Hoehnea 36(4): 551-595, 3 tab., 41 ig., 2009
551
Parmotrema s.l. (Parmeliaceae, lichenized Ascomycota) from Serra
Geral slopes in central Rio Grande do Sul State, Brazil
Adriano Afonso Spielmann1,2 and Marcelo Pinto Marcelli1
Received: 08.12.2008; accepted: 10.09.2009
ABSTRACT - (Parmotrema s.l. (Parmeliaceae, lichenized Ascomycota) from Serra Geral slopes in central Rio Grande do
Sul State, Brazil). A survey of the parmotremoid lichens occurring in the central area of Rio Grande do Sul State (Brazil)
revealed 31 species of Parmotrema A. Massal., distributed in three groups corresponding to the former genera Canomaculina
Elix & Hale, Parmotrema s. str. and Rimelia Hale & Fletcher. All species are described, illustrated, commented and special
notes compare similar species as an aid to identiication.
Key words: Canomaculina, Parmotrema, Rimelia, Rio Grande do Sul
RESUMO - (Parmotrema s.l. (Parmeliaceae, lichenized Ascomycota) de escarpas da Serra Geral na região central do Rio
Grande do Sul State, Brasil). Em um levantamento das espécies de Parmotrema A. Massal. que ocorrem nos barrancos
e peraus de uma região central do Rio Grande do Sul, 31 espécies foram encontradas, as quais podem ser divididas em
três grupos morfológicos, que correspondem a Canomaculina Elix & Hale, Parmotrema s. str. e Rimelia Hale & Fletcher.
Descrições, ilustrações e comentários sobre espécies próximas são fornecidos, baseando-se no material encontrado.
Palavras-chave: Canomaculina, Parmotrema, Rimelia, Rio Grande do Sul
Introduction
Continuing the studies on the diversity of
Parmeliaceae from Southern Brazil, focused on
species found in roadsides and slopes in the central
Rio Grande do Sul State (Spielmann 2005, Spielmann
& Marcelli 2008), we are presenting here the most
diverse group found: Parmotrema s.l.
Parmotrema was proposed a long time ago
(Massalongo 1860) to delimit some species belonging
to the genus Parmelia Ach., and Parmotrema
perforatum (Wulfen) A. Massal. was chosen as the
type. The name Parmotrema reports the perforate
apothecia of this species, from the Greek parmos =
cup, and trema = perforation (Feige 1998). The genus
was largely neglected by the contemporary workers,
maybe with the exception of Krempelhuber (Hale
1984), and the species belonging to Parmotrema
were dealt with in the large genus Parmelia Ach. by
Zahlbruckner (1926a, 1930). Vainio (1890) assembled
most species at present placed in Parmotrema in
his Parmelia section Amphigymnia Vainio, and his
delimitation was largely followed, sometimes with
slight changes. Dodge (1959), for example, proposed
Parmelia subgenus Amphigymnia (Vainio) Dodge,
and with this name, the species were monographically
treated by Hale (1965). In this meantime, some
combinations in Parmotrema were proposed by M.
Choisy, as can be seen in Lamb (1963). Later, Hale
(1974a) recognized Parmotrema and placed in it the
species studied in his monograph of Amphigymnia
(Hale 1965). Therefore, Parmotrema began to be
widely used. Nevertheless, some workers (e.g.
Hawksworth, Dey, Krog, and Swinscow) do not
accept this delimitation (Culberson 1991). Krog
& Swinscow (1983) changed their mind, yet until
recently (see Purvis et al. 1992) some Europeans still
do not used Parmotrema (and no one of the genera
proposed by Hale). Today the generic delimitation
in Parmeliaceae is very controversial, but several
Hale’s propositions are more widely used (see, e.g.,
Hawksworth et al. 2008).
The genus Canomaculina was proposed by
Elix & Hale (1987) to accommodate three species
previously classiied within Parmelina Hale (Hale
1974b, 1976): C. consors (Nyl.) Elix & Hale, C.
muelleri (Vainio) Elix & Hale and C. pilosa (Stiz.)
Elix & Hale. Then the characteristic features of
Canomaculina were adnate to strongly adnate thalli
1. Instituto de Botânica, Caixa Postal 3005, 01061-970 São Paulo, SP, Brazil
2. Autor para correspondência: adrianospielmann@yahoo.com.br
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Hoehnea 36(4): 551-595, 2009
(Hale 1976), typically narrow lobes, (1–) 2–4 (–5)
mm, and dimorphic rhizines (Elix & Hale 1987).
Some years later, Kurokawa (1991a) proposed
Rimeliella Kurok., to embrace several species
formerly classiied within Parmotrema, with
adnate to loosely adnate thalli, wide lobes (5–15
mm) and dimorphic rhizines. Afterwards, Elix
(1997) concluded that lobe’s width (narrow in
Canomaculina, wide in Rimeliella) was not enough
to distinguish the two genera, and since both have
dimorphic rhizines, he synonymized Rimeliella
under Canomaculina.
Rimelia was proposed by Hale & Fletcher
(1990), based on Parmelia section Hypotrachyna
*Irregularis (Vainio 1890). It shares several features
with Parmotrema, but differs by a combination of
characters: upper surface reticulate cracked, cilia
almost always present [lacking only in Rimelia
ruminata (Zahlbr.) Hale & Fletcher], lower surface
generally rhizinate to margins (with several
exceptions), and rhizines often squarrose (Hale
& Fletcher 1990). However, there are species of
Parmotrema with reticular maculae (particularly
those with salazinic acid) and rhizinate to the margin,
as well as species of Rimelia with a wide bare zone
on lobes undersurface, so that the distinction of the
genera is, in many instances, dificult. Canomaculina
Elix & Hale, by their turn, has species with broad or
narrow lobes, ciliate, upper surface with efigurate
maculae, lower surface rhizinate to the margins and
dimorphic rhizines (Elix 1997).
Previously we treated Canomaculina and
Rimelia separately (Spielmann & Marcelli 2008),
but here they are included under Parmotrema,
following the proposed circumscription of Blanco
et al. (2005). Although few Parmotrema s. str. were
included in this paper, P. perforatum, the type of
the genus, was shown to be closely related to the
others representatives. Maybe future studies, based
on a higher amount of species, mainly from South
America, could change this picture.
Nowadays about 350 species of Parmotrema s.l.
are known (Blanco et al. 2005), although much more
remain to be discovered or correctly delimited, based
on modern taxonomic features.
Material and methods
Detailed description and maps of the studied area,
material and methods employed and considerations
on general morphology can be found in Spielmann
(2005) and Spielmann & Marcelli (2008). Spot
tests were performed by the use of K (potassium
hydroxide), C (sodium hypochlorite) and P (paraphenylenediamine). TLC analysis followed Huneck
& Yoshimura (1996), Orange et al. (2001) and
Bungartz (2001). Several specimens were collected
“on the roadside”. So this information is not repeated
in the Specimens examined. Just different data were
kept. Brazilian States abbreviations follow Marcelli
(1998).
Results and Discussion
In the following lora we tried to put in evidence
old and modern generic concepts, both in the key
and in the disposition of the species, to facilitate the
identiication. The igures, however, are arranged in
alphabetical order.
Key to Parmotrema from Serra Geral slopes in
central Rio Grande do Sul State, Brazil
1. Upper surface clearly maculate, underside rhizinate up to the margin or more rarely nude
2. Maculae efigurate, cilia generally short, tapered and thick, underside often brown throughout
or black only in the center (formerly Canomaculina; Group I below)
3. Thallus with soredia
4. Medulla K–, KC+ reddish, P- (norlobaridone) ..................................................... P. conferendum
4. Medulla K+ yellow, KC-, P+ orange (stictic acid) ...................................................... P. muelleri
3. Thallus without soredia
6. Medulla K+ yellow → red (salazinic acid) ......................................................... P. subcaperatum
6. Medulla K–
7. Medulla KC+ rose (norlobaridone) .................................................................. P. recipiendum
7. Medulla KC– (fatty acids) ....................................................................................... P. consors
2. Maculae reticulate, cilia long and thin, underside black (formerly Rimelia; Group III below)
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553
8. Thallus with soredia or pustules
9. Thallus with soredia raised from pustules and soralia; medulla UV+ yellow-orange
(lichexanthone) ..................................................................................................... P. diffractaicum
9. Thallus with soredia raised only from soralia; medulla UV–
10. Medulla K+ yellow → red (salazinic acid) ..................................................... P. clavuliferum
10. Medulla K–
11. Medulla KC+ reddish rose (norlobaridone) ........................................ P. commensuratum
11. Medulla KC– (caperatic acid) ......................................................................... P. simulans
8. Thallus without soredia or pustules
12. Medulla K+ yellow → red, KC– (salazinic acid) ........................................................ P. cetratum
12. Medulla K–, KC+ rose or red (norlobaridone) ...................................................... P. homotomum
1. Upper surface emaculate (rarely inconspicuously maculate), underside with a distinct bare zone
around the margins (Parmotrema s. str. ; Group II below)
13. Thallus with soredia, isidia or pustules
14. Thallus with isidia (occasionally disintegrating into soredia with the age)
15. Lobes eciliate; medulla C+ red (lecanoric acid) ........................................................ P. tinctorum
15. Lobes ciliate; medulla C–
16. Medulla K+ yellow, KC+ reddish, UV– (stictic acid and norlobaridone) ......... P. internexum
16. Medulla K–, KC+ rose → quickly orange, UV+ greenish blue (alectoronic acid) ...... P. mellissii
14. Thallus with soredia or pustules
17. Thallus with pustules, soredia present or lacking
18. Lobes eciliate ................................................................................................. P. alidactylatum
18. Lobes ciliate
19. Medulla yellowish to orange ............................................................... P. lavomedullosum
19. Medulla white
20. Medulla K+ yellow, UV– (stictic acid) ......................................................... P. bangii
20. Medulla K+ yellow → red (salazinic acid), UV+ yellow-orange
(lichexanthone) ....................................................................................... P. spinibarbe
17. Thallus without pustules, soredia always present
21. Lobes eciliate
22. Medulla C+ red (lecanoric acid) .............................................................. P. austrosinense
22. Medulla C–
23. Medulla K+ yellow (atranorin) ................................................................. P. mordenii
23. Medulla K– .................................................................................... P. praesorediosum
21. Lobes ciliate
24. Medulla K+ yellow, P+ orange (stictic acid) .................................................. P. perlatum
24. Medulla K–, P–
25. Medulla KC+ rose or reddish, UV– (gyrophoric acid)
26. Thallus membranaceous to subcoriaceous, of medium size (8.5−11.0
cm in diameter); conidia sublageniform, 5.0−9.0 µm long ........ P. sancti-angeli
26. Thallus subcoriaceous, robust (20–28 cm in diameter); conidia
iliform, 8.0–12.5 µm long .................................................................. P. indicum
25. Medulla KC+ rose → quickly orange, UV+ greenish blue
27. Soredia often rising from arbuscular structures like a caulilower;
with an orange medullar pigment K+ red (unknown anthraquinone)
present at the soralia and lobe apices ........................................ P. hypomiltoides
27. Soredia rising from marginal to submarginal soralia and usually linear;
pigment orange K+ red (skyrin) present only in the older parts, next
to the lower cortex ..................................................................... P. rampoddense
13. Thallus without vegetative propagules
28. Medulla K+ yellow or K+ yellow → red
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29. Medulla K+ yellow → red (salazinic acid) ....................................................... P. mantiqueirense
29. Medulla K+ yellow (stictic acid) ................................................................................ P. eciliatum
28. Medulla K–
30. Medulla KC–, UV– (protopraesorediosic acid) ..................................................... P. melanothrix
30. Medulla KC+ rose → quickly orange, UV+ greenish blue (alectoronic acid)
31. Lower surface with a white margin; conidia unciform 4.0−5.5 µm long .............
............................................................................................................................ P. subrugatum
31. Lower surface with a brown margin; conidia bacillar to iliform 6.0–12.5 µm
long ........................................................................................................................... P. wainioi
Group I – Canomaculina-like lichens (species
with dimorphic rhizines, efigurate or punctiform
maculae, and rhizinate up to the margin, belonging
to Canomaculina sensu Elix 1997)
This group of species is distinguished by the
usually thickened, furcated and spiky cilia, maculate
efigurate upper cortex, lower surface often brown
and usually rhizinate up to the margin and the
presence of dimorphic rhizines, the later feature
being the more distinctive one.
To make easier the comparison between the
known species of canomaculinoid lichens, a table
(1) was created, with data obtained from literature:
Adler & Elix (1987), Canêz (2005), Chen et al.
(2003), Eliasaro (2001), Elix (1997), Ferraro & Elix
(1993, 2000), Fleig (1997), Hale (1959, 1965, 1976,
1977, 1986), Krog (1974), Krog & Swinscow (1981),
Kurokawa (1974, 1991a), Kurokawa & Lai (2001),
Marcelli & Ribeiro (2002), Mata García (1994),
Ribeiro (1998) and Swinscow & Krog (1988).
Parmotrema conferendum Hale, Mycotaxon 5(2):
433. 1977 ≡ Rimeliella conferenda (Hale) Kurok.,
Annals of the Tsukuba Botanical Garden 10: 4.
1991 ≡ Canomaculina conferenda (Hale) Elix,
Mycotaxon 65: 476. 1997. Type: VENEZUELA.
Estado dE Mérida: El Valle, on rocks along stream,
2,600 m alt., 8-II-1974, M.E. Hale 43291 (holotype
US), ide Hale (1977).
Figure 9
Thallus olivaceous gray, brownish in herbarium,
lobate, membranaceous to subcoriaceous, loosely
adnate, saxicolous, 6.5-13.5 cm broad. Lobes
irregularly branched, laterally overlapping to
contiguous, 5-15 mm wide, surface smooth to
irregular, lustrous, becoming rugose and scrobiculate
in the center; apical zone plane to subconcave,
margin plane to ascendant or revolute, crenate to
incise-crenate (sublacinulate); marginal zone plane
to ascendant or revolute, margin undulate, crenate to
30518004 miolo.indd 554
lacinulate, sometimes involute. Lacinules simple, 0.51.0 × 0.5-2.0 mm, marginal, plane to canaliculated,
evenly distributed, apex rounded or truncate. Maculae
weak to distinct, punctiform, irregular or reticulate,
laminal. Cilia black, simple to furcated or sometimes
squarrose, 0.10-1.20 × 0.02-0.10 mm, frequent,
evenly distributed. Pustules and isidia absent. Soralia
dirty white, usually capitate, coalescing at maturity,
marginal to submarginal, turning the lateral margin
involute; soredia subgranular. Medulla white. Lower
surface pale brown, lustrous, sometimes strongly
veined and depressed, rugose; marginal zone pale
brown, sometimes white variegated, lustrous, 0.51.0 mm wide, with short or immature rhizines, bare,
smooth, rugose or veined, without a sharp limit.
Rhizines black to dark brown, dimorphic, the short
ones simple, straight or curly, 0.10-0.50 × 0.010.05 mm, abundant, evenly distributed, the long ones
usually simple, sometimes furcated or irregularly
branched, 0.50-3.00 × 0.05-0.20 mm, frequent, in
disperse groupings. Apothecia unknown. Pycnidia
submarginal, little conspicuous, without prominent
margin, ostiole black. Conidia iliform, straight or
arcuate, 10-16 × ca. 1 µm.
Color tests: cortex K+ yellow, UV- ; medulla K-,
C- or C+ rose, KC+ reddish, P-, UV-.
tLC: atranorin, norlobaridone, loxodin and
unidentiied fatty acids.
Distribution: Africa (Winnem 1975), North America
(Esslinger 2008) and South America (Hale 1977). In
South America it is known to Argentina (Hale 1977,
Calvelo & Liberatore 2002), Brazil (Marcelli 2004),
Uruguay (Osorio 2003) and Venezuela (Hale 1977).
In Brazil it was recorded to PR (Eliasaro & Donha
2003) and RS (Spielmann 2006).
Specimens examined: BRAZIL. rio GrandE do suL:
Sinimbu, Cava Funda, 29°27′41.8″S, 52°31′11.7″W,
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500 m alt., 4-II-2003, A.A. Spielmann 24 (SP); idem,
29°27′33.4″S, 52°31′05.1″W, 520 m alt., with mosses,
5-I-2004, A.A. Spielmann & L.S. Canêz 668 (SP);
idem, Linha Almeida, 29°23′20.2″S, 52°30′21.9″W,
5-I-2004, A.A. Spielmann & L.S. Canêz 1297 (SP).
Parmotrema conferendum is distinguished by
the usually capitate, marginal to submarginal soralia,
and medulla with norlobaridone and loxodin (K–,
KC+ reddish). The morphologically similar species
differ mostly in chemical constituents (table 1): P.
subsumptum has salazinic and consalazinic acids,
reacting K+ yellow → blood red (Fleig 1997), and P.
reitzii Hale produces salazinic acid and norlobaridone
(Hale 1977), reacting K+ yellow → blood red and
KC+ reddish. This species was placed as synonym
of Rimeliella conferenda by Kurokawa (1991a),
and as synonym of Rimeliella subsumpta by Fleig
(1997). However, Elix (1997) accepted P. reitzii as
distinct, an opinion kept here. Parmotrema larense
López-Figueiras has, in addition to norlobaridone,
lichexanthone in the medulla (Mata García 1994),
being UV+ yellow-orange.
The specimen A.A. Spielmann & L.S. Canêz
668 (SP) has the lower surface strongly veined and
sometimes depressed, as well as a white variegated
margin. All specimens were found growing on the
roadsides, in open places.
Parmotrema consors (Nyl.) Krog & Swinscow, The
Lichenologist 15(2): 129. 1983 ≡ Parmelia consors
Nyl., Flora 68: 613. 1885 ≡ Canomaculina consors
(Nyl.) Elix & Hale, Mycotaxon 29: 239. 1987.
Type: BRAZIL. Minas GErais: Weddell (lectotype
selected by Hale 1976, H-NYL35277), ide Hale
(1976).
Figure 10
Thallus greenish gray, lobate, adnate,
corticicolous or saxicolous, 8-20 cm broad.
Lobes irregularly branched, laterally overlapping,
2-15 mm wide, surface smooth to rugose and
scrobiculate, usually pruinose, becoming strongly
rugose in the center; margin smooth to incise-crenate
(sublacinulate). Lacinules, pustules soredia and isidia
absent. Maculae distinct, punctiform to irregular,
laminal. Cilia black, simple to furcated or irregularly
branched, spiked and robust or slender, 0.103.00 × 0.05-0.20 mm, usually turned downwards,
abundant, evenly distributed. Medulla white. Lower
surface black, lustrous, smooth to rugose; marginal
zone brown, lustrous, 1-3 mm wide, rhizinate,
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555
with attenuate limit, smooth to papillate. Rhizines
black, simple to irregularly branched, 0.20-3.00
× 0.02-0.30 mm, dimorphism not always evident,
but when discernible the short rhizines abundant
and evenly distributed, the larger ones frequent and
assembled. Apothecia concave to urceolate, 1.59.0 mm in diameter, adnate to substipitate, laminal,
margin smooth to crenate, amphithecium smooth,
disc brown, perforate at maturity; epithecium 3-8
µm; hymenium 35-60 µm; subhymenium 15-30
µm; ascospores ellipsoid, 11.0-16.5 × 7.0-10.0
µm, episporium 1.0-1.5 µm. Pycnidia laminal to
submarginal, conspicuous, usually with prominent
margin, ostiole black; conidia bacillar to iliform,
straight or curved, 7-15 × ca. 1 µm.
Color tests: cortex K+ yellow, UV-; medulla K-, C-,
KC-, P-, UV-.
TLC: atranorin and an unidentiied fatty acid.
Distribution: Oceania (Elix 1994) and South America
(Hale 1976). In South America it is known to Argentina
(Osorio 1970d, 1976; Hale 1976; Adler 1992; Calvelo
& Liberatore 2002), Brazil (Zahlbruckner 1930, Hale
1976, Marcelli 2004), Paraguay (Hale 1976), Uruguay
(Osorio 1970b/c, 1972, 1992a; Hale 1976) and
Venezuela (Feuerer 2005). In Brazil it was recorded to
MG (Hale 1976, Ribeiro 1998), MS (Osorio 1992b),
MT (Hale 1976), PR (Osorio 1977b, Eliasaro 2001,
Eliasaro & Adler 2000, Eliasaro & Donha 2003), RJ
(Hale 1976), RS (Spielmann 2006), SC (Hale 1976)
and SP (Hale 1976, Marcelli 1991).
Specimens examined: BRAZIL. rio GrandE do suL:
Boqueirão do Leão, Linha Sinimbuzinho, Perau da
Nega, 29°20′02.2″S, 52°26′27.9″W, 440 m alt., in
the stream’s margin and close to the street, in a little
shaded place, 24-II-2004, A.A. Spielmann & L.S.
Canêz 1288 (SP); idem, 29°18′81.1″S, 52°26′88.5″W,
on the forest border, 7-IX-2000, L.S. Canêz & A.A.
