Taxonomic treatment of the Polytrichaceae from Brazil

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
URL: http://www.bioone.org/doi/full/10.1639/0007-2745-113.3.646

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 Taxonomic treatment of the Polytrichaceae from Brazil

Denilson Fernandes Peralta1 and Olga Yano

Instituto de Botânica, Caixa Postal 3005, 01031-970 São 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.

¤ ¤ ¤

The Polytrichaceae are characterized by formation of (5 Polytrichadelphus pseudopolytrichum (Raddi) G.L.

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 Hyvönen 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
Smith) from Serra do Frade (today Serra dos Órgãos),
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.

1
Corresponding author e-mail: MATERIALS AND METHODS
denilsonfp@yahoo.com.br Materials were obtained both from field
DOI: 10.1639/0007-2745-113.3.646 collection and loans from herbaria (ALCB, B, BM, C,

The Bryologist 113(3), pp. 646–672 0007-2745/10/$2.75/0

Copyright E2010 by The American Bryological and Lichenological Society, Inc.
 Peralta & Yano: Taxonomic treatment of the Polytrichaceae from Brazil
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.
TAXONOMIC TREATMENT
Polytrichaceae Schwägr. in Willd., Sp. Pl. ed. 5(2): 1.
1830. Type: Polytrichum Hedw.
KEYS TO THE SPECIES OF POLYTRICHACEAE 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
647
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 (Müll. Hal.) A. Jaeger, Ber.
Thätigk. St. Gallischen Naturwiss. Ges. 1873–74:
241 (Gen. et Sp. Musc. 1: 709). 1875 ;
Catharinea androgyna Müll. 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.
(syntypes MEL? hb. Kunzeanum, BM?). Fig. 1
Catharinea ulei Müll. Hal., Hedwigia 38 (Beibl. 1):
57. 1899. nom. nud.
Atrichum mosenii (Broth.) Paris, Index Bryol. Suppl.
17. 1900 ; Catharinea mosenii Broth., Bih.
Kongl. Svenksa Vetensk.-Akad. Handl. 21 Afd.
648 The Bryologist 113(3): 2010

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).
 Peralta & Yano: Taxonomic treatment of the Polytrichaceae from Brazil 649

