J Hattori Bot. Lab. No. 100: 41- 52 (A ug. 2006)

NEW SYNONYMS AND COMBINATIONS IN THE GENUS PTYCHOMITRlUM (: )*

TONG CA01,2 , BENRONG ZUOI,2 , SHUI-LIANG GUO I,3 , JAAKKO HYVONEN3 AND VnVI VIRTANEN4

ABSTRACT. Based on a comparison of type and other specimens, patens (Miill. Hal.) Par. and P. obtusifolium (Borth.) Par. are reduced to synonymy of P. sellowianum (Miill. Hal.) Jaeger, and P. papil/osum (Herz.) Card. is reduced to synonymy of P. chimborazense (Mitt.) Jaeger. Three new combinations are proposed: P. gardneri var. angustifolium (Nog.) Cao & Guo, P. sellowianum var. angustifolium (Broth.) Cao & Guo, and P. hieronymi var. obtusifolium (Ther.) Cao & Guo. Pty­ chomitriumferrugineum (Herz.) Broth. is described in detail based on the holotype.

KEy WORDS : Bryopsida, New combination, Ptychomitricaeae, Ptychomitrium, Synonym, Typifica­ tion.

INTRODUCTION Ptychomitrium was elevated to the generic level by Fiirnrohr (1829). Since then, 145 taxa have been recorded (Cao & Guo 2001, Crosby, Magill & Bauer 1992, Fiirnrohr 1829, Guo et al. 2000, Schiavone & Biasuso 1997, Wijk et al. 1964, 1969). In our previous stud­ ies, we have reported two new species, 19 new synonyms, and the geographical distribution pattern of the genus (Cao & Guo 2001; Cao & Koponen 1997; Cao & Vitt 1994; Cao, Guo & Vitt 1995, 2001 , Cao, Guo & Yu 2003a b; Guo & Cao 1999, 2000a, b; Guo et at. 2000). In the present paper, we report three synonyms and three combinations of Ptychomitrium based on our study of herbarium specimens from BM, H, lE, NICH, and S. Ptychomitrium ferrugineum (Herz.) Broth. is also described in detail based on the type material from lE.

NEW SYNONYMS Ptychomitrium sellowianum (Miill. Hal.) laeger., Ber. S. Gall. Naturw. Ges. 1872-1873: 104.1874. (Ad.l: 382). (Figs. 1- 19) ==Brachysteleum sellowianum Miill. Hal., Syn. 1: 769. 1849. Type: Patria. Brasilia, in Campis Mon­ tevidensibus: Sellow (isotype: H!). = Brachysteleum depressum Miill. Hal., Hedwgia 38 : 153. 1899. == Ptychomitrium depressum (Miil!. Ha!.) Par. , Ind. Bryo!. Supp!. 289. 1900. Type: Natal, Jammeriappen, 1. Dittrich 1898 19. Hb. Ar­ boreti Zoeschen-Dieck. (isotype: H!) (fide Guo & Cao, 2000a). * The project was supported by National Natural Science Foundation of China (Project No. 30370111 , 30570121). I College of Life and Environmental Sciences, Shanghai Normal University, Shanghai 200234, China. 2 Institute of Applied Ecology, Chinese Academy of Sciences, Shenyang 110016, China. 3 Biology, P. O. Box 65 , FIN-00014 University of Helsinki, Finland. 4 Botanical Museum, P. O. Box 7, FIN-00014 University of Helsinki, Finland. 42 1. Hattori Bot. Lab. No. 100 2 0 0 6

