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The 2018 classification and checklist of lichenicolous fungi, with 2000 non-
lichenized, obligately lichenicolous taxa
Author(s): Paul Diederich, James D. Lawrey and Damien Ertz
Source: The Bryologist, 121(3):340-425.
Published By: The American Bryological and Lichenological Society, Inc.
URL: http://www.bioone.org/doi/full/10.1639/0007-2745-121.3.340
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The 2018 classification and checklist of lichenicolous fungi,
with 2000 non-lichenized, obligately lichenicolous taxa
Paul Diederich
1,5
, James D. Lawrey
2
and Damien Ertz
3,4
1
Mus´
ee national d’histoire naturelle, 25 rue Munster, L–2160 Luxembourg, Luxembourg;
2
Department of Biology,
George Mason University, Fairfax, VA 22030-4444, U.S.A.;
3
Botanic Garden Meise, Department of Research,
Nieuwelaan 38, B–1860 Meise, Belgium;
4
F´
ed´
eration Wallonie-Bruxelles, Direction G´
en´
erale de l’Enseignement non
obligatoire et de la Recherche scientifique, rue A. Lavall´
ee 1, B–1080 Bruxelles, Belgium
ABSTRACT.Lichenicolous fungi represent a highly specialized and successful group of organisms that live
exclusively on lichens, most commonly as host-specific parasites, but also as broad-spectrum pathogens,
saprotrophs or commensals. We present here the most recent update to the classification of lichenicolous
fungi in the Ascomycota and Basidiomycota to genus level, arranged phylogenetically according to
published classifications. For each genus, all known lichenicolous taxa (obligately lichenicolous taxa,
lichenicolous lichens, and facultatively lichenicolous taxa) are listed, along with information about types,
synonyms, pertinent literature and whether or not molecular data are available for any of the listed
species. The number of accepted lichenicolous fungi is now 2319, with 2000 obligately lichenicolous
species, subspecies or varieties, 257 lichenicolous lichens and 62 facultatively lichenicolous taxa. These
species are found in 10 different classes of Fungi (Ascomycota and Basidiomycota), 55 orders, 115 families
and 397 genera. The 2319 total taxa is an increase from the 1559 total species reported in the last
published catalogue in 2003, and a larger number than the approximately 1800 reported in the most
recent online checklist (www.lichenicolous.net) posted in January 2018. Of the total number of taxa, 2219
(96%) are ascomycetes and 100 (4%) are basidiomycetes. Of the 397 genera containing lichenicolous
species, c. 50% (198) are entirely lichenicolous. In addition, six families (Abrothallaceae, Adelococcaceae,
Cyphobasidiaceae, Obryzaceae, Polycoccaceae, Sarcopyreniaceae) and two orders (Abrothallales,
Cyphobasidiales) are entirely lichenicolous. Sequence information is available for lichenicolous species
in 128 (32%) of the 397 genera containing lichenicolous species, and in 56 (28%) of the 198 entirely
lichenicolous genera. Many species are known from only one host lichen, but it is likely that broader host
ecologies will be discovered as new sequence information is obtained from ongoing microbiome studies.
Phaeopyxis Rambold & Triebel is considered as a new synonym of Bachmanniomyces D.Hawksw., resulting
in five new combinations B. australis (Rambold & Triebel) Diederich & Pino-Bodas (”P. australis), B.
carniolicus (Arnold) Diederich & Pino-Bodas (”Biatora carniolica), B. muscigenae (Alstrup &
E.S.Hansen) Diederich & Pino-Bodas (”P. muscigenae), B. punctum (A.Massal.) Diederich & Pino-
Bodas (”Nesolechia punctum) and B. varius (Coppins, Rambold & Triebel) Diederich & Pino-Bodas (”
P. varia). As a consequence of a phylogenetic analysis including new sequences, Dactylospora K¨
orb. is
regarded as a new synonym of Sclerococcum Fr. : Fr., resulting in one new name (S. acarosporicola Ertz &
Diederich) and 46 new combinations. Sclerococcaceae R´
eblova
´, Unter. & W.Gams is considered as a new
synonym of Dactylosporaceae Bellem. & Hafellner. The new Sclerococcum ophthalmizae Coppins is
described. Sclerophyton occidentale Herre is lectotypified on the lichenicolous fungus present in the type
specimen and becomes a younger synonym of Sclerococcum parasiticum. A replacement name is Arthonia
polydactylonis Diederich & Ertz (”A. ceracea). Further new combinations are Abrothallus lobariae
(Diederich & Etayo) Diederich & Ertz (”Phoma lobariae), A. psoromatis (Zhurb. & U. Braun) Diederich
& Zhurb. (”P. psoromatis), Asteroglobulus pyramidalis (Etayo) Diederich (”Cornutispora pyramidalis),
Didymocyrtis grumantiana (Zhurb. & Diederich) Zhurb. & Diederich (”Phoma grumantiana),
5
Corresponding author’s e-mail: paul.diederich@education.lu
DOI: 10.1639/0007-2745-121.3.340
The Bryologist 121(3), pp. 340–425 Published online: October 18 2018 0007-2745/18/$8.75/0
Copyright Ó2018 by The American Bryological and Lichenological Society, Inc.
Epithamnolia atrolazulina (Etayo) Diederich (”Hainesia atrolazulina), Gyalolechia epiplacynthium
(Etayo) Diederich (”Fulgensia epiplacynthium), Nesolechia doerfeltii (Alstrup & P.Scholz) Diederich (”
Phacopsis doerfeltii), N. falcispora (Triebel & Rambold) Diederich (”P. falcispora), N. oxyspora var. fusca
(Triebel & Rambold) Diederich (”P. oxyspora var. fusca), Preussia peltigerae (Brackel) Diederich (”
Sporormiella peltigerae), Scutula curvispora (D.Hawksw. & Mia
˛dl.) Diederich (”Libertiella curvispora), S.
didymospora (D.Hawksw. & Mia
˛dl.) Diederich (”L. didymospora), Stigmidium haesitans (Nyl.) Diederich
(”Verrucaria haesitans), and S. parvum (Henssen) Diederich (”Pharcidia parvum).
KEYWORDS.Endolichenic fungi, lichenicolous lichens, microbiome, mycoparasites, phylogeny.
^^^
Lichenicolous fungi represent a highly specialized
and successful group of organisms that live exclu-
sively on lichens, most commonly as host-specific
parasites, but also as broad-spectrum pathogens,
saprotrophs and commensals. Although they have
been collected and described for centuries, little is
known about the basic biology of most species,
including modes of cellular contact, nutrient ex-
change, antibiosis, or virulence. Most are assumed to
be restricted to a single host genus, but the degree of
host-specificity is usually not known because of
uncertainties about species boundaries and distribu-
tions. Very few lichenicolous fungi have been
brought into culture, and little is known about their
cultural requirements.
Lichenicolous fungi are assumed to represent an
important source of new species in many groups of
fungi (Hawksworth & Rossman 1997), but their
overall diversity remains uncharted territory. This
classification provides a snapshot of the phylogenetic
diversity of lichenicolous fungi based on information
updated continuously in the online website www.
lichenicolous.net. It is being published here in the
classification series in The Bryologist to provide a
permanent baseline survey to which future classifi-
cations can be compared as new information
accumulates.
LICHENICOLOUS FUNGI
Lichen parasites.All lichenicolous fungi are
technically lichen parasites (they acquire fixed
carbon from living lichen thalli), but many have
no noticeable effects on the lichen host and some
have the same nutrition as lichens. Most of the
currently recognized lichenicolous species are weak
parasites of either the lichen mycobiont (mycopar-
asites) or photobiont (phycoparasites) or sometimes
both, recognizable by the presence of galls, lesions
and discolorations on the lichen thallus. Some
species are broad-spectrum pathogens that cause
extensive damage capable of killing the host, but this
is very rare among lichenicolous species (Hawks-
worth 1982c). Contacts between the parasite and the
lichen host have been observed microscopically
(Grube & Hafellner 1990; Grube & de los R´
ıos
2001; de los R´
ıos & Grube 2000; Diederich 1996),
but the actual nature of most associations is not
known with certainty.
Lichenicolous lichens.In addition to lichen
myco- and phycoparasites, some described lichen-
icolous fungi are known to be derived from lichens
and have the same nutrition (Rambold & Triebel
1992). They are therefore lichen fungi in search of a
photobiont, facultative lichens usually referred to in
the literature, and in this classification, as ‘‘lichen-
icolous lichens.’’ They commonly begin growth as a
free-living lichen mycobiont that becomes lichenized
by associating with the photobiont of an unrelated
lichenized thallus. Over time, the parasitic myco-
biont becomes independent, either continuing to use
the original host’s photobiont or obtaining another
photobiont later. Other lichenicolous lichens have
their own photobiont but are typically collected on
the surfaces of other ‘‘host’’ lichens. In this case, they
are not biologically parasites (they are nutritionally
lichens), but routinely associate with other lichens,
growing over the surface of the host and clearly
affecting its health and fitness. In this classification,
we include all of these fungi in a separate category
(lichenicolous lichens) and mention in the species
listing following each genus what hosts they are
known to associate with.
Host specificity.In this classification, obligate
parasites and pathogens will be the predominant
category to which species are assigned, and the main
known hosts will be listed. A few species are
Diederich et al.: 2018 classification and checklist of lichenicolous fungi 341
commonly collected from lichens but are also known
to be capable of living on non-lichen substrates of
various sorts. These will be referred to in this
classification as ‘‘facultatively lichenicolous’’ species.
The host ecology of most species is assumed to be
remarkably narrow for parasitic organisms. Most
described species, perhaps as many as 95% (Lawrey
& Diederich 2003), associate with a single lichen
species or genus and are never found elsewhere.
Similarly, some lichens harbor far more described
lichenicolous fungi than others (Table 2 in Lawrey &
Diederich 2003), indicating that suitability as hosts
varies among lichen species probably for a wide
variety of reasons (discussed by Hawksworth 1982d;
Lawrey & Diederich 2003).
The close association of lichen parasites and
lichen hosts would argue for a potential coevolu-
tionary relationship among the associates. Until
recently, lichen-parasite interactions have not been
studied with regard to coevolution despite the fact
that high levels of host-specificity are assumed to
indicate cospeciation resulting from coevolution
(Page 2003). In a very interesting first look at this,
Millanes et al. (2014b) found that parasitism can be
an important driver of speciation in lichen parasites
and may help to explain the high numbers of both
species and levels of host specificity in certain groups
of lichenicolous fungi. The parasite studied was
Biatoropsis usnearum, a basidiomycete known to
infect a variety of Usnea lichens. The morphological
variability in B. usnearum had been noted before and
suggested to the authors that more than one parasite
species might actually be involved in these interac-
tions. The authors found that this was indeed the
case—Biatoropsis usnearum is made up of seven
lineages, some of which are more host specific than
others. This diversity apparently arose by host
switching, especially in the most host-specific
lineages. This interesting first look at possible
lichen-parasite coevolution suggests that host spec-
ificity and diversity in lichen parasites may therefore
be a by-product of a number of ecological interac-
tions that have rarely been studied in detail in these
organisms.
Although lichenicolous fungi have been collected
for centuries, our understanding of their actual
distributions in nature, including the levels of host
specificity and host ranges, will likely change over
time as more information becomes available. This
new information will include not only new collec-
tions, but also data from lichen microbiome studies
(discussed in the next section) that indicate the
presence of these fungi in asymptomatic lichens. For
example, Fernandez-Mendoza et al. (2017) discov-
ered using 454 pyrosequencing approaches that
sequences of lichenicolous fungi are found not only
in lichens with symptoms, but also in the same
lichen species without symptoms, and (most sur-
prisingly) in other lichen species never known to
harbor the parasites. It may therefore eventually be
discovered that the apparent high levels of host
specificity in these fungi are not as high as they
appear now. However, far more high-throughput
sequencing of lichens is needed to establish this with
certainty.
Effects of lichenicolous fungi on lichen health
and fitness.Very little is known about the effects
lichen parasites have on lichen hosts, but it is
assumed that in many cases the association is more
or less commensalistic with little or no effects on the
lichens. In other cases, there is obvious discoloration
or thallus damage, sometimes extensive, but the
extent to which this affects lichen growth and
reproduction is still largely unknown. The broad-
spectrum pathogen Athelia arachnoidea is known to
cause damage in European lichen communities,
especially those affected by air pollution (e.g.,
Arvidsson 1976, 1978), but it does not appear to
alter lichen community composition (Hawksworth
1982c). Few studies have measured the direct effect
of parasites on lichen fitness, but a recent study
(Merinero & Gauslaa 2017) showed experimentally
that the gall-forming parasites Plectocarpon lichenum
and P. scrobiculatae reduced the growth and fitness
of the Lobaria lichens on which they grow. Effects on
lichens may also involve degradation of lichen
compounds, something lichenicolous fungi are
known to do (Lawrey 2000). This can result in
exposure of lichens to damaging UV radiation
(Edwards et al. 2017) or to increased competition
and predation (Asplund et al. 2016, 2018), ultimately
leading to reduced growth and fitness.
Lichenicolous fungi as members of lichen
mycobiomes.Lichens have long been known to
harbor a diverse but largely invisible mycobiota in
addition to the lichen-forming mycobiont (see e.g.,
Girlanda et al. 1997; Petrini et al. 1990; Suryanar-
ayanan et al. 2005; Li et al. 2007; Harutyunyan et al.
2008; Fleischhacker et al. 2015; Muggia et al. 2016;
Muggia & Grube 2010; Spribille et al. 2016;
Fernandez-Mendoza et al. 2017; Wang et al. 2016;
342 The Bryologist 121(3): 2018
Zhang et al. 2015). Until recently, these fungi were
assumed to be mainly opportunistic and facultative
plant- and soil-inhabiting saprotrophs. Some are
undoubtedly described lichenicolous forms that have
not developed recognizable structures, but there are
indications that additional groups of obligately
lichen-associated fungi may occur in lichens. Detec-
tion techniques were originally culture-based, but
recent uses of molecular methods have expanded our
knowledge of the diversity, composition and spec-
ificity of these fungi, and it is now apparent that
many are not generalized randomly-distributed
contaminants but members of recognizable, more-
or-less obligate, lichen-associated communities.
Hofstetter et al. (2007) were first to suggest that
nonlichen fungal sequences commonly amplified
from lichens may represent a previously unrecog-
nized source of lichen-specific fungal diversity. These
were later referred to as ‘‘endolichenic’’ forms by
Arnold et al. (2009), who viewed them as phyloge-
netically distinct from both lichen-forming myco-
bionts and lichenicolous fungi and an important
evolutionary link to plant-associated endophytes
(e.g., Chagnon et al. 2016; Fleischhacker et al.
2015; Muggia et al. 2016; U’Ren et al. 2010, 2012,
2014).
Arnold et al. (2009) originally considered
endolichenic and lichenicolous fungi to be distin-
guishable by: (1) presence of symptoms—lichen-
icolous fungi show outward symptoms of infection
and endolichenics lack any obvious outward symp-
toms; (2) phylogeny—endolichenic fungi were
found to be most common among primarily non-
lichenized lineages of euascomycetes (Sordariomy-
cetes, Dothideomycetes, Leotiomycetes and
Pezizomycetes) but absent among the lichen-dom-
inated clades (Lecanoromycetes, Arthoniomycetes,
Lichinomycetes), while lichenicolous fungi were
thought to be closely related to lichens and found
mainly in lichen-forming clades. As for the first
difference (symptoms), it is now evident that lichens
with and without obvious symptoms of infection
harbor numerous described lichenicolous species in
their mycobiomes (Fernandez-Mendoza et al. 2017).
Since very few described lichenicolous species have
been sequenced until very recently, the ability to
identify endolichenics as lichenicolous species is
limited at present. However, as more sequences of
described species are published, it will be possible to
identify those previously assigned to endolichenic
fungi that actually belong to described lichenicolous
species. As for the second difference (phylogeny),
Lawrey & Diederich (2003) showed earlier, and
many studies done since (incorporated into this
classification), continue to support the idea that
lichenicolous fungi are well-represented in both
lichen-forming and non-lichenized clades of fungi.
Since phylogenetic placement helps to elucidate
origins of these fungi, it is apparent that lichen-
icolous fungi have multiple origins from lichen-
forming and non-lichenized ancestors while endo-
lichenic fungi are most likely descended from non-
lichenized ancestors (Arnold et al. 2009). Given what
we know about these lichen-associated groups now,
we would argue that they represent components of
the same communities and the distinction between
them is likely to become less clear-cut as more lichen
mycobiomes are studied and the inhabitants iden-
tified.
DIVERSITY AND CLASSIFICATION HISTORY
Lichenicolous fungi have been studied and
described since the mid-18
th
century, long before
the symbiotic nature of lichens was recognized
(Acharius 1795, 1810; Dillenius 1742). Illustrated
andremarkablythoroughtreatmentsappearedin
the 19
th
and early 20
th
centuries (Keissler 1930;
Lindsay 1869; Olivier 1905–1907; Vouaux 1912–
1914; Zopf 1896), drawing the attention of both
lichenologists and other mycologists to these fungi.
Modern compilations were produced by Clauzade
& Roux (1976 – with 457 species) and Clauzade et
al. (1989 – with 686 species) (Fig. 1). Hawksworth
(1983) published keys to 218 lichenicolous species
known from the British Isles and Ihlen & Wedin
(2008) keys to 298 species known from Sweden. The
most recent published compilation (Lawrey &
Diederich 2003) listed 1559 total species worldwide
and the most recent online worldwide checklist
includes around 1800 described species, including
in some cases infraspecific taxa (Lawrey & Die-
derich 2003; http://www.lichenicolous.net). How-
ever, taking into account Hawksworth’s (1991b,
2001) global estimates of fungal diversity and
average levels of host specificity, and the total
number of 19,409 lichen species in the most current
classification (L¨
ucking et al. 2016, 2017), the actual
number of lichenicolous fungal species should be
much higher. Estimates are that 3000–5000 species
will eventually be described. In the present cata-
Diederich et al.: 2018 classification and checklist of lichenicolous fungi 343
logue, we list a total of 2319 lichenicolous species or
infraspecific taxa.
MATERIALS AND METHODS
We produced a hierarchical listing of genera
based on the most recent data available in www.
lichenicolous.net [posted January 3, 2018]. For all
genera, the latest phylogenetic and taxonomic
papers have been consulted, and the resulting
classification compared with other recent classifi-
cations of fungi (e.g., Jaklitsch et al. 2016, L ¨
ucking
et al. 2016, Lumbsch & Huhndorf 2007; Wijaya-
wardene et al. 2017, Zhao et al. 2017) to detect
errors or omissions.
As is the case for the online checklist, we list for
each genus all lichenicolous species or infraspecific
taxa (obligately lichenicolous taxa, lichenicolous
lichens and facultatively lichenicolous taxa), along
with information about types, synonyms, numbers
of each category of lichenicolous species or infra-
specific taxa, pertinent literature and whether or not
molecular data are available.
Molecular techniques and phylogenetic analy-
ses.For the phylogenetic tree of the Sclerococcales
(Fig. 4), a total of five nuLSU rDNA and four
mtSSU sequences were newly amplified and
sequenced for three specimens of Dactylospora
lobariella and two specimens of D. parasitica,
using the techniques described in Ertz et al.
(2018a,b). Members of Sclerococcales for which the
nuLSU and/or the mtSSU sequences were avail-
able were selected from Diederich et al. (2013),
Muggia et al. (2016, 2017), R´
eblova
´et al. (2017)
and Pino-Bodas et al. (2017), and their sequences
retrieved from GenBank (Table 6). Penicillium
javanicum, Pyrgillus javanicus and Sphinctrina
turbinata were chosen as outgroup. The sequences
were aligned using MAFFT v6.814b (Katoh et al.
2002) on the CIPRES Web Portal (Miller et al.
2010) and improved manually using Mesquite
3.04 (Maddison & Maddison 2015). Terminal
ends of sequences and ambiguously aligned
regions were delimited manually and excluded
from the datasets.
Best-fit evolutionary models were estimated
using Akaike Information Criterion (AIC; Akaike
1973) as implemented in jModelTest2 2.1.6 (Darri-
ba et al. 2012). The TIM1þIþGandtheGTRþG
models were selected respectively for the nuLSU
and the mtSSU datasets. The combined nuL-
SUþmtSSU dataset consisted of 29 specimens and
1610 unambiguously aligned sites. A bayesian
analysis was carried out on the nuLSUþmtSSU
dataset using the Metropolis-coupled Markov chain
Monte Carlo (MCMCMC) method in MrBayes v.
3.2.6 (Ronquist & Huelsenbeck 2003) on the
CIPRES Web Portal (Miller et al. 2010). Two
parallel MCMCMC runs were performed, each
using four independent chains and 40 million
generations, sampling trees every 1000
th
generation.
Posterior probabilities (PP) were determined by
calculating a majority-rule consensus tree generated
from 60002 post-burnin trees of the 80002 trees
sampled by the two MCMCMC runs using the sumt
option of MrBayes. PP 0.95 were considered to
be significant. The phylogenetic tree was visualized
using FigTree v. 1.4.2 (Rambaut 2012). The dataset
and tree were deposited in Treebase as submission
23081.
RESULTS AND DISCUSSION
At the time of this publication, the total number
of lichenicolous fungi, including in certain cases
subspecific taxa (Table 1,Supplementary Table S1)
is 2319, with 2000 obligately lichenicolous species,
257 lichenicolous lichens and 62 facultatively lichen-
icolous species. This is a significant increase from the
1559 total species reported in the last published
checklist (Lawrey & Diederich 2003), and a larger
number even than the approximately 1800 reported
in the most recent online checklist posted in January
earlier this year. Of the total number of species, 2219
(96%) are ascomycetes (Fig. 2) and 100 (4%) are
Figure 1. Number of obligate non-lichenized lichenicolous taxa (species,
subspecies or varieties) accepted by Lindsay (1869), Vouaux (1912–14),
Clauzade & Roux (1976), Clauzade et al. (1989), Lawrey & Diederich
(2003) and the present classification (2018).
344 The Bryologist 121(3): 2018
basidiomycetes (Fig. 3), and this ratio did not
change from 2003.
Of the 397 genera in the list, 198 (c. 50%) are
entirely lichenicolous (Table 2); there are also six
families and two orders, some with large numbers
of species, that are entirely lichenicolous (Table 3).
Table 2. Number of genera containing varying levels of lichenicolous
species.
Level of representation
Ratio of
lichenicolous species to
all species in the genus
Number of
genera
All lichenicolous species 100% 198
80% 210
Some lichenicolous species 50% 227
20% 270
At least one lichenicolous
species
.0% 397
Table 1. Current total numbers of lichenicolous fungi (obligate
lichenicolous, lichenicolous lichens and facultative lichenicolous)
represented in various taxonomic ranks.
Taxonomic rank Category Numbers
Species obligate lichenicolous 2000
lichenicolous lichens 257
facultative lichenicolous 62
Total 2319
Genera obligate lichenicolous 341
lichenicolous lichens 63
facultative lichenicolous 34
Total 397
Families obligate lichenicolous 98
lichenicolous lichens 28
facultative lichenicolous 21
Total 115
Orders obligate lichenicolous 45
lichenicolous lichens 16
facultative lichenicolous 17
Total 55
Classes obligate lichenicolous 8
lichenicolous lichens 4
facultative lichenicolous 6
Total 10
Number of species
Basidiomycota obligate lichenicolous 93
lichenicolous lichens 0
facultative lichenicolous 7
Total 100
Ascomycota obligate lichenicolous 1907
lichenicolous lichens 257
facultative lichenicolous 55
Total 2219
Table 3. Families and orders of fungi containing only lichenicolous
species.
Taxonomic category Number of species
Families with only lichenicolous species
Abrothallaceae 57
Polycoccaceae 72
Adelococcaceae 14
Sarcopyreniaceae 11
Obryzaceae 3
Cyphobasidiaceae 2
Orders with only lichenicolous species
Abrothallales 57
Cyphobasidiales 2
Table 4. Genera with the largest number of described lichenicolous fungi
(all three categories, obligate, facultative and lichenicolous fungi). For
genera indicated in bold, all known species are lichenicolous.
