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The Pyxine pungens complex in São Paulo State, Brazil

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The Pyxine pungens complex comprises species producing atranorin in the upper cortex, lacking vegetative propagules and reacting K+ blackish purple in the pigmented medulla. Based mainly on the development of apothecia, pigmentation and development of the internal stipe and chemistry, three new species morphologically similar to P. pungens are described. Pyxine rhodesiaca and P. schechingeri are also treated and compared to this complex of species. © 2011 by the American Bryological and Lichenological Society, Inc.
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The Pyxine pungens complex in São Paulo State, Brazil
Author(s): Patrícia Jungbluth and Marcelo Pinto Marcelli
Source: The Bryologist, 114(1):166-177. 2011.
Published By: The American Bryological and Lichenological Society, Inc.
DOI: 10.1639/0007-2745-114.1.166
URL: http://www.bioone.org/doi/full/10.1639/0007-2745-114.1.166
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The Pyxine pungens complex in Sa
˜o Paulo State, Brazil
Patrı
´cia Jungbluth
1
, and Marcelo Pinto Marcelli
Instituto de Bota
ˆnica, Nu
´cleo de Pesquisa em Micologia, Caixa Postal 3005,
Sa
˜o Paulo-SP, 01061-970, Brazil
ABSTRACT.The Pyxine pungens complex comprises species producing atranorin in the upper cortex,
lacking vegetative propagules and reacting K+blackish purple in the pigmented medulla. Based
mainly on the development of apothecia, pigmentation and development of the internal stipe and
chemistry, three new species morphologically similar to P. pungens are described. Pyxine
rhodesiaca and P. schechingeri are also treated and compared to this complex of species.
KEYWORDS.Taxonomy, Physciaceae,Pyxine rhodesiaca.
¤¤¤
Pyxine Fr. (Physciaceae) comprises more than 65
species, distributed mainly in tropical and
subtropical regions of the world. Only P. sorediata
(Ach.) Mont. and P. subcinerea Stirt. have a relatively
broad distribution extending to temperate Europe
(Moberg 1983). This genus is characterized by a
foliose thallus, the pigmented hypothecium, the
epihymenium reacting K+purple, asci with two- to
rarely three-celled brown Dirinaria-type ascospores,
and 3 to 4 mm long bacillary to sublageniform
conidia. The morphologically (Awasthi 1975; Kalb
1987) and phylogenetically (Helms et al. 2003) most
closely related genus is Dirinaria differs in lacking
rhizines, a K2epihymenium and the presence of
divaricatic or sekikaik acid in the medulla as major
metabolites. In addition, differing from Dirinaria,
many Pyxine species are UV+bright golden yellow
due to lichexanthone in the upper cortex and may
have pigmentation in the upper medulla.
An important character to distinguish the
species in Pyxine is the chemistry – more specifically
the thin layer chromatographic profile –, the
presence, localization and development of vegetative
propagules and of maculae, the color of the
medullary layers, the characterization of the different
parts of the apothecium, such as the carbonization
process of the exciple and the pigmentation and
degree of development of the internal stipe. Some
publications on Pyxine with interesting points of
view and discussions of morphology, anatomy and/
or chemistry are Malme (1897), Stirton (1898),
Imshaug (1957), Swinscow and Krog (1975), Kalb
(1987) and Scutari (1990, 1995) are essential for the
comprehension of this genus.
Twenty-nine species and two varieties of Pyxine
are recorded from Brazil and little was added to the
knowledge of this genus in this country since Kalb’s
monograph (1987). Six of these species produce
atranorin in the upper cortex and lack vegetative
propagules: P. endolutea Kalb, P. primaria Kalb, P.
pungens Zahlbr., P. rhizophorae Kalb, P. rhodesiaca
Vain. ex Lynge and P. schechingeri Kalb. From these
species, Pyxine endolutea and P. rhizophorae the
upper medulla is cream to sulphur yellow and reacts
K+yellow to orange, whereas P. primaria has a K2
cream colored upper medulla. The remaining species
1
Corresponding author’s e-mail:
pjungbluth@yahoo.com
DOI: 10.1639/0007-2745-114.1.166
The Bryologist 114(1), pp. 166–177 0007-2745/11/$1.35/0
Copyright E2011 by The American Bryological and Lichenological Society, Inc.
have at least the upper layer of the medulla yellow to
orange or ochre reacting K+blackish purple.
This group of species without vegetative
propagules and without lichexanthone in the upper
cortex but with a K+blackish purple upper medulla is
commonly considered well understood. It is intended
to show that this group is more diversified than
expected and to reaffirm the importance of characters
as the upper cortex maculation, apothecial margin,
internal stipe and chemistry as well to introduce some
others characters useful in discerning Pyxine species.
MATERIALS AND METHODS
The specimens studied here were collected in
southeastern Brazil, more specifically in the States of Sa
˜o
Paulo and Minas Gerais. Type-specimens were kindly
loaned to us by the curators of BM,S,TUR and Wor were
studied during visits to the herbaria KALB and UPS.