3(3): 32. 1895. TYPE: BRAZIL, MINAS GERAIS: Caldas, The type specimens of Atrichum mosenii
VII-1823, Mosén 332 (syntype H-BR!). (Broth.) Paris (H!) and A. riograndense (Broth.) Paris
Atrichum riograndense (Broth.) Paris, Index Bryol. (H! R!) cannot be distinguished from A. androgynum
281: 70. 1904 ; Catharinea rio-grandensis and the names should be considered synonyms of A.
Broth., Bih. Kongl. Svenksa Vetensk.-Akad. androgynum as suggested by Nyholm (1971).
Handl. 26 Afd. 3(7): 32. 1900. TYPE: BRAZIL, RIO Selected specimen examined. BRAZIL. MINAS
GRANDE DO SUL: Silveira Martins ad, Terram GERAIS: Caldas, Mosén 331 (H); Caparaó Novo, Parque
marginis viae declivum umbrosam, 19-III-1893, Nacional do Caparaó, Vital & Buck 11630 (SP).
Lindman 210 (syntypes H-BR!, R!). PARANÁ: Dorizon, Hotel Água Dorizon, Wasum et al.
3745 (MO, SP); Guarapuava, Fazenda Campo Real,
Description. Plants medium-sized, dark-green
Hatschbach 21025 (MBM, MO, PACA); Palmas, O. Yano
to brown, 50–100 mm long. Stems erect, forming
& Pirani 6551 (SP). RIO DE JANEIRO: Itatiaia, Ule 38 (S);
tufs. Rhizome prostrate. Rhizoids pale-brown. Leaves
Nova Friburgo, Kurtz & Dihlmann 27 (RB); Rio de
lanceolate, 4–7 mm long, widespreading, quadratic-
Janeiro, Costa & Molinaro 243 (RB); Teresópolis,
hexagonal cells, 10–15 3 10–15 mm, thick-walled,
Parque Nacional da Serra dos Órgãos, O. Yano &
smooth, sheathing base inconspicuous, plane
Costa 13549 (SP). RIO GRANDE DO SUL: Bom Jesus,
margins, double dentate, bordered, costa narrow 1/
Sehnem 220 (PACA, SP); Cambará do Sul, Fortaleza dos
5–1/3, percurrent, lamellae occupying only the
Aparados, Vianna 615 (ICN); Canela, Farias 181 (ICN);
costae, 4–6 lamellae, 2–4 cells high, distal cells simple,
Caxias do Sul, Criúva, 17-IX-1988, Rossato et al. s.n.
rhombic, smooth, lamella cells in lateral view
(MO); Dois Irmãos, Morro Reuter, Sehnem 12513
quadratic-hexagonal. Monoicous, perichaetium and
(PACA, SP); Esmeralda, Vianna 735 (ICN); Flores da
perigonium terminal, leaves not differentiated.
Cunha, 19-II-1992, Brinker et al. s.n. (MO);
Sporophyte 1(–3), seta 30 mm long. Capsule
Jaquirana, Wasum et al. 775 (MBM, ALCB);
asymmetric, urn cylindric, stomata absent,
Montenegro, Sehnem 4038 (PACA, R, RB, SP); idem, São
operculum conic, long-rostrate, 32 teeth. Calyptra
Francisco de Paula, Vila Oliva, Sehnem 2736 (PACA,
cucullate, glabrous.
SP); São Leopoldo, Feitoria, Sehnem 10 (PACA, SP);
Distribution. Africa, Mexico, Guatemala,
Silveira Martins, Lindman 191 (H). SANTA CATARINA:
Honduras, Costa Rica, Colombia and Brazil (States
Serra Geral, V-1890, Ule s.n. (R); idem, Ule 38 (SP);
of Minas Gerais, Paraná, Rio de Janeiro, Rio Grande
São José, Serra da Boa Vista, Reitz & Klein 9926, 9927
do Sul, Santa Catarina and São Paulo).
(SP). SÃO PAULO: Bananal, Nova Suiça, O. Yano 1264
Habitat. On soil, at margins of submontane and
(MO, SP); Campos da Cunha, Peralta et al. 3740 (SP);
montane forests, in the Bioma Mata Atlântica and
Campos do Jordão, Parque Estadual de Campos do
Pampa, between 600–2,000 m alt.
Jordão, O. Yano & Marcelli 19434 (SP); Cunha, Vital
Discussion. This species is diagnosed by the
7768 (NY, SP); Moji das Cruzes, Parque Municipal da
bordered leaves that are crisped when dry. Müller
Serra de Itapety, Peralta et al. 2754 (SP);
(1848–1849) described the species as either dioicous
Pindamonhangaba, Pico Itapeva, Vital 1012 (SP);
or monoicous, but only bisexual gametophytes, with
Santo André, Reserva Biológica do Alto da Serra de
many antheridia around the seta, were found among
Paranapiacaba, O. Yano & Kirizawa 9385 (SP); São
the material from Brazil.
José do Barreiro, Vattimo 3–112 (R); São Paulo,
Nyholm (1971) uses the lengh of lamina cells to
Parque Estadual das Fontes do Ipiranga, O. Yano et
separe the varieties of A. androgynum. But among the
al. 15650 (SP).
Brazilian material, we found a large variation on this
character even within a single population. Hence we Itatiella G.L. Smith, Mem. New York Bot. Gard.
treated all specimens as A. androgynum. 21(3): 52. 1971. TYPE: I. ulei (Müll. Hal.) G.L.
The collections from Brazil identified as Smith.
Atrichum oerstedianum (Müll. Hal.) Mitt. don’t have Itatiella ulei (Broth. ex Müll. Hal.) G.L. Smith, Mem.
the antheridia surrounded by the differentiated New York Bot. Gard. 21(3): 52. 1971 ;
perigonial leaves, as is clearly the case in the type (K!). Psilopilum ulei Broth. ; Psilopilum ulei Broth.
650 The Bryologist 113(3): 2010
ex Müll. Hal., Bull. Herb. Boissier 6: 26. 1898.
TYPE: BRAZIL. SERRA DE ITATIAIA, 2100 m, in campo,
III-1894, Ule 1905 (Bryotheca Brasiliensis 151)
(syntypes H-BR!, R!, FH (photo!), PC!, R!, S!, W!).
Polytrichum ulei Broth., Rodriguésia 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 10–15 mm, 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 Espı́rito Santo, Minas
Gerais, Paraná, Rio de Janeiro and São Paulo).
Habitat. On soil at the margins of montane
forest and high Campos in the Bioma Mata Atlântica,
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 in R
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 ESPı́RITO
SANTO: Iúna, Parque Nacional do Caparaó, Vital &
Buck 11764 (B, E, MO, MG, NY, RB, SP). MINAS GERAIS: Alto
Caparaó, Buck et al. 27004 (SP, MO, NY); Itamonte,
Schiffner 755 (NY, S); Lima Duarte, Parque Estadual
do Itibipoca, Krieger 16189 (PACA). PARANÁ: Dusén
2558 (B, E, S). RIO DE JANEIRO: Itatiaia, 1894, Ule s.n.
(W); Resende, Schiffner s.n (B, W).; Teresópolis,
Parque Nacional da Serra dos Órgãos, Costa et al.
4168 (RB); Visconde de Mauá, Schäfer-Verwimp &
Verwimp 8325 (MO, SP). SÃO PAULO: Campos do
Jordão, Parque Estadual de Campos do Jordão, 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 8–10 mm, 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 Atlântica Bioma at ca.
1,100 m alt.
Discussion. Notoligotrichum minimum is
differentiated from the other species of
 Peralta & Yano: Taxonomic treatment of the Polytrichaceae from Brazil 651

Figure 2. Itatiella ulei. a. Female gametophyte, b. Male gametophyte, c–e. Leaves, f–h. Perichaetial leaves, i–j. Perigonial leaves, k.
Apical cells of the leaf, l. Marginal 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 lamella, r. Epidermal cells of
the capsule, s. Transverse section of the epidermal cells of the capsule, t. Calyptra (Yano & Marcelli 15742-SP).
652 The Bryologist 113(3): 2010

