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Brianaria (Psoraceae), a new genus to accommodate the Micarea sylvicola group

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The new genus Brianaria S. Ekman & M. Svensson is introduced for the Micarea sylvicola group, with the new combinations Brianaria bauschiana (Körb.) S. Ekman & M. Svensson, B. lutulata (Nyl.) S. Ekman & M. Svensson, B. sylvicola (Flot. ex Körb.) S. Ekman & M. Svensson and B. tuberculata (Sommerf.) S. Ekman & M. Svensson. The new genus is characterized by a chlorococcoid, non-micareoid photobiont, small, convex apothecia without an excipulum, an ascus of the ‘Psora-type’, 0–1-septate ascospores, dimorphic paraphyses, and immersed pycnidia containing bacilliform conidia. Brianaria is shown to form a monophyletic group in the Psoraceae, where it is probably the sister group to Psora and Protoblastenia.
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The Lichenologist 46(3): 285–294 (2014) 6British Lichen Society, 2014
doi:10.1017/S0024282913000613
Brianaria (Psoraceae), a new genus to accommodate the
Micarea sylvicola group
Stefan EKMAN and Ma
˚ns SVENSSON
Abstract: The new genus Brianaria S. Ekman & M. Svensson is introduced for the Micarea sylvicola
group, with the new combinations Brianaria bauschiana (Ko¨rb.) S. Ekman & M. Svensson, B.lutulata
(Nyl.) S. Ekman & M. Svensson, B. sylvicola (Flot. ex Ko¨rb.) S. Ekman & M. Svensson and B.tuber-
culata (Sommerf.) S. Ekman & M. Svensson. The new genus is characterized by a chlorococcoid,
non-micareoid photobiont, small, convex apothecia without an excipulum, an ascus of the ‘Psora-
type’, 0–1-septate ascospores, dimorphic paraphyses, and immersed pycnidia containing bacilliform
conidia. Brianaria is shown to form a monophyletic group in the Psoraceae, where it is probably the
sister group to Psora and Protoblastenia.
Key words: lichens, micareoid, Pilocarpaceae,Protoblastenia,Psora
Accepted for publication 22 May 2013
Introduction
The genus Micarea, described by Fries (1825)
and conserved with M. prasina as the type
species (Coppins 1989, ICBN appendix
III), was resurrected from oblivion at the
end of the 19th century ( Hedlund 1892). In
his precocious revision of small crustose li-
chens, Hedlund (1892) emended Micarea to
include 20 species. As circumscribed by him,
the genus included species with unicellular
or transversely septate ascospores, branched
and anastomosing, apically unthickened par-
aphyses, an excipulum composed of para-
physis-like hyphae, a small-celled photobiont
[4–8(–9) mm, later known as ‘micareoid’]
and immarginate, often tuberculate apothecia.
Several decades later, Zahlbruckner (1921–
40) introduced an artificial but highly influ-
ential taxonomy in his Catalogum Lichenum
Universalis, in which lecideoid lichens were
sorted according to spore septation. Conse-
quently, the genus Micarea was split and its
species again transferred to other genera,
mainly Lecidea,Catillaria,andBacidia.When
the Zahlbrucknerian ice sheet slowly melted
in the 1960s and 70s, the pursuit for a more
natural classification was revitalized and fol-
lowed by the revival of previously described
genera, as well as the description of numer-
ous new ones.
One of the first to yet again re-establish
Micarea was Anderson (1974), who trans-
ferred Lecidea tuberculata Sommerf. to this
genus. The inclusion of a species with a non-
micareoid photobiont was in effect an emen-
dation of Hedlund’s generic delimitation,
although Anderson did not discuss this. Sub-
sequently, Ve
ˇzda & Wirth (1976) published a
new key to the genus, accepting Hedlund’s
work but also adding several species, such as
Micarea bauschiana (Ko¨rb.) Ve
ˇzda & Wirth,
M.lutulata (Nyl.) Coppins (as M.umbrosa
Ve
ˇzda & Wirth) and M.sylvicola (Flot.)
Ve
ˇzda & Wirth. They noted that M.bauschi-
ana and M.lutulata were probably closely re-
lated, but did not comment on the similari-
ties between these species and M.sylvicola
and M.tuberculata.
In his breakthrough revision of the Euro-
pean species of Micarea, Coppins (1983)
noted that the genus included several small
S. Ekman: Museum of Evolution, Uppsala University,
Norbyva¨gen 16, SE-75236 Uppsala, Sweden.
Email: stefan.ekman@em.uu.se
M. Svensson: Department of Ecology, Swedish Univer-
sity of Agricultural Sciences, P. O. Box 7044, SE-75007
Uppsala, Sweden.
groups of apparently closely related species.
He recognized 11 infrageneric groups in
Micarea, one of which (group ‘I’, hereafter
known as the Micarea sylvicola group) con-
sisted of M.bauschiana, M.lutulata,M.sylvi-
cola, and M.tuberculata. Coppins concluded
that this group was ‘almost worthy of sub-
generic status’.
With the advent of molecular methods in
taxonomy came the opportunity to test this
hypothesis. Andersen & Ekman (2005), in a
phylogeny based on mitochondrial ribosomal
DNA, showed that Micarea was paraphyletic,
and that several of the infrageneric groups
identified by Coppins probably deserved
generic recognition. In this phylogeny, the
Micarea sylvicola group was represented by
M.bauschiana and M.sylvicola, and formed
a highly supported group together with Psora
decipiens (Hedw.) Hoffm. in the Psoraceae.
Subsequent studies based on three or more
loci and better taxon sampling confirmed
that the family Psoraceae consists of Protoblas-
tenia,Psora and the Micarea sylvicola group
(Ekman et al. 2008; Ekman & Blaalid 2011;
Schmull et al. 2011). In these analyses, the
M.sylvicola group has been represented by
M. sylvicola only (Ekman et al. 2008; Schmull
et al. 2011) or M. bauschiana and M. sylvicola
(Ekman & Blaalid 2011).
As no name at genus level appears to be
available for Micarea sylvicola and its close
relatives, a new genus is described here to ac-
commodate it. Furthermore, although there
is ample evidence to show that the M.sylvi-
cola group belongs in the Psoraceae, previous
phylogenetic analyses did not include M. lu-
tulata and M. tuberculata. In order to demon-
strate that these species are unequivocally
close relatives of M. sylvicola and M. bauschi-
ana, we present a new phylogeny that in-
cludes all four species in the same analysis.
Materials and Methods
Taxon sampling
We selected 27 species of the Pilocarpaceae in the sense
of Andersen & Ekman (2005) and Ekman et al. (2008),
and Psoraceae in the sense of Andersen & Ekman (2005)
and Ekman & Blaalid (2011) (i.e. including Psora,Proto-
blastenia, and the ‘Micarea sylvicola group’ described
here as Brianaria, but excluding Eremastrella,Glyphopeltis,
and Psorula). We did not include the genus Protomicarea
in the Psoraceae, as this genus was shown by Schmull
et al. (2011) not to belong in this family. Protomicarea
was tentatively referred to the Psoraceae by Hafellner &
Tu
¨rk (2001) and Lumbsch & Huhndorf (2010) but was
not included in the phylogenetic analysis by Ekman &
Blaalid (2011). Two species (Brianaria sylvicola and B.
tuberculata) were represented by two terminal units, re-
sulting in an ingroup with 29 members. We used Sphaer-
ophorus globosus as outgroup, the selection of which was
based on the phylogeny by Mia˛ dlikowska et al. (2006),
in which the Sphaerophoraceae is sister to the Psoraceae
and Ramalinaceae.