Anatomical studies were made by examinations
of sections cut by hand with a razor blade and
mounted in water under light microscope. Apothecia
whose thalline margins become excluded by age are
named cocoe
¨s-type, whereas apothecia without thalline
margins at any stage of their development are referred
to as the obscurascens-type, following the
nomenclature proposed by Kalb (1987). Except from
the types, which had only one apothecium sectioned,
at least five apothecia in different states of maturity
were diametrically sectioned to study the development
and pigmentation process of the internal stipe.
Concerning to morphological terms, the term
internal stipe was first proposed by Swinscow and
Krog (1975) and is here adopted. Stipital lacuna is
proposed here as the cavity occupied by an
invagination of the lower cortex when no internal
stipe is developed.
We define maculae as discolorations in the
thallus upper surface caused by the absence of algae
cells in the algal layer in the sense of Awasthi (1982)
and Scutari (1990); hence, the faint discolorations
present in some of the Pyxine species here studied
over a continuous algal layer are not considered true
maculae; they are most probably consequence of
thickenings of the upper cortex that appear as
elevated veins and are here named as that. In many
Pyxine species, the older maculae may become
fissured and these have been called pseudocyphellae
by some authors (Awasthi 1982; Kalb 1987). Veins
never develop into cracks and are mainly restrict to
young parts of the laciniae, disappearing upon
ageing. The descriptions presented below are based
on all specimens, including the types, available.
Significant differences between the types and other
specimens are discussed.
The chemistry of the thalli was determined by
traditional color reactions and thin layer
chromatography (TLC) in solvent A and C, following
Bungartz (2001). Types were tested only when there
was sufficient material to allow such studies.
Despite broadly used, the term ‘‘isolectotype’’
never was included in the International Code of
Botanical Nomenclature. We understand this is
because it is misleading, i.e., depending the author it
can be understood as possible to be applied to any
specimen of the type collection from where the
lectotype was chosen (what our experience
demonstrated absolutely dangerous, at least for lichens)
or restricted to real pieces of the chosen lectotype (what
we consider to be a ‘‘true’’ isolectotype). Most of the
times, it is not possible to know what sense was used by
the author of a given publication. We therefore prefer
using ‘‘duplicate of the lectotype’’ when referring to a
real piece of the lectotype.
RESULTS AND DISCUSSION
Key to the Brazilian species with orange to ochre
upper medulla, lacking lichexanthone and
vegetative propagules
1. Medulla completely orange to ochre, norstictic acid present
in epihymenium ............................................. P. schechingeri
1. Medulla with a white lower layer, without norstictic acid in
epihymenium ........................................................................ 2
2. Maculae very evident at the entire upper surface,
effigurate to subreticulate .......................... P. rhodesiaca
2. Maculae, if present, pale and restricted to distal parts of
the thallus, resembling veins, ........................................ 3
3. Apothecia of obscurascens-type .............. P. mantiqueirensis
3. Apothecia of cocoe
¨s-type ...................................................... 4
4. Apothecia without internal stipe ................ P. astipitata
4. Apothecia with a internal stipe which is at least partially
tinged orange .................................................................. 5
5. Internal stipe orange with a white lateral layer, norstictic
acid present in medulla .................................... P. exoalbida
5. Internal stipe orange throughout, sometimes with a basal
white layer below, norstictic acid absent ........... P. pungens
Jungbluth & Marcelli: Pyxine pungens complex 167
THE SPECIES
1. Pyxine astipitata Jungbluth & Marcelli sp. nov.
Figs. 1 & 2
MB 518614
Similis Pyxine pungentis, sed terpenis aliiset
stipitibus absentibus differt.
TYPE:BRAZIL.SA
˜O PAULO STATE: Municipality of
Altino
´polis, Fazenda da Gruta, 21u04908.90S,
47u26914.50W, 650 m alt., on trunk of a thin tree
at the border of a secondary forest, 6 June 2008, P.
Jungbluth &M.J. Kitaura 2078 (holotype: SP).
Description. Thallus orbicular, corticolous, grey
to brownish grey, laciniate, 2.5–7.0 cm in diam.,
Figures 1–2. Holotype of Pyxine astipitata.1. Habit (Bar 55.0 mm). 2. Apothecia without internal stipe (Bar 50.5 mm).
168 The Bryologist 114(1): 2011
closely adnate. Proximal upper surface continuous to
cracked, smooth to rugose, dull, convex. Distal upper
surface continuous, smooth, some parts with
depressions, slightly shiny, convex to slightly convex
to plane, without a darker zone near the tips.
Laciniae sublinear, irregularly branched, contiguous,
rarely laterally overlapping, 0.5–0.8 (–1.2) mm wide;
apices mainly rounded, flat to concave, adnate; lateral
margin irregular to sinuous; axils acute. Pruina
farinaceous or glistening in very small patches,
subapical. True maculae absent or very rare, mainly
restricted to the margins, sometimes with elevated
veins paler than the upper cortex in distal parts,
laminal. Medulla very strong yellow to vivid orange
above, the lower layer white and very thin. Distal
lower surface grey to black, slightly shiny, smooth to
slightly rugose. Proximal lower surface black, slightly
shiny, smooth to irregular. Rhizines concolorous
with the lower cortex, simple or a few irregularly
branched, abundant, evenly distributed, up to 0.3 mm
long. Apothecia of cocoe
¨s-type, frequent, plane to
convex, sessile, laminal, up to 1.0 mm in diam.;
margin sometimes with an orange or brownish tone
due to the color of the inner tissues, smooth to
slightly undulate, not visible in very convex
apothecia; disc black, slightly shiny, epruinose.