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).
 Peralta & Yano: Taxonomic treatment of the Polytrichaceae from Brazil
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.
CONCEPCIÓN: 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 15–20 mm,
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 Atlântica and Pampa.
Discussion. This species is diagnosed by its
obtuse and inflexed leaf apex. Müllen & 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, Sangão do Macaco Branco, Bueno 3329,
3340 (ICN); Garibaldi, Schäfer-Verwimp & Verwimp
10703 (SP). SANTA CATARINA: Urubici, Serra do Corvo
Branco, Schäfer-Verwimp & Verwimp 13507 (SP).
653
Oligotrichum denudatum Merrill, Novon 1(3): 107.
1991. TYPE: BRAZIL. MINAS GERAIS: São 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
(holotype NY!; isotype SP!). 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 15–20 mm, 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 Atlântica
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: São
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.
(holotype NY!). Fig. 6
Polytrichum leucodontium Müll. Hal., Bot. Zeitung
(Berlin) 2: 726. 1844. Polytrichum nigricans J.D.
Hook. & Wilson, London J. Bot. 3: 151. 1844.
TYPE: BRAZIL. Gardner 10 (syntypes NY!, E!,
MANCH!).
Description. Plants medium-sized, dark-green
to brown, 30–40 mm long. Stems erect, forming tufs.
Rhizome prostrate. Rhizoids pale-brown. Leaves
lanceolate-ligulate, 3.5–5.0 mm long, widespreading,
654 The Bryologist 113(3): 2010

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 (Schäfer-Verwimp & Verwimp 13507-SP).
 Peralta & Yano: Taxonomic treatment of the Polytrichaceae from Brazil 655

Figure 5. Oligotrichum denudatum. a. Female gametophyte, b. Male gametophyte, c–e. Leaves, f–g. Perichaetial leaves, h–i.
Perigonial leaves, j. Apical 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. Epidermal cells of the capsule, with stomata, p. Transverse
section of the epidermal cells of the capsule, q. Calyptra, r. Peristome teeth (isotype, Vital & Buck 11941-SP).
656 The Bryologist 113(3): 2010

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).
 Peralta & Yano: Taxonomic treatment of the Polytrichaceae from Brazil
quadratic cells, 10 3 15–20 mm, 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, Paraná, Rio de Janeiro, Rio Grande do Sul,
Santa Catarina and São Paulo).
Habitat. On soil along trails and on banks in
montane forests at the Mata Atlântica Bioma;
between ca. 1,800–2,200 m alt.
Discussion. Similar to Oligotrichum denudatum,
see differentiation under this species. Polytrichum
leucodontium Müll. 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, Schäfer-Verwimp & Verwimp 7521 (SP);
Caparaó, Parque Nacional do Caparaó, Vital & Buck
11605 (SP, NY); Delfim Moreira, Schäfer-Verwimp &
Verwimp 10042 (SP); Itamonte, O. Yano et al. 22628
(SP); Marmelópolis, Peralta 3568 (SP); Passa Quatro,
III-1921, Zikán s.n. (B, MANCH); Santa Bárbara, Vital
& Favalli 10830 (SP). PARANÁ: 1902, P. Dusén s.n
(MANCH, W, S); Castro, Weir 115 (NY); Morretes, Serra
da Graciosa, Schwacke 1726 (RB); União da Vitória, O.
Yano & Pirani 6484 (SP). RIO DE JANEIRO: Engenheiro
Passos, O. Yano & Santos 7670 (SP); Itatiaia, I-1902,
Dusén s.n. (S); Nova Friburgo, VII-1835, Lusihnatt
s.n. (S); Parati, Vital 15401 (SP); Resende, Dusén 2555
(B, W); Teresópolis, Serra dos Órgãos, X-1896, Ule s.n
(W). RIO GRANDE DO SUL: São Francisco de Paula, Serra
do Faxinal, Sehnem 5304 (PACA, ICN, SP). SANTA
CATARINA: Bom Retiro, Sehnem 7039 (PACA). SÃO PAULO:
Bananal, Nova Suiça, O. Yano 1244 (MO, SP); Campos
do Jordão, Parque Estadual de Campos do Jordão, 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);
657
São José 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 (Müll. Hal.) Mitt., J.
Linn. Soc., Bot. 12: 618. 1869 ; Polytrichum
campylocarpum Müll. Hal., Syn. Musc. Frond. 1:
209. 1849. TYPE: VENEZUELA. CARACAS: Galipan,
4000 (5000?) ft., I-1846, Funck & Schlim 379
(lectotype BM; isotypes GOET, JE, L, NY),
designated by Menzel (1985[1986]). Fig. 7
Pogonatum camptocaulon (Müll. Hal.) Paris, Index
Bryol. 978. 1898 ; Polytrichum camptocaulon
Müll. Hal., Acta Soc. Sci. Fenn. 19(5): 17. 1891.
TYPE: BRAZIL. MINAS GERAIS: Caraça, in terra
arenosa, IV-1885, Wainio s.n. (lectotype H-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 10–15 mm, 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 São Paulo).
Habitat. On soil, along trails and on banks at
the edge of montane forests at the Mata Atlântica
Bioma; between ca. 600–2,000 m alt.
Discussion. This species differs from the
Brazilian congeners by the lamellae with with two
pyriform distal cells, a unique feature among South
American species (Hyvönen 1989). The species is
similar to Pogonatum comosum (Müll. Hal.) Mitt.,
658 The Bryologist 113(3): 2010