= Ptychomitrium weberbaueri Broth., Bot. Jahrb. 56 (Beibl. 123): 10. 1920. Type: Prov. Sandia: San­ dia; Felsen; 2100m (615) (holotype: H!) (fide Guo & Cao, 2000a). = Brachysteleum vernicosum Miill. Hal. , Linnaea 43 : 450. 1882. '" Ptychomitrium vernicosum (Mull. Ha!. ) Kindb., Enum. Bryin Exot. 94. 1888. Type: Argentinia Tucumanensis, Siambon. et in Cuestis borealibus supra Oran cum Leptodontio rhacomitrioidi (isotype: JE!, S! , BM! , lectotype S!) (fide Schiavone & Biasuso, 1997). = Brachysteleum patens Mul!. Hal., Bull. Herb. Boiss. 6: 108. 1898. '" Ptychomitrium patens (Miill. Hal.) Par. Ind. Bryol Suppl. 289, 1900, syn. novo Type : Brazil, Serra Itatiaia, 1800m, altum, ad rupes, Martio 1894: E. Ole, 1829 (isotype: H!). = Glyphomitrium obtusifolium Broth., Bih. K. Svensk. Vet. Ak. Hand!. 26 Afd. 3(7): 20. 1900. '" Pty­ chomitrium obtusifolium (Borth.) Par., Ind. Bryol. ed. 2, 4: 132. 1905, syn. novo Type: Brazil, Rio Grande do SuI.: Porto Alegre, Pedras Brancas, ad saxa caespites parvos efformans (n. 95 ex p) (holotype: H!). medium-sized, 1- 3 cm high. Leaves crisped and strongly keeled above when dry, spreading to erect-spreading when moist, linear-lanceolate from an oblong base, 3-4.5 mm long, plicate below; leaf margins entire, thickened to 3 or 4-stratose, revolute on both sides below; costa single, strong, up to the apex; lamina cells smooth, bistratose above, unistratose below; upper lamina cells small, irregularly quadrate, 5- 7 pm wide; me­ dial lamina cells short-rectangular to rectangular, 8- 14 pm long, 5-6 pm wide; basallami­ na cells near margins short-rectangular to rectangular, 9- 30 /1m long, 8-12 /1m wide, those along costa 30- 81 /1m long, 9- 16 /1m wide, with thin and straight walls. Setae short, erect, 2-4 mm long. Capsules elongate-ovoid to ovoid; peristome teeth lanceolate, 2 or 3 clefts nearly to base, densely papillose; annulus transparent, well developed; opercula with long erect beak. Spores finely papillose, 12- 15 pm. Calyptra mitrate, plicate, covering middle or lower part of capsule. When Brotherus (1900) reported P obtusifolium (as Brachysteleum obtusifolium) from Brazil, he considered P obtusifolium similar to P sellowianum, and pointed out that the for­ mer differed from the latter only by its obtuse leaf apices and smaller capsules. Our exami­ nation of these two types and other specimens from H, S, and FH showed that the differ­ ences in leaf apex and capsule size are not obvious. Ptychomitrium patens was reported by Muller (1898) (as Brachysteleum patens) based on the collection of E. Ule from Brazil. Having compared the type specimens of P patens and P sellowianum in H, we found that P patens is identical to P sellowianum except that its leaf bases are not distinctly plicate and its basal lamina cells are slightly sinuous. We consider that Ptychomitirum obtusifolium (Broth.) Par. and P patens (Miill. Hal.) Par. are both synonymous with P sellowianum (Mull. Hal.) Jaeger. Ptychomitrium sellowianum (Miill. Hal.) Jaeger. has been recorded from Argentina,

Figs. 1- 19. Ptychomitrium sellowianum (Mull. Ha!.) Jaeger.- 1- 5: Leaves; 6- 7, 9: Transverse sections of upper part of leaf; 8: Transverse section of basal part of leaf; 10: Ce ll s on the lower part of lea f; 11 - 13 : Basal leaf cells along costa; 14- 16: Basal leaf cells near margins; 17- 19: Capsules. (1 - 2, 6, 11 , 14, 19 drawn from isotype of P sellowianum; 3, 7, 12, 15 , 17 from holotype of P obtusifolium; 4, 5, 8- 10, 13 , 16, 18 from isotype of P patens). Line scale: A=0.5 mm (figs. 1- 5, 18); B= 0.5 mm (figs. 17, 19); C=20 ,um (figs. 6- 16). T. C AO ET AL.: New synonyms and combinations in Ptychomitrium 43

UUUUU w ~~ Dc nn o OoOOODOOO'?,QO ~6 ~ ~~n 0 (\ " 9 OOOooooOOOQJ UUUl 11 00 11 I ABC UODC 13 12 D DC 14 00nO [ n nr UUUL Dooe OOOt n~qr 15 19 18

