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Anais da Academia Brasileira de Ciências (2019) 91(1): e20180313
(Annals of the Brazilian Academy of Sciences)
Printed version ISSN 0001-3765 / Online version ISSN 1678-2690
http://dx.doi.org/10.1590/0001-3765201920180313
www.scielo.br/aabc | www.fb.com/aabcjournal
An Acad Bras Cienc (2019) 91(1)
BIOLOGICAL SCIENCES
A new species and new records of Leptogium (Ach.) Gray (Collemataceae, Peltigerales)
from Rio Grande do Sul State with an identication key for the genus
MARCOS J. KITAURA1, NATÁLIA M. KOCH1, FABIANE LUCHETA2, MÁRCIA
I. KÄFFER2, JAIRO L. SCHMITT2, JULIANA PEDROSO3, SUZANA A.
MARTINS4, ANDRESSA S. RODRIGUES1 and LUCIANA S. CANÊZ1
1Laboratório de Evolução e Biologia Evolutiva, Programa de Pós-Graduação em Biologia Vegetal, Universidade
Federal de Mato Grosso do Sul, Avenida Costa e Silva, s/n, 79070-900 Campo Grande, MS, Brazil
2Laboratório de Botânica, Programa de Pós-Graduação em Qualidade Ambiental, Universidade
Feevale, Rodovia ERS 239, 2755, 93525-075 Novo Hamburgo, RS, Brazil
3Instituto de Ciências Biológicas, Universidade Federal do Rio Grande, Avenida Itália, Km 8, 96201-900 Rio Grande, RS, Brazil
4Museu de Ciências Naturais, Fundação Zoobotânica do Rio Grande do Sul, Rua
Dr. Salvador França, 1427, 90690-000 Porto Alegre, RS, Brazil
Manuscript received on March 27, 2018; accepted for publication on July 24, 2018
How to cite: KITAURA MJ, KOCH NM, LUCHETA F, KÄFFER MI, SCHMITT JL, PEDROSO J, MARTINS SA,
RODRIGUES AS AND CANÊZ LS. 2019. A new species and new records of Leptogium (Ach.) Gray (Collemataceae,
DOI 10.1590/0001-3765201920180313.
Abstract: Specimens of Leptogium collected in ten localities from the State of Rio Grande do Sul were
studied. Sixteen species were found of 28 records mentioned to the state, which represents around 57%
Grande do Sul. Leptogium exaratum is proposed as a new
species.
state. The lectotype of L. javanicum is reported here and detailed descriptions are provided to the examined
this genus in Brazil.
Key words:
Correspondence to: Marcos Junji Kitaura
ORCid: http://orcid.org/0000-0002-2175-3533
INTRODUCTION
Leptogium (Ach.) Gray is a genus of jelly lichens
that has been studied since Micheli (1729) and
Dillenius (1741). Nowadays, the number of species
180 species (Otálora et al. 2014), but a taxonomical
review of the genus is still necessary. Leptogium
by Eschweiler (1833), through of the species: L.
bullatum (Ach.) Nyl., L. rivulare (Ach.) Mont. and
L. tremelloides (L.f.) Gray.
Leptogium puiggarii Müll. Arg. and L.
dimorphum Müll. Arg. were the first species
described from Brazil and they were reported for
the municipality of Apiaí, São Paulo in Müller
Argoviensis (1879, 1881), respectively.
MARCOS J. KITAURA et al. Leptogium FROM RIO GRANDE DO SUL STATE
An Acad Bras Cienc (2019) 91(1) e20180313 2 | 26
Leptogium species
reported was L. phyllocarpum
Almost 20 years after, Malme (1924) mentioned
to the state L. cyanescens var. austroamericanum
Malme, L. marginellum (Sw.) Mont., L.
megapotamicum Malme, L. moluccanum (Pers.)
Vain., L. phyllocarpum (Pers.) Nyl., L. pichneum
(Ach.) Malme, L. puiggarii Arg., L tremelloides (L.
f.) Gray, and L. ulvaceum (Pers.) Vain.
During the decades 1970-2000, a series of
contributions was published about lichens from
Rio Grande do Sul (Fleig 1988, 1995, Fleig and
Grüninger 2000a, 2000b, Mazzitelli et al. 1999,
Osorio 1981, 1985, Osorio and Fleig 1982, 1984a,
b, Fleig 1985, 1987, 1988, 1989, 1990a, b, 1991,
1994, Osorio et al. 1981, 1983, 1997, Zanette et al.
1981); and posteriorly, all information published
with lichens have been developed by the team of
Fundação Zoobotânica do Rio Grande do Sul, which
2016) and new lichen species have been described
each year (Kitaura and Marcelli 2012, Aptroot et al.
et al. 2014, Kitaura et al. 2014,
Leptogium
from Rio Grande do Sul State mainly through
the HAS and HURG herbaria materials, updating
species, include two new citations and the first
Leptogium for Rio
Grande do Sul.
MATERIALS AND METHODS
The specimens were collected during lichenological
Grande do Sul: Campo Bom, Caraá, Maquiné,
Parobé, Pelotas, Rio Grande, Rolante, Santo
Antônio da Patrulha, São Leopoldo and Taquara.
Forest fragments, urban and rural areas
constitute the main landscape of these
municipalities, with average altitude varying from
(Alvares et al. 2013). The original vegetation in
most of the municipalities is characterized by
semideciduous forests (Anschau 2016) - Campo
Bom, Taquara, São Leopoldo, Parobé, Rolante,
and Santo Antônio da Patrulha, while some
municipalities are composed of Atlantic rainforests,
with areas of semideciduous and mixed Araucaria
forests (Sevegnani and Baptista 1996) - Caraá and
Maquiné; and others are mainly represented by
restinga forests and grasslands (Burger and Ramos
2007) - Rio Grande and Pelotas.
Most of the collected specimens are deposited
in HAS herbarium and some of them in the HURG
and the CGMS herbaria. All studied material was
described according to the protocol of Kitaura
(2012) and Kitaura et al. (2015).
The term duplicate of lectotype was used
to design a true piece of the lectotype specimen,
according to Kitaura et al. (2013).
Neither the type material nor the material
reported to Rio Grande do Sul of Leptogium
coralloideum (Meyen and Flot.) Vain., L .
diaphanum (Sw.) Mont., L. moluccanum, L.
pichneum, L. puiggarii and L. ulvaceum were
revised during the current study, but the original
descriptions (protologues) were used to prepare the
5590 and with a Canon RebelT3i coupled on an
Olympus Sz stereomicroscopy and on an Olympus
CX22LED microscopy.
RESULTS
All Leptogium species names previously reported
were grouped in a list, along with the other species
found by the authors. The type specimens from
MARCOS J. KITAURA et al. Leptogium FROM RIO GRANDE DO SUL STATE
An Acad Bras Cienc (2019) 91(1) e20180313 3 | 26
Known distribution to RS: It is mentioned
Santa Catarina and São Paulo States.
Specimens examined: Brazil, Rio Grande
do Sul: Rio Grande municipality, 01 Apr. 2016,
J. Pedroso 245 (HURG); Ibid., 07 Apr. 2016, J.
Pedroso 392 (HURG); Ibid., 08 Apr. 2016, J.
Pedroso 249 pr. min. p. (HURG); Ibid., Taquara
municipality, Fazenda dos Búfalos, Bacia
Notes: Leptogium atlanticum is characterized
by the upper surface with irregular to longitudinal
rounded on the lamina and margin of the thallus
(Figure 1a). Helicoidal columnar hyphae were
observed in the material from Rio Grande do Sul
as in the specimens studied by Kitaura et al. (2015).
This species is common in forests near to the
coast in São Paulo State (Kitaura et al. 2015), but it
is rare in Rio Grande do Sul. Leptogium atlanticum
was commonly confused with L. denticulatum,
but it does not have lobules on the thallus and the
apothecia are ornamented by denticules (Kitaura et
al. 2015).
2. Leptogium austroamericanum (Malme) C.
Leptogium cyanescens var. austroamericanum
Grande do Sul State, Santo Ângelo prope Cachoeira
[Agudo municipality], 3 Sep. 1819, G.O.A. Malme
950 B (lecto-: S!, designated in Benatti et al. 2013),
duplicates of lecto-: S! Malme 950 A e D.
