Journal of Bryology (2008) 30: 264–270

Macromitrium cavaleriei, a little known from China with four new synonyms (Musci: )

SHUI-LIANG GUO1 and SI HE2

Shanghai Normal University, China and 2Missouri Botanical Garden, USA

SUMMARY

Macromitrium cavaleriei Cardot & The´r. in The´r. is a little known Chinese moss of the family Orthotrichaceae. The species, which, forms dense mats with creeping stems, varies much in its branch leaf shape and in the laminal cell papillosity of the branch and perichaetial leaves. In the present paper, Macromitrium cavaleriei is re-evaluated taxonomically, including the new synonymizations of M. gebaueri Broth., M. sinense E. B. Bartram, M. syntrichophyllum The´r. & P. de la Varde, and M. syntrichophyllum var. longisetum The´r. & Reimers from China. The synonymizations extend the distribution of M. cavaleriei from Guizhou to 15 provinces and one municipality, mainly in the area to the South of the Yangtze River, Southwest China, and Taiwan. Additionally, M. cavaleriei is reported as new to the moss flora of Japan.

KEYWORDS: , nomenclature, synonymization, , variation.

INTRODUCTION TAXONOMIC TREATMENT

The genus Macromitrium Brid. contains 44 valid species in Macromitrium cavaleriei Cardot & The´r. in The´r., Bull. China according to recent studies (Redfearn, Tan & He, Acad. Int. Geogr. Bot. 16: 40. 1906. Type: China. Kouy 1996; Xiong, 2000; Guo et al., 2007). Among them, Teche´ou (Guizhou), November 1902, J. Cavalerie 833 Macromitrium cavaleriei Cardot & The´r. in The´r. has been (lectotype, PC-0083631, designated here; isolectotypes, virtually unknown since its inception although it was the PC-0083629, BM-000919466) (Figs. 1 and 2). first species of Macromitrium described from the mainland Macromitrium gebaueri Broth., Symb. Sin. 4: 72. pl. 1 f. China (The´riot, 1906). During the course of our study on 10. 1929, syn. nov. Type: China. Yunnan, An Ba¨umen in der Chinese Macromitrium, we examined the type specimens of wtp. St. des Schweli-Salween-Scheidegebirge, alt. 2000– this taxon as well as the types of relevant names, and came 2800 m, 1914, Gebauer s.n. (holotype, H-BR-2572001). to the conclusion that the following four taxa are new Macromitrium sinense E. B. Bartram, Ann. Bryol. 8: 13. f. synonyms of M. cavaleriei: M. gebaueri Broth., M. sinense 7. 1936, syn. nov. Type: China. Kweichow Prov. (Guizhou), E. B. Bartram, M. syntrichophyllum The´r. & P. de la Varde, Fan Ching Shan, alt. 2250 m, on bark of tree, 7 September and M. syntrichophyllum var. longisetum The´r. & Reimers. 1931, S.-Y. Cheo 430 (lectotype, PC-0083730, designated An extensive study of many specimens under various names here; isolectotype, F). of Macromitrium (Guo, Enroth & Koponen, 2007) has Macromitrium syntrichophyllum The´r. & P. de la Varde, allowed us to re-evaluate the taxonomy and distribution Rev. Bot. Bull. Mens. 30: 347. 1918, syn. nov. Type: China. of M. cavalerieri. In this paper, a detailed taxonomic Leoufang, alt. 900 m, October 1910, Courtois 332 (lectotype description, illustrations, new synonymy and a short PC-0083731, isolectotype BM-000919468) – Tchan kia po discussion of related species are presented. The study shows (prov. Nagan hoi), October 1910, Courtois 335 (syntype, that M. cavaleriei has a wide distribution range in China, not seen). including 16 provinces/municipalities: Anhui, Chongqing, Macromitrium syntrichophyllum var. longisetum The´r. & Fujian, Guangdong, Guangxi, Guizhou, Henan, Hubei, Reimers, Hedwigia 71: 55. 1931, syn. nov. Type: China. Hunan, Jiangsu, Jiangxi, Sichuan, Taiwan, Xizang, Prov. Kwangsi (Guangxi), Yao-shan, alt. 2000 m., 5 Yunnan and Zhejiang. January 1929, S. S. Sin 2075 (holotype, PC-0083734).