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).
 Peralta & Yano: Taxonomic treatment of the Polytrichaceae from Brazil
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); Caparaó, Parque Nacional do
Caparaó, Vital & Buck 11607 (SP); Catas Altas, Peralta
et al. 6358 (SP); Itamonte, O. Yano & Visnadi 9904
(SP); Lima Duarte, Caraça, Ule 1318 (R),
Marmelópolis, 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); Teresópolis, Mautone
& Guimarães 118 (RB). RIO GRANDE DO SUL: Cambará do
Sul, Peralta & Gonçalves 3314, 3319, 3323 (SP); São
Francisco de Paula, Farias 185 (ICN). SANTA CATARINA:
Bom Retiro, Schäfer-Verwimp & Verwimp 10490 (SP);
Serra Geral, Vital & Buck 12425 (SP). SÃO PAULO:
Bananal, O. Yano 1226 (SP); Campos do Jordão,
Parque Estadual de Campos do Jordão, O. Yano &
Watanabe 9226 (SP); Cubatão, Vital 13986 (SP);
Cunha, Vital 7761 (SP); Pindamonhangaba, Pico do
Itapeva, Vital 642 (SP); Piquete, D.F. Peralta 3614
(SP); São Bernardo do Campo, O. Yano et al. 30337,
30351 (SP); São José do Barreiro, Loefgren & Edwal
537 (SP); São Luis do Paraitinga, O. Yano et al. 7791
(SP); São Paulo, Hoehne 128 (SP, MANCH).
Pogonatum pensilvanicum (Hedw.) P. Beauv.,
Mém. 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: Mühlenberg
(syntypes BM, G). Fig. 8
Pogonatum gardneri (Müll. Hal.) Mitt., J. Proc. Linn.
Soc., Bot. 8: 156. 1864 ; Polytrichum gardneri
Müll. Hal., Syn. Musc. Frond. 2: 560. 1851. TYPE:
BRAZIL. Tijuca, clay bank, XI-1836, Gardner 15
(syntype E!).
Pogonatum glaziovii (Hampe) A. Jaeger, Ber. Thatigk.
St. Gallischen Naturwiss. Ges. 1: 716. 1875 ;
659
Polytrichum glaziovii Hampe, Vidensk. Meddel.
Dansk Naturhist. Foren. Kjøbenhavn ser. 3 4: 54.
1872. TYPE: BRAZIL. Glaziou 5156 (lectotype BM,
isotype FH), designated by Hyvönen (1989).
Polytrichum campophilum Müll. 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 (Lectotype H-BR),
designated by Hyvönen (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 15 mm, 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,
Espı́rito Santo, Goiás, Minas Gerais, Paraná, Rio de
Janeiro, Rio Grande do Sul, Santa Catarina and São
Paulo.
Habitat. On soil along trails and on banks at
margins of montane forests at the Mata Atlântica,
Cerrado and Pampa Biomes; between ca. 400–
2,200 m alt.
Discussion. The persistent protonema
differentiates this species from other Brazilian taxa.
Two other species develop persitent protonemata
(Hyvönen 1989): Pogonatum brachyphyllum (Michx.)
P. Beauv. and P. spinulosum Mitt. the former is
endemic to North America and is diagnosed by a
truncate leaf apex and well developed lamellae.
Pogonatum spinulosum is endemic to Asia and is
characterized by the many teeth on the ventral side of
660 The Bryologist 113(3): 2010

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).
 Peralta & Yano: Taxonomic treatment of the Polytrichaceae from Brazil
the costa, and well developed lamellae with the
verrucose distal cells.
Selected specimen examined. BRAZIL. DISTRITO
FEDERAL: Mumunhas, Vital 13498 (SP). ESPı́RITO SANTO:
Domingos Martins, Schäfer-Verwimp & Verwimp
10189 (SP); Santa Teresa, O. Yano 3593 (SP). GOIÁS:
Alto Paraı́so de Goiás, 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). PARANÁ: 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, Dusén s.n. (e, s);
Nova Friburgo, 22-II-1903, P. Dusén s.n. (S); Parati,
Vital 7745 (MO, NY, SP). RIO GRANDE DO SUL: Cambará
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 Cecı́lia,
Vital 5594 (SP); Porto Belo, O. Yano & Yano 10038
(SP); Tijucas, Sehnem 3251 (PACA). SÃO PAULO: Atibaia,
Peralta et al. 5598 (SP); Barra do Turvo, Vital 3221
(SP); Brotas, Vital 11249 (SP); Campos do Jordão,
Parque Estadual de Campos do Jordão, O. Yano &
Watanabe 9221 (SP); Ibiúna, O. Yano & Yano 10074
(SP); Piquete, Peralta 3630 (SP); São Bernardo do
Campo, O. Yano et al. 15233 (SP); Santo André,
Schiffner 793 (S); Serra Negra, O. Yano et al. 18516
(SP); Ubatuba, Vital 4781 (SP).
Pogonatum perichaetiale (Mont.) A. Jaeger subsp.
oligodus (Kunze ex Müll. Hal.) Hyvönen, Acta
Bot. Fenn. 138: 15. 1989 ; Polytrichum oligodus
Kunze ex Müll. Hal., Syn. Musc. Frond. 1: 206.
1848. TYPE: CHILE. Poeppig 227 (lectotype BM,
isotypes G, H-BR, JE, HBG, PC), designated by
Menzel (1985[1986]). Fig. 9
Pogonatum itatiaiae (Müll. Hal.) Paris, Index Bryol.
Suppl.: 278. 1900 ; Polytrichum itatiaiae Müll.
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) (lectotype H-BR, isotypes, HBG,
R!), designated by Menzel (1985[1986]).
661
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 5–10 mm, 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 Atlântica Biome; between ca.
2,300–2,400 m alt.
Discussion. Hyvönen (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:
Cambuı́, Peralta 267 (SP, SJRP). RIO DE JANEIRO: Itatiaia,
Ule 1764 (R); Resende, O. Yano et al. 22618 p.p. (SP).
Polytrichadelphus (Müll. 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 Órgãos, ‘‘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. (syntype FI (photo!)). Fig. 10
Polytrichadelphus croceus Mitt., J. Proc. Linn. Soc.,
Bot. 4: 98. 1859. TYPE: BRAZIL. RIO DE JANEIRO:
Organ Mts. (Serra dos Órgãos), common on
clayey banks, 3000 feet, Gardner 12 (syntype E!).
Polytrichadelphus semiangulatus (Brid.) Mitt., J. Linn.
Soc., Bot. 12: 609. 1869 ; Pogonatum
semiangulatum Brid., Bryol. Univ. 2: 744. 1827.
662 The Bryologist 113(3): 2010

