Dulong River, 3500 m, M.Zang 969 (IFP, KUN), Bingzhongluo Xiang, E slope of 530

7 Gaoligong Shan, Nu Jiang (Salween) catchment, valley on SW slope of Gawagapu Mountain, c m, 20 Aug 2006, D.G.Long (KUN). Acknowledgements We appreciate the National Natural Science Foundation of China (Project: ) and Chinese Academy of Sciences for support of the present research. We acknowledge to Herbarium of Kunming Institute of Botany, Chinese Academy of Sciences (KUN), Herbarium of Institute of Botany, Chinese Academy of Sciences (PE), Herbarium Haussknecht of Friedrich-Schiller-Universität Jena (JE), Natural History Museum (BM) and The Hattori Botanical Laboratory (NICH) for providing specimens for our study. References AMAKAWA, T. 1975: Jungermanniaceae. In OHASHI, H. (ed.): The Flora of Eastern Himalaya (Third report). Bull. Univ. Mus. Univ. Tokyo 8: GROLLE, R. 1964: Miscellanea hepaticologica (11 20). J. Jap. Bot. 39: GROLLE, R. 1966: Die Lebermoose Nepals. Ergeb. Forsch.-Unternehm. Nepal Himalaya 1(4): HATTORI, S. 1975: Anthocerotae and Hepticae. In: Ohashi H, The Flora of Eastern Himalaya (Third Report). Bull. Univ. Mus. Univ. Tokyo 8: HATTORI, S. & T. AMAKAWA 1971: A new genus Horikawaella (Jungermanniaceae; Hepaticae) from Nepal. Misc. Bryol. Lichenol. 5: HERZOG, T. 1939: Zwei Bryophytensammlungen aus dem Sikkim-Himalaya. Ann. Bryol. 12: LONG, D.G. & R. GROLLE 1990: Hepaticae of Bhutan II. J. Hattori Bot. Lab. 68: LONG, D.G. 2005: Notes on Himalayan Hepaticae 2: New records and extensions of range for some Himalayan Leafy Liverworts. Crypt. Bryol. 26: MIEHE, G. 1990: Langtang Himal. Flora und Vegetation als Klimanzeiger und -zeugen im Himalaya. Diss. Bot. 158: MÜLLER, K : Die Lebermoose Europas. In: RABENHORTS s Kryptogamen-Flora Vol. 6, Teil 2. Geest & Portig, Leipzig. SCHUSTER, R.M. 1969: Mylia S.F. Gray emend. Carr. In: SCHUSTER, R.M.: The Hepaticae and Anthocerotae of North America Vol. 2: Columbia Univ. Press, New York. SUN, J., T. CAO & C. GAO. 2002: A liverwort species of Horikawaella (Hepaticae) is found in China. Acta Bot. Yunnan. 24(3): VAŇA, J. & D.G. LONG 2009: Jungermanniaceae of the Sino-Himalayan region. Nova Hedwigia 89: YI, Y.-J., J.-Y. LIU & C. GAO. 1998: Horikawaella (Hepaticae), new to China with a new species from Yunnan Province. Acta Phytot. Sinica 36(3): Manuscript received November 20, 2011; accepted August 10,