2 3 4 5 44 1. Hattori Bot. Lab. No. 100 2 0 0 6

Brazil, Bolivia, Colombia, Uruguay, Paraguay, Peru, and South Africa, with a disjunction between South America and South Africa (Guo & Cao 2000a). This species is somewhat similar to P. crispatum (Hedw.) Jaeger., which can, however, be distinguished from P. sell­ owianum by having: I) leaves not revolute on both sides below; 2) short-rectangular to rec­ tangular basal lamina cells; and 3) bistratose leaf margins on the upper part. Specimens examined: Argentina- Tucuman, Camion Taf'del valle. no. 7615 , Dec. 1946. (FH). Brazil- Catharina, Turbarao ad rupes prope Conconhaz, E. Ule, no. 11 , Oct., 1889 (S); Catharina, E. Ule, no. 32902, 1889 (JE); Catharina, E. Ule, no. 845 (H); Sao Pauo, V. Schiffiner, 22. 5. 1901 (S); Sao Paulo, V. Shiffner, no. 960 (H); Theresopolis, F. Holnal, I!. 3. 95. (H); Theresaopolis, F. V. Hokual 1809 (H); Theresaopolis, Duson, 221711902 (H); Sede parque Zateoual Orgair nits, C. Hizziui, 1952.9.8 (FH); Serra dos orgaos ad rupes oktober, E. Ule, 1896 (S); Estado Rio de Janeiro, Mozi de Curces, aufFelesn, E. Froehlich, no. 14 (S); Serra Geral, E. Ule no. 538 (H); Sarra Geral. E. Ule, no. 539, Apr. 1889, (H); Sarra dos Orgaos E. Ule, no. 2057. (H); Bonssisusao, J. Weir, no. 54 (H); Uruguay - N. Garcia Zorr6n, no. 3270 (PC); W. G. Heata, PI. uruguayensis, no. 81047 (JE); Mowberiveo, no. 46, 1898 (JE); Sierra dr lao Animas Terram. Felippone no. 1250, 1358, 1393, 928 (H). Santa Clara no. 426 (FH); Peru - Sandia; Felsen; 2100m (615), Holotype of Ptychomitrium we­ berbaueri Broth (H!); Paraguay - Balanse 1878-84 (H); South Africa- Natal, JammerJappen: J, Dit­ trich 1898 Ig. Hb. Arboreti Zoeschen-Dieck (Isotype of Brachysteleum depressum Mull. Hal. : H); J. M. Wood no. 7123 (H); Swaziland: Mababane, E. S. Keny, 897 (H); Bolivia - Au Fels blaikeu im bergwall bei famaipata (ortcord) Th. Herzog, Dec., 07 (JE).

Ptychomitrium chimborazense (Spruce ex Mitt.) Jaeger. Ber. St. Gall. Naturw. Ges. 1872- 73: 103. (1874) (Ad.l: 377) (Figs. 20-37) ,=Glyphomitrium chimborazense Spruce ex Mitt., J. Linn. Soc. Bot. 12: 107. 1869. Type: Ins. Sti Vin­ centis, Herb. Hook. Andes Quitenses, Chimborazo, Jameson, et ad pedem orientalem pracipuse­ cus secus oppidum Guano in Saxis (8500-11000 ped.), Spruce, no. 136 (lectotype: NY! selected here, isotypes: BM! E!). = Ptychomitrium buchtienii Broth., Mitt. Inst. Allg. Bot. Hamberg, 8: 403. 1931. Type: Sud-Peru: Arequipa, 2000-2500 m (E. Guenther et O. Buchfield no. 1976) (holotype: H!) (fide Deguchi 1987). = Glyphomitrium cochabambae Herz., Beih. Bot. Centralbl. 26(2): 63 . 1909.,=Ptychomitrium cochabambae (Herz.) Card. Rev. Bryol. 40: 44.1913. Type: Bolivia: An Felsen bei Cochabamba, ca. 2800 m, January 1908, leg. Th. Herzog (holotype: JE!; isotype: FI!). (fide Deguchi, 1987). = Brachysteleum emersum Miill. Hal., Linnaea 43: 448. 1882. ,=Ptychomitrium emersum (Miill. Hal.) Kindb., Enum. Bryin. Exot. 94. 1888. Type: Argentinia subtropica, Cuesta de pinos, ubi forma elegans nitidoviridis viget; in Monte Nevado prepe Salta, 11,000 ped. altum, ubi forma Sordida habitat. Catamarea, in convalle excelsa Granadillas in Valles (Vayas) altos supra yakutula (form sordida), inito Febr. 1872. (lectotype: BM! selected here, isotype: BM! S! JE!) (fide Crum