# British Bryological Society 2008 Received 17 February 2008. Revision accepted 14 August 2008 DOI: 10.1179/174328208X300705 MACROMITRIUM CAVALERIEI A LITTLE KNOWN MOSS FROM CHINA 265

Figure 1. Macromitrium cavaleriei. 1–16: branch leaves. 17–26: inner perichaetial leaves. 27–28: dry capsules. 29–33: wet capsules. 34–35: calyptrae. 36–37: stem leaves. (1–3, 17–20, 28–31, 34, from lectotype of M. cavaleriei, J. Cavalerie 833, PC-0083631; 4–6, 21–22, 36–37, from lectotype of M. syntrichophyllum, Courtois 332, PC-0083731; 7–10, 25–26, 33, from holotype of M. gebaueri, Gebauer s.n. H; 11–16, 24, 31–32, 35, from lectotype of M. sinense, S.-Y. Cheo 430, PC). Scales: A51.00 mm (1–35); B50.55 mm (36–37). 266 S.-L. GUO AND S. HE

Figure 2. Macromitrium cavaleriei. 1–4: lower cross-sections of leaf. 5: middle cross-sections of leaf. 6–9: upper cross-sections of leaf. 10: api- cal laminal cells of perichaetial leaf. 11–14: basal laminal cells of inner perichaetial leaf. 15–17: lower laminal cells of inner perichaetial leaf. 18–21: median laminal cells of inner perichaetial leaf. 22–23: upper laminal cells of inner perichaetial leaf. 24–27: basal laminal cells of branch leaf. 28–30: lower laminal cells of branch leaf. 31–32: median laminal cells of branch leaf. 33–36: upper laminal cells of branch leaf. 37: basal laminal cells of stem leaf. 38: median laminal cells of stem leaf. 2, 5–6, 10–11, 17–18, 22, 24, 28–29, 31, 33, from lectotype of M. cavaleriei, PC-0083631; 1, 7, 8, 12, 16, 19, 23, 25, 30, 32, 34, 37–38, from lectotype of M. syntrichophyllum, PC-0083731; 3-4, 9, 13, 20, 26, 35, from lec- totype of M. sinense, PC; 14–15, 21, 27, 36, from holotype of M. gebaueri,H).Scales:A568 mm (1–2, 5–8, 10–12, 16–19, 22–23, 25, 30,32– 34, 37–38); B5100 mm (3–4, 9, 13–15, 20–21, 24, 26–29, 31, 35–36). MACROMITRIUM CAVALERIEI A LITTLE KNOWN MOSS FROM CHINA 267

