J Hattori Bot. Lab. No. 94: 159-177 (Aug. 2003)

THE (HEPATICAE) IN ASIA

M. L. SOl

ABSTRACT. Ten of Metzgeria are represented in Asia: M. consanguinea Schiffn., M. crassip­ ilus (Lindb.) A. Evans, M. Joliicola Schiffn., M. Jurcata CL.) Dumort. , M. leptoneura Spruce, M. lind­ bergii Schiffn., M. pubescens (Schrank.) Raddi, M. robinsonii Steph. , M. scobina Mitt., and M. tem­ perata Kuwah. Twenty synonyms are proposed and a key to the species in Asia is provided. Metzge­ ria albinea Spruce and M. vivipara A. Evans are excluded from this region. K EY WORDs: Hepaticae, Metzgeria, Asia.

INTRODUCTION Metzgeria is a genus with about 240 binomials listed in the Index Hepaticarum (Geissler & Bischler 1985). Kuwahara (1986, 1987) in his life long study had created quite a number of new species from Asia, and the Neotropics. But as noted by Schuster (1992), the number of species may probably be grossly inflated and since there is not yet a world monograph of this genus, the total number of species could be drastically reduced. Most of the specimens examined in the present study include recent collections from China, India, Nepal, the Philippines, Malaysia, and . A total of 52 species names have been recorded from this region, and of the 36 species recognized by Kuwahara (1986), only 10 are accepted here. Twenty new synonyms are proposed. M etzgeria albinea Spruce and M . vivipara A. Evans are excluded from this region. For some unknown reasons, holo­ types (and even isotypes) of many species described by Kuwahara were not present in vari­ ous herbaria where they were supposedly deposited, and, at most only the isotypes were ex­ amined. Due to the small number of species recognized in this region, the many sections and subsections created by Kuwahara (1978b) are not adopted here.

TAXONOMIC TREATMENT Metzgeria Raddi, Jungermanniogr. Etrusca: 34. 1818. Mem. Mat. Fis. Sci. Ital. Sci. Mode­ na, Pt. Mem. Fis. 18 : 45. " 1818"1820. Lectotype, designated by Kuwahara 1966: M. glabra Raddi nom. illeg. (= M.furcata (L.) Dumort.)

Key to species of Metzgeria in Asia 1. Thallus with hairs on both dorsal and ventral surfaces ...... 2 2. Dorsal surface with hairs on costa only ...... M. Joliicola 2. Dorsal surface very hairy ...... 3 3. Both surfaces very hairy; costa dorsally of7- 1O cells, ventrally of7- 1O cells ...... M. pubescens 3. Dorsal surface very hairy, hairy only on ventral costa; costa dorsally of 2 cells, ventrally of 2 cells ...... M. robinsonii

1 Biology Department, Hong Kong Baptist University, 224 Waterloo Road, Hong Kong. 160 1. Hattori Bot. Lab. No. 94 2 003

I. Thallus with hairs on ventral surface only ...... 4 4. Gemmae on dorsal surface ...... M. erassipilus 4. Gemmae, if present, on margin of thallus ...... 5 5. Autoecious ...... M. lindbergii 5. Dioecious ...... 6 6. Each laminal cell with a large papilla ...... M. seobina 6. Each laminal cell without a large papilla ...... 7 7. Hairs weakly to distinctly falcate, mostly paired ...... M. leptoneura 7. Hairs not falcate, paired and/or single ...... 8 8. Thallus with obtuse apex ...... M. fureata 8. Thallus often with tapered apex ...... 9 9. Lamina 13- 28(40) cells wide; median laminal cells 25-43(50)X 18- 30(35) .urn; gemmae produced at apex and margin of thallus ...... M. eonsanguinea 9. Lamina 11 - 18(26) cells wide; median laminal cells 33- 58(70)X 25-43 .urn; gemmae produced at margin of thallus ...... M. temperata 1. Metzgeria consanguinea Schiffn., Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur. 60 (2): 271 . 1893.- Type: Indonesia. Java: "in summo apice montis Pangerango," 20 Feb 1890, G. Karsten s.n. (holotype: FH!; isotypes: BM!, FH!, G-00I087!, S[B37897]!, W-9093 !). Metzgeria longilexla Steph., Sp. Hepat. 6: 54. 1917; nom illeg. (Art. 52.1 ).- Original material: India. Sikkim: Himalaya, Darjeeling, Re'ang, 650 ft, 8 Nov 1907, Sal Dhoj 28, pro E. Long, herb. Levier 5658 (BM! , G-000095!). Metzgeria sinensis Chen, Fedders Repert. Spec. Nov. Regni Veg. 58: 38. 1955 ; fide Zhang 1991.­ Type : China. "Prov. Szetschwan, Nan-tschwan, Ching-fu-schan, ... Aug 1945, C. C. Jao 7" (holo­ type: PE). Melzgeria harae Kuwah. , J. Hattori Bot. Lab. 28 : 296. 1965; syn. nov.- Type: Nepal. Walunchung Gola-Zongi, 2700- 3800 m, 11 Nov 1963, H. Kanai et al. 236481 (holotype: not located). Melzgeria aeula Kuwah. , Rev. Bryol. Licheenol. 36: 531. '1969' 1970; fide So 2002.- Type : Philip­ pines. Luzon Is: Mountain Province, Mt Pulog, 2700-2900 m, Pro! & Mrs A. 1. Sharp and Z. Iwatsuki s.n. (holotype: NICH-267429 [not located]). Metzgeria maerospora Kuwah., J. Hattori Bot. Lab. 32: 17. 1969; syn. nov.- Type: India. Sikkim: Migothang-Nayathang, 3300m, on twig of tree, I June 1960, M. Togashi s.n. (holotype: NICH- 20 1680 [not located]). Metzgeria darjeelingensis Kuwah. , 1. Hattori Bot. Lab. 39: 365. 1975; syn. nov.-Type: India. West Bengal: Darjeeling area, Tongloo, around Dak Bangalow, alt. ca. 1000 ft , on branch, light-open, mesophilous, Pro! & Mrs A. 1. Sharp and Z. Iwatsuki s.n. (holotype: NICH-258622 [not locat­ ed]). Melzgeria macrocellulosa Kuwah ., 1. Hattori Bot. Lab. 39: 367. 1975 ; syn. nov.- Type : India. West Bengal: Ranbi forest near Darjeeling, Darjeeling area, alto ca. 7400 ft, on cliff, Pro! & Mrs A. 1. Sharp and Z. 1walsuki S.n. (holotype: NICH-258076 [not located]). Melzgeria nilgiriensis Srivastava & Udar, New Bot. 2: 24. 1975; syn. nov.-Type : India. Government Botanic Garden, Ootacamund (Nilgiri hills), 7000-7500 ft, 22 Oct 1962, R. Udar & V. Chandra 182162 (holotype: LWU!).