Thallus foliose, 2.0–10.0 cm broad, grayish
to fluorescent light, opaque, matt, gray with
yellowish regions under stereomicroscope. Lobes
2.0–5.0 mm wide, agglomerated, attached in
apices rounded to irregular, ascending, smooth
to ornamented; lateral margins ascending and
more than 70% of the found species were revised.
Most of the mentioned type species was collected
originally in tropical regions and all names were
considered as good species, except for Leptogium
tremelloides.
One new species (L. exaratum) and three new
records including L. atlanticum, L. milligranum
and L. vesiculosum to Rio Grande do Sul State were
found from revised ca. 120 specimens deposited in
HAS and HURG herbaria.
1. Leptogium atlanticum Marcelli and
Kitaura, The Bryologist, 118 (1): 12. 2015. Type:
Brazil, São Paulo, municipality of Peruíbe, Reserva
Ecológica Juréia-Itatins. Núcleo Guarauzinho,
1993, M.P. Marcelli and O. Yano 23710 (holo-:
SP!, iso-: COLO!, H!).
Thallus foliose, ca. 2 cm broad, gray to
under stereomicroscope. Lobes 1.5–2.5 mm wide,
agglomerated, adpressed, adnate, upper surface
rugulose under 20× magnification; apices
usually rounded, plane to ascending, smooth to
ornamented; lateral margins irregular, ornamented,
crispate; lower side bluish to yellowish, irregular
Lobules usually rounded, laminal and marginal,
granular to claviform when young, simple to
branched, 0.05–0.625 × 0.05–0.75 mm, firm,
erect, concolorous with the thallus, grouped,
frequent to abundant. Thallus attached by hapters,
frequent, homogeneous. Rhizinae and hairs absent.
Apothecia not observed. Anatomy: Thallus ca. 90
diam. Medulla with helicoidal columnar hyphae,
5–7 cells, frequent; cyanobacteria blue to green,
frequent, elliptic cells, 3.0 × 2.5 µm diam., ca.
colorless. Pycnidia not observed.
MARCOS J. KITAURA et al. Leptogium FROM RIO GRANDE DO SUL STATE
An Acad Bras Cienc (2019) 91(1) e20180313 4 | 26
Figure 1 - Leptogium species (a) ornamented thallus of Leptogium atlanticum. (bL. austroamericanum.
(c) pedicellate apothecia of L. azureum. (d) submarginal apothecia of L. chloromelum. Bars = 2.0 mm.
MARCOS J. KITAURA et al. Leptogium FROM RIO GRANDE DO SUL STATE
An Acad Bras Cienc (2019) 91(1) e20180313 5 | 26
involute, usually ornamented, crispate; lower
side grayish to yellowish, rugulose under 20×
appearance, simple to irregular branched, laminal
caducous, erect, grouped, abundant. Lobules
rounded, usually simple, laminal and marginal,
0.3 × 0.5 mm, scarce, grouped. Thallus attached
by hapters, frequent, homogeneous. Rhizinae and
hairs absent. Apothecia not observed. Anatomy:
cells, 5.0 µm diam. Medulla with columnar hyphae
sinuous, ca. 5 cells, frequent; cyanobacteria green,
5.0 µm diam.; gelatinous matrix frequent, colorless
and yellowish near the upper cortex. Pycnidia
absent.
Known distribution to RS: See Spielmann
et al.
Specimens examined: Brazil, Rio Grande do
Sul: Rio Grande municipality, Restinga APP6, 01
Apr. 2016. J. Pedroso 221, 224, 227, 230, 232, 243,
244, 247, 249 252 (HURG); Ibid., 07 Apr. 2016
J. Pedroso 257, 259, 260, 261 (HURG); Ibid., 19.
Apr. 2016, J. Pedroso 391 (HURG); Ibid., 22 Apr.
2016, J. Pedroso 411 (HURG); Ibid., 05 May 2016,
J. Pedroso 513 (HURG); Ibid., Santo Antônio da
Patrulha municipality, Sítio Estrada Arroio Grande,
Notes: Leptogium austroamericanum is
characterized by the upper surface irregularly
isidia usually cylindrical with withered appearance.
This species is common in Rio Grande do Sul
State, but the specimens were collected sterile.
3. Leptogium azureum (Sw.) Mont., Historie
Lichen
azureus Sw., Lichenographiae Suecicae Prodromus:
137. 1798. Type: Jamaica, on montain, leg. O.
Swartz s/n (lecto-: H-ACH!, duplicate of lecto-:
BM!, designated by Jørgensen and James (1983)).
Thallus foliose, ca. 6 cm broad, bluish gray
under fluorescent light, opaque, matt, bluish
under stereomicroscope. Lobes up to 8.5 mm
wide, agglomerated, attached in points, adnate,
lateral margins smooth, irregular, undulated;
lower side smooth, bluish. Isidia and lobules
absent. Thallus attached by hapters, frequent,
homogeneous distribution. Rhizinae and hairs
absent. Apothecia up to 3.0 mm diam., submarginal
to laminal, pedicellate, disc reddish, usually plane;
margin cream, smooth; amphithecia beige, smooth;
corona absent; pedicel 0.3–1.5 mm high, originated
by the thallus, smooth. Anatomy: Thallus 50–60
µm diam. Medulla with columnar hyphae straight,
2–3 cells, frequent; cyanobacteria blue, ca. 10
gelatinous matrix scarce, colorless. Apothecia
with hymenium 150–170 µm high; subhymenium
irregular cells; parahymenium tissue continuous
with hypothecium, prosoplectenchymatous;
proper exciple absent; thalline exciple cortex with
paraplectenchymatous cells, 45 µm (4 cells) at the
apices, 25–40 µm (2–4 cells) at the mid-height,
paraplectenchymatous tissue 100–115 µm (8–10
7.5 µm, apices acute, submuriform to muriform.
Pycnidia not observed.
Known distribution to RS: See Spielmann
Alegre (Käffer et al. 2011) and Santana do
Specimens examined: BRAZIL:
Caraá, Localidade Fraga, APA Municipal.
03 Apr. 2008. S.A. Martins 2140 pr. p. (HAS -
MARCOS J. KITAURA et al. Leptogium FROM RIO GRANDE DO SUL STATE
An Acad Bras Cienc (2019) 91(1) e20180313 6 | 26
sterile); Ibid., 19 Jun. 2008. S.A. Martins 2210;
Ibid., Rio Grande municipality, Restinga APP6, 05
May 2016. J. Pedroso 528 (HURG).
Notes: Leptogium azureum is characterized
by the smooth thallus and apothecia with pedicel
originated from the thallus, with 0.3–1.5 mm high
(Figure 1c).
This species has wide distribution (Aragón et
al. 2004), but it is a complex of species and further
studies are necessary.
4. Leptogium brebissonii Mont., Histoire
Naturelle des Iles Canaries 3(2): 130. 1840. Type:
Jørgensen (1995).
Description: See Galloway and Jørgensen
(1995) and Kitaura (2012).
Known distribution to RS: See Spielmann
(2006).
Notes: Leptogium brebissonii has ridged
upper surface, isidia granular to short cylindrical,
7–10 cells (Galloway and Jørgensen 1995, M.J.
Kitaura, unpublished data).
5. Leptogium chloromelum (Sw.) Nyl., Mém
Lichen
chloromelus Sw., Fl Ind Occid 3: 1892. 1806. Type:
Jamaica, leg. Swartz s/n (holo-: H-NYL 41252!
(?)), Mexico, Borrejo, leg. Fr. Müller s /n (epi-:
H-NYL 41247!).
Thallus foliose, 2.0–4.0 cm broad, grayish
yellowish under stereomicroscope. Lobes 1.5–4.0
mm wide, agglomerated, adnate to attached in
apices rounded, ascending, irregular to crispate,
smooth; lateral margins ascending, smooth,
crispate; lower side yellowish gray, ridged under
20× magnification. Isidia and lobules absent.
