J Hallori 8 01. Lab. No. 95: 263- 270 (Jan. 2004)

ON SEVEN SPECIES OF THE (HEPATICAE) IN CHINA, INCLUDING ONE NEW SPECIES AND ONE NEW COMBINATION

CHIEN GAOl AND YU-HUAN WU I

ABSTRACT. The distributions of seven species of Schistochilaceae in China are discussed. A new species, minor C. Gao & Y.-H. Wu, is described from Taiwan, China. A new combina­ tion, Gottschea macrodonta (Nichols.) C. Gao & Y.-H. Wu, is proposed based on Schistochila macrodonta Nichols. owing to its lacking of underleaves.

INTRODUCTION The family Schistochilaceae of Buch (1928) was originally proposed as a segregate from the Scapaniaceae, founded for taxa supposed consistently lacking perianths, including only the genus Schistochila Dumort. which comprised about 70 species (Stephani 1909), with a number of others subsequently described, chiefly by Buch (1939) and Grolle (1966). Schuster (1963) established a new genus Paraschistochila R.M. Schust. (=Gottschea) with Paraschistochila pinnatifolia (Hook.) R.M. Schust. as type (Basionym: Jungermannia pin­ natifolia Hook.) and pointed out that Paraschistochila differs from Schistochila by: I) in­ sertion and orientation of leaves clearly incubous; and 2) underleaves almost lacking. Grolle & Zijlstra (1984) transferred the species of Schistochila without underleaves to Gottschea Nees ex Mont. (1843) of the family Schistochilaceae. There are three genera in the family Schistochilaceae, most of the species are distributed in Pacific tropical islands both in the northern and southern hemispheres, but more common in the latter (Buch 1939; Schuster & Engel 1977). Inoue (1985), Lin et al. (1992) and C. Gao (2000) recorded five, three, and three species from Taiwan, Hainan, and Yunnan, respectively. Altogether, two genera and six species have been reported from China till now. In this paper one new species is described and one new combination proposed.

Key to the genera of Schistochilaceae in China 1. Underleaves large, usually present ...... 2. Schistochila 1. Underleaves absent or sometimes minute ...... 1. Gottschea

Genus 1. Gottschea Nees ex Mont., Ann. Sci. Nat., Bot. Ser. 2, 19: 245. 1843. Lectotype: Gottschea hombroniana Mont. Bonner (1966) listed 46 species names of Gottschea. Most of the taxa are distributed in tropic areas of the southern hemisphere. In the northern hemisphere, the known range of Gottschea now includes only Indonesia, Philippines, India, Japan, Bhutan, Malaysia, Sri Lanka, Thailand, Micronesia, Papua New Guinea, and South China (Hainan, Taiwan, and Yunnan). Three species of Gottschea have been reported from China, including G.

I Institute of Applied Ecology, Chinese Academy of Sciences, Shenyang, 110016, China. 264 1. Hattori Bot. Lab. No. 95 2 004 macrodonta (Nichols.) C. Gao & Y-H. Wu, novo comb. due to its lacking of under leaves.

Key to the species of Gottschea in China I. Margins of leaves entire or rarely with small angulations...... 3. G. nuda I. Margins of leaves spinose or dentate...... 2 2. Margins ofleaves dentate...... 2. G. macrodonta 2. Margins of leaves spinose...... 3 3. Dorsal leaf-lobe subquadrate, with a triangular tooth, forming a truncate apex. . .. I. G. aligera 3. Dorsal leaf-lobe asymmetrically oblong-ovate, gradually narrowed to the end of carina ...... 4. G. philippinensis

I. Gottschea aligera (Nees & Blume) Nees in Gottsche et aI., Synop. Hepat. 17. 1844; C. Gao in C. Gao & T. Cao, Flora Yunnanica Tom. 17: 323. 2000. Jungermannia aligera Nees & Blume in Nees, Nova Acta Acad. Leop.-Caes. 11(1): 135 . 1823. Schistochila aligera (Nees & Blume) Jack & Steph., Hedwigia 31: 12. 1892. Scapania subnuda Steph., Spec. Hepat. 6: 504. 1924. Illustration: H. Inoue 1985. fig. 3: 7- 14; S. Piippo 1984. fig. 13. Habitat: On old trunks and decayed wood. Chinese specimens examined: TAIWA N: Xinzhu County, Yuanyang Lake Natural Reserve, C. Gao & T Cao 980318a (IFSBH). HAI NAN: Tongshi County, Wuzhi Shan Natural Reserve, all. 800- 1200 m, T Cao 200122 (IFSBH); Ml. Jianfengling, Pc. Chen 398a, 370b, 485a, 623a, 440e (IFSBH). Distribution: China (Taiwan, Hainan), India. Sri Lanka, Philippines, Thailand, Malaysia, Mi ­ cronesia, Papua New Guinea, Indonesia. GOffschea aligera is distinguished from other Gottschea species by 1) larger size; 2) leaf margins distinctly spinose; 3) dorsal lobe coherent with leaf in one side; 4) apex truncate; and 5) cilia short and divided.