Illustrations: Kuwahara (1965a: fig. I: h- n as M. harae; 1969b: fig. 1 as M. aeula; 1969a: fig. M. L. So: The genus Metzgeria in Asia 161 a- o as M. macrospora; 1975 : fig. I as M. consanguinea; fig. II as M. darjeelingensis, fig. IlI, IV as M. macrocellulosa); Srivastava & Udar 1975: fig. IX, X, XI as M. nilgiriensis); Stephani (1985: Icon. n. 7646 as M. cuspidata Steph. nom. in sched. , n. 7664 as M. longitexta). Descriptions: Kuwahara (1966: 232 as M. consanguinea; 1975: 365 as M. darjeelingensis). Distribution: Worldwide. Specimens examined: Bhutan. Forested slope between Sengor and Sheridrang, NW of Mongar, 2980m, 1979, D. G. Long 8631 (E). China. Taiwan: Northern part, 24 Oct 1899, K. Miyake 187 (G- 001080). Jiangxi : Shanqingshan, 1550m, 1987, X M. Shao 256,296, 445, 1059 (HSNU). Yunnan: Yulongshan, small valley S of Ha Lan Gon Lake, Wen Hai Mountain, 3200 rnn, 1990, D. G. Long 18990 (E). Zhejiang: Wuyangleng, 1290m, 1987, R. L. Zhu 294,313, 452 (HSNU); July 2001 , R. L. Zhu 72 1-10 (HSNU). India. North District: below Takche, N of Lachung, gully in scrubby cleared Tsuga forest, on trunk, 3040 m, 1996, D. G. Long 26476 (E); 1923, R. P Foreau s.n. (E). Indonesia. Java: Buitenzorg, 1897, E. Nyman S.n. (S); Tjibodas, Mt Gedeh, July 1898, M. Flesicher s.n., herb. Levier 4139 (G-001089); Geger Bentang, 1520m, 1949, Noerta & Soekar 1010, 102 7 (L); Korndang Badok, 1895, J. Massart 1731 (FH); Pangerango, 2985 m, 1894, V. Schiffner 328 (FH, G-OO I 088); Priagan, G. Gede, 2150m, 1930, F Verdoorn 2809 (NY). Lesser Sunda Islands: Flores, Prov. Mang­ garai, Ruteng, Lake Mese, 1230m, 1988, Fr. E. Schmutz SVD 7093 (L); Manggarai Prov., N slope of Ngando Napu, S of Ruteng, 1780 m, 1988, A. Touw & M. Snoek 23237 CL). Sumatra: 1952, W. v.d. Wijk 1808 (L). Japan. Shikoku: Tosa, Mt Yokugura, Aug 1980, S. Okamura 380 as M. cuspidata (G); Omata-dani, Yatsuo-cho, Nei-gun, Toyama-ken, 1974, M. Mizutani s.n. (Hattori: Hepat. Jap. Exs. ser. 20, no. 986) (G, M, S). Honshu: Mt Oyama, Is Miyake, lzu lsls, 700 m, 1976, H. Inoue s.n. (Inoue: Bryoph. Se\. Exs. no. 364) (EGR, G, MELU, S); Gifu Pref., Mt Gozenyama, on rock, 900 m, 1970, T. Magohuku s.n. (S); Yezo, Mt Shiubeshi, 1905, U Faurie 1822 (BM, FH); Yamanashi Pref., Mt Fuji, Shokin-guchi-tozan-dou, 1450- 1600 m, on trunk, 1971 , M. Shimizu 260 (NY). Kyushu: Fukuoka Pref., Mt Shaka, on bark, 1000 m, 1970, T. Amakawa 6999 (NY). Nepal. N-facing slopes of Dobala Danda above Kabeli Khola, ravine in wet mossy oak/laurel forest, 2350 m, 1989, D. G. Long 17329, 17365 (E); Rasuwa District, near Sing Gompa, on trunk of Rhododendron barbatum, 3320m, 1992, D. G. Long 22174 (E); near Gupha Pokharu, by stream in open degraded Rhododendron forest, 2920 m, 1989, D. G. Long 17541 (E); Sankhuwasabha District, Pemthang Kharka, S side of Barun Khola, 341Om, 1991 , D. G. Long 20747 (E); E Nepal, 5th Bot. Exped. to E Himalaya by Univ. of Tokyo, in a mossy forest below Lamo Pokhhari (pond), on soil, 2600-2800 m, 1972, Iwatsuki 617 as M. maerospora (JE, TNS). Philippines. Luzon: Semper S.n. (FH). Vietnam. Vinh-Phuc Province: Tam-Dao Mts, 950- 1330 m, 18 Nov 1998, T. Poes & T. Ninh 9896/AL (EGR). Metzgeria consanguinea is here treated in a broad sense to include species with ta­ pered and obtuse apices and gemmae are usually produced around the tapered apices as well as along the margins. However, gemmae may be lacking in some collections. The ta­ pered apices may vary in length, and are sometimes tinged with blue at the very tip. Several species of Metzgeria with tapered thalli CM. darjeelingensis, M . macrocellu­ /osa, M. macrospora, and M. harae) from the Himalayan region were created by Kuwahara (1969, 1975). These species all have tapered and obtuse forms, producing marginal and apical gemmae, with a lamina of 12- 34 cells and costal cells of 2- 3 dorsally and 2-3(-4) ventrally. The distinctions between these species are extremely narrow and fall within the morphological variations of M. consanguinea, which is widespread in ElSE Asia. These species, together with M. nilgiriensis from India, are all placed under its synonymy. 2. Metzgeria crassipilus (Lindb.) A. Evans, Rhodora 11: 188. 1909. Metzgeriafurcata 162 1. Hattori Bot. Lab. No. 94 2 003

Fig. I. Photomicrographs of Metzgeria. Metzgeria robinsonii Steph. a, b, dorsal sur­ face showing hairs. Metzgeria fo liicola Schiffn. c: dorsal surface of thallus, showing epiphyl­ lous habit and hairs on dorsal costa, d: dorsal surface of thallus showing gemmae and hairs on some portions of the thallus. (a, b from holotype in G; c from holotype of M. cristata Steph. in G; d from holotype of M. hispidissima Steph. in G). All X20.

ssp. crassipilis Lindb., Acta Soc., Fauna Fl. Fenn. 1(2): 42. '1877' I 878.- Lectotype, designated by Schuster 1992: U.S.A. "Pennsilvania, Laurel Hill", 23 Jun 1843, W S. Sullivant s.n. (not located). Fig. 2: a, b Metzgeria novicrassipilis Kuwah., 1. Hattori Bot. Lab. 20: 138. 1958; fide Kuwahara 1984.-Type: Japan. Fukuoka: Mt Hiko-san, ca. 700 m, Y. Kuwahara 6120 (holotype: NICH [not located]). Metzgeria indica Udar & Srivastava, Rev. Bryol. Lichenol. 37: 361. ' 1970' 1971 ; syn. nov.- Type: India. Kodaikanal, Coaker's Walk, Palni Hills, ca. 6500 ft , 4 Jan 1966, R. Udar & s. C. Srivasta­ va 53/66 (holotype: LWU!). Metzgeria pandei Srivastava & Udar, New Bot. 2: 16. 1975; syn. nov.- Type: India. Kodaikanal , Palni Hills, 6500 ft, Bryant Park, 18 Oct 1962, R. Uda r & V Chandra 52/62 (holotype: LWU!).

Illustrations: Kuwahara (1958: fig. VII: 1- 3 as M. novicrassipi/us); Srivastava & Udar (1975: fig. V, VI as M. pandei). Description: Kuwahara (1958: 138 as M. novicrassipi/us). Distribution: Worldwide. M. L. So: The genus Melzgeria in Asia 163

Fig. 2. SEM micrographs of laminal cells of Metzgeria. Metzgeria crassipilus. (Lindb.) . Evans. a, b: dorsal surface showing gemmae. Metzgeria pubescens (Schrank) Raddi. c: ventral surface and margin, showing hairs, d: laminal cells, e: dorsal surface, show­ ing hairs, f: ventral surface. (a, b from Glenny 10481 in WELT; c-ffrom J Dransfield 1910 in L).

Specimens examined: China. Sichuan: Mt Omei, SW of Chengdu, 650-950 m, 1980, A. Touw 23823 (L). India. India-orient., Sudaion Mt, Pfiederer K8 as M. caledonica Steph. (G-000947). In­ donesia. Java: Kampong Rarakan, 1300m, 1949, Noerta 810,814 (L); Kampong Dawoean, 1360m, 1949, Noerta & Soekar 913, 925 (L). Sumatra: Fort de Kok, 1899, K. Geisenhagen 122 (M). Japan. Kyushui: Kumamota Pref., Mt Ichifusa, Kumagun Kozakiyama, 600-1000m, 1973, r Magofuku 9121 (S). Sri Lanka. Poondeloya, 4-8000 ft , 1868, J Nietner 155, det. Muller as M. latifrons Steph. (EGR, G-009877, M) ; Nuwara Eliya District, Nuwara Eliya, 1850 m, 1976, M. Onraedt 76.L.2735 as M. vivipara (BR). 164 J. Hattori Bot. Lab. No. 94 2 0 0 3