Apothecia up to 3.0 mm diam., submarginal,
subpedicellate, disc reddish, usually plane;
margin concolorous with the thallus, circular
ridges, smooth; amphithecia concolorous with the
thallus, irregular ridges, without ornamentation;
corona absent; pedicel very short, originated by
one layer of isodiametric cells of cortical layers,
2.5–5.0 × 5.0–7.5 µm. Medulla with columnar
hyphae not observed; cyanobacteria green, 12–20
spherical to elliptical cells per filaments, 2.5–
5.0 × 5.0 µm cell; gelatinous matrix colorless,
abundant. Apothecia with hymenium 125 µm high;
subhymenium 25–37.5 µm, colorless; hypothecium
yellowish; proper exciple paraplectenchymatous,
parahymenial tissue continuous with proper
exciple, paraplectenchymatous, 50 µm (7 cells) at
the apices; thalline exciple cortex with one layer of
isodiametric cells, 5.0–7.5 µm diam. Pycnidia not
observed.
Known distribution to RS: See Spielmann
(2006).
Specimens examined: BRAZIL:
Caraá municipality, Área de Proteção
Ambiental, Nascente do Rio dos Sinos, BHRS,
Ibid., Rio Grande municipality, Restinga APP6, 01
Apr. 2016, J. Pedroso 255 (HURG); Ibid., 14 Apr.
2016, J. Pedroso 325, 327, 329 (HURG); Ibid.,
19 Apr. 2016, J. Pedroso 374, 400, 401 (HURG);
Ibid., 20 Apr. 2016, J. Pedroso 436, 470 (HURG);
Ibid., 22 Apr. 2016. J. Pedroso 408 pr. min. p., 467
(HURG); Ibid., 05 May 2016, J. Pedroso 505, 506,
507 (HURG); Ibid., 27 Sep. 2016, J. Pedroso 491
(HURG); Ibid., São Leopoldo municipality, ecology
base, BHRS, riparian forest, area 1, 29° 45’50.6”S,
1029 (HAS); Ibid., Pelotas municipality, Dunas das
Acácias, corticícola, árvore na beira da estrada de
acesso as dunas, 03 Apr. 2017, A.S. Rodrigues 150,
152.
MARCOS J. KITAURA et al. Leptogium FROM RIO GRANDE DO SUL STATE
An Acad Bras Cienc (2019) 91(1) e20180313 7 | 26
Notes: Leptogium chloromelum is characterized
by ridged thallus, and apothecia submarginal and
subpedicellate (Figure 1d).
It is frequently found in the Brazilian savanna
(cerrado) in São Paulo and Mato Grosso do Sul
States.
6. Leptogium cochleatum
and P. James, Lichenologist, 15(2): 113. 1983.
Lichen cochleatum
LINN-SM, designed by Jørgensen and James
(1983)).
Description: See Aragón et al. (2005).
Known distribution to RS: See Spielmann
et al. 2015).
Notes: Leptogium cochleatum is characterized
James 1983).
The apothecia have euparaplectenchymatous
proper margin (Jørgensen and James 1983) that
gives a yellowish color to the amphithecia.
The image of Leptogium cochleatum published
by Jorgensen and James (1983) shows one specimen
L. azureum that is constituted by pedicellate
apothecia. Aragón et al. (2005), however, described
L. cochleatum as having sessile apothecia, but it is
indeed subpedicellate, as shown by Jørgensen and
James (1983).
The type is part of Linnaeus collection from
LINN herbarium, and the material must be studied
on the museum.
7. Leptogium coralloideum (Meyen and Flot.)
Vain., Ann Acad Sci Fenn, Ser A, 6(7): 110. 1915.
Leptogium diaphanum f. coralloideum Meyen
and Flot., Nova Acta Acad Caes Leop - Carol Germ
Nat Cur 19: 226. 1843. Type: not traced (Jørgensen
(1994), Galloway and Jørgensen (1995)).
Description: See Verdon (1992).
Distribution known to RS: See Spielmann
(2006).
Notes: Leptogium coralloideum is
surface. The apothecia have paraplectenchymatous
subhymenial tissue with various layers and thalline
exciple cortex constituted by one layer of cells
(Verdon 1992).
8. Leptogium corticola
Lea, Catalogue of Plants Native and Naturalized
collected in the Vicinity of Cincinnati: 47. 1847.
≡Collema corticola Taylor, London J Bot 5: 195.
1847. Type: U.S.A., Ohio, Leg. T.G. Lea s/n (holo-:
US, iso-: BM!).
Description: See Kitaura (2012).
Known distribution to RS: Santana do
Notes: Leptogium corticola is characterized
amphithecia concolorous with the thallus due
to cortex of thalline exciple with 15–25 µm
they are also constituted by proper exciple of
colloplectenchymatous cells (M.J. Kitaura,
unpublished data).
9. Leptogium cyanescens (Rabenh.) Körb.,
Systema Lichenum Germaniae: 420. 1855.
Collema cyanescens Rabenh. Deutsch. Krypt.
Fl.: 50, 1845. Type: Switzerland, Ticino, Schaerer
- Lich. Helv. N. 409 (lecto-: BM!, designed by
Jørgensen and James (1983)).
Thallus foliose, 3.0–7.0 cm broad, bluish gray
bluish under stereomicroscope. Lobes 2.0–6.0 mm
wide, overlapping, adnate, adpressed, upper surface
plane, smooth; lateral margins plane to ascending,
undulated and sinuous, smooth to ornamented;
lower side bluish gray, smooth to slightly ridged
under 20× magnification. Isidia cylindrical,
withered appearance, 0.05–0.2 × 0.025–0.1 mm,
MARCOS J. KITAURA et al. Leptogium FROM RIO GRANDE DO SUL STATE
An Acad Bras Cienc (2019) 91(1) e20180313 8 | 26
laminal. Lobules rounded, 0.15–0.4 mm diam.,
marginal. Thallus attached by hapters, frequent,
homogeneous. Rhizinae and hairs absent. Apothecia
not observed. Anatomy: Thallus 35.0–50.0 µm
Medulla with columnar hyphae usually straight,
2–3 cells, frequent; cyanobacteria yellow to blue,
frequent, spherical cells 5.0 µm diam., number of
scarce, hyaline. Pycnidia not observed.
Known distribution to RS: See Spielmann
et al. 2015).
Specimens examined: Brazil, Rio Grande
do Sul: Campo Bom municipality, Chácara
dos Tachettos, BHRS, riparian forest, area 2,
Ibid., Caraá municipality,
04 Jun. 2013, M. Käffer 985; Ibid., Localidade
Fraga, APA municipal, 19 Nov. 2008, S.A. Martins
2216 (HAS); Ibid., Área de Proteção Ambiental,
Nascente do Rio dos Sinos, BHRS, riparian forest,
Ibid., 01
Apr. 2016, J. Pedroso 216, 217, 218 (HURG); Ibid.,
07 Apr. 2016, J. Pedroso 278 (HURG); Ibid., 08
Apr. 2016, J. Pedroso 286 (HURG); Ibid., 14 Apr.
2016, J. Pedroso 311, 324 (HURG); Ibid., 22 Apr.
2016, J. Pedroso 439, 463 (HURG); Ibid., 27 Apr.
2016, J. Pedroso 478, 479 (HURG); Ibid., Parobé
2015, F. Lucheta 336 (HAS); Ibid., 29°37’48.5”S,
(HAS); Ibid.
Jul. 2015, F. Lucheta 339 (HAS); Ibid., Rolante
2016, F. Lucheta 342 (HAS); Ibid., F. Lucheta
343; Ibid., Santo Antônio da Patrulha municipality,
Fazenda da Marca, BHRS, riparian forest, area
Ibid., area
Ibid., São Leopoldo
municipality, Ecology base, BHRS, riparian forest,
Ibid., Taquara
municipality, Fazenda dos Búfalos, BHRS, riparian
Ibid.,
Pelotas municipality, mata do Totó, mata ciliar, 26
Mar. 2017. A.S. Rodrigues 098.
Notes: Leptogium cyanescens is characterized
by the upper surface smooth to slightly ridged under
20× magnification (Figure 2a), and cylindrical
isidia with withered appearance.
Leptogium cyanescens differs from L.
austroamericanum by the type of upper surface.
The upper surface of L. cyanescens is usually
smooth whereas the upper surface is irregularly
L. austroamericanum. Both species
have cylindrical isidia with withered appearance.
10. Leptogium denticulatum Nyl., Ann
Sci Nat, Bot Biol Vég 7: 302. 1867. Type: Nova
Granada [Colombia], San Jil, 1300 m. alt., 1863, A.
Lindig sub no. 48 (holo-: H-NYL 41427!).
Description: See Kitaura et al. (2015).
