A new species of Acanthostigma (Tubeufiaceae, Dothideomycetes) from the southern hemisphere

Abstract

A new species belonging to the Dothideomycete genus Acanthostigma is described from bark of two Nothofagus species from Argentina. Its identity as a new species is based on both morphology and molecular sequence data. Acanthostigma patagonica differs from other species in the genus by having larger ascomata and setae and wider, asymmetrical ascospores. An amended key to Acanthostigma species is provided along with a discussion of other species previously described from South America.

Full text

A new species of
Acanthostigma
(Tubeufiaceae, Dothideomycetes)
from the southern hemisphere
R.M. Sa´nchez
1
Laboratorio de Estudios Ba´ sicos y Biotecnolo´gicos de
Algas y Hongos, CERZOS-CCT-CONICET y U.N.S.,
Camino La Carrindanga Km7, B8000FWB Bahı
´a
Blanca, Buenos Aires, Argentina
A.N. Miller
University of Illinois, Illinois Natural History Survey,
1816 South Oak Street, Champaign, Illinois 61820-
6970, USA
M.V. Bianchinotti
Laboratorio de Estudios Ba´ sicos y Biotecnolo´gicos de
Algas y Hongos, CERZOS-CCT-CONICET y U.N.S.,
Camino La Carrindanga Km7, B8000FWB Bahı
´a
Blanca, Buenos Aires, Argentina
Abstract
:A new species belonging to the Dothideo-
mycete genus
Acanthostigma
is described from bark of
two
Nothofagus
species from Argentina. Its identity as
a new species is based on both morphology and
molecular sequence data.
Acanthostigma patagonica
differs from other species in the genus by having
larger ascomata and setae and wider, asymmetrical
ascospores. An amended key to
Acanthostigma
species
is provided along with a discussion of other species
previously described from South America.
Key words:
ITS, LSU,
Nothofagus
, phylogenetics,
southern hemisphere, systematics
INTRODUCTION
In South America and particularly in Argentina
several groups of ascomycetes have been insufficiently
documented. The Andean Patagonian forests are
considered one of the most interesting biogeograph-
ical formations of Argentina, however they have been
categorized as one of the most poorly studied fungal
reservoirs of the world (Lodge et al. 1995). Since 2005
the principal aim of our work has been to increase the
knowledge of the biodiversity of ascomycetes on trees
native to the Andean Patagonian forests. Many of the
species we have found are related to North American
and Australasian species, which occupy habitats with
similar environmental conditions. In this work we
considered the genus
Acanthostigma
, in which almost
all the species have been described from the northern
hemisphere. However, due to the small size of the
ascomata, specimens of
Acanthostigma
and generally
all Tubeufiaceae have been overlooked and their
known distribution reflects more of the collecting
activities of mycologists instead of their actual
geographical distribution (Rossman 1987).
The systematic position of
Acanthostigma
de Not.
(de Notaris 1863) has a long history that has yet to be
resolved. After being transferred in and out of the
family Tubeufiaceae several times (Saccardo 1883;
Ellis and Everhart 1892; von Arx and Mu¨ller 1975;
Barr 1980, 1990, 1993; Crane et al. 1998) it finally was
redescribed and placed in this family by Re´blova´ and
Barr (2000). This placement was supported by
molecular phylogenetic analyses (Tsui et al. 2006,
2007; Promputtha and Miller 2010).
The genus
Acanthostigma
was known previously
only from the northern hemisphere (TABLE I) be-
cause most of the species have been described from
Asia, Europe and North America. The distribution of
Acanthostigma
was extended to the southern hemi-
sphere with the report of
Acanthostigma minutum
(Fuckel) Sacc. from Argentina (Sa´nchez and Bian-
chinotti 2010).
Through our work on the biodiversity of ascomy-
cetes on trees native to the Andean Patagonian forests
in Argentina a putative new species of
Acanthostigma
was found and herein is described, illustrated and
compared morphologically and genetically to other
known species in the genus. We also include in the
phylogenetic analyses the other species reported from
Argentina,
A. minutum
, and discuss its phylogenetic
relationships with North American species. The key to
all accepted species in the genus provided by
Promputtha and Miller (2010) is amended to include
the new species.
MATERIALS AND METHODS
Morphological characterization.—
Samples were collected in
forests of Los Alerces National Park (Chubut) and Lanı´n
National Park (Neuque´n) in the Andes of Patagonia
(Argentina). The vegetation is composed mostly of native
Nothofagus
species together with some species of
Cupressa-
ceae
,
Proteaceae
, ferns and mosses. Leaves, small branches
and bark showing fungal growth when observed with a field
magnifying lens were placed in paper bags and transported
to the laboratory. The samples were dried at room
temperature and deposited at Bahı´a Blanca Biology
Herbarium (BBB). For collections made in 2009 a GPS
eTrex Legend (Garmin Co., USA) was used to obtain map
Submitted 29 Apr 2011; accepted for publication 27 Jun 2011.
1
Corresponding author. E-mail: rsanchez@uns.edu.ar
Mycologia,
104(1), 2012, pp. 223–231. DOI: 10.3852/11-105
#2012 by The Mycological Society of America, Lawrence, KS 66044-8897
223