Metzgeria crassipilus is the only species in this region with discoid or oval-shaped gemmae arising form the dorsal surface of the thallus. Gemmae, if present, in other species arise from the margin of the thallus. Features pertaining to M. indica, especially the pres­ ence of gemmae on the dorsal surface of the thallus, indicate that this species is conspecific with M. crassipilus as already suggested by Kuwahara (1984a: 278). 3. Metzgeriafoliicola Schiffn., Denkschr. Kaiser!' Akad. Wiss., Math.-Naturwiss. Kt. 67: 181. 1898. Sp. Hepat. 6: 51. 1917.-Lectotype, designated by So 2002: Indonesia. Java: Prov. Preanger, montis Pangerango; Tjibodas, 1640 m, 24 Apr 1894, Schiffner 320 (FH!; isolectotypes: G-00I024!, L!, M! , STR!, W!). Fig. I: c, cl, 3: a- c Metzgeria eristata Steph., Sp. Hepat. 6: 49. 1917; syn. nov.- Type: Indonesia. Java: Tjibodas, epi­ phyllous, E. Nyman s. n. (holotype: G-001007!). Metzgeria eristatissima Herzog, Mem. Soc. Fauna F!. Fennica 26: 38. 1951 ; .fide Kuwahara \966.­ Type: Indonesia. Java: 'uf ei nem Farnblatt, ohne nahere Fundortsnotiz, 1930, F Verdoorn sub. no. 210' (holotype: JE! ; isotype [210] : JE!). Metzgeria dubia Herzog, Ann. Naturhist. Mus. Wien 1942 , 53: 361. 1943 ; nom. inva l. (Art. 32.1 (c).- Original material : Indonesia. Sumatra: Padang, F S. Child 8, herb. Herzog 18 (lE!). Illustrations: Herzog (1951: fig. 22 as M. eristatissima); Kuwahara (1966: fig. 20: a, b as M. foli­ ieola , 20 : e-i as M. eristata; 1978a: fig. I: I, 6: 2 as M. foliieola; Stephani (1985: Icon. n. 7644 as M. eristata). Distribution: China, Indonesia, Papau and Vanuatu. Specimens examined: China. Hunan: Yizhang Co., Mt Mangshan, Langpanhu Swamp, 1235 m, 1997 , T. Koponen, S. Huttunen & P. C. Rao 51183 as M. albinea (H). Taiwan: Chitou, on rock, 1200 m, 1979, Y. Yuzawa H 16688 (hb Yamada). Indonesia. Java: Prov. Preanger, montis Pangerango, Tjibodas, 1600 m, 24 Apr 1894, V. Sehiffner 319, 321 (FH; isosyntypes of M. foliieola); Tjiburrum, 2 May 1894, V. Sehitrner 323 (FH ; isosyntype). Lesser Sunda Islands: Flores, Ruteng, Poco Ranaka, epiphyte, 1950 m, 1975 , 1. F Veldkamp 7120C (L). Molucca: Ternate, slope of Mt Ternate, on wood, 1951 , P. Groenharl8016 (L). Sumatra: in monte Singalang, 26 July 1894, V. Sehiffner 324,325 (FH ; isosyntypes); Preanger, montis Pangerango, Tjibodas, 1894, 1600 m, V. Sehiffner 322 (FH , WRSL; isosyntypes); Tjibodas, 0. Renner 429 (S); Tjibodas, Dec 1936, 0. Renner 34g (lE); Tjibodas, 1898, E. Nyman s. n. (L, W); Tjibodas, Botanic Garden, 1952, R. v.d. Wijk 928 (L); Res. Priangan, G. Gegerbentang, 1500- 2000 m, 1930, F Verdoorn s.n. (Verdoorn: Hepat. Se!. et Crit. ser. VII, no. 341) (BM, G, JE, L, M, S, UC, W). For a discussion of MetzgeriaJoliicola, see So (2002). Metzgeria cristata was described as having two rows of hairs along the ventral midrib. However, an examination of the type material in G (epiphyllous on a leaf) shows that the hairs are actually on the dorsal surface of the midrib (Fig. I: c) similar to those seen in M. cristatissima and M. hispidissima Steph. and most specimens of M. Joliicola. An examina­ tion of material of M. Joliicola shows that the dorsal midrib of most has scanty bris­ tle-like hairs similar to material identified as M. hispidissima. Three specimens of M. cristatissima all labell ed as "n. sp." by Herzog are present in lE and one of which, "sub. no. 210," was designated as the type in the protologue. The other two are no. 210. All three specimens are epiphyllous on fern leaves, apparently from the same . Kuwahara (1966) placed M. cristatissma under the synomy of M. Joliicola, based on its epiphyllous habit. M. L. So: The genus Metzgeria in Asia 165

Fig. 3. SEM micrographs of laminal cells of Metzgeria. Metzgeria Joliicola Schiffn. a: dorsal surface showing marginal gemmae and hairs on dorsal costa, b: ventral surface showing involucre, c: laminal cells. Metzgeria leptoneura Spruce. d: ventral surface showing long falcate hairs. Metzgeria scobina Mitt. e: marginal cells, f: laminal cells showing papilla. (a-c from Schiffner 321 in M; d from A. Touw 15730 in L; e, f from L. J Brass 24868a in FH).

4. Metzgeriafurcata (L.) Dumort., Recueil Observ. Jungerm.: 26. Tournay. 1835. Junger­ manniafurcata L., Sp. PI. (ed. 1) 2: 1136. 1753.- Lectotype, designated by Grolle & So 2002: [icon.] "Lichenastrum tenuifolium,furcatum, thecis globosis pilosis" in Dille­ nius, Historia Muscorum t. 74, f. 45 A- C, FG (excl. DE). 1742"1741". Epitype, desig­ nated by Grolle & So 2002: OXF (Herb. Dillenius fol. 163, n. 45); isoepitype: H-SOL 166 1. Hattori Bot. Lab. No. 94 200 3

(ex Herb. Dillenius fol. 163, n. 45)! Metzgeria decipiens (c. Massal.) Schiffn. in Engler, Forschungr. Gazelle 4 (4), Bot.: 43 . '1889' 1890. MetzgeriaJurcata var. f3 decipiens C. Massal., Nuovo Giorn. Bot. Ital. 17: 256. I 885;fide Grolle 2002.- Type: Tierra del Fuego, Isla de los Estados [insula Statuum (Staten Is)] : Basil Hall, Port Vancouver, Spegazzini s.n. (holotype: YU). Metzgeria quadriseriata A. Evans, Proc. Wash. Acad. Sci. 8: 142. I 906; fide Kuwahara I 966.-Type: Japan. Shikoku: Prov. Tosa, Aki-gun, "Ioki-mura, Tosa, Mar 1903, T. Yoshinaga 11" (holotype: VU! ; isotype: G-011881 i). Metzgeriajauriana Steph., Sp. Hepat. 6: 50. 1917 "feuriana"; syn. nov.- Type: Korea. Prope Gen­ san, in Mt Diamonts, 1906, Faurie 36 [incorrectly listed as Taiwan. "Formosa";fide Kuwahara 1986: 208] (holotype: G-001022!; isotypes: BM!, FH!, FI! , NY!). Metzgeria lutescens Steph., Sp. Hepat. 6: 54. 1917; fide Kuwahara 1983.- Type: India. "India ori­ ent.", 1913, Pfleiderer 23 (holotype: G-009907!). Metzgeria planifrons Steph., Sp. Hepat. 6: 59. 1917 ;fide Hattori 1952; nom. illeg. (Art. 52.1)-Orig­ inal material: same as M. quadriseriata Steph. (G-O 11881 i). Metzgeria crispula Herzog, Ann. Bryol. 12: 72. 1939; fide Kuwahara 1983.-Lectotype, designated here: India. Sikkim: "Himalaya, Tsomgo Lake, zwischen Gangtok und Natu La, .. . ca. 3800 m", Aug 1937, C. Troll 39 (lE!; syntypes [29/d, 29/e]: JE!). Metzgeria philippinensis Kuwah. , J. Hattori Bot. Lab. 31: 168. 1968; syn. nov.- Type: Philippines. Luzon Is: Mt Banahao, between Puroy to Duronganwan Ill, 2000- 2300 m, on tree trunk, May/June 1965, Pro! & Mrs A. J. Sharp and Z. Iwatsuki s.n. (holotype: NICH-265496 [not lo­ cated]). Metzgeria amakawae Kuwah. , Rev. Bryol. Lichenol. 36: 534. ' 1969' 1970; synonymized by Kuwa­ hara 1983 with M. fauriana.-Type: Japan. "Yamanashi Pref., Mt Kitadake, 2200 m", Amakawa 5407 (holotype: NICH [not located]). Metzgeria involvens S. Hatt. in Hara, Flora ofE Himalaya Third Report, Uni. Mus. Univ. Tokyo Bull. no. 8: 240. 1975; syn. nov.- Type : Nepal. Between Topke Gola and Shewaden, on rock, 2850m, 28 June 1972, Z. Iwatsuki 1914 (holotype: not located; isotype: TNS!). Metzgeria mitrata Kuwah. , J. Hattori Bot. Lab. 39: 371. 1975; syn. nov.- Type : India. West Bengal: around Sandakphu Bangalow, Darjeeling area, on trunk, alt. ca. 11900 ft, Pro! & Mrs A. 1 Sharp and Z. Iwatsuki s.n. (holotype: NICH [not located]). Metzgeria orientalis (Kuwah.) Kuwah., Rev. Bryol. Lichenol. 14: 394. 1978. Apertithallus orientalis Kuwah., 1. Hattori Bot. Lab. 31: 260. 1968; syn. nov.- Type: Philippines. Luzon: Mt Bu1usan, between junction to Masacot Spring, 450- 900 m, on tree branch, 31 May, 1965, Z. Iwatsuki & E. Sharp s.n. (holotype: NICH-265932 [not located]). Metzgeria liaoningensis Gao in Fl. Hepat. Chin. Boreali-Orient.: 175. 1981 ; syn. nov.-Type: China. Liaoning: Distr. Fengchengxian, Mt Fenghuangshan, on rock, 24 VII 1961, Gao Chien 6931 (holotype: IFSBH'). Illustrations: Kuwahara (1958: fig. Ill, fig. V: 1-9 as M. quadriseriata; 1960: fig. 10: h- j as M. quadriseriata; 1966: fig. 11: cl, 21: e-g as M. decipiens; 1968a: fig. 5 as M. philippinensis; 1968b: fig. a- p as M. orientalis); Srivastava & Udar (1975: fig. XIX as M. crispula); Stephani (1985 : Icon. n. 7666 as M. lutescens, n. 7674 as M. planifrons). Description: Kuwahara (1966: 236). Distribution: Worldwide. Additional specimens examined: Bhutan. Upper Pho Chu, W of Tarina Lakes, 4380 m, 30 Aug 2000, G. & S. Miehe 00-338-06, mixed with Plagiochila retusa Mitt. (herb. G. & S. Miehe). China. M. L. So: The genus Melzgeria in Asia 167