Spielmann 1384 (HCB); Herveiras, near to “Balneário
Tio Juba”, shaded place, 6-II-2004, A.A. Spielmann
& L.S. Canêz 690 (SP); Sinimbu, Cava Funda,
29°27’33.4”S, 52°31′05.1″W, 520 m alt., open place,
5-I-2004, A.A. Spielmann & L.S. Canêz 665 (SP), A.A.
Spielmann & L.S. Canêz 666 (SP), A.A. Spielmann &
L.S. Canêz 687 (SP), A.A. Spielmann & L.S. Canêz
1290 (SP), A.A. Spielmann & L.S. Canêz 1294
(SP), A.A. Spielmann & L.S. Canêz 1296 (SP), A.A.
Spielmann & L.S. Canêz 1300 (SP), A.A. Spielmann
& L.S. Canêz 1328 (SP).
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substances
ChLoroatranorin
without vegetative
propagules
sorediate
isidiate
P. cristobaliae
pustulate
lobulate
P. ventanicum
P. lobuliferum
P. laciniellum
CortiCaL
P. tandilensis
usniC aCid
P. laciniellum (trace)
P. subsumptum (trace)
P. lobuliferum (trace)
P. recipiendum (trace)
P. subcaperatum (trace)
without substanCEs
P. consors
P. pilosum
saLaziniC and
ConsaLaziniC aCids
P. cristobaliae
P. leucosemothetum
P. erubescens
P. reitzii
P. neotropicum
P. laciniellum
P. spinibarbe
P. subtinctorium
P. subcaperatum
P. subsumptum
P. ultralucens
P. ventanicum
P. lobuliferum
P. tandilensis
P. uruguensis
protoCEtrariC aCid
P. laciniellum (trace)
P. fumarprotocetraricum
P. ventanicum
P. tandilensis (trace)
fumarprotocetraric
acid
P. fumarprotocetraricum
stictic acid and allied
P. muelleri
norstictic acid
Loxodin
P. kwalense
P. cristobaliae
P. conferendum
P. recipiendum
norLobaridonE
P. kwalense
P. haitiense
P. subtinctorium
P. cristobaliae
P. conferendum
P. haitiense
P. recipiendum
P. larense
P. subtinctorium
P. reitzii
P. subsumptum (trace)
lecanoric acid
P. lobuliferum (trace)
P. melanochaetum
medullar
GyrophoriC aCid
virEnsiC aCid
lichexanthone
P. ventanicum
P. tandilensis
P. larense
P. spinibarbe
P. ultralucens
Hoehnea 36(4): 551-595, 2009
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Table 1. Reproductive structures and chemistry in Canomaculina sensu Elix (1997) [atranorin is always present in the cortex].
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Parmotrema consors is characterized by the
absence of vegetative propagules and medulla
containing only an unidentiied fatty acid (negative
reactions). The similar P. pilosum (Stizenb.) Krog
& Swinscow produces soredia, being regarded the
sorediate pair of P. consors (Hale 1976).
In the specimens here examined the rhizines
range from abundant and covering the entire surface,
to frequent, sometimes leaving some parts with few
rhizines. As to the dimorphism, it ranges from sharply
evident in some specimens to hardly distinguishable
in others.
Parmotrema muelleri (Vainio) Blanco, Crespo,
Divakar, Elix & Lumbsch, Mycologia 97(1): 157.
2005 ≡ Parmelia muelleri Vainio, Acta Societatis
pro Fauna et Flora Fennica 7(1): 49. 1890 ≡
Canomaculina muelleri (Vainio) Elix & Hale,
Mycotaxon 29: 240. 1987. Type: BRAZIL. Minas
GErais: Sítio (nowadays Antônio Carlos), Vainio in
Lichenes Brasiliensis exsiccati n° 948 (lectotype
selected by Hale 1976, TUR, Vainio herbarium n°
2677, isolectotypes BM, FH, M, UPS), ide Hale
(1976).
Figure 25
Thallus greenish gray to brownish in herbarium,
lobate, adnate, subcoriaceous, corticicolous or
saxicolous, 11.5-15.0 cm broad. Lobes irregularly
branched, laterally overlapping, 1-11 mm wide,
surface smooth to scrobiculate, becoming rugose
and cracked in the center; margin smooth to
sublacinulate. Maculae distinct, punctiform to
irregular, laminal. Cilia black, simple to furcated or
irregularly branched, robust to spiky, 0.20-1.20 ×
0.04-0.20 mm, usually downwards turned, frequent,
present mainly on the crenae axils. Pustules and
isidia absent. Soralia capitate, often coalescing,
laminal or frequently reaching the margin; soredia
granular. Medulla white. Lower surface black,
lustrous, rugose; marginal zone brown, 1.0-3.0 mm
wide, with rhizines or nude area up to 1.0 mm wide,
lustrous, with attenuate limit, rugose and papillate.
Rhizines black, simple to irregularly branched, 0.102.00 × 0.02-0.10 mm, abundant, evenly distributed.
Apothecia absent [according to Vainio (1890), 2–5
mm in diameter, cupulate or subpeltate, subssessile,
disc testaceous, margin very thin, subentire or
sorediate; hymenium ca. 80 µm; ascospores shortellipsoid to ellipsoid or subglobose, 11-14 ×
(7-) 8-10 µm]. Pycnidia laminal to submarginal,
30518004 miolo.indd 557
557
conspicuous, usually with prominent margin, ostiole
black; conidia iliform, 10-15 × ca. 1 µm.
Color tests: cortex K+ yellow, UV-; medulla K+
yellow, C-, KC-, P+ orange, UV-.
TLC: atranorin, stictic acid, cryptostictic acid, and an
unidentiied substance with Rf 48 in solvent C.
Distribution: Asia (Awasthi 1976), North and South
America (Hale 1976). In South America it is known
to Argentina (Hale 1976, Calvelo & Liberatore 2002),
Brazil (Zahlbruckner 1930, Hale 1976, Marcelli
2004), Peru (Hale 1976), Uruguay (Osorio 1992a) and
Venezuela (Hale 1976). In Brazil it was recorded to
MG (Hale 1976, Ribeiro 1998), MS (Osorio 1992b),
PR (Osorio 1977a, Eliasaro 2001, Eliasaro & Adler
2000, Eliasaro & Donha 2003), RS (Spielmann 2006)
and SP (Jungbluth 2006).
Specimens examined: BRAZIL. rio GrandE do suL:
Herveiras, near to “Balneário Tio Juba”, shaded place,
6-II-2004, A.A. Spielmann & L.S. Canêz 685 (SP);
Sinimbu, Cava Funda, 29°27’33.4”S, 52°31’05.1”W,
520 m alt., open place, 5-I-2004, A.A. Spielmann &
L.S. Canêz 1298 (SP).
Parmotrema muelleri is recognizable by the capitate,
laminal soralia and medulla with stictic acid (K+
yellow, P+ orange). The sorediate P. pilosum is
destitute of demonstrable medullar substances
(Hale 1976). Another similar canomaculinoid
lichen is P. kwalense (Krog & Swinscow) Krog
& Swinscow, but it has isidia and in addition to
stictic acid, norstictic acid is present (Swinscow
& Krog 1988).
In the specimen A.A. Spielmann & L.S. Canêz
1298 the rhizines apparently have different sizes, but
they surely are stages of development of the same
type of rhizines, i.e., they’re not dimorphic. This
conirms the observations of Hale (1976).
Parmotrema recipiendum (Nyl.) Hale, Phytologia
28(4): 338. 1974 ≡ Parmelia recipienda Nyl., Flora
68(24): 609. 1885 ≡ Rimeliella recipienda (Nyl.)
Kurok., Annals of the Tsukuba Botanical Garden
10: 7. 1991 ≡ Canomaculina recipienda (Nyl.)
Elix, Mycotaxon 65: 477. 1997. Type: BRAZIL.
s.l., s.c. (holotype H-NYL35212), ide Kurokawa
(1991a).
Figure 29
08/04/10 11:19
558
Hoehnea 36(4): 551-595, 2009
Thallus pale grey or brownish in herbarium,
lobate, subcoriaceous, loosely adnate, saxicolous,
8.5-14.0 cm broad. Lobes irregularly branched,
laterally overlapping, 5-20 mm wide, surface smooth
to scrobiculate, lustrous, becoming scrobiculate,
rugose or cracked in the thallus center; apical zone
plane to subconcave; lateral marginal zone undulate,
plane to ascendant and revolute or involute, margin
crenate to incise-crenate (sublacinulate). Lacinules,
pustules, soredia and isidia absent. Maculae distinct,
punctiform to irregular, laminal. Cilia black, usually
simple, sometimes furcated, irregularly branched or
squarrose, 0.10-2.00 × 0.04-0.10 mm, abundant,
evenly distributed. Medulla white. Lower surface
dark brown, lustrous, smooth to rugose or papillate,
sometimes veined; marginal zone brown to pale
brown, sometimes white variegated, lustrous to
sublustrous, rhizinate or with small rhizines in
development, rarely nude (up to 0.5 mm wide),
without a sharp limit, smooth, rugose or papillate.
Rhizines usually black, sometimes dark brown,
dimorphic, the short ones simple to squarrose or
irregularly branched, raised or curly, 0.10-0.50
× 0.01-0.04 mm, abundant, evenly distributed,
the larger ones simple to furcated or irregularly
branched, 0.50-2.50 × 0.04-0.30 mm, frequent,
dispersed in groups. Apothecia urceolate, 5-14 mm
in diameter, stipitate, laminal to submarginal, margin
smooth to crenate, or sometimes dentate or shortdentate, amphithecium maculate, smooth, disc
brown, epruinose, perforate; epithecium 7-10 µm;
hymenium 55-65 µm; subhymenium 13-20 µm;
ascospores ellipsoid to ovoid, 12.5-15.0 × 6.5-9.0
µm, episporium 0.5–1.0 µm. Pycnidia submarginal
to laminal, conspicuous, usually with prominent
margin, ostiole black; conidia iliform straight or
arcuate, 10-16 × ca. 1 µm.
Color tests: cortex K+ yellow, UV-; medulla K-, C-,
KC+ rose, P-, UV-.
TLC: atranorin, norlobaridone and loxodin.
Distribution: Oceania (Kurokawa 1991a, Elix 1994)
and South America (Kurokawa 1991a). In South
America it is known to Argentina (Hale 1965,
Kurokawa 1991a, Calvelo & Liberatore 2002), Brazil
(Marcelli 2004), Paraguay (Lynge 1914, as Parmelia
annae Lynge; Hale 1965), Peru (Hale 1965, Kurokawa
1991a) and Uruguay (Osorio 1980). In Brazil it was
recorded to MG (Hale 1965), MT (Lynge 1914, as
30518004 miolo.indd 558
Parmelia annae; Kurokawa 1991a), PR (Kurokawa
1991a, Eliasaro 2001, Eliasaro & Donha 2003), RJ
(Kurokawa 1991a), RS (Spielmann 2006) and SP
(Hale 1965, Jungbluth 2006).
Specimens examined: BRAZIL. rio GrandE do suL:
Sinimbu, Cava Funda, 29°27′33.4″S, 52°31’05.1”W,
520 m alt., 5-I-2004, A.A. Spielmann & L.S. Canêz 696
(SP); Sobradinho, margin of highway RST-481, near
the crossroads, 29°24’20.2”S, 53°01’25.9”W, 375 m
alt., 17-VII-2003, A.A. Spielmann 352 (SP).
Parmotrema recipiendum is distinguished by
the lack of vegetative propagules and the presence
of norlobaridone and loxodin (medulla K–, KC+
reddish). Parmotrema subcaperatum (Kremp.) Hale
is very similar morphologically, so much that the
two species were already synonymous (Hale 1965).
The last one, however, has salazinic and consalazinic
acids (medulla K+ yellow → blood red). On the
other hand, Parmotrema cristobaliae (Ferraro &
Elix) Blanco, Crespo, Divakar, Elix & Lumbsch and
P. erubescens (Stirton) Krog & Swinscow have in
medulla both norlobaridone and loxodin as well as
salazinic acid (Krog & Swinscow 1981, Ferraro &
Elix 1993). Finally, Parmotrema conferendum has
the same chemical pattern of P. recipienda, but is a
sorediate species.
The specimen A.A. Spielmann 352 (SP) has fertile
lobes with several submarginal, grouped apothecia
(igure 29), very like the holotype photograph of
P. recipiendum (Kurokawa 1991a: igure 2). All
specimens were collected on the roadsides in open
places.
Parmotrema subcaperatum (Kremp.) Hale, Phytologia
28(4): 339. 1974 ≡ Parmelia subcaperata Kremp.,
Vidensk. Meddel. Dansk Naturhist. Foren.
Kjøbenhavn. 25: 10. 1874 ≡ Rimeliella subcaperata
(Kremp.) Kurok., Annals of the Tsukuba Botanical
Garden 10: 7. 1991 ≡ Canomaculina subcaperata
(Nyl.) Elix, Mycotaxon 65: 477. 1997. Type:
BRAZIL. Minas GErais: Caeté, Serra da Piedade,
Lagoa Santa [sic?], E. Warming 297 (holotype M),
ide Kurokawa (1991a).
Figure 34
Thallus greenish gray to brownish in herbarium,
lobate, subcoriaceous, loosely adnate, saxicolous,
6-12 cm broad. Lobes irregularly branched, laterally
overlapping to crowded, 4–20 mm wide, surface
smooth to rugose and scrobiculate, lustrous, becoming
08/04/10 11:19
A.A. Spielmann & M.P. Marcelli: Parmotrema from central Rio Grande do Sul State, Brazil
rugose and cracked in the center; apical zone plane to
subconcave, margin plane to ascendant or revolute,
regular to undulate, crenate to sublacinulate; lateral
marginal zone usually ascendant, regular to undulate,
sometimes involute, margin crenate to lacinulate.
Lacinules simple to furcated or dichotomous, 0.52.5 × 0.5-1.5 mm, plane, apex acute to rounded or
truncate, full dispersed in the thallus, lower surface
brown or white variegated; maculae weak to distinct,
punctiform to irregular, laminal. Cilia black, simple
to furcated, sometimes dichotomous or irregularly
branched, 0.20-1.20 × 0.04-0.10 mm, abundant,
evenly distributed. Pustules, soredia and isidia
absent. Medulla white. Lower surface dark brown
to black-brown, especially in the center, lustrous,
rugose; marginal zone brown, sublustrous, 1-2 mm
wide, with rhizines or rarely nude up to 1 mm wide,
limit absent, smooth to rugose or sometimes veined.
Rhizines black, dimorphic, the short ones 0.10-0.50
× 0.01-0.03 mm, straight or curly, simple, abundant,
evenly distributed, the larger ones 0.40-3.00 ×
0.05-0.30 mm, simple to furcated or irregularly
branched, frequent. Apothecia urceolate, 1-7 mm
in diameter, stipitate, laminal to submarginal,
margin smooth to crenate or incise, amphithecium
smooth, disc epruinose, perforate; epithecium 7-10
µm; hymenium 40-50 µm; subhymenium 13–20
µm; ascospores ellipsoid, 11-14 × 7.5-9.0 µm,
episporium 0.5-1.0 µm. Pycnidia submarginal
to laminal, conspicuous, usually with prominent
margin, ostiole black; conidia bacilliform to iliform,
straight or arcuate, 8.5-15.0 × ca. 1.0 µm.
559
(SP); idem, 29°27’33.4”S, 52°31’05.1”W, 520 m alt.,
open place, 5-I-2004, A.A. Spielmann & L.S. Canêz
710 (SP), A.A. Spielmann & L.S. Canêz 976 (SP), A.A.
Spielmann & L.S. Canêz 1295 (SP).
Parmotrema subcaperatum is distinguished by
the absence of vegetative propagules, medulla with
salazinic (K+ yellow → red) and the dark brown
lower surface. The similar P. recipiendum presents
just norlobaridone and loxodin (medulla K–, KC+
reddish) while P. reparatum (Stirton) Blanco, Crespo,
Divakar, Elix & Lumbsch has a distinct black lower
surface (Kurokawa 2001).
Kurokawa (1991a) and Fleig (1997) reported
the presence of usnic acid in P. subcaperatum. This
substance was not found in this study or by Eliasaro
(2001) to specimens from Paraná State.
It’s interesting to note that both Hale (1965),
Kurokawa (1991a) and Fleig (1997) regarded
Parmelia imperforata Nyl. as synonym of P.
subcaperatum. However, according to Zahlbruckner
(1909), P. imperforata (as suggested by the name)
has imperforate apothecia, and conidia 10–12 µm
long, while P. subcaperatum has perforate apothecia
slightly longer conidia, 8.5–15 µm long.
Actually P. subcaperatum is part of a complex of
species, which is being studied through the types and
will be published elsewhere (Spielmann & Marcelli,
unpublished data).
Distribution: Oceania and South America (Kurokawa
1991a). In South America it is known to Argentina
(Calvelo & Liberatore 2002), Brazil (Zahlbruckner
1930, Kurokawa 1991a, Marcelli 2004), Paraguay
(Osorio 1970a), Uruguay (Osorio 1972) and Venezuela
(Vareschi 1973). In Brazil it was recorded to MG
(Kurokawa 1991a), MS (Osorio 1992b), PR (Osorio
1977b, Kurokawa 1991a), RJ (Kurokawa 1991a), RS
(Spielmann 2006) and SP (Kurokawa 1991a).
Group II – Parmotrema-like lichens (Species
without dimorphic rhizines and rarely maculate, with
a broad naked rim below, belonging to Parmotrema
s. str.)
Parmotrema s. str. is characterized by the
generally broad and loosely adnate to adnate thalli,
relatively wide lobes, lower surface often with a
wide bare zone along the margins and rhizines from
simple to furcated or irregularly branched, but never
dimorphic (Elix 1993).
To facilitate the comparison among the species
of Parmotrema s. str. found, the main morphological
and chemical data were summarized in the table 2.
In some species of Parmotrema the rhizines reach
the margin, mainly in the form of “rhizinal papillae”
(Awasthi 1976) or “rhizines in development” (Fleig
1997). Of the species here dealt with, Parmotrema
internexum shows this feature.
Specimens examined: BRAZIL. rio GrandE do suL:
Sinimbu, Cava Funda, 29°27’41.8”S, 52°31’11.7”W,
500 m alt., open place, 4-II-2003, A.A. Spielmann 74
Parmotrema alidactylatum Estrabou & Adler,
Mycotaxon 66: 132. 1998. Type: ARGENTINA.
C órdoba : Departamento Sobremonte, Cerro
Color tests: cortex K+ yellow, UV-; medulla K+
yellow → red, C+ slowly yellowish, KC+ orange, P+
yellow → orange, UV–.
TLC: atranorin, salazinic and consalazinic acids.
30518004 miolo.indd 559
08/04/10 11:19
WITHOUT
VEGETATIVE
PROPAGULES
ATRANORIN
ALECTORONIC AND
α-COLLATOLIC ACIDS
SOREDIATE
P. mordenii
P. subrugatum
P. wainioi
P. hypomiltoides
P. rampoddense
SKYRIN
P. hypomiltoides
P. rampoddense
ANTHRAQUINONE
P. hypomiltoides
CAPERATIC ACID
P. mordenii
P. praesorediosum
PRAESOREDIOSIC ACID
P. praesorediosum
PROTOPRAESOREDIOSIC
ISIDIATE
P. melanothrix
PROTOLICHESTERINIC ACID
SALAZINIC AND CONSALAZINIC ACIDS
P. mantiqueirense
STICTIC ACID AND ALLIED
P. eciliatum
PUSTULATE
P. alidactylatum
P. mellissii
P. mordenii
P. alidactylatum
P. mordenii
P. praesorediosum
P. alidactylatum
P. spinibarbe
P. perlatum
NORLOBARIDONE
P. internexum
P. bangii
P. internexum
LECANORIC ACID
P. austrosinense
GYROPHORIC ACID
P. indicum
P. sancti-angeli
P. tinctorum
P. lavomedullosum
SECALONIC ACID A
P. lavomedullosum
LICHEXANTHONE
P. spinibarbe
Hoehnea 36(4): 551-595, 2009
MEDULLAR SUBSTANCES
560
30518004 miolo.indd 560
Table 2. Morphological and chemical features in the species of Parmotrema s. str. found (atranorin is full present in the cortex of all species).
08/04/10 11:19
A.A. Spielmann & M.P. Marcelli: Parmotrema from central Rio Grande do Sul State, Brazil
Colorado, 30º08’S, 64º54’W, on granite, V-1995,
Estrabou (holotype BAFC37877, isotypes BM,
CANB, H, O, S, TUR, Estrabou Herbarium 648),
ide Estrabou & Adler (1998).