Figs. 20- 37. Ptychomitrium chimborazense (Mitt.) Jaeger.-20: Transverse section of upper part of leaf; 21: calyptra and capsule; 22- 23: Leaves; 24: Basa11eaf cells near margins; 25 : Basal leaf cells along costa; 26: Spores; 27 : Capsule (figs. 20-27 were drawn from lecto­ type of P. chimborazense in BM). 28 : Spores; 29, 32: Upper transverse section of leaf; 30: Middle transverse section of leaf; 31, 33: Basal laminal cells; 34: Calyptra; 35: Capsule; 36- 37: Leaves (figs. 28-37 were drawn from holotype of P. papillosum in JE). Line scale: A=0.5 mm (figs. 21-23, 27, 34-37); B=50 J.1m (figs. 20, 24- 26, 28- 33). T. CAO ET AL.: New synonyms and combinations in Ptychomitrium 45

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27

33

11 AB 46 J. Hattori Bot. Lab. No. 100 200 6

1984). = G/yphomitrium papillosum Herz., Beih. Bot. Centralbl. 26(2): 64. 1909. == Ptychomitrium papillo­ sum (Herz.) Card., Rev. Bryol. 40: 44. 1913, syn. novo Type: Bolivia, An Felsen bei Huaillas (Cordillera real), ca. 3800m; lanuar, 08 (holotype: lE! ; isotype: H!). Plants small, 1- 2 cm high. Leaves irregularly crisped and keeled when dry, usually concave when moist, oblong-lanceolate, with an oblong, weakly plicate and sheathing base, gradually narrowed via long subulate portion into a sharp tip; leaf apex cucullate or some­ what cucullate; lamina bistratose above, unistratose below; leaf margins incurved above, re­ curved on both sides below at base; lamina cells distinctly papillose on both surfaces ex­ cept at leaf base. The upper lamina cells obscure, 7- 811m wide, basal lamina cell enlarged, 25- 3511m long, 6- 811m wide, thin-walled and hyaline. Capsules ovate to oblong, with wide orifice, slightly contracted below the orifice; operculum with a long beak; annulus well developed, with ovate-oblong cells; peristome teeth 16, imperfectly and irregulately forked, densely papillose throughout on both surfaces. Spores 10-1411m, minutely papil­ lose. Calyptra mitrate, lobed at base. Mitten (1869) reported P chimborazense (as G/yphomitrium chimborazense) based on the collection of R. Spruce from Andes Quitenses (Ecuador). The species is distinguished by having: (I) leaves lanceolate from an oblong sheathing base, with an acuminate tip and cucullate apex; (2) papillose lamina cells on both surfaces, except at leaf base; (3) basal lamina cells hyaline with rather thin walls; (4) capsule ovate with a wide orifice and con­ stricted below the orifice; and (5) peristome teeth irregularly forked and much perforated. Crum (1984) has already speculated that P papi/losum (Herz.) Card., a taxon recorded from Bolivia, might only be a synonym of P chimborazense. After examining the type specimens we found no differences between the types of these two species, and thus con­ firmed Crum's speculation. Specimens examined: Argentina - Tucuman, Bartellanos, no. 1669, 1935. 1. 29 (1E); Pategani­ ca, P. G. Lomtz (H); Uruguay - La Par. Felippone, no. 829 (H); O. S. Lomtz (H); De lujug. Fr. Clares, no. 11335, 11432. (3/1 , 190 I) (H); De Buenos Aires., H. Roivaine, no. 395 (H); Prov. de Cor­ doba, Sierra Achala, F. Kurtz. No. 8334 (H); Peru - Sandia, Weberbaue, no. 829 (H); Ca lea, P. E. Hegewald, no. 8639 (H); Chile - Bojin de Arrayun, Breuily, no. 1919 (H).

NEW COMBINATIONS Ptychomitrium gardneri var. angustifolium (Nog.) Cao & Guo, comb. novo ==Ptychomitrium /ongisetum Reim. et Sak. var. angustifolium Nog., 1. Hattori Bot. Lab. 12: 24. 1954.