Plants in dense mats, medium-sized, brownish green, bulging, thin-walled and hyaline, with 2–4 small papillae dark brown below, yellowish green to dark green above. per cell or almost smooth, and the lower laminal cells are Stems creeping, with erect branches, 5–25 mm long, densely rectangular, irregularly rectangular, or rhomboidal, and foliate, often with several short branchlets at upper part of smooth or unipapillose. The inner perichaetial leaves are branches. Stem leaves suberect to strongly reflexed when oblong-lanceolate, narrowly to long acuminate and often moist, oblong-lanceolate to triangular-lanceolate, 0.6–1.4 aristate at apex. Finally, the calyptrae are campanulate and 6 0.2–0.3 mm, acuminate at apex; upper and median hairy. laminal cells subquadrate or rounded quadrate, 8.0– 10.0 mm wide, bulging, smooth or slightly papillose; lower DISCUSSION laminal cells slightly elongate; basal laminal cells rectan- gular, 20.0–30.0 6 4.0–6.0 mm, moderately thick-walled. Macromitrium sinense E. B. Bartram, M. syntrichophyllum Branch leaves strongly crisped, somewhat contorted, some- The´r. & P. de la Varde, M. syntrichophyllum var. longisetum times with upper portion strongly involute when dry, widely The´r. et Reimers. and M. gebaueri Broth. were originally spreading, occasionally squarrose-recurved when moist, described as new from China, and little information could linear-lanceolate to ovate-lanceolate, (1.5–)2.5–4.2 6 0.3– be found about them after they were published. Having 0.5 mm, mostly acuminate or narrowly acuminate, some- checked their types, we found that all four taxa could be times broadly acuminate, often keeled, incurved at apex, accommodated within M. cavaleriei. Compared with those revolute at one side of margin near base, plane or slightly of the types of M. cavaleriei, the branch leaves of the plicate to plicate, occasionally with short, oblong-lanceo- lectotype of M. sinense are slightly shorter, with more late leaves, more or less obtuse at apex; leaves of secondary broadly acuminate apices and moderately pluri-papillose branches becoming smaller toward base; margins entire, median and upper laminal cells. The branch leaves of the sometimes finely or indistinctly serrulate in upper part due lectotype of M. syntrichophyllum are somewhat shorter with to extruding papillae; costae single, yellowish brown, acuminate-obtuse apices, while in the type of M. syntricho- ending a few cells below apex, occasionally percurrent or phyllum var. longisetum, the branch leaves are similar in shortly excurrent; median and upper laminal cells quadrate length to those of M. cavaleriei but some leaves have to subquadrate, 8.0–12.0 6 8.0–10.0 mm, slightly to acuminate-obtuse apices. The holotype of M. gebaueri is moderately bulging, thin-walled, hyaline or slightly rather similar to the lectotype of M. cavaleriei except that obscure, mostly with 2–4 small, low papillae per cell, its inner perichaetial leaves have moderately strongly occasionally almost smooth; lower laminal cells rhomboi- unipapillose laminal cells in their upper and median dal, rectangular to sublinear, 15.0–30.0 6 4.0–6.0 mm, portions. The variation of the types of M. cavaleriei, M. nearly hyaline, somewhat thick-walled, smooth or unipa- syntrichophyllum var. longisetum, M. syntrichophyllum, M. pillose; basal laminal cells near margins at one side often gebaueri and M. sinense is mainly found in branch leaf differentiated, regularly rectangular, pellucid with thinner shape, laminal cell papillosity of the branch and perichae- walls, occasionally basal juxtacostal cells irregularly rec- tial leaves, but the variations are overlapping. Therefore, we tangular, thin-walled, pellucid. Autoicous. Perichaetia sub- consider the latter four names as new synonyms of M. terminal on branches. Inner perichaetial leaves oblong- calaveriei Cardot & The´r. lanceolate, often plicate in basal part, to 2.5 mm long, Chinese collections of M. cavaleriei show considerable narrowly acuminate to long acuminate or aristate at apex; morphological variation. When moist, the branch leaves are costae excurrent; laminal cells mostly longer than wide, mostly widely spreading, but occasionally recurved-squar- hyaline, smooth, sometimes unipapillose in median and rose ones are found. The leaf margin is often narrowly upper portions. Calyptrae campanulate, medium-sized, revolute below on one side, but sometimes revolute also with numerous yellowish or yellowish brown hairs. Setae above (Zhejiang, P.-C. Wu, 810, MO). The leaves are (2–)4.025.0(210.0) mm long, smooth, purple or brown, usually oblong- to linear-lanceolate and acuminate or twisted to straight. Capsules erect, ovoid, oblong-ovoid to narrowly so, but sometimes they are shortly ovate- shortly cylindrical, 1.4–1.8 6 0.7–0.9 mm, slightly con- lanceolate with acuminate-obtuse or obtuse to acute apices stricted at mouth when old; annuli weakly differentiated, (Guizhou, M. Bourell, 4775, MO), or shortly lanceolate with a row of purplish, smaller exothecial cells surrounding with gradually acuminate apices in the lower part of the mouth; opercula conic-rostrate, 0.7–0.8 mm long; exostome secondary branches (Anhui, Wan Zong-Ling & Luo Jian- teeth lanceolate, with obtuse apices, up to 120 mm long, Xin, 9026, MO). The upper margins are usually entire, but densely papillose throughout. Spores minutely papillose, sometimes finely serrate from extruding papilliae (Yunnan, 15–30 mm in diameter. Redfearn et al., 693, MO), the median and upper laminal Macromitrium cavaleriei is characterized by its suberect cells are often inflated or conically bulging and slightly to strongly reflexed, oblong- to triangular-lanceolate stem pluripapillose or mammillose (Sichuan, J. Giraldi accepit leaves with acuminate apices. The branch leaves are linear- 1896, F), but may be small, flat, obscure and distinctly lanceolate to ovate-lanceolate and mostly acuminate or pluripapillose (Guizhou, Cavaleriei, no. 1931, PC). The narrowly acuminate, but sometimes broadly acuminate at basal laminal cells are mostly smooth or weakly unipapil- the apex. Its median and upper leaf laminal cells are lose, but sometimes more distinctly unipapillose (Guangxi, 268 S.-L. GUO AND S. HE