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).
 Peralta & Yano: Taxonomic treatment of the Polytrichaceae from Brazil 663

Figure 10. Polytrichadelphus pseudopolytrichum. a. Female gametophyte, b. Male gametophyte, c–d. Leaves, e–f. Perichaetial
leaves, g–h. Perigonial 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, with verrucose cuticle, n. Transverse section of the sheathing base,
o. Lateral view of the lamella, p. Epidermal cells of the capsule, with stomata, q. Transverse section of the epidermal cells of the
capsule, s. Calyptra, t. Peristome teeth (Peralta 3618-SP).
664 The Bryologist 113(3): 2010

TYPE: BRAZIL. Paris 1825, Persoon (lectotype B!), (SP); Caldas, Pietrobom-Silva 1978 (MO, SJRP, SP);
designated by Smith (1976). Caratinga, Vital 5858 (NY, SP); Catas Altas, Parque
Polytrichadelphus juniperaceus Müll. Hal., Gen. Musc. Natural do Caraça, Peralta et al. 6818 (SP); Itamonte,
Frond.: 168. 1900. nom. nud. O. Yano & Visnadi 9878 (SP); Lima Duarte, Parque
Estadual do Ibitipoca, O. Yano et al. 21546 (SP).
Description. Plants large-sized, reddish or dark-
PARANÁ: Bocaiúva do Sul, Hatschbach 23456 (MBM;
green, 80–100 mm long. Stems erect, forming tufs.
PACA); Jaguariaı́va, O. Yano et al. 5230, 5232 (SP);
Rhizome prostrate. Rhizoids pale-brown. Leaves
Laranjeiras do Sul, Vital 5786 (NY, SP); Mallet, Wasum
lanceolate, 6–8 mm long, widespreading, quadratic
2814, 2833 (SP; MBM, SP); Tijucas do Sul, O. Yano &
cells, 10 3 10 mm, thick-walled, smooth, sheathing
Shirata 27872 (SP). RIO DE JANEIRO: Angra dos Reis, O.
base conspicuous, plane margins, dentate, not
Yano et al. 23793 (SP); Itatiaia, Ule 1768 (R); Nova
bordered, costa broad 3/4–4/5, percurrent, lamellae
Friburgo, Sehnem 6744 (PACA; SP). RIO GRANDE DO SUL:
occupying costae and laminae, 50–60 lamellae, 6 cells
Bom Jesus, Sehnem 221 (NY, PACA, SP); São Francisco
high, distal cells simple, pyriform, rugose, lamella
de Paula, Sehnem 5344 (PACA; SP). SANTA CATARINA:
cells in lateral view quadratic. Dioicous, perichaetium
Bom Retiro, Schäfer-Verwimp & Verwimp 10491 (SP);
lateral, leaves long, perigonium lateral, leaves short
Negro, Vital 5602 (NY, SP). SÃO PAULO: Apiaı́, Vital
and broad. Sporophyte 1(–2), seta 50–60 mm long.
2302 (SP); Campos do Jordão, Froehlich 17 (S);
Capsule asymmetric, urn cylindric, stomata present
Caraguatatuba, Sucre et al. 7007 (MO, RB); Cunha,
in all urn surface, operculum conic, long-rostrate, 64
Custódio Filho 549 (SP); Iporanga, Joly 1235 (SP);
teeth. Calyptra cucullate, glabrous.
Itapecirica da Serra, Eiten & Godland 6317 (NY, SP);
Distribution. Endemic to the Southeast and
Jambeiro, O. Yano & Yano 22683 (SP); Natividade da
South of Brazil (States of Espı́rito Santo, Minas
Serra, O. Yano et al. 7734 (SP); Piquete, Peralta 3618
Gerais, Paraná, Rio de Janeiro, Rio Grande do Sul,
(SP); Rio Grande da Serra, 7-VII-1901, Schiffner s.n.
Santa Catarina and São Paulo).
(NY); Santo Antonio do Pinhal, O. Yano 1156 (SP);
Habitat. On soil along trails and on banks at the
São Paulo, Schiffner 937 (NY, S); Ubatuba, O. Yano et
margin of submontane and montane forests at the
al. 8109 (SP).
Mata Atlântica and Pampa Biomes; between ca. 600–
2,200 m alt.
Polytrichum Hedw., Sp. Musc. Frond.: 88. 1801.
Discussion. This is the only Brazilian species TYPE: P. commune Hedw.
with perichaetia developed on small branches. Others
Polytrichum angustifolium Mitt., J. Linn. Soc., Bot.
diagnostic features are the red colour of sheathing
12: 622. 1869. TYPE: BRAZIL. Brasilia Tropical,
base, pyriform and verrucose distal cells of lamella
Burchell 3768 (syntypes BM!, E!). Fig. 11
and imbricate and acute leaf apex.
Polytrichum brasiliense Hampe, Vidensk. Meddel.
Some samples are erroneously identified as P.
Dansk Naturhist. Foren. Kjøbenhavn. ser. 3, 10:
magellanicus (Hedw.) Mitt. This species has robust
53. 1872 ; Pogonatum brasiliense (Hampe) A.
habit and the agude leaf apex, and is restricted to the
Jaeger, Ber. Thätigk. St. Gallischen Naturwiss.
southern South America (i.e., below the 40u parallel;
Ges. 1873–74: 266. (Gen. Sp. Musc. 1: 728).
Smith 1976). Some samples are identified as
TYPE: BRAZIL. RIO DE JANEIRO: Rio de Janeiro,
Polytrichastrum formosum (Hedw.) G.L. Smith but
Glaziou 5199 (syntype PC!, B!). syn. nov.
this species has the terminal perichaetia and hairy
Polytrichum angustifolium Schimp., Bull. Herb.
calyptrae, this species have the distal cells of lamella
Boissier 5: 178. 1897. nom. illeg. non Mitten
verrucose in common with Polytrichadelphus
(1869).
pseudopolytrichum (Raddi) G.L. Smith.
Polytrichum alticaule Müll. Hal., Hedwigia (Beibl.)
Selected specimen examined. BRAZIL. ESPı́RITO
38(1): 59. 1899. nom. nud
SANTO: Conceição do Castelo, O. Yano et al. 4949 (SP);
Muniz Freire, O. Yano et al. 4953 (SP). MINAS GERAIS: Description. Plants large-sized, reddish to dark-
Itabira do Campo, Mattos 18 (R); Alto Caparaó, green, 150–800 mm long. Stems erect, forming tufs.
Parque Nacional do Caparaó, Visnadi & Vital 2632 Rhizome prostrate. Rhizoids pale-brown. Leaves
 Peralta & Yano: Taxonomic treatment of the Polytrichaceae from Brazil 665