Anhui: Qingliangfeng, W F. Zheng 1779 (HSNU). Guizhou: collector unknown (HSNU-3668). Hong Kong: Tai Mo Shan, 880 m, 1995, M. L. So 95424 (HKBU); Tai Mo Shan, 700 m, 1994, M. L. So & R. L. Zhu 94727L14 (HKBU). Jiangxi: Sanqingshan, 1520m, 1987, X M. Shao 1025 (HSNU). Sichuan: Mt Erlang, 1850m, 1978, Q. Li 2706 (HSNU). Yunnan: Diqing Prefecture, Weixi County, W side of Litiping Plateau, 3120 m, 1993, D. G. Long 24531 (E). Zhejiang: Fenyangshan, 1600 m, 1992, R. L. Zhu 92227 (HSNU); July 2001, R. L. Zhu 721-5 (HSNU). Indonesia. Irian Jaya: Western Sudirman Range, Carstensz Mountains, Bacopa Valley, alpine meadow, 3840 m, 25 Jan 1991, H. A. Miller 21918,21919 (MU). Java: Tjibodas, Mountain Gardens, 1450m, 1952, W Meijer 174 as M. decipiens (L); Anonymous s.n. (S[B22738]); Prov. Buitenzorg, Anonymous s.n. (S[B22740]). Sulawe­ si (Celebes): Anonymous s.n. (S[B22618]). Japan. Shikoku: Tosa, Kuromori, 12 Jun 1910, Collector unknown 4, 38 (MU). Honshu: Hando-yama, Minaguchi, Shiga Pref., on moist granite rocks in shade, 380m, 1963, T. Kodama s.n. as M. quadriseriata (Hattori: Hepat. Jap. Exs. ser. 14, no. 680) (BM, EGR, M, S); Yaezi Pass, Kamikatsu-cho, Katsuura-gun, Tokushima-ken, 1973, T. Kodama s.n. as M. decipiens (Hattori: Hepat. Jap. Exs. ser. 20, no. 987) (M). According to Kuwahara (1968a: 170), the essential taxon characteristics of M. philip­ pinensis are 2 dorsal costal cells, 3---4 ventral costal cells, lamina of 12- 20 cells and no gemmae. However, all these features indicate that specimens assignable to M. philippinen­ sis belong to M.Jurcata. Metzgeria orientalis was created based on a single collection from the Philippines and placed in a new genus Apertithallus (Kuwahara 1968b: 260), which was later reduced to a section within the genus Metzgeria (Kuwahara 1978b: 389). The sole diagnostic feature of this new species is the midrib not reaching the apex. However, all fea­ tures are within the variability of the widespread M. Jurcata. Furthermore, in M . orientalis, branches have quite normal midribs, reaching the apex (Kuwahara 1968b: fig. b). Based on specimens upon Metzgeria Jurcata examined in the present study, midribs that do not reach the apex are not unusual. Especially when gemmae and lateral branches are numerous, the "punctuated appearance" of the midribs is common, and there are even branches without a midrib. As already noted by Kuwahara in his discussion on reduction of gametophytes (1 978a: 132), "midribs are often reduced to only 1 medullary cell and midrib was only partly developed." The apices of the thallus in M. mitrata was described as being hooded. However, such hooded apices can also be seen in some specimens of M. Jurcata. Metzgeria planifrons was described by Stephani (1917) based on a specimen provided by Evans (as inscribed on the specimen label in 0). However, the same collection had al­ ready been described by Evans (1906) as M. quadriseriata, and even the specimen number is the same. Hence, M. planifrons is here regarded as nomen illegitimum. Kuwahara (1958) admitted he did not study the type material. 5. Metzgeria leptoneura Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15 : 555. 1885.­ Type: Peru. "in monte Campana andium, Spruce s.n." (holotype: MANCH-19520!; isotypes: F!, 0-017844!, W!). Fig. 3: d Metzgeria hamata Lindb., Acta Soc. Fauna Fl. Fenn. I (2): 10, 25. '1877' 1878; nom. illeg. (Art. 52 .1); fide Engel I 976.-0riginal material: N. America.-Jamaica.-Chile.-New Zealand.­ India: Sikkim. Metzgeria hamatiformis Schiffn., Nova Acta Acad. Caes. Leop-Carol. German. Nat. Cur. 60: 272. 1893; fide Kuwahara 1966.- Type: Indonesia. "Amboina: Tolepoel, prope Ambon," 1889, G. Karsten s.n. (holotype: FH! ; isotypes: BM!, G-001053, S[B-37923]!). 168 J. Hattori Bot. Lab. No. 94 2 0 0 3

Metzgeria sandei Schiffn., Denkschr. Kaiserl. Akad. Wiss., Math.-Naturwiss. KI. 67: 181. 1898; syn. nov.-Lectotype, designated here: Indonesia. Sumatra: In monte Singalang, 2390 m, 25 Jul 1894, V Schiflner 332 (FH! ; isolectotypes: FH! , M!). Metzgeria fuscescens Mitt. ex Steph., Bull. Herb. Boissier 7: 945. Sp. Hepat. 1: 293. 1899; fide Kuwahara 1966.-Lectotype, designated by Kuwahara 1966: Indonesia. Java: In monte Mega­ mendong, 6000 ft, Motley s.n. (NY; isolectotypes: FH! , G-OOI 031 !). Metzgeria curviseta Steph., Hedwigia 44: 72 (31 Jan). Renauld & Cardot, Bull. Soc. Roy. Bot. Bel­ gique 41, Mem.: 120 (8 May). 1905; fide Kuwahara 1965a.-Lectotype, designated by Kuwa­ hara 1965a: India. "Simla" (Panjab Himalaya), Mahassu, on rocks, 8000 ft, I Nov 1900, J Doulea s.n. , herb. Levier 2981 (G-009851!; isolectotypes: BM! , FI!). Metzgeria subhamata S. Hatt. in Herzog & Noguchi, 1. Hattori Bot. Lab. 14: 30. I 955 ;fide Kuwaraha I 969c.- Type: Taiwan. "Im Gebirge, an Biiumen, 1400- 1500m, 21 Aug 1947, G. H. Schwabe 100" [listed as 101 in the protologue] (holotype: NICH [not located]; isotype: JE!). Metzgeria iwatsukii Kuwah., 1. Hattori Bot. Lab. 31: 162. I 968; fide Kuwahara I 984a.- Type: Philip­ pines. Luzon Is.: Mountain Province, Mt Pulog, . .. ca. 2200 m, . .. May-June 1965, Prof & Mrs. Sharp and Z. lwatsuki S. n. (holotype: NICH-266418 [not located]). Metzgeria borneensis Kuwah., 1. Hattori Bot. Lab. 28: 167. 1965; syn. nov.- Type: Malaysia. Bor­ neo: South slope of Mt Kinabalu, around Paca Cave, 2980 m, on rock, M. M. 252881 (holotype: NICH [not located]). Metzgeria luzonensis Kuwah., 1. Hattori Bot. Lab. 31: 164. 1968; syn. nov.-Type: Philippines. Luzon Is.: Mt Pulog, ca. 2200 m, ... , May/June 1965, Pro! & Mrs A. J Sharp and Z. Iwatsuki s.n. (holotype: NICH-266388 [not located]). Metzgeria sharpii Kuwah., 1. Hattori Bot. Lab. 31: 171. 1968; syn. nov.- Type: Philippines. Luzon Is: Mountain Provi nce, Mt Pulog, .. . ca. 2200 m, May- June, 1965, Pro/ & Mrs A. J Sharp & z. 1watsuki s.n. (holotype: NICH-267324 [not located]).