Known distribution to RS: Porto Alegre
Notes: Leptogium denticulatum has smooth
denticules restrict only to the margin of apothecia.
11. Leptogium diaphanum (Sw.) Mont., Ann
Sci Nat, Bot Biol VéLichen
diaphanus Sw., Nov Gen Sp [HBK]: 147. 1788.
Type: Jamaica, India Occidentalis, in montobus
altissinus, inter musco. (PC (I.P.R. Cunha,
unpublished data)).
Thallus foliose, 3–7 cm broad, bluish to gray
to fluorescent light, opaque, matt, bluish under
stereomicroscope. Lobes 2.5–4.0 mm wide,
MARCOS J. KITAURA et al. Leptogium FROM RIO GRANDE DO SUL STATE
An Acad Bras Cienc (2019) 91(1) e20180313 9 | 26
Figure 2 - Leptogium species (a) thallus of L. cyanescens. (b) thallus of L. diaphanum. (c) thallus of L. exaratum. (d)
transversal section of the thallus of L. exaratum
MARCOS J. KITAURA et al. Leptogium FROM RIO GRANDE DO SUL STATE
An Acad Bras Cienc (2019) 91(1) e20180313 10 | 26
overlapping, adpressed, adnate, upper surface
plane to ascending, smooth to ornamented;
lateral margins ascending, undulate and sinuous,
ornamented; lower side yellowish gray to bluish,
Isidia absent. Lobules elongated, simple to
erect, concolorous with the thallus to yellowish,
laminal and marginal. Thallus attached by hapters,
abundant, homogeneous. Rhizinae and hairs absent.
Apothecia not observed. Anatomy: Thallus 25–35
columnar hyphae straight to slightly inclined, 2
cells, scarce; cyanobacteria green, spherical cells
5.0 µm diam., cells per filament undetermined;
gelatinous matrix scarce, hyaline. Pycnidia not
observed.
Known distribution to RS: Santana do
Specimens examined: Brazil, Rio Grande
do Sul: Caraá Municipality, APA do Caraá, 03
Dec. 2008, S.A. Martins 2258 (HAS); Ibid., 12
Ibid., 04 Jun.
p. min. pr. (HAS); Ibid., Rio
Grande municipality, Restinga APP6, 31 Mar.
2016, J. Pedroso 203 (HURG); Ibid., 07 Apr. 2016,
J. Pedroso 273, 274, 279 (HURG); Ibid., Santo
Antônio da Patrulha municipality, BHRS, riparian
Ibid.,
Caraá municipality, 29º 47’4”S, 50º25’38”, 15 Nov.
2014, N.M. Koch 927 (HAS); Ibid., N.M. Koch
925 (HAS). Ibid., Fraga locality, APA Municipal.
19 Jun. 2008. S.A. Martins 2211 (HAS); Ibid.,
Área de Proteção Ambiental, Nascente do Rio dos
Sinos, BHRS, riparian forest, area 7, 29º42’05.8”S,
1035 a (HAS); Ibid., Santo Antônio da Patulha
municipality, Fazenda Marca, BHRS, riparian
Notes: Leptogium diaphanum is characterized
surface with minutely irregular ridges under 20×
magnification, and elongated lobules, simple to
irregularly branched (Figure 2b).
The lobules are flatted and differ from the
cylindrical isidia of L. austroamericanum and L.
cyanescens.
Leptogium diaphanum was mentioned to São
Paulo and Paraná States (I.P.R. Cunha, unpublished
data), but the new citation was not published.
The type is probably deposited in PC
with its curator.
12. Leptogium exaratum M.J. Kitaura and F.
Lucheta, sp. nov.
Similar to L. granulans, but apothecia
ornamented with isidia usually cylindrical, that can
be simple to irregular branched. Type: Brazil, Rio
Grande do Sul, Parobé municipality, 29°37’48.5”S,
and iso-: CGMS).
Thallus foliose, ca. 7 cm broad, muscicolous,
to bluish gray under stereomicroscope. Lobes 2.0–
4.5 mm, agglomerated, attached in points, loosely
apices rounded, ascending to revolute, smooth to
ornamented; lateral margins ascending, sinuous
and undulated, ornamented; lower side brownish,
0.1 mm, usually cylindrical, simple to irregular
grouped, abundant, on the laminal, marginal,
pedicel and apothecia. Lobules absent. Thallus
attached by hapters, frequent, homogeneous.
Apothecia up to 1.5 mm diam., pedicellate, disc
MARCOS J. KITAURA et al. Leptogium FROM RIO GRANDE DO SUL STATE
An Acad Bras Cienc (2019) 91(1) e20180313 11 | 26
concave, reddish; margin smooth, yellowish,
isidiate or not ornamented; amphithecia constituted
by thallus, concolorous with the thallus, which is
inserted on the lateral of amphithecia, ornamented
by isidia; corona absent; pedicel ca. 1.0 mm high,
inflated, longitudinal ridges, isidiate. Anatomy:
cells, 5.0 µm diam. Medulla with columnar
hyphae inclined, ca. 5 cells, scarce, usually near
determined, spherical, ca. 2.5–5.0 µm diam.,
greenish; gelatinous matrix scarce. Apothecia with
colorless, irregular cells prosoplectenchymatous;
parahymenium tissue continuous with the
hypothecium, elongated to paraplectenchymatous
cells, ca. 50 µm at the base and the apices; proper
exciple absent; thalline exciple covered by thallus;
basal paraplectenchymatous tissue present, cortex
with paraplectenchymatous cells, 25 µm (4 cells)
mid-height and base. Ascospores rare, usually
immature, fusiform, 15–20 × 5.0–7.5 µm, apices
acute, submuriform. Pycnidia not observed.
Known distribution to RS: Locality type.
Notes: Leptogium exaratum (Figure 2c, d and
a) is characterized by the presence of apothecia
ridges. The thallus, pedicel and apothecia are
usually covered by cylindrical isidia (Figure 2c).
Leptogium granulans as well as L. exaratum
are isidiate species, but L. granulans have isidia
restrict to the margin and lamina of the thallus,
whereas L. exaratum have isidia on the thallus,
pedicel and apothecia. Furthermore, the form of
the isidia is distinct between L. granulans and L.
exaratum. The isidia of L. granulans are granular,
moniliform (piled up) to irregular branched (M.J.
Kitaura, unpublished data), whereas the isidia of L.
exaratum is usually cylindrical, simple to irregular
branched.
Leptogium javanicumL.
exaratum and L. granulans, but L. granulans and L.
exaratum are isidiate species whereas L. javanicum
does not have isidia.
The epithet exaratum, from Latin, refers to
on the pedicel of L. exaratum, L. granulans and L.
javanicum.
13. Leptogium hibernicum M.E. Mitch. ex
P.M. Jørg., Herzogia 2: 462. 1973. Type: Ireland,
Kerry, Killarney, Torc cascade, on mossy trees, 4
Aug.1933, G. Degelius s/n (holo-: UPS!).
Thallus up to 10 cm broad, bluish gray under
gray to bluish gray under the stereomicroscope.
Lobes 2–7 (–9) mm wide, overlapping, adpressed,
adnate; upper surface smooth with longitudinal
apices rotund, involute, smooth; lateral margin
usually smooth, plane to involute, straight to
usually cylindrical, ca. 0.05–0.40 × 0.05 mm,
whitish or concolorous with the thallus base and
proximal part. Lobules absent. Thallus attached by
hairs these are unbranched, simple to agglutinated,
up to 50 µm (ca. 10 spherical cells) long, beige,
on lower cortex, abundant to dense. Hapters and
rhizinae absent. Apothecia absent. Anatomy:
cortex 2.5 × 5.0 µm, one layer, quadratic cells of
lower cortex, 5.0 × 5.0 µm, one layer. Medulla with
long hyphae that connect the cortices, branched,
columnar hyphae absent; cyanobacteria greenish,
frequent to scarce, filament with 10–20 elliptic
cells, 5.0 × 2.5 µm; gelatinous matrix abundant,
colorless. Pycnidia not observed.
MARCOS J. KITAURA et al. Leptogium FROM RIO GRANDE DO SUL STATE
An Acad Bras Cienc (2019) 91(1) e20180313 12 | 26
Known distribution to RS: See Spielmann
(2006).