coordinates. For microscopic examinations sections were
cut freehand under a Leica EZ4 stereo microscope and
mounted in water or 5%KOH with phloxine. A Leica
DM2000 dissecting microscope with a Samsung NV10 digital
camera was used to capture micromorphological images. At
least 10 measurements were taken for each structure
mounted in tap water. Averages for asci and ascospores
are in parentheses. Material was mounted in calcofluor
0.5%for examination under a Nikon Eclipse 80i fluores-
cence microscope with a Nikon DXM 1200F camera system.
DNA extraction and sequencing.—
The extraction, amplifi-
cation and sequencing of DNA followed that of Promputtha
and Miller (2010). The 5.8S and internal transcribed spacer
2 (ITS) region along with the first 600 bp of the 59end of
28S large subunit (LSU) nrDNA, which contains the
variable D1 and D2 regions, were sequenced.
Phylogenetic analyses.—
The ITS and LSU datasets, which
were amended from Promputtha and Miller (2010), were
aligned in Sequencher 4.9 (Gene Codes Corp., Ann Arbor,
Michigan) and manually corrected by eye. Six and two
ambiguous regions consisting of 82 and 25 bp were delimited
by eye and excluded from the ITS and LSU datasets
respectively. The first 66 bp in the 59end and the last 11 bp
in the 39end of LSU also were excluded due to missing
characters in most taxa. The Akaike information criterion
(AIC) (Posada and Buckley 2004) as implemented in
Modeltest 3.7 (Posada and Crandall 1998) was used to
determine the best-fit model of evolution for each dataset,
which was the HKY model (Hasegawa et al. 1985) for the ITS
dataset and the SYM model (Zharkikh 1994) for the LSU
dataset. Both models included a proportion of invariable
sites with the remaining sites subjected to a gamma
distribution shape parameter. Maximum likelihood (ML)
analyses were conducted with PhyML (Guindon and Gascuel
2003) as implemented in SeaView 4.1 (Gouy et al. 2010)
under these parameters: The above models were implement-
ed, invariable sites and across site variation were optimized
and 1000 bootstrap replicates were performed from a BioNJ
starting tree employing the best of nearest neighbor
interchange (NNI) and subtree pruning and regrafting
(SPR) branch swapping. Bayesian analyses employing Mar-
kov chain Monte Carlo (MCMC) were performed on each
dataset with MrBayes 3.1.2 (Huelsenbeck and Ronquist
2001) as an additional means of assessing branch support.
Constant characters were included, the above models of
evolution were implemented and 10 000 000 generations
were sampled every 1000th generation resulting in 10 000
total trees. The Markov chain always achieved stationarity
after the first 10 000 generations, so the first 1000 trees,
which extended well beyond the burn-in phase of each
analysis, were discarded. Posterior probabilities were deter-
mined from a 95%consensus tree generated in PAUP 4.0b10
(Swofford 2002) using the remaining 9000 trees.
TABLE I. Ecological information on the accepted species of
Acanthostigma
Taxa Known hosts Distribution
A. ellisii
On
Abies alba
and on stromata of
Diatrype
stigma
.
Europe:
Germany, Norway, Poland, Slovakia (Robak
1976).
North America
: USA (Re´blova´ and Barr
2000).
A. filiforme
Decorticated wood on ground of
coniferous-deciduous forest.
North America:
USA (Promputtha and Miller 2010).
A. longisporum Calluna vulgaris
,
Dryas octopetala
,
Rhododendron ferrugineum
,
R. hirsutum
,
Rhodothamnus chamaecistus
,
Vaccinium myrtillus
,
V. uliginosum
.
Europe
: Austria, Norway (Re´blova´ and Barr 2000).
A. minutum Fagus sylvatica
,
Gaultheria shallon
,
Nothofagus pumilio
,
Populus
sp.,
Quercus
sp. and on old xylariaceous stromata on
Nothofagus dombeyi
.
Asia:
China, Taiwan.
Europe:
France, Germany,
Switzerland.
North America:
Canada, USA
(Re´blova´ and Barr 2000).
South America:
Argentina (Sa´nchez and Bianchinotti 2010).
A. multiseptatum
Decorticated wood on ground of
coniferous-deciduous forest.
North America:
USA (Promputtha and Miller 2010).
A. patagonica
On bark of
Nothofagus alpina
and
N. pumilio
.
South America:
Argentina (this paper).
A. perpusillum
On inner side of bark of
Cerasus
sp., on
Elletaria
sp. and on decayed wood of
Quercus
sp.
Asia:
Java.
Europe
: Austria, France, Italy,
Switzerland.
North America:
Canada, USA.
South
America:
Grenada, Venezuela (Re´blova´ and Barr
2000).
A. revocatum Quercus
sp. and
Salix vitellina
.
Europe:
Italy (Re´blova´ and Barr 2000).
A. scopulum Fagus sylvatica
,
Pinus
sp.,
Tsuga
sp.
Asia:
China, Taiwan (Chang 2003, Kodsueb et al.
2004).
Europe:
Austria.
North America
: USA
(Re´blova´ and Barr 2000).
A. septoconstrictum
Decorticated wood on ground of
coniferous-deciduous forest.
North America:
USA (Promputtha and Miller 2010).
224 MYCOLOGIA