Figure 1
Thallus pale gray, lobate, membranaceous,
loosely adnate, saxicolous, 10 cm broad. Lobes
irregularly branched, laterally overlapping, 3.5–8.0
mm wide, surface smooth to lightly rugose, opaque
to sublustrous, becoming rugose and cracked; apical
zone subconcave, margin undulate, entire to broadly
crenate, plane or slightly involute; lateral marginal
zone undulate, plane or ascendant and involute,
margin entire to crenate. Lacinules, maculae, cilia
and isidia absent. Pustules simple, short-claviform
or irregular, marginal, submarginal or laminal, more
abundant in the proximal areas, with a lax medulla,
becoming sorediose or not, arising from wrinkles
or from capitate structures, often starting as black
points in the thallus. Soredia granular, coming
from pustules. Medulla white. Lower surface black,
lustrous, smooth or more often rugose; marginal
zone pale brown or ochraceous, nude or with few
rhizines, smooth to papillate or sometimes rugose,
opaque to sublustrous, with a sharp limit, 2–4 mm
wide. Rhizines black or concolor to the marginal
zone, simple or branched, sometimes with the apex
latted or coalescing in the distal areas, 0.20–0.70
× 0.05–0.10 mm, dispersed in groups, frequent.
Apothecia and pycnidia unknown.
Color tests: cortex K+ yellow, UV–; medulla K+
yellow, C–, KC–, P–, UV–.
TLC: atranorin (cortex and medulla), protolichesterinic
acid and protopraesorediosic acid.
Distribution: Argentina (Calvelo & Liberatore 2002).
Specimens examined: BRAZIL. rio GrandE do suL:
Boqueirão do Leão, Linha Sinimbuzinho, Perau da
Nega, 29°20’02.2”S, 52°26’27.9”W, 440 m alt.,
left side of the stream, open place, 23-II-2004, A.A.
Spielmann & L.S. Canêz 1205 (SP).
Parmotrema alidactylatum is characterized by
the adnate and eciliate thallus with pustules, and the
presence of atranorin in the medulla (K+ yellow).
Parmotrema soredioaliphaticum Estrabou & Adler
is very close, but its pustules become sorediose
(Estrabou & Adler 1998). This subtle difference could
30518004 miolo.indd 561
561
place P. soredioaliphaticum in the synonymy of P.
alidactylatum (a hypothesis admitted by Estrabou &
Adler 1998). In fact, the only specimen described here
show some sorediate pustules. Additional collections
from Argentina and Brazil, together with the study of
the types, could contribute to this question.
We are naming here the structures of this lichen
as pustules, while in the original concept Estrabou &
Adler (1998) called them dactyls. These terms are not
clearly delimited in the current literature, and in this
respect, there are two other species morphologically
close to P. alidactylatum: P. tsavoense (Krog &
Swinscow) Krog & Swinscow and P. dactylosum
Fleig. The irst one has physodic acid (medulla KC+
purple), oxyphysodic acid, and iliform conidia with
18–20 µm long (Krog & Swinscow 1981, Swinscow
& Krog 1988), while P. dactylosum produces
caperatic acid and unidentiied fatty acids, the
conidia being sublageniform to almost bifusiform,
5.0–7.5 µm long (Fleig 1999).
Parmotrema austrosinense (Zahlbr.) Hale, Phytologia
28(4): 335. 1974 ≡ Parmelia austrosinensis
Zahlbr., Symbolae Sinicae 3: 192. 1930. Type:
CHINA. K wEitsChou : Gwanyinschen near
Guiyang, Setschwan, Handel-Mazzetti 10580
(lectotype BPI, selected by Hale 1959, isosyntypes
BPI), ide Hale (1959).
Figure 2
Thallus pale greenish gray, lobate, loosely
adnate, membranaceous to subcoriaceous in some
places, saxicolous, 11 cm broad. Lobes irregularly
branched, laterally overlapping, 3.0–7.5(–9.0)
mm wide, surface smooth, opaque to sublustrous,
becoming rugose and cracked in the center; apical
zone plane to subconcave, margin little to strongly
undulate, usually ascendant, smooth to crenate;
lateral marginal zone little or strongly undulate,
ascendant, margin entire. Lacinules, pustules, isidia
and cilia absent. Maculae weak. Soralia white or
with black points, marginal, linear interrupted,
lightly wider in the older parts; soredia farinose.
Medulla white. Lower surface black, opaque,
rugose; marginal zone brown or white variegated,
lustrous, 2–10 mm wide, with weak to sharp limit,
smooth to usually rugose. Rhizines black to concolor
with the lower surface, usually simple, 0.2–0.5 ×
0.1–0.3 mm, few, dispersed in groups. Apothecia
absent [according with Hale (1965), substipitate,
to 10 mm. in diameter, amphithecium maculate,
08/04/10 11:19
562
Hoehnea 36(4): 551-595, 2009
sparsely sorediate, disc perforate; hymenium 75 µm;
ascospores 10–16 × 6–10 µm, episporium 1,5 µm].
Pycnidia submarginal, conspicuous, with or without
prominent margin, ostiole black; conidia iliform,
(11.0–) 14.0–16.0 (–17.5) × ca. 0.5 µm.
Color tests: cortex K+ yellow, UV–; medulla K–, C+
red, KC+ red, P–, UV–.
TLC: atranorin, lecanoric acid.
Distribution: Oceania (Feuerer 2005), Australia (Hale
1965), Asia (Zahlbruckner 1932, Hale 1965), Europe
(Hale 1965), Africa (Hale 1965, Swinscow & Krog
1988) and Americas (Hale 1965). In South America
it is known to Argentina (Hale 1965, Calvelo &
Liberatore 2002), Brazil (Marcelli 2004), Colombia
(Hale 1965), Paraguay (Hale 1959), Uruguay (Hale
1965, Osorio 1992a) and Venezuela (Hale 1965,
Vareschi 1973); in Brazil it was recorded to MG
(Ribeiro 1998), MS (Osorio 1992b), PR (Eliasaro
2001), RS (Spielmann 2006) and SP (Marcelli 1991,
Ribeiro 1998).
Specimens examined: BRAZIL. rio GrandE do suL:
Sinimbu, Cava Funda, 29°27’41.8”S, 52°31’11.7”W,
500 m alt., 5-I-2004, A.A. Spielmann & L.S. Canêz
1088 (SP).
Parmotrema austrosinense is distinguished by
the linear soralia, the lacking cilia, and the presence
of lecanoric acid (medulla C+ red) and atranorin
(cortex greenish gray, K+ yellow). The white
variegated lower surface also seems to be a constant
feature, agreeing with the literature (Hale 1965,
1979, Krog & Swinscow 1981, Hale & Cole 1988,
Elix 1994, Fleig 1997, Brodo et al. 2001).
Parmotrema defectum (Hale) Hale share the
above presented features, but has an adnate to strongly
appressed thallus, and lightly shorter conidia, 10–12
µm long (Krog & Swinscow 1981) [yet Swinscow
& Krog (1988) gave a different size: 16–20 (24) µm
long]. It is the sorediate counterpart of P. soyauxii
(Müll. Arg.) Hale, while P. austrosinense is the
counterpart of P. andinum (Müll. Arg.) Hale (Hale
1965, Swinscow & Krog 1988).
Two other similar sorediate species, P. cooperi
(Steiner & Zahlbr.) Sérus. and P. sancti-angeli
(Lynge) Hale, can be easily identiied by the presence
of marginal cilia (Hale 1965).
The illustrations of P. austrosinense in Tavares
(1945, as Parmelia meridionalis), Awasthi (1976),
30518004 miolo.indd 562
Hale (1979), Hale & Cole (1988) and Brodo et al.
(2001) are very alike the specimen here studied.
Notwithstanding, the picture from Swinscow & Krog
(1988) seems to suggest the presence of capitate,
sorediate structures.
It is interesting to note that Hale (1965) and
Awasthi (1976) asserted that P. austrosinense has
“widely perforated” apothecia, while Krog &
Swinscow (1981), Elix (1994) and Nash & Elix
(2002a) stated that the apothecia vary from perforate
to imperforate, unfortunately not specifying if
in the same thalli or not. Species pairs in which
one counterpart has perforate apothecia and the
other not are, as far as we are aware, unreported in
Parmotrema. If both P. austrosinense and P. andinum
present perforate apothecia, then the species pair is
well deined. Nevertheless, if a “hidden” species,
very similar to P. austrosinense, with imperforate
apothecia, in fact exist, its counterpart would be
another one, for example, some of the specimens
reported by Awasthi (1976) as Parmelia andina,
with apothecia imperforate or perforate in the center.
Moreover, there are differences in the soralia and
soredia between the authors that describe perforate
and imperforate apothecia. Since there are several
accepted synonyms in both P. austrosinense and P.
andinum, only a revisionary work could solve this
question, because more than one taxon seems to be
implied.
The typiication of this species is also a little
confused. Firstly, Hale (1965) asserted that the
lectotype is located in WU, and the “isotypes” in
BPI and W. Elix (1994), Fleig (1997) and Eliasaro
(2001) wrote “isolectotypes” instead of isotypes. On
the other hand, Fleig (1997) stated that the lectotype
is in US, not BPI. As some authors (e.g. Elix 1994,
Eliasaro 2001) used “ide Hale 1965”, and Hale
(1965) sends back to his previous paper, we are
here using the data from Hale (1959), who made the
typiication.
Ribeiro (1998) described the rhizines of
Parmotrema austrosinense as “dimorphic, black,
simple and dichotomous”. With this features, his
lichen is more properly a Canomaculina, and
deserves further conirmation.
Parmotrema bangii (Vainio) Hale, Phytologia
28(4): 335. 1974 ≡ Parmelia bangii Vainio, in
Schmidt, Botanisk Tidskrift 29: 104. 1909. Type:
BOLIVIA. La Paz, 1891, M. Bang 13 (holotype
H-NYL35500), ide Hale (1965).
08/04/10 11:19
A.A. Spielmann & M.P. Marcelli: Parmotrema from central Rio Grande do Sul State, Brazil
Figure 3
Thallus greenish gray, lobate, membranaceous,
loosely adnate, corticicolous, 13.5 cm broad.
Lobes irregularly branched, crowded, 3.5–9.0 mm
wide, surface smooth to slightly rugose, opaque to
sublustrous, becoming reticulate cracked in the old
parts, cortex disintegrating in large submarginal
areas and forming sorediate pustules; apical zone
plane to subconcave, margin undulate, broadly
crenate to sublacinulate; lateral marginal zone
undulate, at irst imbricate, then ascendant and
involute, margin sublacinulate. Lacinules, maculae
and isidia absent. Cilia black, simple or rarely with
a branched, penicillate apex, 0.50–3.00 × 0.03-0.05
mm, abundant to frequent, present mainly in the lobe
axils. Pustules verruciform to capitate, laminal to
more commonly submarginal, often coalescing and
covering large areas, turning the margins undulate
and involute, fragmenting in soredia. Soralia
concolor to the thallus, pustular or formed from the
disintegration of the cortex, diffuse, coarse, usually
laminal; soredia granular. Medulla white. Lower
surface black, lustrous, smooth to rugose; marginal
zone pale or dark brown, sometimes white variegated
or beige under the sorediate areas, or even black in
some points, lustrous, nude or with sparse rhizines,
1.5–3.0 mm wide, with an attenuate or sharp limit,
smooth, rugose or veined. Rhizines black, simple
or branched, 0.50–2.00 × 0.03–0.05 mm, frequent,
evenly distributed. Apothecia absent [according to
Vainio (1909), up to 5 mm in diameter, cupulate,
subsessile, imperforate, excipulum smooth or with
a few soredioid or isidioid granules, ascospores
ellipsoid-oblong, 26–36 × 11–16 µm, and episporium
3–4 µm (Hale 1965)]. Pycnidia absent [according to
Vainio (1909), conidia sub-bifusiform, 5 µm long].
Color tests: cortex K+ yellow, UV–; medulla K+
yellow, C–, KC–, P+ slowly orange, UV–.
TLC: atranorin, stictic acid, constictic acid, cryptostictic
acid, hypoconstictic acid, hypostictic acid, and an
unidentiied substance similar to rhizocarpic acid.
Distribution: Africa (Østhagen & Krog 1976, Krog
& Swinscow 1981, Sérusiaux 1984, Swinscow &
Krog 1988) and South America (Hale 1965). In
South America it is known to Bolivia (Vainio 1909,
Zahlbruckner 1930, Hale 1965, Feuerer et al. 1998),
Brazil (Marcelli 2004), Colombia (Hale 1965) and
Venezuela (Feuerer 2005).
30518004 miolo.indd 563
563
Specimens examined: BRAZIL. rio GrandE do suL:
Boqueirão do Leão, Linha Sinimbuzinho, Perau da
Nega, 29°20’02.2”S, 52°26’27.9”W, 440 m alt., open
place, 23-II-2004, A.A. Spielmann & L.S. Canêz 700
(SP).
Parmotrema bangii is characterized by the
sorediose pustules and the granular soredia originated
from the cortex disintegration, together with the
presence of stictic acid (medulla K+ yellow, P+
slowly orange). Parmotrema perlatum is a similar
sorediate species, also with stictic acid, but in this
species the soralia are linear and marginal, never
forming pustules. P. madylinae Fletcher has pustules,
but instead of stictic acid, it has protocetraric acid
(medulla K–) and shorter ascospores, 23–25 × 14–15
µm, episporium 2 µm (Hale 1986).
Unfortunately, we were unable to ind pycnidia
in our specimen. However, it must be noted that
Krog & Swinscow (1981) reported the conidia as
being bacillar to iliform, 8–10 µm long, so different
from that described by Vainio (1909).
Parmotrema eciliatum (Nyl.) Hale, Phytologia 28(4):
336. 1974 ≡ Parmelia crinita Ach. var. eciliata
Nyl., Flora 52: 291. 1869. Type: MEXICO.
orizaba: Bourgeau s.n. (holotype H-NYL35295,
isotype P), ide Hale (1965).
Figures 13, 37
Thallus greenish gray, lobate, membranaceous
to subcoriaceous, loosely adnate, saxicolous,
12 cm broad. Lobes irregularly branched, laterally
overlapping to crowded, 4.0−8.0 mm wide, median
axis extended to subascendant, surface smooth to
irregular, opaque to lustrous, becoming rugose and
cracked in the center; apical zone subplane to concave,
margin undulate, subplane or ascendant, sometimes
involute, crenate to incise-crenate (sublacinulate);
lateral marginal zone undulate, suberect to erect,
subimbricate and involute. Lacinules, pustules,
soredia and isidia absent. Maculae present only close
to the apothecia. Cilia black, simple to furcated or
sometimes irregularly branched, 0.50–3.00 × 0.02–
0.10 mm, frequent, evenly distributed. Medulla white.
Lower surface black, lustrous, rugose; marginal zone
brown to beige, narrow (0.5–2.0 mm) or with large
parts white variegated (1–6 mm), lustrous to opaque,
nude, smooth to rugose, with a sharp limit. Rhizines
black, simple to rarely irregularly branched, 0.20-1.50
× 0.01–0.02 mm (with two well distinct sizes in
some lobes), sometimes twisted or coalescent, or
08/04/10 11:19
564
Hoehnea 36(4): 551-595, 2009
slightly prostrate and entangled, abundant, more
or less evenly distributed. Apothecia urceolate to
patent, 1–13 mm in diameter, stipitate, laminal
to submarginal, base constricted, rugose, margin
smooth to dentate, eciliate, amphithecium maculate,
strongly rugose, disc brown, imperforate, plane to
involute, entire or splitted; ascospores ellipsoid to
reniform, 22.5–29.0 (–31.5) × 12.5–16.5 (–19.0)
µm, episporium 2.5−3.5 µm. Pycnidia submarginal
to laminal, abundant, ostiole black; conidia bacillar
to iliform, rarely with a brief apical thickening, 5–9
× ca. 1 µm.
Color tests: cortex K+ yellow, UV–; medulla K+
yellow, C–, KC–, P+ orange, UV–.
TLC: atranorin, stictic acid, constictic acid,
cryptostictic acid and hypostictic acid.
Distribution: Asia (Kurokawa 1991b, Kurokawa &
Lai 2001, Feuerer 2005), Oceania (Elix 1994), Africa
(Hale 1965, Krog & Swinscow 1981, Swinscow &
Krog 1988), North and Central America (Hale 1965,
Feuerer 2005) and South America (Hale 1965). In
South America, it is known to Argentina (Lynge
1914, Hale 1965, Adler 1992, Calvelo & Liberatore
2002), Brazil (Hale 1965, Marcelli 2004) and Uruguay
(Osorio 1972, 1992a). In Brazil, it was recorded to
PR (Eliasaro & Donha 2003), RJ (Hale 1965) and RS
(Spielmann 2006).
Specimens examined: BRAZIL. rio GrandE do suL:
Sobradinho, margin of highway RST-481, near the
crossroads, 29°24’20.2”S, 53°01’25.9”W, 375 m alt.,
open place, 17-VII-2003, A.A. Spielmann 410 (SP).
Parmotrema eciliatum is distinguished by the
ciliate lobes, presence of stictic acid (medulla K+
yellow, P+ orange), apothecia imperforate, and the
absence of vegetative propagules. Parmotrema
aldabrense (C.W. Dodge) Hale, reported to Africa,
sometimes presents stictic acid as an accessory
substance to the norstictic acid (Medulla K+ yellow
→ red), his main substance. However, it has perforate
apothecia, maculae distinct, ascospores 13–18 ×
5–7 µm (Hale 1965) and iliform conidia, 12–15
µm long (Swinscow & Krog 1988). Parmotrema
preperforatum (W.L. Culb.) Hale, known in the
United States, also has norstictic acid and stictic
acid, together with perforate apothecia (Culberson
1973), but this is the only information we were
able to ind about this species, so that we don’t
30518004 miolo.indd 564
know more differences between P. aldabrense and
P. preperforatum, except that one is recorded to
Africa and the other to North America. Finally,
P. blanchetianum (Müll. Arg.) Kalb, has similar
morphology, but produces protocetraric acid instead
stictic acid. In the course of the present work we
noted that P. eciliatum could also be distinguished
by the cilia, usually furcated or irregularly branched
(igure 37).
Parmotrema flavomedullosum Hale, Mycotaxon
1(2): 110. 1974. Type: BRAZIL. santa Catarina:
Campos Novos, Colony Santa Catarina, saxicolous,
9-VIII-1963, R. Reitz & R.M. Klein 15051
(holotype US), ide Fleig (1997).
Figures 14, 40
Thallus pale grey, lobate, membranaceous
to subcoriaceous in some areas, loosely adnate,
corticicolous or saxicolous, 15.0-18.5 cm broad.
Lobes irregularly branched, laterally overlapping to
crowded, 4-20 mm wide, surface smooth, rugose or
with the cortex disintegrating in patches, sublustrous
to opaque, becoming rugose in the center; apical zone
plane to concave or involute, often curled or apically
straighten and, combining with the involution of the
lateral margins, assuming the aspect of a “T” when
the lobe is observed from the lower side (igure
40), margin entire to crenate; lateral marginal zone
undulate, plane to ascendant and involute, margin
entire to crenate or incise-crenate (sublacinulate).
Lacinules, maculae and isidia absent. Cilia black,
cylindrical or latted, simple to dichotomous,
0.40-3.00 × 0.05-0.10 mm, few, present on the
axillar crenae, more abundant in some areas than
others. Pustules developed from wrinkles, laminal
to marginal, present mainly in the apices and lobe
margins, breaking up in soredia. Soralia capitate to
extensive, marginal to submarginal, approaching
the lamina, usually developed from pustules, or
sometimes directly from the fragmentation of
the surface, commonly causing the “T” aspect of
the lobes; soredia from subgranular to granular,
sometimes isidioid and coalescing. Medulla yellow
to orange. Lower surface black, lustrous, smooth,
rugose or rugose-reticulate; marginal zone brown to
beige or chestnut, opaque to lustrous, 1.5-6.0 mm
wide, nude, limit from sharp to attenuate, smooth,
papillate, rugose to strongly rugose-reticulate or
with veins, usually with cracks. Rhizines black or
sometimes with a whitish apex, simple to furcated
08/04/10 11:19
A.A. Spielmann & M.P. Marcelli: Parmotrema from central Rio Grande do Sul State, Brazil
or irregularly branched, 0.20-3.00 × 0.02-0.10 mm,
few to frequent, dispersed in small groups. Apothecia
and pycnidia unknown.
Color tests: cortex K+ yellow, UV-; medulla K+
yellow, C+ orange, KC+ yellow, P- (fading the
pigment yellow), UV-.
TLC: atranorin, gyrophoric acid, secalonic acid A and
C and unidentiied substance of Rf 51 in solvent C.
Distribution: Known only to South America, where
it was reported to Argentina (Hale 1974c), Brazil
(Marcelli 2004), Paraguay and Venezuela (Hale 1974c).
In Brazil it was recorded to MG (Hale 1974c), MS
(Osorio 1992b), PR (Hale 1974c, Eliasaro 2001), SC
(Hale 1974c, Fleig 1997) and RS (Spielmann 2006).
Specimens examined: BRAZIL. r io G randE do
s uL : Boqueirão do Leão, Cascata do Gamelão,
29°18’13.7”S, 52°26’51.7”W, 500 m alt., on the steep
slope near the waterfalls, shaded place (inside the
forest), 1-II-2004, A.A. Spielmann & M.A. Sulzbacher
743 (SP); idem, in the right side of the stream, open
place, 1-II-2004, A.A. Spielmann & M.A. Sulzbacher
1037 (SP).