Genus Order
Number of
described
species
Arthonia Arthoniales (Asco) 140
Stigmidium Verrucariales (Asco) 94
Opegrapha Arthoniales (Asco) 72
Sclerococcum Sclerococcales (Asco) 64
Polycoccum Trypetheliales (Asco) 60
Tremella Tremellales (Basidio) 60
Lichenochora Phyllachorales (Asco) 44
Endococcus Verrucariales (Asco) 44
Pronectria Hypocreales (Asco) 44
Abrothallus Abrothallales (Asco) 42
Cercidospora Dothideomycetes inc. sed. (Asco) 40
Plectocarpon Arthoniales (Asco) 40
Lichenopeltella Dothideomycetes inc. sed. (Asco) 39
Rhizocarpon Rhizocarpales (Asco) 39
Sphaerellothecium Dothideomycetes inc. sed. (Asco) 35
Zwackhiomyces Collemopsidiales (Asco) 35
Chaenothecopsis Mycocaliciales (Asco) 33
Skyttea Helotiales (Asco) 30
Taeniolella Asterotexiales (Asco) 30
Table 5. Genera of lichenicolous fungi with published sequences. In
some cases the type species of the genus has been sequenced, but the
lichenicolous members of the genus have not been sequenced.
Category Numbers
Number of genera with lichenicolous sequences 128
Number of genera with type sequenced 171
Number of genera with type and lichenicolous sequenced 102
Diederich et al.: 2018 classification and checklist of lichenicolous fungi 345
Figure 2. Morphological diversity of lichenicolous ascomycetes. A. Arthonia diorygma on Graphidaceae (Arthoniales, Arthoniaceae). B. A. graphidicola on
Graphis (id.). C. Plectocarpon lichenum on Lobaria (id., Lecanographaceae). D. Mixtoconidium insidiens (sexual stage) on Ramalina (id., gen. inc. sed.). E.
id. (pycnidial stage). F. Lichenostigma maureri on Usnea (Lichenostigmatales, Phaeococcomycetaceae). G. Pronectria robergei on Peltigera (Hypocreales,
Bionectriaceae). H. Trichonectria rubefaciens on Pleurosticta (id.). I. Lichenobarya usneae on Usnea (id., Hypocreaceae). J. Llimoniella ramalinae on
Ramalina (Helotiales, Corderitidaceae). K. Skyttea pertusariicola on Pertusaria (id.). L. Unguiculariopsis manriquei on Lobaria (id.). M. Hemigrapha
asteriscus on Peltigera (Asterotexiales, gen. inc. sed.). N. Nigromacula uniseptata on Hypotrachyna (gen. inc. sed.). O. Neolamya peltigerae on Peltigera
(id.). All photographs by Paul Diederich.
346 The Bryologist 121(3): 2018
Figure 3. Morphological diversity of lichenicolous basidiomycetes. A. Chionosphaera lichenicola on Fellhanera (Agaricostilbales, Chionosphaeraceae). B.
Cyphobasidium hypogymniicola on Hypogymnia (Cyphobasidiales, Cyphobasidiaceae). C. C. usneicola on Usnea (id.). D. Heterocephalacria physciacearum
on Physcia (Filobasidiales, Filobasidiaceae). E. Biatoropsis minuta on Usnea (Tremellales, Tremellaceae). F. Tremella caloplacae s.l. on Xanthoria (id.). G.
Arrhenia peltigerina on Peltigera (Agaricales, Hygrophoraceae). H. Athelia arachnoidea (sexual stage) on Xanthoria (Atheliales, Atheliaceae). I. id.
(sclerotial stage). J. Leucogyrophana lichenicola on Cladonia (Boletales, Coniophoraceae). K. Erythricium aurantiacum on Physcia (Corticiales,
Corticiaceae). L. Marchandiomyces corallinus on Xanthoparmelia (id.). M. Burgellopsis nivea on a sterile crustose lichen (Cantharellales, gen. inc. sed.). N.
Burgella lutea on Sticta (id.). O. Minimedusa pubescens on Scoliciosporum chlorococcum (id.). All photographs by Paul Diederich.
Diederich et al.: 2018 classification and checklist of lichenicolous fungi 347
Many genera that are entirely lichenicolous contain
only one species, but some contain over 30 species
(Table 4). Sequence information is available for
lichenicolous species in 128 (32%) of the 397 genera
containing lichenicolous species, in 56 (28%) of the
198 entirely lichenicolous genera (Table 5), and the
generic type has been sequenced in 171 genera
(some of which include lichenicolous species that
have not yet been sequenced). Many species have
also been brought into culture (listed in www.
lichenicolous.net).
THE 2018 CLASSIFICATION AND CHECKLIST OF
LICHENICOLOUS FUNGI IN THE ASCOMYCOTA AND
BASIDIOMYCOTA
In the following compilation, the 391 fungal
genera containing lichenicolous species in all three
categories (obligately lichenicolous, lichenicolous
lichens, facultatively or doubtfully lichenicolous)
are listed hierarchically by subfamily (where
applicable), family, order, subclass (where applica-
ble), class, subphylum and phylum. Following the
style of the lichen classification (L¨
ucking et al. 2016
[2017]), all categories are listed in strict alphabet-
ical order. For each genus entry, the following
information is given: type, synonyms (only when
the type species is lichenicolous), data on the
existence of sequences [e.g., in ‘‘Mol.: yes (T, L)’’ ,
‘‘yes’’ means that sequences exist, ‘‘T’’ that the
generic type has been sequenced and ‘‘ L’’ that at
least one lichenicolous species has been sequenced],
nomenclatural or taxonomic notes, and number of
taxa in each category [e.g., ‘‘23-1-2 (80)’’ means
that 23 species, subspecies or varieties are obligately
lichenicolous (non-lichenized), 1 taxon is a lichen-
icolous lichen, 2 taxa are facultatively (or doubt-
fully) lichenicolous, while the genus includes a total
of c. 80 species (estimation based on recent
literature and online databases); the latter number
is omitted when all species are lichenicolous].
Species isolated from lichens in culture are included
and are treated as doubtfully lichenicolous. Endo-
lichenic species have sometimes been cultured but
often are known only from DNA sequences. When
they have not been named, they are excluded from
the classification. Species names are given in bold
regular font when obligately lichenicolous,inbold
italic font for lichenicolous lichens,andinbold
regular font underlined when facultatively (or
doubtfully) lichenicolous. An alphabetical listing
of all species follows, along with the most common
knownlichenhosts.Insomeinstances,poorly
known species are included within the genus where
they most likely belong, without formally combin-
ing them there. References cited after each genus
mainly refer to taxonomic treatments. In author
citations of genera and species, ‘‘ ex’’ citations are
omitted.
Phylum ASCOMYCOTA Caval.-Sm.
Subphylum PEZIZOMYCOTINA O.E.Erikss. &
Winka
Class ARTHONIOMYCETES O.E.Erikss. & Winka
Order Arthoniales D.Hawksw. & O.E.Erikss.
Family Arthoniaceae Rchb.
Arthonia Ach. s.l.; type: A. radiata (Pers.) Ach. (lichenized) [¼
Asterotrema M¨
ull.Arg.; type: A. parasiticum;¼Celidiopsis
A.Massal.; type: C. gyrolophii;¼Celidium Tul.; type: C.
fuscopurpureum;¼Charcotia Hue; type: C. rufidula;¼
Conida A.Massal.; type: C. clemens;¼Conida subgen.
Conidella Elenkin; type: C. urceolata;¼Helicobolomyces
Matzer; type: H. lichenicola (asexual), i.e. A. cinnabarinula;
¼Spilodium A.Massal.; type: S. fuscopurpureum;¼Sub-
hysteropycnis Wedin & Hafellner; type: S. maculiformans
(asexual), i.e. A. maculiformis]. Mol.: yes (T, L). Currently
polyphyletic.
Species: 139-1-0 (500). abrothallina Nyl.: on Pseudocyphel-
laria;agelastica R.C.Harris & Lendemer: on Lecanora
louisiana;albotrachynae Etayo: on Hypotrachyna endo-
chlora;almquistii Vain.: on Amygdalaria,Koerberiella,
Porpidia and Trapelia;amandineicola van den Boom &
Ertz: on Amandinea efflorescens;amylospora Almq.: on
Porpidia and Farnoldia;anatolica Halıcı& Candan: on
Aspicilia contorta;anjutae S.Y.Kondr. & Alstrup [as
‘anjutii’]: on Teloschistes;apotheciorum (A.Massal.) Almq.
[¼?A. galactinaria Leight.; ¼?A. subvarians Nyl.]: on
Myriolecis;arthoniicola Diederich & Aptroot: on Arthonia
catenulata;aspiciliae Alstrup & E.S.Hansen: on Aspicilia;
atropunctata Vain.: on sterile foliicolous cf. Echinoplaca;
ayseniae Halıcı& Candan [as ‘aysenae’]: on Acarospora;
Celidium bacidiosporum Steiner: on Parmotrema andinum
[a poorly known species possibly belonging to Arthonia
s.l.]; badia Wedin & Hafellner: on Pseudocyphellaria
granulata;biatoricola Ihlen & Owe-Larss.: on Biatora
efflorescens;brussei Egea & Torrente [”A. capensis Egea &
Torrente,nom.illeg.,nonA. capensis Stizenb.]: on
Lecanographa;caerulescens (Almq.) R.Sant.: on Lecanora
varia;calicii Kantvilas & Wedin [as ‘caliciae’]: on Calicium
tricolor;catillarioides Etayo: on Sticta;cinnabarinula
M¨
ull.Arg. [¼Helicobolomyces lichenicola Matzer]: on
foliicolous Porina and Trichothelium;circinata Th.Fr.: on
Umbilicaria;clemens (Tul.) Th.Fr. [”Conida clemens
(Tul.) A.Massal.]: on Rhizoplaca;cohabitans Coppins: on
Arthothelium reagens;colombiana Etayo: on Cladonia;
coniocraeae Brackel: on Cladonia coniocraea;coriifoliae
348 The Bryologist 121(3): 2018
Wedin & Hafellner: on Pseudocyphellaria coriifolia;coro-
nata Etayo: on Flavoparmelia caperata;cryptotheciae
Matzer & R.Sant.: on foliicolous Myriostigma candida;
darbishirei Follmann & Werner: on Roccella nigerrima;
destruens Rabenh. var. destruens:onPhyscia;destruens
var. nana Grube & Hafellner: on Xanthomendoza fallax;
digitatae Hafellner: on Cladonia [often co-occurring with
Milospium lacoizquetae]; digitispora Etayo [as ‘digito-
spora’]: on Sticta;diorygmae S.Joshi, D.K.Upreti &
S.Nayaka: on Diorygma;diploiciae Calat. & Diederich: on
Diploicia canescens;epicladonia (Nyl.) Alstrup & Zhurb. [”
Scutula epicladonia (Nyl.) Sacc.]: on Cladonia;epifarinosa
Etayo: on Pannaria;epimela (Almq.) I.M.Lamb: on
Amandinea punctata;epiphyscia Nyl.: on Physcia;epito-
ninia Halıcı& Candan: on Toninia;excentrica Th.Fr.: on
Lepraria;farinacea (H.Olivier) R.Sant.: on Ramalina,
especially R. farinacea;flavicantis Wedin & Hafellner: on
Pseudocyphellaria flavicans;follmaniana Diederich: on
Roccella;fuscopurpurea (Tul.) R.Sant. [”Celidium fusco-
purpureum Tul.; ”Spilodium fuscopurpureum (Tul.)
A.Massal.]: on Peltigera;garajonayi Etayo: on Porina;
gelidae R.Sant.: on Placopsis gelida;glacialis Alstrup &
E.S.Hansen: on Rhizoplaca melanophthalma;glaucomaria
(Nyl.) Nyl.: on Lecanora rupicola;graphidicola Coppins: on
Graphis scripta;griseopruinosa F.Berger & E.Zimm.: on
Pertusaria pseudocorallina;Celidiopsis gyrolophii A.Mas-
sal.: on Cora [a poorly known species pertaining to
Arthonia s.l.]; haematommatum Kalb & Hafellner: on
Haematomma;haematostigma Grube & Hafellner: on
Ramalina;hawksworthii Halıcı:onDimelaena;hertelii
(Calat., Barreno & V.J.Rico) Hafellner & V.John: on
Aspicilia;heterodermiae Etayo: on Heterodermia;hodgesii
Lendemer & R.C.Harris: on Graphis lineola;infectans Egea
& Torrente: lichen, juvenile lichenicolous on Lecanographa
hypothallina and Sparria cerebriforme;ingaderiae Foll-
mann: on Ingaderia;insularis Kantvilas & Wedin: on
Sirenophila eos;intermedia Matzer: on foliicolous Porina;
intexta Almq.: on Lecidella;invadens Coppins: on
Schismatomma quercicola;japewiae Grube & Holien: on
Japewia;lecaniicola van den Boom & M.Haji Moniri: on
Lecania triseptata;lecanoricola Alstrup & Olech [as
‘lecanoriicola’]: on Lecanora populicola;lecanorina (Almq.)
R.Sant.: on Lecanora albella;lepidiota (H.Olivier) R.Sant.:
on Fuscopannaria praetermissa;Abrothallus lepidophilus
Anzi: on Cladonia [a poorly known species possibly
belonging to Arthonia s.l.]; lethariicola Alstrup &
M.S.Cole: on Letharia;leucomelodis F.Berger & E.Zimm.:
on Leucodermia leucomelos;lobariellae Etayo: on Lobariella;
lobothalliae Etayo: on Lobothallia alphoplaca;maculifor-
mis Wedin & Hafellner [¼Subhysteropycnis maculiformans
Wedin & Hafellner]: on Pseudocyphellaria glabra and P.
homoeophylla; marginalis Etayo: on Pseudocyphellaria;
mazosiicola Matzer & R.Sant. [¼A. mazosiae R. Sant.,
nom. inval., nom. illeg., non A. mazosia (A.Massal.)
Willey]: on foliicolous Mazosia;microsticta Vain.: on
foliicolous cf. Fellhanera bouteillei [a species of doubtful
application]; minuta Wedin & Hafellner: on Pseudocyphel-
laria coriifolia;molendoi (Frauenf.) R.Sant.: on Calogaya
and Rusavskia;navasiana Etayo: on Lepraria;neglectula
Nyl.: on a sterile, sorediate crust (Lepraria neglecta?);
nephromiaria Nyl.: on Nephroma;nideri (J.Steiner)
Clauzade, Diederich & Cl.Roux, comb. inval.: on Pyreno-
desmia;obscurior Triebel: on Pilophorus dovrensis;oligo-
spora Vˇ
ezda: on Myriolecis crenulata;pannariae Zhurb. &
Grube: on Protopannaria pezizoides and Psoroma hypnorum;
pantherina Etayo: on Pertusaria pustulata;Asterotrema
parasiticum M¨
ull.Arg.: on Athallia pyracea [probably
belongs to Arthonia, fide Aptroot 1991; incorrectly cited
as Arthonia parasitica (M¨
ull.Arg.) Aptroot by Sundin et al.
2012 and Frisch et al. 2014b]; parietinaria Hafellner &
A.Fleischhacker: on Xanthoria parietina;parmeliae
(C.W.Dodge & G.E.Baker) D.Hawksw. & Iturr.: on Physcia
caesia;peltigerea Th.Fr.: on Peltigera and Solorina;
peltigerina (Almq.) H.Olivier: on Peltigera and Solorina;
pelvetii (Hepp) H.Olivier: on Crocodia aurata;pepei Etayo
&P
´
erez-Ortega: on Parmelina;phaeophysciae Grube &
Matzer: on Phaeophyscia;physcidiicola Frisch & G. Thor:
on Physcidia wrightii;plectocarpoides (S.Y.Kondr. &
D.J.Galloway) Wedin & S.Y.Kondr.: on Pseudocyphellaria
scabrosa;polia Etayo & R.Sant.: on Diploicia canescens;
polydactylonis Diederich & Ertz, nom. nov. [”A. ceracea
Etayo & Breuss nom. illeg.]: on Peltigera polydactylon [a
younger homonym of Arthonia ceracea F´
ee]; pragensis
Hora
´kova
´:onCaloplaca teicholyta;prominens Follmann:
on Pentagenella gracillima;protoparmeliae Etayo: on
Protoparmelia badia;protoparmeliopseos Etayo & Die-
derich: on Protoparmeliopsis muralis;pseudocyphellariae
Wedin: on Pseudocyphellaria multifida;pseudopegraphina
Matzer: on foliicolous Mazosia;punctella Nyl.: on
Diplotomma;rangiformicola Brackel & Etayo: on Cladonia
rangiformis;rimeliicola Diederich: on Parmotrema;rino-
dinicola Candan & Halıcı:onRinodina oleae;rubescens
(Arnold) Clauzade, Diederich & Cl.Roux: on Buellia
epipolia;rufidula (Hue) D.Hawksw., R.Sant. & Øvstedal
[”Charcotia rufidula Hue]: on Umbilicaria;sagenidii
Vˇ
ezda & Kantvilas: on Lecanactis mollis;sampaianae
(Diederich & Etayo) Ertz & Diederich [”Plectocarpon
sampaianae Diederich & Etayo]: on Nevesia sampaiana;
santessoniana Wedin & Hafellner: on Pseudocyphellaria
scabrosa;santessonii Matzer: on foliicolous Porina;semi-
immersa Wedin & Hafellner: on Pseudocyphellaria;sher-
parum Grube & Matzer: on Lecanora sherparum;squamar-
inae Etayo: on Squamarina;stereocaulina (Ohlert) R.Sant.:
on Stereocaulon;stevensoniana R.C.Harris & Lendemer: on
Haematomma accolens;stictaria Nyl.: on Pseudocyphellaria;
stipitata M.M.E.Alves, Aptroot & M.Ca
´ceres: on uniden-
tified sterile, sorediate, corticolous lichen (?Bacidina,Fell-
hanera); subaggregata Wedin & Hafellner: on
Pseudocyphellaria hirsuta;subconveniens Nyl. [¼?A.
ricasoliae M¨
ull.Arg.]: on Lobaria;subfuscicola (Linds.)
Triebel: on Lecanora albella and L. carpinea;subvelutinae
(Vain.) R.Sant.: on foliicolous Sporopodium;sytnikii
S.Y.Kondr.: on Dufourea;tabescens (Anzi) Jatta: on
Parmelia saxatilis;tavaresii Grube & Hafellner: on
Anthracothecium and Pyrenula;tetraspora S.Y.Kondr. &
K¨
arnefelt: on Caloplaca chilensis;thelotrematis Coppins:
on Thelotrema;toensbergii Holien & Frisch: on Mycoblas-
tus affinis;tremelloides Etayo: on Heterodermia;triebeliae
Zhurb.: on Dactylina;urceolata (Elenkin) Calat., Barreno &
V.J.Rico [”Conida urceolata Elenkin]: on Aspicilia;varians
(Davies) Nyl.: on Lecanora bicincta,L. helicopis,L. rupicola
and L. swartzii;versoeensis Alstrup [¼A. aggregata Alstrup,
M.S.Christ. & Hafellner]: on Amandinea punctata;xantho-
parmeliarum Etayo: on Xanthoparmelia.
Lit.: Alstrup 1993a; Alstrup & Cole 1998; Alstrup & Hansen 2001;
Alstrup & Olech 1996; Alves et al. 2014; Aptroot et al. 1995,
1997; Berger & Zimmermann 2016; Brackel 2010a, 2015;
Calatayud et al. 1995, 2004a; Candan & Halıcı2009; Diederich
1995; Ertz et al. 2005; Etayo 1993, 1996, 2002a, 2008, 2010b,
2010c, 2017, 2018b; Etayo & Breuss 1998; Etayo & Diederich
2009; Etayo & P´
erez-Ortega 2016; Etayo & Sancho 2008;
Fleischhacker et al. 2016; Follmann & Werner 2003; Frisch &
Holien 2018; Frisch et al. 2014a; Grube & Matzer 1997; Grube et
Diederich et al.: 2018 classification and checklist of lichenicolous fungi 349
al. 1995; Hafellner 2009; Halıcı2008; Halıcı&Candan2009,
2011; Hawksworth 1991a; Hawksworth & Iturriaga 2006; Hertel
1969; Hora
´kova
´1994; Ihlen et al. 2004b; Joshi et al. 2013; Kalb et
al. 1995; Kantvilas & Vˇ
ezda 1992; Kantvilas & Wedin 2015;
K¨
arnefelt et al. 2002; Kocourkova
´et al. 2012; Kondratyuk 1996a;
Kondratyuk & Zelenko 2002; Lendemer et al. 2016; Matzer 1996;
Santesson 1993a; Santesson et al. 2004; Triebel et al. 1991; van
den Boom & Haji Moniri 2018; van den Boom et al. 2017; Wedin
& Hafellner 1998; Wedin & Kondratyuk 1997; Zhurbenko 2002;
Zhurbenko & Grube 2010.
Briancoppinsia Diederich, Ertz, Lawrey & van den Boom; type:
B. cytospora. Mol.: yes (T, L). Asexual, pycnidial.
Species: 1-0-0. cytospora (Vouaux) Diederich, Ertz, Lawrey
&vandenBoom[”Phoma cytospora (Vouaux)
D.Hawksw.]: on parmelioid lichens, Cladonia,Lecanora
conizaeoides,Lepra, etc..
Lit.: Diederich et al. 2011b.
Family Lecanographaceae Ertz, Tehler, G.Thor &
Frisch
Lecanographa Egea & Torrente; type: L. lyncea (Sm.) Egea &
Torrente (lichenized). Mol.: yes (T).
Species: 3-1-0 (38). dialeuca (Cromb.) Egea & Torrente:
lichen, facultatively lichenicolous on Roccella;imitans
Werner & Follmann: on Roccella humboldtiana;occidenta-
lis Egea & Torrente: on Roccellina;rinodinae (Vˇ
ezda)
R.Sant.: on Phaeorrhiza nimbosa.
Lit.: Egea & Torrente 1994; Follmann & Werner 2003; Gonnet et al.
2017; Hafellner 2009; van den Boom & Etayo 2006; Vˇ
ezda 1969.
Phacographa Hafellner; type: P. glaucomaria. Mol.: yes (T, L).
Species: 3-0-0. glaucomaria (Nyl.) Hafellner [¼Opegrapha
maculans (Arnold) Poelt & T¨
urk]: on Lecanora rupicola s.l.
(incl. L. bicincta); protoparmeliae Hafellner: on Proto-
parmelia badia;zwackhii (Zwackh) Hafellner [”Opegra-
pha zwackhii (Zwackh) Hafellner]: on Phlyctis.
Lit.: Hafellner 2009.
Plectocarpon F´
ee; type: P. pseudosticta [”Delisea F´
ee, nom.
illeg.; ¼Epiphora Nyl.; type: E. encaustica;¼Lichenomyces
Trevis.; type: L. parasiticus Trevis., nom. illeg., i.e. P.
lichenum]. Mol.: yes (L).
Species: 40-0-0 (41). aequatoriale Etayo: on Sticta;
arthonioides Diederich: on Lobaria isidiophora;bunodo-
phori Wedin, Ertz & Diederich: on Bunodophoron patago-
nicum;cladoniae R.Sant.: on Cladonia;concentricum Ertz,
Diederich & Wedin: on Pseudocyphellaria homeophylla;
coppinsii Ertz & Diederich: Pseudocyphellaria crocata;
cristalliferum Christnach, Ertz & Diederich: on Sticta;
diedertzianum Y.Joshi, Upadhyay & Chandra: on Myelo-
chroa;dimorphosporum Etayo, Flakus & Kukwa: on
Ricasolia patinifera;dirinariae Ertz & van den Boom: on
Dirinaria;encausticum (Nyl.) R.Sant. [”Epiphora encaus-
tica Nyl.]: on Brodoa intestiniformis;gallowayi (S.Y.Kondr.)
Ertz & Diederich: on Pseudocyphellaria dissimilis;gayanum
Etayo: on Siphula;hypogymniae Zhurb. & Diederich: on
Hypogymnia bitteri;lambinonii Diederich & Etayo: on
Lobaria;latisporum Ertz, Diederich & Wedin: on Pseudo-
cyphellaria coriifolia;leuckertii (S.Y.Kondr. & D. J.Gallo-
way) Ertz & Diederich: on Pseudocyphellaria intricata and P.
norvegica;lichenum (Sommerf.) D.Hawksw.: on Lobaria
anomala,L. anthraspis and L. pulmonaria;linitae (R.Sant.)