Figure 11. Polytrichum angustifolium. 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 (Vital 7063-SP).
666 The Bryologist 113(3): 2010
lanceolate, 6–8 mm long, widespreading, quadratic
cells, 10 3 10 mm, 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 Espı́rito Santo, Minas Gerais, Paraná, Rio
de Janeiro, Rio Grande do Sul, Santa Catarina and
São Paulo).
Habitat. On soil, along trails in margins of
montane forests at the Mata Atlântica 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. ESPı́RITO
SANTO: Caparaó, Schäfer-Verwimp & Verwimp 11489
(SP); Santa Teresa, O. Yano et al. 4922 (SP). MINAS
GERAIS: Alto Caparaó, Schäfer-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, Caraça, 1885,
Wainio s.n. (PC); Ouro Preto, II-1890, Ule s.n. (B; BM;
SP); Santa Bárbara, Ule 1317 (R). PARANÁ: 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); Petrópolis, Luetzelburg 15457
(B); Resende, Parque Nacional do Itatiaia,. Eiten &
Eiten 7631 (SP); Teresópolis, Serra dos Órgãos, Brade
12538 (R). RIO GRANDE DO SUL: Cambará do Sul, Serra
da Rocinha, Sehnem 7805 (PACA, ICN, R, RB, SP); Dois
Irmãos, Morro Reuter, Sehnem 13827 (PACA, SP); São
Francisco de Paula, Taimbé, Sehnem 5272 (PACA, SP).
SANTA CATARINA: Bom Retiro, Sehnem 6995 (PACA, SP);
Irineópolis, Reitz & Klein 11568 (SP). SÃO PAULO:
Campos do Jordão, 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); Salesópolis, 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
(syntype BM?), ex descr.
Polytrichum subremotifolium Geh. & Hampe, Flora
64: 377. 1881. TYPE: BRAZIL. SÃO PAULO: Apiaı́,
sterile, V-1879, prope Faxina, Puiggari 634
(syntype H-BR!). syn. nov.
Polytrichum regnellii Broth., Gen. Musc. Frond.: 184.
1900. nom. nud.
Description. Plants medium-sized, dark-green,
5–300 mm long. Stems erect, forming tufs. Rhizome
prostrate. Rhizoids pale-brown. Leaves lanceolate, 6–
7 mm long, widespreading, quadratic cells, 10 3
10 mm, thick-walled, smooth, sheathing base
conspicuous, plane margins, dentate, not bordered,
costa broad 3/4–4/5, percurrent, lamellae occupying
costae and laminae, 30–40 lamellae, 4–5 cells high,
distal cells single, U-shaped, smooth, lamella cells in
lateral view quadratic. Dioicous, perichaetium
terminal, leaves long, perigonium terminal, leaves
short and broad. Sporophyte single, seta 60–80 mm
long. Capsule symmetric, urn cylindric, apophysis
 Peralta & Yano: Taxonomic treatment of the Polytrichaceae from Brazil 667

