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Taxonomic treatment of the Polytrichaceae from Brazil

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Author(s) :Denilson Fernandes Peralta and Olga Yano

Source: The Bryologist, 113(3):646-672. 2010.

Published By: The American Bryological and Lichenological Society, Inc.

DOI:

http://dx.doi.org/10.1639/0007-2745-113.3.646

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Taxonomic treatment of the Polytrichaceae from Brazil

Denilson Fernandes Peralta

1

and Olga Yano

Instituto de Botaˆnica, Caixa Postal 3005, 01031-970 Sa˜o Paulo, SP, Brazil

ABSTRACT. Thirteen species of Polytrichaceae belonging to seven genera are hereby reported from

Brazil: Atrichum androgynum, Itatiella ulei, Notoligotrichum minimum, Oligotrichum

canaliculatum, O. denudatum, O. riedelianum, Pogonatum campylocarpum, P. pensilvanicum, P. perichaetiale subsp. oligodus, Polytrichadelphus pseudopolytrichum, Polytrichum angustifolium, P. commune and P. juniperinum. Polytrichadelphus organensis and Polytrichum reflexifolium are placed in synonymy with Polytrichum juniperinum, Polytrichum brasiliense with P. angustifolium and P. subremotifolium with P. commune. Keys, descriptions and illustrations are presented for all species. KEYWORDS. Atrichum, Itatiella, Notoligotrichum, Oligotrichum, Pogonatum, Polytrichadelphus,

Polytrichum, Bryophyta, taxonomy.

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The Polytrichaceae are characterized by formation of an epiphragm from the columella, the frequent presence of longitudinal lamellae on the upper leaf surface and, when present, the multicelular peristome appearing single and commonly with 32 ou 64 teeth (Gradstein et al. 2001). It is the only family in Polytrichales and Polytrichopsida sensu Goffinet et al. (2009). The family comprises between 218 (Crosby et al. 1999) and 300 species (Churchill & Linares C. 1995) accommodated in 23 genera (Goffinet et al. 2009), of which seven are monospecific (Crosby et al. 1999). The family including Dawsonia, is monophyletic and according Hyvo¨nen et al. (1998, 2004) forms the sister group to other peristomate mosses.

Gradstein et al. (2001) reported ten genera and 40 species from the Neotropics and of these, 32 species are thought to occur in Brazil (Yano 1996). The first citation of Polytrichaceae from Brazil was by Raddi (1820). He described Catharinea pseudopolytrichum

(5 Polytrichadelphus pseudopolytrichum (Raddi) G.L. Smith) from Serra do Frade (today Serra dos O´ rga˜os), in the State of Rio de Janeiro. The main Brazilian work was made by Farias (1982, 1984) studying seven species to State of Rio Grande do Sul. Although the Polytrichaceae are included in many floristic reports, no critical taxonomic revisions has been completed for Brazil or another Neotropical region.

We hereby present the result of a taxonomic revision based on extensive collections of Brazilian specimens, including many types. Eighty-one names have been applied to specimens of Polytrichaceae from Brazil. Thirteen species are accepted here; six are valid names, but are listed as doubtful names; eight others are valid names, but the collections from Brazil were misidentified and the species should be excluded from the flora; 18 are nomen nudum; 19 are used as synonym (four of these are listed for the first time), and 17 are herbarium names.

MATERIALS ANDMETHODS

Materials were obtained both from field collection and loans from herbaria (ALCB,B,BM,C, 1Corresponding author e-mail:

denilsonfp@yahoo.com.br DOI: 10.1639/0007-2745-113.3.646

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CESJ,E,EGR,FH,H,ICN,MANCH,MBM,MG,MO, NY,OXF, PACA,PC,R,RB,SJRP, S, SP,SPF, SPSF,UPCB,W) with

special attention to type specimens. Specimens were examined with stereo and lightmicroscopes. In total, 2594 specimens of Polytrichaceae from Brazil were analysed, including 20 types, six and 13 original collections to which a nomen nudum or a herbarium name, respectively, was applied. We used the DELTA (DEscription Language for TAxonomy) software to make keys and descriptions.

TAXONOMICTREATMENT

Polytrichaceae Schwa¨gr. in Willd., Sp. Pl. ed. 5(2): 1. 1830. Type: Polytrichum Hedw.

KEYS TO THESPECIES OFPOLYTRICHACEAE IN BRAZIL

1. Sporophyte present ... 2

1. Sporophyte absent ... 14

2. Calyptrae hairy ... 3

2. Calyptrae glabrous or with few trichomes ... 8

3. Capsule with stomata and apophysis; peristome 64 teeth ... 4

3. Capsule without stomata and apophysis; peristome 32 teeth ... 6

4. Margins involute ... Polytrichum juniperinum 4. Margins plane ... 5

5. Distal cell of lamellae pyriform ... Polytrichum angustifolium 5. Distal cell of lamellae U-shaped ... Polytrichum commune 6. Protonema persistent ... Pogonatum pensilvanicum 6. Protonema not persistent ... 7

7. Distal cell of lamellae double, pyriform ... ... Pogonatum campylocarpum 7. Distal cell of lamellae single, rhombic ... ... Pogonatum perichaetiale subsp. oligodus 8. Peristome 64 teeth; sporophyte lateral arising in small branches; sheathing base red ... ... Polytrichadelphus pseudopolytrichum 8. Peristome 32 teeth or absent; sporophyte terminal; sheathing base not red ... 9

9. Margins bordered by very long cells (10:1) ... ... Atrichum androgynum 9. Margins not bordered ... 10

10. Capsule without stomata and peristome ... Itatiella ulei 10. Capsule with stomata and peristome; 32 teeth ... 11

11. Lamellae occupying costae and laminae; cells papillose .. ... Notoligotrichum minimum 11. Lamellae occupying only costae or absent; cells smooth ... 12

12. Lamellae absent ... Oligotrichum denudatum 12. Lamellae over costae only ... 13

13. Leaves lanceolate, apex acute or acuminate, plane ... ... Oligotrichum riedelianum 13. Leaves ligulate, apex obtuse or conchiform, inflexed ... ... Oligotrichum canaliculatum 14. Protonema persistent; leaf gametophyte restrict to perichaetium and perigonium ... ... Pogonatum pensilvanicum 14. Protonema not persistent; leaf gametophyte well developed ... 15

15. Lamellae occupying costae; leaves crisped when dry ... 16

15. Lamellae occupying costae and laminae; leaves imbricate or widespreading when dry ... 19

16. Lamellae absent ... Oligotrichum denudatum 16. Lamellae well developed ... 17

17. Margin bordered by very long cells (10:1) ... ... Atrichum androgynum 17. Margin not bordered ... 18

18. Leaves ligulate; apex obtuse or conchiform ... ... Oligotrichum canaliculatum 18. Leaves lanceolate; apex acute or acuminate ... ... Oligotrichum riedelianum 19. Cells papillose ... Notoligotrichum mininum 19. Cells smooth ... 20

20. Margins involute ... Polytrichum juniperinum 20. Margins plane ... 21

21. Sheathing base orange or red ... 22

21. Sheathing base whitish or green ... 23

22. Distal cell of lamellae rugose; leaf apex acute ... ... Polytrichadelphus pseudopolytrichum 22. Distal cell of lamellae smooth; leaf apex acuminate ... ... Polytrichum angustifolium 23. Distal cell of lamellae double and pyriform ... ... Pogonatum campylocarpum 23. Distal cell of lamellae single, rhombic or U-shaped .... 24

24. Distal cell of lamellae U-shaped ... ... Polytrichum commune 24. Distal cell of lamellae rhombic ... 25 25. Distal cell of lamellae thick-walled; leaf apex acuminate,

erect ... Pogonatum perichaetiale subsp. oligodus 25. Distal cell of lamellae thin-walled; leaf apex acute,

incurved ... Itatiella ulei

Atrichum P. Beauv., Mag. Encycl. 5: 329. 1809, nom. cons. TYPE: A. undulatum (Hedw.) P. Beauv.

Atrichum androgynum (Mu¨ll. Hal.) A. Jaeger, Ber. Tha¨tigk. St. Gallischen Naturwiss. Ges. 1873–74: 241 (Gen. et Sp. Musc. 1: 709). 1875 ; Catharinea androgyna Mu¨ll. Hal., Syn. Musc. Frond. 1: 193. 1849. TYPE: AFRICA.PIOMONT: bon

spec. prope swollendam in nemoribus ubi. m.octobr.c. Thecis operculatis, Ecklon s.n. ex. (syntypesMEL? hb. Kunzeanum,BM?). Fig. 1

Catharinea ulei Mu¨ll. Hal., Hedwigia 38 (Beibl. 1): 57. 1899. nom. nud.

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Figure 1. Atrichum androgynum. a. Gametophyte, b–c. Leaves, d. Perigonial leaves, e. Apical cell of the leaf, f. Marginal cells of leaf, g. Cells of the sheathing base, h. Transverse section of the apex of the leaf, i. Transverse section of the median leaf, j. Transverse section of the sheathing base, k. Lateral view of lamella, l. Epidermal cells of the capsule, m. Transverse section of the epidermal cells of the capsule, n. Calyptra, o. Peristome teeth (Yano & Marcelli 19434-SP).

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3(3): 32. 1895. TYPE: BRAZIL,MINAS GERAIS: Caldas,

VII-1823, Mose´n 332 (syntypeH-BR!).