2. Gottschea macrodonta (Nichols.) C. Gao & Y-H.Wu, novo comb. (Fig. 1: 1- 9) Schistochila macrodonta W.E. Nicholson in H. Handel-Mazzetti, Sym. Sin. 5: 29. 1930; C. Gao in C. Gao & T. Cao, Flora Yunnanica Tom. 17: 323. 2000. Type: China. NW-Yunnan, Tschiang. all. 2800-3450m. 5. Jul 1916, H. Handel-Mazzetti 9366 (not seen). Habitat: On old trunks. Specimen examined: BHUTAN, C-Mongar, near large waterfall above Namning, SE of Sengor, all. 2730 m, D. G. Long 8665 (lE). Distribution: China (Yunnan), Bhutan. Although we failed to borrow the type material from herbaria lE and H, R. Grolle (lE) sent us a duplicate specimen of Schistochila macrodonta from Bhutan with characteristics

Fig. 1. Gottschea macrodonla (Nichols.) C. Gao & Y-H. Wu. I. Plant shoot, dorsal view. 2. Plant shoot, ventral view. 3- 5. Leaves. ventral and dorsal view. 6. Leaf in silU, dorsal view. 7. Leaf in situ, showing scattered rhizoids. 8. Cells of ventral lobe margin. 9. Cells of ventral lobe base. 10. Median cells of ventral lobe. All drawn from D.G. Long 8665. JE. Scales bar: A= I cm (1 - 2), B= I mm (3 - 7), C=50.um (9- 10), D= 100.um (8). C. GAD & Y-H . WU: Schistochilaceae in China 265 266 J. Hattori Bot. Lab. No. 95 2 004

according with the type description. This specimen has no underleaves, but sometimes pa­ raphyllia are present along leaf insertion. Grolle & Zijlstra (1984) placed the species with­ out underleaves in Gottschea. Owing to its lacking of underleaves, Schistochila macrodon­ ta Nichols. is transferred here to Gottschea, as Gottschea macrodonta (Nichols.) C. Gao & Y-H. Wu.

3. Gottschea nuda (Horik.) Grolle & Zijlstra, Taxon 33: 89. 1984; Piippo, lourn. Hattori Bot. Lab. 68: 46. 1990; C. Gao in C. Gao & T. Cao, Flora Yunnanica Tom. 17 : 322. 2000. Schistochila nuda Horik., 1. Sci. Hiroshima Uniy. ser. B., diy. 2, 2: 215 . 1934. Text. fig . 38. 1934. Type: China. Taiwan: Between Shinsuiei and Shuchokyokai, Taito, Horikawa 10626 (HIRO!) Paraschistochila nuda (Horik.) Inoue, Illustrations of Japanese Hepaticae. 2: 179. 1974; M.J. Lai & J.R. Wang-Yang, Index bryoflorae formosensis, Taiwania 21: 159- 203 . 1976. Illustration: Horikawa 1934. Text - fig. 38 (1--4) as Schistochila nuda Horik., C. Gao 2000: fig. 165 (l 0- 14) as Gottschea nuda. Habitat: On old tree trunks. Chinese specimens examined: TAIWAN : Between Shinsuiei and Shuchokyokai, Horika wa 10626 (HIRO). YUNN AN: Deqin, L.5o Wang n. s. (KUN). Distribution: China, Japan, Philippines. large, green or yellow-green, dry specimens in herbarium brown-green. Leaf margin entire or with cilia formed of a single cell.

4. Gottschea philippinensis Mont., Ann. Sci. Nat. Bot. ser. 2, 19: 224. 1843. Schistochila philippinensis (Mont.) Jack & Steph., Bot. Centralbl. 60: 98. 1894. Schistochilaster philippinensis (Mont.) H.A. Miller, Phytologia 20: 319. 1970. Paraschistochila philippinensis (Mont.) R.M . Schust., Bull. Nat. Sci. Mus. Tokyo 14: 647. 1971; Lai , Publ. Dep. Bot. Uniy. Helsinki 9: 1--40. 1981. Illustration: S. Piippo 1984: 40, fig . 15: a- c as Schistochila philippinensis (Mont.) Jack & Steph.; H. Inoue 1985: 114, fig. 3: 7- 14. Habitat: On trunks of trees or rocks. Chinese specimens examined: HAI NAN: Mt. Jianfengling, Pc. Chen 125, 259, 260, 86a (I8SC). Distribution: China (Hainan, Taiwan), Philippines, Indonesia, Sri Lanka, Papua New Guinea. Stephani (1909), Horikawa (1934) and So (2003) treated Schistochila philippinensis Mont. as synonym of Schistochila aligera (Nees & Blume) Jack & Steph. However, Buch (1939) as well as Grolle & Zijlstra (1984) accepted them as two independent species. In G. aligera (Nees & Blume) Nees the leaf is narrowly lanceolate-ovate or long oblong and the dorsal lobe apex truncate, but in G. philippinensis the leaf is broad-lanceolate, the dorsal lobe broad-lanceolate, and its apex acute, gradually narrowed.