Figure 12. Polytrichum commune. a. Female gametophyte, b. Male gametophyte, c–e. Leaves, g–i. Perichaetial leaves, j–l.
Perigonial leaves, m. Apical cells of the leaf, n. Margin cells of the leaf, o. Cells of the sheathing base, p. Transverse section of the
apex of the leaf, q–r. Transverse section of the median leaf, s. Transverse section of the sheathing base, t. Lateral view of the lamella,
u. Epidermal cells of the capsule, v. Transverse section of the epidermal cells of the capsule, w. Calyptra, x. Peristome teeth
(Baranov 78-SP).
668 The Bryologist 113(3): 2010
present, stomata present at urn base, operculum
conic, long-rostrate, 64 teeth. Calyptra cucullate,
hairy.
Distribution. Cosmopolitan. Brazil (States of
Amazonas, Bahia, Distrito Federal, Espı́rito Santo,
Goiás, Minas Gerais, Paraná, Rio de Janeiro,
Roraima, Rio Grande do Sul, Santa Catarina and São
Paulo).
Habitat. On soil, along trails and on banks at
the margin of low, submontane, montane forests and
high fields at the Amazonia, Mata Atlântica, 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
G and 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:
São 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:
Brası́lia, Vital 6285 (SP, NY). ESPı́RITO SANTO: Iúna,
Parque Nacional do Caparaó, Vital & Buck 11768 (B,
SP, NY). GOIÁS, Alto Paraı́so de Goiás, Irwin et al. 32845
(MO, NY, S); Cristalina, Irwin et al. 9851 (NY, SP). MINAS
GERAIS: Barão 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). PARANÁ:
Antonina, Hatschbach 14136 (MBM, PACA, UPCB);
Bocaiúva do Sul, Poliquesi & Cordeiro 151 (MBM);
Curitiba, J. Weir 99 (E, NY); Desvio, Ypiranga, 28-V-
1909, P. Dusén 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); Teresópolis,
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: Caracaraı́, Prance et
al. 21441 (NY). SANTA CATARINA: Bom Retiro, Reitz 2431
(NY); Papanduva, Vital 5645 (SP, NY); Ponte Alta, Vital
9415 (SP); Urubici, Schäfer-Verwimp & Verwimp
10493 (SP). SÃO PAULO: Barra do Turvo, O. Yano 2289
(SP); Campo Grande, Schiffner 355 (S, W); Campos do
Jordão, Peralta 191 (SP, SJRP); Cunha, Peralta et al.
3764 (SP); Guarulhos, O. Yano 312 (SP); Ilhabela, O.
Yano 5156 (SP); Moji Guaçu, Visnadi &Vital 729 (SP);
Paraibuna, Vital 7078 (SP, NY); Peruı́be, O. Yano &
Mello 11445 (SP); Rio Grande da Serra, Baranov 78
(SP); São José do Barreiro, Vattimo 3–101 (R); São
Paulo, Schiffner 667 (B); São Simão, Vital 2453 (SP);
Ubatuba, Visnadi &Vital 1037 (SP).
Polytrichum juniperinum Hedw., Sp. Musc. Frond.:
89. 1801. TYPE: SUIçA. locus montanus
subalpestris, delineata exemplo ex Helvetia,
s.col. Fig. 13
Polytrichum reflexifolium Herzog, Arch. Bot. São
Paulo 1(2): 71. 1924. TYPE: BRAZIL. SÃO PAULO:
Tremembé, 560 m alt., 6-II-1921, Soares s.n.
(holotype SP!, isotype MANCH!). syn. nov.
Polytrichadelphus organensis Herzog, Repert. Sp. Nov.
Regni Veg. 21: 31. 1925. TYPE: BRAZIL. RIO DE
JANEIRO: Serra dos Órgãos, P. Luetzelburg 7260
(syntype B!). syn. nov.
Polytrichum aristiflorum Mitt., J. Linn. Soc., Bot. 12:
620. 1869. TYPE: BRAZIL. Weir 11 (holotype NY!,
isotypes E!, MANCH!).
Polytrichum vulgare Hedw., Rodriguésia 54: 266.
1980. nom. nud.
Polytrichum pallidicaule Müll. Hal., Hedwigia (Beibl.)
38(1): 57. 1899. nom. nud.
Polytrichum brachypyxis Müll. Hal., Hedwigia (Beibl.)
38(1): 57. 1899. nom. nud.
Description. Plants large-sized, green to dark-
green, 50–100 mm long. Stems erect, forming tufs.
Rhizome prostrate. Rhizoids pale-brown. Leaves
 Peralta & Yano: Taxonomic treatment of the Polytrichaceae from Brazil 669