Figs. 38-50. Ptychomitrium hieronymi var. obtusifolium (Ther.) Guo & Cao. - 38- 39: Leaves; 40: Upper lamina cells; 41: Middle lamina cells; 42 : Capsule; 43 : Transverse sections of upper part of leaf; 44: annulus; 45: Calyptra; 46, 48: Leaf apex; 47 : basal lamina cells along costa; 49: Trans­ verse sections of basal part of leaf; 50: Spores (drawn from isosyntype in H). Line scale: A=0.5mm (figs. 38- 39,42,45); B=200.um (figs. 46); C=50.um (figs. 40-44, 47- 50). Figs. 51-60. Ptychomitrium hieronymi Besch. - 51 - 52: Transverse sections of middle part of leaf; 53, 55-56: Leaves; 54: Spores; 57: Capsule; 58: Basal lamina cells along the costa; 59: Basal lamina cells near margins, 60: Upper lamina cells (drawn from holotype in BM). Line scale: D=0.5 mm (figs. 53, 55- 57); E=50.um (figs. 51 - 52, 54, 58-60). 47 d combinations In· ptychomitrium 1. CAD ET AL. New synonyms an

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Type: Japan, Honshu. Gumma Pref.: Mt. Kanaiyama, ca. I lOO m, on limestone (Nagano 1817) (holotype: NICH!). Noguchi (1954) published the variety Ptychomitrium longisetum Reim. et Sak. var. angustifolium Nog. based on the collection of Nagano. This variety is characterized by its leaves, from an oblong base sharply narrowing to a linear upper part, and being crisped when dry. Because P longisetum was shown to be a synonym of P polyphylloides (c. Muell.) Par (Iwatsuki & Noguchi, 1973), and the latter was synonymous with P gardneri Lesq. (Cao & Vitt 1994), the new combination P gardneri var. angustifolium (Nog.) Cao & Guo is made here. Ptychomitrium longisetum var. angustifolium is rather similar to P lin­ earifolium Reim., but the latter has shorter setae «7 mm), while the former has longer setae (> 10 mm).

Ptychomitrium sellowianum var. angustifolium (Broth.) Cao & Guo, comb. novo =.P weberbaueri var. angustifolium Broth., Bot. Jahrb. 56. (Beibl. 123): 10. 1920. Type: Prov. Sandia, Sandia Felsen; 2400 m (829) (holotype: H!). Brotherus (1920) described this variety as P weberbaueri var. angustifolium because of its narrower leaf base with plane margins. Guo and Cao (2000a) treated P weberbaueri Broth. as a synonym of P sellowianum (Mull. Hal.) Jaeger. The new combination P sell­ owianum var. angustifolium is proposed here.

Ptychomitrium hieronymi var. obtusifolium (Ther.) Cao & Guo, comb. novo (Figs. 38- 50) =. Brachysteleum chimborazense var. obtusifolium Ther., Rev. Chil. Hist. Nat. 27: 11. 1923. =. Pty­ chomitrium chimborazense var. obtusifolium (Ther.) Broth. , Nat. PfI . ed. 2. 11 : 9. 1925. Type: Chile, La Ermita, Las Condes, alt. 1200 m (Bertho & Jaffuel, X. 1919); Caj6n del Arragan, alt. 1400m (A. Breuil, XI. 1919 (isosyntype: H!). Ptychomitrium chimborazense var. obtusifolium (Ther.) Broth. was reported by Theriot (1923) as Brachysteleum chimborazense var. obtusifolium. This variety is charac­ terized by: I) long linear-lanceolate leaves having an obtuse or slightly obtuse apex; 2) a sheathing leaf base with transparent lamina cells; 3) distinctly papillose lamina cells on both surfaces; 4) ovate capsules with a wide orifice. We examined the type specimens of P chimborazense var. obtusifolium, P chimborazense and P hieronymi (Figs. 51-60) and found P chimborazense var. obtusifolium to be closer to P hieronymi than to P chimbo­ razense, especially in its leaf shape, leaf length and leaf set when dry. Therefore, P chimbo­ razense var. obtusifolium should be treated as a variety of P hieronymi. In the taxonomic treatment of the genus, Brotherus (1925) placed both P hieronymi and P chimborazense in Sect. Macrostoma based on their papillose lamina cells. The leaves of P chimborazense are 2- 3.5 mm long, short lanceolate, and irregularly crisped when dry, while those of Pty­ chomitrium hieronymi and P hieronymi var. obtusifolium are much longer (4- 6.5 mm in length), linear-lanceolate, and distinctly and regularly crisped when dry. Ptychomitrium hi-