Marker selection and sequence acquisition
For 26 of the 30 included terminals, we downloaded
sequence data from GenBank (http://www.ncbi.nlm.
nih.gov/genbank/) representing three different genes,
viz. the largest subunit of the RNA polymerase II gene
(RPB1), the internal transcribed spacer ( ITS) region
(including ITS1, 5.8S, and ITS2) of the nuclear riboso-
mal RNA gene, and the small subunit of the mitochon-
drial ribosomal RNA gene (referred to here as mrSSU).
For the remaining four terminals, we produced new
sequence data as described below. Most terminals were
composed of sequence data from a single herbarium
specimen. To avoid excessive amounts of missing data
in the resulting matrix, we accepted two cases with
terminals composed of sequence data from more than
one herbarium specimen, viz.Psora decipiens and P.
rubiformis. The sequence data used for this study is sum-
marized in Table 1.
PCR amplification and DNA sequencing
New ITS and mrSSU sequences were generated from
four specimens representing three species of Brianaria,
namely B. lutulata,B. sylvicola, and B. tuberculata. In ad-
dition, several unsuccessful attempts were made to obtain
RPB1 sequences. Laboratory methods generally con-
formed to Ekman & Blaalid (2011), except that we used
the Hot StarTaq Mastermix Kit (Qiagen) and QIAquick
PCR Purification Kit (Qiagen). In a few cases, we per-
formed direct PCR from apothecial sections without
preceding DNA extraction (Wolinski et al. 1999).
Sequence alignment
Sequences were aligned using MAFFT version 6.935
(Katoh & Toh 2008a). The three ITS components, ITS1,
5.8S and ITS2, were aligned separately using the X-INS-i
algorithm with MXSCARNA pairwise structural align-
ments and Contrafold base-pairing probabilities (Katoh
& Toh 2008b). A structural euascomycete mrSSU refer-
ence alignment was downloaded from the Comparative
RNA Web Site (http://rna.ccbb.utexas.edu; Cannone et
al. 2002). This alignment was used as a profile, to which
our mrSSU sequences were added using the L-INS-i
algorithm. This choice was motivated by the fact that
the reference alignment included complete or near-
THE LICHENOLOGIST286 Vol. 46
complete mrSSU sequences that were much longer than
our sequences. Subsequently, the reference alignment
was removed and gap-only sites stripped. RPB1 sequences
were aligned using the G-INS-i algorithm at the amino
acid level and subsequently back-translated into DNA
sequences. The choice of algorithm was motivated by
the very few expected gaps in the RPB1 sequences once
introns had been removed.
Ambiguous alignments were filtered out using Ali-
score version 2.0 (Misof & Misof 2009). All pairs of
taxa were used to calculate the consensus profile. Gaps
were treated as ambiguities and window size was set to
4. These are the most conservative options available in
Aliscore.
Phylogenetic analysis
Maximum likelihood (ML) estimation of phylogeny
was performed using GARLI version 2.0 (Zwickl 2006)
under a single GTR model with rate heterogeneity across
sites, modelled as a discretized gamma distribution with
six categories and a proportion of invariable sites. Phy-
logeny estimates were produced for 1) each of the three
genes for the purpose of assessing potential gene tree
conflicts, and 2) the complete concatenated but unparti-
tioned data, primarily for the purpose of generating an
empirical branch-length prior for downstream Bayesian
inferences. For the concatenated data, we performed
1000 optimizations from starting trees generated by
stepwise random addition of taxa, every possible attach-
ment point being evaluated. For all data sets, branch
support was assessed using 1000 non-parametric boot-
strap replicates. Majority-rule consensus trees from the
single genes were subsequently input into Compat.py
(Kauff & Lutzoni 2002) for conflict identification above
70% bootstrap support.
The concatenated data were divided into seven poten-
tial character subsets, ITS1, 5.8S, ITS2, mrSSU, as well
as RPB1 first, second and third codon positions. These
subsets were subsequently input into PartitionFinder
version 1.0.1 (Lanfear et al. 2012) for a simultaneous ex-
haustive search for thebest-fitting partitioning scheme and
the best-fitting model of each partition under the con-
straint that only models with one, two, or six substitution
Table 1. GenBank accession numbers for DNA sequences included in this study. Newly obtained sequences are in bold.
Dashes represent missing data
ITS mrSSU RPB1
Bapalmuia palmularis AY756457 AY567781
Brianaria bauschiana — AY567770
B. lutulata JX983582 JX983586
B. sylvicola I JX983583 JX983587
B. sylvicola II AY567769 AY756392
B. tuberculata I JX983584 JX983588
B. tuberculata II JX983585 JX983589
Byssolecania variabilis AY756458 AY567780
Byssoloma leucoblepharum AY756459 AY567778 AY756380
Micarea adnata AY756468 AY567751 AY756388
M. alabastrites AY756469 AY567764 AY756389
M. assimilata AY756470 AY567739
M. byssacea AY756485 AY567749
M. erratica AY756475 AY567737 AY756390
M. lignaria var. lignaria AY756481 AY567748
M. lithinella AY756482 AY567734
M. melaena AY756483 AY567743
M. misella AY756486 AY567752
Protoblastenia calva EF524319 DQ986904 EF524338
P. rupestris EF524318 — EF524329
Psora californica EF524322 EF524292 EF524334
P. cerebriformis EF524325 EF524293 EF524335
P. decipiens EF524326 AY567772 EF524337
P. globifera EF524323 EF524294 EF524331
P. nipponica EF524312 — EF524336
P. pacifica EF524314 EF524297 EF524332
P. rubiformis HQ650620 AY756374 DQ986831
P. tuckermanii EF524317 —
P. vallesiaca EF524324 EF524291
Sphaerophorus globosus AY256769 AY256751 AY756424
2014 BrianariaEkman & Svensson 287
rate categories could be selected. We used the Bayesian
Information Criterion to select among models and parti-
tioning schemes.
We performed Bayesian phylogenetic inference using
Markov chain Monte Carlo (MCMC) as implemented
in PHYCAS version 1.2.0 (Lewis et al. 2010). Five dif-
ferent analyses were performed for the purpose of quan-
tifying the support for the Psoraceae and the new genus
Brianaria under a variety of model assumptions and
branch-length priors.
The first analysis assumed independent best-fitting
models for each of the partitions inferred by Partition-
Finder. Rate heterogeneity was, when applicable, mod-
elled as a discretized gamma distribution with six catego-
ries. We used flat Dirichlet priors on state frequencies, as
well as the substitution rate matrix for six-rate models, a
beta prime (1, 1) distribution on the transition and trans-
version rates for two-rate models, a uniform (0001,
200) on the gamma distribution shape parameter, a uni-
form (0, 1) on the proportion of invariable sites, and a
flat relative rate distribution, a transformed Dirichlet
distribution described by Fan et al. (2011), on the subset
rate multipliers. The prior on branch lengths was set to
an exponential with rate parameter 25. This rate param-
eter was estimated by fitting an exponential distribution
to the branch lengths obtained from the ML analysis.
Curve-fitting was performed with EasyFit Professional
version 5.5 (MathWave Technologies).
The second analysis was identical to the first, except
that the prior on branch lengths was set to an exponen-
tial seeded by an exponential hyperprior with rate 10.
This is a hierarchical model on branch lengths, in which
the mean of the prior distribution is not fixed but treated
as a parameter to be estimated. As a hierarchical model
on branch lengths does not impose a specific exponen-
tial distribution, it should have less influence on pos-
terior parameter distributions (Ekman & Blaalid 2011;
Rannala et al. 2012).