Genus 2. Schistochila Dumort., Recueil Observ. lung.: 15 . 1835. Lectotype: 5. appendiculata (Hook.) Dumort. (designated by Grolle 1966). Plant large, forming pure populations or growing with other bryophytes on tree or rock. Schistochila is distinguished by I) underleayes present; 2) whole leaf margin ciliate; and 3) dorsal leaf margin densely ciliate, with long, spreading, slender cilia formed of 1- 5 (6) narrow, highly elongated uniseriate cells. Engel (1990) established a number of 192 species of Schistochila. Most of them are C. GAO & Y.-H. Wu: Schistochilaceae in China 267 distributed in South America and Asian Pacific tropical islands. The genus Schistochila has been carefully studied especially by Schuster (1971), Schuster & Engel (1977), Grolle (1966) and So (2003). Two species, S. acuminata Steph. and S. blumei (Nees) Trevis. (s. macrodonta Nichols. belongs to Gottschea as G. macrodonta (Nichols.) C. Gao & Y.-H. Wu), have been reported from China. During our recent study, we found a further species, Schistochila minor C. Gao & Y.-H. Wu; there are thus three species of Schistochila known from China.

Key to species of Schistochila in China I. Ventral lobe of leaves large; leaf with a single wing ...... 2 I. Ventral lobe ofleaves small; leaf with paired wings...... 2. S. minor 2. Ventral lobe of leaves with several to many linear lamellae on surface; apex of dorsal lobe of leaves rounded or nearly truncate; underleaves 1/3-1 /2-bifid ...... I. S. blumei 2. Ventral lobe of leaves lacking lamellae on surface; dorsal lobe of leaves with triangularly acuminate apex; underleaves retuse or I14-bifid ...... 3. S. acuminata

I. Schistochila blumei (Nees) Trevis., Mem. Real. Istit. Lombard. Sci. Mat. Nat. ser. 3, 4: 392. 1877; Piippo, Journ. Hattori Bot. Lab. 68 : 121. 1990. Jungermannia blumei Nees in 8\ume, Nova Acta Acad. Leop.-Carol. 11(1): 136. 1823. Schistochila loriana Steph., Spec. Hep. 4: 79. 1909. Schistochilaformosana Horik., Ann. Bryol. 6: 59. 1933. Type: China. Taiwan (Formosa): Prov. Tainan, Mt. Morrison, Horikawa 9218 (HIRO!) Illustration: Horikawa 1933: 60, fig . I. as S. formosana; Piippo 1984: 35, fig . 11: d- i; Inoue 1985: Ill. fig. 1: 1- 7. Habitat: On old tree trunks and decayed wood. Chinese specimens examined: TAIWAN: Mt. Morrison, Y. Horikawa 9218 (HIRO); Xinzhou County, Yuanyang Lake Nature Reserve, a1t. 1670m, C. Gao & T. Cao 9880790, 981488, 980757, 980275, 980294,980757 (IFSBH). Distribution: China (Taiwan), Philippines, Indonesia (Java), Malaysia, Thailand, Papua New Guinea. This species is distinguished by I) the large size of the green-yellow plant; 2) leaf nar­ rowly ovate; 3) dorsal lobe ovate, apex obtuse, margins with 1- 5 (6) long cilia formed of uniseriate cells, cilia distinctly longer near the base; 4) leaf cells thin-walled, trigones small or indistinct; and 5) underleaves divided to 1/3- 1/4 their length, lobes triangular, margin with long cilia.