Wedin & Hafellner: on Lobaria linita;macaronesiae
Diederich, Etayo & S´
erus.: on Lobaria immixta and L.
macaronesiae;melanohaleae Christnach, Ertz & Diederich:
on Melanohalea ushuaiensis;nephromeum (Norman)
R.Sant.: on Nephroma bellum;obtectum Ertz & Diederich:
on Pseudocyphellaria encoensis;opegraphoideum Christ-
nach, Ertz, Diederich & Wedin: on Pseudocyphellaria
homeophylla and P. multifida;parmotrematis Etayo, Flakus
& Kukwa: on Parmotrema reticulatum;peltigerae Zhurb.,
Ertz, Diederich & Mia
˛dl.: on Peltigera leucophlebia;
pseudocyphellariae Diederich: on Pseudocyphellaria gilva;
pseudoleuckertii Diederich, Ertz & Wedin: on Pseudocy-
phellaria coriifolia;pseudosticta (F´
ee) F´
ee: on Pseudocy-
phellaria freycinetii and P. glabra;ramalinae Etayo &
Flakus: on Ramalina celastri;scrobiculatae Diederich &
Etayo: on Lobarina scrobiculata;serusiauxii Ertz & Die-
derich: on Sticta;stereocaulicola Kukwa, Etayo & Flakus:
on Stereocaulon;sticticola Ertz, Wedin & Diederich: on
Sticta;tibellii Ertz & Diederich: on Pseudocyphellaria
rubella;triebeliae Diederich & Ertz: on Ricasolia quercizans;
usneae Diederich & Etayo: on Usnea exasperata;usneaus-
tralis Etayo: on Usnea;venustum Ertz, Coppins &
Diederich: on Nephroma papillosum;violaceum Ertz,
R.Sant., Diederich & Wedin: on Nephroma antarcticum.
Lit.: Aptroot et al. 1997; Diederich & Ertz 2018; Diederich & Etayo
1994; Ertz & Diederich 2007; Ertz & van den Boom 2012; Ertz et al.
2003, 2005; Etayo 2007, 2017; Etayo & Sancho 2008; Etayo et al.
2018; Hawksworth & Galloway 1984; Joshi et al. 2016b; Kukwa et
al. 2012; Santesson 1993a, 1994a; Scholz 1998; Wedin & Hafellner
1998; Zhurbenko et al. 2008.
Family Opegraphaceae Stizenb.
Opegrapha Ach. s.l.; type: O. vulgata (Ach.) Ach. (lichenized) [¼
Kalaallia Alstrup & D.Hawksw.; type: K. reactiva;¼
Leciographa A.Massal.; type: L. parasitica;¼Lecoglyphis
Clem.; type: L. centrifuga (see under O. rupestris); ¼
Mycopegrapha Vain.; type: M. hymeniicola;¼Opegraphoidea
Fink; type: O. staurotheliicola]. Mol.: yes (T, L). Currently
polyphyletic.
Species: 72-0-0 (300). agelaea F´
ee: on pyrenocarpous
lichen; anomea Nyl. [¼Leciographa pertusariae Vouaux; ¼
O. wetmorei M.S.Cole & D.Hawksw.]: on Lepra,Ochro-
lechia,Pertusaria and Varicellaria;bacidiae R.Sant.: on
Brasilicia brasiliensis;blakii Ertz & Diederich: on cf.
Ochrolechia;brevis Coppins: on Thelotrema petractoides;
brevissima Kalb & Hafellner: on Haematomma;briancop-
pinsii S.Y.Kondr., L.L}
ok¨
os&J.-S.Hur:onPorpidia
albocaerulescens;brigantina Hafellner: on Brigantiaea;
bryoriae Zhurb.: on Bryoria;buelliae Zhurb.: on Buellia
insignis;chapsae Etayo: on Chapsa;cladoniicola Ertz &
Diederich: on Cladonia;cryptotheciae Matzer: on foliic-
olous Myriostigma candidum;diffracticola R.C.Harris &
Ladd: on Bacidia diffracta;ectolechiacearum Matzer &
R.Sant.: on foliicolous Calopadia,Loflammia and Sporopo-
dium;encephalographoidea Diederich & Aptroot: on
Pyrenula;enterographae Etayo & R.Sant.: on Enterogra-
pha;epiporina Matzer: on foliicolous Porina;foreaui
(Moreau) Hafellner & R.Sant. [¼O. trassii S.Y.Kondr. &
Coppins]: on Heterodermia;Leciographa furfuracea
(Anzi) Rehm: on Lecanora rupicola and Rhizocarpon [a
poorly known species pertaining to Opegrapha s.l.];
geographicola (Arnold) Hafellner [”Weddellomyces geo-
graphicola (Arnold) Alstrup & D.Hawksw.]: on Rhizocar-
pon;gregalis Vain.: on pyrenocarpous lichen; grossulina
M¨
ull.Arg.: on Pyrenulaceae; Leciographa gyrolophii
350 The Bryologist 121(3): 2018
(A.Massal.) Vouaux: on Cora [a poorly known species
pertaining to Opegrapha s.l.]; hafellneri E.Zimm., Etayo &
F.Berger: on Placopsis gelida;hellespontica Vondra
´k&
Kocourk.: on Variospora aurantia;Mycopegrapha hyme-
niicola Vain.: on Lecanora [a poorly known species
pertaining to Opegrapha s.l.]; insidens (J.Steiner)
S.Y.Kondr.: on Athallia pyracea,Pyrenodesmia variabilis
and Verrucaria buschirensis;invadens Etayo: on Pannaria
farinosa;kalbii Matzer: on foliicolous Byssoloma;lamyi
(Nyl.) Triebel: on corticolous Lecanora;lepidella Nyl.: on
cf. Pertusaria;lichenicola Thor, L¨
ucking & Tat.Matsumo-
to: on Porina;lopezariae Etayo & Sipman: on Lopezaria;
maligna Triebel: on Lecidea fuscoatra;matzeri L¨
ucking: on
foliicolous Amazonomyces sprucei;mazosiae Matzer: on
foliicolous Mazosia;melanospila M¨
ull.Arg.: on Parmotre-
ma;Leciographa nephromatis Stein: on Nephroma [a
poorly known species pertaining to Opegrapha s.l.];
nigeriensis C.W.Dodge: on Cryptothecia;parasitica
(A.Massal.) H.Olivier [”Leciographa parasitica A.Massal.]:
on Aspicilia calcarea;perturbans Follmann: on Ingaderia
pulcherrima;pertusariicola Coppins & P.James: on
Pertusaria leioplaca;phaeophysciae R.Sant., Diederich,
Ertz & Christnach: on Phaeophyscia hispidula;phlyctidi-
cola (Vouaux) Etayo: on Phlyctis;phyllobathelii Matzer &
R.Sant.: on foliicolous Phyllobathelium;phylloporinae
M¨
ull.Arg.: on foliicolous Porina;physcidiae Kalb & Elix:
on Physcidia australasica;pigozziana Etayo & Aptroot: on
Thelotrema;plectocarpoidea Diederich: on Phaeographis;
porinicola Matzer: on foliicolous Porina;pulvinata Rehm
var. pulvinata [¼?Homostegia dermatocarponis Alstrup &
M.Cole (as ‘dermatocarpae’); ¼?Phaeospora dermatocarpo-
nis (R.G.Werner) S.Y.Kondr.]: on Dermatocarpon, possibly
also Catapyrenium and Endocarpon;pulvinata var. plac-
idiicola Zhurb.: on Placidium;reactiva (Alstrup &
D.Hawksw.) Etayo & Diederich [”Kalaallia reactiva
Alstrup & D.Hawksw.]: on Hymenelia lacustris;reinkellae
Follmann: on Roccella lirellina;rotunda Hafellner: on
Physconia distorta;rouxiana Nav.-Ros. & Hladun: on
Polyblastia nidulans;rupestris Pers. [¼?O. centrifuga
A.Massal.; ”Lecoglyphis centrifuga (A.Massal.) Clem.]:
on Verrucaria;sipmanii Matzer: on foliicolous Porina;
sphaerophoricola Isbrand & Alstrup: on Sphaerophorus
globosus;Opegraphoidea staurotheliicola Fink [¼?Ope-
grapha romsiae S.Y.Kondr. & Kudratov (as ‘romsae’)]: on
Staurothele [a poorly known species pertaining to Ope-
grapha s.l.]; stellanigra Etayo: on Sticta;stereocaulicola
Alstrup & D.Hawksw.: on Stereocaulon;stigmodes Nyl.: on
Porina;strigulae Matzer & R.Sant.: on foliicolous Strigula;
submaritima Hafellner: on Tephromela atra;tenuior Stirt.:
on Thelotrema;thelotrematis Coppins: on Thelotrema
lepadinum and T. macrosporum;uniseptata Matzer: on
foliicolous Strigula;velata (M¨
ull.Arg.) Vain.: on foliicolous
Actinoplaca,Calenia,?Echinoplaca and Gyalectidium;
verseghyklarae S.Y.Kondr., L.L}
ok¨
os & J.-S.Hur: on
Ochrolechia pallescens;vulpina Vondra
´k, Kocourk. &
Tretiach: on Caloplaca s.l., incl. Leproplaca cirrochroa,
Pyrenodesmia erodens and Variospora flavescens.
Lit.: Alstrup & Cole 1998; Alstrup & Hawksworth 1990; Aptroot el
al. 1997; Awasthi 1991; Berger & Zimmermann 2016; Clauzade et
al. 1989; Coppins & Kondratyuk 1998; Ertz 2009; Ertz & Diederich
2003; Ertz et al. 2004, 2005; Etayo 1996, 2007, 2010b, 2017; Etayo &
Aptroot 2006; Etayo & Sancho 2008; Follmann & Werner 2003;
Hafellner 1985b, 1994a, 2007, 2009; Harris & Ladd 2007; Isbrand &
Alstrup 1992; Kalb & Elix 1995; Kalb et al. 1995; Kondratyuk &
Galloway 1995c; Kondratyuk & Kudratov 2002; Kondratyuk &
Zelenko 2002; Kondratyuk et al. 2015, 2016b; L¨
ucking 1998; Matzer
1996; Navarro-Rosin´
es & Hladun 1995; Orange 2002; Triebel 1989;
Vondra
´k & Kocourkova
´2008; Zhurbenko 2009b, 2013a; Zhurben-
ko & Vershinina 2014.
Paralecanographa Ertz & Tehler; type: P. grumulosa. Mol.: yes
(T, L).
Species: 1-1-0. grumulosa (Dufour) Ertz & Tehler [”
Opegrapha grumulosa Dufour; incl. Lecanographa hemi-
sphaerica (Laundon) Egea & Torrente]: initially lichen-
icolous on Roccella,rarelyonDirina,Gorgadesia or
Opegrapha, eventually developing an own thallus; thelop-
sisocia (B. de Lesd.) Cl.Roux, comb. provis. [”Opegrapha
grumulosa var. thelopsisocia B. de Lesd.; ”O. thelopsisocia
(B. de Lesd.) B. de Lesd.]: on Thelopsis isiaca.
Lit.: Egea & Torrente 1994, Roux 2014.
Family Roccellaceae Chevall.
Cresponea Egea & Torrente; type: C. premnea (Ach.) Egea &
Torrente (lichenized). Mol.: yes.
Species: 1-0-0 (21). lichenicola Aptroot & M.Ca
´ceres: on
Pyrenula.
Lit.: Ca
´ceres et al. 2014.
Enterographa F´
ee; type: E. crassa (DC.) F´
ee (lichenized). Mol.:
yes (T).
Species: 9-0-1 (55). bagliettoae F.Seavey & J.Seavey: on
Bagliettoa;brezhonega Sparrius & Aptroot: on Porina rosei
and possibly other lichens with Trentepohlia;confusa Ertz
& Van den Broeck: on Arthonia ochraceella;epigraphis
Etayo & Sipman: on Graphis;epiphylla (S´
erus.) Ertz,
Diederich & Sparrius: on foliicolous Coenogonium geralense
[¼C. flavicans]; fellhaneroides Yeshitela, Eb.Fisch., Kill-
mann & S´
erus.: on Coenogonium;mazosiae Matzer &
R.Sant.: on foliicolous Mazosia;meklitiae Yeshitela,
Eb.Fisch., Killmann & S´
erus.: probably lichenicolous over
Coenogonium;osagensis C.A.Morse: on Bacidina;punctata
Ertz & Diederich: on Lobaria.
Lit.: Ca
´ceres et al. 2001; Ertz el.2005; Etayo 2017; Matzer 1996;
Morse 2013; Seavey et al. 2017; Sparrius & Aptroot 2007; Van den
Broeck & Ertz (2018).
Mazosia A.Massal.; type: M. rotula (Mont.) A.Massal. (liche-
nized). Mol.: yes.
Species: 1-0-0 (27). adelphoparasitica Matzer: on foliico-
lous Mazosia rotula.
Lit.: Matzer 1996.
Schismatomma A.Massal. s.l.; type: S. pericleum (Ach.) Branth &
Rostr. (lichenized). Mol.: yes (T).
Species: 1-0-0 (10). physconiicola Ertz & Diederich: on
Physconia.
Lit.: Ertz & Diederich 2006.
Sigridea Tehler; type: S. californica (Tuck.) Tehler (lichenized).
Mol.: yes (T).
Species: 1-0-0 (6). labyrinthica (Follmann) Ertz & Die-
derich [”Plectocarpon labyrinthicum Follmann]: on
Roccella portentosa.
Lit.: Ertz et al. 2005.
Order Arthoniales genera incertae sedis
Arthophacopsis Hafellner; type: A. parmeliarum. Mol.: no.
Species: 1-0-0. parmeliarum Hafellner: on Parmelia s.str.
Lit.: Hafellner 1998a.
Diederich et al.: 2018 classification and checklist of lichenicolous fungi 351
Arthothelium A.Massal.; type: A. spectabile A.Massal. (liche-
nized). Mol.: yes (T). Currently polyphyletic.
Species: 1-0-0 (140). hymeniicola Ertz & Fryday: on
Bacidia (hymenium).
Lit.: Ertz & Fryday 2017.
Mixtoconidium Etayo; type: M. canariense. Mol.: no.
Species: 2-0-0. insidiens (Vouaux) Etayo & van den Boom
[¼M. canariense Etayo]: on Ramalina;nashii (Hafellner)
Etayo & van den Boom [”Plectocarpon nashii Hafellner]:
on Niebla robusta.
Lit.: Ertz & Diederich 2007; Etayo 1995a; van den Boom & Etayo
2017.
Paradoxomyces Matzer; type: P. nymanii. Mol.: no.
Species: 1-0-0. nymanii Matzer: on foliicolous Strigula.
Lit.: Matzer 1996.
Perigrapha Hafellner [a hemihomonym of Perigrapha Lederer
(Lepidoptera)]; type: P. superveniens. Mol.: no.
Species: 5-0-0. cetrariae Zhurb.: on Cetraria laevigata;
lobariae Zhurb.: on Lobaria japonica agg.; nitida Ertz,
Diederich, Christnach & Wedin: on Pseudocyphellaria glabra;
phaeophysciae Etayo & P´
erez-Vargas: on Phaeophyscia cf.
orbicularis;superveniens (Nyl.) Hafellner: on Parmelia s.str.
Lit.: Ertz et al. 2005; Hafellner 1996a; P´
erez-Vargas et al. 2013;
Zhurbenko & Ohmura 2018; Zhurbenko et al. 2015a.
Phacothecium Trevis.; type: P. varium. Mol.: no.
Species: 1-0-0. varium (Tul.) Trevis. [”Arthonia varia
(Tul.) Jatta; ¼Opegrapha physciaria (Nyl.) D.Hawksw. &
Coppins]: on Xanthoria.
Lit.: Hafellner 2009.
Trichophyma Rehm; type: T. bunchosiae. Mol.: no.
Species: 2-0-0. bunchosiae Rehm: on foliicolous Strigula;
similis Matzer: on foliicolous Strigula.
Lit.: Matzer 1996.
Order Lichenostigmatales Ertz, Diederich & Lawrey
Family Phaeococcomycetaceae McGinnis & Schell
Etayoa Diederich & Ertz; type: E. trypethelii. Mol.: yes (T, L).
Species: 1-0-0. trypethelii (Flakus & Kukwa) Diederich &
Ertz [”Phaeosporobolus trypethelii. Flakus & Kukwa]: on
Graphidaceae, Lecanora,Pertusaria,Trypetheliaceae and
other crustose lichens.
Lit.: Ertz et al. 2014b; Flakus & Kukwa 2012.
Lichenostigma Hafellner; type: L. maureri [¼Diederimyces Etayo
(not validly published); type: D. fuscideae Etayo; ¼
Phaeosporobolus D.Hawksw. & Hafellner; type: P. usneae
D.Hawksw. & Hafellner]. Mol.: yes (T, L). Many lichen-
icolous Lichenostigma species do not belong to Lichen-
ostigma s.str., but obviously pertain to Lichenotheliaceae;
they are treated under that family within Lichenostigma s.l.
Species: 5-0-0. alpinum (R.Sant., Alstrup & D.Hawksw.)
Ertz & Diederich [”Phaeosporobolus alpinus R.Sant.,
Alstrup & D.Hawksw.; incl. Diederimyces fuscideae Etayo,
D. microsporus Etayo, P. minutus Etayo]: on Lepra,
Ochrolechia,Varicellaria and other lichens; chlaroterae
(F.Berger & Brackel) Ertz & Diederich [”P. chlaroterae
F.Berger & Brackel]: on Lecanora and other lichens;
fellhanerae (R.C.Harris & Lendemer) Ertz & Diederich
[”P. fellhanerae R.C.Harris & Lendemer]: on Fellhanera;
hyalosporum Kalb & Hafellner [as ‘hyalospora’]: on
Haematomma eremaeum;maureri Hafellner [¼P. usneae
D.Hawksw. & Hafellner]: mainly on fruticose lichens.
Lit.: Ertz et al. 2014b; Etayo 1995b; Etayo & Sancho 2008;
Hawksworth & Hafellner 1986; Kalb et al. 1995.
Class DOTHIDEOMYCETES O.E.Erikss. & Winka
Order Abrothallales P´
erez-Ort. & Suija [¼
Lichenoconiales Diederich, Lawrey & K.D.Hyde]
Family Abrothallaceae P´
erez-Ort. & Suija [¼?
Lichenoconiaceae Diederich & Lawrey]
Abrothallus De Not.; type: A. bertianus [¼Epinephroma Zhurb.;
type: E. kamchatica;¼Pseudolecidea Marchand; type: P.
parmeliarum, i.e. A. parmeliarum;¼Vouauxiomyces Dyko &
D.Hawksw.; type: V. truncatus, i.e. A. microspermus]. Mol.:
yes (L).
Species: 42-0-0. acetabuli Diederich: on Pleurosticta
acetabulum;bertianus De Not. [¼A. glabratulae I.Kotte]:
on Melanelixia,alsoreportedfromMelanohalea and
other parmelioid lichens; boomii P´
erez-Ortega & Suija:
on Nephroma;brattii (S.Y.Kondr.) Suija & P´
erez-Ortega:
on Pseudocyphellaria faveolata;bryoriarum Hafellner: on
Bryoria;buellianus De Not.: on Parmelina;caerulescens
I.Kotte [¼?A. tulasnei M.S.Cole & D.Hawksw.]: on
Xanthoparmelia;canariensis P´
erez-Ortega, van den
Boom & Suija: on Crocodia aurata;chrysanthus Stein:
on Usnea;cladoniae R.Sant. & D.Hawksw.: on Cladonia;
doliiformis P´
erez-Ortega & Suija: on Sticta;eriodermae
Suija, Etayo & P´
erez-Ortega: on Erioderma;ertzii Suija &
P´
erez-Ortega: on Lobaria pulmonaria;etayoi P´
erez-
Ortega & Suija: on Sticta;granulatae Wedin: on
Pseudocyphellaria;halei P´
erez-Ortega, Suija, D.Hawksw.
&R.Sant.:onLobaria;heterodermiicola Etayo &
F.Berger: on Leucodermia leucomelos;hypotrachynae
Etayo&Diederich:onHypotrachyna;kamchatica
(Zhurb. & Stepanchikova) P ´
erez-Ortega & Suija [”
Epinephroma kamchatica Zhurb. & Stepanchikova]: on
Nephroma;lobariae (Diederich & Etayo) Diederich &
Ertz, comb. nov. [”Phoma lobariae Diederich & Etayo]:
on Lobaria;macrosporus Etayo & R.Sant.: on Lobariella;
microspermus Tul. [¼Vouauxiomyces truncatus (B. de
Lesd.) Dyko & D.Hawksw.]: on Flavoparmelia caperata;
nephromatis Suija & P´
erez-Ortega: on Nephroma;niger
Etayo: on Hypotrachyna,andpossiblyParmotrema and
Usnea;parmeliarum (Sommerf.) Arnold [”Pseudoleci-
dea parmeliarum (Sommerf.) Marchand; ¼Abrothallus
cetrariae I.Kotte]: on Nesolechia (mainly N. oxyspora,
rarely N. oxyspora var. fusca or N. thallicola,withor
without visible galls) over Parmelia,Parmotrema,Plati-
smatia,Xanthoparmelia,etc.;parmotrematis Diederich:
on Parmotrema;peyritschii (Stein) I.Kotte: on Cetraria;
pezizicola Diederich & R.C.Harris: on Cladonia pezizi-
formis;prodiens (Harm.) Diederich & Hafellner: on
Hypogymnia;psoromatis (Zhurb. & U.Braun) Diederich
& Zhurb., comb. nov. [”Phoma psoromatis Zhurb. &
U.Braun]: on Psoroma;puntilloi Brackel: on Lobaria
pulmonaria;ramalinae Diederich: on Ramalina;Vouaux-
352 The Bryologist 121(3): 2018
iomyces santessonii D.Hawksw.: on Platismatia glauca;
secedens Wedin & R.Sant.: on Pseudocyphellaria;stereo-
caulorum Etayo & Diederich: on Stereocaulon;stictarum
Etayo: on Sticta;stroblii Hafellner: on Menegazzia
terebrata;suecicus (Kirschst.) Nordin [¼Vouauxiomyces
ramalinae (Nordin) D.Hawksw.; ”Phoma ramalinae
Nordin]: on Ramalina;teloschistis Brackel, P ´
erez-Ortega
&Suija:onSeirophora and Teloschistes;usneae Rabenh.
var. usneae:onUsnea;usneae var. tetraspora Etayo &
Osorio: on Usnea;welwitschii Mont.: on Sticta.
Lit.: Brackel 2015; Brackel & Puntillo 2016; Diederich 1990,
2003, 2004a, 2011; Diederich et al. 2017; Etayo 2002, 2010b,
2017; Etayo & Diederich 1995; Etayo & Osorio 2004; Etayo &
van den Boom 2006; Hafellner 1994a; Hafellner et al. 2008;
Hawksworth 1990a; Ihlen & Wedin 2008; Kondratyuk 1996b;
Nordin 1964; P ´
erez-Ortega et al. 2014; Suija 2006; Suija et al.
2011, 2015a; Wedin 1994; Zhurbenko & Braun 2013; Zhurbenko
et al. 2012.
Lichenoconium Petr. & Syd.; type: L. lichenicola. Mol.: yes (L).
Species: 15-0-0. aeruginosum Diederich, M.Brand, van
den Boom & Lawrey: on Cladonia;cargillianum (Linds.)
D.Hawksw.: on Letharia,Parmotrema,Platismatia,Ram-
alina and Usnea;christiansenii M.S.Cole & D.Hawksw.:
on Nodobryoria abbreviata;echinosporum D.Hawksw.:
on Cladia muelleri;edgewoodensis Alstrup & M.S.Cole:
on Parmelia sulcata;erodens M.S.Christ. & D.Hawksw.:
on various lichens; follmannii S.Y.Kondr. & D.J.Gallo-
way: on Lobaria pulmonaria;laevisporum Kalb &
Hafellner: on Haematomma;lecanorae (Jaap)
D.Hawksw.: on various lichens; lichenicola (P.Karst.)
Petr.&Syd.:onPhyscia;plectocarpoides S.Y.Kondr. &
D.J.Galloway: on Pseudocyphellaria;pyxidatae (Oudem.)
Petr.&Syd.:onCladonia;reichlingii Diederich: on
Ramalina fraxinea;usneae (Anzi) D.Hawksw.: on
various lichens; xanthoriae M.S.Christ.: on various
lichens.
Lit.: Alstrup & Cole 1998; Christiansen 1956, 1980; Cole &
Hawksworth 2004; Diederich 1986; Hawksworth 1977b, 1981; Kalb
et al. 1995; Kondratyuk & Galloway 1995b; Kondratyuk et al. 1994;
Lawrey et al. 2011.