Atrichum riograndense (Broth.) Paris, Index Bryol. 281: 70. 1904 ; Catharinea rio-grandensis Broth., Bih. Kongl. Svenksa Vetensk.-Akad. Handl. 26 Afd. 3(7): 32. 1900. TYPE: BRAZIL,RIO GRANDE DO SUL: Silveira Martins ad, Terram

marginis viae declivum umbrosam, 19-III-1893, Lindman 210 (syntypesH-BR!,R!).

Description. Plants medium-sized, dark-green to brown, 50–100 mm long. Stems erect, forming tufs. Rhizome prostrate. Rhizoids pale-brown. Leaves lanceolate, 4–7 mm long, widespreading, quadratic-hexagonal cells, 10–15 3 10–15mm, thick-walled, smooth, sheathing base inconspicuous, plane margins, double dentate, bordered, costa narrow 1/ 5–1/3, percurrent, lamellae occupying only the costae, 4–6 lamellae, 2–4 cells high, distal cells simple, rhombic, smooth, lamella cells in lateral view quadratic-hexagonal. Monoicous, perichaetium and perigonium terminal, leaves not differentiated. Sporophyte 1(–3), seta 30 mm long. Capsule asymmetric, urn cylindric, stomata absent, operculum conic, long-rostrate, 32 teeth. Calyptra cucullate, glabrous.

Distribution. Africa, Mexico, Guatemala, Honduras, Costa Rica, Colombia and Brazil (States of Minas Gerais, Parana´, Rio de Janeiro, Rio Grande do Sul, Santa Catarina and Sa˜o Paulo).

Habitat. On soil, at margins of submontane and montane forests, in the Bioma Mata Atlaˆntica and Pampa, between 600–2,000 m alt.

Discussion. This species is diagnosed by the bordered leaves that are crisped when dry. Mu¨ller (1848–1849) described the species as either dioicous or monoicous, but only bisexual gametophytes, with many antheridia around the seta, were found among the material from Brazil.

Nyholm (1971) uses the lengh of lamina cells to separe the varieties of A. androgynum. But among the Brazilian material, we found a large variation on this character even within a single population. Hence we treated all specimens as A. androgynum.

The collections from Brazil identified as Atrichum oerstedianum (Mu¨ll. Hal.) Mitt. don’t have the antheridia surrounded by the differentiated perigonial leaves, as is clearly the case in the type (K!).

The type specimens of Atrichum mosenii (Broth.) Paris (H!) and A. riograndense (Broth.) Paris

(H!R!) cannot be distinguished from A. androgynum

and the names should be considered synonyms of A. androgynum as suggested by Nyholm (1971).

Selected specimen examined. BRAZIL.MINAS GERAIS: Caldas, Mose´n 331 (H); Caparao´ Novo, Parque

Nacional do Caparao´, Vital & Buck 11630 (SP). PARANA´: Dorizon, Hotel A´gua Dorizon, Wasum et al.

3745 (MO,SP); Guarapuava, Fazenda Campo Real,

Hatschbach 21025 (MBM,MO,PACA); Palmas, O. Yano

& Pirani 6551 (SP).RIO DE JANEIRO: Itatiaia, Ule 38 (S);

Nova Friburgo, Kurtz & Dihlmann 27 (RB); Rio de

Janeiro, Costa & Molinaro 243 (RB); Tereso´polis,

Parque Nacional da Serra dos O´ rga˜os, O. Yano & Costa 13549 (SP).RIO GRANDE DO SUL: Bom Jesus,

Sehnem 220 (PACA,SP); Cambara´ do Sul, Fortaleza dos

Aparados, Vianna 615 (ICN); Canela, Farias 181 (ICN);

Caxias do Sul, Criu´va, 17-IX-1988, Rossato et al. s.n. (MO); Dois Irma˜os, Morro Reuter, Sehnem 12513

(PACA, SP); Esmeralda, Vianna 735 (ICN); Flores da

Cunha, 19-II-1992, Brinker et al. s.n. (MO); Jaquirana, Wasum et al. 775 (MBM,ALCB);

Montenegro, Sehnem 4038 (PACA,R,RB,SP); idem, Sa˜o

Francisco de Paula, Vila Oliva, Sehnem 2736 (PACA, SP); Sa˜o Leopoldo, Feitoria, Sehnem 10 (PACA, SP);

Silveira Martins, Lindman 191 (H).SANTA CATARINA:

Serra Geral, V-1890, Ule s.n. (R); idem, Ule 38 (SP);

Sa˜o Jose´, Serra da Boa Vista, Reitz & Klein 9926, 9927 (SP).SA˜ O PAULO: Bananal, Nova Suic¸a, O. Yano 1264 (MO,SP); Campos da Cunha, Peralta et al. 3740 (SP);

Campos do Jorda˜o, Parque Estadual de Campos do Jorda˜o, O. Yano & Marcelli 19434 (SP); Cunha, Vital

7768 (NY,SP); Moji das Cruzes, Parque Municipal da

Serra de Itapety, Peralta et al. 2754 (SP);

Pindamonhangaba, Pico Itapeva, Vital 1012 (SP);

Santo Andre´, Reserva Biolo´gica do Alto da Serra de Paranapiacaba, O. Yano & Kirizawa 9385 (SP); Sa˜o

Jose´ do Barreiro, Vattimo 3–112 (R); Sa˜o Paulo,

Parque Estadual das Fontes do Ipiranga, O. Yano et al. 15650 (SP).

Itatiella G.L. Smith, Mem. New York Bot. Gard. 21(3): 52. 1971. TYPE: I. ulei (Mu¨ll. Hal.) G.L.

Smith.

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ex Mu¨ll. Hal., Bull. Herb. Boissier 6: 26. 1898. TYPE: BRAZIL.SERRA DE ITATIAIA, 2100 m, in campo,

III-1894, Ule 1905 (Bryotheca Brasiliensis 151) (syntypesH-BR!,R!,FH(photo!),PC!,R!,S!,W!).

Polytrichum ulei Broth., Rodrigue´sia 54: 266. 1980.

nom. nud. Fig. 2

Description. Plants medium-sized, dark-green to brown, 10–25 mm long. Stems erect, forming tufs. Rhizome prostrate. Rhizoids pale-brown. Leaves oblong-lanceolate, 2.5–4.0 mm long, widespreading, hexagonal-quadratic cells, 20–25 3 10–15mm, thick-walled, smooth, sheathing base conspicuous, involute margins, dentate, not bordered, costa narrow 1/5–1/ 3, percurrent, lamellae occupying costae and laminae, 30–40 lamellae, 5–6 cells high, distal cells simple, rhombic, smooth, lamella cells in lateral view hexagonal to quadratic. Dioicous, perichaetium terminal, leaves long, perigonium terminal, leaves short and broad. Sporophyte 1(–3), seta 5–10 mm long. Capsule symmetric, urn globose, stomata absent, operculum conic, long-rostrate, teeth absent. Calyptra cucullate, glabrous.

Distribution. Endemic to Southeastern and Southern Brazil (States of Espı´rito Santo, Minas Gerais, Parana´, Rio de Janeiro and Sa˜o Paulo).

Habitat. On soil at the margins of montane forest and high Campos in the Bioma Mata Atlaˆntica, between 900–2,890 m alt.

Discussion. This species is caracterized by its small size and the inflexed leaf apex. Some specimens of Pogonatum campylocarpum growing directly exposed to sun at high elevations present a similar aspect, but can be distinguished based on the distal lamella cells, single and rhombic in Itatiella ulei and double and pyriform in P. campylocarpum.

Species of Psilopilum resemble Itatiella but differ by the presence of a peristome and stomata. The original publication lists a specimen by Ule numbered ‘‘1905’’. Only one specimen held inR

bears this number, but it is written in pencil, and unlikely by the original collector or author. All other specimens from Bryotheca Brasiliensis are labeled ‘‘151’’ and are therefore consider type material. Vattimo-Gil & Vattimo (1980) reported Polytrichum ulei Broth. from Brazil, this name is a nomen nudum and the specimen (Souza s.n.) is Itatiella ulei.

Selected specimen examined. BRAZIL ESPı´RITO SANTO: Iu´na, Parque Nacional do Caparao´, Vital &

Buck 11764 (B,E,MO,MG,NY,RB,SP).MINAS GERAIS: Alto

Caparao´, Buck et al. 27004 (SP,MO,NY); Itamonte,

Schiffner 755 (NY,S); Lima Duarte, Parque Estadual

do Itibipoca, Krieger 16189 (PACA).PARANA´: Duse´n

2558 (B,E,S).RIO DE JANEIRO: Itatiaia, 1894, Ule s.n.

(W); Resende, Schiffner s.n (B,W).; Tereso´polis,

Parque Nacional da Serra dos O´ rga˜os, Costa et al. 4168 (RB); Visconde de Maua´, Scha¨fer-Verwimp &

Verwimp 8325 (MO,SP).SA˜ O PAULO: Campos do

Jorda˜o, Parque Estadual de Campos do Jorda˜o, O. Yano & M.P. Marcelli 15742 (SP); Piquete, Peralta

3621, 3625 (SP).

Notoligotrichum G.L. Smith, Mem. New York Bot. Gard. 21(3): 50. 1971. TYPE: N. australe (J.D.