Parmotrema lavomedullosum is recognizable by
the ciliate lobes, pustules fragmenting in soredia and
the yellow to orange medulla with gyrophoric acid
and secalonic acids A and C (K+ yellow, C+ orange,
KC+ yellow). Some other species of Parmotrema
with yellow medulla can be distinguished as follow:
P. araucariarum (Zahlbr.) Hale has eciliate lobes
and the gyrophoric acid absent (Hale 1974c), P.
endosulphureum (Hillm.) Hale has eciliate lobes
and produces isidia instead of soredia (Hale 1974c).
Finally, Parmelia elabens Kurok., described in 1974
based on specimens from Paraná State, is regarded
a synonym of P. lavomedullosum by Fleig (1997),
Hale & DePriest (1999) and Eliasaro (2001).
Parmotrema hypomiltoides (Vainio) Kurok., Bulletin
of the Botanic Gardens of Toyama 3: 18. 1998 ≡
Parmelia hypomiltoides Vainio, Acta Societatis
pro Fauna et Flora Fennica 7: 35. 1890. Type:
BRAZIL. Minas GErais: Sítio (nowadays Antônio
Carlos), Vainio, Lichenes Braziliensis exsiccati s.n.
(holotype TUR–V2543), ide Fleig (1997).
565
irregularly branched, laterally overlapping, 2–17 mm
wide, median axis extended to subascendant,
surface smooth to scrobiculate, becoming rugose
and reticulate cracked in the old parts; apical zone
rounded, subconcave, margin usually ascendant
and involute, sometimes revolute, undulate, entire
to crenate or incise-crenate (sublacinulate); lateral
marginal zone undulate and usually involute over
the median axis and turning the lobes convex and
almost canaliculate, margin crenate to sublacinulate.
Lacinules, pustules and true isidia absent; maculae
weak to distinct, punctiform, laminal. Cilia black,
simple or rarely irregularly branched, 0.50–2.00
× 0.02–0.05 mm, from few to frequent, evenly
distributed. Soralia concolor to the thallus, capitate
or irregular, usually marginal and in the sublacinules
apices, sometimes laminal, often forming arbuscular
structures with sorediate apices (like a caulilower);
soredia granular or sometimes isidioid. Medulla
white with orange areas K+ red close to the soralia
or in the older parts. Lower surface black, lustrous,
smooth to slightly rugose or with veins, sometimes
papillate; marginal zone brown to dark brown, often
white or beige variegated, or completely black,
opaque to lustrous, 2–6 mm wide, nude, with a
sharp or attenuate limit, smooth to slightly rugose.
Rhizines black, simple to irregularly branched,
0.20–3.00 × 0.02–0.05 mm, frequent, dispersed in
groups. Apothecia absent [according to Fleig (1997),
submarginal, imperforate, urceolate to patent and
splitted when old, substipitate, with a wide base,
rugose, amphithecium rugose, sorediate; ascospores
18.0–23.0 × 9.0–12.5 µm, episporium 1.5–2.5
µm]. Pycnidia absent [according to Fleig (1997),
submarginal, sterile]; conidia unknown.
Color tests: cortex K+ yellow, UV–; medulla K–, C–,
KC+ rose → quickly orange, P–, UV+ greenish blue.
TLC: atranorin, alectoronic acid, α-collatolic acid,
skyrin and unidentiied anthraquinone of Rf 49 in
solvent C.
Figures 16-17
Distribution: Oceania (Louwhoff & Elix 1999) and
South America (Hale 1965). In South America it
is known to Brazil (Zahlbruckner 1930, Marcelli
2004) and Venezuela (Feuerer 2005). In Brazil it was
recorded to MG (Hale 1965), PE (Kurokawa & Moon
1998), RS (Spielmann 2006), SC (Fleig 1997) and SP
(Hale 1965, Ribeiro 1998).
Thallus pale gray, lobate, membranaceous,
loosely adnate, saxicolous, 7.5–11.0 cm broad. Lobes
do
30518004 miolo.indd 565
Specimens examined: BRAZIL. rio GrandE
suL: Boqueirão do Leão, Cascata do Gamelão,
08/04/10 11:19
566
Hoehnea 36(4): 551-595, 2009
29°18’13.7”S, 52°26’51.7”W, 500 m alt., on the steep
slope near the waterfalls, shaded place (inside the
forest), 1-II-2004, A.A. Spielmann & M.A. Sulzbacher
1164 (SP), A.A. Spielmann & M.A. Sulzbacher 1216
(SP), A.A. Spielmann & M.A. Sulzbacher 1261 (SP),
A.A. Spielmann & M.A. Sulzbacher 1262 (SP).
Parmotrema hypomiltoides is distinguished by
the production of soredia, often formed in arbuscular
structures (like a caulilower), the presence of
alectoronic and a-collatolic acid (medulla KC+ rose
→ quickly orange, UV+ greenish blue) and the orange
pigment K+ red (unidentiied anthraquinone) close to
the soralia and lobe apices. In P. rampoddense (Nyl.)
Hale the soralia are usually marginal and linear, and
the orange pigment K+ red (skyrin) is present only
close to the lower cortex, in the old parts. Finally, P.
mellissii (C.W. Dodge) Hale develops true isidia.
Fleig (1997) irst proposed the new combination
of Parmelia hypomiltoides in Parmotrema. At that
time, the status of nomenclatural propositions in
theses were still somewhat obscure in the St. Louis
Code (Greuter et al. 2003). Nevertheless, according to
the Vienna Code (McNeill et al. 2007), Fleig’s thesis
is not effectively published. So the following author
to propose the new combination, Kurokawa (1998,
March) must be credited. Interesting to note that
DePriest & Hale (1998, April-June) also proposed
this new combination, since Hale already recognized
this good species although not validly published.
Later (Hale & DePriest 1999) they correctly used
Kurokawa’s combination.
Parmotrema indicum Hale, Mycotaxon 5(2): 436.
1977. Type: INDIA. taMiL nadu: Kodaikanal,
forest below Silver Cascade, on rocks in stream,
elevation 1,800 m, M.E. Hale & P.G. Patwardhan
43874 (holotype US), ide Hale (1977).
Figures 18, 39
Thallus greenish gray or mineral gray, lobate,
loosely adnate, subcoriaceous, saxicolous, 20–28
cm broad. Lobes irregularly branched, laterally
overlapping, 4–20 (–25) mm wide, surface smooth,
opaque to lustrous, becoming rugose and cracked
or reticulate cracked in the center; apical zone
plane to subconcave, margin plane to undulate,
often ascendant, entire to crenate; lateral marginal
zone undulate to strongly undulate, subascendant
to ascendant, involute in the older parts, sometimes
revolute, margin entire to crenate. Lacinules, maculae,
pustules and isidia absent. Cilia simple to branched,
30518004 miolo.indd 566
0.50–3.50 (–4.50) × 0.05–0.15 mm, frequent to
abundant, evenly distributed. Soralia marginal,
rarely submarginal, linear-continuous or interrupted,
slightly wider when dense, sometimes orbicular and
laminal in the old parts; soredia granular, sometimes
dense and coalescing, sometimes darkened in the
central areas. Medulla white, in some points with
orange patches K+ wine red near the lower cortex.
Lower surface black, opaque or lustrous, papillate,
smooth or rugose; marginal zone brown, sometimes
ochraceous (especially in the older, sorediate parts),
black or brown and black variegated, nude, lustrous,
with an attenuate or sharp limit, smooth, papillate or
rugose, (2–) 4–9 mm wide. Rhizines black, simple,
0.20–2.00 (–4.00) × 0.05–0.15 mm, frequent to
abundant, dispersed in groups. Apothecia unknown.
Pycnidia submarginal, ostiole black; conidia iliform,
straight or arcuate, 8.0–12.5 × ca. 1.0 µm.
Color tests: cortex K+ yellow, UV–; medulla K–, C+
rose, KC+ rose, P–, UV–.
TLC: atranorin, gyrophoric acid.
Distribution: Africa (Krog & Swinscow 1981, as
Parmelia indoafra; Swinscow & Krog 1988), Asia
(Hale 1977), and South America, where it is known
to Brazil (Marcelli 2004); in Brazil it was recorded to
RS (Spielmann 2006).
Specimens examined: BRAZIL. rio GrandE do suL:
Boqueirão do Leão, Linha Sinimbuzinho, Perau da
Nega, 29°20’02.2”S, 52°26’27.9”W, 440 m alt., left
margin of a stream, shaded by a shrub and grasses,
23-II-2004, A.A. Spielmann & L.S. Canêz 931 (SP);
Sinimbu, Cava Funda, 29°27’41.8”S, 52°31’11.7”W,
500 m alt., open place, 12-II-2003, A.A. Spielmann
61 (SP).
Parmotrema indicum is characterized by the
robust thallus with ciliate lobes, marginal, linear
soralia, the presence of gyrophoric acid (medulla
C+ rose, KC+ rose) and iliform conidia, 8.0–12.5
× ca. 1.0 µm. P. sancti-angeli (Lynge) Hale has
sublageniform conidia, 5.0−7.5 × ca. 1.0 µm. P.
permutatum (Stirton) Hale presents a yellow lower
medulla (secalonic acid A) and P. rampoddense
produces alectoronic acid (medulla UV+ greenish
blue).
However, usually the thalli are not pycnidiate,
and so the conidia type is useless in the distinction of
P. indicum and P. sancti-angeli. During the present
08/04/10 11:19
A.A. Spielmann & M.P. Marcelli: Parmotrema from central Rio Grande do Sul State, Brazil
study, we realized other accessory differences between
these species. P. indicum often has furcated or more
branched cilia, also slightly thickened (0.05–0.15
mm wide), reminding the cilia from Canomaculina
(igure 39), wide marginal soralia, together with
orbicular and laminal ones in the older lobes, and
a robust, usually coriaceous and adnate thallus. In
P. sancti-angeli, the cilia are usually simple (rarely
furcated) and thinner (0.05–0.10 mm). the marginal
soralia are narrow and the orbicular, laminar ones
absent. The thallus is usually membranaceous, with
ascendant lobes
There is also the option to verify if the medulla
of one specimen has norlobaridone, present only in
P. indicum (Hale 1977, Krog & Swinscow 1981,
Swinscow & Krog 1998). Yet this substance was not
found in our specimens.
Although Fleig (1997) stated that P. indicum
occurs in North America, we were unable to conirm
this.
Parmotrema internexum (Nyl.) Hale ex DePriest &
B. Hale, Mycotaxon 67: 204. 1998 ≡ Parmelia
internexa Nyl., Flora 68(24): 609. 1885. Type:
BRAZIL. são pauLo: Serra near to Santos, 1844,
Weddel s.n. (holotype H-NYL35210), ide Fleig
(1997).
Figure 19
Thallus greenish gray, lobate, adnate,
membranaceous, saxicolous, 7–10 cm broad. Lobes
irregularly branched, laterally overlapping, 2.5–7.0
mm wide, surface smooth to lightly scrobiculate,
opaque to lustrous, becoming rugose and cracked
in the center; apical zone plane to lightly concave
or convex, plane to undulate; margin crenate or
sometimes dentate; lateral marginal zone plane and
imbricate to ascendant and involute, usually curling
in the axils, undulate, margin entire to crenate.
Lacinules, pustules and soralia absent. and maculae
absent (but sometimes with irregular white patches).
Cilia black, simple, few or rare, evenly distributed yet
more abundant in the lobe axils, narrow, 0.20–0.80 ×
0.02–0.10 mm. Isidia concolor to the thallus or with
black or brown apices, cylindrical, simple, granular,
branched or coralloid, erect, irm, more dense in
the ascendant areas, eciliate, laminal to marginal,
0.05–0.40 × 0.05–0.20 mm. Medulla white. Lower
surface black, lustrous, rugose; marginal zone brown
or sometimes white variegated, nude or with young
rhizines, lustrous, with an attenuate limit, smooth
30518004 miolo.indd 567
567
to subrugose or sometimes with veins, 0.7–3.5 mm
wide. Rhizines black, simple or branched, dense and
abundant, widely dispersed, 0.10–1.00 × 0.02–0.05
mm. Apothecia absent [according to Fleig (1997),
laminal, imperforate, 3–4 mm in diameter, urceolate,
substipitate, entire or splitted, amphithecium
isidiate, eciliate; ascospores 19.0–31.0 × 9.5–15.0
µm, episporium 3 µm]. Pycnidia rare, submarginal,
with black ostiole; conidia bacillar to slightly
sublageniform, 5.0–7.0 × ca. 1.0 µm.
Color tests: cortex K+ yellow, UV–; medulla K+
yellow, C–, KC+ rose-orange, P+ slowly orange, UV–.
TLC: atranorin, norlobaridone, stictic acid, constictic
acid, cryptostictic acid and hypoconstictic acid.
Distribution: Africa (Dodge 1959, as Parmelia
meiosperma), North America (Hale 1979, Esslinger
2008) and South America, where it is known to
Brazil (Zahlbruckner 1930, Marcelli 2004). In Brazil
it was recorded to MG (Ribeiro 1998), PR (Fleig
1997, Eliasaro 2001, Eliasaro & Donha 2003), RS
(Spielmann 2006), SC (Fleig 1997) and SP (Marcelli
1991, 1992, Fleig 1997).
Specimens examined: BRAZIL. rio GrandE do suL:
Boqueirão do Leão, Linha Sinimbuzinho, Perau da
Nega, 29°20’02.2”S, 52°26’27.9”W, 440 m alt.,
close to a stream, 24-II-2004, A.A. Spielmann & L.S.
Canêz 1252 (SP); Sinimbu, Cava Funda, 29°27′1.6″S,
52°31′02.9″W, 520 m alt., open place, 5-I-2004, A.A.
Spielmann & L.S. Canêz 1167 (SP).
Parmotrema internexum is characterized by
the laminal isidia, ciliate lobes and the presence
of norlobaridone (medulla KC+ rose-orange) and
stictic acid (medulla K+ yellow, P+ slowly orange).
Parmotrema crinitum (Ach.) M. Choisy, has a robust
thallus (8-25 cm broad), wider lobes (6–12 mm),
often ciliate isidia, and norlobaridone lacking (Hale
1965).
In solvent C, the norlobaridone was “masked”
inside the dot of stictic acid, but in solvent A the two
dots were sharply distinct. However, the presence of
this substance seems not universal, since it was not
detected by Eliasaro (2001) and, according to Fleig
(1997) and Eliasaro & Donha (2003), it was present
only in some specimens.
The synonymy of this species is unclear. Hale
(1974d) placed Parmelia internexa f. meiosperma
Hue in Pseudoparmelia Lynge. Fleig (1997) listed
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568
Hoehnea 36(4): 551-595, 2009
Parmelia internexa var. meiosperma Hue, P.
meiosperma (Hue) C.W. Dodge and P. catharinesis
Müll. Arg. f. isidiosa Müll. Arg. as synonyms of
Parmotrema internexum. However, in Hale &
DePriest (1999) the irst two names are synonyms of
Hypotrachyna bogotensis (Vainio) Hale and the last
one of Parmotrema crinitum (Ach.) M. Choisy.
Parmotrema mantiqueirense Hale, Bibliotheca
Lichenologica 38: 113. 1990. Type: BRAZIL.
Minas GErais: Serra da Mantiqueira, zwischen
1,880 m und der Gipfelregion des Pedra Redonda
in 2,000 m, 3-VII-1979, K. Kalb s.n. (holotype
Kalb herbarium, isotype US), ide Hale (1990).
Figure 20
Thallus pale gray, lobate, subcoriaceous,
fragile, loosely adnate, saxicolous, ca. 20 cm broad.
Lobes irregularly branched, laterally overlapping,
7.0−27.0 mm wide, surface smooth, lustrous,
reticulate cracked and slightly rugose in the older
parts, usually with several scars; apical zone
subconcave, undulate, ascendant to involute, margin
sublacinulate, margin crenate; lateral marginal zone
undulate, plane to ascendant, subimbricate, margin
entire to sublacinulate. Lacinules and maculae
absent. Cilia black, usually simple, sometimes
furcated, usually thickened and spiky, 0.50–3.50
× 0.05−0.10 mm, abundant, evenly distributed.
Pustules, soredia and isidia absent. Medulla white.
Lower surface black, lustrous, smooth, papillate,
with scars or cracks; marginal zone dark brown or
sometimes black, 2.0−9.0 mm wide, nude, lustrous,
smooth or papillate, having a wide area with young
rhizines, with a sharp limit. Rhizines black, simple
or sometimes furcated, rarely squarrose, narrow to
thickened, 0.30−3.00 × 0.02−0.10 mm, abundant,
dispersed in groups. Apothecia absent [according to
Eliasaro (2001), laminal, adnate, disc dark brown,
imperforate, 2–5 mm in diameter; ascospores 13–18
× 6–10 µm]. Pycnidia submarginal to rarely marginal,
ostiole black; conidia bacillar to iliform, 6.5−11.0 ×
ca. 1.0 µm.
Color tests: cortex K+ yellow, UV−; medulla K+
yellow → blood red, C+ yellow, KC+ orange, P+
yellow → orange, UV−.
TLC: atranorin, salazinic and consalazinic acids.
Distribution: known only to South America, where it
was reported to Brazil (Hale 1990, Marcelli 2004). In
30518004 miolo.indd 568
Brazil it was recorded to MG (Hale 1990, Fleig 1997,
Ribeiro 1998), PR (Fleig 1997, Eliasaro 2001) and RS
(Spielmann 2006).
Specimens examined: BRAZIL. rio GrandE do suL:
Barros Cassal, Nascente do Rio Pardo, 28°57’38.7”S,
52°38’53.9”W, 700 m alt., open place, close to a
stream, 27-I-2004, A.A. Spielmann et al. 938 (SP).
Parmotrema mantiqueirense is distinguished by
the wide lobes (7–27 mm), emaculate upper surface,
the presence of salazinic acid (medulla K+ yellow →
blood red, P+ yellow → orange) and the absence of
vegetative propagules. P. eurysacum (Hue) Hale has
subpalmate lacinules, often branched and longer cilia
(1.5–5.0 mm) and perforate apothecia.
Although Fleig (1997) asserted that P.
mantiqueirense occurs in North America, we were
unable to conirm this information.
Parmotrema melanothrix (Mont.) Hale, Phytologia
28(4): 337. 1974 ≡ Parmelia urceolata Eschw. var.
melanothrix Mont., Ann. Sci. Nat., Bot. ser. 2(2):
372. 1834. Type: BRAZIL. Gaudichaud 89 bis
(holotype P), ide Hale (1965).
Figure 21
Thallus olivaceous gray, lobate, subcoriaceous,
loosely adnate, saxicolous, 7.5 cm broad. Lobes
irregularly
branched,
laterally
overlapping,
(3.0−) 5.0−10.0 mm wide, median axis plane to
subascendant, surface smooth, lustrous, becoming
slightly rugose towards the center; apical zone
plane to undulate, margin crenate; lateral marginal
zone undulate, plane to ascendant, margin crenate.
Lacinules, pustules, soredia and isidia absent.
Maculae distinct, punctiform, laminal. Cilia black
or cupreous (then K+ purplish), simple or rarely
furcated, 1.00−4.00 × 0.05−0.10 mm, abundant,
evenly distributed. Medulla white. Lower surface
black, lustrous, veined; marginal zone beige or white
variegated, usually with brown patches, sometimes
with purplish patches, 4.0−8.0 (−15.0) mm, nude,
lustrous, smooth to rugose, with a sharp limit. Rhizines
black or brown, simple to irregularly branched,
narrow to thickened, 0.10−0.30 × 0.10−0.20 mm,
frequent, dispersed in groups, sometimes reaching
the margin. Apothecia urceolate, laminal, 1.0−2.0
mm in diameter, stipitate, with a wide base, margin
crenate, amphithecium strongly maculated, ciliate,
disc concave, epruinose, imperforate or rarely
perforate (one from seven apothecia seen), immature;
08/04/10 11:19
A.A. Spielmann & M.P. Marcelli: Parmotrema from central Rio Grande do Sul State, Brazil
hymenium little developed and ascospores absent
[according to Hale (1965), hymenium 80–100 µm;
ascospores 20–26 × 10–16 µm, episporium 2–3
µm]. Pycnidia submarginal or touching the margin,
conspicuous, without prominent margin, ostiole
black; conidia bacillar to iliform, 6.5−10.0 × ca. 0.5
µm.
Color tests: cortex K+ yellow, UV−; medulla K−, C−,
KC−, P−, UV−.
TLC: atranorin, protopraesorediosic acid.
Distribution: Oceania (Feuerer 2005), Africa (Réunion
Island) and South America (Hale 1965). In South
America it is known to Brazil (Marcelli 2004), Peru
(Feuerer 2005) and Venezuela (Vareschi 1962). In
Brazil it was recorded to MA (Hale 1965), MG (Lynge
1914, Hale 1965, Ribeiro 1998), MS (Osorio 1992b),
MT (Lynge 1914, Fleig 1997, Ribeiro 1998), PR (Hale
1965, Fleig 1997, Eliasaro 2001), RJ (Hale 1965),
RS (Spielmann 2006), SC (Fleig 1997) and SP (Hale
1965; Marcelli 1991, 1992, Ribeiro 1998).