Specimens examined: Brazil, Rio Grande
do Sul: Caraá municipality, Fraga locality, APA
Municipal, 19 Nov. 2008, S.A. Martins 2206
(HAS); Ibid., 03 Jul. 2008, S.A. Martins 2142
(HAS).
Notes: Leptogium hibernicum (Figure 3b) is
isidia and hairs constituted by spherical cells.
Specimens from Rio Grande do Sul are
attached mainly by agglutinated hyphae (rhizine-
type specimen (Degelius s/n) is attached only by
hairs on the substrate. The rhizinae are constituted
by cylindrical hyphae, while the hairs are composed
species constituted by spherical cells, with rhizinae.
hairs constituted by cylindrical and spherical cells,
L. velutinum (Kitaura et al. 2014).
The development of isidium was considered an
important taxonomical characteristic in Leptogium
saturninum
when compared to L. hibernicum type, but further
studies are necessary. The isidia from Rio Grande
do Sul specimens are spherical when young and
cylindrical with age, branching on the apices,
whereas the isidia of L. hibernicum are granular
when young, remaining granular to branching with
age.
The type of Leptogium hibernicum also has
lobuloid or cerebroid structures on the lamina that
are not found in Rio Grande do Sul specimens.
Leptogium mantiqueirense Kitaura and
Marcelli and L. digitatum (A. Massal.) Zahlbr. have
Leptogium burgessii (L.) Mont. and L. ethiopicum
whereas L. laceroides B. de Lesd. and L. nylanderi
Kitaura and Marcelli have smooth upper surface
and ornaments (Kitaura and Marcelli 2013).
14. Leptogium involutum
S.M. Martins, Herzogia 2: 462. 1973. Type: Brazil,
Rio Grande do Sul State, municipality of Santana
do Livramento, APA do Ibirapuitã, Estância São
Maurício, riparian forest in a native grassland
Description: see Kitaura et al. (2014).
Known distribution to RS: Santana do
Livramento, only to type locality (Kitaura et al.
Notes: Leptogium involutum has involute
lobe margins, ridged upper surface and hairs with
cylindrical cells.
Leptogium hibernicum is an isidiate species
L. involutum that is a non-isidiate
species.
15. Leptogium isidiosellum (Riddle)
Bryologist Leptogium
marginellum var. isidiosellum
Jon, 1913, Britton and Shafer 276 (holo-: FH, iso-:
MICH!).
Thallus foliose, ca. 4.0 cm broad, bluish gray
to green under stereomicroscope. Lobes 2.5–5.0
mm wide, agglomerated, adpressed, adnate, upper
to longitudinal ridges under 20× magnification;
apices irregular, ascending, ornamented; lateral
margins plane to ascending, crenulated, smooth
to ornamented; lower side bluish, ridged. Isidia
irregular, withered appearance, simple to irregular
branched, marginal and laminal on the ridges, 0.075–
1964, Kitaura, 2012) not observed. Anatomy:
cortices, 2.5 × 2.5–5.0 µm. Medulla with columnar
MARCOS J. KITAURA et al. Leptogium FROM RIO GRANDE DO SUL STATE
An Acad Bras Cienc (2019) 91(1) e20180313 13 | 26
Figure 3 - Leptogium species (a
tissue). (b L. hibernicum (white arrow = hairy lower side). (c) ridged thallus with
irregular isidia of L. isidiosellum. (dLeptogium javanicum. Bars a = 100 µm, b, c and d = 2 mm.
MARCOS J. KITAURA et al. Leptogium FROM RIO GRANDE DO SUL STATE
An Acad Bras Cienc (2019) 91(1) e20180313 14 | 26
hyphae not observed; cyanobacteria green, 12–17
gelatinous matrix abundant, colorless. Pycnidia
absent.
Known distribution to RS: São Francisco
de Paula (Käffer et al. 2007) and Santana do
Specimens examined: Brazil, Rio Grande
do Sul: Campo Bom municipality, Chácara dos
Tachettos, riparian forest, (BHRS) - Area 2, 29º
M. I. Käffer 1025 (HAS); Caraá municipality,
APA do Caraá, 04 Nov. 2013. S.A. Martins 2852
(HAS); Ibid., Parobé municipality, 29°37’48.5”S,
338
(HAS); Ibid., Rolante municipality, 29°39’5.5”S,
341 (HAS); Ibid., Santo Antônio da Patrulha
municipality, Fazenda Marca, BHRS, riparian
Ibid.,
Ligustrum
Koch 924 p.p. (HAS).
Notes: Leptogium isidiosellum is characterized
by the ridged upper surface with irregular isidia
(Figure 3c). Apothecia were not found in Rio
Grande do Sul material, but they are constituted by
unpublished data).
Leptogium isidiosellum and L. coralloideum
the tissues of apothecia. The type of L. coralloideum
found. Leptogium isidiosellum
on the thalline exciple whereas L. coralloideum has
a thalline exciple constituted by one layer of cells
as the apothecia of L. phyllocarpum.
16. Leptogium javanicum Mont., Syll Gen Sp
Crypt: 379. 1856. Type: Java, Sindoro, ad truncos
et ramos Gnaphalii sp. (Nec aliorum) in summon,
Thallus foliose, ca. 7 cm broad, bluish gray
bluish under stereomicroscope. Lobes 3–5 mm
wide, agglomerated, attached in point, slightly
lateral margins ascending, smooth, undulate and
sinuous; lower side bluish to yellowish, irregular
lobules absent. Thallus attached by hapters,
frequent, homogeneous. Rhizinae and hairs
absent. Apothecia up to 2.5 mm diam., laminal,
pedicellate, disc usually plane, reddish; margin
smooth, beige; amphithecia concolorous with
the thallus, covered by pedicel, when young with
bullate projections on the apex; corona slightly
present, usually on mature apothecia; pedicel 0.5–
young and more wider than apothecia, irregular
longitudinal ridges. Anatomy: Thallus 35–40 µm
Medulla with columnar hyphae straight, frequent,
2 cells; cyanobacteria blue, number of cells per
matrix scarce, yellow. Apothecia with hymenium
110–140 µm high; subhymenium 62.0–75.0 µm
prosoplectenchymatous cells, hyaline; parahymenial
tissue continuous with hypothecium, collo- to
paraplectenchymatous; proper exciple absent;
thalline exciple cortex with paraplectenchymatous
base; basal paraplectenchymatous tissue 100–140
apices, 32.5–45.0 × 7.5–12.5 µm, 4–6 × 2–3 cells,
submuriform to muriform. Pycnidia not observed.
Known distribution to RS: See Spielmann
(2006).
MARCOS J. KITAURA et al. Leptogium FROM RIO GRANDE DO SUL STATE
An Acad Bras Cienc (2019) 91(1) e20180313 15 | 26
Specimens examined: Brazil, Rio Grande do
Sul: Caraá, Localidade Fraga, APA Municipal. 03
Sep. 2008. S.A. Martins 2140 pr. p. (HAS); Ibid.,
08 Oct. 2008, S.A. Martins 2184 (HAS).
Notes: Leptogium javanicum has inflated
pedicel with longitudinal ridges, and bullate
projections on the apices of young apothecia
(Figure 3d).
The material Junghuhn s/n, designated as
holotype of Stephanephorus javanicus Mont. is a
Leptogium,
and one lectotype must be selected according to
the ICN. The specimens in type collection of L.
javanicum were reported as 1 and 2. The fragment
1 is designated as the lectotype of L. javanicum,
whereas the fragment 2 is L. cf. cyanescens.
Aino Henssen, on the label of the material,
selected specimen 1 as the lectotype in 1966, but
she did not publish her choice.
The specimens from Rio Grande do Sul
have prosoplectenchymatous hypothecium and
ascospores 32.5–45.0 × 7.5–12.5 µm, whereas
the Java specimen has hypothecium with irregular
colloplectenchymatous cells and ascospores 15–
Rio Grande do Sul were considered Leptogium
javanicum, but further studies are required.
17. Leptogium marginellum (Sw.) Gray, Nat
Lichen marginellus Sw.,
Nov Gen Sp Pl [HBK]: 147. 1788. Type: Jamaica,
Blue Ridge Mt., (musci intermixtis innascitur, in
jugo montium caeruleorum), leg. Swartz s/n (lecto-:
H-ACH 1917!).