Order Acrospermales Minter, Peredo & A.T.Watson
Family Acrospermaceae Fuckel
Gonatophragmium Deighton; type: G. mori (Sawada) Deighton.
Mol.: yes. Asexual, hyphomycetous.
Species: 1-0-0 (8). lichenophilum F.Berger & U.Braun: on
Xanthoria parietina.
Lit.: Berger et al. 2015.
Order Asterotexiales Firmino, O.L.Pereira & Crous
Liu et al. (2017) considered Asterotexiales to be a
younger synonym of Asterinales D.Hawksw.& O.E.Er-
ikss. However, the genus Asterina appeared to be
polyphyletic in a phylogenetic analysis (Hyde et al.
2016). While most Asterina species clustered with
Asterotexiales, sequences of two species, including the
generic type, belonged to a distinct, non-related lineage.
Hyde et al. (2016) concluded that sequences of the
generic type may be from a contaminant and that
Asterotexiales should consequently be synonymized
under Asterinales, a view followed by Liu et al. (2017).
However, this synonymy can only be established after
proving that those type sequences represent a contam-
inant, and obtaining new, reliable sequences of the type
of Asterina.
Order Asterotexiales genera incertae sedis
Buelliella Fink; type: B. minimula (lectotypified by Hafellner
1979). Mol.: yes (T). Currently polyphyletic.
Species: 12-0-0. colombiana Etayo: on Phlyctis andensis;
dirinariae Diederich: on Dirinaria picta;eximia Kalb &
Hafellner: on Pyxine cocoes;heppiae van den Boom: on
Heppia arenacea;inops (Triebel & Rambold) Hafellner: on
Caloplaca s.l.; lecanorae Suija & Alstrup: on Lecanora;
minimula (Tuck.) Fink: on Pertusaria;physciicola Poelt &
Hafellner: on Phaeophyscia and Physcia;poetschii Hafell-
ner: on Endocarpon;protoparmeliopseos Etayo & P´
erez-
Ortega: on Protoparmeliopsis;pusilla Hafellner: on Brig-
antiaea leucoxantha;trypethelii (Tuck.) Hafellner: on
Trypethelium.
Lit.: Aptroot et al. 1997; Ertz & Diederich 2015; Etayo 2002a;
Hafellner 1979, 1985b; Hafellner et al. 2008; Kalb 1990; P ´
erez-
Ortega & Etayo 2010; Suija & Alstrup 2004; van den Boom
2010.
Hemigrapha (M¨
ull.Arg.) D.Hawksw.; type: H. asteriscus (lecto-
typified by Ertz & Diederich 2015). Mol.: yes (T).
Species: 8-0-0. asteriscus (M ¨
ull.Arg.) D.Hawksw.: on
Peltigera;atlantica Diederich & Wedin: on Sticta canar-
iensis;nephromatis Wedin & Diederich: on Nephroma
australe;phaeospora Matzer & L¨
ucking: on foliicolous
Strigula phyllogena;pilocarpacearum M.Ca
´ceres &
L¨
ucking: on Byssoloma and Fellhanera;pseudocyphellariae
Diederich & Wedin: on Pseudocyphellaria argyracea;
strigulae Matzer: on foliicolous Strigula;tenellula
(M¨
ull.Arg.) Matzer: on foliicolous Porina.
Lit.: Ca
´ceres & L¨
ucking 2000; Diederich & Wedin 2000; Eriksson et
al. 2001; Ertz & Diederich 2015; Hawksworth 1975; Matzer 1996.
Karschia K¨
orb.; type: K. talcophila (lectotypified by Hafellner
1979). Mol.: yes (T).
Species: 3-0-0 (4). alpicolae D.Hawksw. & Alstrup: on
Allantoparmelia alpicola;santessonii Hafellner: on Porpi-
dia;talcophila (Ach.) K¨
orb.: on Diploschistes.
Lit.: Alstrup & Hawksworth 1990; Ertz & Diederich 2015; Hafellner
1979; Santesson 1994a.
Labrocarpon Etayo & P´
erez-Ortega; type: L. canariense. Mol.:
yes (T, L).
Species: 1-0-0. canariense (D.Hawksw.) Etayo & P´
erez-
Ortega: on Pertusaria.
Lit.: Ertz & Diederich 2015; Hawksworth 1982b; P´
erez-Ortega &
Etayo 2010.
Melanographa M¨
ull.Arg.; type: M. tribuloides (lectotypified by
Ertz & Diederich 2015). Mol.: no.
Species: 1-0-0. tribuloides (Tuck.) M¨
ull.Arg.: on Anthra-
cothecium and Pyrenula.
Lit.: Ertz & Diederich 2015.
Melaspilea Nyl. s.l.; type: M. arthonioides Nyl. (lichenized).
Mol.: yes (T, L). The type belongs to Melaspileaceae
Diederich et al.: 2018 classification and checklist of lichenicolous fungi 353
(Eremithallales). All lichenicolous species need transfer to
other genera, probably within Asterotexiales.
Species: 10-0-0 (150). amarkantakensis S.Joseph & G.P.Si-
nha: on Pertusaria amarkantakana;cupularis M¨
ull.Arg.: on
Pyrenula;epigena M¨
ull.Arg.: on Reimnitzia santensis;
epigraphella (Nyl.) M¨
ull.Arg.: on Acanthothecis consocians;
galligena Zhurb. & I.Zhdanov.: on Pertusaria;insitiva
Stirt.: on Pertusaria leioplaca;leciographoides Vouaux: on
Verrucaria;lekae Brackel & Kalb: on Sarcographa;nitido-
chapsae Pushpi Singh, Y.Joshi & Kr.P.Singh: on Nitido-
chapsa leprieurii;tucumana Flakus, Etayo & Kukwa: on
Pertusaria.
Lit.: Clauzade et al. 1989; Ertz & Diederich 2015; Flakus et al. 2014;
Joseph & Sinha 2015; Kalb et al. 2012; Singh et al. 2017; Zhurbenko
& Zhdanov 2013.
Melaspileopsis (M¨
ull.Arg.) Ertz & Diederich; type: M. diplasio-
spora (Nyl.) Ertz & Diederich (saprotrophic). Mol.: yes (T, L).
Species: 1-0-0. cf. diplasiospora (Nyl.) Ertz & Diederich: on
Graphis elegans.
Lit.: Ertz & Diederich 2015.
Stictographa Mudd; type: S. lentiginosa. Mol.: yes (T).
Species: 2-0-0. dirinariicola Diederich & Ertz: on Dirinaria
picta;lentiginosa (Leight.) M¨
ull.Arg.: on Phaeographis
dendritica.
Lit.: Ertz & Diederich 2015; Diederich et al. 2017.
Taeniolella S.Hughes s.l.; type: T. exilis (P.Karst.) S.Hughes
(saprotrophic). Mol.: yes (T, L). The type belongs to
Kirschsteiniotheliaceae (Dothideomycetes). Most lichen-
icolous species need transfer to other genera within
Asterotexiales.
Species: 29-0-1 (45). arctoparmeliae Heuchert & Zhurb.: on
Arctoparmelia separata;arthoniae (M.S.Christ. &
D.Hawksw.) Heuchert & U.Braun [”Cladosporium arthoniae
M.S.Christ. & D.Hawksw.]: on Dendrographa decolorans,
Lecanactis abietina,Lecanographa lyncea and Pachnolepia
pruinata;atricerebrina Hafellner: on Tephromela atra;
breviuscula (Berk. & M.A.Curtis) S.Hughes: facultatively
lichenicolous; caespitosa M.S.Cole & D.Hawksw.: on Pertu-
saria;christiansenii Alstrup & D.Hawksw.: on Stereocaulon
(incl. on Arthonia lichenicolous on Stereocaulon); chryso-
thricis Diederich: on Chrysothrix candelaris;cladinicola
Alstrup: on Cladonia subgen. Cladina and on the C. uncialis
group; delicata M.S.Christ. & D.Hawksw.: on various lichens;
diederichiana Etayo & Calat.: on Placopsis;friesii (Hepp)
Hafellner: on Strigula stigmatella;hawksworthiana Heuchert,
Ertz & Common: on Phaeographis;ionaspisicola Alstrup &
E.S.Hansen: on Ionaspis odora;lecanoricola Heuchert &
Diederich: on Lecanora rupicola;pertusariicola D.Hawksw. &
H.Mayrhofer: on Lecanora,Lepra,Ochrolechia,Ophioparma,
Pertusaria and Varicellaria;phaeophysciae D.Hawksw.:
mainly on Phaeophyscia,alsoonHyperphyscia,Pectenia,
Pertusaria,Physcia and Physconia;pseudocyphellariae Etayo:
on Nephroma antarcticum and Pseudocyphellaria hillii;
punctata M.S.Christ. & D.Hawksw.: on Graphis,morerarely
on Arthonia,Fissurina,Pertusaria and Phaeographis;pyrenu-
lae Heuchert & Diederich: on Pyrenula;rolfii Diederich &
Zhurb.: on Cetraria;santessonii Etayo & Heuchert: on
Roccellina cerebriformis;serusiauxii Diederich: on Dendrog-
rapha decolorans,Tylophoron and other sterile crustose
lichens with Trentepohlia;strictae Alstrup: on Cladonia
stricta;thelotrematis Heuchert & Brackel: on Thelotrema;
toruloides Heuchert & Diederich: on Thelotrema;trapeliop-
seos Diederich: on Trapeliopsis;umbilicariae Heuchert &
Etayo: on Umbilicaria;umbilicariicola Heuchert & Etayo: on
Umbilicaria;verrucosa M.S.Christ. & D.Hawksw.: on Aspic-
ilia,Micarea,Pachnolepia and Rhizocarpon;weberi Heuchert
&Sparrius:onThelotrema weberi.
Lit.: Alstrup 1993b; Alstrup & Hansen 2001; Alstrup & Hawks-
worth 1990; Cole & Hawksworth 2001; Diederich 1989, 1990,
1992b; Diederich & Zhurbenko 1997, 2001; Ertz et al. 2016; Etayo
2010b; Etayo & Calatayud 2005; Etayo & Sancho 2008; Hafellner
1998b, 2007; Hawksworth 1979a; Heuchert et al. 2018; Zhurbenko
& Alstrup 2004.
Order Botryosphaeriales C.L.Schoch, Crous &
Shoemaker
Family Phyllostictaceae Fr.
Phyllosticta Pers.; type: P. convallariae Pers. Mol.: yes. Asexual,
pycnidial.
Species: 1-0-0 (100). galligena F.Moreau: on Parmotrema
[a poorly known species that might not belong here].
Lit.: Moreau 1951.
Order Capnodiales Woron.
Family Cladosporiaceae Nann.
Cladosporium Link; type: C. herbarum (Pers.) Link. Mol.: yes
(T, L).
Species: 1-0-2 (300). antarcticum K.Schub., Crous &
U.Braun: isolated from Gondwania regalis, probably lichen-
icolous; licheniphilum Heuchert & U.Braun: on macro-
lichens; macrocarpum Preuss: facultatively lichenicolous
on Cetraria islandica and Xanthoria parietina.
Lit.: Bensch et al. 2012; Brackel 2014; Heuchert & Braun 2006;
Zhurbenko 2012.
Verrucocladosporium K.Schub., Aptroot & Crous; type: V.
dirinae. Mol.: yes (T).
Species: 1-0-1. dirinae K.Schub., Aptroot & Crous: isolated
from Dirina massiliensis, probably lichenicolous; roccellae
U.Braun, Diederich & Heuchert: on Roccella.
Lit.: Braun et al. 2009; Crous et al. 2007.
Family Dissoconiaceae Crous & de Hoog
Ramichloridium de Hoog; type: R. apiculatum (J.H.Mill.,
Giddens & A.A.Foster) de Hoog. Mol.: yes (T).
Species: 2-0-0 (35). cladoniicola U.Braun & Heuchert: on
Cladonia portentosa; tropicum U.Braun & Diederich: on
crustose lichen with Trentepohlia.
Lit.: Braun et al. 2009; Diederich et al. 2017.
Family Mycosphaerellaceae Lindau
Pseudocercospora Speg.; type: P. vitis (L´
ev.) Speg. Mol.: yes (T).
Species: 1-0-0 (1500). lichenum (Keissl.) D.Hawksw.: on
various lichens.
Lit.: Etayo 2002; Hawksworth 1979a.
354 The Bryologist 121(3): 2018
Sphaerulina Sacc. s.l.; type: S. myriadea (DC.) Sacc. Mol.: yes
(T). The two lichenicolous species are poorly known and
might not belong here.
Species: 2-0-0 (65). dubiella (Nyl.) W.Wats.: on a crustose
lichen;lepidiotae(Anzi)Vain.: on Fuscopannari a praetermissa.
Lit.: Clauzade et al. 1989.
Wernerella Nav.-Ros.,Cl.Roux & Giralt; type: W. maheui. Mol.: no.
Species: 2-0-0. acarosporae Etayo: on Acarospora nodulosa;
maheui (Werner) Nav.-Ros., Cl.Roux & Giralt: on Rinodina.
Lit.: Etayo 2010c; Navarro-Rosin´
es et al. 1996a, 1998d.
Family Teratosphaeriaeae Crous & U.Braun
Austrostigmidium P´
erez-Ortega & Garrido-Benavent; type: A.
mastodiae. Mol.: yes (T, L).
Species: 1-0-0. mastodiae P´
erez-Ortega & Garrido-Bena-
vent: on Mastodia tessellata.
Lit.: P´
erez-Ortega et al. 2015.
Hobsoniopsis D.Hawksw.; type: H. santessonii. Mol.: yes (T, L).
Species: 1-0-0. santessonii (Lowen & D.Hawksw.)
D.Hawksw.: on Peltigera, mainly P. scabrosa.
Lit.: Lowen et al. 1986; Sikaroodi et al. 2001.
Xanthoriicola D.Hawksw.; type: X. physciae. Mol.: yes (T, L).
Species: 1-0-0. physciae (Kalchbr.) D.Hawksw.: on Xan-
thoria parietina.
Lit.: Hawksworth 1979a; Hawksworth & Punithalingam 1973;
Ruibal et al. 2011.
Order Collemopsidiales P´
erez-Ort., Garrido-Ben. &
Grube
Family Xanthopyreniaceae Zahlbr.
Collemopsidium Nyl.; type: C. iocarpum (Nyl.) Nyl. (liche-
nized). Mol.: yes.
Species: 1-0-0 (27). cephalodiorum (Triebel & Grube)
Grube [”Cercidospora cephalodiorum Triebel & Grube]: on
Pilophorus cephalodiorum.
Lit.: Grube 2005.
Didymellopsis (Sacc.) Clem. & Shear; type: D. latitans. Mol.: no.
Species: 8-0-0. Pharcidia collematis Dughi: on Lempho-
lemma chalazanum [a poorly known taxon placed
provisionally here]; collematum (J.Steiner) Grube &
Hafellner: on Collema;latitans (Nyl.) Clem. & Shear: on
Lichinella and Lempholemma;nephromatis Zhurb. &
Etayo: on Nephroma cellulosum;perigena (Nyl.) Grube
& Hafellner: on Placidium squamulosum;pulposi (Zopf)
Grube & Hafellner: on Collema,Enchylium,Lathagrium,
Lempholemma and Scytinium;Verrucaria symbiotica Nyl.
[¼?Didymella pulposi var. garovaglii Vouaux; ¼?Lichen-
ochora hypanica S.Y.Kondr., L.L}
ok¨
os & J.-S.Hur]: on
Endocarpon [poorly known taxa in need of a revision,
placed provisionally here]; viridireagens Etayo: on
Leptogium.
Lit.: Etayo 2017; Grube & Hafellner 1990; Roux 2014; Zhurbenko et
al. 2015b.
Zwackhiomyces Grube & Hafellner; type: Z. coepulonus. Mol.:
yes (T, L).
Species: 35-0-0. arenicola R.C.Harris: on unidentified
areolate, saxicolous thallus; argentinae (R¨
as¨
anen)
D.Hawksw. & V.Atienza: on Acarospora;aspiciliae Halıcı
& Candan: on Aspicilia;berengerianus (Arnold) Grube &
Triebel: on Lecidea berengeriana;calcariae (Flagey)
Hafellner & Nik.Hoffm.: on Aspicilia;calcisedus Cl.Roux:
on Verrucariaceae;cervinae Calat., Triebel & P ´
erez-
Ortega: on Acarospora;cladoniae (C.W.Dodge) Diederich:
on Cladonia;coepulonus (Norman) Grube & R.Sant.: on
Calogaya,Caloplaca s.str. and Rusavskia elegans;die-
derichii D.Hawksw. & Iturr.: on Cladonia;dispersus
(K¨
orb.) Triebel & Grube: on Protoblastenia rupestris;
echinulatus Brackel: on Physconia;euplocinus Hafellner,
Grube & Egan: on Speerschneidera euploca;heppiae van
den Boom: on Heppia arenacea;inconspicuus Grube &
Hafellner: on Myriolecis dispersa;kantvilasii S.Y.Kondr.:
on Parmotrema;kiszkianus D.Hawksw. & Mia
˛dl.: on
Peltigera;lacustris (Arnold) Orange: on Ionaspis lacustris;
lecanorae (Stein) Nik.Hoffm. & Hafellner: on Myriolecis;
lithoiceae (B. de Lesd.) Hafellner & V.John: on Verru-
caria;macrosporus Alstrup & Olech: on Protopannaria;
martinatianus (Arnold) Triebel & Grube: on Porpidia;
melanohaleae Etayo & van den Boom: on Melanohalea
exasperata;namibiensis Diederich & M.Schultz: on
Psorotichia;peltigerae Mia
˛dl. & Alstrup: on Peltigera;
physciicola Alstrup: on Physcia caesia;polischukii Dar-
mostuk & Khodos.: on Bacidia;rolfii (Etayo) Etayo: on
Candelina;sipmanii Diederich & Zhurb.: on Phaeorrhiza
sareptana;socialis (K¨
orb.) Cl.Roux [¼Z. immersae
(Arnold) Grube & Triebel]: on Clauzadea, perhaps also
on Bacidia bagliettoana;solenopsorae van den Boom: on
Solenopsora holophaea;sphinctriniformis Grube & Ha-
fellner: on Romjularia lurida;sphinctrinoides (Zwackh)
Grube & Hafellner: on Lecanora campestris;sulcatus
P´
erez-Ortega & Etayo: on Austroparmelina macrospora;
turcicus Kocakaya, Halıcı& Aksoy: on Physcia magnusso-
nii.
Lit.: Alstrup 1993a; Alstrup & Olech 1993; Aptroot et al. 1997;
Diederich & Zhurbenko 2009; Etayo 2010b; Grube & Hafellner
1990; Halıcı& Candan 2009; Harris 1995; Hawksworth &
Atienza 1994; Hawksworth & Iturriaga 2006; Hawksworth &
Mia
˛dlikowska 1997a; Hoffmann & Hafellner 2000; Kocakaya et
al. 2011; Khodosovtsev & Darmostuk 2017; Kondratyuk 1996b;
Mia
˛dlikowska & Alstrup 1995; Orange 2002; P ´
erez-Ortega et al.
2011a; Roux 2014; van den Boom 2010; van den Boom & Etayo
2014.
Order Dothideales Lindau
Family Dothideaceae Chevall.
Plowrightia Sacc.; type: P. ribesia (Pers.) Sacc. Mol.: yes.
Species: 1-0-0 (50). mereschkowskyi Vouaux: on Aspicilia
hispida [a poorly known species].
Lit.: Clauzade et al. 1989.
Family Saccotheciaceae Bonord.
Pseudoseptoria Speg.; type: P. donacicola Speg. Mol.: yes.
Species: 1-0-0 (8). usneae (Vouaux) D.Hawksw.: on Usnea.
Lit.: Hawksworth 1981.
Diederich et al.: 2018 classification and checklist of lichenicolous fungi 355
Order Lichenotheliales K.Knudsen, Muggia &
K.D.Hyde
Family Lichenotheliaceae Henssen
Lichenothelia D.Hawksw.; type: L. scopularia (Nyl.) D.Hawksw.
(saxicolous, non-lichenized). Mol.: yes (L).
Species: 3-0-3 (30). convexa Henssen: over siliceous rocks,
facultatively lichenicolous; renobalesiana D.Hawksw. &
V.Atienza: on endolithic Verrucariaceae (Bagliettoa,Theli-
dium,Verrucaria) over calcareous rocks; rugosa (G.Thor)
Ertz & Diederich [”Lichenostigma rugosum G.Thor, as
‘rugosa’]: on Diploschistes;scopularia (Nyl.) D.Hawksw.:
over siliceous rocks, facultatively lichenicolous; spiralis-
pora Etayo [as ‘spiratispora’]: on Acarospora sphaerospora;
umbrophila Muggia, Kocourk. & K.Knudsen: over rocks,
facultatively lichenicolous.
Lit.: Atienza & Hawksworth 2008; Etayo 2010c; Ertz et al. 2014b;
Kocourkova
´& Knudsen 2011b; Muggia et al. 2015b.
Lichenostigma Hafellner s.l. [incl. Lichenostigma subgen.
Lichenogramma Nav.-Ros. & Hafellner; type: L. elongatum;
see under Lichenostigmatales for Lichenostigma s.str.].
Mol.: yes (L). All species treated here have been described
in Lichenostigma, but certainly do not belong there. They
obviously belong to Lichenotheliaceae (L.cf.elongatum
and several unidentified specimens have been sequenced
byErtzetal.2014),butthegenericaffinitiesareyet
unknown.
Species: 25-0-1. amplum Calat. & Hafellner: on Buellia;
anatolicum Halıcı& Kocakaya: on brown Acarospora;
bolacinae Nav.-Ros., Calat. & Hafellner: on Polycauliona
bolacina;canariense Etayo & van den Boom: on Aspicilia
calcarea;cosmopolites Hafellner & Calat.: on Xanthopar-
melia;dimelaenae Calat. & Hafellner: on Dimelaena
oreina;diploiciae Calat.,Nav.-Ros.&Hafellner:on
Diploicia subcanescens;elongatum Nav.-Ros. & Hafellner
[as ‘elongata’]: on Aspicilia and Lobothallia;epipolinum
Nav.-Ros., Calat. & Hafellner [as ‘epipolina’]: on Dip-
lotomma epipolium;epiporpidiae S.Y.Kondr., L.L}
ok¨
os &
J.-S.Hur: on Porpidia;epirupestre P´
erez-Ortega & Calat.:
on Pertusaria pertusa;episulphurella Etayo & van den
Boom: on Lecanora sulphurella;gracile Calat., Nav.-Ros.
& Hafellner: on Acarospora fuscata;heterodermiae
S.Y.Kondr., L.L}
ok¨
os et J.-S.Hur: on Heterodermia;
iranicum Brackel & Valadbeigi: on Lobothallia praeradio-
sa;lecanorae Calat. & Nav.-Ros.: on Lecanora farinacea;
radicans Calat. & Barreno: on Aspicilia;rouxii Nav.-Ros.,
Calat. & Hafellner: on Squamarina;rupicolae Fern.-Brime
&Nav.-Ros.:onPertusaria rupicola;saxicola K.Knudsen
& Kocourk.: saxicolous, facultatively lichenicolous on
Polysporina simplex;semi-immersum Hafellner: on Buel-
lia;subradians Hafellner, Calat. & Nav.-Ros.: on Acar-
ospora;supertegentis Ihlen & R.Sant.: on Aspicilia
supertegens;svandae Vondra
´k&ˇ
Soun: on Acarospora
cervina;triseptatum Halıcı&D.Hawksw.:onAspicilia;
verrucosum Brackel & Valadbeigi: on Pleopsidium go-
biense.
Lit.: Alstrup & Hawksworth 1990; Calatayud & Barreno 2003;
Calatayud & Navarro-Rosin´
es 2000; Calatayud et al. 2002,
2004b; Etayo & Sancho 2008; Ferna
´ndez-Brime et al. 2010;
Hafellner 1982a, 1999, Hafellner & Calatayud 1999; Halıcı&
Hawksworth 2007; Halıcıet al. 2009d; Harris & Lendemer
2009; Ihlen 2004; Kondratyuk et al. 2013, 2017; Knudsen &
Kocourkova
´2010b; Navarro-Rosin´
es & Hafellner 1996; P´
erez-
Ortega & Calatayud 2009; Thor 1985; van den Boom &
Etayo 2006; Valadbeigi & Brackel 2011; Vondra
´k&ˇ
Soun
2007.