Figure 3. Macromitrium ferriei. 1–8: branch leaves. 9–13: inner perichaetial leaves. 14: stem leaf. 15: branch leaf apex. 16: basal cross-section of branch leaf. 17: upper cross-section of branch leaf. 18: middle cross-section of branch leaf. 19: upper laminal cells of branch leaf. 20: basal laminal cells of branch leaf. 21: lower laminal cells of branch leaf. 22: upper laminal cells of inner perichaetial leaf. 23: median laminal cells of inner perichaetial leaf. 24: basal laminal cells of inner perichaetial leaf. 25: calyptra (all from lectotype of M. ferriei, PC-0083650). Scales: A50.40 mm (1–14); B51.0 mm (25); C568 mm (16–24); D50.2 mm (15). MACROMITRIUM CAVALERIEI A LITTLE KNOWN MOSS FROM CHINA 269

S. Y. Cheo, 2754, MO). Even in the same specimen, branch Many specimens previously identified as M. ferriei in leaves with differently papillose laminal cells could be found China are actually M. cavaleriei. Therefore, M. cavaleriei is (Taiwan, James R. Shevock, 18153, MO). Inner perichaetial rather common in China, while M. ferriei is rarer than leaves show relatively stable characters, but although the earlier thought. Macromitrium cavaleriei is new to Japan laminal cells are mostly smooth, they are occasionally (cf., Noguchi, 1967), where it is probably widely distrib- unipapillose in the median and upper parts. The setae are uted. Populations of M. cavaleriei are exclusively epiphytic frequently 4.0–5.0 mm long but may be up to 10 mm or and saprophytic in China, while in Japan, some are shorter than 2 mm (Fhjiang, H. H. Chung, B351,PC). saxicolous. Macromitrium cavaleriei is closely related to M. ferriei Selected specimens examined (Macromitrium cavaleriei): Cardot & The´r. and is probably included within the latter in CHINA: Anhui: Mt Huangshan, Z.-L. Wan & J.-X. Luo Japan by Noguchi (1967, 1989), who considered M. ferriei a 9026 (MO, PE); Jinzhai Co., Z.-M. Zeng T0272 (MO). highly variable species, especially in its branch leaves. Chongqing: Nanchuan Co., M.-Z. Wang 860390 (MO, FH, According to Noguchi’s description, Macromitrium ferriei as M. ferriei). Fujian: Fuzhou (Foochow) City, H.-H. is characterized by a large-celled area in the lower part of its Chung B351 (PC); Buong Kang, Yenping, H.-H. Chung leaves, where the cells have thick walls and few or no B90-b (PC, as M. syntrichophyllum). Guangdong: Fengshun papillae. This area extends upwards with cells size gradually Co., S. He 40329 (MO); Sha Lo Shan, Y. W. Taam (H). decreasing and the number of papillae gradually increasing. Guangxi: Sanjiang (San Kiang Hsien) Co., S.-Y. Cheo 2754 In fact, the large-celled area in the lower part of the branch (MO); Rongshui Co., Z.-Y. Li 89125 (MO). Guizhou: leaves exists also in M. cavaleriei. Noguchi (1967) once Fanjingshan Nature Reserve, Bourell 4775 (MO); Jiangkou noted that the median and upper leaf laminal cells of M. Co., C. Gao 32530 (MO). Henan: Xixia Co., J.-X. Luo 233 ferriei collections from Kinosaki are hexagonal, with thin (MO). Hubei: Mt. Shennongjia, P.-C. Wu 195, 393 (MO). walls, several papillae and somewhat obscure, as in the type Hunan: Linling & Sinning, Handel-Mazzetti 11308 (H-BR, of M. ferriei, while a specimen from Mt. Yanaze has rather as M. japonicum); Badagong shan, Virtanen 50043 (H, as hyaline median laminal cells with thick walls and few M. ferriei). Jiangxi: Mt Lushan, P.