Internal stipe (Fig. 2) absent or poorly developed, if
a vestige is present, yellow to orange, sometimes
white above, yellow to orange and sometimes white
above when a vestige is present; the stipital lacuna is
occupied by an invagination of the lower cortex.
Epihymenium 10 mm high; hymenium 60–80 mm
high; subhymenium 50–80 mm high. Ascospores 2-
septate, ellipsoid, (12–) 13–18 36–7 mm. Pycnidia
frequent, subapical. Conidia sublageniform, 3.0–
4.5 mm ca. 1.0 mm.
Chemistry. Upper cortex K+yellow, UV2;
upper medulla K+violet, C+reddish, KC+reddish
brown, P+blackish violet, UV2; atranorin,
triterpenes (Fig. 3).
Comments. Pyxine astipitata is characterized by
the cocoe
¨s-type apothecia, the poorly developed to
absent internal stipe (hence the name ‘‘astipitata’’)
and the K+violet, P+blackish violet upper medulla.
It is common to observe that the lower cortex turns
inward where the internal stipe would normally
develop (Fig. 2).
True maculae are absent or very rare and
restricted to the margins of the laciniae. Elevated
veins can be present in the upper cortex and
sometimes resemble maculae when they are slightly
paler than the upper cortex. These elevated veins,
Figure 3. Type material TLC array of the species of Pyxine pungens complex in solvents A and C. 1: P. astipitata;2:P. exoalbida;3:
P. mantiqueirensis;4:P. pungens;5:P. rhodesiaca. Black ellipses: purple tone stains after charring and heat; gray ellipses: rose tone
stains after charring and heat. Control: atranorin (black circle) and norstictic acid (gray circle).
Jungbluth & Marcelli: Pyxine pungens complex 169
when present, are restricted to distal parts of the
thallus.
The type specimen of Pyxine rhodesiaca differs
by its abundant effigurate to irregularly linear laminal
and marginal maculae, dispersed throughout the
upper surface and the well-developed white internal
stipe. Pyxine pungens differs by having a almost
totally orange well developed internal stipe that
reacts K+rose. Pyxine exoalbida Jungbluth & Marcelli
has a well-developed orange internal stipe and
norstictic acid in the medulla. Pyxine elixii Kalb
(holotype: herb. KALB!), P. mantiqueirensis and P.
schechingeri differ by the obscurascens-type apothecia
with a well-developed internal stipe. Moreover, P.
elixii and P. schechingeri have smaller ascospores (up
to 13 mm) and a medulla that is pigmented
throughout. Pyxine elixii is only known from
Australia and has distinct maculae on the upper
surface. Pyxine schechingeri contains norstictic acid
present in the epihymenium.
Distribution. This new species was found in
Altino
´polis municipality in Sa
˜o Paulo State, near the
border with Minas Gerais State.
Additional material examined. BRAZIL.SA
˜O
PAULO STATE: Municipality of Altino
´polis, 21u04908.90
S, 47u26914.50W, 650 m alt., 6 June 2008, P.
Jungbluth 2079; idem, M.J. Kitaura 1063, 1065 (SP).
Pyxine exoalbida Jungbluth & Marcelli, sp. nov.
Figs. 4 & 6
MB 518615
Similis Pyxine pungentis, sed acidum norsticticum
continenti et anatomia stipitum differt.
TYPE:BRAZIL.SA
˜O PAULO STATE: Municipality of Bauru,
Tibiric¸a
´District, secondary forest managed by
Instituto Florestal, 22u13.45.00S, 49u04950.30W,
575 m alt., on tree branch, 4 June 2008, M.J.
Kitaura &P. Jungbluth 1021 (holotype: SP).
Description. Thallus orbicular, corticolous, grey
to brownish grey, laciniate, 3.5–6.0 cm in diam.,
adnate. Proximal upper surface continuous, smooth
or rugose to smooth, slightly shiny, convex. Distal
upper surface continuous, smooth, sometimes
scrobiculate because of the ‘‘maculae’’, slightly or
distinctly shiny, mostly convex but sometimes plane,
without a darker zone near the tips. Laciniae
sublinear, irregularly branched, contiguous to rarely
laterally overlapping, 0.5–1.0 (–1.4) mm wide; apices
subrounded, convex, loosely adnate; lateral margin
smooth to irregular to sinuous; axils acute. Pruina
rare, farinaceous, dispersed, subapical. True maculae
rare, originated in the bottom of the axils and mainly
restrict to the margins of the laciniae, elevated
irregular or linear to netlike veins, present mainly in
distal parts of the thallus, sometimes paler than the
upper cortex, give to the upper surface a scrobiculate
appearance. Medulla orange to ochre to pale brown,
lower layer white. Distal lower surface grey, slightly
shiny, rugose. Proximal lower surface grey to black,
slightly shiny, smooth to rugose. Rhizines
concolorous with the lower cortex, the apices
becoming paler, irregularly branched, mainly at the
tips, frequent, evenly distributed, up to 0.4 mm long.