Hook.) G.L. Smith

Notoligotrichum minimum (Cardot) G.L. Smith, Mem. New York Bot. Gard. 21(3): 51. 1971 ; Polytrichadelphus minimus Cardot, Bull. Herb. Boissier, ser. 2, 5: 1008. 1905. TYPE: ARGENTINA. TERRE-DE-FEU: Bahia Tekenika, M.C. Skottsberg

s.n. 135 (isotype S! hb. Cardot). Fig. 3

Description. Plants medium-sized, red or green to brown, 10–20 mm long. Stems erect, forming tufs. Rhizome prostrate. Rhizoids pale-brown. Leaves lanceolate, 8–12 mm long, widespreading, quadratic cells, 8–12 3 8–10mm, thick-walled, papillae numerous, sheathing base conspicuous, plane margins, dentate, not bordered, costa narrow 1/5–1/ 3, percurrent, lamellae occupying costae and laminae, 40–46 lamellae, 1–2 cells high, distal cells simple, rhombic, papillose, lamella cells in lateral view quadratic. Dioicous, perichaetium terminal, leaves long, perigonium terminal, leaves short and broad. Sporophyte single, seta 10–20 mm long. Capsule asymmetric, urn cylindric, stomata present at urn base, operculum conic, long-rostrate, 32 teeth. Calyptra cucullate, glabrous.

Distribution. Argentina, Chile and Brazil (State of Rio de Janeiro).

Habitat. On soil along trails and on banks in montane forest at the Mata Atlaˆntica Bioma at ca. 1,100 m alt.

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Figure 3. Notoligotrichum minimum. a. Gametophyte, b–e. Leaves, f–g. Perichaetial leaves, h. Apical cells of the leaf, i. Marginal cells of the leaf, j. Cells of the sheathing base, k. Transverse section of the apex of the leaf, l. Transverse section of the median leaf, m. Transverse section of the sheathing base, n. Lateral view of lamella, o. Epidermal cells of the capsule, p. Transverse section of the epidermal cells of the capsule, q. Calyptra, r. Peristome teeth (Yano & Visnadi 9908-SP).

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Polytrichaceae by its lamina cells with papillate cuticle.

Specimen examined. BRAZIL.RIO DE JANEIRO:

Resende, Parque Nacional do Itatiaia, O. Yano & Visnadi 9908 (SP).

Oligotrichum D.C., Fl. Franc. (Lamarck), ed. 2: 491. 1805. nom. cons. TYPE: O. hercynicum (Hedw.)

Lamarck & D.C.

Oligotrichum canaliculatum (J.D. Hook. & Arnott) Mitt., J. Linn. Soc., Bot. 12: 606. 1869 ; Polytrichum canaliculatum J.D. Hook. & Arnott, Bot. Beechey Voy.: 54. 1832. TYPE: CHILE. CONCEPCIO´ N: Lay & Collie s.n. (?). Fig. 4

Description. Plants small-sized, dark-green or green to brown, 4–5 mm long. Stems erect, forming tufs. Rhizome prostrate. Rhizoids pale-brown. Leaves ligulate, 2.5–3.0 mm long, widespreading when wet, crisped when dry, quadratic cells, 15–20 3 15–20mm, thick-walled, smooth, sheathing base conspicuous, plane margins, dentate, not bordered, apex obtuse, costa narrow 1/5–1/3, percurrent, lamellae occupying only the costa, 8–10 lamellae, 7–9 cells high, distal cells simple, rhombic, smooth, lamella cells in lateral view quadratic. Dioicous, perichaetium terminal, leaves long, perigonium terminal, leaves short and broad. Sporophyte single, seta 15–20 mm long. Capsule symmetric, urn globose, stomata present at urn base, operculum conic, long-rostrate, 32 teeth. Calyptra cucullate, glabrous.

Distribution. Argentina, Chile and Brazil (States of Rio Grande do Sul and Santa Catarina*).

Habitat. On banks at the margins of submontane forests in the Bioma Mata Atlaˆntica and Pampa.

Discussion. This species is diagnosed by its obtuse and inflexed leaf apex. Mu¨llen & Frye (1947) distinguished it from O. riedelianum (Mont.) Mitt. and O. erosum (Hampe) Lindb. by its broad costa (1/ 3 of the leaf width) and short seta (23 mm). The seta length in the other species is 30–45 mm long. The plants are smaller than others species of the genus and is related to the temperate areas of Brazil.

Specimen examined. BRAZIL.RIO GRANDE DO SUL:

Cacequi, Sanga˜o do Macaco Branco, Bueno 3329, 3340 (ICN); Garibaldi, Scha¨fer-Verwimp & Verwimp

10703 (SP).SANTA CATARINA: Urubici, Serra do Corvo

Branco, Scha¨fer-Verwimp & Verwimp 13507 (SP).

Oligotrichum denudatum Merrill, Novon 1(3): 107. 1991. TYPE: BRAZIL.MINAS GERAIS: Sa˜o Roque de

Minas, mesic forest and riverside at base of waterfall, Parque Nacional da Serra da Canastra, Cachoeira Casca D’Anta, 20u189S, 46u319W, 1,000 m alt., 19-IX-1984, Vital & Buck 11941 (holotypeNY!; isotypeSP!). Fig. 5

Description. Plants medium-sized, dark-green to brown, 35 mm long. Stems erect, forming tufs. Rhizome prostrate. Rhizoids pale-brown. Leaves lanceolate-ligulate, 2.5–3.0 mm long, widespreading, long-quadratic cells, 15–20 3 15–20mm, thin-walled, smooth, sheathing base conspicuous, plane margins, dentate, not bordered, costa narrow 1/5–1/3, percurrent, lamellae absent. Dioicous, perichaetium terminal, leaves long, perigonium terminal, leaves short and broad. Sporophyte single, seta 30–40 mm long. Capsule symmetric, urn globose, stomata present at urn base, operculum conic, long-rostrate, 32 teeth. Calyptra cucullate, glabrous.

Distribution. Endemic to Southeastern Brazil (State of Minas Gerais).

Habitat. On soil along trails and on banks near waterfalls in montane forests at the Mata Atlaˆntica Bioma; at ca. 1,000 m alt.

Discussion. This species is similar to O. riedelianum but by the absence of lamellae, although we found some leaves with a small lamella of 1–2 cell high.

Specimen examined. BRAZIL.MINAS GERAIS: Sa˜o

Roque de Minas, Parque Nacional Serra da Canastra, Vital & Buck 11931b (SP,NY).

Oligotrichum riedelianum (Mont.) Mitt., J. Linn. Soc., Bot. 12: 606. 1869 ; Polytrichum riedelianum Mont., Ann. Sci. Nat., Bot., ser. 2, 16: 269. 1841. TYPE: BRAZIL. Montagne s.n.

(holotypeNY!). Fig. 6

Polytrichum leucodontium Mu¨ll. Hal., Bot. Zeitung (Berlin) 2: 726. 1844. Polytrichum nigricans J.D. Hook. & Wilson, London J. Bot. 3: 151. 1844. TYPE: BRAZIL. Gardner 10 (syntypesNY!,E!, MANCH!).

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Figure 4. Oligotrichum canaliculatum. a. Gametophyte, b–d. Leaves, e–f. Perichaetial leaves, g. Apical cells of the leaf, h. Marginal cells of the leaf, i. Cells of the sheathing base, j. Transverse section of the apex of the leaf, k. Transverse section of the median leaf, l. Transverse section of the sheathing base, m. Lateral view of the lamella, n. Epidermal cells of the capsule, with stomata, o. Transverse section of the epidermal cells of the capsule, p. Calyptra, q. Peristome teeth (Scha¨fer-Verwimp & Verwimp 13507-SP).

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Figure 6. Oligotrichum riedelianum. a. Female gametophyte, b. Male gametophyte, c–e. Leaves, f–h. Perichaetial leaves, i–j. Perigonial leaves, k. Apical cells of the leaf, l. Margin cells of the leaf, m. Cells of the sheathing base, n. Transverse section of the apex of the leaf, o. Transverse section of the median leaf, p. Transverse section of the sheathing base, q. Lateral view of the lamella, r. Epidermal cells of the capsule, with stomata, s. Transverse section of the epidermal cells of the capsule, t. Calyptra, u. Peristome teeth (Yano 1244-SP).

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quadratic cells, 10 3 15–20mm, thin-walled, smooth, sheathing base conspicuous, plane margins, dentate, not bordered, costa narrow 1/5–1/3, percurrent, lamellae occupying only the costa, 12–15 lamellae, 5–7 cells high, distal cells simple, rhombic, smooth, lamella cells in lateral view quadratic. Dioicous, perichaetium terminal, leaves long, perigonium terminal, leaves short and broad. Sporophyte single, seta 40–60 mm long. Capsule symmetric, urn globose, stomata present at urn base, operculum conic, long-rostrate, 32 teeth. Calyptra cucullate, glabrous.

Distribution. Endemic to Brazil (States of Minas Gerais, Parana´, Rio de Janeiro, Rio Grande do Sul, Santa Catarina and Sa˜o Paulo).

Habitat. On soil along trails and on banks in montane forests at the Mata Atlaˆntica Bioma; between ca. 1,800–2,200 m alt.

Discussion. Similar to Oligotrichum denudatum, see differentiation under this species. Polytrichum leucodontium Mu¨ll. Hal. and P. nigricans J.D. Hook. & Wilson are homotypic synonyms and are here confirmed to represent a species identical to O. riedelianum as proposed by Wijk et al. (1967).