Illustrations: Kuwahara (1958: fig. V: 10- 20 as M. hamata; 1966: fig. I: a-g as M. hamata; 1968a: fig. 2 as M. iwatsukii; fig. 7 as M. sharpii); Stephani (1985: Icon. n. 7645 as M. curviseta, n. 7676 as M. sandei). Description: Kuwahara (1966: 223 as M. hamata). Distribution: Worldwide. Representative specimens: China. Anhui: Qingliangfeng, W F Zh eng 1516, 1532 (HSNU); Ma­ jiahe, 1986, K. F Cai 2095 (HSNU); Huangshan, 1400m, 1983, R. L. Wu 165 (HSNU). Xizang: Motuo, Nalungla, Anonymous s.n. (KUN). Yunnan: Tsang Shan, Shu Ching Qan valley NW of Hua­ dianba, 2950m, 1990, D. G. Long 19246 (E); Lijiang District, between Baishui and Heishui, N ofLi­ jiang, 2820 m, 1990, D. G. Long 18890 (E). Zhejiang: Jiulongshan, 1420 m, 1981 , Z. L. Liu 511, 611 (HSNU). India. Nainital Di strict: N facing cliffs below Tiffin Top, Nainital, 2250 m, I Feb 2002, D. G. Long 20294 (E); N side of Snow View ridge, E of Kilbury Road, Nainital, 2250 m, 2 Feb 2002, D. G. Long 20944 (E); between Land's End and Tiffin Top, Nainital, 2210m, I Feb 2002, D. G. Long 30925 (E). NW Himalaya (Simla): Mashobra, on ground, 8000 ft, 1906, E. Long 108, herb. Levier 5806 (BM, FI, G; syntypes of M. curviseta) ; NW Himalaya, Tihri Garhnival Kidarkanta, 10000 ft , 1904, Bahadru s.n., herb. Levier 4702 (S, UC). West Di strict: Chhurong Chhu Valley between Tsoka and Jamlinghang, 3165m, 1992, D. G. Long 222471 (E); Nagteeb, 7900 ft, 1864, W Bell 241, 242 (E). Sikkim Himalaya: prope Kurseong, Sonada, 1200 ft, 1899, Decoly & Schaul s.n. , herb. Levier 405 (M); Eastern Himalaya, 1938, Sherifls.n. (E). Indonesia. Java: Prov. Preanger, montis Pangeran­ go, Tjiburrum, 1685 m, 2 May 1894, V Schiffner 318 (BM, FH , M; isosyntypes of M. hamata var. an­ gustior Schiffn.); Buitenzorg, 1897, M. Treub s.n. (E, NSW); Batavia, in monte Megamendong, Tela- M. L. So: The genus Metzgeria in Asia 169 ga Wama, 1400m, 1894, V. Schiffner 255 (M); 1000m, 1894, V. Schiffner 254 (UC); V Schiffner 253 (EGR, WRSL); Prav. Preanger, Pangerango, Tjibodas, 1630m, 1894, V Schiffner 262 (E, M, NSW); V Schiffner 260 (UC); Prav. Preanger, montis Pangerango, 1545 m, 24 Apr 1894, V Schiffner 329, 330, 331 (FH; syntypes of M. sandei); Res. Priangan, G. Gegerbantang, 1500-2000m, 1940, F Ver­ doorn s.n. (Verdoom: Hepat. Se!. et Crit. ser. VIII, no. 382) (M, UC); Res. Priangan, G. Gede, Tjibo­ das, 1420-1650m, 1930, F Verdoorn S.n. (Verdoom: Hepat. Se!. et Crit. ser. VIII, no. 381) (M, UC); Tjibodas, 1835m, V. Schiffner 277 (UC); Tjibodas, 1899, Geisenhagen 46 (M); Tjibodas, 1910, M. Fleischer s.n., herb. Levier 4135 (EGR, NSW); Tjibodas, 1898, E. Nyman s.n. (BM, CHR, EGR, F, FH, NY, S, W); Tjibodas, 1560 m, 1894, V Schiffner 351 (BM); V. Schiffner 352 (FH, WRSL); Tjibo­ das, 1500m, 1894, V. Schiffner 348 (E, EGR, W); 349 (FH, W); 353 (FH, W); 1120m, V Schiffner 347 (FH); 1500m, V Schiffner 343, 350, 351 (FH); Tjibodas, 1927, H. Burgeff 8133/a (S); Salak, 700 m, 1897, E. Nyman s.n. (BM, FH, S, W); Salabintana, near waterfall, 1500 m, 1910, M. Fleischer 118 (M); Idjenp1ateau, an Tjemarabiium, 200 m, 1911 , J Froehlich s.n. (S); Prav. Buitenzorg, Anony­ mous s.n. as M./urcata (S[B22739]); Prav. Buitenzorg, 1843, Anonymous s.n. (S[B22737]); Zollinger s.n. (S[B22743]); Natural Reserve, Border 2- 3, 1770 m, 1949, Noerta & Soekar 1704 (L); Geger Bentang-Hondje-Warak, 1685 m, 1949, Noerta & Soekar 1978 (L). Kalimantan: Peak of Balikpapan, terrace Andau, 1000m, 1952, W. Meijer B-1975A, B-2498/a (L). Lesser Sunda Islands: Flores, Mang­ garai Prov. , Go10 Lusang, S of Ruteng, 1470-1500m, 1988, A. Tou w & M. Snoek 22873,22928 (L). Moluccas: Ke1i, NW Buru, S of Bara, Waeduna River, 350-400 m, 1984, M. M. J v. Balgooy 50- 25, 4892A (L). Sulawesi: Tana Toraka, forest remnant between Lempo & Lokomata, 1150m, 1990, A. Touw & M. Snoek 24492 (L); Sopu valley, 80 km SSE of Palu, 1000 m, 1979, M. M. J v. Ba/gooy 3070C (L); 2000 m, 1979, E. Hennipman 5515 (L); Subdiv. Enrekang, Tinabang, W side of the Rante Mario, 3000 m, 1937, P. J Eyma 803 (L). Sumatra: In monte Singalang, 1800m, 1894, V Schiffner 285 (G-1028, M); "in monte Singalang, .. .", 2390 m, 24/25 lul 1894, V. Schiffner 333, 334 (FH; syn­ types of M. sandei); montis Merapi, V. Schiffner 293 (WRSL); Fort de Kok, 1899, K. Geisenhagen 26 (M); Palembang, Bukit Besar, 1899, K. Geisenhagen 144 (M); Merapi, 1760m, 1894, V Schiffner 355 (FH); G. Dempo, 1949, W. C. Verboom s.n. (L); Prov. Lampung, Mt Tanggamus, 1968, M. Jacobs B8778 (L); Gouv. S.O.K., G. Sibajak, 1500m, 1913, M. Fleischer s.n. (Verdoom: Hepat. Se\. et Crit. ser. VIII, no. 383) (BM, L, M, S, W). Japan. Honshu: Fujisan, 1897,J Bisset s. n. (E). Shikoku: Prav. Awa, 1932, T. 1shikawa s.n. (M); Kumamoto, Midzukami, 800 m, 1950, M. Mayebara S.n. (Hattori: Hepat. lap. ser. 4, no. 175) (M, VC); Kochi-ken, Mt Honokawa, on moist rock, 1940, M. Kamimura 7064 (CAHUP). Malaysia. Ma1ay Peninsula: Pahang. Genting Highlands, 1310 m, 1997, N. Klazenga 34 (L). Sabah: Tawau, Tawau River Forest Reserve, 1959, W. Meijer 8-12975 (L); Tawau River Forest Reserve, 1960, W. Meijer 738A (L); Mt Trusmadi, 1000m, 1969, H. P. Nooteboom 1362E (L); West Coast Reserve, Mt Kinaba1u, 1960, W. Meijer B10531 as M. sharpii (TNS). Sarawak: 4th Division, Gunong Mulu National Park, 430m, 1978, A. Touw 21249, 21258 (L). Nepal- Rasuwa Distrct: be­ tween Shin Goimpa and Phorang, 3150m, 10 Oct 2001, D. G. Long 30519 (E); Langtang Khola be­ tween Chunama and Ghora Tabe1a, 2720m, 12 Oct 2001, D. G. Long 30575 (E); between Bamboo Lodge and Hot Springs, 1920 m, 17 Oct 200 I, D. G. Long 30667 (E); between Thu10 Syabru and Brabal, 2190m, 18 Oct 2002, D. G. Long 30701 (E); Sankhuwasabha District, forested edge between Tashigaon and Kauma, 2670 m, 1991, D. G. Long 20587 (E); Vorhimalaya, westlich Rauje, 4000-4500 m, 1968, J Poelt H724 (M); S slopes of Pawa Khola Valley, 2420 m, 1991 , D. G. Long 21296 (E); 2483 m, 1991 , D. G. Long 20567 (E); near Sete, 2500 m, 1965, J A. Noordijk 108, 109 (L); Langtang Himal: Trekking route from Dunche to Kyangjing; valley of Langtang Khola between Ghora Tabela (3300 m) and Lama Hotel (3050 m), 1991 , W. A. Weber 8-99326 (UC); Rasuwa Dis­ trict, near Syabru, 2300 m, 1992, D. G. Long 22022 (E); wood below Taskshinder, 2000-3000 m, 170 1. Hattori Bot. Lab. No. 94 2 003