Specimens examined: BRAZIL. rio GrandE do suL:
Boqueirão do Leão, Linha Sinimbuzinho, Perau da
Nega, 29°20’02.2”S, 52°26’27.9”W, 440 m alt.,
close to a stream, slightly shaded, 24-II-2004, A.A.
Spielmann & L.S. Canêz 1033 (SP).
Parmotrema melanothrix is distinguished by the
long, abundant cilia (1–4 mm), maculate upper cortex,
conidia bacillar to iliform, 6.5−10.0 µm long, just
protopraesorediosic acid in the medulla (spot tests
negative) and the absence of vegetative propagules.
P. catarinae Hale has gyrophoric acid (Hale 1986).
P. glaucocarpoides (Zahlbr.) Hale has sublageniform
conidia 5–6 µm long and protolichesterinic acid
(Krog & Swinscow (1981).
According to Hale (1965), Fleig (1997), Ribeiro
(1998) and Eliasaro (2001), the medullar substance
typical of P. melanothrix is protolichesterinic acid, yet
in the specimen here studied just protopraesorediosic
acid was found.
Parmotrema mellissii (C.W. Dodge) Hale, Phytologia
28(4): 337. 1974 ≡ Parmelia mellissii C.W. Dodge,
Annals of the Missouri Botanical Garden 46(1-2):
134. 1959. Type: AFRICA. Santa Helena, J.C.
Melliss 23 (holotype K, isotype US), ide Hale
(1965) and Fleig (1997).
Figures 22-23, 41
30518004 miolo.indd 569
569
Thallus greenish gray, lobate, membranaceous
to subcoriaceous, loosely adnate, saxicolous, 15
cm broad. Lobes irregularly branched, laterally
overlapping, (3.0−)4.0−10.0 mm wide, surface
smooth or cracked, opaque, becoming reticulate in
the center; apical zone plane to subconcave, undulate,
margin crenate; lateral marginal zone undulate,
ascendant and involute or subimbricate, margin
crenate. Lacinules, pustules and maculae absent.
Cilia black, simple or irregularly branched, 0.50−3.00
× 0.02−0.05 mm, abundant, evenly distributed.
Soredia originated from isidia. Isidia concolor to the
upper cortex, apices brown or black, usually simple
or with two to four branches, cylindrical or granular,
sometimes coarsely granular and coalescing,
rarely coralloid, 0.10–0.30 × 0.05–0.10 mm, often
raised from papillae, wrinkles or from the cortex
fragmentation, submarginal, laminal or marginal,
erect, irm or caducous, apex often ciliate, giving rise
to soredia or not. Medulla white, except for some
orange old areas (K+ wine red). Lower surface black,
opaque to lustrous, smooth to rugose; marginal zone
brown or rarely black, sometimes white variegated,
2.0−5.0 mm wide, nude, lustrous or sometimes
opaque, smooth to rugose, with a sharp or attenuate
limit. Rhizines black, simple or irregularly branched,
1.00−3.50 × 0.05−0.10 mm, abundant, dispersed in
groups. Apothecia and pycnidia unknown.
Color tests: cortex K+ yellow, UV−; medulla K−, C−,
KC+ rose → quickly orange, P−, UV+ greenish blue.
TLC: atranorin, alectoronic acid, α-collatolic acid
and skyrin.
Distribution: Oceania (Malcolm & Galloway 1997,
Louwhoff & Elix 1998, 1999, Kantvilas et al. 2002),
Asia (Hale 1965, Awasthi 1976, Kurokawa 1991b,
1993, Kurokawa & Lai 2001, Louwhoff & Elix
2002), Africa (Krog & Swinscow 1981, Swinscow
& Krog 1988), North America (Hale 1965, Nash &
Elix 2002a), Central America (Hale 1965) and South
America (Hale 1965). In South America it is known
to Brazil (Marcelli 2004), Chile (Feuerer 2005),
Colombia (Hale 1965), Guyana (Feuerer 2005) and
Uruguay (Osorio 1992a). In Brazil it was recorded to
MG (Hale 1965, Ribeiro 1998), PR (Eliasaro 2001),
RS (Spielmann 2006), SC (Fleig 1997) and SP (Pereira
& Marcelli 1989, Ribeiro 1998).
Specimens examined: BRAZIL. rio GrandE do suL:
Sinimbu, Cava Funda, 29°27’41.8”S, 52°31’11.7”W,
08/04/10 11:19
570
Hoehnea 36(4): 551-595, 2009
500 m alt., open place, 12-II-2003, A.A. Spielmann
111 (SP).
Parmotrema mellissii is characterized by the
isidiate thallus with alectoronic acid (medulla KC+
rose → quickly orange, UV+ greenish blue). The
similar P. hypomiltoides do not form isidia, but
soredia, often in arbuscular structures. The cilia are
simple to irregularly branched (igure 41).
Parmotrema mordenii (Hale) Hale, Phytologia
28(4): 337. 1974 ≡ Parmelia mordenii Hale,
Smithsonian Contributions to Botany 4: 19. 1971.
Type: DOMINICA. North of Coulibistri, ca. 30 m
alt. M.E. Hale 35649 (holotype US, isotypes BM,
UPS), ide Hale (1971b).
Figure 24
Thallus greenish gray or with some gray
patches, especially in the lobe apices, lobate,
membranaceous, adnate, saxicolous, 15 cm broad.
Lobes irregularly branched, laterally overlapping,
2–10 mm wide, surface smooth or sometimes
cracked, opaque to sublustrous, becoming rugose
and cracked in the old parts; apical zone plane to
subconcave, undulate, often involute, margin entire
to crenate; lateral marginal zone undulate, imbricate
to ascendant, especially in the axils, giving a coiled
look to the lobe, sometimes ascendant with the apex
involute. Lacinules, maculae, isidia, pustules and
cilia absent. Soralia whitish, usually marginal and
linear, or sometimes submarginal to laminal and
with a pustular look, rarely capitate, turning the
lateral margins ascendant and involute, or frequently
sinuous; soredia subgranular. Medulla white. Lower
surface black, lustrous, rugose; marginal zone brown,
lustrous, 2.0–6.0 mm wide, nude, smooth, rugose or
papillate, with attenuate limit. Rhizines concolor to
the lower surface, simple although usually joined
and forming disks of adherence to the substrate,
0.10–0.50 × 0.05–0.10 mm, frequent, dispersed in
small groups. Apothecia unknown. Pycnidia laminal
to submarginal, barely conspicuous, ostiole black;
conidia sublageniform, 4.0-6.5 × 1.0 µm.
Color tests: cortex K+ yellow, UV-; medulla K+
yellow, C-, KC-, P+ slowly weak yellow, UV-.
TLC: atranorin (cortical and medullar), caperatic acid,
protopraesorediosic acid and protolichesterinic acid.
Distribution: Africa (Winnem 1975), Central America
(Hale 1971b), North America (Jackson & Hopkins
30518004 miolo.indd 570
1980, Esslinger 2008) and South America (Feuerer
2005). In South America it is known to Brazil
(Marcelli 1991) and Venezuela (Feuerer 2005). In
Brazil it was recorded to SP (Marcelli 1991).
Specimens examined: BRAZIL. rio GrandE do suL:
Sinimbu, Cava Funda, 29°27’41.8”S, 52°31’11.7”W,
500 m alt., open place, 12-II-2003, A.A. Spielmann
110 (SP).
Parmotrema mordenii is recognizable by the
adnate thallus, eciliate lobes, presence of soralia
and medulla with atranorin (K+ yellow, P+ slowly
weak yellow). In P. praesorediosum the soralia are
usually crescent-shaped and atranorin is lacking in
the medulla. Although Sipman (2005) also treat these
two species as distinct, Krog & Swinscow (1981)
putted P. mordenii as synonym of P. praesorediosum.
In the protologue of P. mordenii, Hale (1971b)
stated that the soralia could occasionally form
coralloid subfatiscent structures. We found in the
specimen collected some pustular, capitate soralia that
remember this description, however, more material
is necessary to prove if this is also a good feature to
take this species apart from P. praesorediosum.
Parmotrema perlatum (Huds.) M. Choisy, Catalogue
des lichens de la region lyonnaise, fasc. 9, Bulletin
mensuel de la Société Linnéenne de Lyon et
des Société Botanique de Lyon 21: 174. 1952 ≡
Parmelia perlata (Huds.) Ach., Methodus qua
omnes detectos lichenes: 216. 1803. Type: in
Dillenius Herbarium (OXF), illustrated in the plate
20, igure 39B in Dillenius, Historia Muscorum. p.
197, 1742, ide Fleig (1997).
= Parmotrema chinense (Osbeck) Hale & Ahti, Taxon
35: 133. 1986.
Figure 26
Thallus greenish gray or brownish in herbarium,
lobate, membranaceous, loosely adnate, saxicolous,
7.0 cm broad. Lobes irregularly branched, laterally
overlapping, 4.0−7.0 mm wide, surface continuous,
smooth to subrugose, opaque to sublustrous,
becoming reticulate cracked in the center; apical
zone subconcave, undulate to plane, margin crenate;
lateral marginal zone undulate, imbricate to ascendant
and involute, margin crenate to incise-crenate
(sublacinulate). Lacinules, maculae, pustules and
isidia absent. Cilia black, simple, (0.20−) 0.50−1.00
× 0.02 (−0.10) mm, frequent, evenly distributed but
08/04/10 11:19
A.A. Spielmann & M.P. Marcelli: Parmotrema from central Rio Grande do Sul State, Brazil
less common in the lobe apices. Soralia concolor
to the thallus, linear, marginal, sometimes wide or
subcontinuous, turning the margins undulate and
involute, or orbicular to capitate in the sublacinules;
soredia farinose to subgranular. Medulla white.
Lower surface black, lustrous, rugose; marginal zone
brown, 1.0−2.0 mm, lustrous, smooth to rugose,
with a sharp limit. Rhizines black, simple, 0.30−1.00
× 0.01−0.02 mm, abundant, evenly distributed.
Apothecia absent [according to Hale (1965), 3–5 mm
in diameter, disc imperforate; hymenium 70–80 µm;
ascospores 22–30 × 13-16 µm, episporium 3 µm].
Pycnidia absent [according to Maas-Gesteranus
(1947, as Parmelia trichotera Hue), pycnidia with
a black ostiole; conidia cylindrical, 5–7 × ca. 1 µm].
Color tests: cortex K+ yellow, UV−; medulla K+
yellow, C−, KC−, P+ orange, UV−.
TLC: atranorin, stictic acid, constictic acid,
cryptostictic acid, hypoconstictic acid, hypostictic
acid, menegazziaic acid and norstictic acid.
Distribution: Oceania (Elix 1994, Malcolm &
Galloway 1997, Louwhoff & Elix 1999, Kantvilas
et al. 2002), Asia (Hale 1965, Kurokawa 1991b,
Kurokawa & Lai 2001), Europe (Hale 1965), Africa
(Hale 1965, Swinscow & Krog 1988), North America
(Hale 1965, Brodo et al. 2001, Nash & Elix 2002a),
Central America (Hale 1965), and South America
(Hale 1965). In South America it is known to
Argentina (Adler 1992, Calvelo & Liberatore 2002),
Bolivia (Feuerer 2005), Brazil (Marcelli 2004), Chile
(Hale 1965), Uruguay (Osorio 1972) and Venezuela
(Feuerer 2005). In Brazil it was recorded to MG
(Ribeiro 1998), PR (Eliasaro 2001, Eliasaro & Donha
2003), RS (Spielmann 2006), SC (Fleig 1997) and SP
(Marcelli 1991).
Specimens examined: BRAZIL. rio GrandE do suL:
Sinimbu, Cava Funda, 29°27’41.8”S, 52°31’11.7”W,
500 m alt., open place, 4-II-2003, A.A. Spielmann 65
(SP).
Parmotrema perlatum is distinguished by
the ciliate lobes, marginal soralia (capitate in
the sublacinules) and the presence of stictic acid
(medulla K+ yellow, P+ orange). Parmotrema bangii
can be differentiated for the formation of sorediose
pustules, while P. hypoleucinum (J. Steiner) Hale
has small maculae and a wide white margin in the
lower surface (Brodo et al. 2001). In P. hypotropum
30518004 miolo.indd 571
571
(Nyl.) Hale there’s also a lower white margin (as
P. hypoleucinum) but it has just norstictic acid, the
medulla being K+ yellow → red (Brodo et al. 2001).
Finally, P. rampoddense, although with a similar
morphology, presents alectoronic acid (medulla KC+
rose → quickly orange, UV+ greenish blue).
Some authors (Hale 1965, Swinscow & Krog
1988, Eliasaro 2001, Sipman 2005) described P.
perlatum with revolute lobes, while Fleig (1997)
called them involute and Hale & Cole (1988)
as “revolute with the apex involute”. This last
observation agrees with the specimen here seen. The
presence of trace of norstictic acid in the solvent
C, already reported by Elix (1994) and Brodo et al.
(2001), was conirmed here too.
Hale & Ahti (1986) discovered an earlier
name, Lichen chinensis that became universally
accepted. Nevertheless recently Hawksworth (2004)
reestablished Parmotrema perlatum as the valid
name.
Parmotrema praesorediosum (Nyl.) Hale, Phytologia
28(4): 338. 1974 ≡ Parmelia praesorediosa Nyl.,
Sert. Lich. Trop. Labuan Singapore: 18. 1891.
Type: SINGAPURA. 30-XI-1879, E. Almquist
(holotype H-NYL35547, isotype S), ide Fleig
(1997).
Figure 27
Thallus aluminium gray or slightly straw
pinkish, lobate, adnate, membranaceous, saxicolous,
6.0−10.5 cm broad. Lobes irregularly branched,
laterally overlapping, 3.0−10.0 mm wide, surface
smooth, opaque to sublustrous, becoming rugose
and cracked in the old parts; apical zone subplane
to slightly concave, undulate, usually ascendant,
margin entire to subcrenate; lateral marginal zone
undulate, imbricate to ascendant, margin entire to
crenate. Lacinules, maculae, pustules, isidia and
cilia absent. Soralia whitish or slightly grayish,
usually crescent-shaped or more wide and dense,
or orbicular to capitate, marginal to submarginal or
laminal, turning the lateral margins ascendant and
sometimes densely sinuous; soredia subgranular.
Medulla white. Lower surface black, lustrous or
opaque, smooth to rugose; marginal zone pale brown
or sometimes black or white variegated, 3.0−7.0
mm wide, lustrous or opaque, with a sharp limit,
smooth to reticulate rugose or with veins, sometimes
papillate. Rhizines black or concolor to the lower
surface, sometimes brown with white apices, usually
08/04/10 11:19
572
Hoehnea 36(4): 551-595, 2009
simple, 0.20−0.80 × 0.05−0.10 mm, few to frequent,
dispersed in small groups. Apothecia laminal, very
immature (asci and ascospores not developed), rare
[according to Hale (1965), apothecia 4–10 mm in
diameter, with a short stipe, amphithecium rugose
and sorediate, disc imperforate; hymenium 70–80
µm; ascospores 15-21 × 7-10 µm, episporium 1.2–
2.0 µm]. Pycnidia submarginal to laminal, ostiole
black; conidia sublageniform, 4.0−7.5 × ca. 1.0 µm.
Color tests: cortex K+ yellow, UV−; medulla K−, C−,
KC−, P−, UV−.
TLC: atranorin, praesorediosic acid, caperatic acid,
protolichesterinic acid and an unidentiied substance
of Rf 54 in solvent C.
Distribution: Oceania (Elix 1994, Louwhoff &
Elix 1999), Africa (Hale 1965, Swinscow & Krog
1988), Asia (Hale 1965), North America (Hale 1965,
Esslinger 2008), Central America (Hale 1965) and
South America (Hale 1965). In South America it is
known to Argentina (Hale 1965, Calvelo & Liberatore
2002), Brazil (Hale 1965, Marcelli 2004), Chile
(Feuerer 2005), French Guiana, Paraguay (Hale 1965),
Uruguay (Osorio 1992a) and Venezuela (Hale 1965,
Vareschi 1973). In Brazil it was recorded to BA (Lynge
1914, as Parmelia capitata), MG (Hale 1965), MS
(Fleig & Riquelme 1991), PA (Brako et al. 1985), RJ
(Hale 1965), RS (Spielmann 2006), SC (Fleig 1997)
and SP (Marcelli 1991, 1992).
Specimens examined: BRAZIL. rio GrandE do suL:
Santa Cruz do Sul, margin from highway RST-287,
km 102, 29°41’03.3”S, 52°25’33.6”W, 150 m alt., 28VII-2003, A.A. Spielmann 386 (SP), A.A. Spielmann
399 (SP).
Parmotrema praesorediosum is characterized
by the adnate thallus, eciliate lobos, crescentshaped soralia and medulla with caperatic acid,
praesorediosic acid and protolichesterinic acid (spot
tests negative). In P. mordenii the soralia are usually
marginal and linear, sometimes pustular and laminal,
and atranorin is present in the medulla (K+ yellow).
P. dilatatum (Vainio) Hale has a different chemistry,
with trace of usnic acid in the cortex and medulla
with protocetraric acid and echinocarpic acid (Fleig
1997).
The caperatic acid was reported by Hale (1965)
and Awasthi (1976), but several authors don’t found
it, like Elix (1994), Fleig (1997), Kurokawa & Lai
30518004 miolo.indd 572
(2001), Louwhoff & Elix (1999) and Nash & Elix
(2002a). Maybe it’s necessary to revise the chemical
identiication of this lichen.
Parmotrema rampoddense (Nyl.) Hale, Phytologia
28(4): 338. 1974 ≡ Parmelia rampoddensis
Nylander, Acta Societatis Scietiarum Fennicae
26(10): 7. 1900. Type: SRI LANKA. Ramboda,
Almquist (holotype H-NYL35555, isotype S), ide
Hale (1965).
Figure 28
Thallus greenish gray ti pale gray, lobate,
membranaceous to subcoriaceous, loosely adnate,
corticicolous or saxicolous, 4.5−20.0 cm broad.
Lobes irregularly branched, laterally overlapping
to crowded, 4.0−20.0 mm wide, surface smooth,
rugose, irregular or scrobiculate, opaque to lustrous,
becoming rugose and cracked towards the center;
apical zone plane to concave, usually ascendant
and sometimes revolute, undulate, margin entire to
subcrenate or sublacinulate; lateral marginal zone
undulate, plane, ascendant, subimbricate or involute,
margin entire to subcrenate or sublacinulate to
lacinulate. Lacinules simple, plane, 0.5–2.5 (−4.0)
× 0.5–2.5 mm, evenly distributed or sometimes
only in the center, apex truncate to rounded, lower
surface black, rarely brown or white variegated;
maculae absent. Cilia black or cupreous and then K+
purplish, simple, furcated or irregularly branched,
0.50−6.0 × 0.02−0.10 mm, frequent to abundant,
evenly distributed. Pustules and isidia absent.
Soralia whitish, marginal to submarginal, linear
interrupted or wider (up to 2.5 mm) and approaching
the lamina, turning the lateral margins involute,
or capitate, laminal or in the lacinules apices;
soredia farinose to grossly granular, coalescing in
the older parts, sometimes developing small cilia.
Medulla white, except by some orange parts K+
red in older parts, close to the lower cortex. Lower
surface black, lustrous, smooth to rugose; marginal
zone brown or sometimes black or with parts white
or beige variegated close to the soralia, 1.0−4.0
mm wide, nude, sublustrous, lustrous or opaque,
smooth to rugose or rugose-reticulate, rarely with
veins, with an attenuate or sharp limit. Rhizines
black, simple, furcated or irregularly branched,
0.20−3.50 × 0.02−0.05 (−0.10) mm, usually longer
in the marginal zone than in the central lobe margins,
frequent to abundant, dispersed in groups by almost
all surface, sometimes some rhizines reaching the
08/04/10 11:19
A.A. Spielmann & M.P. Marcelli: Parmotrema from central Rio Grande do Sul State, Brazil
margin. Apothecia absent [according to Hale (1965),
3–10 mm in diameter, amphithecium sorediate, disc
imperforate; hymenium 65–75 µm; ascospores 10–12
× 6–7 µm, episporium 1 µm]. Pycnidia submarginal
to laminal, barely conspicuous, without prominent
margin, ostiole black or brown; conidia bacillar to
iliform, 5.0−9.0 × ca. 1.0 µm.
Color tests: cortex K+ yellow, UV−; medulla K−, C+
rose → quickly orange or C– (rarely C+ yellowish
in some areas), KC+ rose → quickly orange, P−,
UV+ greenish blue. Soredia sometimes UV+ slightly
yellowish . The orange pigment (skyrin) is lacking in
some specimens.
TLC: atranorin, alectoronic acid, α-collatolic acid
and skyrin.
Distribution: Oceania (Hale 1965, Elix 1994,
Louwhoff & Elix 1999); Asia (Hale 1965, Awasthi
1976, Kurokawa 1991b, Kurokawa & Lai 2001, Elix
& Schumm 2001, Louwhoff & Elix 2002), Africa
(Hale 1965, Winnem 1975), North America (Hale
1965, Brodo et al. 2001, Esslinger 2008), Central
America (Hale 1965) and South America (Hale 1965).