Thallus up to 5.0 cm broad, gray under
fluorescent light, matt, opaque, yellowish gray
under stereomicroscope. Lobes 1.5–2.5 mm wide,
agglomerated, attached in points, adnate, upper
ascending to revolute, smooth to ornamented;
lateral margins ascending, undulated and sinuous,
usually ornamented; lower side gray to yellowish,
erect, concolorous with the thallus, usually on
apothecia. Thallus attached by hapters, frequent,
homogeneous. Apothecia 0.1–0.4 mm diam.,
subpedicellate, disc plane to slightly concave,
reddish; margin of apothecia beige, smooth or
ornamented; amphithecia beige, ornamented;
corona absent; pedicel very short. Anatomy:
cortices 5.0 µm diam. Medulla with columnar
hyphae straight to inclined, frequent, ca. 5 cells;
5.0 µm diam.; gelatinous matrix scarce to frequent,
colorless. Apothecia with hymenium 110–125 µm
hypothecium 25–33 µm, colloplectenchymatous,
colorless; parahymenial tissue continuous with
hypothecium, colloplectenchymatous of irregular
cells; proper exciple absent; thalline exciple cortex
with paraplectenchymatous cells, 5.0–10.0 µm
at the base, with cyanobacteria duct that originate
the lobules. Ascospores and pycnidia not observed.
Known distribution to RS: Spielmann
Specimens examined: Brazil, Rio Grande do
Sul: Caraá municipality, APA do Caraá, 04 Nov.
2013. S.A. Martins 2851 (HAS); Ibid., Santo
Antônio da Patrulha municipality, Fazenda Marca,
Ibid., São Leopoldo municipality, Ecology base,
BHRS, riparian forest, Area 1, 29º45’50.6”S,
1028 (HURG).
Notes: Leptogium marginellum has upper
surface longitudinally slightly ridged under 20×
(Figure 4a).
MARCOS J. KITAURA et al. Leptogium FROM RIO GRANDE DO SUL STATE
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The apothecia of L. marginellum are abundant,
but the ascospores are usually not observed.
18. Leptogium megapotamicum
Bot 19: 8 (1924). Type: Brazil, Rio Grande do Sul,
Porto Alegre, Parthenon, Malme 574 (holo-: S!).
to yellowish gray under stereomicroscope. Lobes
1.5–7.0 mm wide, irregular, adnate, adpressed,
longitudinal ridges under stereomicroscope;
apices irregular, plane to ascending, inflated or
not inflated, smooth; lateral margins plane to
ascending, undulated, smooth. Isidia and lobules
absent. Apothecia up to 4 mm, marginal to laminal,
subpedicellate to pedicellate, disc reddish, plane
when young to convex; margin concolorous with
the thallus and yellowish, smooth to slightly
crenulated; amphithecia concolorous with the
thallus and yellowish or only concolorous with the
thallus, slightly crenulated to ridged; corona absent;
pedicel 0.5–3.0 mm long, originated by the ridges
(smaller) or margin of the lobes. Anatomy: Thallus
5.0 µm diam. Medulla with columnar hyphae
absent; cyanobacteria green, near the cortices,
spherical to ellipsoid, 2.5–5.0 × 2.5 µm; gelatinous
matrix abundant, colorless. Apothecia with
hymenium 80 µm high; subhymenium 20.0–37.5
proso- to colloplectenchymatous; parahymenial
tissues continuous with hypothecium, proso- to
colloplectenchymatous, 37.5 µm at the base, 25 µm
at the apex; proper exciple absent; thalline exciple
cortex with one layer, 5.0 µm (1 cell) at the apices,
µm (6 cells) at the base. Ascospores transversely
septate, 60–75 × 5 µm, 9–12 × 1 cells, apices acute,
acicular. Pycnidia not observed.
Known distribution to RS: See Spielmann
(2006).
Specimens examined: Brazil, Rio Grande do
Sul: Caraá, Localidade Fraga, APA Municipal, near
the trip, sunny place, 03 Sep. 2008. S.A. Martins
2140 (HAS); Ibid., 08 Oct. 2008. S.A. Martins
2198 pr. p.; Ibid., 03 Dec. 2008, S.A. Martins
2233, 2234, 2267 (HAS); Ibid., 27 Apr. 2009, S.A.
Martins 2276 (HAS).
Notes: Leptogium megapotamicum has ridged
transversely septate ascospores.
In Rio Grande do Sul, L. brebissonii and
L. megapotamicum have transversely septate
ascospores. However, L. brebissonii has isidia and
L. megapotamicum is a non-isidiate species.
19. Leptogium milligranum
67: 290. 1964. Type: U.S.A., New Jersey, Pasadena,
stereomicroscope. Lobes 2.0–4.0 mm wide,
agglomerated, attached in point, slightly adnate,
under 20× magnification; apices rounded to
irregular, ascending, ornamented; lateral margins
ascending, irregular, ornamented; lower side light
gray to yellowish, irregular ridges. Isidia granular,
moniliform to grouped, laminal. Lobules absent.
Thallus attached by hapters, frequent, homogeneous.
Rhizinae and hairs absent. Apothecia not observed.
cells of the cortices 5.0 µm diam. Medulla without
columnar hyphae; cyanobacteria green, spherical,
ca. 5 µm diam.; gelatinous matrix abundant,
colorless. Pycnidia not observed.
Known distribution to RS: New record to
Rio Grande do Sul, but previously mentioned to
São Paulo State.
Specimens examined: Brazil, Rio Grande
do Sul:
Ligustrum
2014. N.M. Koch 924 pr.p. (HURG).
MARCOS J. KITAURA et al. Leptogium FROM RIO GRANDE DO SUL STATE
An Acad Bras Cienc (2019) 91(1) e20180313 17 | 26
Figure 4 - Leptogium species (a) marginal apothecia with lobules of L. marginellum (white arrow). (b) ridged upper surface
of Leptogium megapotamicum. (c) granular isidia of L. milligranum. (d) subpedicellate apothecia of L. moluccanum. Bars
= 2 mm.
MARCOS J. KITAURA et al. Leptogium FROM RIO GRANDE DO SUL STATE
An Acad Bras Cienc (2019) 91(1) e20180313 18 | 26
Notes: Leptogium milligranum is characterized
simple, moniliform to grouped.
Leptogium brebissonii and L. milligranum can
Leptogium
brebissonii has transversely septate ascospores,
whereas L. milligranum has submuriform
ascospores.
Therefore, Leptogium brebissonii and L.
milligranum are usually found sterile and they can
Leptogium
milligranum has isidia granular, simple, moniliform
to grouped whereas L. brebissonii has isidia
granular to short cylindrical, simple to branched at
the apices.
Leptogium milligranum can be confused as L.
isidiosellum
isidia that has cylindrical appearance, whereas the
last is constituted by cylindrical to irregular isidia.
Leptogium milligranum was recently
mentioned to Brazil and São Paulo State (Kitaura
et al. 2017).
20. Leptogium moluccanum (Pers.) Vain.,
Étude sur la Classification Naturelle et la
Morphologie des Lichens du Brézil 1: 223. 1890.
Collema moluccanum Pers., Voy. Uranie, Bot 203
(1826). Type: Mullucann Island [],
PC).
Thallus foliose, up to 10 cm broad, light gray
under stereomicroscope. Lobes 2.0–5.0 µm wide,
agglomerated to overlapping, attached in point,
rounded, plane to ascending, smooth; lateral margins
smooth, plane to ascending, sinuous to undulated;
lower side bluish to gray, smooth to slightly
absent. Thallus attached by hapters, frequent,
homogeneous. Rhizinae and hairs absent. Apothecia
1.0–2.0 mm diam., laminal, subpedicellate, disc
plane, concave or convex, reddish; margin and
amphithecia smooth, beige; corona absent; pedicel
cortices with isodiametric cells 5.0 µm diam.
Medulla with columnar hyphae straight, 2–3 cells,
frequent to abundant; cyanobacteria blue, number
of cells per filament undetermined, 2.5–5.0 µm
diam.; gelatinous matrix scarce, hyaline. Apothecia
with hymenium ca. 125 µm high; subhymenium
ca. 50 µm, hyaline, prosoplectenchymatous;
parahymenial tissue continuous with hypothecium,
subparaplectenchymatous; proper exciple absent;
thalline exciple cortex paraplectenchymatous 5.0–
fusiform, apices obtuse and acuminated, 25–30 ×
7.5–12 µm (4–6 × 2–4 cells), muriform. Pycnidia
not observed.