1965 , J A. Noordijk 75 (L). Philippines. Luzon: Mt Pulog, 2500 m, 1968, M. Jacobs B299, B258 (L); Mt Pulog, trail to summit, 1978, Alvarez & Saprid 0-78566 (CAHUP); Mt Tabayoc, Lake Ingcolos, 2420- 2450 m, 1968, M. Jacobs B21 8, B390 (L); Mt Tabayoc, 1968, M. Ja cobs B698 (CANB, FH) ; Mt Makiling, 1949, Dequi/la 16pp, 22 (L); Mt Data, 7000 ft , 1931 , F. C. Hadden s. n. (FH). Mindoro: Mt Tabaco, Ramayam, 1988, E. Salgado 88.? 1205 7 as M. luzonensis (BR). Sri Lanka. Hunasgirias Peak, 2500 m, 9 Feb 1898, M. Fleischer 2082 as M. papulosa (EOR). Thailand. Payap, summit of Doi Inthanon, 1969, v. Beusekom B205, D215 (L); Doi Inthanon, 2500 m, 1965, A. Touw 9736 (EOR, L); Udawn, Phu Luang, 1300-1350m, 1966, A. Touw 10717 (L); Nokornsrithamarat, Khao Luang, 700 m, 1966, A. Touw 12005 (EOR, L) . From the descriptions and illustrations of Kuwahara (1968a), P sharpii and P luzo­ nensis, both from the Philippines, and P borneensis from Borneo, fall within the variability of the common M. leptoneura, especially the paired marginal weakly falcate hairs. The hair lengths were thought to be of taxonomic value in separating different species. These hairs, ranging from (50)70-170 ,Urn, were first described as bristle-like, but later they were de­ scribed as flexuous. Hence the length of hairs is no longer a diagnostic feature separating these species from M. leptoneura. As already noted by Kuwahara (1968a: 164), M. iwat­ sukii, also described from the Philippines and later synonymized with M. leptoneura, is a member of the large group of Metzgeria hamata-complex whose taxonomic nature is very flexible. The hairs in M. iwatsukii are 70- 200,Um. Metzgeria sandei, a species described from Java, was described as having a costal anatomy of 2/2-4 cells, otherwise, all other features are similar to that of the most common M. leptoneura. Based on the numerous specimens cited above, the ventral costal cells are usually two in number, but four is not unusual. Metzgeria leptoneura with its wide geographical range is very variable indeed. Plants from the Hilamayas and Japan often have the typicaly short paired hamate hairs (150,Um long) whereas many collections from New Guinea have much longer, falcate and flexuous hairs, reaching a maximum length of 400 ,Urn. Species identified as M. nitida Mitt. from Australia and New Zealand usually have a mixture of short and long flexuous or even straight hairs. 6. Metzgeria lindbergii Schiffn., Denkschr. Kaiser!. Akad. Wiss., Math.-Naturwiss. K!. 67: 182. I 898.- Lectotype, designated by So 2002: Indonesia. Java: Prov. Batavia, 200- 260m, V Schiffner 340 (FH!; isolectotypes: L-912.295. 1228! , M!, S[B37942-3]! , UC!, W! , WRSL! ; syntype [Sumatra: 1600- 1680, Schiffner 345]: FH!). Metzgeria himalayensis Kashyap, J. Bombay Nat. Hist. Soc. 25: 280. 191 7;fide Kuwahara 1958.­ Type : India. 1912/4, Kashyap s.n. (holotype: not located). Metzgeria conjugata var.japonica S. Hatt. , 1. Hattori Bot. Lab. 15: 80. I 955;fide Kuwahara 1984b.­ Type : Japan. Hokkaido: Hidaka, Mt Apoi, 1954, S. Shimizu s.n. (holotype: NICH [not located]). Metzgeria minuta Kuwah. , 1. Hattori Bot. Lab. 31: 166. 1968; fide Kuwahawa 1984b.- Type : Philip­ pines. Luzon Is: Mt Pu10g, ... 2200- 2700 m, ... , May/June 1965, Pro! & Mrs A. J Sharp and Z. 1watsuki s.n. (holotype: NICH-265588 [not located]). Metzgeria assamica Srivastava, 1. Indian Bot., Soc. 55: 194. 1976; syn. nov.-Type: India. Assam: Mausmai, Cheerapunji, 4500 ft , Oec- Jan 1975- 76, D. D. Awasthi & N. K. Mehrotra & Party 8//75- 76 (holotype: LWU!). Metzgeria minor (Schiffn.) Kuwah. , J. Jap. Bot. 53: 269. 1978. Metzgeria conjugata var. minor M. L. SO: The genus Metzgeria in Asia 171