Order Patellariales D.Hawksw. & O.E.Erikss.
Family Patellariaceae Corda
Stratisporella Hafellner; type: S. episemoides. Mol.: no.
Species: 1-0-0. episemoides (Nyl.) Hafellner: on Enter-
ographa.
Lit.: Aptroot et al. 1997; Hafellner 1979; Kutorga & Hawksworth
1997.
Tryblidaria (Sacc.) Rehm; type: T. fenestrata (Cooke & Peck)
M.E.Barr. Mol.: no. The lichenicolous species is poorly
known.
Species: 1-0-0 (9). capensis (Stizenb.) Vouaux: on Buellia
callaina.
Lit.: Clauzade et al. 1989.
Order Pleosporales M.E.Barr.
Family Dacampiaceae K¨
orb.
Dacampia A.Massal.; type: D. hookeri. Mol.: yes (T).
Species: 14-1-0. caloplacicola Halıcı, Candan & Etayo:
on Blastenia crenularia;cladoniicola Halıcı&A.
¨
O.T¨
urk:
on Cladonia;cyrtellae Brackel: on Lecania cyrtella;
engeliana (Saut.) A.Massal.: on Solorina;hookeri
(Borrer) A.Massal.: lichenicolous lichen on Solorina;
lecaniae Kocourk. & K.Knudsen: on Lecania fuscella;
leptogiicola (D.Hawksw.) D.Hawksw.: on Leptogium;
muralicola Halıcı&D.Hawksw.:onProtoparmeliopsis
muralis;peltigericola D.Hawksw. & C.J.B.Hitch: on
Peltigera;pentaseptata Etayo: on Parmotrema;rhizocar-
picola D.Hawksw.: on Rhizocarpon;rubra Halıcı,
Candan & Calat.: on Aspicilia;rufescentis (Vouaux)
D.Hawksw.: on Peltigera;thamnoliicola Zhurb. ad int.:
on Thamnolia;xanthomendozae Etayo & Halıcı:on
Xanthomendoza.
Lit.: Brackel 2010a, 2010b; Earland-Bennett et al. 2006; Ertz et al.
2015; Etayo 2017; Halıcı& Hawksworth 2008; Halıcıet al. 2008,
2009a, 2009b; Hawksworth 1980b, 1986; Henssen 1995; Khodo-
sovtsev & Darmostuk 2016; Kocourkova
´& Knudsen 2010b;
Zhurbenko 2012.
Family Didymellaceae Gruyter
Ascochyta Lib.; type: A. pisi Lib. Mol.: yes (T).
Species: 2-0-0 (400). candelariellicola D.Hawksw. & Kalb:
on Candelariella aurella;santessonii Alstrup & D.Hawksw.:
on Phaeophyscia sciastra.
Lit.: Alstrup & Hawksworth 1990; Hawksworth & Kalb 1992.
Epicoccum Link; type: E. nigrum. Mol.: yes (T, L).
Species: 0-0-1 (10). nigrum Link: facultatively lichenicolous
over various lichens.
Lit.: David & Hawksworth 1995; Diederich 2004b.
356 The Bryologist 121(3): 2018
Leptosphaerulina McAlpine; type: L. australis McAlpine. Mol.:
yes (T).
Species: 1-0-0 (30). peltigerae (Fuckel) Riedl: on Peltigera.
Lit.: Hawksworth 1980b.
Phoma Sacc. s.l.; type: P. herbarum Westend. Mol.: yes (T).
Asexual, pycnidial. Most lichenicolous Phoma species from
which molecular data are available belong to Didymocyrtis
(Phaeosphaeriaceae, Pleosporales), and some more pertain
to that genus on the basis of morphological characters. The
remaining species, of unknown phylogenetic affinities, are
enumerated here.
Species: 10-0-0 (100). aggregata Calat. & Etayo: on
Diploschistes ocellatus;biformis Vouaux: on Anisomeridium
biforme [a poorly known taxon]; dubia (Linds.) Sacc. &
A.Trotter: on Usnea;everniae D.Hawksw.: on Evernia
prunastri;fistulata Etayo & Osorio: on Parmotrema
fistulatum [does not belong to Didymocyrtis]; lecanorina
Diederich: on Lecanora;maculiformans Ihlen: on Dibaeis
baeomyces;melanohaleicola D.Hawksw. & Earl.-Benn.: on
Melanohalea exasperata;pisutii S.Y.Kondr., A.Lackoviˇ
cova
´,
E.Lisicka
´& A.Guttova
´:onXanthomendoza ulophyllodes;
recepii Halıcı& Candan: on Caloplaca s.str.
Lit.: Brackel 2011a; Calatayud & Etayo 2001; Diederich 1986;
Earland-Bennett et al. 2006; Etayo & Osorio (2004); Hafellner &
Wieser 2000; Halıcıet al. 2014; Hawksworth 1980b, 1981, 1994;
Ihlen 1998; Mart´
ınez & Hafellner 1998; Oksner 2010.
Family Didymosphaeriaceae Munk
Sporidesmiella P.M.Kirk; type: S. claviformis P.M.Kirk. Mol.:
yes. Asexual, hyphomycetous.
Species: 2-0-0 (32). lichenophila U.Braun & Heuchert: on
Ramalina stenospora; physconiicola Zhurb., U.Braun &
Kobzeva: on Physconia distorta.
Lit.: Braun & Heuchert 2010; Zhurbenko et al. 2015d.
Family Leptosphaeriaceae M.E.Barr
Leptosphaeria Ces. & De Not.; L. doliolum (Pers.) Ces. & De
Not. Mol.: yes (T).
Species: 3-0-0 (150). clarkii D.Hawksw.: on Peltigera;
oxneriae Cl.Roux & S.Y.Kondr.: on Xanthomendoza ulo-
phyllodes;pycnostigma (Nyl.) Sacc. & D.Sacc.: on Baeomyces.
Lit.: Clauzade et al. 1989; Hawksworth 1980b; Kondratyuk et al.
2017.
Family Massarinaceae Munk
Stagonospora (Sacc.) Sacc.; type: S. paludosa (Sacc. & Speg.)
Sacc. Mol.: yes (T). Asexual, pycnidial.
Species: 1-0-0 (220). exasperatulae Brackel: on Melanoha-
lea exasperatula.
Lit.: Brackel 2009.
Family Microsphaeropsidaceae Q.Chen, L.Cai &
Crous
Microsphaeropsis H¨
ohn.; type: M. olivacea (Bonord.) H¨
ohn.
Mol.: yes (T).
Species: 3-0-0 (37). caloplacae Etayo & Yazici: on Calogaya;
lichenicola Etayo: on Pannaria;physciae Brackel: on
Physcia.
Lit.: Brackel 2014; Etayo & Sancho 2008; Etayo & Yazici 2009.
Family Periconiaceae Nann.
Periconia Tode; type: P. lichenoides Tode. Mol.: yes (L). Asexual,
hyphomycetous.
Species: 0-0-1 (45). digitata (Cooke) Sacc.: facultatively
lichenicolous on Cladonia,Peltigera and Xanthoria parie-
tina.
Lit.: Brackel 2014.
Family Phaeosphaeriaceae M.E.Barr
Didymocyrtis Vain.; type: D. consimilis (lectotypification by Ertz
et al. 2015) [¼Diederichia D.Hawksw.; type: D. pseude-
verniae;¼Diederichomyces Crous & Trakunyingcharoen;
type: D. xanthomendozae]. Mol.: yes (T). Several Phoma
and one Leptosphaeria species obviously belong here, but no
new combinations are proposed for some, as they might
represent synonyms of other species.
Species: 25-0-0 (26). bryonthae (Arnold) Hafellner [¼?
Phoma denigricans Hafellner]: on Lecanora epibryon;canar-
iensis van den Boom & Etayo: on Ramalina subgeniculata;
Phoma candelariellae Z.Kocakaya & Halıcı:onCandelariella
aurella [similar to D. consimilis]; cladoniicola (Diederich,
Kocourk. & Etayo) Ertz & Diederich [”Ph. cladoniicola
Diederich, Kocourk. & Etayo]: on Cladonia,Flavoparmelia,
Parmelina,Ramalina and Squamarina;consimilis Vain. [¼
Ph. caloplacae D.Hawksw.]: on Caloplaca s.str., possibly also
on Cladonia,Heterodermia and Melanohalea;epiphyscia
Ertz & Diederich [”Ph. physciicola Keissler; ¼?Ph. epiphyscia
Vouaux, described from Xanthoria and Phaeophyscia,
neotype on Phaeophyscia sciastra]: on Physcia aipolia,
probably also on Physcia adscendens,P. stellaris,P. tenella
and Xanthoria parietina;foliaceiphila (Diederich, Kocourk.
& Etayo) Ertz & Diederich [”Ph. foliaceiphila Diederich,
Kocourk. & Etayo]: on Cladonia,Parmelia and Pseudevernia;
Phoma fuliginosa M.S.Cole & D.Hawksw.: on Xanthomen-
doza trachyphylla [similar to D. consimilis]; graphidacearum
van den Boom & Ertz: on Pallidogramme;grumantiana
(Zhurb. & Diederich) Zhurb. & Diederich, comb. nov. [”
Ph. grumantiana Zhurb. & Diederich]: on Cladonia;Phoma
heterodermiae S.Y.Kondr., L.L}
ok¨
os & J.-S.Hur: on Hetero-
dermia [similar to D. consimilis]; infestans (Speg.) Hafellner:
on Teloschistes chrysophthalmus and T. flavicans;kaernefeltii
(S.Y.Kondr.) Hafellner [”Polycoccum kaernefeltii
S.Y.Kondr.]: on Teloschistes chrysophthalmus;melanelixiae
(Brackel) Diederich, Harris & Etayo [”Ph. melanelixiae
Brackel]: on Canoparmelia,Cetraria,Cetrelia,Evernia,
Hypotrachyna,Melanelixia,Parmelia,Parmotrema,Platisma-
tia,Pseudevernia,Punctelia and Usnea;micropunctum
Etayo: on Parmotrema;Phoma peltigerae (P.Karst.)
D.Hawksw.: on Peltigera;Leptosphaeria protousneae Etayo:
on Protousnea [similar to D. ramalinae]; physciae (Brackel)
Hafellner [”Merismatium physciae Brackel; ”Pleospora
physciae (Brackel) Hafellner & E.Zimm.]: on Physcia;
pseudeverniae (Etayo & Diederich) Ertz & Diederich [”
Diederichia pseudeverniae (Etayo & Diederich) D.Hawksw.;
”Macrophomina pseudeverniae Etayo & Diederich]: on
Pseudevernia;ramalinae (Desm.) Ertz, Diederich & Hafellner
Diederich et al.: 2018 classification and checklist of lichenicolous fungi 357
[¼Ph. ficuzzae Brackel; ¼?Pyrenidium ucrainicum
S.Y.Kondr., L.L}
ok¨
os & J.-S.Hur]: on Ramalina;Phoma
rozziana Etayo: on Pseudocyphellaria [similar to D. clado-
niicola]; slaptoniensis (D.Hawksw.) Hafellner & Ertz [”
Polycoccum slaptoniense D.Hawksw.]: on Xanthoria parietina;
trassii Suija, Darmostuk & Khodos.: on Cetraria aculeata;
xanthomendozae (Diederich & Freebury) Diederich &
Freebury [”Diederichomyces xanthomendozae (Diederich &
Freebury) Crous & Trakunyingcharoen; ”Ph. xanthomen-
dozae Diederich & Freebury]: on Xanthomendoza;Phoma
thamnoliae Zhurb. ad int.: on Thamnolia.
Lit.: Brackel 2008, 2010a; Diederich et al. 2007; Etayo 1996, 2017;
Etayo & Diederich 1996a; Etayo & Sancho 2008; Hafellner 1994a,
2015; Hafellner & Zimmermann 2012; Hawksworth 1981, 1994,
2003; Hawksworth & Cole 2004; Khodosovtsev et al. 2018;
Kocakaya et al. 2015; Kondratyuk 2008; Kondratyuk et al. 2012,
2013; Lawrey et al. 2012; van den Boom et al. 2017; Zhurbenko
2012.
Ophiobolus Riess; type: O. disseminans Riess. Mol.: yes (T).
Both lichenicolous species probably do not belong here.
Species: 2-0-0 (350). aspiciliae (H.Olivier) Vouaux: on
Aspicilia;thallicola (Ces. & De Not.) Sacc.: on Peltigera
[often considered a synonym of Neolamya peltigerae, but
following Brackel 2014 possibly a distinct species].
Lit.: Brackel 2014; Clauzade et al. 1989.
Phaeoseptoria Speg.; type: P. papayae Speg. [¼Phaeosphaeria
I.Miyake; type: P. orizae I.Miyake]. Mol.: yes (T).
Species: 1-0-0 (90). peltigerae Punith. & Spooner: on
Peltigera.
Lit.: Punithalingam & Spooner 1997.
Xenophoma Crous & Trakunyingcharoen; type: X. puncteliae.
Mol.: yes (T, L).
Species: 1-0-0. puncteliae (Diederich & Lawrey) Crous &
Trakunyingcharoen [”Phoma puncteliae Diederich &
Lawrey]: on Punctelia.
Lit.: Lawrey et al. 2012; Trakunyingcharoen et al. 2014.
Family Pleomassariaceae M.E.Barr
Lichenopyrenis Calat., Sanz & Aptroot; type L. galligena. Mol.:
no.
Species: 1-0-0. galligena Calat., Sanz & Aptroot: on
Leptochidium albociliatum.
Lit.: Calatayud et al. 2001b.
Family Pleosporaceae Nitschke
Pleospora Ces. & De Not.; type: P. herbarum (Pers.) Ces. & De
Not. Mol.: yes (T).
Species: 6-0-0 (65). bernadetiae van den Boom [as
Plaeospora bernadetae]: on Protoparmeliopsis muralis;col-
lematum Zukal: on Collemataceae; Verrucaria congestula
Cromb.: on Aspiciliopsis macrophthalma [a taxon of
uncertain position, provisionally placed here]; crozalsii
Vouaux: on Sticta sylvatica;tretiachii Hafellner [¼P.
aquatica Tretiach & Nimis, non Griffiths]: on Aspicilia;
xanthoriae Khodos. & Darmostuk: on Xanthoria parietina.
Lit.: Clauzade et al. 1989; Khodosovtsev & Darmostuk 2016;
Tretiach & Nimis 1999; van den Boom 2015.
Family Pyrenochaetopsidaceae Valenzuela-L ´
opez,
Crous, Cano, Guarro & Stchigel
Pyrenochaetopsis Gruyter; type: P. leptospora (Sacc. & Briard)
Gruyter. Mol.: yes (T, L).
Species: 0-0-1 (7). microspora (Gruyter & Boerema) Gruyter,
Aveskamp & Verkley: facultatively lichenicolous on Buellia.
Lit.: de Gruyter et al. 2010.
Family Sporormiaceae Munk
Preussia Fuckel [a hemihomonym of Preussia Karsch, Orthop-
tera]; type: P. funiculata (Preuss) Fuckel [¼Sporormiella Ellis
& Everh.; type: S. nigropurpurea Ellis & Everh.]. Mol.: yes (T).
Species: 1-0-0 (50). peltigerae (Brackel) Diederich, comb.
nov. [”Sporormiella peltigerae Brackel]: on Peltigera rufescens.
Lit.: Brackel 2009.
Family Trematosphaeriaceae K.D.Hyde, Y.Zhang
ter, Suetrong & E.B.G.Jones
Trematosphaeria Fuckel; type: T. pertusa (Pers.) Fuckel. Mol.:
yes (T).
Species: 1-0-0 (20). lophiostoma M.Choisy & Werner: on
Staurothele [a poorly known species].
Lit.: Clauzade et al. 1989.
Order Pleosporales genera incertae sedis
Berkleasmium Zobel; type: B. cordeanum Zobel. Mol.: yes.
Species: 1-0-0 (36). parmeliellae Etayo & Diederich: on
Parmeliella testacea.
Lit.: Etayo & Diederich 1995.
Ellisembia Subram.; type: E. coronata (Fuckel) Subram. Mol.: yes.
Species: 1-0-0 (60). lichenicola Heuchert & U.Braun: on
various lichens.
Lit.: Heuchert & Braun 2006.
Pseudopyrenidium Nav.-Ros., Zhurb. & Cl.Roux; type: P.
tartaricola. Mol.: no.
Species: 1-0-0. tartaricola (Linds.) Nav.-Ros., Zhurb. &
Cl.Roux [”Weddellomyces tartaricola (Linds.) Alstrup &
D.Hawksw.]: on Ochrolechia.
Lit.: Alstrup & Hawksworth 1990; Navarro-Rosin´
es et al. 2010.
Pyrenochaeta De Not.; type: P. nobilis De Not. Mol.: yes (T).
Species: 2-0-0 (5). collematis Vouaux: on Enchylium tenax
[not a Pyrenochaeta, fide Roux 2014]; xanthoriae Die-
derich: on Xanthoria.
Lit.: Clauzade et al. 1989; Diederich 1990; Hafellner 1982a.
Order Strigulales L¨
ucking, M.P.Nelsen & K.D.Hyde
Family Strigulaceae Zahlbr.
Strigula Fr.; type: S. smaragdula Fr. (lichenized). Mol.: yes (T).
Species: 1-0-0 (70). dichosporidii Etayo: on Dichospori-
dium nigrocinctum.
Lit.: Etayo 2002.
358 The Bryologist 121(3): 2018
Order Trypetheliales L¨
ucking, Aptroot & Sipman
Family Polycoccaceae Ertz, Hafellner & Diederich
Clypeococcum D.Hawksw.; type: C. cladonema. Mol.: yes (L).
Species: 12-0-0. amylaceum Etayo: on Parmotrema;bispo-
rum Zhurb.: on Cetraria and Nephromopsis;cajasense
Etayo: on Hypotrachyna;cetrariae Hafellner: on Cetraria
and Nephromopsis;cladonema (Wedd.) D.Hawksw.: on
Cetrelia and Xanthoparmelia;epimelanostolum
(D.Hawksw. & Øvstedal) Grube & Hafellner: on Buellia
melanostola;galloides Etayo: on Lecidea s.l.; grossum
(K¨
orb.) D.Hawksw.: on Umbilicaria;hypocenomycis
D.Hawksw.: on Hypocenomyce;placopsiiphilum Øvstedal
& D.Hawksw.: on Placopsis;psoromatis (A.Massal.) Etayo
[¼C. epicrassum (H.Olivier) Hafellner & Nav.-Ros.]: on
Squamarina;rugosisporum Etayo & Zhurb.: on Parmo-
trema.
Lit.: Ertz et al. 2015; Etayo 2010b, 2017; Etayo & Triebel
2010; Grube & Hafellner 1990; Hafellner 1994a, 1995a;
Hawksworth 1977a, 1980a, 1982a; Navarro-Rosin´
es et al.
1994; Øvstedal & Hawksworth 1986; Zhurbenko 2009a;
Zhurbenko et al. 2016.
Polycoccum K¨
orb.; type: P. sauteri, i.e. P. trypethelioides. Mol.:
yes (T, L).
Species: 60-0-0. acarosporicola Halıcı&D.Hawksw.:on
Acarospora cervina;aksoyi Halıcı& V.Atienza: on Aspic-
ilia;alboatrae (Vouaux) Etayo: on Diplotomma venus-
tum;amygdalariae F.Berger & Triebel: on Amygdalaria;
anatolicum Halıcı&E.Akg
¨
ul: on Lepraria incana;
arnoldii (Hepp) D.Hawksw.: on Diploschistes and
Rhizocarpon;atrostriatae van den Boom: on Physcia
atrostriata;cartilaginosum (Arnold) D.Hawksw.: on an
unidentified saxicolous lichen?; cladoniae Diederich &
D.Hawksw.: on Cladonia;clauderouxii S.Y.Kondr.,
L.L}
ok¨
os & J.-S.Hur: on Zeroviella laxa;clauzadei Nav.-
Ros. & Cl.Roux: on Rusavskia elegans;crassum Vˇ
ezda: on
Peltigera;crespoae Va
´czi & D.Hawksw.: on Xanthopar-
melia;decolorans Calat. & Triebel: on Immersaria;
deformans R.Sant. & Brackel: on Placopsis;dictyonema-
tis Etayo & R.Sant.: on Cora;diederichianum Nav.-Ros.,
nom. inval. (Art. 30.8): on Placynthium;dzieduszyckii
(Boberski) D.Hawksw.: on Bagliettoa;epiphyllum Mat-
zer: on foliicolous Trichothelium;epizoharyi Calat. &
V.Atienza: on Buellia zoharyi;evae Calat. & V.J.Rico: on
Dimelaena;follmannii (C.W.Dodge) Alstrup: on Lecidea
cremoricolor;heterodermiae Calat.: on Heterodermia;
hymeniicola (Berk. & Broome) Zhurb. [”Endococcus
hymeniicola (Berk. & Broome) D.Hawksw.]: on Lobaria
[generic position uncertain]; ibericum Etayo & van den
Boom: on Rinodina;innatum (M ¨
ull.Arg.) D.Hawksw.: on
an unidentified corticolous lichen; islandicum Brackel &
F.Berger: on Placopsis gelida;jamesii D.Hawksw.: on
Psoromidium;kerneri J.Steiner: on Lecidea fuscoatra;
laursenii Zhurb.: on Cladonia;longisporum Etayo: on
Pseudocyphellaria;marmoratum (Kremp.) D.Hawksw.:
on crustose lichens on calcareous rocks (Clauzadea,
Polyblastia,Thelidium,Verrucaria); microcarpum Die-
derich & Etayo: on Cladonia;microstictum (Leight.)
Arnold: on Acarospora,alsoreportedfromBuellia,
Ionaspis and Rhizocarpon;minutulum Kocourk. &
F.Berger: on Trapelia placodioides;montis-wilhelmii
Diederich: on Hypotrachyna;nigrosporum Etayo: on
Buellia aethalea;ochvarianum Y.Joshi: on Ochrolechia,
Pertusaria and Varicellaria;opulentum (Th.Fr. & Almq.)
Arnold: on Polyblastia;peltigerae (Fuckel) V ˇ
ezda: on
Peltigera;perrugosae Brackel: on Placopsis perrugosa;
psorae Zhurb. & Triebel: on Anamylopsora pulcherrima;
pulvinatum (Eitner) R.Sant.: on Physcia;rinodinae van
den Boom: on Rinodina;rubellianae Calat. & V.Atienza:
on Caloplaca rubelliana;rugulosarium (Linds.)
D.Hawksw.: on Austroplaca cirrochrooides and Gondwa-
nia;serusiauxii Matzer: on foliicolous Byssolecania;
sporastatiae Arnold: on Sporastatia;squamarioides
(Mudd) Arnold: on Placopsis;stellulatae (Vouaux)
Hafellner: on Buellia stellulata;stictaria (Linds.) D.J.Gal-
loway: on Pseudocyphellaria;superficiale D.Hawksw. &
Mia
˛dl.: on Peltigera;teresum Halıcı& K.Knudsen: on
Xanthoria;thallicola Arnold: on cf. Aspicilia candida;
tinantii Diederich: on Blastenia ferruginea;trypethe-
lioides (Th.Fr.) R.Sant. [¼P. sauteri K¨
orb., fide
Santesson 1960]: on Stereocaulon;umbilicariae (Linds.)
D.Hawksw.: on Lasallia;ventosicola Zhurb.: on Ophio-
parma ventosa;vermicularium (Linds.) D.Hawksw.: on
Thamnolia;versisporum (Bagl. & Carestia) D.Hawksw.:
on Rhizocarpon geographicum and an unidentified
saxicolous lichen.
Lit.: Alstrup 2002; Aptroot el al.1997; Atienza & Hawksworth 2008;
Atienza et al. 2003; Berger 2000; Brackel & Berger 2010; Calatayud
2004; Calatayud & Atienza 2000; Calatayud & Rambold 1998;
Calatayud & Rico 1995; Clauzade et al. 1989; Diederich 1990; Ertz
et al. 2015; Etayo 2010b, 2010c; Etayo & Diederich 1998; Etayo &
Sancho 2008; Galloway 2004; Gaya & Navarro-Rosin´
es 2008;
Hafellner 2015; Halıcıet al. 2007a, 2009c, 2013; Hawksworth
1985b; Hawksworth & Diederich 1988; Hawksworth & Mia
˛dli-
kowska 1997a; Joshi et al. 2017a; Kocourkova
´& Berger 1999;
Kondratyuk et al. 2016b; Lord et al. 2013; Matzer 1996; Navarro-
Rosin´
es 1992; Navarro-Rosin´
es & Roux 1990, 1998; Triebel 1989;
Va
´czi & Hawksworth 2001; van den Boom 2010, 2012; van den
Boom & Etayo 2014; Zhurbenko 2007b, 2010b; Zhurbenko &
Dillman 2010.