-C. Chen 273 (MO, PE); papillae. Noguchi (1967) thought that this character divides P.-C. Wu 003 (MO, PE). Sichuan: Wenchuan Co., Allen (M. ferriei) into two species. He also noted that, contrasting 6722 (MO); Nanping Co., Allen 7194 (MO). Taiwan: with the short branches of the type specimen and many Taichung Co., Shevock 18153 (MO, as M. ferriei); Ilan other collections, specimens from Mt. Yanagi-take, Co., C.-C. Chuang 2024 (NY, as M. comatum). Xizang: Miyazaki Prefecture and other high elevation localities in Linchih Co., Y.-G. Su 2573 (MO); Meto Co., Y.-G. Su 4782 Japan have branches up to 10 mm long, and seem to belong (MO). Yunnan: Kunming City, Redfearn et al. 1983 (MO); to a different taxon. However, Noguchi thought these Yangbi Co., Redfearn 693 (MO). Zhejiang: Deqing Co., P.- differences are merely sporadic variations. Therefore, we C. Wu 1265 (MO); Lin-An Co., P.-C. Wu 1151 (MO, PE). believe that Noguchi’s circumscription of M. ferriei JAPAN: Honshu: Mt Rakan, Hiroshima, 7 May 1961. H. includes M. cavaleriei. Based on our examination of the Ando (H3090253); Taishaku-Kyo, Hiroshima, 29 March type and a large number of Chinese and Japanese 1968, H. Ando (H3090254). Kyushu: Miyazaki-ken, Nishi- specimens of M. ferriei, the latter is a distinct species morokata-gun, Mt, Shickikuma, 24 November 1980, Z. (Fig. 3) that could be distinguished from M. cavaleriei Iwatsuki (H3090257); Higo, Aida, 28 October 1928. K. (Figs. 1 and 2) by its obscure upper and median laminal Mayebara (H3090194). Okinawa: Kunigami-mura, cells with several distinct papillae, its shorter and oblong- Yonaha-dake, K. Saito (H3090258). Shizouka: Pref. lanceolate inner perichaetial leaves, and its branch leaves Honkawane-cho, Kanzo, Z. Iwatsuki (MO). with obtuse, subacute or broadly acuminate apices (Table 1) ACKNOWLEDGEMENTS Macromitrium ferriei Cardot & The´r., Bull. Acad. Int. Geogr. Bot. 18: 250. 1908. Type: Japan. Oshima, Yowan We acknowledge financial support by the State Dake, 18. Oct. 1899, J. B. Ferrie s.n. (Lectotype, PC- Scholarship Fund of China for the senior author’s visit 0083650, designated here; isolectotypes, PC-0083649, PC- to the University of Helsinki in 2003–2004, the Scientific 0083651). Research Foundation for the Returned Overseas Chinese

Table 1. Distinction between M. cavaleriei and M. ferriei.

Character M. cavaleriei M. ferriei

Branch leaves To 3.2 mm long, narrowly acuminate to acuminate, To 2.0 mm long, obtuse, subacute, occasionally broadly acuminate to broadly acuminate Median and upper laminal Pellucid, collenchymatous, bulging, with 3–5 small Rather obscure, strongly pluripapillose cells of branch leaves papillae or sometimes smooth Inner perichaetial leaves To 2.5 mm long, abruptly constricted from an oblong To 1.5 mm long, ovate- or triangular-lanceolate, base to a long acuminate or aristate apex gradually narrowed to a shortly acuminate apex 270 S.-L. GUO AND S. HE

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SHUILIANG GUO, College of Life and Environmental Sciences, Shanghai Normal University, Shanghai 200234, China. E-mail: [email protected] SI HE, Missouri Botanical Garden, PO Box 299, St Louis, MO 63166, USA. E-mail: [email protected]