The third analysis was identical to the first except we
allowed trees with polytomies to be sampled according
to the model of Lewis et al. (2005). The purpose of this
analysis was to investigate whether the forced sampling
of fully resolved trees in other analyses could cause ex-
cessive branch support (Lewis et al. 2005). We chose to
set the polytomy prior (C) to 1. Thereby every tree to-
pology was treated as a priori equally probable, regard-
less of the number of internal nodes. As there are c.1000
times more unique topologies with at least one polytomy
than there are fully resolved 30-taxon trees (Felsenstein
2004), our prior amounts to treating the class of polyto-
mous trees as a priori far more likely than the fully re-
solved ones.
The fourth analysis was identical to the first, except
that we used a fixed branch-length prior drawn from a
gamma distribution with shape 0647 and scale 0 062.
The parameterization of the gamma distribution was
taken from the above-mentioned EasyFit curve-fitting
procedure. The rationale behind this analysis was that a
branch-length prior violating the true distribution of
branch lengths may bias posterior probabilities (Kolacz-
kowski & Thornton 2007).
The fifth analysis was identical to the first, except that
independent GTR+I+Gmodels were used for the sub-
sets. This analysis was carried out to safeguard against
potential overestimates of branch support, in case of hid-
den inadequacies in the best-fitting models (Huelsenbeck
& Rannala 2004). It should be noted, however, that the
potential adequacy of the GTR+I+Gmodel is restricted
to temporally reversible and homogeneous processes.
Each analysis included three runs, each with one cold
and three heated chains, the hottest with power 05. We
ran each analysis for 200 000 generations, sampling every
100th generation. The reason for the smaller number of
generations compared to the commonly used MrBayes
(Ronquist & Huelsenbeck 2003) is that a generation is
defined differently in PHYCAS, one generation in this
software corresponding to c.100generationsinMrBayes.
Average standard deviations of splits (with frequency
b01) between runs, identical to the default measure
used to diagnose MrBayes runs, were calculated from
summaries provided by the ‘showsplits’ command in
AWTY online (Wilgenbush et al. 2004) after having re-
moved the first half of each tree sample as burn-in. Mar-
ginal likelihoods of the data were calculated with Tracer
version 1.5 (Rambaut & Drummond 2009) using the
importance sampling estimator originally suggested by
Newton & Raftery (1994) and modified by Suchard et al.
(2003). Importance sampling, as well as the widely used
harmonic mean, have, however, been shown to be unreli-
able when comparing models with high dimensionality
(Lartillot & Philippe 2006). Therefore, we also calculated
marginal likelihoods using the stepping-stone procedure
described by Fan et al. (2011) and implemented in PHY-
CAS. This implementation requires a fixed tree topology,
for the purpose of which we used the majority-rule con-
sensus trees with all compatible groups. We took 1000
samples from each of 21 stepping stones, with the excep-
tion that 2000 samples were taken from the posterior.
The fixed-topology requirement rendered stepping-stone
estimation impossible under the polytomy model.
Results
The resulting filtered alignment consisted of
1731 sites, 352 of which belonged to the
ITS, 807 to the mrSSU, and 642 to the
RPB1. The number of variable alignment
positions (assuming that gaps are treated as
missing data) was 166, 300, and 289, respec-
tively in the ITS, mrSSU, and RPB1. The
total amount of missing data, including
gaps, was 28%, RPB1 being clearly over-
represented due to technical difficulties am-
plifying this gene successfully.
We did not record any conflicts between
the three genes. The best-fitting partitioning
scheme, given the seven potential partitions,
THE LICHENOLOGIST288 Vol. 46
included five partitions: ITS1 + ITS2, 5.8S,
mrSSU, RPB1 first and second codon posi-
tions, and RPB1 third codon positions. The
best-fitting models for each of these partitions
was found to be SYM+G, K80+I, GTR+I+G,
K80+I+G, and K80+G, respectively. The
total number of free parameters in this model,
including the subset rate multipliers, was 26,
compared to the 54 free parameters in the
analysis with five independent GTR+I+G
models. Kolmogorov-Smirnov tests of good-
ness-of-fit of preliminary maximum-likeli-
hood branch lengths did not reject an expo-
nential distribution (D ¼013, P¼030),
although a gamma distribution had better fit
(D ¼009, P¼075). Average standard de-
viations of split frequencies between MCMC
runs ranged from 0007 to 0 009 depending
on the analysis, which is low enough to con-
clude that MCMC analyses had converged
and that our tree samples represent valid
samples from the posterior distributions.
The five Bayesian analyses are summar-
ized in Table 2. Depending on model and
branch-length prior, the posterior probability
of the branch uniting the Psoraceae ranges
from 098 to 100, whereas the posterior
probability of the branch uniting the genus
Brianaria ranges from 095 to 0 98. By com-
parison, ML bootstrap proportions were
073 for the Psoraceae and 089 for Brianaria.
Stepping-stone estimation of marginal likeli-
hoods seems to provide better resolution to
discriminate between models and priors than
importance sampling. A majority-rule con-
sensus tree with all compatible groups from
the first of the Bayesian inferences, the one
with independent best-fitting model for each
partition and an exponential (25) branch-
length prior, is shown in Fig. 1.
Discussion
The phylogenetic analysis indicates that Bria-
naria, including B. bauschiana,B. lutulata, B.
sylvicola and B. tuberculata, forms a monophy-
letic group within the Psoraceae, which is in
agreement with the findings of Andersen &
Ekman (2005), Ekman & Blaalid (2011) and
Schmull et al. (2011). Evidence also suggests
that Brianaria is the sister group to the rest of
the currently known members of the Psora-
ceae,viz. Psora and Protoblastenia.Conversely,
there is no indication that Brianaria belongs
in Micarea or the Pilocarpaceae, where its spe-
cies have previously been included.
Differences in tree lengths and support for
the Psoraceae and Brianaria between analyses
based on best-fitting models for each parti-
tion are minuscule, whether or not allowing
for polytomies and irrespective of the partic-
ular branch-length prior. Marginal likelihood
differences are modest as estimated by im-
portance sampling, whereas the more reliable
stepping-stone estimation indicates reason-
ably strong support for a gamma distributed
branch-length prior. This is not surprising,
as the initial fitting of ML branch lengths
provided better fit to a gamma distribution
than to an exponential distribution. The
analysis based on independent GTR+I+G
models, on the other hand, resulted in a
much worse marginal likelihood than other
analyses, whether estimated by importance
Table 2. Overview of Bayesian phylogenetic analyses performed with PHYCAS. The ‘‘best’ model refers to the combination
of partition models selected by PartitionFinder. Polytomies were modelled according to Lewis et al. (2005). Stepping-stone
estimation of the marginal likelihood was not possible because of a fixed-topology requirement in the implementation. Support
for the Psoraceae refers to the posterior probability of the node uniting Brianaria, Protoblastenia, and Psora
Model Branch-length prior
Branch-length
hyperprior
Importance
sampling lnL
Stepping-stone
lnL
Average
tree length
Psoraceae
support
Brianaria
support
Best Exponential (25) None -- 10960308 -- 11837 908 2517 0 98 100
Best Exponential Exponential (10) -- 10960513 -- 11833 221 2554 0 98 099
Best+polytomies Exponential (25) None -- 10967942 — 2519 0 95 098
Best Gamma (0647, 0 062) None --10959854 -- 11824 099 2558 0 97 099
5(GTR+I+G) Exponential (25) None -- 11149 117 -- 12088786 2 524 097 1 00
2014 BrianariaEkman & Svensson 289
sampling or stepping stones, suggesting that
it suffers from severe overfitting. However,
whereas underfitting is known to cause severe
topological bias, overfitting seems to be much
less of a problem ( Huelsenbeck & Rannala
2004; Lemmon & Moriarty 2004). Conse-
quently, the overfitted analysis provides some
indication that support for the Psoraceae and
Brianaria is not overestimated. Ekman &
Blaalid (2011), based on more characters
but fewer taxa in Brianaria, found support
for a Psoraceae including Brianaria to be
096 and 095 for independent best-fitting
and GTR+I+Gpartition models, respec-
tively, when integrating over a wide interval
of exponential branch-length priors. They
did not, however, investigate the effect of
allowing for polytomies or other than expo-
nential branch-length priors. Nominal ML
bootstrap support values for the Psoraceae
and Brianaria nodes (073 and 089, respec-
tively) seem to be in line with the posterior
probabilities, given previous suggestions that
bootstrap support at 070 corresponds to a
095 probability of a clade being real (Hillis
& Bull 1993). This approximation assumes
that rates of change are moderate and more
or less equal across the tree, which may hold
true in our case. Altogether, support for the
monophyly of Psoraceae and Brianaria appears
to be high and does not seem to be affected by
any of the known causes of inflated posterior
probabilities in Bayesian inference of phylog-
eny, viz. model underfitting, branch-length
prior misspecification, or the forcing of fully
resolved trees.