Apothecia of cocoe
¨s-type, frequent, plane to slightly
convex, sessile, laminal, up to 1.5 mm in diam.;
margin sometimes with an orange or brown tone due
to the color of the inner tissues, smooth to undulate,
not visible in old convex apothecia; disc black, dull,
epruinose. Internal stipe (Fig. 6) well developed,
orange, circled by a white outside layer.
Epihymenium 5–10 mm high; hymenium 60–65 mm
high; subhymenium 25–70 mm high. Ascospores 2-
septate, ellipsoid, (13–) 14–16 (–20) 35–7 mm.
Pycnidia frequent, submarginal or subapical. Conidia
sublageniform, 4.0–5.5 mm ca. 1.0 mm.
Chemistry. Upper cortex K+yellow, UV2;
upper medulla K+blackish violet, C+faintly reddish,
KC+reddish, P+blackish, UV2; internal stipe K+
faintly rose; atranorin, norstictic acid, triterpenes
(Fig. 3).
Comments. Pyxine exoalbida is characterized by
cocoe
¨s-type apothecia with an orange internal stipe
reacting K+faintly rose, surrounded by a white
outside layer (hence the name ‘‘exoalbida’’), and the
presence of norstictic acid in the medulla, making it
the only known species without vegetative propagules
with norstictic acid in South America. It is important
to note that norstictic acid is present in lower
concentrations in distal parts of the thallus and
cannot be detected if only tips of laciniae are taken
for chemical analysis. Moreover, the strong K+
blackish violet and P+blackish reactions in the upper
medulla mask the typical reactions of norstictic acid.
170 The Bryologist 114(1): 2011
Figures 4–7. 4. Holotype of P. exoalbida (Bar 55.0 mm). 5. Holotype of P. mantiqueirensis (Bar 51.0 cm). 6. The orange
internal stipe of P. exoalbida (holotype), surrounded by a white layer (Bar 50.5 mm). 7. The internal stipe of P. mantiqueirensis
(holotype), orange above and white below (Bar 50.5 mm).
Jungbluth & Marcelli: Pyxine pungens complex 171
It is strongly advised to rely on TLC to confirm the
identity of this species.
Pyxine exoalbida is very similar in morphology
with P. pungens, which, however, lacks norstictic acid
and does not have a white layer surrounding the
orange internal stipe. Pyxine africana Kalb, P.
schechingeri and P. vermiformis Swinscow & Krog
(holotype: BM!) are similar species with pigmented
internal stipe and norstictic acid in medulla, but the
three differ from P. exoalbida in having obscurascens-
type apothecia. Moreover, P. africana has a K+
orange, C2,KC2and P+orange medulla (Kalb
1987), P. schechingeri has norstictic acid in the
epihymenium, smaller ascospores (up to 13 mm) and
a medulla pigmented throughout, without a white
layer below, and finally, P. vermiformis has markedly
convex laciniae, smaller ascospores (12–15 36–
7mm). This species is recorded only from lava rocks
in discrete areas of eastern Africa. Furthermore, each
of these species has a different triterpene array (see
Kalb 1987, fig. 10–12 for P. africana and P.
schechingeri).
Distribution. Brazil, Sa
˜o Paulo State,
Municipalities of Altino
´polis, Bauru, Cachoeira de
Emas, Luizia
ˆnia and Mogi-Guac¸u.
Additional material examined. BRAZIL.SA
˜O
PAULO STATE: Municipality of Altino
´polis, 21u04908.90
S, 47u26914.50W, alt. 650 m, M.J. Kitaura 989, 990
(SP); idem, Municipality of Cachoeira de Emas,
21u589S, 47u209W, alt. 530m, 20 August 1976, M.P.
Marcelli 15876 (SP); idem, Municipality of Luizia
ˆnia,
21u42926.80S, 50u08928.00W, 358 m alt., 1 June
2008, M.J. Kitaura 882, 887 (SP); idem, Municipality
of Mogi-Guac¸u, 22u159S47u109W, 590 m alt., 7
December 1976, M.P. Marcelli 15876 (SP).
Pyxine mantiqueirensis Marcelli & Jungbluth, sp.
nov. Figs. 5 &7
MB 518616
Similis Pyxine pungentis, sed tipo apotheciorum
differt.
TYPE:BRAZIL.MINAS GERAIS STATE: Municipality of Sa
˜o
Tome
´das Letras, 21u449360S, 44u589390W,
959 m alt., on rock in open woodland next to
the Fla
´vio Waterfall, 19 January 2009, M.J.
Kitaura,M.P. Marcelli &B.R. da Hora 1361
(holotype: SP).