Schiffn., Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur. 60: 271. I 893; fide Schiffner 1900.- Type: Indonesia. Java: 1889, Goebel s.n. (holotype: not located). MetzgeriaJukuokana Kuwah., 1. Jap. Bot. 53 : 264. 1978; syn. nov.-Type: Japan. Fukuoka Pref., Ohnojo-shi, Ushikubi, shrine wood, on tree trunk, ca. 30m, 30 Dec 1977, Y Kuwahara 9014 (holotype: not located; isotype: NY!). Illustrations: Kuwahara (1958: fig. VI as M. conjugata var. japonica; 1968a: fig. 4: a- h as M. minuta; 1978c: fig. 1- 18 as M.fukuokana). Description: Kuwahara (1966: 229 as M. conjugata Lindb. subsp.japonica (S. Hatt.) Kuwah.). Distribution: Asia and Australasia. Representative specimens: Bhutan. Above Lami Gompa, Bumtang, on shady rocks, 3000 m, 1979, D. G. Long 8377, 8419 (E). China. Anhui: Qingliangfeng, W F. Zheng 1155, 1225 (HSNU); Majiahe, 1985, K. H. Cai 1128 (HSNU); 1986, K. H. Cai 2054 (HSNU). Jiangxi: Shanqingshan, 1987, X M. Shao 250 (HSNU). Sichuan: Mt Omei, SW of Chengdu, 1000 m, 1980, A. Tou w 23979 (L). Yunnan: Chungtien, 4350-4450m, 1915, Handel-Mazzetti 6925 (W). Zhejiang: Jiulongshan, 920m, 1981, Z. L. Liu 945, 1808 (HSNU); Jiulongshan, Beiping, 1130m, 1981, R. L. Hu 710 (HSNU); Wuyangleng, 1285m, 1987, R. L. Zhu 334,1016 (HSNU). Indonesia. Java: Prov. Batavia, 250 m, 1893, V. Sciffner 341 (E, FH, NSW; isosyntypes); Prov. Batavia, 250 m, 1893, V. Schiffner 343 (BM, FH, M, W; isosyntypes); prope Gadok, 500m, 1894, V. Schiffner 344 (FH, S, W; isosyntypes); Prov. Batavia, 200-260 m, Schiffner 33 7, 338, 339 (FH); Tjibodas, Mt Garden, 1953, W Meijer 8025 (L); Buitenzorg, 1897, E. Nyman s.n. (CHR, FH, L); Prov. Preanger, in decliv. austral. montis Pangerango, 1500 m, 1894, V. Schiffner 348 (M, S); Geger Bentang-Hondje-Warak, 1690 m, 1949, Noerta & Soekar 2045, 2777 (L); Bogor, Kebon Raya, 200 m, 1952, M. Weijer 70, B5316 (S); Bogor, Botanic Gardens, 1952, R. v.d. Wijk 884B, 918 (L); Kandang Badak, 1700 m, 1952, R. v.d. Wijk 440B (L). Kalimantan: G. Sentuli, north of Kuala-Kuajan, Sampit District, lOOm, 1953, Kostermans 20B (L). Lesser Sunda Islands: Flores, Prov. Manggarai, Ruteng, Lake Mese, 1230 m, 1988, Fr. E. Schmutz SVD 6812, 6834 (L). Sulawesi: 1100-1250m, 1979, E. Hennipman 5525C (L). Japan. Iwashiro, 1910, A. Yasuda 22 (W). Honshu: Gifu-ken, Motosu-gun, Neo-mura, 1- 2 km SW of Shimo-psu, 300-350 m, 1984, M. Mizutani 11729 (S); Sasayama-cho, Taki-gun, Hyogo-ken, 200 m, 1965, M. Mizutani s.n. (Hattori: Hepat. Jap. Exs. ser. 18, no. 882) (M, S) ; Prov. Musashi, Chichibu, 1922, K. Sakurai H 7483 (CHR); Osaka Pref., Hara, Takatsuki, 1951 , T. Kodama s.n. (Hattori: Hepat. Jap. Exs. ser. 13, no. 632) (EGR, M, MU, S); Hyogo Pref., Tsukihara, Kobe City, 130 m, 1964, T. Ko­ dama s.n. (Hattori: Hepat. Jap. Exs. ser. 14, no. 678) (EGR, M, S). Kyushu: Kumamoto Pref., Ishochi, lOOm, 1950, K. Mayebara H1 8508 (M); Miyazaki, Minaminaka, Kitago, 300m, 1947, S. Hattori & T. Kurata s. n. as M. himalayensis (Hattori: Hepat. Jap. ser. 2. No. 93) (NSW); Fukuoka Pref., Dazaifu, 40m, 1953, Y Kuwahara s. n. (Hattori: Hepat. Jap. ser. 7, no. 325) (EGR, MU, S, UC) . Hokkaido: IsI Rishiri, Yamunaisawa, 100 m, 1954, T. Shimizu H5361 7 (M). Malaysia. Malay Penin­ sula: Pahang, Genting Highlands, 1310m, 1997, N Klazenga 57, 150 (L). Nepal. N-facing slopes of Dobala Danda above Kabeli Khola, ravine in wet mossy oakllaural forest, on log, 2350 m, 1989, D. G. Long 17352 (E); Tinjure Danda above Chauki, 2900 m, 1989, D. G. Long 17576 (E). Philippines. Mindanao: Butuan, 1911 , 0. M. Weber 1347 (BM, E). Luzon: Infanta, Province of Tayabas, Mt Bin­ uang, epiphyttous, 1909, C. B. Robinson s.n. (Bureau of Science no. 9589) (BM, EGR); Mt Pulog, 2500 m, 1968, M. Jacobs B262 (L); Locality unknown, 1906, A. Loher 5498 (BM). Sri Lanka. New­ era-ElIia, 1865,0. Beccari 60 (L); Newara Eliya District, Jard. Bot. de Hakgala, 23 Mar 1976, M. Onraedt 76.L.2756 as M. /uzonensis (BR). M etzgeria lindbergii, as the only autoicous species in this region, is easily distin­ guished from other species, and plants are usually fertile. Hairs are a mixture of single and 172 1. Hattori Bot. Lab. No. 94 2 0 0 3 paired ones. 7. Metzgeria pubescens (Schrank) Raddi, Jungermanniogr. Etrusca: 35. [= Mem. Mat. Fis. Soc. Ita!' Sci. Modena, Pt. Mem. Fis. 18: 46]. 1818. Jungermannia pubescens Schrank, Primit. FI. Salisb.: 231. 1792. Apometzgeria pubescens (Schrank) Kuwah., Rev. Bryo!. Licheno!. 34: 214. 1966.- Type: Austria. "An feuchten, schattigen Kalk­ felsen der Gebirge urn Salzburg." Sauter [Rabenh., Hep. Europ. (Exs.) no. 84] (Neo­ type, designated by Grolle & So 2002: Wl). Fig. 2: c-f Metzgeria duricosta Steph., Sp. Hepat. 6: 50. 1917;fide Hattori 1955 .- Type: Korea. Insula Quel­ paert, 1907, U Faurie 223 (holotype: G-009716!). Metzgeria longifrondis Gao in Gao & Chang, Fl. Hepat. Chin. Boreali-Orient.: 210. 1981 ; fide Zhang 1991.- Type: China. Prov. Liaoning, Distr. Benxi Shi, on moist rock surface,S VII 1973, Gao Chien & Chang Kuang-chu 8340 (holotype: IFSBH!). Illustrations: Kuwahara (1966: fig . 13); Stephani (1985: Icon. n. 7648 as M. duricosta, n. 7695 as M. pubescens). Description: Kuwahara (1966: 212). Distribution: Europe, N America, Hawaii and Asia. Additional specimens examined: China. Anhui: Majiahe, 1985, K. H. Cai 1122, 1188, /189 (HSNU); 1986, K. H. Cai 2055 (HSNU). Schen-si (Shaanxi): prope Ta-sche-tseun, 1896, J. Giraldi s.n., herb. Levier 192 (M). Yunnan: Deqen Co. , Meili Shan, by the Dokar La River, 2845 m, 23 May 1994, D. F Chamberlain s.n. (E). Hawaii. Exact locality unknown, 1876, J. Barly 19 (MU-42357). India. W Himalayas, Dalhousie, Chamba Road, on moist rocks, 7000 ft, 1928, R. S. Chopra H7490, H7491 (CH R-524645-6). Indonesia. Sumatra: Atjeh, G. Kemiri, 2600 m, 1971 , J. Dransfield 1910 (L). Japan. Shikoku: Ehime Pref., Mt Higashiakaishi, ca. 1600m, 1950, M. Hara 13362 (CAHUP); Kochi-ken, Mt Irazu, on moist rock, ca. 1000 m, 1949, T Yamanaka-35 9796 (CAHUP); Prov. Awa, 1932, T lshikawa s.n. (M). Hokkaido: Antaroma, Kamikawa District, 1856, T Sasaki s.n. (Hattori: Hepat. Jap. ser. 11 , no. 541) (M, MU). Honshu: Gifu Pref., Miyagawa, 900m, 1958, N. Kitagawa s.n. (Hattori: Hepat. Jap. Exs. ser. 12, no. 584) (M). Hokkaido: Kushiro-gun, Kushiro-cho, 15 m, 16 Nov 1980, K. Takita s.n. (Iwatsuki & Mizutani: Bryophy. Esc. Fasc. 3, no. 136) (EGR). Nepal. Wood below Takshinder, 2000-3000 m, 1965, J. A. Noordijk 74 (L). Metzgeria pubescens was placed in a separate genus, Apometzgeria, by Kuwahara (1966) based on the dense hairy surfaces of the thallus and numerous costal cells across the midrib. However, the presence of hairs on the dorsal surface occurs in quite a number of species, viz. M. robinsonii, M. foliicola and M. bartletii Kuwah. The number of costal cells is also variable, from as few as 2-3(4) in most species to as many as 9-16 in M. alpina R. M. Schuster & J. J. Enge!. Metzgeria pubescens is here treated as a member of Metzgeria. 8. Metzgeria robinsonii Steph., Sp. Hepat. 6: 60. 1917.-Type: Philippines. Luzon, Province of Laguna, Mt Banajao, Jan 1909, C. B. Robinson s.n., Merrill 6591 (holo­ type: G-00I008l; isotypes: BMl, NYl). Fig. 1: a, b Illustrations: Kuwahara (1966: fig. 12: d; 18: c, d; 1968a: fig. 6); Stephani (1985 Icon. n. 7675). Description: Kuwahara (1966: 221). Distribution: Endemic to the Philippines. Metzgeria robinsonii is known only from the type. This species bears superficial re­ semblance to M. pubescens, with numerous hairs on the dorsal surface (fig. I: a, b), a fea- M. L. So: The genus Metzgeria in Asia 173 ture not seen in other species of Metzgeria in this region. However, the two dorsal and two ventral costal cells, together with hairs confined to the ventral costa will serve to distin­ guish itself from M. pubescens. 9. Metzgeria scobina Mitt., J. Linn. Soc., Bot. 22: 243. '1887' I 886.-Type: Borneo. Sarawak, on Mt Matang, Everett s.n. (holotype: NY!; isotype: 0-001006!). Fig. 3: e, f Metzgeria subscobina Kuwah, 1. Hattori Bot. Lab. 31: 173. I 968 ;jide So 2002.-Type: Philippines. Luzon Is: between Bulusan Lake & Second Lake, Mt Bulusan, 600-1500 m, May-June, 1965, Pro! & Mrs A. J Sharp and Z. Iwatsuki s.n. (holotype: NICH-265918 [not located]). Description: Kuwahara (1966, p. 221). Illustrations: Kuwahara (1966: fig. 17: b-e as M. scobina; 1968a: fig. 8 as M. subscobina); Stephani (1985 : Icon. n. 7677 as M. scobina). Distribution: The Philippines, Indonesia, Malaysia, New Zealand and New Guinea. Additional specimens examined: Indonesia. Sumatra: Kehding s. n. (FH, G-I005). Malaysia. Borneo: Sansubong Bay, Collector unknown s.n. (NY). Sarawak: Mt Matang, 1866, 0. Beccari 54 (L). Selangor: Ridley 470 (NY). For a discussion on M. scobina see So (2002). The presence of a prominent papilla in each laminal cell is a unique character which separates this species from all others in this region. Also, it is quite rare and only a few collections are available for study. 10 . Metzgeria temperata Kuwah., J. Hattori Bot. Lab. 40: 219. 1976.- Type: Japan. Pref. Mie, Komono-cho, Yunoyama-onsen, 450m alt., on trunk of Quercus, 1955, T Ko­ dama 9999 (holotype: NICH-61032 [not located]). Illustration: Kuwahara (1958: fig. IV as M.fruticulosa (Dick.) A. Evans). Description: Kuwahara (1976: 219). Distribution: Japan & N America. Additional specimens examined: Japan. Honshu: Shiga-ken, Mt Ryozen, Nyutani, 260-300 m, on tree trunk, M. Mizutani 12739 (L, S); Gifu-ken, Itadorimura, Otani valley, 300--400 m, 1987, M. Mizutani 13109 (L, S); Gifu-ken, Hachiman-cho, Hachiman-jo, 250- 350 m, 1988, M. Mizutani 13636 (L, S); Nekko-gawa, Amagi-yugashima-cho, Tagata-gun, Shizuoka-ken, 1980, M. Mizutani s.n. (Hattori: Hepat. Jap. Exs. ser. 22, no. 1084) (L, S); Nagano, Kose-spa, 8 June 1983, J G. Gar­ diner K10 (JE). Kyushu: Miyazaki Pref., Kushima-shi, Ootsuka, 500m, 1950, S. Hattori H16584 (M). U.S.A. Virginia: Giles County, vicinity of Mountain Lake Biological Station, on trunk, 3900 ft, 1978, R. Busk 2092 (CAHUP). When the extreme variability and malleability of most species of Metzgeria is consid­ ered, the differences between M. temperata and M. consanguinea mentioned in the key by Kuwahara (1976) are exceedingly small. It is highly likely that M. temperata is merely a form of the latter species. In fact, all specimens listed as M. temperata by Kuwahara (1976) were originally assigned to M. consanguinea.