Selected specimen examined. BRAZIL.MINAS GERAIS: Caldas, Regnell 35b, 36b (S); Camanducaia,

Monte Verde, Scha¨fer-Verwimp & Verwimp 7521 (SP);

Caparao´, Parque Nacional do Caparao´, Vital & Buck 11605 (SP, NY); Delfim Moreira, Scha¨fer-Verwimp &

Verwimp 10042 (SP); Itamonte, O. Yano et al. 22628

(SP); Marmelo´polis, Peralta 3568 (SP); Passa Quatro,

III-1921, Zika´n s.n. (B,MANCH); Santa Ba´rbara, Vital

& Favalli 10830 (SP).PARANA´: 1902, P. Duse´n s.n

(MANCH,W,S); Castro, Weir 115 (NY); Morretes, Serra

da Graciosa, Schwacke 1726 (RB); Unia˜o da Vito´ria, O.

Yano & Pirani 6484 (SP).RIO DE JANEIRO: Engenheiro

Passos, O. Yano & Santos 7670 (SP); Itatiaia, I-1902,

Duse´n s.n. (S); Nova Friburgo, VII-1835, Lusihnatt

s.n. (S); Parati, Vital 15401 (SP); Resende, Duse´n 2555

(B,W); Tereso´polis, Serra dos O´ rga˜os, X-1896, Ule s.n

(W).RIO GRANDE DO SUL: Sa˜o Francisco de Paula, Serra

do Faxinal, Sehnem 5304 (PACA,ICN,SP).SANTA CATARINA: Bom Retiro, Sehnem 7039 (PACA).SA˜ O PAULO:

Bananal, Nova Suic¸a, O. Yano 1244 (MO,SP); Campos

do Jorda˜o, Parque Estadual de Campos do Jorda˜o, O. Yano & Watanabe 9223, 9227 (SP); Cunha, Vital 7760

(NY, SP); Moji das Cruzes, Serra do Mar, Fonseca &

Kirizawa 13, 15 (SP); Piquete, Peralta et al. 4076 (SP);

Sa˜o Jose´ do Barreiro, Serra da Bocaina, IX-1879, Glaziou & Schwacke s.n. (R).

Pogonatum P. Beauv., Mag. Encycl. 5: 329. 1804. TYPE: P. aloides (Hedw.) P. Beauv.

Pogonatum campylocarpum (Mu¨ll. Hal.) Mitt., J. Linn. Soc., Bot. 12: 618. 1869 ; Polytrichum campylocarpum Mu¨ll. Hal., Syn. Musc. Frond. 1: 209. 1849. TYPE: VENEZUELA.CARACAS: Galipan,

4000 (5000?) ft., I-1846, Funck & Schlim 379 (lectotypeBM; isotypesGOET,JE,L,NY),

designated by Menzel (1985[1986]). Fig. 7 Pogonatum camptocaulon (Mu¨ll. Hal.) Paris, Index

Bryol. 978. 1898 ; Polytrichum camptocaulon Mu¨ll. Hal., Acta Soc. Sci. Fenn. 19(5): 17. 1891. TYPE: BRAZIL.MINAS GERAIS: Carac¸a, in terra

arenosa, IV-1885, Wainio s.n. (lectotypeH-BR),

designated by Menzel (1985[1986]).

Description. Plants medium-sized, dark-green to blackish, 20–30 mm long. Stems erect, forming tufs. Rhizome prostrate. Rhizoids pale-brown. Leaves lanceolate, 4–5 mm long, widespreading, quadratic cells, 10 3 10–15mm, thick-walled, smooth, sheathing base conspicuous, plane margins, dentate, not bordered, costa broad 3/4–4/5, percurrent, lamellae occupying costae and laminae, 64–70 lamellae, 5–6 cells high, distal cells double, pyriform, smooth, lamella cells in lateral view quadratic. Dioicous, perichaetium terminal, leaves long, perigonium terminal, leaves short and broad. Sporophyte single, seta 30–40 mm long. Capsule asymmetric, urn cylindric, stomata absent, operculum conic, long-rostrate, 32 teeth. Calyptra cucullate, hairy.

Distribution. America: Costa Rica, El Salvador, Panama, Dominican Republic, Bolivia, Colombia, Ecuador, Peru, Venezuela and Brazil (States of Minas Gerais, Rio de Janeiro, Rio Grande do Sul, Santa Catarina and Sa˜o Paulo).

Habitat. On soil, along trails and on banks at the edge of montane forests at the Mata Atlaˆntica Bioma; between ca. 600–2,000 m alt.

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Figure 7. Pogonatum campylocarpum. a. Female gametophyte, b. Male gametophyte, c–e. Leaves, f–g. Perichaetial leaves, h–i. Perigonial leaves, j. Apical cells of the leaf, k. Margin cells of the leaf, l. Cells of the sheathing base, m. Transverse section of the apex of the leaf, n. Transverse section of the median leaf, o. Transverse section of the sheathing base, p. Lateral view of the lamella, q. Epidermal cells of the capsule, r. Transverse section of the epidermal cells of the capsule, s. Calyptra, t. Peristome teeth (Peralta 3614-SP).

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which differs by the entire leaf margin, conspicuous constriction betwen sheathing base and laminae and distal cells of lamella (but this species has two cells too! Clarify) double and thick-walled. Vattimo-Gil & Vattimo (1980), reported Pogonatum tortile (Sw.) Brid. from Brazil based on the specimen Bandeira s.n. (RB!). However, this specimen belongs to P.

campylocarpum, and hence, P. tortile does not occur in Brazil.

Selected specimen examined. BRAZIL.MINAS GERAIS: Caldas, Regnell s.n. (E); Camanducaia,

21-II-1979, Martins s.n. (SP); Caparao´, Parque Nacional do

Caparao´, Vital & Buck 11607 (SP); Catas Altas, Peralta

et al. 6358 (SP); Itamonte, O. Yano & Visnadi 9904

(SP); Lima Duarte, Carac¸a, Ule 1318 (R),

Marmelo´polis, Peralta 3555 (SP); Ouro Preto, Ule

1319 (R).RIO DE JANEIRO: Itatiaia, I-1896, Ule s.n. (S);

Nova Friburgo, 1-V-1923, Bandeira s.n. (RB); Parati,

Vital 7734, 7737, 7746 (SP,NY); Tereso´polis, Mautone

& Guimara˜es 118 (RB).RIO GRANDE DO SUL: Cambara´ do

Sul, Peralta & Gonc¸alves 3314, 3319, 3323 (SP); Sa˜o

Francisco de Paula, Farias 185 (ICN).SANTA CATARINA:

Bom Retiro, Scha¨fer-Verwimp & Verwimp 10490 (SP);

Serra Geral, Vital & Buck 12425 (SP).SA˜ O PAULO:

Bananal, O. Yano 1226 (SP); Campos do Jorda˜o,

Parque Estadual de Campos do Jorda˜o, O. Yano & Watanabe 9226 (SP); Cubata˜o, Vital 13986 (SP);

Cunha, Vital 7761 (SP); Pindamonhangaba, Pico do

Itapeva, Vital 642 (SP); Piquete, D.F. Peralta 3614

(SP); Sa˜o Bernardo do Campo, O. Yano et al. 30337,

30351 (SP); Sa˜o Jose´ do Barreiro, Loefgren & Edwal

537 (SP); Sa˜o Luis do Paraitinga, O. Yano et al. 7791

(SP); Sa˜o Paulo, Hoehne 128 (SP,MANCH).

Pogonatum pensilvanicum (Hedw.) P. Beauv., Me´m. Soc. Linn. Paris 1: 461. 1823 ; Polytrichum pensilvanicum Hedw., Sp. Musc. Frond. 96. 1801 ; Polytrichum brevicaule Brid., J. Bot. (Schrader) 1800(1): 284. 1801. nom. superfl. TYPE: USA.PENNSYLVANIA: Mu¨hlenberg

(syntypesBM,G). Fig. 8

Pogonatum gardneri (Mu¨ll. Hal.) Mitt., J. Proc. Linn. Soc., Bot. 8: 156. 1864 ; Polytrichum gardneri Mu¨ll. Hal., Syn. Musc. Frond. 2: 560. 1851. TYPE:

BRAZIL. Tijuca, clay bank, XI-1836, Gardner 15

(syntypeE!).

Pogonatum glaziovii (Hampe) A. Jaeger, Ber. Thatigk. St. Gallischen Naturwiss. Ges. 1: 716. 1875 ;

Polytrichum glaziovii Hampe, Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn ser. 3 4: 54. 1872. TYPE: BRAZIL. Glaziou 5156 (lectotypeBM,

isotypeFH), designated by Hyvo¨nen (1989).

Polytrichum campophilum Mu¨ll. Hal., Hedwigia 38(Beibl. 1): 58. 1899. nom. nud.

Pogonatum subabbreviatum Broth., Denkschr. Kaiserl. Akad. Wiss., Wien. Math.-Naturwiss. Kl. 83: 350. 1924. TYPE: BRAZIL.MINAS GERAIS: Serra do

Itatiaia, Macieiras, ad margines viarum, 16-I-1925, Vaughan Bandeira 91 (LectotypeH-BR),

designated by Hyvo¨nen (1989).

Description. Plants small-sized, protonema persistent, dark-green, 2–3 mm long. Stems erect, forming tufs. Rhizomes absent. Rhizoids pale-brown. Leaves lanceolate, 2–3 mm long, widespreading, quadratic cells, 15 3 15mm, thick-walled, smooth, sheathing base conspicuous, plane margins, dentate, not bordered, costa narrow 1/5–1/3, percurrent, lamellae occupying costae and laminae, 10–20 lamellae, 3–4 cells high, distal cells single or double, rhombic, smooth, lamella cells in lateral view quadratic. Dioicous, perichaetium terminal, leaves long, perigonium terminal, leaves short and broad. Sporophyte single, seta 11–15 mm long. Capsule symmetric, urn cylindric, stomata absent, operculum conic, long-rostrate, 32 teeth. Calyptra cucullate, hairy.