TAXONOMY
Acanthostigma patagonica R.M. Sa´nchez, A.N. Mill. et
Bianchin., sp. nov. FIGS. 1–2
MycoBank MB561057
Ascomata superficialia vel immersa, globosa, subglobosa
ad conica, atro-brunnea, 100–390 mm diam, 140–490 alta;
superficies setosa, setis brunneis, rectis vel curvis, 0–4-
septatis, 49–125 mm longis, 5–13 mm in basi latis, 1–5 mmin
apice latis. Papillae 30–113 350–100 mm. Peridium textura
angulari, cellulis brunneis, polyedricis, intra lutescentibus.
Pseudoparaphyses tenues, septatae, hyalinae, ramosae, 1–
3mm, reticulum formantes. Periphyses undulatae, hyalinae.
Asci bitunicati, cylindrico–clavati, 8-spori, 85–139 315–
33 mm. Ascosporae fusiformes, ad extremitates decrescentes
et dissimiles, hyalinae, laeves, 8–18-septatae, septa con-
stricta, 1–2 cellulis mediis latioribus 62–111 35–10 mm.
Type.
ARGENTINA. NEUQUE
´N PROVINCE: Pque.
Nacional Lanı´n, on Provincial Route Number 48,
before reaching Playas de Yuco, on bark of
Nothofagus
alpina
, 17-V-2007,
M.V. Bianchinotti
,
R.M. Sa´nchez
,
MVB573
(HOLOTYPE designated here, BBB).
Etymology.
Refers to the geographical place in
Argentina where it was found.
Ascomata superficial to semi-immersed, globose,
subglobose to conical, dark brown to bright black
when dried, surface densely setose, 100–390 mm diam,
140–490 mm high. Setae mostly distributed on entire
ascoma but sometimes only on the upper half,
straight or curved, with acute tip, 0–4-septate, dark
brown, 49–125 mm long, 5–13 mm wide at the base, 1–
5mm wide at the apex. Ostiole papillate, 30–113 350–
100 mm. Ascomatal wall of textura angularis, two-
layered in longitudinal section, inner layer composed
of 4–6 rows of thin walled, flattened to angular, pale
ochreous cells, 5–13 33–8 mm; outer layer composed
of 5–7 rows of thick walled, elongated, dark brown
cells, 8–15 34–10 mm. Pseudoparaphyses narrow,
ramified, septate, forming a strong mesh, hyaline, 1–
3mm thick. Periphyses filling the entire papilla,
undulated, hyaline. Asci bitunicate, cylindrical to
clavate, short stipitate, eight-spored, 85–139 315–
33 mm (110 320 mm, n 516). Ascospores elongate
fusiform, straight or slightly curved, ends asymmetri-
cal, tapering at both ends, with one or two middle
cells near the apical end slightly broader than the
others, 9–18 septate, slightly constricted at the septa,
smooth, hyaline, 62–111 35–10 mm (83 37.5 mm, n
5100). Anamorph not known.
Habitat.
Scattered on bark of
Nothofagus alpina
in a
mixed forest of
N. alpina
,
N. dombeyi
and
N. obliqua
and on bark of
N. pumilio
in a pure forest.
Specimens examined
. ARGENTINA. CHUBUT PROV-
INCE: Pque. Nacional Los Alerces, on the way to Bagguilt
Lake, on bark of
Nothofagus pumilio
, 15-V-2007,
M.V.
Bianchinotti
,
R.M. Sa´ nchez
,