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Fig. 1. Majority-rule consensus tree with all compatible groups, average branch lengths, and posterior probabilities
of nodes resulting from Bayesian MCMC using PHYCAS under independent best-fitting models for five partitions
and an exponential branch-length prior with rate parameter 25. Familial affiliations are indicated.
THE LICHENOLOGIST290 Vol. 46
Taxonomy
Brianaria S. Ekman & M. Svensson gen.
nov.
MycoBank No.: MB803358
Distinguished from Micarea s. str. by having a non-
micareoid photobiont, dimorphic paraphyses, and a
wider tube structure in the tholus, from Psora by the
crustose thallus, lack of oxalate and anthraquinone
crystals in the apothecia, and from Protoblastenia by the
absence of anthraquinones in the apothecia and the lack
of an excipulum.
Type species: Brianaria sylvicola (Flot. ex Ko¨rb.) S.
Ekman & M. Svensson.
Thallus scurfy-granular-verruculose areo-
late, grey-greyish green. Photobiont of two
types, 1) chlorococcoid, 5–12(–15) mmor2)
irregularly ellipsoid and up to 15 10 mm.
Ascomata immarginate, convex-hemispher-
ical, often becoming tuberculate, 015–070
(–120) mm diam. Excipulum absent. Hy-
pothecium 80–200 mm tall, composed of
interwoven hyphae 1–3 mm thick. Hymenium
30–75 mm. Paraphyses dimorphic, either
evenly distributed, sparingly branched, often
anastomosing below, 08–15mm wide or
fewer in number, single; or in fascicles, sim-
ple or occasionally forked above, distinctly
septate, 15–30mm wide, up to 4 mm api-
cally. Ascospores non-septate (sometimes 1-
septate in B. tuberculata), 55–12015–
50mm. Asci 8-spored, cylindrical-clavate,
25–45 7–12 mm. Tholus with a wide, dark
tube structure that expands towards the top,
without a pale axial body (‘Psora-type’ sensu
Ekman et al. 2008).
Pycnidia immersed in thallus, 004–020
mm diam., black. Conidiogenous cells ecylin-
drical, 5–10 10–15mm. Conidia bacilli-
form to oblong to obovoid, 3–7 1–2 mm.
Chemistry. No lichen substances detected
by TLC. Three different pigments occur in
the apothecia of Brianaria,viz. blue-green,
K-- , N+ red (‘Pigment A’ sensu Coppins
1983, ‘Cinereorufa-green’ sensu Meyer &
Printzen 2000), brown, K-- ,N
-- or N+
orange-brown (‘Pigment F’ sensu Coppins
1983) and purple, K+ green, N+ red (‘Pig-
ment B’ sensu Coppins 1983, ‘Melaena-red’
sensu Meyer & Printzen 2000).
Etymology. The genus is named in honour
of Brian Coppins, in recognition of his out-
standing contribution to the taxonomy of
crustose lichens in general, and to the genus
Micarea in particular.
Ecology. All four species of Brianaria are
essentially saxicolous and prefer shaded acid
rock, often in rain-protected situations. Other
substrata (e.g., wood, rusted iron) are occa-
sionally inhabited, especially by B.sylvicola.
Notes. As delimited by Lumbsch & Huhn-
dorf (2010), the Psoraceae consists of the
genera Eremastrella,Glyphopeltis,Protoblastenia,
Psora, Psorula and possibly also Protomicarea.
Of these, Eremastrella,Glyphopeltis, Psorula
and Protomicarea have subsequently been
shown not to belong in this family ( Ekman
& Blaalid 2011; Schmull et al. 2011). Psora,
the type genus of the family, differs from
Brianaria in having a well-developed squa-
mulose thallus, anthraquinones in the epi-
thecium and calcium oxalate in the hypo-
thecium (Timdal 1984, 2002). Protoblastenia
differs in having anthraquinones in the
apothecial tissues, in having an exciple com-
posed of parallel-radiate hyphae, and in hav-
ing a preference for calcareous substrata
(Kainz 2004; Kainz & Rambold 2004). Bria-
naria,Psora and Protoblastenia are similar in
having asci of the ‘Psora-type’ and immersed
pycnidia containing bacilliform conidia.
Although not closely related, Brianaria
is anatomically similar to Micarea, differing
from Micarea s. str. (i.e. M.prasina Fr. and
closely related species) primarily by having a
non-micareoid photobiont, dimorphic para-
physes, and a slightly different tholus. Al-
though similar, tube structures in members
of the Pilocarpaceae tend to be thin without
or with a slight tendency to expand near the
apex. This appearance contrasts with the thick
tube that expands near the apex found in the
Psoraceae, including Brianaria (see Fig. 1 in
Ekman et al. 2008).
In spite of the exclusion of Brianaria, the
genus Micarea still includes species with a
non-micareoid photobiont, such as M. lyn-
ceola (Th. Fr.) Palice and M.myriocarpa V.
Wirth & Ve
ˇzda ex Coppins, as well as species
2014 BrianariaEkman & Svensson 291
with dimorphic paraphyses, such as M.
botryoides (Nyl.) Coppins and M. lithinella
(Nyl.) Hedl. (Andersen & Ekman 2005;
Coppins 2009).
The photobiont of Brianaria remains un-
identified to genus. The photobiont in Psora
decipiens and P. globifera has been identified
as Myrmecia biatorellae (Geitler 1963; Galun et
al. 1971; Tschermak-Woess 1988), although
Schaper & Ott (2003) claimed to have found
a species of Asterochloris (Schaper & Ott 2003)
in Psora decipiens. Both Myrmecia and Astero-
chloris are members of the Trebouxiaceae in
the Trebouxiales (Guiry & Guiry 2012). The
primary ‘micareoid’ photobionts in Micarea
prasina,M. peliocarpa, and M. misella,on
the other hand, appear to be Elliptochloris
bilobata,E. reniformis, and E. subsphaerica
(Voytsekhovich et al. 2011). The genus Ellip-
tochloris has an unsettled position in the Pra-
siolales (Guiry & Guiry 2012).
Descriptions of the species of Brianaria, as
well as heterotypic synonyms, can be found
in Coppins (1983) and Czarnota (2007).
Brianaria bauschiana ( Ko
¨rb.) S. Ekman
& M. Svensson comb. nov.