Description. Thallus orbicular, saxicolous, grey
to brownish grey, laciniate, 5.5–7.0 cm in diam.,
adnate. Proximal upper surface continuous to
somewhat cracked, smooth to slightly rugose, slightly
shiny, convex. Distal upper surface continuous,
smooth to irregularly rugose and veined, shiny,
convex, plane to slightly convex near the tips, with a
pale brown zone near the tips. Laciniae linear to
sublinear, irregularly branched, contiguous to
overlapping laterally, (0.5–) 0.7–1.0 (–1.3) mm wide;
apices rounded, flat to concave, when concave with
apical margins ascendant; lateral margin sinuous to
crenate; axils acute. Pruina farinaceous, faint,
disperse, subapical. True maculae rare, originated in
the bottom of the axils and mainly restricted to the
margins, elevated veins sometimes paler than the
upper cortex, abundant in distal parts, irregularly
linear to reticulate, laminal. Medulla dark yellow to
orange above, lower layer thin and white. Distal
lower surface brown to slightly greenish brown,
slightly shiny, smooth to rarely papillate. Proximal
lower surface black, shiny, smooth. Rhizines
concolorous with the lower cortex, sometimes with
white apices, simple to subdichotomously and
irregularly branched, frequent, evenly distributed, up
to 0.7 (–1.0) mm long. Apothecia common of
obscurascens-type, slightly concave to plane, sessile,
laminal, up to 1.5 mm in diam.; margin sometimes
with an orange tone due to the orange color of the
inner tissues, smooth to slightly undulate, not visible
in very convex apothecia; disc black, slightly shiny,
epruinose. Internal stipe (Fig. 7) developed, orange,
sometimes orange above and white below.
Epihymenium 10 mm high; hymenium 60–70 mm
high; subhymenium 80–120 mm high. Ascospores 2-
septate, ellipsoid, 14–17 36–8 mm. Pycnidia
frequent, laminal. Conidia sublageniform, 3–4 mm3
ca. 1.0 mm.
Chemistry. Upper cortex K+yellow, UV2;
upper medulla K+blackish purple, C2,KC2,P+
blackish purple, UV2; internal stipe K+faintly rose
in pigmented parts. Atranorin, triterpenes (Fig. 3).
Comments. Pyxine mantiqueirensis is
characterized by apothecia of the obscurascens-type
with orange internal stipe reacting K+faintly rose.
This species lacks obvious effigurate to irregularly
linear maculae as P. rhodesiaca, but develops elevated
172 The Bryologist 114(1): 2011
veins limited to distal parts of the thallus, as do P.
astipitata,P. exoalbida and P. pungens. However,
unlike these species, P. mantiqueirensis has apothecia
of obscurascens-type.
Pyxine mantiqueirensis resembles P. schechingeri
and P. vermiformis, which differ by the smaller
ascospores (10–13 35–6 mminP. schechingeri and
12–15 36–7 mminP. vermiformis) and a different
triterpene array. Moreover, the medulla of P.
schechingeri is pigmented throughout and norstictic
acid is restricted to the epihymenium, whereas P.
vermiformis and P. africana have norstictic acid in the
medulla. Also, P. africana has color tests different
from the other species, namely a medulla reacting K+
orange, C2,KC2and P+orange medulla and its
own triterpene array (Kalb 1987).
One specimen very similar to Pyxine
mantiqueirensis was found in Herb. KALB (no. 12283)
but its chemistry was not studied. If its identity is
confirmed, the distribution of this species will be
extended southward into Sa
˜o Paulo State. For now P.
mantiqueirensis is known only from the type locality.
The epithet mantiqueirensis refers to the Serra da
Mantiqueira, a great range that represents one of the
major geological Brazilian formations, from where
the specimens were collected.
Additional material examined. BRAZIL.MINAS GERAIS
STATE: Municipality of Sa
˜oTome
´das Letras, 21u439S,
44u589W, 1300 m alt., 18 January 2009, M.J. Kitaura
1250 (SP); SA
˜O PAULO STATE: between Municipalities of
Monteiro Lobato and Sa
˜oBentodoSapucaı
´,22u509S,
45u459W, 24 February 1980, K. Kalb 12283(SP).
Pyxine pungens Zahlbr., Ann. Crypt. exot. 1(2):
210. 1928. Figs. 8 & 9
TYPE:INDONESIA.JAVA: Hortus Bogoriensis, ramicola,
C. van Overeem 23 (holotype: W!; isotype: O).
Pyxine meissneri Tuck. var. subobscurascens Malme,
Bihang Kongl.Svenska Vet. -Akad.Handl. 23, afd.
3(13): 37. 1897. TYPE:PARAGUAY.COLONIA RISSO: pr.
Rio Apa, 21.10.1893, Malme 1950 C [lectotype:
UPS!; isolectotype: S! (upper specimen, typification
made by Kalb 1987)]. – Pyxine berteriana Fe
´e var.
subobscurascens (Malme) Imshaug, Trans. Amer.
Micros. Soc. 76: 256. 1957.
Description. Thallus orbicular, corticolous,
brownish grey, laciniate, 2.0–8.0 cm in diam., adnate.
Proximal upper surface continuous, strongly rugose
and irregular, slightly shiny, slightly convex. Distal
upper surface continuous, smooth to rugose,
sometimes with cavities, slightly shiny, plane to
slightly convex, without a darker zone near the tips.
Laciniae sublinear, irregularly branched, contiguous
to slightly overlapping laterally, (0.3–) 0.5–1.0 mm
wide; apices rounded, usually flat and adnate, rarely
ascendant; lateral margin smooth to sinuous,
occasionally lacinulate; axils acute. Pruina absent or
farinaceous, sparse, subapical. True maculae rare,
pale, laminal, elevated veins present, sometimes paler
than the upper cortex, abundant in distal parts,
irregularly linear to network, laminal. Medulla vivid
orange to ochre above, lower layer thin and white.