Distribution. Widespread in the New World (Canada, Mexico, Cuba, Bolivia, Peru, Paraguay and Brazil); in Brazil known from Bahia, Distrito Federal, Espı´rito Santo, Goia´s, Minas Gerais, Parana´, Rio de Janeiro, Rio Grande do Sul, Santa Catarina and Sa˜o Paulo.

Habitat. On soil along trails and on banks at margins of montane forests at the Mata Atlaˆntica, Cerrado and Pampa Biomes; between ca. 400– 2,200 m alt.

Discussion. The persistent protonema

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Figure 8. Pogonatum pensilvanicum. a. Female gametophyte, b. Male gametophyte, c–e. Leaves, f–h. Perichaetial leaves, i. apical cells of the leaf, j. Margin cells of the leaf, k. Cells of the sheathing base, l. Transverse section of the apex of the leaf, m. Transverse section of the median leaf, n. Transverse section of the sheathing base, o. Lateral view of the lamella, p. Epidermal cells of the capsule, q. Transverse section of the epidermal cells of the capsule, r. Calyptra, s. Peristome teeth (Yano 15233-SP).

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the costa, and well developed lamellae with the verrucose distal cells.

Selected specimen examined. BRAZIL.DISTRITO FEDERAL: Mumunhas, Vital 13498 (SP).ESPı´RITO SANTO:

Domingos Martins, Scha¨fer-Verwimp & Verwimp 10189 (SP); Santa Teresa, O. Yano 3593 (SP).GOIA´S:

Alto Paraı´so de Goia´s, Chapada dos Veadeiros, Costa et al. 2280 (RB); Cristalina, Vital 6273 (SP).MINAS GERAIS: Belo Horizonte, Serra do Curral, Chaos s.n.

(RB); Caldas, Regnell s.n., 36, 969 (S); Catas Altas,

Peralta et al. 6826 (SP); Diamantina, Mexia 5831 (MO, S); Lima Duarte, Parque Estadual do Ibitipoca, O.

Yano et al. 20081 (SP); Santana do Riacho, O. Yano &

Wanderley 10664 (SP).PARANA´: Antonina, Hatschbach

21713 (MBM;PACA); Pato Branco, O. Yano et al. 5444

(SP); Tijucas do Sul, O. Yano & Shirata 27958 (SP).RIO DE JANEIRO: Angra dos Reis, Ilha Grande,

Oliveira-e-Silva 2429 (SP); Itatiaia, V-1902, Duse´n s.n. (e, s);

Nova Friburgo, 22-II-1903, P. Duse´n s.n. (S); Parati,

Vital 7745 (MO,NY,SP).RIO GRANDE DO SUL: Cambara´

do Sul, Guerra et al. 719 (alcb, mbm); Caxias do Sul, O. Yano & Bordin 29144 (SP); Silveira Martins,

4-III-1893, Lindman s.n. (R).SANTA CATARINA: Santa Cecı´lia,

Vital 5594 (SP); Porto Belo, O. Yano & Yano 10038

(SP); Tijucas, Sehnem 3251 (PACA).SA˜ O PAULO: Atibaia,

Peralta et al. 5598 (SP); Barra do Turvo, Vital 3221

(SP); Brotas, Vital 11249 (SP); Campos do Jorda˜o,

Parque Estadual de Campos do Jorda˜o, O. Yano & Watanabe 9221 (SP); Ibiu´na, O. Yano & Yano 10074

(SP); Piquete, Peralta 3630 (SP); Sa˜o Bernardo do

Campo, O. Yano et al. 15233 (SP); Santo Andre´,

Schiffner 793 (S); Serra Negra, O. Yano et al. 18516

(SP); Ubatuba, Vital 4781 (SP).

Pogonatum perichaetiale (Mont.) A. Jaeger subsp. oligodus (Kunze ex Mu¨ll. Hal.) Hyvo¨nen, Acta Bot. Fenn. 138: 15. 1989 ; Polytrichum oligodus Kunze ex Mu¨ll. Hal., Syn. Musc. Frond. 1: 206. 1848. TYPE: CHILE. Poeppig 227 (lectotypeBM,

isotypesG,H-BR,JE,HBG,PC), designated by

Menzel (1985[1986]). Fig. 9 Pogonatum itatiaiae (Mu¨ll. Hal.) Paris, Index Bryol.

Suppl.: 278. 1900 ; Polytrichum itatiaiae Mu¨ll. Hal., Bull. Herb. Boissier 6: 25. 1898. TYPE:

BRAZIL.MINAS GERAIS: Serra Itatiaia, 2,000 m alt.,

in terra, III-1894, Ule 1764 (Bryotheca Brasiliensis 345) (lectotypeH-BR, isotypes,HBG, R!), designated by Menzel (1985[1986]).

Description. Plants small-sized, dark-green to blackish, 10–15 mm long. Stems erect, forming tufs. Rhizomes absent. Rhizoids pale-brown. Leaves lanceolate, 5–6 mm long, widespreading, quadratic cells, 5–10 3 5–10mm, thick-walled, smooth, sheathing base conspicuous, plane margins, dentate, not bordered, costa broad 3/4–4/5, percurrent, lamellae occupying costae and laminae, 40–45 lamellae, 5–6 cells high, distal cells single, rhombic, smooth, lamella cells in lateral view quadratic. Dioicous, perichaetium terminal, leaves long, perigonium terminal, leaves short and broad. Sporophyte single, seta 20–25 mm long. Capsule symmetric, urn cylindric, stomata absent, operculum conic, long-rostrate, 32 teeth. Calyptra cucullate, hairy.

Distribution. Africa and Tropical America. Brazil (States of Minas Gerais and Rio de Janeiro).

Habitat. On soil between rocks in sun exposed places at the Mata Atlaˆntica Biome; between ca. 2,300–2,400 m alt.

Discussion. Hyvo¨nen (1989) recognizes three subspecies, which differ in subtle characters and are in need of a critical revision. A single variety is tentativily accepted for Brazil.

Specimen examined. BRAZIL.MINAS GERAIS:

Cambuı´, Peralta 267 (SP,SJRP).RIO DE JANEIRO: Itatiaia,

Ule 1764 (R); Resende, O. Yano et al. 22618 p.p. (SP).

Polytrichadelphus (Mu¨ll. Hal.) Mitt., J. Proc. Linn. Soc., Bot. 4: 97. 1859. TYPE: P. magellanicus

(Hedw.) Mitt.

Polytrichadelphus pseudopolytrichum (Raddi) G.L. Smith, Bryologist 79(4): 521. 1976 (1977) ; Catharinea pseudopolytrichum Raddi, Mem. Mat. Fis. Soc. Ital. Sci. Modena, Pt. Mem. Fis. 18: 343. 1820. TYPE: BRAZIL:RIO DE JANEIRO: Serra

dos O´ rga˜os, ‘‘Trovasi sulle montagne che costituiscono la cosi detta Serra do Frade, ove e comunissimo, matura e sour frutti’’, nel mese de Maggio, Raddi s.n. (syntypeFI(photo!)). Fig. 10

Polytrichadelphus croceus Mitt., J. Proc. Linn. Soc., Bot. 4: 98. 1859. TYPE: BRAZIL.RIO DE JANEIRO:

Organ Mts. (Serra dos O´ rga˜os), common on clayey banks, 3000 feet, Gardner 12 (syntypeE!).

Polytrichadelphus semiangulatus (Brid.) Mitt., J. Linn. Soc., Bot. 12: 609. 1869 ; Pogonatum

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Figure 9. Pogonatum perichaetiale subsp. oligodus. a. Female gametophyte, b–d. Leaves, e–f. Perichaetial leaves, g. Apical cells of the leaf, h. Margin cells of the leaf, i. Cells of the sheathing base, j. Transverse section of the apex of the leaf, k. Transverse section of the median leaf, l. Transverse section of the sheathing base, m. Lateral view of the lamella, n. Epidermal cells of the capsule, o. Transverse section of the epidermal cells of the capsule, p. Calyptra, q. Peristome teeth (Peralta 267-SP).

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TYPE: BRAZIL. Paris 1825, Persoon (lectotypeB!),

designated by Smith (1976).

Polytrichadelphus juniperaceus Mu¨ll. Hal., Gen. Musc. Frond.: 168. 1900. nom. nud.

Description. Plants large-sized, reddish or dark-green, 80–100 mm long. Stems erect, forming tufs. Rhizome prostrate. Rhizoids pale-brown. Leaves lanceolate, 6–8 mm long, widespreading, quadratic cells, 10 3 10mm, thick-walled, smooth, sheathing base conspicuous, plane margins, dentate, not bordered, costa broad 3/4–4/5, percurrent, lamellae occupying costae and laminae, 50–60 lamellae, 6 cells high, distal cells simple, pyriform, rugose, lamella cells in lateral view quadratic. Dioicous, perichaetium lateral, leaves long, perigonium lateral, leaves short and broad. Sporophyte 1(–2), seta 50–60 mm long. Capsule asymmetric, urn cylindric, stomata present in all urn surface, operculum conic, long-rostrate, 64 teeth. Calyptra cucullate, glabrous.