Known distribution to RS: See Spielmann
(2006).
Specimens examined: Brazil, Rio Grande do
Sul: Caraá municipality, APA do Caraá. 03 Sep.
2008, S.A. Martins 2248 (HAS); Ibid., 08 Nov.
2008, S.A. Martins 2198 pr. p. (HAS); Ibid., 12
Dec. 2012, N.M. Koch 407 (HAS); Ibid., 04 Jun.
Ibid., Área de
Proteção Ambiental, Nascente do Rio dos Sinos,
BHRS, riparian forest, Area 7, 29º42’05.8”S,
Ibid., Mata de restinga APP6,
14 Apr. 2016. J. Pedroso 317 (HURG).
Notes: Leptogium moluccanum is characterized
by the upper surface smooth to slightly ridged under
1.0–2.0 mm diam. (Figure 4d).
Leptogium azureum has pedicellate apothecia
and differs from L. moluccanum that has
subpedicellate apothecia.
MARCOS J. KITAURA et al. Leptogium FROM RIO GRANDE DO SUL STATE
An Acad Bras Cienc (2019) 91(1) e20180313 19 | 26
Leptogium cochleatum has subpedicellate
apothecia as well as L. moluccanum
and James 1983), whereas L. moluccanum has
slightly ridged and apothecia and amphithecia with
smooth margins.
21. Leptogium phyllocarpum (Pers.) Mont.,
Ann Sci Nat, Bot Biol Vég 3, 10: 134. 1848.
Collema phyllocarpum Pers., Voy Uranie, Bot:
204. 1826. Type: Brazil, Rio Grande do Sul, Serra
dos Vallos per Cruz Alta, in Arbore solitária ripae
rivuli, 21 Apr. 1893, G.A. Malme 1265 (neo-: S!,
designated by Jørgensen (2002)).
Thallus foliose, 4.0–10.0 cm broad, gray
stereomicroscope. Lobes up to 2.0 mm wide,
agglomerated, adpressed, adnate, upper surface
under 20× magnification; apices rounded,
ascending, smooth; lateral margins ascending,
sinuous and undulated, smooth; lower side gray,
Lobules without constrict base, ca. 0.25–0.5
mm diam., firm, erect, only on amphithecia,
concolorous with the thallus. Thallus attached by
hapters, frequent, homogeneous. Rhizinae and
hairs absent. Apothecia (only young) 0.3–0.4
mm diam., laminal, subpedicellate, disc concave,
reddish to brown; margin smooth, concolorous
with the thallus; amphithecia ornamented with
collar, concolorous with the thallus; corona
absent; pedicel very short, concolorous with the
isodiametric cells of the cortices 5.0 µm diam.
Medulla without columnar hyphae; cyanobacteria
green, 5–10× 2.5–5 µm, 7–12 elongated cells per
filament; gelatinous matrix abundant, colorless.
Apothecia (young) with hymenium 125 µm
hypothecium 25 µm, prosoplectenchymatous;
proper exciple subparaplectenchymatous, 100–
continuous with the proper exciple, 100 µm (ca.
25 cells) at the base, 25 µm (10 cells) at the apices,
subparaplectenchymatous; thalline exciple cortex
mid-height and base. Ascospores not observed.
Pycnidia not observed.
Known distribution to RS: See Spielmann
et al. 2015).
Specimens examined: Brazil, Rio Grande do
Sul: Caraá, Localidade Fraga, APA Municipal, 08
Oct. 2008. S.A. Martins 2197 (HAS); Ibid., 04 Jun.
Ibid., 04 Nov.
2013. S.A. Martins 2847 (HAS).
Notes: Leptogium phyllocarpum is constituted
subpedicellate, laminal and ornamented (Figure
5a).
Apothecia ornaments were found of various
sizes and forms (Martins 2197 and 2847). They
vary from high ridges to collar structures (lobules
with fused base on the apices of the apothecia).
Probably, the specimens of Rio Grande do Sul
are not L. phyllocarpum stricto sensu, but further
studies are still necessary.
22. Leptogium pichneum
Collema tremelloides var.
pichneum Ach., Syn Meth Lich: 343. 1814. Type:
Nova Hollandia and America Borealis habitats
(H-ACH).
Description: See Acharius (1814), Malme
(1924).
Known distribution to RS: See Spielmann
2007).
Notes: Leptogium pichneum is characterized
by the upper surface slightly smooth, lower side
distinctly rugulose, and isidia simple and cylindrical
MARCOS J. KITAURA et al. Leptogium FROM RIO GRANDE DO SUL STATE
An Acad Bras Cienc (2019) 91(1) e20180313 20 | 26
when young to branched with age (Acharius 1814,
Malme 1924).
Leptogium pichneum differs from L.
cyanescens mainly by the lower surface. Leptogium
pichneum has lower side distinctly rugulose and
L. cyanescens has lower cortex smooth to slightly
ridged under 20× magnification. The type of L.
pichneum was still not described by our protocol
23. Leptogium puiggarii Müll. Arg., Flora
(Regensburg) 62: 161. 1879. Type: Brazil,
São Paulo State, Apiahy (Apiaí) municipality,
Meridional Region mountain, J.J. Puiggari 145
Description: See Müller Argoviensis (1879).
Known distribution to RS: See Spielmann (2006).
Notes: Leptogium puiggarii is characterized
(Müller Argoviensis 1879).
24. Leptogium sessile Vain. Ann Acad Sci
Fenn, Ser A, IV, Biol 6(7): 108. 1915. Type:
Dominica (Ocidental India), Shawford State, ad
(holo-: TUR-V 10791!, iso-: TUR-V 10792!, FH?).
Description: See Vainio (1890), Kitaura
(2012).
Known distribution to RS: See Spielmann
(2006).
Notes: Leptogium sessile is characterized
The amphithecia is concolorous with the
thallus and the thalline exciple is constituted by
one layer of isodiametric cells.
Figure 5 - Leptogium species (a) ornamented apothecia of L. phyllocarpum with. (b) pedicel originated by the thallus of L.
vesiculosum. Bars = 2 mm.
MARCOS J. KITAURA et al. Leptogium FROM RIO GRANDE DO SUL STATE
An Acad Bras Cienc (2019) 91(1) e20180313 21 | 26
Leptogium sessile is the only species
reported to Rio Grande do Sul with immerse to
sessile apothecia. Leptogium cochleatum and L.
moluccanum have subpedicellate apothecia and L.
azureum has pedicellate apothecia, all species with
25. Leptogium subjuressianum Marcelli and
Kitaura, Mycotaxon 120: 218. 2012. Type: Brazil,
Rio Grande do Sul State, municipality of Tapes,
Marcelli 26459 (holo-: SP!).
Description: See Kitaura and Marcelli (2012).
Known distribution to RS: Tapes, only type
locality (Kitaura and Marcelli 2012).
Notes: Leptogium subjuressianum is
characterized by the presence of hairs constituted
by cylindrical cells with arachnoid appearance.
Leptogium subjuressianum has rounded lobules
L.
juressianum that has lacinules, elongated ornaments
(Kitaura and Marcelli 2012).
Leptogium involutum has usually hairs on
the lower side with velvety appearance, whereas
L. subjuressianum has hairy upper surface with
arachnoid appearance.
The hairs of L. subjuressianum are constituted
of L. hibernicum that are constituted by spherical
cells.
26. Leptogium tuckermanii
Missouri Bot Gard 20: 436. 1933. Type: Cuba,
Description: See Dodge (1933).
Known distribution to RS: See Spielmann
(2006).
Notes: Leptogium tuckermanii is characterized
0.5–0.9 mm wide and ascopores 18–22 × 10–12
µm (Dodge 1933).
The specimens of Leptogium moluccanum
sensu Vainio (1890) were included in a synonymy
list of L. tuckermanii (Dodge 1933). Both species
have almost the same measures of apothecia
and ascospores: Leptogium moluccanum sensu
Vainio has apothecia with 0.2–0.7 mm diam. and
ascospores with 18–22 × 6–12 µm, whereas L.
tuckermanii has apothecia 0.5–0.9 mm diam. and
ascospores 18–22 × 10–12 µm (Dodge 1933).
Therefore, Leptogium moluccanum (Pers.)
we also considered them as two distinct species.