MycoBank No.: MB803360
Biatora bauschiana Ko¨rb., Parerga lich.: 157 (1860).—
Lecidea bauschiana (Ko¨rb.) Lettau in Hedwigia 55: 28
(1914).—Micarea bauschiana (Ko¨ rb.) Ve
ˇzda & Wirth in
Folia Geobot. Phytotax.11: 95 (1976); type: Germany,
Baden-Wu
¨rttemberg, ‘‘auf Porphyr bei Baden,’Bausch,
distributed as Rabenhorst: Lich. Europ. 648 (M—lecto-
type, selected by Ve
ˇzda & Wirth 1976, not seen; UPS—
isotype, seen).
Brianaria lutulata (Nyl.) S. Ekman &
M. Svensson comb. nov.
MycoBank No.: MB803361
Lecidea lutulata Nyl. in Flora Jena 56: 297 (1853).—
Micarea lutulata (Nyl.) Coppins in D. Hawksw., P.
James & B. Coppins, Lichenologist 12: 107 (1980); type:
British Isles, ‘‘Jersey, Rozel meadow, bases of rocks,’
1873, Larbalestier (H-NYL 10696 —lectotype, selected
by Coppins 1983, seen).
Brianaria sylvicola (Flot. ex Ko
¨rb.)
S. Ekman & M. Svensson comb. nov.
MycoBank No.: MB803359
Lecidea sylvicola Flot., Lich. Schles.: 171 (1829), nom. in-
val. (Art. 32.1d, 34.1a).—Lecidea sylvicola Flot. ex Ko¨rb.,
Syst. Lich. German.: 254 (1855).— Micarea sylvicola
(Ko¨rb.) Ve
ˇzda & Wirth in Folia Geobot. Phytotax. 11: 99
(1976); type: Czech Republic/Germany/Poland, Lich.
Schles. 171 (UPS —lectotype, selected by Hertel 1975,
seen).
Nomenclatural note. Flotow (1829) intro-
duced the name Lecidea sylvicola, but did not
himself consider this name valid (‘Lecidea syl-
vicola ad int.’) and did not provide a diagnosis
or reference to a validly published diagnosis.
The taxon was validated by Ko¨rber (1855),
who provided a diagnosis and made explicit
reference to nr. 171 in Flotow’s exsiccate.
The earliest known synonyms of L.sylvicola
are L.aggerata Mudd and L.incincta Nyl.,
both of which were published in 1861 (Cop-
pins 1983; Czarnota 2007).
Brianaria tuberculata (Sommerf.)
S. Ekman & M. Svensson comb.nov.
MycoBank No.: MB803362
Lecidea tuberculata Sommerf., Suppl. Fl. Lapp.: 160
(1826).—Micarea tuberculata (Sommerf.) R. A. Ander-
son in Bryologist 77: 46 (1974); type: Norway, Nordland,
Saltdalen, Fiskevaagmo¨llen, March 1822, Sommerfelt
(O—lectotype, selected by Coppins 1983, not seen;
UPS—isotype, seen).
We thank Katja Fedrowitz, Mattias Lif and Veera Tuovi-
nen for laboratory assistance.
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THE LICHENOLOGIST294 Vol. 46
... and Lecidella), which also fit within the circumscription of Lecidea s. lat. were excluded for the following reasons: the systematics of the Micarea is still not entirely resolved and many species included in this genus form a polyphyletic assemblage based on recent molecular studies (Andersen & Ekman 2005, Ekman & Svensson 2014, and the delimitation of species of Lecidella are based mainly on their secondary metabolites as key taxonomical characters but most of those substances can be only detected by sophisticated chemical techniques e.g., mass spectrometry, high-performance liquid chromatography, etc. (Leuckert & Knoph 1993, Knoph & Leuckert 1997. For that reason, these genera were excluded from this study. ...
... Micarea doliiformis differs in having slightly larger pycnidia (140-300 μm), much smaller conidia (3.5-4.7 × 1.5-2 μm), an effuse, pale greyish-green, granular thallus, and a reduced proper exciple. Also, other members of Brianaria and Micarea can be very similar to Bryobilimbia pallida but they can be easily separated by small convex immarginate apothecia (Coppins 1983, Ekman & Svensson 2014. Besides, three species of unknown systematic position, "Lecidea" betulicola (Kullh.) ...
Article
Lecidea Ach. in its broad sense, is one of the largest and most heterogeneous genus of lichenized fungi with a worldwide distribution and with diversity hotspots located in the temperate and polar regions. The genus belongs to a crust-like microlichen group and inhabits many different substrates (e.g., bark, rock, wood, soil, mosses). Lecidea does not form a coherent systematic entity, and previous studies have revealed it as a polyphyletic assemblage with species spread across various families within the Lecanoromycetes. The present study is a modern taxonomic revision of southern South America non-saxicolous lecideoid lichens based on morphological, anatomical and chemical characters. A total of 27 species belonging to ten genera are recognized. The current study reveals a substantial, previously hidden, diversity of lichens in Valdivian temperate and Magellanic subpolar forests; increasing the number of known lecideoid lichens in the studied area. Many new regional records are also reported including six species new to South America (Bryobilimbia hypnorum, Hertelidea botryosa, H. eucalypti, Japewiella tavaresiana, Placynthiella oligotropha, and Ramboldia brunneocarpa). The following species are here described as new to science: Bryobilimbia flakusii Rodr. Flakus sp. nov. (Argentina), B. pallida Rodr. Flakus sp. nov. (Argentina, Chile), Hertelidea printzenii Rodr. Flakus sp. nov. (Argentina), H. stipitata Rodr. Flakus sp. nov. (Argentina, Chile), “Lecidea” vobisii Rodr. Flakus sp. nov. (Argentina), and Ramboldia australis Rodr. Flakus sp. nov. (Argentina, Chile). All species are described and illustrated in detail, and an identification key to the species is provided. In addition, as a result of a revision of available type material, a list of 48 additional species excluded from this study, including brief remarks on their taxonomical affiliations, is provided.
... Characters that define the genus include its green photobiont comprising small, often-paired cells, commonly referred to as 'micareoid', the often-reduced apothecial excipulum, the 8-spored, Pilocarpaceae-type asci where the well-developed, amyloid tholus is penetrated by a narrow channel, with or without a darker staining tube structure, and simple to branched paraphyses; the hyaline ascospores vary from filiform to fusiform to ovoid-ellipsoid, and from simple to transversely multiseptate. The taxonomic history of Micarea has been summarized by several authors, for example Czarnota (2007) and Ekman & Svensson (2014). The modern framework for the species-rank taxonomy of Micarea was essentially erected by Coppins (1983) in his review of European taxa, and was based largely on thallus chemistry, apothecial anatomy and pigmentation, and ascospore and conidial characters. ...
... Coppins (1983) had already recognized that Micarea included several, well-defined species groups, some of which, such as Psilolechia (Coppins & Purvis 1987), were worthy of generic rank. Supported by anatomical, morphological and, more recently, DNAsequence data, this has seen the segregation out of Micarea of Szczawinskia (Funk 1983), Brianaria (Ekman & Svensson 2014) and Leimonis (Harris 2009). However, the genus remains a challenge. ...