Distribution. Endemic to the Southeast and South of Brazil (States of Espı´rito Santo, Minas Gerais, Parana´, Rio de Janeiro, Rio Grande do Sul, Santa Catarina and Sa˜o Paulo).

Habitat. On soil along trails and on banks at the margin of submontane and montane forests at the Mata Atlaˆntica and Pampa Biomes; between ca. 600– 2,200 m alt.

Discussion. This is the only Brazilian species with perichaetia developed on small branches. Others diagnostic features are the red colour of sheathing base, pyriform and verrucose distal cells of lamella and imbricate and acute leaf apex.

Some samples are erroneously identified as P. magellanicus (Hedw.) Mitt. This species has robust habit and the agude leaf apex, and is restricted to the southern South America (i.e., below the 40u parallel; Smith 1976). Some samples are identified as Polytrichastrum formosum (Hedw.) G.L. Smith but this species has the terminal perichaetia and hairy calyptrae, this species have the distal cells of lamella verrucose in common with Polytrichadelphus pseudopolytrichum (Raddi) G.L. Smith.

Selected specimen examined. BRAZIL.ESPı´RITO SANTO: Conceic¸a˜o do Castelo, O. Yano et al. 4949 (SP);

Muniz Freire, O. Yano et al. 4953 (SP).MINAS GERAIS:

Itabira do Campo, Mattos 18 (R); Alto Caparao´,

Parque Nacional do Caparao´, Visnadi & Vital 2632

(SP); Caldas, Pietrobom-Silva 1978 (MO,SJRP,SP);

Caratinga, Vital 5858 (NY,SP); Catas Altas, Parque

Natural do Carac¸a, Peralta et al. 6818 (SP); Itamonte,

O. Yano & Visnadi 9878 (SP); Lima Duarte, Parque

Estadual do Ibitipoca, O. Yano et al. 21546 (SP). PARANA´: Bocaiu´va do Sul, Hatschbach 23456 (MBM; PACA); Jaguariaı´va, O. Yano et al. 5230, 5232 (SP);

Laranjeiras do Sul, Vital 5786 (NY,SP); Mallet, Wasum

2814, 2833 (SP;MBM,SP); Tijucas do Sul, O. Yano &

Shirata 27872 (SP).RIO DE JANEIRO: Angra dos Reis, O.

Yano et al. 23793 (SP); Itatiaia, Ule 1768 (R); Nova

Friburgo, Sehnem 6744 (PACA;SP).RIO GRANDE DO SUL:

Bom Jesus, Sehnem 221 (NY, PACA,SP); Sa˜o Francisco

de Paula, Sehnem 5344 (PACA;SP).SANTA CATARINA:

Bom Retiro, Scha¨fer-Verwimp & Verwimp 10491 (SP);

Negro, Vital 5602 (NY,SP).SA˜ O PAULO: Apiaı´, Vital 2302 (SP); Campos do Jorda˜o, Froehlich 17 (S);

Caraguatatuba, Sucre et al. 7007 (MO,RB); Cunha,

Custo´dio Filho 549 (SP); Iporanga, Joly 1235 (SP);

Itapecirica da Serra, Eiten & Godland 6317 (NY,SP);

Jambeiro, O. Yano & Yano 22683 (SP); Natividade da

Serra, O. Yano et al. 7734 (SP); Piquete, Peralta 3618

(SP); Rio Grande da Serra, 7-VII-1901, Schiffner s.n.

(NY); Santo Antonio do Pinhal, O. Yano 1156 (SP);

Sa˜o Paulo, Schiffner 937 (NY,S); Ubatuba, O. Yano et

al. 8109 (SP).

Polytrichum Hedw., Sp. Musc. Frond.: 88. 1801. TYPE: P. commune Hedw.

Polytrichum angustifolium Mitt., J. Linn. Soc., Bot. 12: 622. 1869. TYPE: BRAZIL. Brasilia Tropical,

Burchell 3768 (syntypesBM!,E!). Fig. 11

Polytrichum brasiliense Hampe, Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn. ser. 3, 10: 53. 1872 ; Pogonatum brasiliense (Hampe) A. Jaeger, Ber. Tha¨tigk. St. Gallischen Naturwiss. Ges. 1873–74: 266. (Gen. Sp. Musc. 1: 728). TYPE: BRAZIL.RIO DE JANEIRO: Rio de Janeiro,

Glaziou 5199 (syntypePC!,B!). syn. nov.

Polytrichum angustifolium Schimp., Bull. Herb. Boissier 5: 178. 1897. nom. illeg. non Mitten (1869).

Polytrichum alticaule Mu¨ll. Hal., Hedwigia (Beibl.) 38(1): 59. 1899. nom. nud

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lanceolate, 6–8 mm long, widespreading, quadratic cells, 10 3 10mm, thick-walled, smooth, sheathing base conspicuous, plane margins, dentate, not bordered, costa broad 3/4–4/5, percurrent, lamellae occupying costae and laminae, 60–70 lamellae, 7–8 cells high, distal cells single, pyriform, smooth, lamella cells in lateral view quadratic. Dioicous, perichaetium terminal, leaves long, perigonium terminal, leaves short and broad. Sporophyte single, seta 40–60 mm long. Capsule symmetric, urn cylindric, apophysis present, stomata present at urn base, operculum conic, long-rostrate, 64 teeth. Calyptra cucullate, hairy.

Distribution. Endemic to the Southern Brazil (States of Espı´rito Santo, Minas Gerais, Parana´, Rio de Janeiro, Rio Grande do Sul, Santa Catarina and Sa˜o Paulo).

Habitat. On soil, along trails in margins of montane forests at the Mata Atlaˆntica Biome between ca. 600–2,200 m alt.

Discussion. Polytrichum angustifolium is characterized by its pyriform distal cells of lamella and the plane leaf margin. The gametophytes are rigid and have a wood-like aspect when dry. The old leaves are falcate and whitish, while in the other species of the genus the leaves are lax and golden-brown. This is the largest species of the family reaching 800 mm high. Some samples are

erroneously identified as Polytrichastrum formosum (Hedw.) G.L. Smith., but this species, has rhombic and verrucose distal cells of lamella and lacks an apophysis.

Polytrichum brasiliense is considered conspecific with P. angustifolium. Polytrichum angustifolium Schimper (1897) is an invalid name (later homonym of P. angustifolium Mitt. [1869]) and the original material belongs in fact to P. juniperinum (Messmer & Frye (1947). Polytrichum xanthopillum Wilson ex Mitt. is similar to P. angustifolium, however, in P. xanthopilum the margin of lamellae in lateral view is denticulate.

Selected specimen examined. BRAZIL.ESPı´RITO SANTO: Caparao´, Scha¨fer-Verwimp & Verwimp 11489

(SP); Santa Teresa, O. Yano et al. 4922 (SP).MINAS GERAIS: Alto Caparao´, Scha¨fer-Verwimp & Verwimp

8903 (SP); Belo Horizonte, Mello-Filho 224, 3446 (R, NY;R); Caldas, A.F. Regnell 35c (S); Camanducaia,

Peralta et al. 3453, 3491, 3512, 3521 (SP); Itamonte, V.

Schiffner 749 (NY,S); Nova Lima, Carac¸a, 1885,

Wainio s.n. (PC); Ouro Preto, II-1890, Ule s.n. (B;BM; SP); Santa Ba´rbara, Ule 1317 (R).PARANA´: Campo

Largo, Hatschbach 19524 (MBM); Ponta Grossa, Vital

5818 (SP,NY).RIO DE JANEIRO: Itatiaia, Serra do Itatiaia,

Ule 1765 (R); idem, Ule 1766 (R): Nova Friburgo,

Luetzelburg 6873a (B); Petro´polis, Luetzelburg 15457

(B); Resende, Parque Nacional do Itatiaia,. Eiten &

Eiten 7631 (SP); Tereso´polis, Serra dos O´ rga˜os, Brade

12538 (R).RIO GRANDE DO SUL: Cambara´ do Sul, Serra

da Rocinha, Sehnem 7805 (PACA,ICN,R,RB,SP); Dois

Irma˜os, Morro Reuter, Sehnem 13827 (PACA,SP); Sa˜o

Francisco de Paula, Taimbe´, Sehnem 5272 (PACA,SP). SANTA CATARINA: Bom Retiro, Sehnem 6995 (PACA,SP);

Irineo´polis, Reitz & Klein 11568 (SP).SA˜ O PAULO: Campos do Jorda˜o, O. Yano & Watanabe 9220 (SP);

Natividade da Serra, O. Yano et al. 7735 (SP);

Pindamonhangaba, D.M. Vital 7063 (SP,NY); Piquete,

O. Yano & Prado 26203 (SP); Saleso´polis, 21-III-1991,

Vital s.n. (SP).

Polytrichum commune Hedw., Sp. Musc. Frond.: 88. 1801. TYPE: vulgaris per totam Europam.

Fig. 12 Polytrichum assimile Hampe, Flora 64: 378. 1881.

TYPE: BRAZIL.RIO DE JANEIRO: Glaziou 11729

(syntypeBM?), ex descr.

Polytrichum subremotifolium Geh. & Hampe, Flora 64: 377. 1881. TYPE: BRAZIL.SA˜ O PAULO: Apiaı´, sterile, V-1879, prope Faxina, Puiggari 634 (syntypeH-BR!). syn. nov.

Polytrichum regnellii Broth., Gen. Musc. Frond.: 184. 1900. nom. nud.