In South America it is known to Argentina (Calvelo
& Liberatore 2002), Bolivia (Hale 1965), Brazil
(Marcelli 2004), Chile (Feuerer 2005), Colombia
(Hale 1965), Guyana (Feuerer 2005), Uruguay (Osorio
1992a) and Venezuela (Feuerer 2005). In Brazil it
was recorded to MG (Hale 1965), PR (Eliasaro 2001,
2006), RS (Spielmann 2006) and SP (Ribeiro 1998).
Specimens examined: BRAZIL. rio GrandE do suL:
Boqueirão do Leão, Linha Sinimbuzinho, Perau da
Nega, 29°20’02.2”S, 52°26’27.9”W, 440 m alt.,
slightly shaded place, 24-II-2004, A.A. Spielmann
& L.S. Canêz 1254 (SP); Herveiras, 29°25’53.7”S,
52°40’19.6”W, 570 m alt., open place, 20-I-2004,
A.A. Spielmann & J. Putzke 1253 (SP); Sinimbu,
Cava Funda, 29°27’41.8”S, 52°31’11.7”W, 500 m alt.,
open place, 4-II-2003, A.A. Spielmann 09 (SP), A.A.
Spielmann 17 (SP), A.A. Spielmann 25 (SP); idem,
12-II-2003 A.A. Spielmann 94 (SP); idem, open place,
12-II-2003, A.A. Spielmann 106 (SP); idem, Linha
Almeida, 29°23’20.2”S, 52°30’21.9”W, wrapping
a wooden stub, close to ground, open place, 4-VII2003, A.A. Spielmann 255 (SP); idem, Linha Almeida,
29°23’20.2”S, 52°30’21.9”W, on bark of Araucaria
angustifolia, open place, 5-I-2004, A.A. Spielmann
& L.S. Canêz 1204 (SP); idem, Salto Rio Pardinho,
29°22’54.8”S, 52°32’33.2”W, 250 m alt., at river’s
30518004 miolo.indd 573
573
margin, shaded place, 2-II-2003, A.A. Spielmann &
M.A. Sulzbacher 471 (SP).
Parmotrema rampoddense is recognizable by
the ciliate lobes, marginal to submarginal soralia, the
presence of alectoronic acid (medulla UV+ greenish
blue), conidia bacillar to iliform, 5.0−9.0 × ca. 1.0
µm and the orange pigment K+ red (skyrin). In P.
pooli (C.W. Dodge) Krog & Swinscow the conidia
are sublageniform, 7−8 µm long (Krog & Swinscow
1981). Parmotrema hypomiltoides develops soredia
in arbuscular structures, while P. mellissii has
ciliate isidia and inally in P. exquisitum (Kurok.)
DePriest & Hale, reported for Uruguay, the lobes
are eciliate (Kurokawa 1987). Nevertheless, Fleig
(1997) regarded this last species as synonym of P.
rampoddense, stating that the type of P. exquisitum
seems an old specimen that suffered weather
conditions.
The K+ purplish reaction of some cilia was also
observed in the type of P. rampoddense (Krog &
Swinscow 1981).
Parmotrema sancti-angeli (Lynge) Hale, Phytologia
28(4): 339. 1974 ≡ Parmelia sancti-angeli
Lynge, Arkiv för botanik 13(13): 35. 1914. Type:
BRAZIL. rio GrandE do suL: Colonia Santo
Angelo prope Cachoeira (nowadays Agudo),
Malme s.n. (holotype S, isotype US), ide Hale
(1965) and Fleig (1997).
Figure 30
Thallus greenish gray, lobate, membranaceous
to subcoriaceous, loosely adnate, corticicolous or
saxicolous, 8.5−11.0 cm broad. Lobes irregularly
branched, laterally overlapping, contiguous or
crowded, 4.0−20.0(−25.0) mm wide, surface
smooth or slightly rugose, opaque to sublustrous,
becoming rugose and cracked in the center; apical
zone plane to subconcave, undulate, margin crenate;
lateral marginal zone undulate or strongly undulate
towards the center, ascendant, sometimes involute,
margin entire to crenate or sublacinulate. Lacinules,
pustules, isidia and maculae absent. Cilia simple or
less frequently furcated, (0.50–) 1.00–5.50 × 0.05–
0.10 mm, usually ascendant, frequent to abundant,
evenly distributed. Soralia white, linear, continuous
or interrupted, wider in the thallus center, marginal;
soredia farinose to subgranular. Medulla white.
Lower surface black, lustrous, smooth to rugose;
marginal zone pale brown, sometimes dark brown,
white variegated or cream color, or still black, 3−15
08/04/10 11:19
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Hoehnea 36(4): 551-595, 2009
mm wide, nude, lustrous, smooth to papillate or
rugose, sometimes strongly veined, with a sharp or
attenuate limit. Rhizines black, cream color, pale
brown, whitish or brown with the apices whitish, or
concolor to the margin, usually simple, sometimes
furcated or rarely more branched, 0.50−2.50 ×
(0.02−) 0.05−0.15 (−0.20) mm, frequent to abundant,
dispersed in groups. Apothecia absent [according to
Hale (1965), adnate, disc imperforate; hymenium 65
µm; ascospores 13–18 × 7–10 µm, episporium 1.0
µm, but see the Notes below]. Pycnidia submarginal,
ostiole black; conidia sublageniform, 5.0−9.0 × ca.
1.0 µm.
Color tests: cortex K+ yellow, UV−; medulla K−, C+
rose or red, KC+ rose or red, P−, UV−.
TLC: atranorin, gyrophoric acid.
Distribution: Oceania (Elix 1994, Louwhoff & Elix
1999), Asia (Hale 1965, Awasthi 1976, Kurokawa
1993, Kurokawa & Lai 2001, Elix & Schumm 2001),
Africa (Hale 1965, Winnen 1975, Krog & Swinscow
1981, Swinscow & Krog 1988) and Americas (Hale
1965). In South America it is known to Argentina
(Hale 1965, Calvelo & Liberatore 2002), Brazil
(Zahlbruckner 1930, Marcelli 2004), Chile, Colombia
(Hale 1965) and Venezuela (Hale 1965, Vareschi
1973). In Brazil, it was recorded to MG (Hale 1965,
Ribeiro 1998), PR (Osorio 1977a,b, Fleig 1997,
Eliasaro 2001), RS (Spielmann 2006), SC (Fleig 1997)
and SP (Hale 1965, Osorio 1989, Pereira & Marcelli
1989, Marcelli 1991, Ribeiro 1998).
Specimens examined: BRAZIL. rio GrandE do suL:
Boqueirão do Leão, Linha Sinimbuzinho, Perau da
Nega, 29°20’02.2”S, 52°26’27.9”W, 440 m alt., in
the stream’s margin and close to the street, in a little
shaded place, 24-II-2004, A.A. Spielmann & L.S. Canêz
1061 (SP); idem, Cascata do Gamelão, 29°18’13.7”S,
52°26’51.7”W, 500 m alt., in the stream’s margin,
slightly shaded, 31-I-2004, A.A. Spielmann & M.A.
Sulzbacher 1110 (SP); Herveiras, 29°25’53.7”S,
52°40’19.6”W, 570 m alt., slightly shaded, 24-I-2004,
A.A. Spielmann et al. 727 (SP); Sobradinho, margin
from RST-481, next to the crossroads, 29°24’20.2”S,
53°01’25.9”W, 375 m alt., open place, 17-VII-2003,
A.A. Spielmann 332 (SP).
Parmotrema sancti-angeli is distinguished by
the ciliate lobes, marginal and linear soralia, the
presence of gyrophoric acid (medulla C+ rose or red)
30518004 miolo.indd 574
and sublageniform conidia 5.0−9.0 × ca. 1.0 µm.
In P. indicum the thallus is robust and the conidia
iliform, 8.0–12.5 × ca. 1.0 µm (see also the Notes
under this species), and P. permutatum (Stirton) Hale
has the lower medulla yellow, because the presence
of secalonic acid A (Fleig 1997).
Parmotrema spinibarbe (Kurok.) Hale ex DePriest
& B. Hale, Mycotaxon 67: 204. 1998 ≡ Parmelia
spinibarbis Kurok., Bulletin of the National
Science Museum Tokyo 17(4): 299. 1974. Type:
BRAZIL. rio dE JanEiro: Petrópolis, ca. 810 m
alt., Kurokawa 8348 (holotype TNS, isotypes G, O,
US, W), ide Kurokawa (1974) and Fleig (1997).
Figure 32
Thallus aluminium gray, lobate, membranaceous,
loosely adnate, saxicolous, 16.5 cm broad. Lobes
irregularly branched, laterally overlapping, 4.0−11.0
mm wide, surface smooth, lustrous or opaque,
becoming reticulate cracked in the center; apical zone
plane to subconcave, usually ascendant, sometimes
involute, undulate, margin strongly crenate;
lateral marginal zone undulate, plane to ascendant,
usually involute, margin crenate to incise-crenate
(sublacinulate). Lacinules and isidia absent. Maculae
weak, punctiform, laminal. Cilia black, simple or
furcated, usually spiky, 0.20−2.00 × 0.02−0.15
mm, frequent, dense to spread, evenly distributed.
Pustules capitate, marginal to submarginal,
fragmenting in soredia. Soralia pustular, marginal
to submarginal, at irst narrow, then becoming wider
and turning the margins thickened and involute, or
subcapitate; soredia granular, sometimes isidioid,
usually coalescing and somewhat stiff, forming a
granular aggregate. Medulla white, sometimes with
orange patches K+ yellow → blood red close to the
lower cortex. Lower surface black, lustrous, smooth
to rugose; marginal zone brown dark, 1.5−4.0 mm
wide, nude, lustrous, smooth to rugose, with attenuate
to sharp limit. Rhizines black, simple or irregularly
branched, 0.20−2.00 × 0.02−0.15 mm, abundant,
covering almost all the surface. Apothecia absent
[according to Kurokawa (1974), stipitate, 3–10 mm
in diameter, disc dark brown, perforate, splitted when
mature; hymenium ca. 50 µm; ascospores 12–14 ×
7–10 µm]. Pycnidia submarginal, without prominent
margin, ostiole black; conidia iliform, straight or
slightly arcuate, 7.5−12.0 × ca. 1.0 µm.
Color tests: cortex K+ yellow, UV−; medulla K+
yellow → blood red, C− or C+ slowly yellowish,
08/04/10 11:19
A.A. Spielmann & M.P. Marcelli: Parmotrema from central Rio Grande do Sul State, Brazil
KC+ orange, P+ yellow → orange, UV+ yelloworange. Some orange patches are present, K+ yellow
→ blood red.
TLC: atranorin, salazinic acid, consalazinic acid,
lichexanthone, and unidentiied substance of Rf 49
in solvent C.
Distribution: Brazil (Marcelli 2004), where it was
recorded to PR (Kurokawa 1974, Fleig 1997), RJ
(Kurokawa 1974), RS (Spielmann 2006), SC (Fleig
1997) and SP (Fleig 1997, Ribeiro 1998).
Specimens examined: BRAZIL. rio GrandE do suL:
Boqueirão do Leão, Linha Sinimbuzinho, Perau da
Nega, 29º20’17.2” S, 52º26’33.6” W, 430 m alt., on
the steep slope in the left side of the stream, partially
shaded, 22-II-2004, A.A. Spielmann & L.S. Canêz
1161 (SP), A.A. Spielmann & L.S. Canêz 1162 (SP).
Parmotrema spinibarbe is distinguished by the
ciliate lobes, sorediose pustules, presence of salazinic
acid and lichexanthone (medulla K+ yellow → blood
red, UV+ yellow-orange), and imperforate apothecia.
Two other similar species share the same chemistry:
P. ultralucens (Krog) Hale has isidia instead of
soredia and longer ascospores,15–17 × 10–12 µm
(Krog 1974), and in P. lichexanthonicum Eliasaro &
Adler the vegetative propagules are lacking (Eliasaro
& Adler 1997).
Eliasaro & Adler (1997) stated that P.
lichexanthonicum could be the parental species of
P. ultralucens. Yet, P. ultralucens has imperforate
apothecia, while in P. lichexanthonicum they are
perforate. According to Hale (1965), species pairs
with these distinct features are not known. So maybe
this is the irst report.
Parmotrema subrugatum (Kremp.) Hale, Phytologia
28(4): 339. 1974 ≡ Parmelia subrugata Kremp.,
Verh. Zool. Bot. Gesell. Wien 18: 320. 1868.
Type: BRAZIL. rio dE JanEiro: Serra dos Órgãos,
Helmreichen s.n. (holotype M, isotype US), ide
Hale (1965) and Fleig (1997).
575
subconcave, undulate, margin entire to sublacinulate;
lateral marginal zone undulate, ascendant, margin
sublacinulate to lacinulate. Lacinules dichotomous
or digitate, subcanaliculate, 1.0–20 × 0.2–3.0
mm, branching several times, evenly distributed,
margin involute, apex truncate or acute; maculae
distinct, punctiform, laminal, marginal or in the
amphithecium. Cilia black, simple, bifurcated or
irregularly branched, 0.5−2.0 × 0.05 mm, frequent,
evenly distributed. Pustules, soredia and isidia
absent. Medulla white. Lower surface black, lustrous,
smooth to rugose; marginal zone ivory white, or
sometimes pale brown, 5−10 mm wide, nude, opaque
to lustrous, with a sharp limit, smooth to rugosereticulate or veined close to the apothecia. Rhizines
black, simple, tortuous, 0.50−1.50 × 0.02−0.05 mm,
abundant, dispersed in groups. Apothecia urceolate
to patent, stipitate, 4−10 mm in diameter, laminal,
frequent, base wide, rugose, amphithecium splitted,
involute, clearly rugose and maculate, sometimes
with digitate projections, margin digitate or imbriate,
without cilia, disc pale ochre, concave to cupulate,
epruinose, imperforate; epithecium 12.5−30.0 µm;
hymenium 60−80 µm; subhymenium 25−40 µm;
ascospores ellipsoid, narrow-ellipsoid or slightly
reniform, (17.5–) 25.0–34.0 × 15–19 µm, episporium
2.5−4.0 µm. Pycnidia laminal or more commonly
submarginal to marginal, conspicuous, ostiole black;
conidia unciform, 4.0−5.5 × ca. 1.0 µm.
Color tests: cortex K+ yellow, UV−; medulla K−, C−,
KC+ rose → quickly orange, P−, UV+ greenish blue.
TLC: atranorin, alectoronic acid and α-collatolic acid.
Distribution: Oceania (Hale 1965, Elix 1994,
Louwhoff & Elix 1999), Asia, Africa (Hale 1965,
Kurokawa & Lai 2001) and Americas (Hale 1965). In
South America it is known to Argentina (Hale 1965,
Calvelo & Liberatore 2002), Brazil (Marcelli 2004)
and Uruguay (Osorio 1972). In Brazil it was recorded
to MG (Hale 1965, Fleig 1997, Ribeiro 1998), PA
(Brako et al. 1985, as Parmotrema cf. subrugatum),
PR (Osorio 1977a/b, Eliasaro 2001), RJ (Hale 1965),
RS (Spielmann 2006) and SP (Ribeiro 1998).
Figure 33
Thallus greenish gray, lobate, subascendant,
coriaceous, corticicolous, 10 cm broad. Lobes
irregularly branched, crowded, 5−7 cm wide, median
axis subascendant or extended, surface smooth to
slightly rugose-scrobiculate, becoming strongly
rugose-scrobiculate in the center; apical zone
30518004 miolo.indd 575
Specimens examined: BRAZIL. r io G randE do
suL: Sobradinho, margin from RST-481, next to the
crossroads, 29°24’′20.2”S, 53°01’25.9”W, 375 m alt.,
open place, 17-VII-2003, A.A. Spielmann 360 (SP).
Parmotrema subrugatum is recognizable by the
unciform conidia, the presence of alectoronic acid
08/04/10 11:19
576
Hoehnea 36(4): 551-595, 2009
(medulla KC+ rose → quickly orange, UV+ greenish
blue), apothecia imperforate, ascospores large (up to
34 µm), the lower ivory white margin and the absence
of vegetative propagules. Parmotrema wainioi (A.L.
Smith) Hale has a brown lower margin, bacillar to
iliform conidia and smaller ascospores (up to 22
µm according to Vainio 1890). In P. argentinum
(Kremp.) Hale, the conidia are bacillar (Fleig 1997),
the apothecia often ciliate, ascospores with up to
22 µm (Hale 1965). P. rigidum (Lynge) Hale has
iliform conidia and perforate apothecia. Finally, in
P. laciniatulum Krog the conidia are sublageniform
(Krog 1991).
Apparently P. subrugatum is the only known
species of Parmotrema with unciform conidia.
However, the literature about is very unclear. Vainio
(1890) described the conidia as cylindrical 4–5 ×
1 µm. Zahlbruckner (1926b) as bacillar, 3.8–6.5 ×
0.5–0.7 µm. In this paper specimens from Brazil
collected by Weddel revealed bacillar and slightly
arcuate conidia, 2.5–5.0 × 0.6–0.7 µm. Based on
the type of conidia found (cylindrical, arcuate and
apical zone, 4–5 × 0.5 µm), Lynge (1914) proposed
Parmelia subrugata var. arcuata. Hale (1965) stated
that the conidia of Parmelia subrugata were not
seen, although before (Hale 1960) he found that the
holotype presented conidia more or less arcuate,
identical to that described by Lynge (1914). Fleig
(1997) found unciform conidia, 5–8 (–9) µm long.
In Elix (1994) and Louwhoff & Elix (1999) they are
bacillar, 4–5 × 1 µm. Finally, Eliasaro (2001) found
cylindrical a sublageniform, straight or slightly
curved conidia, 5–8 µm long.
The specimen studied here is very similar to the
holotype picture published by Hale (1965), where
the strongly rugose apothecia and lacinulate lobes
are sharply distinct.
rugose and cracked in the center; apical zone plane
to subconcave, undulate; margin entire to crenate;
lateral marginal zone undulate, subascendant to
ascendant, more or less imbricate, involute, margin
entire to crenate. Lobules sometimes present, laminal
or marginal, 0.20–4.00 × 0.20–4.00 mm. Lacinules,
cilia, pustules and soralia absent. Maculae usually
absent, when present weak or distinct, laminal. Isidia
with base concolor to the thallus or rarely brownish,
apex pale or dark brown, simple to branched or
sometimes forming coralloid aggregates 1.0–1.5
mm in diameter, narrow, granular or coalescing and
coarsely granular, sometimes becoming sorediose, or
papillary at young, erect, irm or caducous, starting
in the crests or issures, then becoming laminal,
usually more dense in the central areas, 0.05–0.50
× 0.05–0.30 mm. Medulla white. Lower surface
black, opaque or lustrous, rugose; marginal zone
brown, brown and black variegated or sometimes
entirely black, nude, lustrous or rarely opaque, with
a sharp or attenuate limit, smooth, rugose, papillate
or veined, (2.5–) 4.0–12.0 mm wide. Rhizines black
or rarely brown at the margin, simple or furcated,
sometimes with as latted apices, 0.10–3.00 × 0.05–
0.20 mm, frequent to few, dispersed in small groups
or almost evenly. Apothecia absent [according to
Hale (1965), up to 20 mm in diameter, excipulum
isidiate-dentate, amphithecium rugose, maculate,
isidiate, disc imperforate; ascospores 13–15 × 7–10
µm, episporium 1.5 µm]. Pycnidia rare, submarginal,
ostiole black; conidia iliform, straight or arcuate,
(7.5–) 9.0–16.5 (–17.5) × ca. 1.0 µm.
Parmotrema tinctorum (Dèspr. ex Nyl.) Hale,
Phytologia 28(4): 339. 1974 ≡ Parmelia tinctorum
Dèspr. ex Nyl., Flora 55: 547. 1872. Type:
CANARY ISLANDS. Dèspréaux s.n. (holotype
H-NYL35365), ide Fleig (1997).
Distribution: Oceania (Hale 1965, Elix 1994, Malcolm
& Galloway 1997, Louwhoff & Elix 1999), Asia
(Hale 1965, Awasthi 1976, Kurokawa 1991b, 1993,
Kurokawa & Lai 2001, Louwhoff & Elix 2000,
2002), Europe (Purvis et al. 1992), Africa (Hale 1965,
Winnem 1975, Krog & Swinscow 1981, Swinscow
& Krog 1988) and Americas (Hale 1965). In South
America it is known to Argentina (Hale 1965, Calvelo
& Liberatore 2002), Brazil (Marcelli 2004), Chile
(Feuerer 2005), Colombia, Ecuador (Hale 1965),
Guyana, French Guiana (Feuerer 2005), Peru (Hale
1965), Paraguay (Lynge 1914, Hale 1965), Uruguay
(Osorio 1972) and Venezuela (Vareschi 1962, 1973,
Figure 35
Thallus pale to mineral gray, lobate,
membranaceous, subcoriaceous or coriaceous,
loosely adnate, corticicolous or saxicolous, 6–28
cm broad. Lobes irregularly branched, laterally
overlapping or slightly crowded, (3–) 4–22 (–25) mm
wide, surface smooth, opaque to lustrous, becoming
30518004 miolo.indd 576
Color tests: cortex K+ yellow, UV–; medulla K–, C+
red, KC+ red, P–, UV–.