Article
Thirty-five species of Micarea are recorded for Tasmania. Ten are described as new to science: M. ceracea Coppins & Kantvilas (also known from Victoria and New South Wales), characterized by a thallus containing perlatolic and didymic acids, pallid apothecia and 3(-4)-septate ascospores, 10-21 × 3·5-6 µm; M. cinereopallida Coppins & Kantvilas (also known from Chile), with a granular to coralloid, goniocyst-like thallus containing superlatolic acid, pallid to piebald apothecia and (0-)1-septate ascospores, 8-15 × 2·5-5 µm; M. micromelaena Kantvilas & Coppins, similar to the widespread M. melaena but with markedly smaller, 0-1-septate ascospores, 8-12·5 × 2·5-4 µm; M. oreina Kantvilas & Coppins, characterized by a thallus of globose areoles containing gyrophoric acid, black, subglobose apothecia, and 1-septate ascospores, 11-16·5 × 4·5-6·5 µm; M. pallida Coppins & Kantvilas, similar to M. ceracea but distinguished by the presence of porphyrilic acid and relatively small, 3-septate ascos-pores, 9·5-15 × 2·5-4 µm; M. prasinastra Coppins & Kantvilas (also known from New Zealand), a member of the M. prasina group with a finely granular-sorediose thallus containing gyrophoric acid, unpigmented apothecia and (0-)1-septate ascospores, 7-11·5 × 1·8-3·5 µm; M. rubiginosa Coppins & Kantvilas (also known from Chile), likewise allied to M. prasina but with apothecia containing Rubella-orange pigment and ascospores 0-1-septate, 9·5-17 × 3·5-5·5 µm; M. sandyana Kantvilas, related to M. ternaria (Nyl.) Vĕ zda but differing by smaller ascospores, 7-13·5 × 3·5-6 µm; M. saxicola Coppins & Kantvilas, characterized by a relatively thick, grey-brown, areolate thallus, convex, black apothecia and 0(-1)-septate ascospores, 7-18 × 4·5-7 µm; and M. tubaeformis Coppins & Kantvilas, related to M. flagellispora and with filiform ascospores, 45-100 × 1-2 µm, but differing by containing 2 ′-O-methylperlatolic acid and having funnel-shaped pycnidia. Ten species of Micarea are reported for Tasmania for the first time: M. almbornii Coppins, M. argopsinosa P. M. McCarthy & Elix, M. byssa-cea (Th. Fr.) Czarnota et al., M. contexta Hedl., M. farinosa Coppins & Aptroot, M. humilis P. M. McCarthy & Elix, M. incrassata Hedl., M. myriocarpa V. Wirth & Vě zda ex Coppins, M. nowakii Czarnota & Coppins and M. pseudocoppinsii Brand et al. Also recorded for the first time for Victoria are M. alabastrites (Nyl.) Coppins and M. cinerea (Schaer.) Hedl. A key to Micarea-like lichens in Tasmania, which includes Micarea itself as well as Brianaria, Psilolechia and Leimonis, is presented. Leimonis erratica (Körb.) R. C. Harris & Lendemer and Brianaria tuberculata (Sommerf.) S. Ekman & M. Svensson are recorded for Tasmania for the first time.
... Recent molecular phylogenies show that Micarea is paraphyletic (Andersen & Ekman 2005;Sérusiaux et al. 2010), even after the introduction of the new genera Brianaria S. Ekman & M. Svensson for the M. sylvicola group (Ekman & Svensson 2014) and Leimonis Harris & Lendemer for the M. erratica group (Harris 2009). The M. prasina group, which includes the type species M. prasina Fr., forms a monophyletic core group in the genus. ...
Article
The genus Micarea was studied for the first time in the Taita Hills, Kenya. Based on new collections and existing data, we reconstructed a phylogeny using ITS, mtSSU and Mcm 7 regions, and generated a total of 27 new sequences. Data were analyzed using maximum likelihood and maximum parsimony methods. Based mainly on new collections, we discovered four undescribed well-supported lineages, characterized by molecular and phenotypic features. These lineages are described here as Micarea pumila , M. stellaris , M. taitensis and M. versicolor . Micarea pumila is characterized by a minutely granular thallus, small cream-white or pale brownish apothecia, small ascospores and the production of prasinic acid. Micarea stellaris has a warted-areolate thallus, cream-white apothecia usually darker at the centre, a hymenium of light grey or brownish pigment that dissolves in K, and intense crystalline granules that appear as a belt-like continuum across the lower hymenium when studied in polarized light. Micarea taitensis is characterized by a warted-areolate thallus and cream-white or yellowish apothecia that sometimes produce the Sedifolia-grey pigment. Micarea versicolor is characterized by a warted-areolate, sometimes partly granular thallus and apothecia varying from cream-white to light grey to blackish in colour. This considerable variation in the coloration of its apothecia is caused by an occasional mixture of the Sedifolia-grey pigment in the epihymenium and another purplish brown pigment in the hymenium. Micarea stellaris , M. taitensis and M. versicolor produce methoxymicareic acid. The main distinguishing characters are presented in a species synopsis. Three of the new species are nested in the M. prasina group, and the fourth one ( M. taitensis ) resolves as a basal taxon to the M. prasina group. The new species inhabit montane cloud forests, which have fragmented dramatically throughout the Eastern Arc Mountains in recent decades.
... One additional species, M. adnata Coppins, was included in Kashiwadani et al. (2011) but is not included in the checklist (Ohmura & Kashiwadani 2018). Eighty-six species are currently known from Europe (including the four species referred to Brianaria by Ekman & Svensson (2014)). Of these, 63 occur in the British Isles (Coppins (2009), with four species added later). ...
Article
An examination of collections from Japan has increased the number of Brianaria and Micarea species known from that country from eight to 19, including one new species, M. rubioides Coppins (also from Malaysia and the Philippines). Eleven species are reported as new to Japan ( M. botryoides (Nyl.) Coppins, M. denigrata (Fr.) Hedl., M. erratica (Körb.) Hertel et al. , M. hedlundii Coppins, M. lithinella (Nyl.) Hedl., M. micrococca (Körb.) Gams ex Coppins and M. misella (Nyl.) Hedl.) or new to Asia: M. byssacea (Th. Fr.) Czarnota et al. , M. deminuta Coppins and M. xanthonica Coppins & Tønsberg (new to Asia; Japan); M. nitschkeana (J. Lahm ex Rabenh.) Harm. (new to Asia; South Korea). The presence of Micarea prasina s. str. from Japan needs to be confirmed; no collection was found in this study. Additional collections from South Korea and Sri Lanka are also reported, including the new species M. ceylanica Coppins from Sri Lanka. The identity of M. synotheoides (Nyl.) Coppins, originally described from Japan, has been resolved, resulting in the renaming of Western European material, previously under that name, as M. longispora Coppins. Micarea coreana Lőkös et al . is reported here as a synonym of M. erratica . The type of Lecidea inopinula Nyl. requires the new combination Micarea inopinula (Nyl.) Coppins & T. Sprib. to replace Micarea prasinella (Jatta) I. M. Lamb.
... haematites, C. xerica, and C. variabilis groups) or even genera (Pyrenodesmia and Kuettlingeria). In the context of the current taxonomy of the lichenized fungi (Crespo et al., 2010;Nordin et al., 2010;Spribille et al., 2011;Ekman & Svensson, 2014;Buaruang et al., 2015), and particularly of the family Teloschistaceae (Fedorenko et al., 2012;Arup et al., 2013;Søchting et al., 2014;Kondratyuk et al., 2017b, etc.), the three groups within Pyrenodesmia sensu lato should be treated at the genus rank, as they show a considerable phylogenetical and morphological differentiation. ...