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present, stomata present at urn base, operculum conic, long-rostrate, 64 teeth. Calyptra cucullate, hairy.

Distribution. Cosmopolitan. Brazil (States of Amazonas, Bahia, Distrito Federal, Espı´rito Santo, Goia´s, Minas Gerais, Parana´, Rio de Janeiro, Roraima, Rio Grande do Sul, Santa Catarina and Sa˜o Paulo).

Habitat. On soil, along trails and on banks at the margin of low, submontane, montane forests and high fields at the Amazonia, Mata Atlaˆntica, Cerrado, Caatinga and Pampa Biomes; between ca. 600– 2,000 m alt.

Discussion. This species is differentiated by the U-shaped distal cells of the lamellae and the plane margin. The Hedwig original collection is housed in

Gand needs evaluation before designation of types.

The type of Polytrichum subremotifolium Geh. & Hampe (H!) cannot be distinguished from this

species and the name is considered synonymous with P. commune. Some samples are erroneously identified as P. brachymitrium Mull. Hal., a species restricted to Venezuela and Colombia and lacking in Brazil. The illustration attachment in the type (FH(photo!))

shows the pyriform distal cells of lamella. The shape of the distal cells in plants identified as P.

brachymitrium are U-shaped and sometimes pyriform, variation reported for P. commune (Crum & Anderson 1981). Polytrichum ericoides Hampe has the distal cells of lamella U-shaped like in P. commune (Smith 1976) but is restricted to the Andes. It differs by its small-sized, imbricate leaves and coarsely serrate leaf margin.

Selected specimen examined. BRAZIL. AMAZONAS:

Sa˜o Gabriel da Cachoeira, Fros 28248 (MO).BAHIA: Rio

de Contas, Harley et al. 19616 (E,NY); Santa

Teresinha, Mello et al. 2689 (SP).DISTRITO FEDERAL:

Brası´lia, Vital 6285 (SP,NY).ESPı´RITO SANTO: Iu´na,

Parque Nacional do Caparao´, Vital & Buck 11768 (B, SP,NY).GOIA´S, Alto Paraı´so de Goia´s, Irwin et al. 32845

(MO,NY,S); Cristalina, Irwin et al. 9851 (NY,SP).MINAS GERAIS: Bara˜o de Cocais, Irwin et al. 29339 (NY);

Caldas, A.F. Regnell s.n. (H); Caratinga, Vital 5855 (SP, NY); Lima Duarte, Parque Estadual do Ibitipoca,

Krieger 13935 (CESJ); Nova Lima, Vital 5517 (SP,NY);

Ouro Preto, III-1892, Ule s.n. (B,NY,S,W).PARANA´:

Antonina, Hatschbach 14136 (MBM,PACA,UPCB);

Bocaiu´va do Sul, Poliquesi & Cordeiro 151 (MBM);

Curitiba, J. Weir 99 (E,NY); Desvio, Ypiranga,

28-V-1909, P. Duse´n s.n. (B,NY, S).RIO DE JANEIRO: Itatiaia,

O. Yano et al. 19079 (SP); Nova Friburgo, Costa et al.

651 (RB); Parati, Costa et al. 1209 (RB); Tereso´polis,

VII-1901, Goeldi s.n. (MG,SP,W).RIO GRANDE DO SUL:

Caxias do Sul, 30-X-1988, Wasum et al. s.n. (B,MO);

Flores da Cunha, Wasum s.n. (HUCS,NY); Vacaria,

Sehnem 5902 (PACA,SP).RORAIMA: Caracaraı´, Prance et

al. 21441 (NY).SANTA CATARINA: Bom Retiro, Reitz 2431

(NY); Papanduva, Vital 5645 (SP,NY); Ponte Alta, Vital

9415 (SP); Urubici, Scha¨fer-Verwimp & Verwimp

10493 (SP).SA˜ O PAULO: Barra do Turvo, O. Yano 2289 (SP); Campo Grande, Schiffner 355 (S,W); Campos do

Jorda˜o, Peralta 191 (SP,SJRP); Cunha, Peralta et al.

3764 (SP); Guarulhos, O. Yano 312 (SP); Ilhabela, O.

Yano 5156 (SP); Moji Guac¸u, Visnadi &Vital 729 (SP);

Paraibuna, Vital 7078 (SP, NY); Peruı´be, O. Yano &

Mello 11445 (SP); Rio Grande da Serra, Baranov 78

(SP); Sa˜o Jose´ do Barreiro, Vattimo 3–101 (R); Sa˜o

Paulo, Schiffner 667 (B); Sa˜o Sima˜o, Vital 2453 (SP);

Ubatuba, Visnadi &Vital 1037 (SP).

Polytrichum juniperinum Hedw., Sp. Musc. Frond.: 89. 1801. TYPE: SUIc¸A. locus montanus

subalpestris, delineata exemplo ex Helvetia,

s.col. Fig. 13

Polytrichum reflexifolium Herzog, Arch. Bot. Sa˜o Paulo 1(2): 71. 1924. TYPE: BRAZIL.SA˜ O PAULO: Tremembe´, 560 m alt., 6-II-1921, Soares s.n. (holotype SP!, isotypeMANCH!). syn. nov.

Polytrichadelphus organensis Herzog, Repert. Sp. Nov. Regni Veg. 21: 31. 1925. TYPE: BRAZIL.RIO DE JANEIRO: Serra dos O´ rga˜os, P. Luetzelburg 7260

(syntypeB!). syn. nov.

Polytrichum aristiflorum Mitt., J. Linn. Soc., Bot. 12: 620. 1869. TYPE: BRAZIL. Weir 11 (holotypeNY!,

isotypes E!,MANCH!).

Polytrichum vulgare Hedw., Rodrigue´sia 54: 266. 1980. nom. nud.

Polytrichum pallidicaule Mu¨ll. Hal., Hedwigia (Beibl.) 38(1): 57. 1899. nom. nud.

Polytrichum brachypyxis Mu¨ll. Hal., Hedwigia (Beibl.) 38(1): 57. 1899. nom. nud.

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lanceolate, 6–7 mm long, widespreading, quadratic cells, 10 3 30–50mm, thick-walled, smooth, sheathing base conspicuous, involute margins, dentate, not bordered, costa broad 3/4–4/5, percurrent, lamellae occupying costae and laminae, 40–44 lamellae, 6 cells high, distal cells single, pyriform, smooth, lamella cells in lateral view quadratic. Dioicous, perichaetium terminal, leaves long, perigonium terminal, leaves short and broad. Sporophyte single, seta 40–50 mm long. Capsule symmetric, urn cylindric, apophysis present, stomata present at urn base, operculum conic, long-rostrate, 64 teeth. Calyptra cucullate, hairy.

Distribution. Cosmopolitan; in Brazil known from States of Bahia, Distrito Federal, Espı´rito Santo, Goia´s, Minas Gerais, Parana´, Rio de Janeiro, Roraima, Rio Grande do Sul, Santa Catarina, Sa˜o Paulo and Tocantins.

Habitat. On soil along trails and on banks and rocks at the edge of low, submontane, montane forests and high fields at the Mata Atlaˆntica, Cerrado, Caatinga and Pampa Biomes; between ca. 600– 2,200 m alt.

Discussion. This species is easily diagnosed among sympatric species by the involute leaf margin. The type specimen is the same as in P. commune and needs a critical revision. The original description of P. juniperinum var. paulense (Geh. & Hampe) Herzog, offers no diagnostic characteristics compared to the type variety of P. juniperinum. We have not

been able to locate the type specimen and we consider the name doubtful name.

Some samples are erroneously identified as Polytrichum appressum Brid, a species otherwise known only from Africa. All of them belong to P. juniperinum. Some samples are erroneously identified as Polytrichum strictum Menzies ex Brid. (5 P. alpestre Hoppe). This species has in common with P. juniperinum an involute margin, (this is not clear: but it is a name used to plants with elongate habit, probably because the association with Sphagnum (Crum & Anderson 1981), this species or subspecies needs a revision. The Brazilian plants identified as P. alpestre are a P. juniperinum environmental variation without morphologic or anatomic differences.

Messmer & Frye (1947) show the wide variation found in P. juniperinum, as reflected by the long list of synonyms and add as synonyms the Brazilian species P. aristifolium Mitt. and P. angustifolium Schimp.

Vattimo-Gil & Vattimo (1980), reported the name Polytrichum vulgare Hedw. based on the specimen Fidalgo & Kauffmann-Fidalgo 5 (MO!,RB!).

This name has never been published and a reexamination of this specimen revealed it to be Polytrichum juniperinum Hedw. The species Polytrichum reflexifolium Herzog (SP!) is a new

synonym of P. juniperinum. Wijk et al. (1967) listed this species as a synonym of P. juniperinum var. paulense (Geh. & Hampe) Herzog, and we did not find any differences besides the morphological variation of P. juniperinum.

Selected specimen examined. BRAZIL.BAHIA:

Abaı´ra, Harley & Campos 51119 (SP); Camac¸ari,

Mattos Silva & Brito s.n. (ALCB); Rio de Contas,

Harley et al. 26272, 26278 (NY,E; E).ESPI´RITO SANTO:

Caparao´, Scha¨fer-Verwimp & Verwimp 8968 (SP);

Muniz Freire, O. Yano et al. 4957 (SP). GOIA´S: Alto

Paraı´so de Goia´s, Hatschbach et al. 36477 (MBM, PACA).MINAS GERAIS: Itabira do Campo, Ule 1323 (R);

Caparao´, Vital & Buck 11602 (SP,NY); Carangola, Y.