TLC: atranorin, lecanoric acid.
08/04/10 11:19
A.A. Spielmann & M.P. Marcelli: Parmotrema from central Rio Grande do Sul State, Brazil
Dennis 1965, Hale 1965). In Brazil it was recorded
to MG (Lynge 1914, Hale 1965, Fleig 1997, Ribeiro
1998), MS (Fleig & Riquelme 1991, Osorio 1992b),
MT, RJ (Lynge 1914, Hale 1965, Fleig 1997), PA
(Brako et al. 1985), PR (Osorio 1977a/b, Eliasaro
2001), RS (Spielmann 2006), SC (Fleig 1997) and
SP (Hale 1965, Osorio 1989, Nagaoka & Marcelli
1989, Pereira & Marcelli 1989, Marcelli 1991, Ribeiro
1998).
Specimens examined: BRAZIL. rio GrandE do suL:
Boqueirão do Leão, Linha Sinimbuzinho, Perau da
Nega, 29°20’02.2”S, 52°26’27.9”W, 440 m alt.,
stream’s margin and close to the street, shaded place,
24-II-2004, A.A. Spielmann & L.S. Canêz 1257 (SP);
Herveiras, 29°25’53.7”S, 52°40’19.6”W, 570 m alt.,
open place, 20-I-2004, A.A. Spielmann & J. Putzke
1260 (SP); idem, 24-I-2004, A.A. Spielmann, L.S.
Canêz & C. Trentin 932 (SP), A.A. Spielmann et al.
970 (SP); Santa Cruz do Sul, margin from RST-287,
Km 102, 29°41’03.3”S, 52°25’33.6”W, 150 m alt.,
open place, 28-VII-2003, A.A. Spielmann 384 (SP);
idem, on Eucalyptus trunk, open place, 28-VII-2003,
A.A. Spielmann 397 (SP); Sinimbu, Cava Funda,
29°27’41.8”S, 52°31’11.7”W, 500 m alt., open
place, 12-II-2003, A.A. Spielmann 62 (SP), A.A.
Spielmann 100 (SP); idem, open place, 12-II-2003,
A.A. Spielmann 98 (SP); idem, open place, 12-II2003, A.A. Spielmann 113 (SP); idem, 29°27’33.4”S,
52°31’05.1”W, 520 m alt., open place, 5-I-2004,
A.A. Spielmann & L.S. Canêz 1255 (SP); idem,
29°27’33.4”S, 52°31’05.1”W, 520 m alt., with mosses,
open place, 5-I-2004, A.A. Spielmann & L.S. Canêz
1256 (SP); Sobradinho, margin from RST-481, next
to the crossroads, 29°24’20.2”S, 53°01’25.9”W, 375
m alt., open place, 17-VII-2003, A.A. Spielmann 354
(SP); idem, 17-VII-2003, A.A. Spielmann 357 (SP).
Parmotrema tinctorum is distinguished by
the isidiate thallus, broad (4–22 mm) and eciliate,
and the presence of lecanoric acid (medulla C+
red). The apothecia are rare and imperforate (Hale
1965). Parmotrema stuhlmanii (C.W. Dodge) Krog
& Swinscow has narrower lobes (6–8 mm) and
perforate apothecia (Krog & Swinscow 1981), while
P. pseudotinctorum (Abbayes) Hale has inlated
isidia 0.5 × 0.2–0.3 mm (Hale 1965). In P. internexum
(Nyl.) Fleig, the medulla presents norlobaridone and
stictic acid (K+ yellow).
This was the more common species found in the
studied area, so commonly reported (see (Spielmann
30518004 miolo.indd 577
577
2006) at the point to be called “weedy” (Hale 1965).
A very wide morphological variation is admitted,
from cylindrical or coralloid isidia to isidia becoming
sorediose. Maybe more taxa are being wrongly
named and a revision of this group would be very
welcome.
Parmotrema tinctorum is often authored as
“(Nyl.) Hale”, i.e., without Dèspréaux. Here we
agree with Lynge (1914), Krog & Swinscow (1981),
Louwhoff & Elix (1999), Hale & DePriest (1999) and
Hawksworth (2004), since Nylander (1872) admitted
the Dèspréaux as the author of this species (se also
the Vienna Code Art. 46.4, in MacNeill et al. 2007).
The specimens Spielmann 113 and Spielmann
354 are maculate. This feature is rarely found in
the literature (just Galloway 1985), as well as the
presence of lobules, reported by Krog & Swinscow
(1981) and also observed in our specimens.
Parmotrema wainioi (A.L. Smith) Hale, Phytologia
28(4): 339. 1974 ≡ Parmelia wainii A.L. Smith,
Journal of the Linnean Society, London, Botany 46:
85. 1922. Type: BRAZIL. Minas GErais: Caraça,
Vainio, Lichenes Braziliensis exsiccati n° 400
(holotype TUR-V2410), ide Fleig (1997).
Figures 36, 38
Thallus pale gray, lobate, membranaceous to
subcoriaceous, loosely adnate, corticicolous or
saxicolous, 8.0–11.5 cm broad. Lobes irregular,
laterally overlapping to crowded, 3–20 mm wide,
surface smooth to irregular or scrobiculate, opaque to
lustrous, becoming rugose or reticulate cracked in the
center; apical zone plane to subconcave, ascendant,
involute or revolute, undulate, margin entire to
crenate or incise-crenate (sublacinulate); lateral
marginal zone undulate, ascendant to imbricate,
margin irregular to crenate or sometimes lacinulate.
Lacinules plane to subcanaliculate, simple or
furcated, evenly distributed, with rounded, truncate or
acute apex, 2–15 × 2–12 mm, lower surface black or
brown; maculae absent. Cilia black or sometimes red
pigmented (then K+ purplish), usually simple, rarely
furcated or irregular-branched (igure 38), 0.20–3.00
× 0.05–0.10 mm, abundant, evenly distributed.
Pustules, soredia and isidia absent. Medulla white.
Lower surface black, lustrous, smooth to rugose;
marginal zone brown to pale brown, lustrous,
1–6 mm wide, with a sharp or attenuate limit, smooth,
rugose, papillate-rhizinate, scrobiculate, with veins
or with small ridges (relatives to the foveolae from
08/04/10 11:19
578
Hoehnea 36(4): 551-595, 2009
the upper surface). Rhizines black, simple, furcated
or irregularly branched, 0.20–2.50 × 0.02–0.10 mm,
abundant, evenly distributed but more dense in some
areas. Apothecia absent [according to Vainio (1890,
as Parmelia proboscidea), 5–20 mm in diameter,
cupulate, sessile to substipitate, imperforate, margin
entire or crenulated, excipulum reticulate-rugose,
ascospores 14–22 × 9–12 µm]. Pycnidia submarginal
to laminal, conspicuous, with prominent margin,
ostiole black; conidia bacillar to iliform, straight or
arcuate, 6.0–12.5 × ca. 1.0 µm.
Color tests: upper cortex K+ yellow, UV–; medulla
K–, C–, KC+ rose → quickly orange, P–, UV+
greenish blue.
TLC: atranorin, α-alectoronic, α-collatolic and an
unidentiied substance of Rf 31 in solvent C.
Distribution: Oceania, Africa (Feuerer 2005) and South
America (Hale 1965). In South America it is known
to Brazil (Marcelli 2004), where it was recorded to
MG (Vainio 1890, as Parmelia proboscidea Taylor,
Hale 1965, Fleig 1997, Ribeiro 1998), PR (Fleig 1997,
Eliasaro 2001), RJ, GO? (Fleig 1997), RS (Spielmann
2006), SC (Fleig 1997) and SP (Ribeiro 1998).
Specimens examined: BRAZIL. rio GrandE do suL:
Herveiras, 29°25’53.7”S, 52°40’19.6”W, 570 m alt.,
on Eucalyptus trunk, open place, 24-I-2004, A.A.
Spielmann & J. Putzke 728 (SP); Sinimbu, Cava
Funda, 29°27’31.6”S, 52°31’02.9”W, 520 m alt.,
open place, 5-I-2004, A.A. Spielmann & L.S. Canêz
986 (SP).
Parmotrema wainioi is recognizable by the
presence of alectoronic acid (medulla UV+ greenish
blue), bacillar to iliform conidia and the absence of
vegetative propagules. P. pseudobreviciliatum Adler,
Elix & Hale has shorter cilia, up to 2.5 mm long
(Adler 1989). And in P. subrugatum the conidia are
unciform.
The epithet “wainii” is in disagreement with the
recommendation 60C.1 of the Vienna Code (McNeill
et al. 2007), since the end should be “oi”. Ribeiro
(1998) proposed “vainioi”, since from 1919 Wainio
changed his last name to Vainio (Alava 1998).
Nevertheless, the article 60.1 asserts that “the original
spelling of a name or epithet is to be retained”, and
the article 60.11, together with the recommendation
60C.1, deal with just the ends of the epithets, the
correct spelling therefore is “wainioi”.
30518004 miolo.indd 578
Group III – Rimelia-like lichens (species with
reticulate maculae and rhizines often reaching the
margin, without dimorphic rhizines, belonging to
Rimelia sensu Hale & Fletcher)
A comparative table to known species was made,
based mainly on the following references: Brodo et
al. (2001), Canêz (2005), Eliasaro (2001), Eliasaro
& Adler (1998), Elix et al. (2002), Esslinger (1972),
Fleig (1997), Hale (1971a), Hale & Fletcher (1990),
Louwhoff & Elix (1998), Marcelli & Ribeiro (2002),
Mata García (1994), Moon et al. (2001) and Ribeiro
(1998)
Parmotrema cetratum (Ach.) Hale, Phytologia 28(4):
335. 1974 ≡ Parmelia cetrata Ach., Synopsis
Methodica Lichenum: 198. 1814 ≡ Rimelia cetrata
(Ach.) Hale & Fletcher, The Bryologist 93(1): 26.
1990. Type: UNITED STATES OF AMERICA.
pEnnsyLvania: Muhlenberg s.n. (lectotype H-ACH,
selected by Hale & Fletcher 1990, isolectotype
UPS), ide Hale & Fletcher (1990).
Figure 4
Thallus greenish gray, lobate, loosely adnate,
corticicolous, saxicolous or muscicolous, 4.5-8.0
cm broad. Lobes irregularly branched, laterally
overlapping to crowded, 2.5-7.0 mm wide, median
axis extended to subascendant, surface smooth to
slightly rugose, usually undulate or subscrobiculate,
opaque to slightly lustrous, becoming rugose and
reticulate cracked towards de center; apical zone
subconcave, plane to ascendant, undulate; margin
entire to incise-crenate (sublacinulate); lateral
marginal zone plane to more often ascendant, margin
lacinulate. Lacinules furcated, dichotomous or
irregularly branched, 2.0–20.0 × 0.5-3.5 mm, plane
to canaliculated and usually revolute, apex acute
or sometimes truncate, lower surface black with
brown margin, evenly distributed; maculae distinct,
reticular, laminal, forming cracks. Cilia black, simple
to squarrose, 0.20-2.00 × 0.02-0.05 mm, frequent
to few, evenly distributed. Pustules, soredia and
isidia absent. Medulla white. Lower surface black,
lustrous, smooth, rugose or veined; marginal zone
brown, 1.5-5.0 mm wide, lustrous, rhizinate, with
attenuate limit, smooth to veined or scrobiculate.
Rhizines black, usually simple, sometimes
irregularly branched, 0.10-1.00 × 0.01-0.05 mm,
abundant, evenly distributed. Apothecia urceolate,
2-15 mm in diameter, stipitate, submarginal,
margin smooth or incise, amphithecium maculatereticulate, smooth to slightly scrobiculate or rugose,
08/04/10 11:19
A.A. Spielmann & M.P. Marcelli: Parmotrema from central Rio Grande do Sul State, Brazil
579
Table 3. Correspondence between morphological structures and the presence of medullar substances in Rimelia-like lichens.
without
with schizidia
medullar substances
vegetative
sorediate
isidiate
pustulate
or dactyls
propagules
saLaziniC and ConsaLaziniC
aCids
P. cetratum
P. ramescens
P. albinata
P. clavulifera
P. commensurata (±)
P. pontagrossensis
P. reticulatum
protoCEtrariC aCid
P. succinreticulatum
fuMarprotoCEtrariC aCid
P. succinreticulatum
stiCtiC aCid and aLLiEd
P. sticticum
Loxodin
P. homotomum
norLobaridonE
P. homotomum
P. commensuratum
GyrophoriC aCid
P. cetratum
(accessory)
P. clavuliferum
(accessory)
LiChExanthonE
P. diffractaicum
P. pontagrossensis
diffraCtaiC aCid
P. diffractaicum
LobariC aCid
P. albinatum (trace)
MEnEGazziaiC aCid
P. sticticum
CapEratiC aCid
P. macrocarpum
fat aCids
P. pustulatum
P. ruminatum
P. austrocetratum
P. hawaiiensis
P. austrocetratum
(± trace)
P. bonplandii
P. bonplandii
P. simulans
P. clavuliferum
P. succinreticulatum
disc brown, epruinose, perforate, usually involute
and splitted; epithecium 5-10 µm; hymenium
40-55 µm; subhymenium 15-30 µm; ascospores
ellipsoid, 8.5-15.0 × 5.0-10.0 µm, episporium ca. 1
µm. Pycnidia laminal to submarginal or sometimes
reaching the margin, present mainly on the lacinules,
conspicuous, usually with prominent margin, ostiole
black; conidia iliform, 10.0-17.5 × ca. 1.0 µm.
Color tests: cortex K+ yellow, UV-; medulla K+
yellow → blood red, C+ slowly yellowish or orange,
KC+ orange, P+ yellow → orange, UV-.
tLC: atranorin, salazinic and consalazinic acids.
Distribution: Oceania (Elix 1994, Galloway 1985,
Malcolm & Galloway 1997, Louwhoff & Elix 1999),
Asia (Awasthi 1976, Kurokawa & Lai 2001), Africa
(Krog & Swinscow 1981, Swinscow & Krog 1988,
Hale & Fletcher 1990, Aptroot 1991), North America
30518004 miolo.indd 579
P. lacteum
P. subisidiosum
(Hale & Fletcher 1990, Brodo et al. 2001, Nash & Elix
2002b, Esslinger 2008), Central America (Feuerer
2005) and South America (Hale & Fletcher 1990).
In South America it is known to Argentina (Lynge
1925, Osorio 1976, Hale & Fletcher 1990, Adler
1992, Calvelo & Liberatore 2002), Bolivia (Lynge
1925, Feuerer et al. 1998), Brazil (Marcelli 2004),
Chile (Feuerer 2005), Paraguay (Lynge 1914), Peru
(Feuerer 2005), Uruguay (Lynge 1925; Osorio 1972,
1992a, 1995, 2003) and Venezuela (Vareschi 1973). In
Brazil it was recorded to MG (Lynge 1914, 1925; Hale
& Fletcher 1990, Ribeiro 1998), MT (Lynge 1914),
PR (Osorio 1977a/b, Fleig 1997, Eliasaro 2001), RJ
(Lynge 1925), RS (Spielmann 2009), SC (Marcelli
1992, Osorio 1997, Fleig 1997) and SP (Marcelli 1991,
1992; Ribeiro 1998).
Specimens examined: BRAZIL. r io G randE do
s uL : Boqueirão do Leão, Cascata do Gamelão,
29°18’13.7”S, 52°26’51.7”W, 500 m alt., right side of
08/04/10 11:19
580
Hoehnea 36(4): 551-595, 2009
the stream, open place, 31-I-2004, A.A. Spielmann &
M.A. Sulzbacher 1289 (SP); Herveiras, 29°25’53.7”S,
52°40’19.6”W, 570 m alt., with mosses, open place,
21-I-2004, A.A. Spielmann et al. 697 (SP); idem,
on Eucalyptus trunk, open place, 24-I-2004, A.A.
Spielmann et al. 717 (SP); Sinimbu, Cava Funda,
29°27’41.8”S, 52°31’11.7W, 500 m alt., open
place, 4-II-2003, A.A. Spielmann 63 (SP); idem,
29°27’33.4”S, 52°31’05.1”W, 520 m alt., open place,
5-I-2004, A.A. Spielmann & L.S. Canêz 752 (SP);
idem, Linha Almeida, 29°23’20.2”S, 52°30’21.9”W,
5-I-2004, A.A. Spielmann & L.S. Canêz 1292 (SP).
Parmotrema cetratum is recognizable by the
absence of vegetative propagules and the production
of salazinic acid (medulla K+ yellow → blood
red). Some similar species can be distinguished:
P. austrocetratum Elix & Johnston is a lacinulate,
develops schizidia and has iliform conidia up to
25 µm long (holotype CHR!); P. hawaiiensis (H.
Magn.) Blanco, Crespo, Divakar, Elix & Lumbsch
also bear schizidia, but the conidia are just up to 15
µm long and the lacinules lacking (holotype S!);
P. homotomum presents norlobaridone and loxodin
(medulla K–, KC+ rose or reddish) and inally P.
macrocarpum (Pers.) Hale has just caperatic acid
(Fleig 1997).
According to the current literature, P. cetratum
shows a wide range of morphological variation,
maybe a consequence of too broad species concepts,
clearly seen by the long list of synonyms attributed
to this species (see Hale & Fletcher 1990).
Galloway (1985) reported several morphological
differences between specimens from coastal rocks and
from wood or bark inland. Fleig (1997) recognized
three distinct groups of species, based on thallus
shape. Interestingly, Moon et al. (2001), studying
material from Hawaii, also established three groups,
adding features from cilia and substrate preferences,
and that correspond to Fleig’s groups. These patterns
maybe are indicative of speciic variation, not just
environmental modiications
Parmotrema clavuliferum (Räs.) Streimann,
Bibliotheca Lichenologica 22: 93. 1986 ≡ Parmelia
clavulifera Räs., Annales Botanici Societatis
Zoologicae Botanicae Fennicae 20(3): 4. 1944
≡ Rimelia clavulifera (Räs.) Kurok., Journal of
Japanese Botany 66(3): 158. 1991. Type: TAHITI.
Vieillard s.n. (lectotype H, selected by Hale &
Fletcher 1990), ide Hale & Fletcher (1990).
Figures 5-7
30518004 miolo.indd 580
Thallus pale gray, lobate, membranaceous,
loosely adnate, saxicolous or muscicolous, 6.522 cm broad. Lobes irregularly branched, laterally
overlapping, subpalmate-lacinulate, 2-12 mm wide,
surface smooth, opaque to sublustrous, usually
pruinose, becoming reticulate cracked in the center;
apical zone plane to subconcave, undulate, margin
crenate to sublacinulate; lateral marginal zone usually
undulate and ascendant, margin usually lacinulate.
Lacinules simple to furcated, 0.5-2.0 × 0.2-1.5 mm,
plane to canaliculated, apex acute to truncate, evenly
distributed, lower surface white variegated under
the soralia; maculae distinctly, reticular, laminal,
forming cracks. Cilia black, simple or sometimes
squarrose, 0.50-2.00 × 0.02-0.05 mm, frequent
to abundant, but infrequent in the sorediate parts,
evenly distributed. Pustules and isidia absent. Soralia
labriform, capitate or orbicular, present in the lacinules
apices or sometimes submarginal; soredia farinose to
granular. Medulla white. Lower surface black, lustrous,
smooth; marginal zone dark brown, white variegated
under the soralia, 1-2 mm wide, lustrous, nude or
with rhizines, with attenuate limit, smooth to papillate
or slightly rugose. Rhizines black, simple, furcated or
squarrose, 0.2-2.5 × 0.01-0.15 mm, abundant, evenly
distributed, sometimes groups of thickened rhizines can
be seen. Apothecia unknown. Pycnidia submarginal to
laminal, few to rare, conspicuous, without prominent
margin, ostiole black; conidia bacillar to iliform, 7.512.5 × ca. 1.0 µm.
Color tests: cortex K+ yellow, UV-; K+ yellow →
blood red, C+ slowly yellowish or C-, KC+ orange,
P+ yellow → orange, UV-.
TLC: atranorin, salazinic and consalazinic acids.
Distribution: Oceania (Lamb 1963, Feuerer 2005),
Asia (Feuerer 2005), Hawaii (Moon et al. 2001) and
Brazil (Barros & Xavier Filho 1972).