Article
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Most lichens of the family Teloschistaceae (Ascomycota) produce yellow‐orange‐red anthraquinone pigments. However, the genus Pyrenodesmia encompasses species in which anthraquinones are absent and replaced by a grey pigment Sedifolia‐grey. It was shown recently that these species are related to taxa with both anthraquinones and Sedifolia‐grey (C. xerica group, C. haematites group and C. cretensis) and to species with a brown pigment instead of both anthraquinones and Sedifolia‐grey (C. demissa, C. obscurella and C. reptans). Nevertheless, relationships between mentioned anthraquinone‐containing and anthraquinone‐lacking species remained unclear. Eight DNA loci from 41 species were used here trying to resolve these uncertainties. We concluded that C. demissa, C. obscurella and C. reptans are rather distant from the core of Pyrenodesmia and we place them outside of Pyrenodesmia sensu lato. Within Pyrenodesmia sensu lato three lineages were revealed and recognized on generic level: the genus Pyrenodesmia sensu stricto (21 species), the genus Kuettlingeria (14 species) which is resurrected here, and the genus Sanguineodiscus (4 species) which is newly described here. The genus Pyrenodesmia includes taxa which never contain anthraquinones, but Sedifolia‐grey. It matches with the former Caloplaca variabilis group. Taxa of the genera Kuettlingeria and Sanguineodiscus have anthraquinones in their apothecia and Sedifolia‐grey in their thalli. The genus Kuettlingeria includes the former C. xerica group plus C. cretensis and C. diphyodes. The genus Sanguineodiscus includes the former C. haematites group and C. bicolor. The identity of Kuettlingeria (Caloplaca) diphyodes was clarified and the name Pyrenodesmia helygeoides was resurrected. 28 new combinations were proposed. This article is protected by copyright. All rights reserved.
... Notes: The genus Micarea itself is highly heterogeneous, with several lineages falling outside the genus in its proper sense and even outside the Pilocarpaceae; the monophyletic core group also possibly represents more than one genus (Czarnota and Guzow-Krzemińska 2010;Schmull et al. 2011;Ekman and Svensson 2014). Here, we introduce a new species with squamulose thallus that forms an isolated lineage within the monophyletic Micarea core clade (Fig. 89). ...
Article
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This article is the tenth series of the Fungal Diversity Notes, where 114 taxa distributed in three phyla, ten classes, 30 orders and 53 families are described and illustrated. Taxa described in the present study include one new family (viz. Pseudoberkleasmiaceae in Dothideomycetes), five new genera (Caatingomyces, Cryptoschizotrema, Neoacladium, Paramassaria and Trochilispora) and 71 new species, (viz. Acrogenospora thailandica, Amniculicola aquatica, A. guttulata, Angustimassarina sylvatica, Blackwellomyces lateris, Boubovia gelatinosa, Buellia viridula, Caatingomyces brasiliensis, Calophoma humuli, Camarosporidiella mori, Canalisporium dehongense, Cantharellus brunneopallidus, C. griseotinctus, Castanediella meliponae, Coprinopsis psammophila, Cordyceps succavus, Cortinarius minusculus, C. subscotoides, Diaporthe italiana, D. rumicicola, Diatrypella delonicis, Dictyocheirospora aquadulcis, D. taiwanense, Digitodesmium chiangmaiense, Distoseptispora dehongensis, D. palmarum, Dothiorella styphnolobii, Ellisembia aurea, Falciformispora aquatic, Fomitiporia carpinea, F. lagerstroemiae, Grammothele aurantiaca, G. micropora, Hermatomyces bauhiniae, Jahnula queenslandica, Kamalomyces mangrovei, Lecidella yunnanensis, Micarea squamulosa, Muriphaeosphaeria angustifoliae, Neoacladium indicum, Neodidymelliopsis sambuci, Neosetophoma miscanthi, N. salicis, Nodulosphaeria aquilegiae, N. thalictri, Paramassaria samaneae, Penicillium circulare, P. geumsanense, P. mali-pumilae, P. psychrotrophicum, P. wandoense, Phaeoisaria siamensis, Phaeopoacea asparagicola, Phaeosphaeria penniseti, Plectocarpon galapagoense, Porina sorediata, Pseudoberkleasmium chiangmaiense, Pyrenochaetopsis sinensis, Rhizophydium koreanum, Russula prasina, Sporoschisma chiangraiense, Stigmatomyces chamaemyiae, S. cocksii, S. papei, S. tschirnhausii, S. vikhrevii, Thysanorea uniseptata, Torula breviconidiophora, T. polyseptata, Trochilispora schefflerae and Vaginatispora palmae). Further, twelve new combinations (viz. Cryptoschizotrema cryptotrema, Prolixandromyces australi, P. elongatus, P. falcatus, P. longispinae, P. microveliae, P. neoalardi, P. polhemorum, P. protuberans, P. pseudoveliae, P. tenuistipitis and P. umbonatus), an epitype is chosen for Cantharellus goossensiae, a reference specimen for Acrogenospora sphaerocephala and new synonym Prolixandromyces are designated. Twenty-four new records on new hosts and new geographical distributions are also reported (i.e. Acrostalagmus annulatus, Cantharellus goossensiae, Coprinopsis villosa, Dothiorella plurivora, Dothiorella rhamni, Dothiorella symphoricarposicola, Dictyocheirospora rotunda, Fasciatispora arengae, Grammothele brasiliensis, Lasiodiplodia iraniensis, Lembosia xyliae, Morenoina palmicola, Murispora cicognanii, Neodidymelliopsis farokhinejadii, Neolinocarpon rachidis, Nothophoma quercina, Peroneutypa scoparia, Pestalotiopsis aggestorum, Pilidium concavum, Plagiostoma salicellum, Protofenestella ulmi, Sarocladium kiliense, Tetraploa nagasakiensis and Vaginatispora armatispora).
... Largely based on molecular methods, the understanding of species boundaries and species diversity has improved (Czarnota and Guzow-Krzemińska 2010;Guzow-Krzemińska et al. 2016;van den Boom et al. 2017). Recent molecular phylogenies show, for instance, that the morphological concept of Micarea is paraphyletic (Andersen and Ekman 2005;Sérusiaux et al. 2010), even after the introduction of a new genus Brianaria S. Ekman & M. Svensson for the M. sylvicola group (Ekman and Svensson 2014). However, as explained below, before a reliable genus-level phylogenetic reconstruction can be proposed, several taxonomic issues concerning the type species, M. prasina Fr., should be addressed (e.g., Czarnota and Guzow-Krzemińska 2010;Launis et al. 2019). ...
Article
Micarea is a lichenized genus in the family Pilocarpaceae (Ascomycota). We studied the phylogeny and reassessed the current taxonomy of the M. prasina group. We focused especially on the taxonomic questions concerning the type species M. prasina and, furthermore, challenges concerning type specimens that are too old for successful DNA barcoding and molecular studies. The phylogeny was reconstructed using nuc rDNA internal transcribed spacer region (ITS1-5.8S-ITS2 = ITS), mitochrondrial rDNA small subunit (mtSSU), and replication licensing factor MCM7 gene from 31 species. Fifty-six new sequences were generated. The data were analyzed using maximum parsimony and maximum likelihood methods. The results revealed four undescribed, well-supported lineages. Three lineages represent new species described here as M. fallax, M. flavoleprosa, and M. pusilla. In addition, our results support the recognition of M. melanobola as a distinct species. Micarea fallax is characterized by a vivid to olive green thallus composed of aggregated granules and whitish or brownish apothecia sometimes with grayish tinge (Sedifolia-gray pigment).Micarea flavoleprosa has a thick, wide-spreading yellowish green, whitish green to olive green sorediate thallus and lacks the Sedifolia-gray pigmentation. The species is mostly anamorphic, developing apothecia rarely. Micarea melanobola is characterized by a pale to dark vivid green granular thallus and darkly pigmented apothecia (Sedifolia-gray). Micarea pusilla is characterized by a whitish green to olive green thinly granular or membranous thallus, numerous and very small whitish apothecia lacking the Sedifolia-gray pigment, and by the production of methoxymicareic acid. Micarea fallax, M. flavoleprosa, and M. melanobola produce micareic acid. The reliability of crystalline granules as a character for species delimitation was investigated and was highly informative for linking the old type specimen of M. prasina to fresh material.