Mexia 4304 (MO,NY); Conceic¸a˜o do Mato Dentro, O.

Yano 513 (SP,NY); Tiradentes, O. Yano et al. 21871

(SP).PARANA´: Bocaiu´va do Sul, O. Yano & F.C. Silva

11303 (SP); General Carneiro, O. Yano & Pirani 6505

(SP); Quedas do Iguac¸u, O. Yano et al. 5413 (SP).RIO

Figure 14. General distribution of Polytrichaceae in Brazil.

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DE JANEIRO: Angra dos Reis, O. Yano et al. 23731 (SP);

Itatiaia, Ule 1229 (R); Nova Friburgo, VI-1925, Lea˜o

s.n. (MO,RB,S); Parati, Vital 7246 (MO,NY,SP).RIO GRANDE DO SUL: Cac¸apava do Sul, O. Yano & Pirani

5858 (SP); Cambara´ do Sul, Sehnem 4778 (PACA,R,RB, SP); Garibaldi, 15-V-1988, Wasum et al. s.n. (MO,RB);

Montenegro, Sehnem 2095 (R,RB,SP); Santa Cruz do

Sul, Sehnem 5264 (PACA,SP); Silveira Martins,

19-III-1893, Lindman s.n. (R).SANTA CATARINA: Bom Retiro,

Reitz 2366 (B,MO,NY); Capivari de Baixo, V-1891, Ule

s.n. (B; NY;S); Curitibanos, Vitt 21037 (NY,S);

Floriano´polis, O. Yano 2281 (SP); Lages, Sehnem 5410

(PACA,SP); Minas, II-1890, Ule s.n. (B,NY,SP,R);

Orleans, IX-1889, Ule s.n. (B,NY,SP;R,SP,S); Palhoc¸a,

Reitz & Klein 2513 (MBM,NY,PACA,B).SA˜ O PAULO: Amparo, O. Yano & Santos 6035 (SP); Apiaı´, Puiggari

s.n. (S); Bananal, O. Yano 1243 (NY,SP); Barra do

Turvo, Vital 2830 (ALCB, MO,SP); Barueri, Vital 6610

(NY, SP); Botucatu, Vital 2523 (SP); Campos do

Jorda˜o, O. Yano & Marcelli 19717 (SP); Ibiu´na, O.

Yano 4564 (SP); Ilhabela, O. Yano 5155 (SP);

Itapecirica da Serra, Eiten & Godland 6312 (MO,NY, SP); Itu, Weir 11 (E,NY,S); Juquitiba, O. Yano 544 (NY, SP); Sa˜o Bernardo do Campo, Vital 5379 (MO,NY,SP);

Sa˜o Paulo, VIII-1901, V. Schiffner s.n. (B); Sa˜o

Sebastia˜o, O. Yano et al. 7931 (SP); Serra Negra, O.

Yano et al. 18496 (SP).TOCANTINS: Paraı´so de

Tocantins, O. Yano & Kida 26753 (SP).

LIST OFDOUBTFULNAMES

The status of the following taxa could not be determined as no type material could be located.

Polytrichum glabrum Brid. ex Schrad., J. Bot. (Schrader) 1801: 196. 1803. TYPE: In insula

Bourbonis montibus altissimis, in ea part, quae plaine des Caffes decitur, et in fossis exiccatis; Commerson 1771, Augusto legit. In Brasilia auctores nennulli indicant; an recte dubii sumus (syntypeB?).

Polytrichum involutum Hampe, Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn. ser. 3, 9– 10: 263. 1878. TYPE: BRAZIL. Minas Gerais, ad

fontes fluvii Rio Preto, ? 9074 (syntypeBM?,G?).

Polytrichum juniperinum Hedw. var. paulense (Geh. & Hampe) Herzog, Arch. Bot. Sa˜o Paulo 1(2): 71. 1924 ; Polytrichum paulense Geh. & Hampe, Flora 64: 376. 1881. TYPE: BRAZIL. Sa˜o Paulo:

Apeai (Apiaı´), prope Faxina, sterile, V-1879, Puiggari 285, 908 (syntypeBM?,B?).

Polytrichum petersianum Mu¨ll. Hal., Linnaea 38: 575. 1874. TYPE: BRAZIL. Nova Friburgo, in solo

argiloso aprico nudo, 20-VII-1871, Peters s.n. (syntypesHBG?,BM?,NY?).

Polytrichum subcarinatum Hampe, Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn. ser. 3, 6: 150. 1874. TYPE: BRAZIL. Glaziou 5201 (syntypes BM?,B?).

LIST OF NOMINA NUDA

Pogonatum distantifolium Mu¨ll. Hal. ex Herzog, Biblioth. Bot. 87: 103. 1916. nom. nud.

Pogonatum serricolum Mu¨ll. Hal., Gen. Musc. Frond.: 176. 1900. nom. nud.

Polytrichum caudatum Mu¨ll. Hal., Gen. Musc. Frond.: 184. 1900. nom. nud.

Polytrichum lanipes Mu¨ll. Hal., Gen. Musc. Frond.: 185. 1900. nom. nud.

Polytrichum micropysix Mu¨ll. Hal., Gen. Musc. Frond.: 181. 1901. nom. nud.

Polytrichum pallidipes Mu¨ll. Hal., Gen. Musc. Frond.: 184. 1900. nom. nud.

Polytrichum pycnocarpum Mu¨ll. Hal., Gen. Musc. Frond.: 182. 1900. nom. nud.

Polytrichum uleanum Mu¨ll. Hal., Gen. Musc. Frond.: 184. 1900. nom. nud.

LITERATURECITED

Churchill, S. P. & E. Linares C. 1995. Prodromus Bryologiae Novo-Granatensis: Introduccio´n a la Flora de Musgos de Colombia. Biblioteca ‘‘Jose Jeronimo Triana’’ 12: 1–924. Crosby, M. R., R. E. Magill, B. Allen & S. He. 1999. A Checklist

of the Mosses. Missouri Botanical Garden, St. Louis. Crum, H. & L. E. Anderson. 1981. Mosses Eastern North

America, vol. 2. Columbia University Press, New York. Edwards, S. R. 1995. Polytrichum piliferum and P. juniperinum.

Bulletin of the British Bryological Society 65: 49–50. Farias, H. C. 1982. A famı´lia Polytrichaceae no Rio Grande do

Sul, Brasil. Dissertac¸a˜o de Mestrado, Universidade Federal do Rio Grande do Sul, Porto Alegre.

———. 1984. A famı´lia Polytrichaceae no Rio Grande do Sul, Brasil. I Taxonomia. Iheringia, Se´rie Botaˆnica 32: 77–89. Goffinet, B., W. R. Buck & A. J. Shaw. 2009. Morphology,

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Gradstein, S. R., S. P. Churchill & N. Salazar-Allen (eds.). 2001, Guide to the Bryophytes to Tropical America. Memoirs of The New York Botanical Garden 86: 1–577.

Hyvo¨nen, J. 1989. A synopsis of genus Pogonatum

(Polytrichaceae, Musci). Acta Botanica Fennica 138: 1–87. ———, T. A. Hedderson, G. L. Smith Merrill, J. G. Gibbings &

S. Korkinen. 1998. On phylogeny of the Polytrichales. The Bryologist 101: 489–504.

———, S. Korkinen, G. L. Smith Merrill, T. A. Hedderson & S. Stenroos. 2004. Phylogeny of the Polytrichales (Bryophyta) based on simultaneous analysis of molecular and morphological data. Molecular Phylogenetic and Evolution 31: 915–928.

Menzel, M. 1985[1986]. Die Gattung Pogonatum P. Beauv. (Polytrichales, Musci) in Lateinamerika 1. Taxonomie und Geographie von Pogonatum campylocarpum (C. Muell.) Mitt. und P. subflexuosum (Lor.) Broth. Lindbergia 11: 134–140.

Messmer, L. & T. C. Frye. 1947. The Polytrichum group between South America and the United States. The Bryologist 50: 259–268.

Mitten, W. 1869. Musci Austro-Americani. Journal of the Linnean Society, Botany 12: 1–659.

Mu¨llen, D. & T. C. Frye. 1947. Middle and South American species of Oligotrichum. The Bryologist 50: 67–79. Mu¨ller, C. 1848–1849. Synopsis Muscorum Frondosorum

omnium hucusque Cognitorum 1. Berlin: A. Foerstner. Nyholm, E. 1971. Studies in the genus Atrichum P. Beauv. A

short survey of the genus and species. Lindbergia 1: 1–33. Raddi, G. 1820. Di alcune specie nuove di rettili, e plante Brasiliane.

Atti della Societa Italiana di Scienze Naturali 18: 3–39. Smith, G. L. 1976. Neotropical Polytrichaceae V. The Bryologist

79: 521–522.

Vattimo-Gil, I. & I. Vattimo. 1980. Bryophyta (Musci) do herba´rio do Jardim Botaˆnico do Rio de Janeiro II. Rodrigue´sia 54: 257–267.

Wijk, R., W. D. Margadant & P. A. Florschu¨tz. 1967. Index Muscorum. 4 (P–S). Regnum Vegetabile 48: 1–604. Yano, O. 1996. A checklist of Brazilian bryophytes. Boletim do

Instituto de Botaˆnica 10: 47–232.

ms received October 28, 2009; accepted May 28, 2010.

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