MVB542
, BBB; NEUQUE
´N
PROVINCE: Pque. Nacional Lanı´n, on Provincial Route
Number 48, before reaching Playas de Yuco, on bark of
N.
alpina
, 17-V-2007,
M.V. Bianchinotti
,
R.M. Sa´ nchez
,
MVB574
, BBB; NEUQUE
´N PROVINCE: Pque. Nacional
Lanı´n, Paso del Co´rdoba, 40u359440S, 71u089370W, 1245 m,
on
N. pumilio
, 18-I-2009,
M.V. Bianchinotti
,
R.M. Sa´ nchez
,
MVB776
, BBB.
KEY TO SPECIES OF
A
CANTHOSTIGMA
1.1 Ascospores less than 5 mm wide .............. 2
1.2 Ascospores $5mm wide . . ................. 6
2.1 Ascospores up to 4 mm wide ................ 3
2.2 Ascospores 4–5 mm wide . . ................. 5
3.1 Ascospores 22–27 33–3.5 mm, 5(–9)-septate ....
............................
A. revocatum
3.2 Ascospores more than 30 mm long ............ 4
4.1 Ascospores (40–)56–78(–95) 3(2–)2.5–3(–3.5)
mm, 10–14-septate ................
A. scopulum
4.2 Ascospores (75–)85–135(–150) 32.5–4 mm, 12–16-
septate .........................
A. filiforme
5.1 Ascospores (29–)38–51 34–4.5(–5) mm, (4–)6–8-
septate ......................
A. longisporum
5.2 Ascospores (38–)48–64.5 34–4.5 mm, 6–12-septa-
te ...............................
A. ellisii
6.1 Ascospores up to 60 mm long ............... 7
6.2 Ascospores .60 mm long . ................. 9
7.1 Ascospores with symmetrical ends, not or slightly
constricted at septa ....................... 8
7.2 Ascospores with asymmetrical ends, constricted at
septa, 40–50 35–6.5 mm, 7–10-septate . ........
........................
A. septoconstrictum
8.1 Ascospores 30.5–35.5(–42) 35–6 mm, (5–)6–7(–8)-
septate ......................
A. perpusillum
8.2 Ascospores 40–55(–63) 3(5–)6–7(–7.5) mm, 10–
14-septate ......................
A. minutum
9.1 Ascospores with symmetrical ends, 60–90 35–
7.5 mm, 14–18-septate .........
A. multiseptatum
9.2 Ascospores with asymmetrical ends, 62–111 35–
10 mm, 9–18-septate ..............
A. patagonica
RESULTS
The final ITS and LSU alignments contained 57 and 75
taxa and consisted of 325 and 536 bp respectively after
the removal of missing data and ambiguous regions.
The ML trees based on ITS and LSU data are provided
(FIGS. 3, 4 respectively). GenBank accession numbers
are given after taxon names in the phylogenetic trees.
In both analyses
A. patagonica
occurs closely related to
Helicoma vaccinii
in the moderately supported
Tubeu-
fia cerea
clade and is highly supported as a sister taxon
to
H. vaccinii
in the LSU tree.
Acanthostigma
patagonica
and
H. vaccinii
possess 14 bp differences
out of 407 bp (3.5%) in the unadulterated ITS dataset
and nine bp differences out of 638 bp (1.4%) in the
LSU dataset before the removal of ambiguous regions.
The ITS data are based on the highly conserved 5.8S
SA
´NCHEZ ET AL.:
A
CANTHOSTIGMA PATAGONICA
225