The type material of both species, Leptogium
tuckermanii and L. moluccanum, were still not
studied and Leptogium moluccanum is probably a
complex of species.
27. Leptogium ulvaceum (Pers.) Vain.,
Lichenes Insularum Philippinarum III: 38. 1921.
Collema ulvaceum Pers., Voy Uranie, Bot.: 203.
1826. Type: Mariana Islands, ad arborum truncos,
C. Gaudichaud s/n (Type not traced).
Description: See Persoon (1826), I.P.R.
Cunha, unpublished data.
Known distribution to RS: See Spielmann
(2006).
Notes: Leptogium ulvaceum is characterized
green, and lobes with crenulate margin.
Leptogium ulvaceum is similar to L.
tuckermanii and L. moluccanum
with crenulate margin and the other two, lobes with
smooth margin.
28. Leptogium vesiculosum
Lichen vesiculosus Sw., Nov
Gen Sp [HBK]: 147. 1788. Type: Jamaica, on tops
of mountains, O. Swartz (Dodge 1933) (Type not
traced, H?).
Description: Thallus foliose, 3.5 cm broad,
gray under stereomicroscope. Lobes 2–6 mm wide,
slightly overlapping to agglomerated, attached in
rounded, ascending, smooth; lateral margins
ascending, smooth, undulate and sinuous; lower
MARCOS J. KITAURA et al. Leptogium FROM RIO GRANDE DO SUL STATE
An Acad Bras Cienc (2019) 91(1) e20180313 22 | 26
attached by hapters, frequent, homomerous.
Rhizinae and hairs absent. Apothecia up to 2.0
mm diam., submarginal to laminal, pedicellate,
disc plane, reddish; margin smooth, concolorous
with the thallus; amphithecia concolorous with
the thallus, without ornaments; corona absent;
pedicel 0.5–3.0 mm high, without constrict base,
to longitudinal ridges. Anatomy: Thallus 75–80
diam. Medulla with columnar hyphae sinuous,
5–7 cells long; cyanobacteria green, 10–15 cells
frequent to scarce, hyaline. Apothecia with
hymenium ca. 225 µm high, subhymenium ca. 75
irregular hyphae with intracellular space, slightly
yellowish; parahymenial tissue continuous with
subhymenium, colloplectenchymatous, ca. 50 µm
basal tissue; basal paraplectenchymatous tissue
with paraplectenchymatous cells, 75 µm (7cells) at
the margin, 140 µm (7–9 cells) at the mid-height,
thallus inserted near the apice of thalline exciple.
Ascospores fusiform, acute, 25–30 × 7.5–10.0
µm, 4–6 × 1–2 (–3) cells, muriform. Pycnidia not
observed.
Known distribution to RS: New record to Rio
Grande do Sul, but previously mentioned to Mato
Grosso, Minas Gerais and Santa Catarina States.
Specimens examined: Brazil, Rio Grande do
Sul: Maquiné Municipality, Encantada locality,
N.M. Koch 980.
Note: Leptogium vesiculosum is characterized
pedicel and the smooth amphithecia that is
concolorous with the thallus (Figure 5b).
Leptogium vesiculosum differs from L.
javanicum by the margin of the apothecia and
the type of amphithecia. Leptogium vesiculosum
has the margin of apothecia concolorous with
the thallus and amphithecia smooth whereas that
L. javanicum has margin of apothecia beige, and
bullate projections on the apex of amphithecia.
EXCLUDED SPECIES
Leptogium tremelloides was excluded from the
species list of Rio Grande do Sul. Nowadays,
Leptogium tremelloides was combined in Scytinium
lichenoides
(Otálora et al. 2014) and is considered a European
species (Otálora et al. 2008).
Leptogium tremelloides, Malme 492 B, Malme 534,
Malme 849 B and Malme 951, were not revised by
us, but they were re-determined as L. azureum by
Marcelli (2002).
Identication key to Leptogium in Rio
Grande do Sul State, Brazil
1a. Thallus with hairs, tomentose 2
1b. Thallus without hairs, not tomentose 4
2a. Hairs constituted by spherical cells L.
hibernicum
2b. Hairs constituted by cylindrical cells 3
3a. Hairs usually on the upper surface with
cobwebby appearance L. subjuressianum
3b. Hairs usually on the lower surface with velvety
appearance L. involutum
5a. Lower surface with cavities (cyphella- or urn-
L. puiggarii
5b. Lower surface without cavities 6
MARCOS J. KITAURA et al. Leptogium FROM RIO GRANDE DO SUL STATE
An Acad Bras Cienc (2019) 91(1) e20180313 23 | 26
6a. Thallus with isidia or lobules 7
6b. Thallus without isidia or lobules 12
7a. Cylindrical isidia present 8
7b. Lobules present 10
8a. Upper surface smooth, thallus bluish L.
cyanescens
8b. Upper surface rugulose, thallus grayish to
9a. Isidia with withered appearance, thallus grayish
to bluish-gray L. austroamericanum
9b. Isidia without withered appearance, thallus
L. pichneum
10a. Lobules only on apothecia L. marginellum
10b. Lobules on the thallus and apothecia 11
11a. Rounded lobules on the lamina L. atlanticum
11b. Flattened and elongated lobules on the lamina
L. diaphanum
12a. Apothecia sessile L. sessile
12b. Apothecia subpedicellate to pedicellate 13
14a. Margin of apothecia smooth L. vesiculosum
14b. Margin of apothecia with bullate projections
L. javanicum
15a. Apothecia pedicellate L. azureum
15b. Apothecia subpedicellate 16
16a. Amphithecia concolorous with the thallus L.
corticola
16b. Amphithecia beige to yellowish 17
17a. Margin of amphithecia ornamented with
denticules L. denticulatum
17b. Margin of amphithecia without ornaments 18
18a. Upper surface notably finely rugulose
(striate) L. cochleatum
crenulate L. ulvaceum
lobes smooth 20
20a. Apothecia 0.5–0.9 mm wide, ascospores 18–
22 × 10–12 µm (Dodge 1933) L. tuckermanii
20b. Apothecia 0.5–2.0 mm wide, ascospores 28–
36 × 7–10 µm (Verdon 1992) L. moluccanum
21a. Thallus with isidia 22
21b. Thallus without isidia 26
23
22b. Isidia others forms, concolorous with the
thallus 24
23a. Ascospores submuriform to muriform L.
milligranum
23b. Ascospores transversely septate L. brebissonii
L. exaratum
exciple, with subhymenial paraplectenchymatous
tissue L. coralloideum
cell) thalline exciple, without subhymenial
paraplectenchymatous tissue L. isidiosellum
26a. Ascospores transversely septate L.
megapotamicum
26b. Ascospores submuriform to muriform 27
27a. Apothecia submarginal to marginal,
L.
phyllocarpum
27b. Apothecia usually submarginal, amphithecia
smooth without lobules L. chloromelum
MARCOS J. KITAURA et al. Leptogium FROM RIO GRANDE DO SUL STATE
An Acad Bras Cienc (2019) 91(1) e20180313 24 | 26
DISCUSSION
Total of 16 species were documented during the
study of Leptogium specimens from HAS and
HURG Herbaria from Rio Grande do Sul, southern
Brazil, representing ca. 57% of Leptogium
diversity of the state. The descriptions of found
species have not a wide morphological variability
as thought previously. The jelly lichens have fragile
appearance, but the taxonomical characteristics
e.g. columnar hyphae and paraplectenchymatous
tissues, during the dehydration and hydration
process.
Leptogium exaratum is proposed as a new
species; L. atlanticum, L. milligranum and L.
vesiculosum
state; and the lectotype of L. javanicum was
determined after being indicated by Aino Henssen
in the label of the type specimen.
Leptogium fleigiae, L. conglutinatum and
L. paulistanum were also recorded in the list of
Leptogium
but these species are not published and are
considered as invalid names, although they are
updated soon.
The State of Rio Grande do Sul has the highest
Leptogium from Brazil with 28
cited species, but further studies are necessary and
the diversity can be higher than showed here.
ACKNOWLEDGMENTS
support, as well as the reviewers. The authors MJK
and NMK received scholarships from Coordenação
de Aperfeiçoamento de Pessoal de Nível Superior
(CAPES) as post-doctorate (PNPD Program) and
ASR as master degree at Universidade Federal de
Mato Grosso do Sul (UFMS).
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