Article
Printzen, C., Brackel, W. v., Bltmann, H., Cezanne, R., Dolnik, C., Dornes, P., Eckstein, J., Eichler, M., John, V., Killmann, D., Nimis, P. L., Otte, V., Schiefelbein, U., Schultz, M., Stordeur, R., Teuber, D. & Ths, H. 2022. Die Flechten, flechtenbewohnenden und flechtenhnlichen Pilze Deutschlands eine berarbeitete Checkliste. Herzogia 35: 193-393. In der vorliegenden Arbeit werden 2051 Flechten, 520 flechtenbewohnende und 55 flechtenhnliche Pilze, insgesamt 2626 Taxa nebst Synonymen aufgelistet, deren Vorkommen bis 31.12.2021 aus dem Gebiet der Bundesrepublik Deutschland gemeldet wurde. Die Liste basiert auf dem letzten im Jahre 2011 verffentlichten Artenverzeichnis und bercksichtigt 326 Neunachweise von Arten sowie 428 nomenklatorische nderungen, die zwischen 2012 und 2021 in 253 Publikationen verffentlicht wurden. Die Liste umfasst auerdem 114 Taxa, zumeist aus den Verrucariaceae, deren Status weiterhin als problematisch angesehen wird. Printzen, C., Brackel, W. v., Bltmann, H., Cezanne, R., Dolnik, C., Dornes, P., Eckstein, J., Eichler, M., John, V., Killmann, D., Nimis, P. L., Otte, V., Schiefelbein, U., Schultz, M., Stordeur, R., Teuber, D. & Ths, H. 2022. Lichens, lichenicolous and allied fungi of Germany a revised checklist. Herzogia 35: 193-393. The present work lists 2051 lichens, 520 lichenicolous and 55 allied fungi, altogether 2626 taxa and their synonyms, whose occurrence has been reported from the territory of the Federal Republic of Germany by the end of 2021. The list is based on the last species list published in 2011 and comprises 326 new records as well as 428 nomenclatural changes published in 253 publications between 2012 and 2021. The list also includes 114 taxa, mostly from the Verrucariaceae, whose status is still considered problematic.
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Diese erste Rote Liste und Checkliste der Flechten, flechtenbewohnenden und flechtenähnlichen Pilze Bayerns umfasst 2.054 Taxa, davon 1.624 Flechten, 399 flechtenbewohnende Pilze und 31 flechtenähnliche Pilze. Sie wurde aufgrund intensiver Recherchen in der Literatur erstellt, die in einem elektronischen Supplement nachvollziehbar niedergelegt sind. Insgesamt kommen von den drei Artengruppen 1.417 Taxa (1.163 Flechten/236 flechten-bewohnende/18 flechtenähnliche Pilze) in der alpinen Region sowie 1.691 (1.341/323/27) in der kontinentalen Region vor. 909 Taxa (44 %) sind als Rote-Liste-Arten der Kategorien 0, 1, 2, 3 und G ausgestorben oder gefährdet (in der alpinen Region 31 %, in der kontinentalen Region 48 %). Dazu treten als „extrem seltene“ Arten in der Kategorie „R“ 395 Taxa (19 %) hinzu – in der alpinen Region 21 %, in der kontinentalen Region 13 %. In der kontinentalen Region mit der intensiveren Flächennutzung finden sich prozentual mehr gefährdete Arten als in der alpinen Region. Da-gegen finden sich in den Alpen besonders viele extrem seltene Arten. Die bayerischen Flechtenbestände waren in den letzten 200 Jahren (dem Zeitraum, zu dem Angaben zu den Beständen vorliegen), etlichen Belastungen ausgesetzt: Die Industrialisierung führte zu erhöhten Schwefeldioxyd-Konzentrationen in der Luft, die aber Ende des 20. Jahr-hunderts durch verstärkte Anstrengungen in der Luftreinhaltung wieder signifikant verringert werden konnten. Neuordnung und Intensivierung der Landwirtschaft vernichtete vor allem in der Zeit nach 1950 zahlreiche Lebensräume in der offenen Landschaft, während in der Forst-wirtschaft bereits ab Mitte des 19. Jahrhunderts Nadelholzmonokulturen in den Wäldern zu massiven Diversitätsverlusten führten. Seit etwa 40 Jahren bedroht zunehmend eine steigende Eutrophierung (vor allem durch Stickstoffverbindungen aus Industrie, Verkehr und Landwirtschaft) umfassend die Flechten und ihre Lebensräume, vor allem indem sie Höhere Pflan-zen und konkurrenzkräftige Moose fördert. Dringend notwendig zum Schutz der Flechtenflora ist • eine Reduktion der Eutrophierung, • die Wiederherstellung eines Netzes von Kleinstrukturen in der Landschaft, • das weitere Zurückdrängen der Nadelholz-Monokulturen, • die Einrichtung von Altholzinseln in den Wäldern, • die Wiederherstellung von Flechten-Kiefernwäldern sowie • gezielte Artenhilfsmaßnahmen, wie die Übertragung von Flechten in renaturierte Biotope.
Article
A new species Micarea fennica is described based on phenotypic and molecular features. The species is characterized by a pale olive green, bright green or greyish green minutely granular thallus composed of goniocysts, and stalked pycnidia that are dark grey to dark brown with Sedifolia-grey pigment in the wall structures (K+ violet, C+ violet), 0.2–0.5(–1) mm tall and covered by a thin white tomentum. The species produces micareic acid. Micarea fennica is similar to M. hedlundii and M. botryoides, but differs by having a paler granular thallus, stalked dark grey to dark brown pycnidia covered by a thin white tomentum and the production of micareic acid. The new species occupies soft lignum of late decay stages and is probably rare.
Article
The Lecanoromycetes includes most of the lichen-forming fungal species (>13 500) and is therefore one of the most diverse class of all Fungi in terms of phenotypic complexity. We report phylogenetic relationships within the Lecanoromycetes resulting from Bayesian and maximum likelihood analyses with complementary posterior probabilities and bootstrap support values based on three combined multilocus datasets using a supermatrix approach. Nine of 10 orders and 43 of 64 families currently recognized in Eriksson’s classification of the Lecanoromycetes (Outline of Ascomycota—2006 Myconet 12:1–82) were represented in this sampling. Our analyses strongly support the Acarosporomycetidae and Ostropomycetidae as monophyletic, whereas the delimitation of the largest subclass, the Lecanoromycetidae, remains uncertain. Independent of future delimitation of the Lecanoromycetidae, the Rhizocarpaceae and Umbilicariaceae should be elevated to the ordinal level. This study shows that recent classifications include several nonmonophyletic taxa at different ranks that need to be recircumscribed. Our phylogenies confirm that ascus morphology cannot be applied consistently to shape the classification of lichen-forming fungi. The increasing amount of missing data associated with the progressive addition of taxa resulted in some cases in the expected loss of support, but we also observed an improvement in statistical support for many internodes. We conclude that a phylogenetic synthesis for a chosen taxonomic group should include a comprehensive assessment of phylogenetic confidence based on multiple estimates using different methods and on a progressive taxon sampling with an increasing number of taxa, even if it involves an increasing amount of missing data.
Article
Electron micrographs of four lichen species of the Lecanoraceae were examined. The contact between the symbionts ranges from active penetration of the algal cell wall by fungal haustoria in a supposedly primitive growth‐form, through intermediate stages, to a kind of hyphal entrapment in an advanced form. Penetration beyond the plasma membrane of intact algal cells has not been observed. Whereas penetrations may occur in all the algal cells composing the algal layer of the primitive thallus, in the more advanced growth‐forms, only those cells which have reached a certain stage of their life cycle are attacked. 1970 The New Phytologist