Lower surface totally black, slightly shiny, smooth to
papillate, sometimes the distal surface paler. Rhizines
concolorous to the lower surface, simple to rarely
irregularly branched, frequent, evenly distributed, up
to 0.5 mm long. Apothecia cocoe
¨s-type, numerous,
plane, rarely convex, sessile, laminal, up to 1.0 mm in
diam.; margin smooth to slightly undulate; disc
black, slightly shiny, epruinose. Internal stipe
developed, quite totally orange, white inferior part
rarely visible. Epihymenium 5–10 mm high;
hymenium 60–80 mm high; subhymenium 40–100 mm
high. Ascospores 2-septate, ellipsoid, (11–) 14–17 3
(4–) 6–8 mm. Pycnidia frequent, subapical. Conidia
sublageniform, 3.0–4.0 mm3ca. 1.0 mm.
Chemistry. Upper cortex K+yellow, UV2;
upper medulla K+violet, C2,KC2,P+blackish
purple, UV2; internal stipe reacting K+faintly
orange to yellow; atranorin, triterpenes (Fig. 3).
Comments. Pyxine pungens is characterized by the
cocoe
¨s-type apothecia with an orange internal stipe.
The lectotype and the isolectotype of Pyxine
meissneri var. subobscurascens differ from the
holotype of P. pungens by the absence of the
rugosities, so frequent in latter. The thin layer
chromatographic results, however, indicate that the
isolectotype of P. meissneri var. subobscurascens and
the holotype of P. pungens do have a very similar
chemistry, differing seemingly only in subtle
difference concentrations of the same substances. The
specimen Jungbluth 1026 has a similar rugose upper
surface as the holotype of P. pungens, but differs in
having elevated veins, absent from the holotype.
Jungbluth & Marcelli: Pyxine pungens complex 173
Figures 8–12. Holotypes of Pyxine.8–9. P. pungens. 8. Habit (Bar 51.0 cm). 9. The totally orange internal stipe (Bar 50.5 mm).
10–11. P. rhodesiaca.10. Habit (Bar 50.5 cm). 11. The totally orange internal stipe (Bar 50.5 mm). 12.P. schechingeri, habit (Bar
51.0 cm).
174 The Bryologist 114(1): 2011
We prefer to maintain P. meissneri var.
subobscurascens as synonym of P. pungens and
therefore keep the Brazilian specimens under this
name until more material is studied, as these
specimens share the same chemistry, the same
apothecia type and similar ascospores measure
interval (14–17 34–8 mm).
Kalb (1987) wrote that Pyxine pungens is
pruinose and the apothecia are of the cocoe
¨s-type, but
lacks maculae, whereas P. rhodesiaca has conspicuous
reticulate maculae, obscurascens-type apothecia and is
epruinose. There is no doubt that the evident
maculation in P. rhodesiaca is not present in P.
pungens. However, a farinaceous, sparse and
subapical pruina can occur in the two taxa.
Moreover, the holotype of P. rhodesiaca has in fact
apothecia of the cocoe
¨s-type.
Pyxine rugulosa Stirt. (holotype: BM!) is a
morphologically similar species from Australia that
differs by the flat to concave laciniae tips (Rogers
1986b, just a few laciniae left in the holotype), the
plates of pruina, the yellow to pale yellow medulla
and the distinct triterpene array.
Pyxine pungens differs from the newly described
species, P. astipitata,P. exoalbida and P.
mantiqueirensis, in apothecial and chemical
characters. Pyxine astipitata lacks a stipe in the
apothecia and produces distinct triterpenes. Pyxine
exoalbida has an orange internal stipe surrounded by
a white layer and produces norstictic acid in the
medulla. Pyxine mantiqueirensis, although with very
similar triterpenes, differs from P. pungens by the
obscurascens-type apothecia and more convex
laciniae.
Distribution. Australia (Elix 2009; Kalb 1987,
1994; Rogers 1986a, b), Indonesia (Zahlbruckner
1928) and South America, where it is known for
Brazil (Brako et al. 1985, Kalb 1987), Costa Rica
(Kalb 1987), Guyana (Aptroot 1987), Paraguay (Kalb
1987; Malme 1897, as Pyxine meissneri Tuck. var.
subobscurascens Malme) and Venezuela (Lo
´pez-
Figueiras 1986, Kalb 1987). In Brazil, it was cited for
the States of Bahia (Kalb 1987), Minas Gerais (Kalb
1987; Krempelhuber 1873, as P. meissneri fide
Kalb1987; Malme 1897, as Pyxine meissneri Tuck. var.
subobscurascens Malme), between Mato Grosso and
Para
´States [Brako et al. 1985, as Pyxine berteriana
var. subobscurascens (Malme) Imshaug], Mato
Grosso do Sul (Kalb 1987) and Sa
˜o Paulo
(Zahlbruckner 1909, as P. retirugella fide Kalb 1987;
Kalb 1987).
Additional specimens examined,B
RAZIL.SA
˜O
PAULO STATE: Municipality of Mogi-Mirim, 22u269S
46u579W, 630m alt., 14 May 2004, P. Jungbluth 1026 (SP).
Pyxine rhodesiaca Vain. ex Lynge, Rev. bryol.
liche
`nol. 10 (N.S.): 90. 1937. Figs. 10 & 11
TYPE:RHODESIA.SALISBURY:onFicus,F. Eyles 3953
(holotype: TUR-Vainio 34628!).