Figs. 61 - 71. Ptychomitrium ferrugineum (Herz.) Broth. - 61 - 65 : Leaves; 66- 69: Transverse sections of basal part of leaf; 70: Transverse section of upper part of leaf; 71 : Basal lamina cells (drawn from holotype in JE). Line scale: A= 0.5 mm (figs. 61 - 65); B= 80.um (figs. 69); C=50.um (figs. 66- 68, 70- 71). 49 T. CAO ET AL.: New synonyms and combinations in ptychomitrium

ABC

71

69

63 50 1. Hattori Bot. Lab. No. 100 200 6 eronymi var. obtusifolium differs from P hieronymi by its slightly obtuse leaf apices. Their main differences are indicated by the following key. I. Leaves 2- 3.5 mm long, short lanceolate ...... Ptychomitrium chimborazens I. Leaves 4-6.5 mm long, linear-lanceolate 2. Apex acuminate ...... P hieronymi 2. Apex obtuse ...... P hieronymi var. obtusifolium

REVISED D ESCRIPTION OF P FERRUGINEUM Ptychomitrium ferrugineum (Herz.) Broth., Nat. Pfl. ed. 2. 11: 8. 1925. (Figs. 61-71) = Glyphomitrium ferrugineum Herz., Beih. Bot. Centralbl. 27: 350. 1910. Type : Bolivia, Prov. Cochabamba: An Felsen in der Blockhalde iiber dem Tunarisee, ca. 4400 m, I. 1908 (holotype: JE!). Herzog (1910) reported P ferrugineum from Bolivia as Glyphomitrium ferrugineum, with only a brief description. Since then, no further information has emerged concerning this species. We studied the holotype of P ferrugineum from lE and found that this species is rather distinct. Its lamina cells have numerous small papillae on both surfaces. The main diagnostic characteristics of this species are described as follows (Figs. 61 - 71): Plants in dense tufts, saxicolous, yellowish-brown. Stems erect, 20- 25 mm tall, central strand pre­ sent. Leaves crisped and somewhat contorted above when dry, spreading or connivent when moist; 2.5- 3.2 mm long; keeled above, somewhat sheathing at base, apex acuminate; mar­ gins plane, entire; costa percurrent, in section flattened, with 5- 9 guide cells, abaxial stereid band 3 or 4 cells thick, adaxial stereid band 2-3 cells thick, adaxial surface cells differ from the abaxial surface cells by having numerous small papillae. Lamina bistratose above base, abaxial and adaxial cells equal in size, quadrate, 7- 9 )1m, with numerous small papillae on exposed surfaces; basal cells lax, oblong, hyaline. The holotype had no cap­ sules. Among the four sections of Ptychamitrium defined by Brotherus (1925), sect. Macras­ tama is characterized by papillose laminal cells. However, some variation exists in this character among the species in the section. All the species except P exaratifolium Robin­ son have papillose lamina cells on both abaxial and adaxial surfaces, while in P exaratifoli­ um the lamina cells are papillose only on the adaxial surface (Magill & Rooy 1998). Among species with papillose laminal cells on both surfaces, only Ptychamitirum ferrug­ ineum has numerous small papillae on its laminal cells. We suggest that this species may belong in the Pottiaceae, but there are no sporophytes in the type material.

A CKNOWLEDGEMENTS Guo Shuiliang gratefully acknowledges support by the State Scholarship Fund of China for his visit to the University of Helsinki in 2003- 2004, and He Xiao-Ian, Juhani Heino, Sanna Huttunen, and the Natural History Museum of the University of Helsinki, for their hospitality during his visit. Thanks are also due to the curators of BM, FH, G, H, JE, NICH, and PC for loan of specimens. T. CAO ET AL.: New synonyms and combinations in Ptychomitrium 51

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