Order Tubeufiales Boonmee & K.D.Hyde
Family Tubeufiaceae M.E.Barr
Lichenotubeufia Etayo; type: L. eriodermae. Mol.: no.
Species: 5-0-0. boomiana Etayo: on Sticta;eriodermae
(Etayo) Etayo: on Erioderma;heterodermiae (Etayo) Etayo:
on Heterodermia and Physcia;pannariae (Etayo) Etayo: on
Pannaria rubiginosa;tafallae Etayo: on Leptogium.
Lit.: Etayo 2002a, 2002b, 2017.
Order Venturiales Yin.Zhang & K.D.Hyde
Family Venturiaceae M.E.Barr.
Spilodochium Syd.; type: S. vernoniae Syd. Mol.: no. Asexual,
sporodochial.
Species: 1-0-0 (4). parmotrematis Diederich & Etayo: on
Parmotrema.
Lit.: Etayo & Diederich 1995.
Order Venturiales genus incertae sedis
Fusicladium Bonord.; type: F. virescens Bonord. (saprotrophic).
Mol.: yes (L).
Diederich et al.: 2018 classification and checklist of lichenicolous fungi 359
Species: 0-0-1 (75). peltigericola Crous & Diederich:
isolated from Peltigera, probably lichenicolous.
Lit.: Crous et al. 2011.
Class Dothideomycetes familiae incertae sedis
Family Corynesporascaceae Sivan.
Corynespora G¨
ussow; type: C. mazei G¨
ussow. Mol.: yes.
Asexual, hyphomycetous.
Species: 2-0-0 (130). laevistipitata (M.S.Cole &
D.Hawksw.) Heuchert & U.Braun [”Taeniolella laevisti-
pitata M.S.Cole & D.Hawksw.]: on Graphis,Lecanora,
Lepra,Pertusaria,Phaeophyscia and Xanthoria;thorii
U.Braun, Zhurb. & Frisch: on Lecanora.
Lit.: Heuchert & Braun 2006; Zhurbenko et al. 2015d.
Family Naetrocymbaceae R.C.Harris
Leptorhaphis K¨
orb.; type: L. oxyspora (Nyl.) K¨
orb. Mol.: no.
Species: 1-0-0 (15). haematommatum Hafellner & Kalb: on
Haematomma.
Lit.: Kalb et al. 1995.
Family Pseudoperisporiaceae Toro
Keratosphaera H.P.B.Upadhyay; type: K. batistae [¼Sporhaplus
H.B.P.Upadhyay; type: S. rondoniensis, i.e. K. batistae;¼
Sphaeromma H.B.P.Upadhyay; type: S. mazosiae, i.e. K.
batistae]. Mol.: no.
Species: 5-0-0 (6). antoniana Flakus, Farkas & L¨
ucking: on
foliicolous Trichothelium;batistae H.B.P.Upadhyay [¼
Sphaeromma mazosiae H.B.P.Upadhyay; ¼Sporhaplus
rondoniensis H.B.P.Upadhyay]: on foliicolous Mazosia;
dimerellae Matzer: on foliicolous Coenogonium;furcatiseta
Matzer [¼Sphaeromma furcatiseta Matzer]: on foliicolous
Porina;porinae Matzer [¼Sphaeromma porinae Matzer]:
on foliicolous Porina.
Lit.: Farkas 2010; Matzer 1996.
Myxophora D¨
obbeler & Poelt; type: M. amerospora D¨
obbeler &
Poelt. Mol.: no.
Species: 5-0-0 (7). apotheciicola Nik.Hoffm. & Hafellner:
on Leptogium azureum;leptogiophila (G.Winter) Ni-
k.Hoffm. & Hafellner: on Collema and Scytinium plicatile;
ovalispora Nik.Hoffm. & Hafellner: on Collema;placyn-
thii Nik.Hoffm. & Hafellner: on Placynthium caesium;
tetraspora Nik.Hoffm. & Hafellner: on Collema flaccidum.
Lit.: Hoffmann & Hafellner 2000.
Neocoleroa Petr.; type: N. sibirica Petr. Mol.: no.
Species: 3-0-0 (6). inundata (Vain.) Diederich: on
Bacidina;lichenicola (Hansf.) M.E.Barr. subsp. lichenicola
[”Wentiomyces lichenicola (Hansf.) D.Hawksw.]: on
foliicolous lichens; lichenicola subsp. bouteillei (Bricaud,
Cl.Roux & S´
erus.) Barr: on Fellhanera bouteillei,Byssoloma
and possibly Bacidina.
Lit.: Barr 1997; Hawksworth 1980b; Matzer 1996; Roux et al. 1994;
S´
erusiaux et al. 1999.
Family Trichothyriaceae Theiss.
Lichenopeltella H¨
ohn.; type: L. maculans [¼Micropeltopsis
Vain.; type: M. cetrariicola;”Microthyris Clem.; type: M.
maculans;¼Trichothyrina (Petr.) Petr.; type: T. alpestris
(Sacc.) Petr.]. Mol.: no.
Species: 39-0-0 (48). arctomiae P´
erez-Ortega & T.Sprib.:
on Arctomia delicatula;biatorae P´
erez-Ortega & T.Sprib.:
on Biatora flavopunctata;bunodophoronis Diederich: on
Bunodophoron;cetrariae (Bres.) H¨
ohn.: on Cetraria;
cetrariicola (Nyl.) R.Sant. [”Micropeltopsis cetrariicola
(Nyl.) Vain.]: on Cetraria;cladoniarum E.S.Hansen &
Alstrup: on Cladonia subgen. Cladina;communis Etayo: on
Hypotrachyna;coppinsii Earl.-Benn. & D.Hawksw.: on
Verrucaria;epiphylla R.Sant.: on foliicolous Mazosia and
Porina;heppiae van den Boom: on Heppia;heterodermiae
Diederich: on Heterodermia;heterodermiicola M.S.Cole &
D.Hawksw. subsp. heterodermiicola:onHeterodermia;
heterodermiicola subsp. endothallina Etayo: on Hetero-
dermia;hydrophila R.Sant.: on Verrucaria;hypogymniae
Diederich: on Hypogymnia zeylanica;hypotrachynae Die-
derich: on Hypotrachyna;leprosulae P´
erez-Ortega &
T.Sprib.: on Myochroidea leprosula;leptogii Diederich: on
Leptogium;lobariae Etayo & Diederich: on Lobaria;
maculans (Zopf) H¨
ohn. [”Microthyris maculans (Zopf)
Clem.]: on Umbilicaria;megalosporae Diederich: on
Megalospora;microspora Diederich: on Parmeliella;min-
uta R.Sant.: on Agonimia foliacea;pannariacearum Die-
derich: on Pannariaceae; peltigericola (D.Hawksw.)
R.Sant.: on Peltigera;physciae Diederich: on Physcia;
pseudocyphellariae Diederich: on Pseudocyphellaria;ram-
alinae Etayo & Diederich: on Ramalina;rangiferinae
Brackel: on Cladonia;santessonii (P.M.Kirk & Spooner)
R.Sant.: on Peltigera;sclerenchymatica Etayo: on Pseudo-
cyphellaria;setifera Matzer: on foliicolous Bacidia;soiliae
Zhurb. & Pino-Bodas: on Cladia;stereocaulorum Zhurb.:
on Stereocaulon;swaminathaniana Harih., Mibey &
D.Hawksw.: on Porina;thalamica Etayo, Flakus & Kukwa:
on Pseudocyphellaria;thamnoliae R.Sant.: on Thamnolia;
thelidii Diederich: on Thelidium;uncialicola Brackel: on
Cladonia uncialis.
Lit.: Aptroot et al. 1997; Brackel 2010c, 2011b; Cole & Hawksworth
2002; Earland-Bennett & Hawksworth 1999b; Etayo 2010b, 2017;
Etayo & Diederich 1996b; Etayo & Sancho 2008; Hansen & Alstrup
1995; Hariharan et al. 1996; Matzer 1996; Molitor & Diederich
1997; P´
erez-Ortega & Spribille 2009; Santesson 1988, 1989, 1998,
2001; van den Boom & Giralt 2012; Zhurbenko 2010a; Zhurbenko
& Pino-Bodas 2015.
Macrographa Etayo; type: M. antarctica. Mol.: no.
Species: 1-0-0. antarctica Etayo: on Nephroma antarcticum.
Lit.: Etayo & Sancho 2008.
Class Dothideomycetes genera incertae sedis
Ampullifera Deighton; type: A. foliicola [¼Ampulliferella Bat. &
Cavalc.; type: A. amoeboides;¼Ampulliferopsis Bat. &
Cavalc.; type: A. myriapoda, i.e. A. amoeboides;¼Terato-
schaeta Bat. & O.M.Fonseca; type: T. rondoniensis, i.e. A.
amoeboides]. Mol.: no.
Species: 6-0-0. amoeboides (Bat. & Cavalc.) D.Hawksw. [”
Ampulliferella amoeboides Bat. & Cavalc.; ¼Ampulliferopsis
myriapoda Bat. & Cavalc.; ¼Teratoschaeta rondoniensis Bat.
& O.M.Fonseca;]: on foliicolous Mazosia and other
360 The Bryologist 121(3): 2018
Roccellaceae; foliicola Deight.: on foliicolous lichens;
hippocrateacearum (Bat. & Caval.) D.Hawksw. [¼A.
brasiliensis Deight.]: on foliicolous Mazosia and other
lichens; leonensis Deight.: on foliicolous lichens; pirozyn-
skii D.Hawksw.: on foliicolous lichens; ugandensis Deight.:
on foliicolous lichens.
Lit.: Hawksworth 1979; Matzer 1996.
Cercidospora K¨
orb.; type: C. ulothii, i.e. C. macrospora. Mol.:
no.
Species: 40-0-0 (100). alpina Ihlen & Wedin: on Stereo-
caulon cumulatum;anomala Etayo: on Pertusaria pertusa;
apiosporoides (Vouaux) Nav.-Ros., Hafellner & Calat. ad
int. [”Didymella epipolytropa var. apiosporoides Vouaux]:
on Aspicilia;barrenoana Calat. & Nav.-Ros.: on Rhizo-
placa peltata;caudata Kernst.: on Xanthocarpia lactea and
other Teloschistaceae; cecidiiformans Grube & Hafellner:
on Rhizocarpon;cladoniicola Alstrup: on Cladonia;
crozalsiana (H.Olivier) Nav.-Ros., Cl.Roux & Casares:
on Squamarina;decolorella (Nyl.) O.Eriksson & Yue: on
terricolous algae, mosses and lichens; epicallopisma
Arnold: on Variospora aurantia;epicarphinea (Nyl.)
Grube & Hafellner: on Usnochroma;epidesertorum
Nav.-Ros. & Calat. ad int.: on Gyalolechia desertorum;
epipolytropa (Mudd) Arnold: on Lecanora;epithamnolia
Zhurb.: on Thamnolia;etayoana Nav.-Ros. & Calat. ad
int.: on Physcia caesia;exiguella (Nyl.) Arnold: on
Rinodina mniaraea;galligena Hafellner & Nav.-Ros.: on
Aspicilia;hypotrachynicola Etayo: on Hypotrachyna;
javalambrensis Calat. & Nav.-Ros. ad int.: on Aspicilia
desertorum;lecidomae Zhurb. & Triebel: on Lecidoma
demissum;lobothalliae Nav.-Ros.&Calat.:onLobothallia;
macrospora (Uloth) Hafellner & Nav.-Ros. [¼C. ulothii
K¨
orb.]: on Protoparmeliopsis;melanophthalmae Nav.-
Ros., Calat. & Hafellner: on Rhizoplaca melanophthalma;
mutabilicola Calat.et al. ad int.: on Megaspora verrucosa
var. mutabilis;ochrolechiae Zhurb.: on Ochrolechia and
Pertusaria coriaceae;parva Hafellner & Ihlen: on Baeo-
myces;pluriseptata (Nyl.) Arnold: on crustose saxicolous
lichens; punctillata (Nyl.) R.Sant. [¼C. lichenicola (Zopf)
Hafellner]: on Solorina and Peltigera,reportedfrommany
other hosts; rinodinae Etayo & van den Boom: on
Rinodina conradii;solearispora Calat., Nav.-Ros. &
Hafellner: on Aspicilia;soror Obermayer & Triebel: on
Arthrorhaphis citrinella;stenotropae Nav.-Ros. & Hafell-
ner ad int.: on Lecanora gr. polytropa;stereocaulorum
(Arnold) Hafellner: on Stereocaulon;thamnogalloides
Zhurb.: on Thamnolia;thamnoliae Zhurb.: on Thamnolia;
thamnoliicola Ihlen: on Thamnolia;trypetheliza (Nyl.)
Hafellner & Obermayer [¼C. arthrorhaphidicola Alstrup]:
on Arthrorhaphis alpina;verrucosaria (Linds.) Arnold: on
Megaspora verrucosa;werneri Nav.-Ros., Calat. & Hafell-
ner: on Aspicilia;xanthoriae (Wedd.) R.Sant.: on
Xanthoria.
Lit.: Alstrup 1997; Alstrup et al. 1994; Calatayud et al. 2013; Clauzade
et al. 1989; Etayo 2010a, 2017; Etayo & van den Boom 2005;
Hafellner 1987b, 1993; Hafellner & Obermayer 1995; Ihlen 1995,
1998; Ihlen & Wedin 2007; Navarro-Rosin´
es et al. 1995, 1996b, 2004,
2009a; Triebel 1989; Triebel & Scholz 2001; Zhurbenko 2010b, 2012;
Zhurbenko & Triebel 2003; Zhurbenko et al. 1995.
Echinothecium Zopf; type: E. reticulatum. Mol.: no.
Species: 2-0-0. aerophilum Alstrup & M.S.Cole: on
Alectoria;reticulatum Zopf [”Sphaerellothecium reticula-
tum (Zopf) Etayo]: on Parmelia, doubtfully reported from
other parmelioid hosts.
Lit.: Alstrup & Cole 1998; Clauzade et al. 1989; Roux & Triebel
1994; Zhurbenko & Pino-Bodas 2015.
Homostegia Fuckel; type: H. adusta,i.e.H. piggottii. Mol.: no. A
synonym of Pyrenidium (fide Navarro-Rosin´
es et al. in prep.).
Species: 2-0-0. hertelii D.Hawksw., V.Atienza & M.S.Cole:
on Flavoparmelia caperata;piggottii (Berk. & Broome)
P.Karst. [¼H. adusta Fuckel; ¼Phaeospora fritzei Stein]: on
Parmelia s.str.
Lit.: Hawksworth et al. 2004.
Koordersiella H¨
ohn.; type: K. javanica H¨
ohn. [¼Ascohansfor-
diellopsis D.Hawksw.; type: A. deightonii;¼Hansfordiellopsis
Deight.; type: H. aburiensis Deight.]. Mol.: no.
Species: 6-0-0. deightonii (D.Hawksw.) D.Hawksw. &
O.E.Erikss. [”Ascohansfordiellopsis deightonii D.Hawksw.;
¼Hansfordiellopsis elongata D.Hawksw.]: on foliicolous
Porina and ?Gomphillaceae; insectivora (Hansf.)
D.Hawksw. & O.E.Erikss. [”A. insectivora (Hansf.)
D.Hawksw.; ¼H. aburiensis Deight.; ¼H. lichenicola (Bat.
& Maya) Deight.; ¼H. deightonii Bat. & Herr.]: on
foliicolous Asterothyrium,Calenia,Gyalectidium and Tri-
charia;javanica H¨
ohn.: on Gomphillaceae; Hansfordiel-
lopsis minuta D.Hawksw.: on foliicolous lichens;
tenuissima (D.Hawksw.) D.Hawksw. [¼H. tenuissima
D.Hawksw.]: on foliicolous lichens; variegata (D.Hawksw.)
D.Hawksw. [¼H. variegata D.Hawksw.]: on foliicolous
lichens, incl. Bullatina aspidota.
Lit.: Eriksson & Hawksworth 1987; Hawksworth 1979a; Matzer 1996.
Lanatosphaera Matzer; type: L. porinicola. Mol.: no.
Species: 2-0-0. dimerellae Matzer: on foliicolous Coenogo-
nium subluteum;porinicola Matzer: on foliicolous Porina
epiphylla.
Lit.: Matzer 1996.
Monodictys S.Hughes; type: M. putredinis (Wallr.) S.Hughes.
Mol.: yes.
Species: 5-0-0 (50). anaptychiae (Lindau) D.D.Hawksw.:
on Anaptychia;cellulosa S.Hughes [M. lepraria (Berk.)
M.B.Ellis]: on various crustose, corticolous lichens; epi-
lepraria Kukwa & Diederich: on Lepraria;fuliginosa Etayo:
on Lobaria and Ricasolia;phlyctidis R.C.Harris: on Phlyctis
speirea.
Lit.: Etayo & Diederich 1996b; Hawksworth 1975, 1979a; Kukwa &
Diederich 2005; Muscavitch et al. 2017.
Pseudonitschkia Coppins & S.Y.Kondr.; type: P. parmotrematis.
Mol.: no.
Species: 1-0-0. parmotrematis Coppins & S.Y.Kondr.: on
Parmotrema.
Lit.: Coppins & Kondratyuk 1995.
Pseudorobillarda M.Morelet; type: P. phragmitis (Cunnell) Nag
Raj, Morgan-Jones & W.B.Kendr. Mol.: yes (T).
Species: 1-0-0 (12). peltigerae Diederich: on Peltigera,
rarely on Cladonia.
Lit.: van den Boom et al. 1998.
Pyrenidium Nyl.; type: P. actinellum [¼Dacampiosphaeria
D.Hawksw.; type: D. rivana]. Mol.: no. Several species are
provisionally placed here, without being formally com-
bined, awaiting the taxonomic revision of the genus.
Species: 13-0-0. actinellum Nyl.: on Scytinium teretiuscu-
lum;aggregatum Knudsen & Kocourk.: on Phaeophyscia
Diederich et al.: 2018 classification and checklist of lichenicolous fungi 361
and Physcia;coccineum Aptroot: on Siphula decumbens;
crozalsii (Vouaux) Nav.-Ros., comb. inval. (Art. 30.8) [”
Leptosphaeria crozalsii Vouaux]: on Aspicilia gr. calcarea;
cryptotheciae Matzer: on foliicolous Myriostigma candida;
Phaeospora granulosae Arnold: on Trapeliopsis granulosa;
hetairizans (Leight.) D.Hawksw.: on Staurothele;Leptos-
phaeria leucomelaria (Mudd) Vouaux: on Leucodermia
leucomelos;octosporum Looman:onanunidentified
lichen; Dacampiosphaeria rivana (De Not.) D.Hawksw.:
on Peltigera;santessonii L¨
ucking: on Bacidia psychotriae
and Bapalmuia;sporopodiorum Matzer: on foliicolous
Sporopodium;zamiae (M¨
ull.Arg.) Matzer: on foliicolous
Porina.
Lit.: Aptroot 2014; Looman 1962; L¨
ucking 1998; Matzer 1996;
Navarro-Rosin´
es 1992; Roux 2014.
Rhabdospora (Sacc.) Sacc.; type: Septoria oleandri Durieu &
Mont. Mol.: no. A poorly known genus.
Species: 1-0-0 (150). haematommatum Kalb & Hafellner:
on Haematomma.
Lit.: Kalb et al. 1995.
Rosellinula R.Sant.; type: R. haplospora. Mol.: no.
Species: 4-0-0. frustulosae (Vouaux) R.Sant.: on Lecanora
frustulosa and L. argopholis;haplospora (Th.Fr.) R.Sant.: on
Aspicilia and Bellemerea;kalbii (Hafellner) Hafellner &
R.W.Rogers: on Haematomma puniceum;lopadii (Vouaux)
D.J.Galloway: on Brigantiaea.
Lit.: Galloway 2004; Hafellner 1985a, 2004.
Sphaerellothecium Zopf; type: S. araneosum. Mol.: no.
Species: 35-0-0. abditum Triebel: on Immersaria and Lecidea;
aculeatae Khodos., Gavrylenko & Klymenko: on Cetraria
aculeata;araneosum (Arnold) Zopf [¼Echinothecium
glabrum M.S.Christ., Alstrup & D.Hawksw.]: on Ochrolechia,
Lepra and Varicellaria;atryneae (Arnold) Cl.Roux & Triebel:
on Lecanora bicincta,L. cenisia,L. rupicola and L. swartzii;
breussii K.Knudsen, Kocourk. & Etayo: on Placidium
lacinulatum;buelliae (C.W.Dodge) D.Hawksw. & Iturr.:
on Buellia;cinerascens Etayo & Diederich: on Cladonia
parasitica;cladoniae (Alstrup & Zhurb.) Hafellner [”S.
araneosum var. cladoniae Alstrup & Zhurb.]: on Cladonia
subgen. Cladonia and Stereocaulon;cladoniicola E.S.Hansen
& Alstrup: on Cladonia subgen. Cladina;conioides (Nyl.)
Cl.Roux & Diederich: on Baeomyces;contextum Triebel: on
Calvitimela,Protoparmelia,Rhizocarpon,Rhizoplaca and
Sporastatia;epilecanora Zhurb.: on Lecanora epibryon;
episoralium Etayo: on soralia of Heterodermia;episquamar-
inae Etayo: on Squamarina lentigera;gallowayi Diederich:
on Heterodermia;giraltiae van den Boom: on Rinodina;
gowardii Alstrup & M.S.Cole: on Acarospora schleicheri;
heterodermiae van den Boom: on Heterodermia;icmado-
philae (R.Sant.) Zhurb.: on Icmadophila ericetorum;leratia-
num Gardiennet & Cl.Roux: on Brodoa;minutum Hafellner:
on Sphaerophorus;pannariacearum Etayo: on Pannaria;
parietinarium (Linds.) Hafellner & V.John: on Xanthoria;
parmeliae Diederich & Etayo: on Parmelia;parmotrematis
van den Boom [as ‘parmotremae’]: on Parmotrema;phaeor-
rhizae Diederich & Zhurb.: on Phaeorrhiza sareptana;
propinquellum (Nyl.) Cl.Roux & Triebel: on Lecanora
carpinea group; pumilum (Lettau) Nav.-Ros., Cl.Roux &
Hafellner [”Stigmidium pumilum (Lettau) Matzer &
Hafellner; ¼Sphaerellothecium aipolium Nav.-Ros. &
Cl.Roux; ¼?Thelidium parvum C.W.Dodge & G.E.Baker]:
on Physcia;siphulae Zhurb.: on Siphula ceratites;soechtingii
Zhurb. & Alstrup: on Arthrorhaphis alpina;stereocaulorum
Zhurb.: on Stereocaulon;subtile Triebel & Rambold: on
Teloschistes;thamnoliae Zhurb. var. thamnoliae:onTham-
nolia;thamnoliae Zhurb. var. taimyricum Zhurb.: on
Thamnolia;usneicola Etayo: on Usnea.
Lit.: Alstrup & Cole 1998; Diederich 2007b; Diederich &
Zhurbenko 2009; Etayo 2008, 2017; Etayo & Diederich 1998; Etayo
& Sancho 2008; Gardiennet & Roux 2013; Hafellner 1993; Hansen
& Alstrup 1995; Hawksworth & Iturriaga 2006; Khodosovtsev et al.
2016; Knudsen et al. 2009b; Navarro-Rosin´
es & Roux 2017;
Navarro-Rosin´
es et al. 2018; Roux & Triebel 1994; Triebel 1989;
Triebel et al. 1991; van den Boom 2010, 2016; Zhurbenko 2007,
2008a, 2015, 2012, 2015; Zhurbenko & Alstrup 2004; Zhurbenko &
Brackel 2013; Zhurbenko & Triebel 2008.
Stromatopogon Zahlbr.; type: S. baldwinii. Mol.: no.
Species: 3-0-0. baldwinii Zahlbr.: on Usnea;cladoniae
Diederich & S´
erus.: on Cladonia;geminatum Etayo &
Breuss: on Usnea.
Lit.: Diederich 1992a; Diederich & S´
erusiaux 2003; Etayo & Breuss
2004.
Trematosphaeriopsis Elenkin; type: T. parmeliana. Mol.: no.