Pyxine retirugella Nyl. var. endoxantha Mu
¨ll. Arg.,
Bull. Herb. Boiss. 4: 91 (1896). TYPE: AMER.
AUSTR.?, Humboldt 129 (holotype: G).
Description. Thallus orbicular, corticolous, grey
to brownish grey, laciniate, 3.0–11.5 cm in diam.,
adnate to closely adnate. Proximal upper surface
continuous, sometimes with cracks, rugose to very
irregular, dull, slightly convex to convex. Distal
upper surface continuous, with irregularities
(cavities) caused by the maculae, rarely smooth,
slightly shiny, convex to plane, without a darker zone
near the tips. Laciniae sublinear, irregularly
branched, contiguous to laterally overlapping, (0.5–)
0.7–1.2 (–1.5) mm wide; apices rounded, flat and
adnate, sometimes slightly concave, then somewhat
ascendant; lateral margins sinuous and irregular,
sometimes becoming sublacinulate in proximal parts;
axils acute. Pruina absent or farinaceous, sparse,
subapical. Maculae distinct, abundant, irregularly
linear to subreticulate, laminal and marginal, rarely
originating cracks. Medulla ochre above, sometimes
orange to dark orange near the tips, lower layer
white. Lower surface totally black or distal lower
surface grey to black, slightly shiny, smooth to
papillate. Proximal lower surface black, slightly shiny
to distinctly shiny, smooth to papillate. Rhizines
concolorous with the lower cortex or with paler
apices, irregularly branched, abundant, evenly
distributed, up to 1.0 mm long. Apothecia of cocoe
¨s-
type, common to numerous, plane, sometimes
becoming convex, sessile, laminal, up to 1.5 mm in
diam.; margin smooth to slightly undulate; disc
black, slightly shiny, epruinose. Internal stipe
developed, white, sometimes upper layer of the same
color as the medulla. Epihymenium 5–10 mm high;
Jungbluth & Marcelli: Pyxine pungens complex 175
hymenium 40–80 (–100) mm high; subhymenium 60
160 mm high. Ascospores 2-septate, ellipsoid, 14–19 (–
21) 36–8 mm. Pycnidia few, subapical. Conidia
sublageniform, 4.0–5.5 mm ca. 1.0 mm [anatomical data
from additional specimens, not from the holotype].
Chemistry. Upper cortex K+yellow, UV2;
upper medulla K+blackish purple, C2,KC2,P+
blackish purple, UV2; internal stipe K2, atranorin,
triterpenes (Fig. 3).
Comments. Pyxine rhodesiaca is characterized by
the abundantly irregularly maculate upper surface,
the ochre upper medulla reacting K+and P+blackish
purple and the white internal stipe. Kalb (1987)
described the apothecia of P. rhodesiaca as
obscurascens-type. But we reexamined the holotype,
which consists of three small fragments of the same
thallus with three apothecia and another half
apothecia glued to the herbarium cardboard. It was
possible to determine that the development of
apothecia is really of the cocoe
¨s-type, as the two
young ones have thalline margins and the other have
carbonized margins. The additional specimens
studied agree with this observation.
For other differences from others species, see
remarks under P. pungens.
Distribution. Africa (Lynge 1937; Swinscow &
Krog 1975, 1988) and South America, where it is
known from Brazil (Kalb 1987) and Venezuela
(Lo
´pez-Figueiras 1986). In Brazil, it was reported
from the States of Mato Grosso, Mato Grosso do Sul
and Sa
˜o Paulo (Kalb 1987), and is here reported as
new to Distrito Federal, in central Brazil.
Additional material examined. BRAZIL.DISTRITO
FEDERAL: Municipality of Brası
´lia, 15u469S, 47u559W,
1150 m alt., 12 August 1991, M.P. Marcelli 30781,
30819,30824 (SP); SA
˜O PAULO STATE: Municipality of
Bauru, 22u13945.00S, 49u04950.30W, 575 m alt., 04-
VI-2008, M.J. Kitaura 1999 (SP); idem, 22u21901.40S,
49u00929.80W, 598 m alt., 04 June 2008, M.J. Kitaura
1951,1971,1972,1973 (SP); idem, Municipality of
Mogi-Guac¸u, 22u159030S47u099250W, 630 m alt., 05
November 2007, P. Jungbluth 1682 (SP).
Pyxine schechingeri Kalb, Bibl. Lichenol. 24: 71.
1987. Fig. 12
TYPE:BRAZIL.ESTADO DA BAHIA: bei Roda Velha, 650 m
alt., an Sandstein, 15-VII-1980, K. Kalb 12313
(holotype: Herb. KALB!).
Description. Thallus orbicular, saxicolous,
brownish grey, laciniate, 3.5 cm in diam., closely
adnate. Proximal upper surface continuous, rarely
cracked, smooth, shiny, convex. Distal upper surface
continuous, smooth, shiny, convex to plane, with a
weak darker zone near the tips. Laciniae linear to
sublinear, irregularly to subdichotomously branched,
contiguous, 0.4–0.8 mm wide; apices rounded, flat to
convex, adnate; lateral margin smooth to slightly
sinuous; axils acute. Pruina farinaceous, dispersed,
subapical. Maculae absent. Medulla reddish orange
to ochre. Distal and proximal lower surface black,
shiny, smooth to rarely irregular. Rhizines (very few
seen) black, simple, frequent, up to 0.3 mm long.