EXCLUDENDA Metzgeria albinea Spruce, [Rev. Bryol. 15 : 34. 1888; nom. inval. (Art. 32.1(c»] Bull. Soc. Bot. France 36 (suppl. Congres Bot. 1889): 201. "1889"1890. Sp. Hepat. 1: 296. I 899.-Type: Brazil. Prope Rio de Janeiro, Glaziou 7378 (holotype: MANCH-7378!). For discussion see So (2002). 174 1. Hattori Bot. Lab. No. 94 2 003

Metzgeria vivipara A. Evans, Ann. Bot. (London) 24: 287. \9IO.- Type: Puerto Rico. Utu­ ado to Adjuntas, 21 Mar 1906, N. L. Britton & J F. Cowell1242 (holotype: YU1; iso­ type: NY1). The only record in this region was from Sri Lanka (Kuwahara & Onraedt 1979), but the collection located in BR belongs to M. crassipilus which has discoid or elongated gem­ mae on the dorsal surface of the thallus. The hairs from the gemmae are not conspicuously curved, but rather straight. According to Evans (1910), M. vivipara has discoid gemmae with distinctly curved hairs.

LIST OF R ECOGNIZED SPECIES OF METZGERIA IN ASIA, AUSTRALASIA AND THE PACIFIC Accepted names in bold, synonyms in italics Metzgeria alpina R. M. Schust. & 1. 1. Engel Metzgeria glaberrima Steph. 1900 1988 (New Zealand) Metzgeria howeana Steph. 1914 Metzgeria bartlettii Kuwah. 1987 (New Metzgeria pauciseta Steph. 1914 Zealand) Metzgeria caledonica Steph. 1917 Metzgeria comata Steph. 1899 (New Caledo­ Metzgeria/auriana Steph. 1917 nia) Metzgeria mauina Steph. 1917 Metzgeria marginata Steph. 1917 Metzgeria dens is eta Steph. 1917 Metzgeria clavipila Steph. 1917 Metzgeria lutescens Steph. 1917 Metzgeria lucens Steph. 1917 Metzgeria pinnata Steph. 191 7 Metzgeria consanguinea Schiffn. 1893 (Asia! Metzgeria crispula Herzog 1939 Austra lasiaiNeotropics) Metzgeria molokaiensis Kuwah. ' 1960' 1961 Metzgeria sinensis Chen 1955 Metzgeria harae Kuwah. 1965 Metzgeria acuta Kuwah. ' 1969' 1970 Metzgeria campbelliensis Kuwah. 1966 Metzgeria macrospora Kuwah. 1969 Metzgeria philippinensis Kuwah. 1968 Metzgeria darjeelingensis Kuwah. 1975 Metzgeria grollei Kuwah. 1968 Metzgeria macrocellulosa Kuwah. 1975 Metzgeria amakawae Kuwah. ' 1969' 1970 Metzgeria nilgiriensis Udar & Srivastava 1975 Metzgeria involvens S. Hatt. 1975 Metzgeria crassipilus (Lindb.) A. Evans 1909 Metzgeria mitrata Kuwah. 1975 (Asia! Australasia/Americas) Metzgeria orientalis Kuwah. 1978 Metzgeria novicrassipilis Kuwah. 1958 Metzgeria vittii Kuwah. 1980 Metzgeria indica Udar & Srivastava '1970' Metzgeria liaoningensis Gao 1981 1971 Metzgeria crassipaginula Kuwah. 1987 Metzgeria pandei Srivastava & Udar 1975 Metzgeria hebridensis Steph. 1917 (New Metzgeria foliicola Schiffn. 1898 (Melanesia) CaledonialNew HebridesINew Guinea) Metzgeria cristata Steph. 1917 Metzgeria innovans Steph. 191 7 Metzgeria hispidissima Steph. 1917 Metzgeria hattorii Kuwah. 1969 Metzgeria cristatissima Herzog 1951 Metzgeria kuwaharae Piippo 199 1 (Papua Metzgeria ftavovirens Colenso 1889 (New New Guinea) Zealand) Metzgeria leptoneura Spruce 1885 (Worldwide) Metzgeria disci[ormis A. Evans 1910 Metzgeria nitida Mitt. ' 1887' 1886 (L.) Dumort. 1835 (World­ Metzgeria hamati[ormis Schiffn. 1893 wide) Metzgeria australis Steph. 1889 Metzgeria decipiens (c. Massa1.) Schiffn. Metzgeria sandei Schiffn. 1898 , 1889'1890 Metzgeria papulosa Steph. 1899 M. L. So: The genus Metzgeria in Asia 175

Metzgeria Juscescens Mitt. ex Steph. 1900 (New Guinea) Metzgeria curviseta Steph. 1905 Metzgeria pubescens (Schrank) Raddi 1818 Metzgeria longipila Steph. 1914 (Europe/N America/Asia) Metzgeria colensoi Steph. 1917 Metzgeria duricosta Steph. 1917 Metzgeria concavula Pearson 1924 Metzgeria longifrondis Gao 1981 Metzgeria subhamata S. Hatt. 1955 Metzgeria rigida Lindb. '1877' 1878 (Aus­ Metzgeria parvipapulosa Kuwah. '1960' tralialNew Zealand) 1961 Metzgeria crassicostata Steph. 1889 Metzgeria nana Kuwah. '1960' 1961 Metzgeria wattsiana Steph. 1917 Metzgeria borneensis Kuwah. 1965 Metzgeria robinsonii Steph. 1917 (Philippines) Metzgeria luzonensis Kuwah. 1968 Metzgeria saccata Mitt. '1887' 1886 (Aus­ Metzgeria sharpii Kuwah. 1969 tralialNew ZealandlNew Caledonia) Metzgeria Iindbergii Schiffn. 1898 (Asia! Aus­ Metzgeria Jrancana Steph. 1917 tralasia!Pacific) Metzgeria scobina Mitt. '1887' 1886 (Melane­ Metzgeria pectinata Steph. 1917 sia) Metzgeria conjugata var. japonica S. Hatt. Metzgeria subscobina Kuwah. 1968 1955 Metzgeria submarginata M. L. So 2002 (Aus­ Metzgeria oceanica Kuwah. '1960' 1961 tralialNew Zealand) Metzgeria minuta Kuwah. 1968 Metzgeria temperata Kuwah. 1976 (Japan/N Metzgeria assamica Srivastava 1976 America) Metzgeria Jukuokana Kuwah. 1978 Metzgeria thomeensis Steph. 1891 (Africa! Metzgeria minor (Schiffn.) Kuwah. 1978 Neotropics/New Guinea) Metzgeria macveanii Kuwah. '1969' 1970

ACKNOWLEDGMENTS The author is grateful to the curators of the following herbaria for the loan of speci­ mens including types: BM, BR, CAHUp, CANB, CHR, E, EGR, F, FH, FI, G, HO, IFSBH, JE, L, LWU, M, MANCH, MELU, MU, NICH, NSW, NY, S, STR, TNS, UC, W, WELT, and WRSL; also to CANB and L for sending large collections for this study. Dr. Z. Iwatsu­ ki, the editor, Dr. J. J. Engel, and an anonymous reviewer, provided useful comments to the manuscript.

LITERATURE CITED Engel, 1. 1. 1976. Metzgeria hamata Lindb., an illegitimate name of Hepaticae. Lindbergia 3: 219- 220. Evans, A. 1910. Vegetative reproduction in Metzgeria. Annales of Botany 24: 271 - 303. Geissler, P. & H. Bischler. 1985. Index hepaticarum. Vo!. 10. Berlin, Cramer. Grolle, R. 2002. The Hepaticae and Anthocerotae of the subantarctic and temperate islands in the eastern Southern Hemisphere (90 0 E to 0°). An annotated catalogue. Journal of Bryology 24: 57- 80. Grolle, R. & M. L. So. 2002. Typifications of three old names of Metzgeria species (Hepaticae): Jungermannia Jurcata L. 1753, J linearis Sw. 1788 and J pubescens Schrank 1792. Cryp­ togamie Bryologie 23(2): 119- 121. Hattori, S. 1952. Notes on little known Japanese species of Hepaticae (1). Journal of Japanese Botany 27: 245- 246. Hattori, S. 1955. On the Hepaticae of southern Hidaka, with special reference to the Hepaticae occur- 176 1. Hattori Bot. Lab. No. 94 2 003

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Schuster, R. M. 1992. The Hepaticae and Anthocerotae of North America. VI. Chicago, Field Muse­ um. So, M. L. 2002. Metzgeria in Australasia and the Pacific. New Zealand Journal of Botany 40: 603- 627. Srivastava, S. C. & R. Udar. 1975. of the Indian Metzgeriaceae. A monographic study. New Botanist 2: I-57. Stephani, F. 1917- 1924. Species Hepaticarum 5. 766 pp. Geneve et Bale. Stephani, F. 1985. !cones Hepaticarum, microfiches. Leiden, International Documentation Company. Zhang, D. C. 1991 . A Revision of Family Metzgeriaceae from South Western China. Acta Botanica Yunnanica 13(3): 283- 289.