Species: 1-0-0. parmeliana (Jacz.) Elenkin: on Xanthopar-
melia.
Lit.: Hafellner 2001.
Weddellomyces D.Hawksw.; type: W. epicallopisma. Mol.: no.
Species: 12-0-0. aspiciicola Alstrup: on Lobothallia;epi-
callopisma (Wedd.) D.Hawksw.: on Protoblastenia,Pyr-
enodesmia and Variospora;erythrocarpae Nav.-Ros. &
Cl.Roux: on Caloplaca erythrocarpa;gasparriniae
(C.W.Dodge) D.Hawksw. & Iturr.: on ?Caloplaca;hetero-
chrous Nav.-Ros. & Cl.Roux: on Aspicilia;macrosporus
D.Hawksw., Renob. & Coppins: on Aspicilia calcarea and
Lobothallia radiosa;pachyosporicola Nav.-Ros. & Cl.Roux:
on Aspicilia;periphericus (Taylor) Alstrup & D.Hawksw.:
on Pertusaria pseudocorallina;pertusariicola Halıcı:on
Varicellaria lactea;protearius Nav.-Ros. & Cl.Roux: on
Leproplaca proteus and Rusavskia elegans;turcicus Halıcı&
Orange: on Acarospora;xanthoparmeliae Calat. & Nav.-
Ros.: on Xanthoparmelia.
Lit.: Alstrup 1992; Calatayud & Navarro-Rosin´
es 1998; Halıcı2010,
Halıcıet al. 2006; Hawksworth 1978a, 1986, 1990a; Hawksworth &
Iturriaga 2006; Navarro-Rosin´
es & Roux 1995, 1997b.
Class EUROTIOMYCETES O.E.Erikss. & Winka
Subclass CHAETOTHYRIOMYCETIDAE Doweld
Order Chaetothyriales M.E.Barr.
Family Epibryaceae S.Stenroos & C.Gueidan
Epibryon D¨
obbeler; type: E. plagiochilae (Gonz.Frag.) D¨
obbeler.
Mol.: yes (T).
Species: 4-0-0 (30). conductrix (Norman) Nik.Hoffm. &
Hafellner [¼Stigmidium catapyrenii Cl.Roux & Triebel]: on
Catapyrenium cinereum and C. daedaleum;lichenicolaMatzer:
on Mazosia;parvipunctum (Stein) Diederich: on Thelidium;
solorinae (Vain.) Nik.Hoffm. & Hafellner: on Solorina.
Lit.: Hoffmann & Hafellner 2000; Matzer 1996; S´
erusiaux et al. 1999.
362 The Bryologist 121(3): 2018
Family Herpotrichiellaceae Munk
Capronia Sacc.; type: C. sexdecimspora (Cooke) Sacc. Mol.: yes.
Species: 21-0-0 (60). amylacea Etayo: on Peltigera;andina
Etayo: on Placopsis gelida;baeomycetis Diederich: on
Baeomyces;epilobarina S.Y.Kondr. & D.J.Galloway: on
Lobaria;etayoi Flakus & Kukwa: on Cora;guatemalensis
Etayo & van den Boom [as ‘guatamalense’]: on Parmo-
trema;hypotrachynae Etayo & Diederich: on Hypotrachyna
and Menegazzia;josefhafellneri Zhurb. & Etayo: on
Septotrapelia triseptata;leopoldiana Etayo: on Leptogium;
leptogii Etayo & Diederich: on Leptogium;magellanica
Etayo: on Fuscopannaria;minutosetosae Halıcı,
D.Hawksw., A.O.T¨
urk & Candan: on Thelenella muscorum;
muellerelloides Etayo: on Heterodermia;normandinae
R.Sant. & D.Hawksw.: on Normandina pulchella;para-
nectrioides Etayo, Flakus & Kukwa: on Erioderma leylandii;
pseudonormandinae Diederich: on Normandina simoden-
sis;santessoniana Etayo: on Cora;solitaria Etayo: on
Heterodermia and perhaps on Lobaria,Lobariella and Sticta;
suijae Tsurykau & Etayo: on Xanthoria parietina;thamno-
liae Zhurb.: on Thamnolia;triseptata (Diederich) Etayo [”
Muellerella triseptata Diederich]: on various lichens.
Lit.: Aptroot el al.1997; Diederich 1986; Etayo 2002a, 2003, 2010b,
2017; Etayo & Diederich 1998; Etayo & Sancho 2008; Etayo & van
den Boom 2006; Etayo et al. 2013b; Flakus & Kukwa 2012; Halıcıet
al. 2010; Hawksworth 1990a; Kondratyuk & Galloway 1995a;
Tsurykau & Etayo 2017; Zhurbenko 2012; Zhurbenko et al. 2016.
Cladophialophora Borelli; type: C. carrionii (Trejos) de Hoog,
Kwon-Chung & McGinnis. Mol.: yes (T, L). Asexual,
hyphomycetous.
Species: 5-0-0 (35). cladoniae (Diederich) Diederich: on
Cladonia;hawksworthii (Etayo & Diederich) Diederich: on
Megalospora tuberculosa;megalosporae Diederich: on
Megalospora pachycheila;normandinae (Diederich &
Etayo) Diederich: on Normandina pulchella [possibly the
asexual stage of Capronia normandinae]; parmeliae (Etayo
& Diederich) Diederich & Unter.: mainly on parmelioid
lichens, also on Lobaria and Pannaria.
Lit.: Diederich 2010; Diederich et al. 2013; Etayo & Diederich 1995,
1996a; Etayo & Sancho 2008.
Family Trichomeriaceae Chomnunti & K.D.Hyde
Knufia L.J.Hutchison & Unter.; type: K. cryptophialidica
L.J.Hutchison & Unter. Mol.: yes (T, L).
Species: 1-0-0 (13). peltigerae (Fuckel) R´
eblova
´& Unter.
[”Capronia peltigerae (Fuckel) D.Hawksw.]: on Peltigera.
Lit.: Hawksworth 1980b; R´
eblova
´et al. 2013; Untereiner et al. 2011.
Order Chaetothyriales genera incertae sedis
Laeviomyces D.Hawksw.; type: L. pertusariicola. Mol.: no.
Asexual, pycnidial.
Species: 1-0-0. pertusariicola (Nyl.) D.Hawksw. [”Lichen-
odiplis pertusariicola (Nyl.) Diederich]: on Pertusaria.
Lit.: Hawksworth 1979a; Zhurbenko & Otte 2012.
Lichenodiplis Dyko & D.Hawksw.; type: L. lecanorae. Mol.: yes
(T, L). Asexual, pycnidial.
Species: 12-0-0. anomala Etayo&P
´
erez-Vargas [as
anomalus]: on Ochrolechia;crespoae P´
erez-Ortega &
V.Atienza: on Calicium notarisii;dendrographae Diederich
& van den Boom: on Dendrographa;fallaciosa (Hafellner &
Kalb) Diederich [possibly the asexual stage of Muellerella
thalamita (Berger & LaGreca 2014)]: on Buellia and
Orcularia;hawksworthii F.Berger & Diederich: on Pertu-
saria;lecanorae (Vouaux)Dyko&D.Hawksw.[¼?
Vouauxiella caloplacae Alstrup] [probably the asexual stage
of Muellerella lichenicola (Muggia et al. 2015a)]: on
Caloplaca s.l., Lecanora s.l. and other crustose lichens;
lecanoricola (M.S.Cole & D.Hawksw.) Diederich: on
Lecanora;lichenicola Dyko & D.Hawksw.: on Rinodina;
ochrolechiae Y.Joshi & M.Tripathi: on Ochrolechia;ope-
graphae (D.Hawksw.) Diederich: on Opegrapha;poeltii
S.Y.Kondr. & D.Hawksw.: on Xanthoria parietina;rinodi-
nicola Kocourk. & K.Knudsen: on Rinodina.
Lit.: Alstrup 1993a; Berger & Diederich 1996; Berger & LaGreca
2014; Cole & Hawksworth 2001; Diederich 2003; Hawksworth
1981; Hawksworth & Dyko 1979; Joshi et al. 2016a; Kalb 1990;
Knudsen & Kocourkova
´2009; Kondratyuk 1996a; Muggia et al.
2015a; P´
erez-Vargas et al. 2013.
Lichenodiplisiella S.Y.Kondr. & Kudratov; type: L. makarevi-
chiae. Mol.: no. Asexual, pycnidial. Morphologically similar
to Lichenodiplis.
Species: 1-0-0. makarevichiae S.Y.Kondr. & Kudratov [as
‘makarevichae’]: on Teloschistes brevior.
Lit.: Kondratyuk et al. 2002.
Minutoexcipula V.Atienza & D.Hawksw.; type: M. tuckerae.
Mol.: no. Asexual, pycnidial-sporodochial.
Species: 7-0-0. beaglei Etayo: on Lecanora;calatayudii
V.Atienza: on Hypogymnia;mariana V.Atienza: on Lepra
and Pertusaria;miniatoexcipula R.C.Harris & Lendemer:
on Pertusaria, mainly P. epixantha;tephromelae V.Atienza,
Etayo & P´
erez-Ortega: on Tephromela;tuckerae V.Atienza
& D.Hawksw.: on corticolous Pertusaria;tuerkii Hafellner:
on Pertusaria glomerata.
Lit.: Atienza 2002; Atienza & Hawksworth 1994; Atienza et al. 2009;
Etayo & Sancho 2008; Hafellner 1994a; Lendemer et al. 2016.
Muellerella M¨
ull.Arg.; type: M. polyspora [”Muellerellomyces
Cif. & Tomas.; type: M. polyspora;¼Pleothelis M.Choisy;
type: P. myriocarpa, i.e. M. pygmaea;¼Polycarpella Theiss.
& Syd.; type: P. cookei, i.e. M. erratica;¼Thelidiola
C.W.Dodge; type: T. eklundii, i.e. Muellerella pygmaea].
Mol.: yes (L). Bryophylous species may not be congeneric.
Species: 12-0-0 (14). antarctica Etayo: on Hypogymnia
antarctica;atricola (Linds.) Sacc. & D.Sacc.: on Tephromela
atra;erratica (A.Massal.) Hafellner & V.John [¼M.
pygmaea var. athallina (M¨
ull.Arg.) Triebel; ¼Polycarpella
cookei (Linds.) Theiss. & Syd.]: on various lichens;
hospitans Stizenb.: on Bacidia fraxinea and B. rubella;
lecanactidis Diederich & van den Boom: on Sigridea
californica;lichenicola (Sommerf.) D.Hawksw. [probably
the sexual stage of Lichenodiplis lecanorae (Muggia et al.
2015a)]: on various lichens; polyspora M¨
ull.Arg. [¼
Muellerellomyces polysporae Cif. & Tomas.]: on diverse
lichens; pygmaea (K¨
orb.) D.Hawksw. [¼Pleothelis myr-
iocarpa M.Choisy; ¼Thelidiola eklundii C.W.Dodge]: on
various lichens; stictinae (H.Olivier) Vouaux: on Sticta;
thalamita (Nyl.) D.Hawksw., F.Berger & LaGreca: on
Baculifer,Orcularia and Hafellia [possibly the sexual stage
of Lichenodiplis fallaciosa (Berger & LaGreca 2014)];
ventosicola (Mudd) D.Hawksw.: on Ophioparma, possibly
Diederich et al.: 2018 classification and checklist of lichenicolous fungi 363
also on Dimelaena,Protoparmelia and Rhizocarpon;vesic-
ularia (Linds.) D.Hawksw.: on Toninia.
Lit.: Berger & LaGreca 2014; Clauzade et al. 1989; Diederich 2003;
Etayo & Sancho 2008; Hawksworth 1985a; Muggia et al. 2015a;
Triebel 1989.
Order Pyrenulales D.Hawksw. & O.E.Erikss.
Family Requienellaceae Boise
Distopyrenis Aptroot; type: D. americana Aptroot. Mol.: no.
Species: 1-0-0 (8). japonica Harada: on Graphis proserpens.
Lit.: Harada 2000.
Lacrymospora Aptroot; type: L. parasitica. Mol.: no.
Species: 1-0-0. parasitica Aptroot: probably lichenicolous
on a sterile corticolous lichen (?Myriotrema).
Lit.: Aptroot 1991.
Parapyrenis Aptroot; type: P. aurora (Zahlbr.) Aptroot. Mol.: no.
Species: 1-0-0 (8). lichenicola Aptroot & Diederich: on
?Pertusaria.
Lit.: Aptroot et al. 1997.
Order Verrucariales D.Hawksw. & O.E.Erikss.
Family Adelococcaceae Triebel
Adelococcus Theiss. & Syd.; type: A. alpestris. Mol.: no.
Species: 4-0-0. alpestris (Zopf)Theiss.&Syd.:on
Acarospora;immersus Etayo & Breuss: on Dermatocarpon
biennense;interlatens (Arnold) Matzer & Hafellner: on
Clauzadea,Hymenelia,Ionaspis,Polysporina and Sarcogyne;
porocyphii S.Zelenko & S.Y.Kondr.: on Porocyphus.
Lit.: Etayo & Breuss 1998; Kondratyuk & Zelenko 2002; Matzer &
Hafellner 1990.
Sagediopsis Vain.; type: S. tartarina, i.e. S. campsteriana. Mol.:
no.
Species: 10-0-0. aquatica (Stein) Triebel: on Koerberiella
wimmeriana;aspiciliae Nik.Hoffm. & Hafellner: on Aspicilia
myrinii;barbara (Th.Fr.) R.Sant. & Triebel: on Porpidia;
bayozturkii Halıcı,G
¨
ull¨
u & Parnikoza: on Acarospora
mycrocyclos;campsteriana (Linds.) D.Hawksw. & R.Sant.
[¼S. tartarina (Nyl.) Vain.]: on Ochrolechia;dissimilis
Triebel: on Paraporpidia leptocarpa;fissurisedens Hafellner:
on Aspicilia myrinii;lomnitzensis (Stein) Orange: on
Ionaspis lacustris and I. odora;pertusariicola Zhurb.: on
Pertusaria;vasilyevae Zhurb.: on Rhizocarpon.
Lit.: Alstrup & Hawksworth 1990; Hafellner 1993; Halıcıet al. 2017;
Hoffmann & Hafellner 2000; Orange 2002; Triebel 1989; Triebel
1993; Zhurbenko 2009c; Zhurbenko & Yakovchenko 2014.
Family Sarcopyreniaceae Nav.-Ros. & Cl.Roux
Sarcopyrenia Nyl.; type: S. gibba [¼Hassea Zahlbr.; type: H.
bacillosa]. Mol.: no.
Species: 11-0-0. acutispora Nav.-Ros. & Cl.Roux: on cf.
Bagliettoa calciseda;bacillosa (Nyl.) Nav.-Ros. & Hladun
[”Hassea bacillosa (Nyl.) Zahlbr.]: on saxicolous lichens;
bacillospora Nav.-Ros. & Hladun: on Thelidium and
Verrucaria;baetica Nav.-Ros. & Hladun: on Lecania cf.
erysibe;beckhausiana (J.Lahm) M.B.Aguirre, Nav.-Ros. &
Hladun: on saxicolous lichens, mainly Verrucariaceae;
cylindrospora (P.Crouan & H.Crouan) M.B.Aguirre:
mainly on Aspicilia;gibba (Nyl.) Nyl. var. gibba:on
saxicolous lichens, incl. Acarospora and Aspicilia;gibba var.
geisleri (Beckh.) Nav.-Ros. & Hladun: on saxicolous
lichens; lichinellae Nav.-Ros. & Hladun: on Lichinella
stipatula;pluriseptata Nav.-Ros. & Cl.Roux: on Pyreno-
desmia variabilis;sigmoideospora Tretiach & Nav.-Ros.: on
Bagliettoa.
Lit.: Knudsen & Lendemer 2006; Navarro-Rosin´
es & Hladun 1990;
Navarro-Rosin´
es & Hladun 2004; Navarro-Rosin´
es et al. 1998b,
2009b; Tretiach & Navarro-Rosin´
es 1996.
Family Verrucariaceae Zenker
Bellemerella Hafellner & Cl.Roux; type: B. trapeliae. Mol.: no.
Species: 4-0-0. acarosporae Calat. & Nav.-Ros.: on
Acarospora fuscata;polysporinae Calat. & Nav.-Ros.: on
Polysporina simplex;ritae P´
erez-Ortega & T.Sprib.: on
Xylographa;trapeliae Nav.-Ros. & Cl.Roux: on Trapelia.
Lit.: Calatayud & Navarro-Rosin´
es 2001; Navarro-Rosin´
es & Roux;
1997a; P´
erez-Ortega & Spribille 2007.
Clauzadella Nav.-Ros. & Cl.Roux; type: C. gordensis. Mol.: no.
Species: 1-0-0. gordensis Nav.-Ros. & Cl.Roux: on
Verrucaria viridula.
Lit.: Navarro-Rosin´
es & Roux 1996.
Endococcus Nyl.; type: E. rugulosus [¼Discothecium Zopf; type:
D. stigma]. Mol.: yes (L). Several poorly known species are
provisionally placed here, without being formally combined
in this genus.
Species: 44-0-0. alectoriae (D.Hawksw.) D.Hawksw.: on
Alectoria;apiciicola (J.Steiner) R.Sant.: on Usnea;brachy-
sporus (Zopf) M.Brand & Diederich: on Porpidia;Micro-
thelia calcaricola Mudd: on Aspicilia;caudisporus
J.C.David & Etayo: on Lathagrium auriforme;cladiae
Zhurb. & Pino-Bodas: on Cladia;complanatae Arnold:
on Miriquidica;exerrans Nyl.: on Rhizocarpon;Verrucaria
fumosaria Leight.: on Lecidea fuscoatra;fusiger Th.Fr. &
Almq.: on Rhizocarpon;incrassatus Etayo & Breuss: on
Placidiopsis;janae K.Knudsen: on Acarospora rhabarbarina;
karlstadtensis Kocourk. & Brackel: on Endocarpon;Ticho-
thecium latzelii Keissl.: on Collema;macrosporus (Arnold)
Nyl.: on Rhizocarpon;matzeri D.Hawksw. & Iturr.: on
Buellia;Mycoporum melacoccum Nyl.: on a sterile,
saxicolous lichen with a sorediate Cþred thallus; nanellus
Ohlert: on Stereocaulon;oreinae Hafellner: on Dimelaena
oreina;oropogonicola Etayo: on Oropogon;pallidosporus
Etayo: on Pseudocyphellaria;parentium Etayo: on Pseudo-
cyphellaria;parmeliarum Etayo: on Parmelia saxatilis;
pellax Nyl.: on Collema;peltigericola Zhurb. & Stepanchi-
kova: on Peltigera;perminutus Vain.: on Bacidia sub-
incompta;perpusillus Nyl.: on Schaereria, also reported
from other saxicolous lichens; propinquus (K ¨
orb.)
D.Hawksw.: on Porpidia,alsoreportedfromother
saxicolous lichens; protoblasteniae Diederich: on Proto-
blastenia;pseudocarpus Nyl.: on Callome,Enchylium,
Lathagrium,Peltula and Scytinium;pseudocyphellariae
Etayo: on Pseudocyphellaria (also Nephroma); ramalinarius
(Linds.) D.Hawksw.: on Ramalina;rugulosus (Leight.)
364 The Bryologist 121(3): 2018
Nyl.: on Verrucaria, also reported from other saxicolous
lichens; sendtneri (Arnold) Hafellner: on Polyblastia
sendtneri;sipmanii Etayo: on Heterodermia;stigma (K¨
orb.)
Stizenb. [”Discothecium stigma (K¨
orb.) Zopf]: on
Acarospora;thamnoliae Etayo & R.Sant.: on Thamnolia;
thelommatis Kocourk. & K.Knudsen: on Thelomma
santessonii;tricolorans Alstrup: on Platismatia glauca;
variabilis Halıcı, Kocourk. & Diederich: on Staurothele;
verrucisporus Alstrup: on Ionaspis lacustris;verrucosus
Hafellner: on Aspicilia;xanthoparmeliae Y.Joshi,
S.Y.Kondr., L.L}
ok¨
os & Hur: on Xanthoparmelia;zahl-
brucknerellae (Henssen) D.Hawksw.: on Zahlbrucknerella.
Lit.: Alstrup 1993b, 1996; Alstrup et al. 1994; Brackel &
Kocourkova
´2006; David & Etayo 1995; Etayo 2002a, 2010b,
2017; Etayo & Breuss 2001; Etayo & Sancho 2008; Hafellner 1994a;
Hafellner et al. 2002, 2008; Halıcıet a1. 2007b; Hawksworth 1979b,
1982a; Hawksworth & Iturriaga 2006; Joshi et al. 2015; Knudsen
2008; Kocourkova
´& Knudsen 2011a; Matzer 1993; Matzer et al.
1996; Santesson 1994b; S ´
erusiaux et al. 1999; Triebel 1989;
Zhurbenko & Pino-Bodas 2015; Zhurbenko et al. 2012.
Haleomyces D.Hawksw. & Essl.; type: H. oropogonicola. Mol.:
no.
Species: 1-0-0. oropogonicola D.Hawksw. & Essl.: on
Oropogon lopezii.
Lit.: Hawksworth & Esslinger 1993.
Heteroplacidium Breuss; type: H. imbricatum (Nyl.) Breuss
(lichenized). Mol.: yes (T, L).
Species: 0-3-0 (12). fusculum (Nyl.) Gueidan & Cl.Roux:
lichen, juvenile parasite on Aspicilia calcarea;transmutans
K.Knudsen, Breuss & Kocourk.: lichen, juvenile parasite on
Acarospora socialis;zamenhofianum (Clauzade & Cl.Roux)
Cl.Roux [”Verrucaria zamenhofiana Clauzade & Roux]:
lichenicolous lichen on Staurothele areolata, eventually
becoming independent.
Lit.: Breuss 1994; Knudsen et al. 2014.
Halospora (Zschacke) Tomas. & Cif.; type: H. deminuta. Mol.:
no.
Species: 4-0-0. deminuta (Arnold) Tomas. & Cif. [¼
Merismatium deminutum (Arnold) Cl.Roux & Nav.-Ros.]:
on Polyblastia,Staurothele,Thelidium and Verrucaria, etc.;
discrepans (Arnold) Hafellner [¼M. discrepans (Arnold)
Triebel]: mainly on Protoblastenia,alsoonAspicilia,
Hymenelia,Verrucaria, etc.; longispora (Cl.Roux & Nav.-
Ros.) Hafellner [¼M. deminutum subsp. longisporum
Cl.Roux & Nav.-Ros.]: on endolithic lichens; scammoeca
(Lettau) Hafellner [¼M. scammoecum Lettau]: on Poly-
blastia,Thelidium and Verrucaria.
Lit.: Hafellner 2011; Roux et al. 2002; Triebel 1989.
Norrlinia Theiss. & Syd.; type: N. peltigericola. Mol.: no.
Species: 2-0-0. medoborensis S.Y.Kondr.: on Peltigera;
peltigericola (Nyl.) Theiss. & Syd.: on Peltigera.
Lit.: Hawksworth 1980b; Kondratyuk & Galloway 1995a.
Phaeospora Stein; type: P. rimosicola. Mol.: no.
Species: 14-0-0. arctica Hora
´kova
´& Alstrup: on Arctopar-
melia,Cetraria and Nephromopsis;catolechiae Zopf: on
Catolechia wahlenbergii;everniae Etayo & van den Boom:
on Evernia;exoriens (Stirt.) A.L.Sm.: on Pannaria;
lecanorae Eitner: on Lecanora and Myriolecis;parasitica
(L¨
onnr.) Arnold: on Rhizocarpon [and Lopadium?]; pelti-
gericola D.Hawksw.: on Peltigera and Pseudocyphellaria
[and Lobaria?]; perrugosaria (Linds.) D.J.Galloway: on
Placopsis;protoblasteniae Alstrup & Olech: on Protoblas-
tenia incrustans;quaterna Eitner: on Rhizocarpon;rimosi-
cola (Mudd) Stein: on Rhizocarpon [also reported on
Aspicilia and Diplotomma]; squamarinae Etayo: on Squa-
marina cartilaginea;subantarctica Øvstedal & D.Hawksw.:
on Lecania gerlachei;verrucariae Alstrup & E.S.Hansen: on
Verrucaria placynthiae.