Figure 13. Polytrichum juniperinum. a. Female gametophyte, b. Male gametophyte, c–e. Leaves, f–g. Perichaetial leaves, h–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, s. Transverse section of the epidermal cells of the capsule, t. Calyptra, u. Peristome teeth (Yano &
Marcelli 19717-SP).
670 The Bryologist 113(3): 2010
Figure 14. General distribution of Polytrichaceae in Brazil.
lanceolate, 6–7 mm long, widespreading, quadratic
cells, 10 3 30–50 mm, 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, Espı́rito Santo,
Goiás, Minas Gerais, Paraná, Rio de Janeiro,
Roraima, Rio Grande do Sul, Santa Catarina, São
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 Atlântica, 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:
Abaı́ra, Harley & Campos 51119 (SP); Camaçari,
Mattos Silva & Brito s.n. (ALCB); Rio de Contas,
Harley et al. 26272, 26278 (NY, E; E). ESPÍRITO SANTO:
Caparaó, Schäfer-Verwimp & Verwimp 8968 (SP);
Muniz Freire, O. Yano et al. 4957 (SP). GOIÁS: Alto
Paraı́so de Goiás, Hatschbach et al. 36477 (MBM,
PACA). MINAS GERAIS: Itabira do Campo, Ule 1323 (R);
Caparaó, Vital & Buck 11602 (SP, NY); Carangola, Y.
Mexia 4304 (MO, NY); Conceição do Mato Dentro, O.
Yano 513 (SP, NY); Tiradentes, O. Yano et al. 21871
(SP). PARANÁ: Bocaiúva do Sul, O. Yano & F.C. Silva
11303 (SP); General Carneiro, O. Yano & Pirani 6505
(SP); Quedas do Iguaçu, O. Yano et al. 5413 (SP). RIO
 Peralta & Yano: Taxonomic treatment of the Polytrichaceae from Brazil
DE JANEIRO: Angra dos Reis, O. Yano et al. 23731 (SP);
Itatiaia, Ule 1229 (R); Nova Friburgo, VI-1925, Leão
s.n. (MO, RB, S); Parati, Vital 7246 (MO, NY, SP). RIO
GRANDE DO SUL: Caçapava do Sul, O. Yano & Pirani
5858 (SP); Cambará 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);
Florianópolis, 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); Palhoça,
Reitz & Klein 2513 (MBM, NY, PACA, B). SÃO PAULO:
Amparo, O. Yano & Santos 6035 (SP); Apiaı́, 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
Jordão, O. Yano & Marcelli 19717 (SP); Ibiúna, 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); São Bernardo do Campo, Vital 5379 (MO, NY, SP);
São Paulo, VIII-1901, V. Schiffner s.n. (B); São
Sebastião, O. Yano et al. 7931 (SP); Serra Negra, O.
Yano et al. 18496 (SP). TOCANTINS: Paraı́so de
Tocantins, O. Yano & Kida 26753 (SP).
LIST OF DOUBTFUL NAMES
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
(syntype B?).
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 (syntype BM?, G?).
Polytrichum juniperinum Hedw. var. paulense (Geh. &
Hampe) Herzog, Arch. Bot. São Paulo 1(2): 71.
1924 ; Polytrichum paulense Geh. & Hampe,
Flora 64: 376. 1881. TYPE: BRAZIL. São Paulo:
671
Apeai (Apiaı́), prope Faxina, sterile, V-1879,
Puiggari 285, 908 (syntype BM?, B?).
Polytrichum petersianum Müll. Hal., Linnaea 38: 575.
1874. TYPE: BRAZIL. Nova Friburgo, in solo
argiloso aprico nudo, 20-VII-1871, Peters s.n.
(syntypes HBG?, 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 Müll. Hal. ex Herzog,
Biblioth. Bot. 87: 103. 1916. nom. nud.
Pogonatum serricolum Müll. Hal., Gen. Musc. Frond.:
176. 1900. nom. nud.
Polytrichum caudatum Müll. Hal., Gen. Musc.
Frond.: 184. 1900. nom. nud.
Polytrichum lanipes Müll. Hal., Gen. Musc. Frond.:
185. 1900. nom. nud.
Polytrichum micropysix Müll. Hal., Gen. Musc.
Frond.: 181. 1901. nom. nud.
Polytrichum pallidipes Müll. Hal., Gen. Musc. Frond.:
184. 1900. nom. nud.
Polytrichum pycnocarpum Müll. Hal., Gen. Musc.
Frond.: 182. 1900. nom. nud.
Polytrichum uleanum Müll. Hal., Gen. Musc. Frond.:
184. 1900. nom. nud.
LITERATURE CITED
Churchill, S. P. & E. Linares C. 1995. Prodromus Bryologiae
Novo-Granatensis: Introducción 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 famı́lia Polytrichaceae no Rio Grande do
Sul, Brasil. Dissertação de Mestrado, Universidade Federal
do Rio Grande do Sul, Porto Alegre.
———. 1984. A famı́lia Polytrichaceae no Rio Grande do Sul,
Brasil. I Taxonomia. Iheringia, Série Botânica 32: 77–89.
Goffinet, B., W. R. Buck & A. J. Shaw. 2009. Morphology,
anatomy, and classification of the Bryophyta. Pages 56–138.
In: B. Goffinet & A. J. Shaw (eds.), Bryophyte Biology.
Cambridge University Press, Cambridge.
672 The Bryologist 113(3): 2010
Gradstein, S. R., S. P. Churchill & N. Salazar-Allen (eds.). 2001,
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ms received October 28, 2009; accepted May 28, 2010.