Specimens examined: BRAZIL. rio GrandE do suL:
Barros Cassal, 29°07’05.4”S, 52°41’19.9”W, open
place, 24-I-2004, A.A. Spielmann et al. 725 (SP);
Boqueirão do Leão, Linha Sinimbuzinho, Perau da
Nega, 29º20’17.2” S, 52º26’33.6” W, 430 m alt., open
place, 23-II-2004, A.A. Spielmann & L.S. Canêz 1347
(SP), A.A. Spielmann & L.S. Canêz 1359 (SP); idem,
29°20’02.2”S, 52°26’27.9”W, 440 m alt., slightly
shaded, 24-II-2004, A.A. Spielmann & L.S. Canêz
722 (SP); Herveiras, 29°25’53.7”S, 52°40’19.6”W,
570 m alt., muscicolous, open place, 24-I-2004,
A.A. Spielmann et al. 683 (SP); idem, open place,
08/04/10 11:19
A.A. Spielmann & M.P. Marcelli: Parmotrema from central Rio Grande do Sul State, Brazil
20-I-2004, A.A. Spielmann & J. Putzke 1303 (SP);
Sinimbu, Cava Funda, 29°27’41.8”S, 52°31’11.7”W,
500 m alt., open place, 12-II-2003, A.A. Spielmann
36 (SP); idem, 29°27’33.4”S, 52°31’05.1”W, 520
m alt., open place, 5-I-2004, A.A. Spielmann & L.S.
Canêz 688 (SP); idem, 5-I-2004, A.A. Spielmann &
L.S. Canêz 745 (SP); idem, 5-I-2004, A.A. Spielmann
& L.S. Canêz 1305 (SP).
Parmotrema clavuliferum is distinguished by
the sorediate lacinules (igure 6), white variegated
lower margin under the soralia (igure 7), iliform
conidia 7.5-12.5 µm long and the presence of
salazinic (K+ yellow → blood red). In P. reticulatum
the soralia are laminal to submarginal (Moon et al.
2001) and the conidia 12-16 µm (Hale & Fletcher
1990). Parmotrema pseudoreticulatum (Tavares)
Hale has a nude and brown lower margin, 2–4 mm
wide (Tavares 1945), P. commensuratum produces
norlobaridone and loxodin (K–, KC+ rose → reddish)
and P. simulans only caperatic acid (negative
reactions).
Parmotrema commensuratum (Hale) Hale. Phytologia
28(4): 335. 1974 ≡ Parmelia commensurata
Hale, Phytologia 22(1): 31. 1971 ≡ Rimelia
commensurata (Hale) Hale & Fletcher, The
Bryologist 93(1): 27. 1990. Type: MEXICO.
vEraCruz: 9 Km E Jalapa, M.E. Hale 19405
(holotype US, isotypes TNS, UPS), ide Hale &
Fletcher (1990).
Figure 8
Thallus greenish gray, lobate, membranaceous,
loosely adnate, corticicolous or saxicolous, 6.5-11.0
cm broad. Lobes irregularly branched, contiguous
to crowded, 2-10 mm wide, surface smooth
to scrobiculate, lustrous, sometimes pruinose,
becoming reticulate cracked in the center; apical zone
subconcave, margin ascendant to revolute, undulate,
crenate to sublacinulate; lateral marginal zone
undulate, ascendant, margin lacinulate. Lacinules
simple to furcated, 0.5-3.5 × 0.4-2.0 mm, plane
to canaliculate, apical zone, truncate or sometimes
acute, evenly distributed; maculae distinct, reticular,
laminal, forming cracks. Cilia black, usually simple,
rarely furcated or irregularly branched, 0.20-1.50 ×
0.02-0.05 mm, few to frequent, evenly distributed.
Pustules and isidia absent. Soralia orbicular to
labriform, submarginal or in the lacinules apices,
often approaching the lamina and turning the margins
involute; soredia subgranular to granular. Medulla
30518004 miolo.indd 581
581
white. Lower surface black, lustrous, smooth to
rugose; marginal zone brown, lustrous, 1.5-3.0
mm wide, usually densely rhizinate, with attenuate
limit, smooth to papillate-rhizinate. Rhizines
black, simple to squarrose, sometimes furcated or
irregularly branched, 0.20-2.00 (-3.00) × 0.020.05 mm, abundant, evenly distributed and covering
almost all the surface. Apothecia absent [according
to Fleig (1997), urceolate, substipitate, submarginal,
amphithecium sorediate]; ascospores unknown.
Pycnidia submarginal, little conspicuous, rare,
without prominent margin, ostiole black; conidia
bacilliform to iliform, 7.5−10.0 × ca. 1.0 µm.
Color tests: cortex K+ yellow, UV-; medulla K-, C-,
KC+ rose → reddish, P-, UV-.
TLC: atranorin, norlobaridone and loxodin.
Distribution: Africa (Hale & Fletcher 1990), North
America (Hale 1971a, Dey 1974, Esslinger 2008),
Central America (Hale 1971a), and South America
(Feuerer 2005). In South America it is known to
Brazil (Marcelli 2004), Colombia (Hale 1971a) and
Venezuela (Feuerer 2005). In Brazil it was recorded
to PR (Fleig 1997, Eliasaro 2001), RS (Spielmann
2006) and SC (Fleig 1997).
Specimens examined: BRAZIL. rio GrandE do suL:
Sinimbu, Cava Funda, 29°27’41.8”S, 52°31’11.7”W,
500 m alt., open place, 4-II-2003, A.A. Spielmann 30
(SP); idem, 29°27’33.4”S, 52°31’05.1”W, 520 m alt.,
open place, 5-I-2004, A.A. Spielmann & L.S. Canêz
1283 (SP); idem, Linha Almeida, 29°23’20.2”S,
52°30’21.9”W, open place, 5-I-2004, A.A. Spielmann
& L.S. Canêz 1299 (SP).
Parmotrema commensuratum is characterized
by the soralia submarginal and in the lacinules apices
and the presence of norlobaridone (medulla K–, KC+
rose → reddish). Parmotrema simulans (Hale) Hale
has just caperatic acid (negative reactions) and P.
clavuliferum presents salazinic acid (medulla K+
yellow → blood red).
Parmotrema diffractaicum (Essl.) Hale, Phytologia
28(4): 335. 1974 ≡ Parmelia diffractaica Essl., The
Bryologist 75(1): 80. 1972 ≡ Rimelia diffractaica
(Essl.) Hale & Fletcher, The Bryologist 93(1): 27.
1990. Type: UNITED STATES OF AMERICA.
tEnnEssEE: Overton County, Cliff Springs, farm of
M.T. Phillips, Phillips 308 (holotype US, isotype
DUKE), ide Esslinger (1972).
08/04/10 11:19
582
Hoehnea 36(4): 551-595, 2009
Figures 11-12
Thallus pale gray, lobate, membranaceous
to subcoriaceous, loosely adnate, corticicolous
or saxicolous, 7-15 cm broad. Lobes irregularly
branched, laterally overlapping to crowded, 3-11
mm wide, surface smooth, lustrous, becoming
reticulate cracked in the center; apical zone
subconcave, undulate, plane or more frequently
ascendant, margin crenate to sublacinulate; lateral
marginal zone undulate, usually ascendant, involute
when sorediate. Lacinules and isidia absent. Maculae
weak to distinct, irregular, laminal. Cilia simple,
“V” branched, cespitose or sometimes irregularly
branched, 0.50-2.50 × 0.02-0.05 mm, evenly
distributed. Pustules capitate, laminal, fragmenting
in soredia. Soralia capitate to extensive, marginal to
submarginal, or forming sorediose laminal pustules;
when dense turning lobe margins involute; soredia
granular. Medulla white. Lower surface black,
lustrous, smooth to slightly rugose; marginal zone
brown dark or rarely brown, lustrous, 1.5-4.5 mm
wide, nude, with attenuate limit, smooth to papillate,
sometimes rugose or with veins. Rhizines black,
simple, squarrose or irregularly branched, 0.204.00 × 0.02-0.10 mm, abundant, dispersed in groups
dense. Apothecia and pycnidia unknown.
Color tests: cortex K+ yellow, UV-; medulla K-, C-,
KC-, CK+ orange, P-, UV+ yellow-orange.
TLC: atranorin, lichexanthone and diffractaic acid.
Distribution: North America (Esslinger 1972, Hale &
Fletcher 1990, Esslinger 2008) and South America,
where it is known just to Brazil (Hale & Fletcher
1990, Marcelli 2004). In Brazil it was recorded to
MG (Marcelli & Ribeiro 2002, as Rimelia luminosa),
PR (Kurokawa 1985), RS (Spielmann 2006) and SC
(Fleig 1997).
Specimens examined: BRAZIL. r io G randE do
s uL : Boqueirão do Leão, Cascata do Gamelão,
29°18’13.7”S, 52°26’51.7”W, 500 m alt., on the
steep slope at the left side of the stream, open place,
31-I-2004, A.A. Spielmann & M.A. Sulzbacher 1089
(SP); Gramado Xavier, Nascente do Rio Pardinho,
29°14’15.0”S, 52°31’14.3”W, 625 m alt., on
Araucaria angustifolia trunk, slightly shaded, 27-I2004, A.A. Spielmann et al. 1163 (SP).
Parmotrema diffractaicum is distinguished by the
pustular to capitate soralia, the lower naked marginal
zone (igure 12) and the presence of diffractaic acid
30518004 miolo.indd 582
(medulla CK+ orange) and lichexanthone (medulla
UV+ yellow-orange). Parmotrema pontagrossense
(Eliasaro & Adler) Blanco, Crespo, Divakar, Elix
& Lumbsch produces salazinic acid (medulla K+
yellow → blood red) and P. bonplandii (Mata)
Blanco, Crespo, Divakar, Elix & Lumbsch is a
similar, but isidiate species (Mata García 1994).
Rimelia luminosa Marcelli (in Marcelli & Ribeiro
2002) was recently recognized as a synonym of P.
diffractaicum (Marcelli & Canêz 2008).
Parmotrema homotomum (Nyl.) Hale, Phytologia
28(4): 336. 1974 ≡ Parmelia homotoma Nyl.,
Flora 68: 613. 1885 ≡ Rimelia homotoma (Nyl.)
Hale & Fletcher, The Bryologist 93(1): 28. 1990.
Type: BRAZIL. rio dE JanEiro: Organ Mountains,
Weddell s.n. (lectotype H-NYL, selected by Hale
& Fletcher 1990), ide Hale & Fletcher (1990) and
Fleig (1997).
Figure 15
Thallus greenish gray, lobate, membranaceous,
loosely adnate, corticicolous or saxicolous, 8.010.5 cm broad. Lobes irregularly branched, laterally
overlapping to crowded, 2.0-15.0 mm wide, surface
smooth to slightly scrobiculate, opaque to lustrous,
becoming reticulate cracked in the center; apical
zone plane to subconcave, ascendant or revolute,
undulate, margin crenate to sublacinulate; lateral
marginal zone usually ascendant and revolute,
undulate, margin sublacinulate to lacinulate.
Lacinules simple, furcated or irregularly branched,
1.0-6.0 × 0.5-2.5 mm, evenly distributed, plane to
canaliculated or concave, with rounded or acute apex;
maculae weak to distinct, reticular, laminal, forming
cracks. Cilia black, simple to squarrose or sometimes
furcated, 0.20-1.50 × 0.02-0.06 mm, frequent to
abundant, evenly distributed. Pustules, soredia and
isidia absent. Medulla white. Lower surface black,
lustrous, smooth to rugose or papillate; marginal zone
brown to dark brown, lustrous, 0.5-5.0 mm wide,
with rhizines, with attenuate limit, smooth. Rhizines
black, simple, squarrose or irregularly branched,
0.10-3.00 × 0.01-0.06 mm, sometimes robust and
slightly thickened (0.50-1.00 × 0.05-0.10 mm),
dense and abundant, evenly distributed. Apothecia
urceolate to slightly concave, usually laterally latted,
1.5-9.5 mm in diameter, submarginal, stipitate,
base constrict, margin smooth to dentate or shortdentate up to sublacinulate, amphithecium maculatereticulate, smooth to rugose or scrobiculate, disc
08/04/10 11:19
A.A. Spielmann & M.P. Marcelli: Parmotrema from central Rio Grande do Sul State, Brazil
brown, epruinose, perforate; epithecium 5.0-15.0
µm; hymenium 35-60 µm; subhymenium 15-35
µm; ascospores ellipsoid, 10.0-16.5 × 6.5-10.0
µm, episporium 0.7-1.2 µm. Pycnidia submarginal
to laminal, conspicuous, usually with prominent
margin, ostiole black; conidia bacillar to iliform,
straight or slightly arcuate, 6.5-12.5 × 1.0 µm.
Color tests: cortex K+ yellow, UV-; medulla K-,
C+ slowly yellowish or C-, KC+ rose or reddish,
P-, UV-.
TLC: atranorin, norlobaridone and loxodin.
Distribution: Argentina (Fleig 1997) and Brazil
(Zahlbruckner 1930, Marcelli 2004). In Brazil it was
recorded to MG (Hale & Fletcher 1990, Fleig 1997),
PR (Eliasaro 2001), RJ (Hale & Fletcher 1990), RS
(Spielmann 2006), SC and SP (Fleig 1997).
Specimens examined: BRAZIL. rio GrandE do suL:
Herveiras, 29°25’53.7”S, 52°40’19.6”W, 570 m alt.,
shaded place, 24-I-2004, A.A. Spielmann et al. 719
(SP); idem, on Eucalyptus trunk, open place, 24-I2004, A.A. Spielmann et al. 735 (SP), A.A. Spielmann
et al. 940 (SP); Santa Cruz do Sul, margin from RST287, Km 102, 29°41’03.3”S, 52°25’33.6”W, 150 m
alt., on Eucalyptus trunk, 28-VII-2003, A.A. Spielmann
390 (SP); Sinimbu, Cava Funda, 29°27’33.4”S,
52°31’05.1”W, 520 m alt., open place, 5-I-2004, A.A.
Spielmann & L.S. Canêz 968 (SP), A.A. Spielmann &
L.S. Canêz 971 (SP); idem, slightly shaded, 5-I-2004,
A.A. Spielmann & L.S. Canêz 1291 (SP).
Parmotrema homotomum is distinguished
by the absence of vegetative propagules and the
presence of norlobaridone (medulla K–, KC+ rose or
reddish). The similar P. cetratum produces salazinic
acid (medulla K+ yellow → blood red) while in P.
macrocarpum only caperatic acid is present (negative
reactions).
Parmotrema simulans (Hale) Hale, Phytologia 28(4):
336. 1974 ≡ Parmelia simulans Hale, Phytologia
22(1): 32. 1971 ≡ Rimelia simulans (Hale) Hale
& Fletcher, The Bryologist 93(1): 29. 1990. Type:
BRAZIL. Minas GErais: Sítio (nowadays Antônio
Carlos), Vainio, Lichenes Braziliensis exsiccati
n° 918 (lectotype TUR-V, selected by Hale 1971,
isolectotype M), ide Fleig (1997) and Hale &
Fletcher (1990).
Figure 31
30518004 miolo.indd 583
583
Thallus pale gray, lobate, membranaceous,
loosely adnate, corticicolous, 15.5 cm broad.
Lobes irregularly branched, contiguous to laterally
overlapping, 2.5-10.0 mm wide, surface smooth,
opaque to lustrous, sometimes pruinose, becoming
reticulate cracked in the center, apical zone usually
plane to slightly ascendant, rarely undulate, margin
plane crenate to incise-crenate (sublacinulate);
lateral marginal zone usually ascendant, undulate,
margin usually lacinulate. Lacinules simple to
irregularly branched, 0.5-3.0 × 0.5-4.0 mm, plane to
slightly canaliculate, apex acute or truncate, evenly
distributed; maculae distinct, reticular, laminal,
forming cracks. Cilia black, simple to squarrose,
0.50-2.50 × 0.02-0.05 mm, few to abundant, evenly
distributed, similar to the rhizines. Pustules and isidia
absent. Soralia in the lacinules, capitate or extensive;
soredia granular. Medulla white. Lower surface black,
lustrous, smooth to rugose; marginal zone brown
dark, lustrous, 1.5-2.5 mm wide, with rhizines, with
attenuate limit, smooth. Rhizines black, simple,
squarrose or irregularly branched, 0.50-2.50 × 0.010.05 mm, abundant, evenly distributed. Apothecia
absent [according to Fleig (1997), urceolate to patent,
5–10 mm in diameter, substipitate, base constrict,
laminal, amphithecium sorediate, margin smooth,
without cilia, disc perforate; ascospores 13.5–16.5
× 6.0–9.5 µm, episporium 1.0–1.3 µm]. Pycnidia
laminal, rare, ostiole brown; conidia unknown.
Color tests: cortex K+ yellow, UV-; medulla K-, C-,
KC-, P-, UV-.
TLC: atranorin, caperatic acid.
Distribution: Africa (Hale 1971a, Krog & Swinscow
1981), North America (Hale 1971a, Nash & Elix
2002b, Esslinger 2008), Central America (Hale 1971a)
and South America (Hale & Fletcher 1990). In South
America it is known to Brazil (Marcelli 2004) and
Venezuela (Feuerer 2005). In Brazil it was recorded
to MG (Hale 1971a, Hale & Fletcher 1990, Ribeiro
1998), PR (Eliasaro 2001), RS (Spielmann 2006), SC
(Fleig 1997) and SP (Ribeiro 1998).
Specimens examined: BRAZIL. rio GrandE do suL:
Herveiras, 29°25’53.7”S, 52°40’19.6”W, 570 m alt.,
on Eucalyptus trunk, open place, 24-I-2004, A.A.
Spielmann & J. Putzke 1306 (SP).
Parmotrema simulans is recognizable by the
formation of capitate to extensive soralia in the
lacinules and the presence of caperatic acid (negative
08/04/10 11:19
584
Hoehnea 36(4): 551-595, 2009
reactions). Parmotrema clavuliferum differs by the
presence of salazinic acid (medulla K+ yellow →
blood red), and P. succinreticulatum, as the name
says, succinprotocetraric and protocetraric acids are
present (medulla K+ weak yellow → orange, KC+
weak yellow (Eliasaro & Adler 1997).
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Figures 1–6. Species of Parmotrema from Serra Geral. 1. Parmotrema alidactylatum (Spielmann & Canêz 1205). 2. Parmotrema
austrosinense (Spielmann & Canêz 1088). 3. Parmotrema bangii (Spielmann & Canêz 700). 4. Parmotrema cetratum (Spielmann &
Canêz 752). 5. Parmotrema clavuliferum (Spielmann & Canêz 1347). 6. Parmotrema clavuliferum, soralia in the lacinules (Spielmann
& Canêz 1359). Scales in millimeters.
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Figures 7–12. Species of Parmotrema from Serra Geral. 7. Parmotrema clavuliferum, lacinules undersurface (Spielmann & Canêz 722).
8. Parmotrema commensuratum (Spielmann 30). 9. Parmotrema conferendum (Spielmann 24). 10. Parmotrema consors (Spielmann &
Canêz 690). 11. Parmotrema diffractaicum (Spielmann et al. 1163). 12. Parmotrema diffractaicum, underside marginal zone (Spielmann
et al. 1163). Scales in millimeters.
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Figures 13–18. Species of Parmotrema from Serra Geral. 13. Parmotrema eciliatum (Spielmann 410). 14. Parmotrema lavomedullosum
(Spielmann & Sulzbacher 743). 15. Parmotrema homotomum (Spielmann & Canêz 971). 16. Parmotrema hypomiltoides (Spielmann
& Sulzbacher 1262). 17. Parmotrema hypomiltoides, soredia (Spielmann & Sulzbacher 1262). 18. Parmotrema indicum (Spielmann &
Canêz 931). Scales in millimeters.
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Figures 19–24. Species of Parmotrema from Serra Geral. 19. Parmotrema internexum (Spielmann & Canêz 1167). 20. Parmotrema
mantiqueirense (Spielmann et al. 938). 21. Parmotrema melanothrix (Spielmann & Canêz 1033). 22. Parmotrema mellissii (Spielmann 111).
23. Parmotrema mellissii, isidiate wrinkles and soredia (Spielmann 111). 24. Parmotrema mordenii (Spielmann 110). Scales in millimeters.
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593
Figures 25–30. Species of Parmotrema from Serra Geral. 25. Parmotrema muelleri (Spielmann 685). 26. Parmotrema perlatum (Spielmann
65). 27. Parmotrema praesorediosum (Spielmann 399). 28. Parmotrema rampoddense (Spielmann 17). 29. Parmotrema recipiendum
(Spielmann 352). 30. Parmotrema sancti-angeli (Spielmann 332). Scales in millimeters.
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Figures 31–36. Species of Parmotrema from Serra Geral. 31. Parmotrema simulans (Spielmann & Putzke 1306). 32. Parmotrema spinibarbe
(Spielmann & Canêz 1161). 33. Parmotrema subcaperatum (Spielmann & Canêz 976). 34. Parmotrema subrugatum (Spielmann 360). 35.
Parmotrema tinctorum (Spielmann et al. 932). 36. Parmotrema wainioi (Spielmann & Putzke 728). Scales in millimeters.
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Figures 37–41. Species of Parmotrema from Serra Geral. 37. Parmotrema eciliatum, lobe with cilia (Spielmann 419). 38. Parmotrema
wainioi, cilia (Spielmann & Putzke 728). 39. Parmotrema indicum, cilia (Spielmann & Canêz 931). 40. Parmotrema lavomedullosum,
“T” like lobe (Spielmann & Sulzbacher 1037). 41. Parmotrema mellissii, cilia (Spielmann 111). Bar = 1 mm.
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