region and the variable ITS2 region. Although the
entire ITS region was successfully PCR amplified for
the new species, the sequence quality of the ITS1
region unfortunately was poor so intraspecific com-
parisons of complete ITS sequences cannot be made at
this time. No anamorph was found associated with
A.
patagonica
on the substrates.
DISCUSSION
The new species of
Acanthostigma
was compared to
all nine species currently accepted in the genus.
Acanthostigma patagonica
is most similar to
A. multi-
septatum
Promputtha & A.N. Mill. in that both species
possess long ascospores with the greatest number of
FIG.1.
Acanthostigma patagonica
(all from Holotype). A. Ascoma on substrate. B. Longitudinal section through ascomatal
wall. C. Ascospore 16-septate (in phloxine). D. Asci. E–F. Ascospores (in water). G. Ascospore viewed with fluorescence
microscopy. Bars: A 5200 mm, B 520 mm, C–G 510 mm.
226 MYCOLOGIA

septa in the genus. However
A. patagonica
has slightly
larger ascomata and setae and possesses wider
ascospores that are asymmetrical.
Acanthostigma
patagonica
also can be compared with
A. filiforme
Promputtha & A.N. Mill. and
A. scopulum
(Cooke &
Ellis) Peck, which overlap with
A. patagonica
in
ascospore length and number of septa. However both
species can be distinguished from
A. patagonica
by
their narrower ascospores.
Acanthostigma patagonica
is not closely related to any other
Acanthostigma
species in the phylogenetic analyses (FIGS. 3, 4).
According to Sa´nchez and Bianchinotti (2010),
from the species of
Acanthostigma
described by
Spegazzini (1884, 1887, 1898, 1909), three (
A.
FIG.2.
Acanthostigma patagonica
. A. Longitudinal section of ascoma. B. Setae. C. Ascus. D. Ascospores. Bars: A 550 mm, B
522 mm, C–D 510 mm.
SA
´NCHEZ ET AL.:
A
CANTHOSTIGMA PATAGONICA
227

FIG. 3. Tubeufiaceae phylogeny (ln L 521883.7) generated from a PhyML analysis of 57 ITS sequences showing the
phylogenetic placement of
A. patagonica
(in boldface). Thickened branches indicate Bayesian posterior probabilities $95%,
while numbers above or below branches refer to PhyML bootstrap values $50%. The tree is rooted with taxa in the
Tubeufia
cerea
clade based on the LSU analysis.
228 MYCOLOGIA

FIG. 4. Tubeufiaceae phylogeny (ln L 522619.3) generated from a PhyML analysis of 75 LSU sequences showing the
phylogenetic placement of the newly sequenced taxa (in boldface) in this study. Thickened branches indicate Bayesian
posterior probabilities $95%, while numbers above or below branches refer to PhyML bootstrap values $50%. Two species of
Botryosphaeria
are outgroups.
SA
´NCHEZ ET AL.:
A
CANTHOSTIGMA PATAGONICA
229

dimerosporioides
,
A. gnaphaliorum
and
A. imperspi-
cuum
) should be excluded because they possess three-
septate ascospores and appear to have unitunicate
asci. The fourth,
A. guaraniticum
, considered species
dubia, differs from
A. patagonica
in having smaller, 5–
6-septate ascospores (35 36–6.5 mm).
Sa´nchez and Bianchinotti (2010) reported
A.
minutum
from Lanı´n National Park, Neuque´n Prov-
ince, Argentina, on
Nothofagus dombeyi
and
N.
pumilio
. They concluded that their specimens mor-
phologically agreed with
A. minutum
(Fuckel) Sacc.,
differing only in a lesser number of septa in the
ascospores. However in the present phylogenetic
analyses the material from South America does not
group with the North American specimens (FIG.4)
suggesting this may be a new species. Additional
specimens of putative
A. minutum
from Argentina
should be included in further phylogenetic analyses
before a new species is described.
ACKNOWLEDGMENTS
The Consejo Nacional de Investigaciones Cientı´ficas y
Tecnolo´gicas (CONICET: PIP 5660 and PIP 80101000),
the Universidad Nacional del Sur (UNS: PGI) and the
National Science Foundation (DEB-08-44722) are thanked
for financial support. We thank Marta Garelli for assistance
with the Latin description.
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´NCHEZ ET AL.:
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CANTHOSTIGMA PATAGONICA
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