Apothecia of obscurascens-type, frequent, plane to
slightly convex, sessile, laminal, up to 0.6 mm in
diam.; margin smooth to slightly undulate; disc
black, slightly shiny, partly whitish pruinose. Internal
stipe developed, white, upper part reddish brown.
Epihymenium 10–15 mm high; hymenium 50–60 mm
high; subhymenium 90–100 mm high. Ascospores 2-
septate, ellipsoid, 10–13 35–6 mm. Pycnidia
frequent, laminal. Conidia not seen.
Chemistry. Upper cortex K+yellow, UV2;
medulla K+purple, C2,KC2,P+blackish purple,
UV2; internal stipe K2; atranorin, norstictic acid
(in the epihymenium), triterpenes (see Kalb 1987, fig.
10–12).
Comments. Pyxine schechingeri is characterized
by the completely orange medulla, the obscurascens-
type apothecia, the presence of norstictic acid in the
epihymenium, the white internal stipe with a reddish
brown upper part and the small ascospores to 13 mm
long.
For differences from other species, see remarks
under the new species presented above, especially P.
exoalbida.
Distribution. This species is known only from
the type locality.
ACKNOWLEDGMENTS
We thank the curators of BM,S,TUR and Wfor loan of type-
material and Dr. Klaus Kalb (Herb. KALB) and UPS for allowing
the first author to visit the herbaria and study the types cited in
this article. We are also grateful to FAPESP (Fundac¸a
˜ode
Amparo a Pesquisa do Estado de Sa
˜o Paulo) for PhD Grant to
P. Jungbluth (process 05/53955–1), CNPq for a research Grant
to M.P. Marcelli, and Instituto Florestal do Estado de Sa
˜o Paulo
176 The Bryologist 114(1): 2011
for a collecting license (process 42.592/2003). We also thank
Drs. Roland Moberg, Klaus Kalb and H. Kashiwadani for
sending critical literature and Dr Bernard Goffinet for
reviewing the English text.
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Jungbluth & Marcelli: Pyxine pungens complex 177
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The monophyletic origin of the ascomycete family Physciaceae, its position within the Lecanorales and the phylogenetic structure within the family were investigated using nuclear rDNA sequence analyses. The common origin of the Caliciaceae and Physciaceae as previously shown (Wedin et al 2000) was confirmed. Further it could be shown that the Caliciaceae are nested within the Physciaceae. A unique region in loop 37 of the SSU rRNA secondary structure model was identified, which characterizes the Physciaceae/Caliciaceae. The SSU rDNA sequence data did not support a particular relationship with any other Lecanoralean family. Analyses of ITS rDNA sequences revealed a bifurcation of the Physciaceae/Caliciaceae clade, which was found to be congruent with the distribution of certain morphological characters. The congruence with the ITS phylogeny demonstrated the phylogenetic significance of ascus type, hypothecium pigmentation, ascospore characters and excipulum type. Fine-structure details of ascospores and the structure of excipula were found to be important in the recognition of convergences in these traits. Other previously used characters, i.e., growth habit, certain ascospore types or structure of the upper cortex, were found to be of multiple origins within the Physciaceae. All monophyletic lineages of noncrustose growth habit exhibit uniform ascospore types, indicating a higher evolutionary age of ascospore types than foliose growth habit. The taxonomic segregation of the Physciaceae into the Physciaceae and Caliciaceae is proposed here.
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Covers 77 genera and 629 species. This is the first substantial study of tropical lichen flora undertaken using modern research methods. -from Publisher
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Following morphological, anatomical and chemical studies, 15 species of Pyxine (Physciaceae, lichenized ascomycetes) are recorded for Australia. Their classification, distribution and habitats are discussed. The following taxa are reduced to synonymy: Pyxine cocoes var. endoxantha Müll. Arg. and P. meissneri var. rinodinoides Vainio with P. berteriana (Fée) Imshaug; P. brachyloba Müll. Arg., P. nitidula Mull. Arg. and P. microspora Vainio with P. minuta Vainio; P. meissneri var. convexula Malme, P. albida Magnusson and P. pringlei Imshaug with P. petricola; P. meissneri var. subobscurascens Malme with P. pungens Zahlbr.; P. retirugella var. laevior Vainio, P. retirugella var. capitata Zahlbr., P. copelandii Vainio, P. asiatica Vainio, and P. patellaris Kurok, with P. retirugella Nyl.; and P. prominula Stirton with P. cocoes (Sw.) Nyl. The new species P. isidiolenta R. W. Rogers is described and the new name P. linearis R. W. Rogers is proposed for P. retirugella var. endoxantha forma sorediosa Mull. Arg.
Article
The collection of material chemically identical to the fragmen- tary type of Pyxine rugulosa Stirton has allowed clarification of the application of that name. P. rugulosa can be distinguished from other species of Pyxine in Australia by its combination of a broad-lobed, heavily pruinose thallus without vegetative propagules, yellow medulla, and presence of atranorin but not lichexanthone in the cortex. It is appar- ently restricted to regions west of the Great Dividing Range in Queensland.