2. Schistochila minor C. Gao & Y-H. Wu sp. novo(subg. Isoschistochila R.M. Schust.) Fig. 2: 1- 13 Species S. sciureo (Nees) Schiffn. similis, sed differt praecipue plantis parvis, 1- 1.5 cm longis (non 3--4 cm longis), ventrali lobo conspicuo minori quam dorsali (non ventrali lobo majouri quam dorsali), atque supris medio cilii; densis copiosis, infero basin sub integer (non versus ciliato). Plants small, soft, fragile, pale green, pellucid, shiny, loosely prostrate. Stem 1- 1.5 cm long, with leaves 2.5- 3 mm wide, sparingly branched, branches usually much inferior to main axis, Radula-type. Rhizoids large for plant size, sparing toward basal portion of axis, 268 J. Hattori Bot. Lab. No. 95 2 004 scattered along ventral surface of stem, purple. Leaves imbricate, oriented incubous, cari­ nal region in section of 4- 8 cells, the leaf with two halves forming a lunate continuum, ventral and dorsal lobes in whole length 2-winged, unistratose throughout, widely laterally patent, not deflexed, perfectly transversely inserted; ventral lobe about 2/3 in size of the dorsal one, subasymmetrically ovate or oblong, apex obtuse, dorsal and ventral surface smooth, margin irregularly ciliate, cilia of 2- 6 elongate pachydermatic soft cells; dorsal lobe asymmetrically broadly ovate or oblong, 1- 1.2 mm long, 0.3- 0.5 mm wide, about 1/3 in size larger than the ventral one, apex obtuse, dorsal and ventral surfaces smooth, margin irregularly ciliate or occasionally entire in basal half; leaf cells almost subisodiametric, me­ dian cells 20- 28 X20- 35 pm, subbasal cells 22-30 X 30-45 pm, walls thin to slightly thick­ ened, trigones large and bulging, cuticle smooth; oil-bodies and chloroplasts sphaerical. Underleaves remote, bilobed to 1/3- 112 their length, about twice wide as the stem, margin with 3- 6-celled cilia. Dioecious. Androecia not seen. Gynoecia (archegonia) terminal on main axis; subgynoecial innovations absent. Mature gynoecia not seen. Type: China. Taiwan: Xinzhu County, Yuanyang Lake Natural Reserve, C. GaD & T. CaD 980318 (Holotype-IFS8H, Isotype-JE), 980455 (Paratype-IFS8H). S. minor is similar to S. siurea (Nees) Schiffn., but differs from it by I) plant small, 1- 1.5 cm long, not 3-4 cm long; 2) ventral lobe conspicuously smaller than the dorsal one, about the 1/3 size of the dorsal lobe; and 3) upper leaf margin dense ciliate, nearly smooth to the base. Distribution: Endemic to China (Taiwan).

3. Schistochila acuminata Steph., Spec. Hep. IV: 81. 1909. Schislochila rigidula Horikawa, Ann. Bryo!. 6: 60. 1933. Type: China. Taiwan: Mt. Taiheizan, Y. Horikawa 24 (HIRO, not seen). Illustration: Y. Horikawa 1933 : 61 , fig. 2. as S. rigidula; Inoue 1985 : 113. fig. 3. Habitat. On old tree trunks and decayed wood. Chinese specimens examined: TAIwA N: Xinzhu County, Yuanyang Lake Natural Reserve, alt. 1670m, C. GaD & T. CaD 980503, 980588, 980465, 980571 , 980920, 9S0337, 980793, 980402, 980283 (IFSBH). Distribution: China (Taiwan), Philippines, Indonesia, Malaysia. Plants large, 4- 6 cm long, with leaves 5- 7 mm wide, pale green to yellow green, shiny. Leaves narrowly oblong, sparingly imbricate, apex acute, margin densely ciliate, near base subentire. This species is distributed in tropical Asia.

Fig. 2. Schistochila minor C. Gao & Y.-H. Wu. I. Apical part of gynoecial shoot, ventral aspect. 2. Apical part of shoot, dorsal view. 3, 4. Leaves of ventral and dorsal view. 5. Underleaf. 6. Cross section of leaf. 7. Apex of underleaf. 8, 9. Cross section of the keel part of leaf. 10. Cilia from middle of ventral leaf-lobe. 11. Cilium from apex ofunderleaf. 12. Apical cells from leaf-lobe, with oil bodies. 13. Median cells from leaf­ lobe, with oil bodies. 14. Median cells from leaf-lobe. All drawn from type Gao & Cao 980318, IFSBH. Scales bar: A= I mm (1 - 2), 8 = 0.1 mm (3-4), C= 0.2mrn (5, 7), D= 50pm (8- 14). C. GAO & Y-H. Wu: Schistochilaceae in China 269 270 1. Hattori Bot. Lab. No. 95 2 004

ACKNOWLEDGMENTS We appreciate the National Nature Science Foundation of China (Project: 30 I 00025, 30270114) and the Chinese Academy of Sciences (Project: YTZ 010506) for support of this research. The senior author gratefully acknowledges Prof. Lai Ming-Jou's hospitality during his visit to Taiwan in 1998. We thank Prof. Lin Pang-Juan, Zhang Li and Zhu Rui­ Liang for their providing specimens and references. Thanks are due to Prof. R. Grolle for helpful comments and suggestions. We also thank Prof. D.H. Vitt for improving the Eng­ lish manuscript.

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