Lit.: Alstrup & Hansen 2001; Alstrup & Olech 1996; Etayo 2010c;
Galloway 2004; Hawksworth 1980b, 1983; Hora
´kova
´& Alstrup
1994; Øvstedal & Hawksworth 1986; S´
erusiaux et al. 1999; van den
Boom & Etayo 2014; Zhurbenko 2014.
Placidiopsis Beltr.; type: P. grappae Beltr. (lichenized).Mol.: yes (T).
Species: 0-1-0 (20). minor var. longispora Etayo & Breuss:
lichenicolous lichen on Diploschistes.
Lit.: Etayo 2017.
Placocarpus Trevis.; type: P. schaereri. Mol.: yes (T, L).
Species: 0-3-0 (5). americanus K.Knudsen, Breuss &
Kocourk.: lichen, juvenile parasite on Protoparmeliopsis
muralis;melanophthalmosus Cl.Roux & Gueidan: lichen,
juvenile parasite on Rhizoplaca melanophthalma;schaereri
(Fr.) Breuss: lichen, juvenile often parasitic on Protoparme-
liopsis muralis.
Lit.: Breuss 1985; Knudsen et al. 2009a; Roux & Gueidan 2011;
Zhurbenko & Notov 2015.
Placopyrenium Breuss; type: P. bucekii (Na
´dv. & Serv´
ıt) Breuss
(lichenized). Mol.: yes (T, L).
Species: 0-6-0 (22). breussii Cl.Roux & Gueidan: lichen,
juvenile parasite on Aspicilia calcitrapa;canellum Nyl.:
lichen, juvenile parasite on Aspicilia;cinereoatrum (Degel.)
Orange: facultatively lichenicolous on Staurothele fissa;
formosum Orange: lichen, juvenile parasite on Aspicilia
aquatica;fuscellum (Turner) Gueidan & Cl.Roux: lichen,
juvenile facultatively parasite on Bagliettoa and Verrucaria;
noxium Breuss: lichenicolous lichen on Staurothele areolata.
Lit.: Breuss 1998, 2009; Orange 2004, 2009; Roux & Gueidan 2011.
Telogalla Nik.Hoffm. & Hafellner; type: T. olivieri. Mol.: no.
Species: 2-0-0. cajasense Etayo: on Leptogium;olivieri
(Vouaux) Nik.Hoffm. & Hafellner: on Xanthoria.
Lit.: Etayo 2017; Hoffmann & Hafellner 2000.
Thelidium A.Massal.; type: T. amylaceum A.Massal. Mol.: yes.
Species: 0-1-0 (100). impressulum Zschacke: on Verrucar-
iaceae with an endolithic thallus.
Lit.: Roux 2014.
Verrucaria Schrad.; type: V. rupestris Schrad. (lichenized). Mol.:
yes (T). Currently polyphyletic.
Species: 1-5-0 (300). bernardinensis Breuss: lichenicolous
lichen on Staurothele;cetera Breuss: lichenicolous lichen on
Staurothele areolata, eventually becoming independent;
conturmatula Nyl.: on Ionaspis lacustris;falcata Breuss:
lichen appearing lichenicolous on or between areoles of
Verrucaria maculicarpa;inficiens Breuss: lichenicolous
lichen on Staurothele areolata and S. elenkinii, eventually
becoming independent; placynthiae Alstrup & E.S.Hansen:
lichenicolous lichen on Placynthium nigrum.
Lit.: Alstrup & Hansen 2001; Breuss 1998, 2007; Orange 2002.
Verrucula J.Steiner; type: V. aegyptiaca. Mol.: yes (L).
Species: 0-22-0. aegyptiaca (M¨
ull.Arg.) J.Steiner: lichen-
icolous lichen on Variospora;ahlesiana (Zwackh) J.Steiner:
Diederich et al.: 2018 classification and checklist of lichenicolous fungi 365
lichen, probably lichenicolous; arnoldaria Nav.-Ros. &
Cl.Roux: lichenicolous lichen on Calogaya arnoldii;biator-
inaria (Zehetl.) Nav.-Ros. & Cl.Roux: lichenicolous lichen
on Calogaya biatorina;clauzadaria Nav.-Ros. & Cl.Roux:
lichenicolous lichen on Caloplaca clauzadeana;coccinearia
(Zehetl.) Nav.-Ros. & Cl.Roux: lichenicolous lichen on
Caloplaca coccinea;coronataria Nav.-Ros. & Cl.Roux:
lichenicolous lichen on Flavoplaca coronata;elegantaria
(Zehetl.) Nav.-Ros. & Cl.Roux: lichenicolous lichen on
Rusavskia elegans;fulvaria Nav.-Ros. & Cl.Roux ad int.:
lichenicolous lichen on Leproplaca proteus;granulosaria
(Clauzade & Zehetl.) Nav.-Ros. & Cl.Roux: lichenicolous
lichen on Flavoplaca granulosa;helvetica (B. de Lesd.)
Nav.-Ros. & Cl.Roux: lichenicolous lichen on Leproplaca
cirrochroa;hladuniana (Nav.-Ros. & Cl.Roux) Nav.-Ros. &
Cl.Roux: lichenicolous lichen on Flavoplaca marina;
inconnexaria Nav.-Ros. & Cl.Roux: lichenicolous lichen
on Caloplaca inconnexa;lactearia Nav.-Ros. & Cl.Roux:
lichenicolous lichen on Xanthocarpia lactea s.str.; latericola
(Erichs.) Nav.-Ros. & Cl.Roux: lichenicolous lichen on
Calogaya pusilla;maritimaria Nav.-Ros. & Cl.Roux:
lichenicolous lichen on Flavoplaca maritima;microspora
Nav.-Ros. & Cl.Roux: lichenicolous lichen on Variospora
velana;navasaria Nav.-Ros. & Cl.Roux: lichenicolous
lichen on Flavoplaca navasiana;polycarparia Nav.-Ros. &
Cl.Roux.: lichenicolous lichen on Flavoplaca polycarpa;
protearia (Zehetl.) Nav.-Ros. & Cl.Roux: lichenicolous
lichen on Leproplaca proteus;pusillaria Nav.-Ros. &
Cl.Roux: lichenicolous lichen on Calogaya pusilla;tarraco-
nensis Nav.-Ros. & Cl.Roux: lichenicolous lichen on
Variospora.
Lit.: Hoffman & Hafellner 2000; Navarro-Rosin´
es et al. 2007; Roux
2014; Zehetleitner 1978.
Verruculopsis Gueidan, Nav.-Ros. & Cl.Roux; type: V. poelti-
ana. Mol.: yes (T, L).
Species: 0-3-0 (6). flavescentaria Gueidan, Nav.-Ros. &
Cl.Roux: lichenicolous lichen on Calogaya flavescens;
irrubescentis (M´
enard & Cl.Roux) Gueidan, Nav.-Ros. &
Cl.Roux: lichenicolous lichen on Caloplaca irrubescens;
poeltiana (Clauzade & Cl.Roux) Gueidan, Nav.-Ros. &
Cl.Roux: lichenicolous lichen on Calogaya aurantia.
Lit.: M´
enard & Roux 1991; Navarro-Rosin´
es et al. 2007.
Order Verrucariales genera incertae sedis
Gemmaspora D.Hawksw. & Halıcı; type: G. lecanorae. Mol.: no.
Species: 1-0-0. lecanorae (Werner) D.Hawksw. & Halıcı:on
Aspicilia.
Lit.: Hawksworth & Halıcı2007.
Merismatium Zopf; type: M. lopadii (Arnold) Zopf, i.e. M.
nigritellum [¼Heterophracta (Sacc. & D.Sacc.) Theiss. &
Syd.; type: Leptosphaerulina heterophracta, i.e. M. hetero-
phractum;Phaeocyrtis Vain.; type: P. consocians, i.e. M.
heterophractum]. Mol.: no.
Species: 10-0-0. cladoniicola Alstrup: on Cladonia ciliata;
coccisporum (Norm.) Vouaux: on Amygdalaria,Euopsis,
etc.; coccotremicola Etayo: on Coccotrema;corae (Pat.)
Etayo & R.Sant.: on Cora;decolorans (Arnold) Triebel
[¼?Sphaerulina dolichotera (Nyl.) Vouaux; ¼?S. inter-
media Vouaux]: on Lopadium,Bilimbia,Placynthiella,
Trapeliopsis,etc.;heterophractum (Nyl.) Vouaux [¼
Leptosphaerulina heterophracta (Nyl.) Sacc. & D.Sacc.; ¼
Phaeocyrtis consocians (Nyl.) Vain.]: on Biatora,Bilimbia,
Lopadium,etc.;lecanorae (H.Olivier) Vouaux: on Ochro-
lechia parella;nigritellum (Nyl.) Vouaux [¼M. lopadii
(Arnold) Zopf]: on Catapyrenium,Leptogium,Lichen-
omphalia,Lopadium,Micarea,Mycobilimbia,Ochrolechia,
Protoblastenia,etc.;peregrinum (Flot.) Triebel: on
Rimularia;thamnoliicola Alstrup & E.S.Hansen: on
Thamnolia.
Lit.: Alstrup 1997; Alstrup & Hansen 2001; Etayo 2010b; Etayo &
Sancho 2008; Hafellner 2011; Hertel & Rambold 1990; Triebel
1989.
Pseudostigmidium Etayo; type: P. nephromiarium. Mol.: no.
Species: 5-0-0. biseptatum Etayo: on Pseudocyphellaria;
confusum Etayo: on Pseudocyphellaria;disparatum Etayo:
on Pseudocyphellaria;fumosum Etayo: on Pseudocyphel-
laria;nephromiarium (Linds.) Etayo: on Nephroma
antarcticum.
Lit.: Etayo & Sancho 2008.
Stigmidium Trevis.; type: S. schaereri [¼Pharcidia K¨
orb.; type:
P. congesta, i.e. S. congestum]. Mol.: no.
Species: 94-0-0 (95). acetabuli Calat. & Triebel: on
Pleurosticta acetabulum;aggregatum (Mudd) D.Hawksw.:
on Aspicilia;ahtii Etayo & Palice: on apothecia of
Cladonia lopezii;alectoriae (Linds.) Etayo: on Usnea;
allogenum (Nyl.) D.Hawksw.: on Rhizocarpon;arthoniae
(Arnold) Hafellner: on Arthonia;arthrorhaphidis Hafell-
ner & Obermayer: on Arthrorhaphis citrinella;aspiciliae
(J.Lahm) Cl.Roux: on Aspicilia gr. calcarea;bellemerei
Cl.Roux & Nav.-Ros.: on Lecania;beringicum Zhurb. &
Triebel: on Stereocaulon;buelliae Zhurb. & Himelbrant:
on Buellia disciformis;californicum K.Knudsen & Ko-
courk.: on Caloplaca stanfordensis;calopadiae Matzer: on
foliicolous Calopadia;cartilagineae Calat. & Triebel: on
Squamarina;cerinae Cl.Roux & Triebel: on Caloplaca
s.str., also reported from other Teloschistaceae; cladonii-
cola Zhurb. & Diederich: on Cladonia;clauzadei Cl.Roux
&Nav.-Ros.:onVerrucaria;coarctatae S.Y.Kondr.,
L.L}
ok¨
os&J.-S.Hur:onTrapelia coarctata [ascomata
setose: Niesslia?]; collematis Cl.Roux & Triebel: on
Collema;congestum (K¨
orb.) Triebel [”Pharcidia congesta
K¨
orb.]: on apothecia of Lecanora gr. chlarotera;conspur-
cans (Th.Fr.) Triebel & R.Sant.: on Psora rubiformis;
croceae (Arnold) Cl.Roux & Triebel: on Solorina;
cupulare (Pat.) D.Hawksw.: on Sticta;degelii R.Sant.:
on Degelia;disconephromeum Etayo: on Nephroma
antarcticum;divaricatae E.Zimm. & F.Berger: on Evernia
divaricata;ephebes (Henssen) D.Hawksw.: on Ephebe;
epinesolechia Etayo: on Nesolechia oxyspora galls over
Punctelia;epiphyllum Matzer: on foliicolous Actinoplaca
and Echinoplaca;epistigmellum (Vouaux) Kocourk. &
K.Knudsen: on marine Polycauliona;epixanthum Hafell-
ner: on Acarospora;eucline (Nyl.) Vˇ
ezda: on Varicellaria;
exasperatum Etayo: on Melanohalea exasperata;frigidum
(Sacc.) Alstrup & D.Hawksw.: on Thamnolia;fuscatae
(Arnold) R.Sant.: on Acarospora;glebarum (Arnold)
Hafellner: on Toninia;grex Alstrup & Olech: on
Ochrolechia cf. frigida;gyrophorarum (Arnold)
D.Hawksw. [”Endococcus gyrophorarum (Arnold) J.C.Da-
vid & D.Hawksw.]: on Umbilicaria;haesitans (Nyl.)
Diederich, comb. nov. [”Pharcidia haesitans Nyl.]: on
Protothelenella sphinctrinoides;hafellneri Zhurb.: on
Nephromopsis cucullata;hageniae (Rehm) Hafellner: on
Anaptychia and Physconia;hesperium Kocourk., K.Knud-
sen & Diederich: on Polycauliona and Wetmoreana;
366 The Bryologist 121(3): 2018
heterodermiae Etayo: on Heterodermia;humidum P´
erez-
Ortega, Halıcı& K.Knudsen: on Thelenella muscorum;
hygrophilum (Arnold) R.Sant.: on Rhizocarpon and
Porpidia;hypotrachynicola Etayo: on Hypotrachyna;
joergensenii R.Sant.: on Agonimia foliacea;johnii Halıcı
&D.Hawksw.:onCaloplaca s.l.; lecidellae Triebel,
Cl.Roux & Le Coeur: on Lecidella elaeochroma;lendemeri
Kocourk. & K.Knudsen: on Aspicilia;leprariae Zhurb.: on
Lepraria neglecta;leptogii Etayo: on Leptogium;leuco-
phlebiae Cl.Roux & Triebel: on Peltigera leucophlebia;
lichenum (Arnold) Triebel & P.Scholz: on Verrucaria
maculiformis;lobariae Zhurb. & Etayo: on Lobaria;
marinum (Deakin) Swinscow: on maritime Verrucaria;
Pharcidia maritima B. de Lesd.: on Verrucaria [a poorly
known taxon provisionally placed here]; micareae van den
Boom: on Micarea alabastrites;microcarpum Alstrup &
J.C.David: on Cetraria and Nephromopsis cucullata;
microspilum (K¨
orb.) D.Hawksw.: on Graphis scripta;
microsporum Etayo & Osorio: on Teloschistes;mitchellii
Cl.Roux & Bricaud: on Pannaria conoplea;mycobilimbiae
Cl.Roux, Triebel & Etayo: on Bilimbia;neofusceliae Calat.
& Triebel: on Xanthoparmelia pulla;parmotrematis
F.Berger & E.Zimm.: on Parmotrema;parvum (Henssen)
Diederich, comb. nov. [”Pharcidia parva Henssen]: on
Ephebe lanata;peltideae (Vain.) R.Sant.: on Peltigera,also
reported from Pseudocyphellaria,Solorina and Sticta;petri
Hafellner: on Clauzadea monticola;placopsidis E.Zimm.
&F.Berger:onPlacopsis;placynthii Cl.Roux & Nav.-Ros.:
on Placynthium nigrum;porinae Matzer: on foliicolous
Porina;pseudopeltideae Cl.Roux & Triebel: on Peltigera;
psorae (Anzi) Hafellner: on Psora and Psorula;ramalinae
(M¨
ull.Arg.) Etayo & Diederich [¼S. epiramalina (Vouaux)
Hafellner]: on Ramalina;rivulorum (Kernst.) Cl.Roux &
Nav.-Ros.: on Verrucaria;rouxianum Calat. & Triebel: on
Acarospora cervina;schaereri (A.Massal.) Trevis.: on
Solorina thalli parasitized by Dacampia hookeri;schizo-
porum Matzer: on foliicolous Porina;seirophorae Etayo
& van den Boom: on Seirophora scorigena;solorinarium
(Vain.) D.Hawksw.: on Solorina;spegazzinii Etayo: on
Pseudocyphellaria;squamariae (B. de Lesd.) Cl.Roux &
Triebel: on Lecanora,Protoparmeliopsis and Rhizoplaca;
squamarinicola Calat. & Triebel: on Squamarina;stereo-
caulorum Zhurb. & Triebel: on Stereocaulon;stygnospi-
lum (Minks) R.Sant.: on Dermatocarpon;subcladoniicola
van den Boom: on Cladonia;superpositum (Nyl.)
D.Hawksw.: on Sporodictyon;tabacinae (Arnold) Triebel:
on Toninia;tetrasporum Etayo: on Verrucariaceae;
trichotheliorum Matzer: on foliicolous Trichothelium;
triebeliae Etayo: on Parmotrema;Pharcidia verrucaria-
rum (Arnold)Sacc.&D.Sacc.:onVerrucaria [a poorly
known taxon provisionally placed here]; vezdae Matzer:
on foliicolous Fellhanera colchica;xanthoparmeliarum
Hafellner: on Xanthoparmelia.
Lit.: Alstrup 1993a; Alstrup & Hawksworth 1990; Alstrup & Olech
1996; Berger & Zimmermann 2016; Calatayud & Triebel 1999, 2001;
Clauzade et al. 1989; Etayo 1994, 2000, 2002a, 2010c, 2017; Etayo &
Sancho 2008; Hafellner 1982c, 1994a, 1994b; Hafellner & Obermayer
1995; Hafellner et al. 2008; Hawksworth 1975; Kihlman 1891;
Kocourkova
´& Knudsen 2009a, 2009b, 2010, 2012; Kondratyuk et al.
2016b; Lord et al. 2013; Matzer 1996; Matzer & Hafellner 1990;
Molitor & Diederich 1997; P´
erez-Ortega et al. 2010; Roux 2014;
Roux & Navarro-Rosin´
es 1994; Roux & Triebel 1994; Roux et al.
1995; Santesson 1989, 1993b; Triebel 1989; Triebel & Scholz 2001;
Roux et al. 1998; van den Boom 2016; van den Boom & Etayo 2017;
Zhurbenko & Etayo 2012; Zhurbenko & Diederich 2008; Zhurbenko
et al. 2012; Zimmermann & Berger 2018.
Subclass MYCOCALICIOMYCETIDAE Tibell
Order Mycocaliciales Tibell & Wedin
Family Mycocaliciaceae A.F.W.Schmidt
Chaenothecopsis Vain.; type: C. rubescens. Mol.: yes (L).
Species: 24-0-9 (40). arthoniae Tibell: on Arthonia vinosa;
australis Tibell: on Lecanactis abietina;brevipes Tibell: on
Arthonia and Lecanactis;cinerea Tibell: on Lecanactis abietina;
consociata (Na
´dv.) A.F.W.Schmidt: on Chaenotheca chrysoce-
phala;dibbleandersoniarum Selva: on Felipes leucopellaeus;
epithallina Tibell: on Chaenothecatrichialis;formosaTitov: on
Chaenotheca sphaerocephala;haematopus Tibell: facultatively
lichenicolous; hospitans (Th.Fr.) Tibell: on Haematomma
ochroleucum and Lecanora;kalbii Tibell & K.Ryman: on
Lecanora caesiorubella;lecanactidis Tibell: on Lecanactis
abietina;leifiana Titov: on Arthonia;lignicola (Na
´dv.)
A.F.W.Schmidt: facultatively lichenicolous, especially on
Chaenotheca;nigra Tibell: on Chaenotheca;ochroleuca
(K¨
orb.) Tibell & K.Ryman: on Haematomma ochroleucum;
parasitaster(Bagl.& Carestia)D.Hawksw.:onCladonia;pilosa
Tibell & Kalb: on Tylophoron;pusilla (Ach.) A.F.W.Schmidt:
onCalicium,Chaenotheca,Psilolechia,algae or most commonly
lignicolous; pusiola (Ach.) Vain.: on Chaenotheca or sapro-
phytic; retinens (Nyl.) Tibell: on Inoderma byssaceum and
Sporodophoron cretaceum;rubescens Vain.: on Inoderma
byssaceum, other lichens with Trentepohlia or saprophytic;
rubina Tibell: on Pyrgidium montellicum;sagenidii Tibell: on
Lecanactismollis;savonica(R¨
as¨
anen)Tibell:on Chaenothecaor
saprophytic; sinensis Titov: on unidentified lichen with
chlorococcoid algae; subparoica (Nyl.) Tibell: on Haematom-
ma ochroleucum;tasmanica Tibell: on diverse lichens, incl.
Chaenotheca chrysocephala;treicheliana (Stein) Kalb: on
corticolous Lecanora;vainioana (Na
´dv.) Tibell: on Arthonia,
Lecanactis abietina,Calicium salicinum, other lichens with
Trentepohlia or saprophytic; vinosa Titov: on lichens; vir-
idialba (Kremp.) A.F.W.Schmidt: on Chaenotheca chryscepha-
la or saprophytic; viridireagens (Na
´dv.) A.F.W.Schmidt: on
Calicium,Chaenotheca or on free-living algae.
Lit.: Clauzade et al. 1989; Gasparyan et al. 2014; Selva 2013; Tibell
1991b, 1998; Tibell & Ryman 1995; Titov 2006.
Mycocalicium Vain.; type: M. parietinum (Ach.) Vain. Mol.: yes.
Species: 3-0-0 (12). chiodectonicola Aptroot & Etayo: on
Chiodecton;enterographicola Aptroot & M.Ca
´ceres: on
Enterographa cf. quassiaecola;rapax Tibell: on Pertusaria.
Lit.: Aptroot et al. 2016; Etayo & Aptroot 2017; Titov 2006.
Sphinctrina Fr.; type: S. turbinata. Mol.: yes (T, L).
Species: 9-0-0. anglica Nyl.: on Protoparmelia;benmargana
Selva: on Pertusaria;intermedia Tibell: on Pertusaria;
leucopoda Nyl.: on Pertusaria and Diploschistes;ophiopar-
mae Kalb: on Ophioparma araucariae;pallidella (Willey)
Selva: on Pertusaria;paramerae D.Mu˜
niz & Hladun: on
Pertusaria paramera;tubaeformis A.Massal.: on Pertusaria;
turbinata (Pers.) de Not.: on Pertusaria.
Lit.: Kalb 2001; L¨
ofgren & Tibell 1979; Mu˜
niz et al. 2013; Purvis et
al. 1992; Tibell et al. 2014; Titov 2006.
Subclass SCLEROCOCCOMYCETIDAE R ´
eblova
´,
Unter. & W.Gams
Diederich et al.: 2018 classification and checklist of lichenicolous fungi 367
Order Sclerococcales R´
eblova
´, Unter. & W.Gams
Family Dactylosporaceae Bellem. & Hafellner [¼
Sclerococcaceae R´
eblova
´, Unter. & W.Gams]
Sclerococcum Fr.; type: S. sphaerale [¼Dactylospora K¨
orb.; type:
D. floerkei, i.e. D. parasitica]. Mol.: yes (T, L).
Species: 60-0-4 (80). acarosporae S.Y.Kondr.: on Acarospora;
acarosporicola Ertz&Diederich,nom.nov.[”Dactylospora
acarosporae (H.Magn.) Hafellner]: on Acarospora;aerugino-
sum (Holien & Ihlen) Ertz & Diederich, comb. nov. [”D.
aeruginosa Holien & Ihlen]: on various lichens, also on bark
or wood; ahtii (Zhurb. & Pino-Bodas) Ertz & Diederich,
comb. nov. [”D. ahtii Zhurb. & Pino-Bodas]: on Cladonia;
allantoideum (Alstrup & D.Hawksw.) Ertz & Diederich,
comb. nov. [”D. allantoidea Alstrup & D.Hawksw.]: on
Xanthoparmelia pulla;amygdalariae (Triebel) Ertz & Die-
derich, comb. nov. [”D. amygdalariae Triebel]: on
Amygdalaria;anziae (Zhurb., Ezhkin, Skirina & Y.Ohmura)
Ertz & Diederich, comb. nov. [”D. anziae Zhurb., Ezhkin,
Skirina & Y.Ohmura]: on Anzia;aptrootii Diederich: on
Fissurina dumastii;areolatum Etayo: probably on Xylogra-
pha;aspiciliicola (Alstrup & D.Hawksw.) Ertz & Diederich,
comb. nov. [”D. aspiciliicola Alstrup & D.Hawksw.]: on
Aspicilia;athallinum (M¨
ull.Arg.) Ertz & Diederich, comb.
nov. [”D. athallina (M¨
ull.Arg.) Hafellner; ¼?Abrothallus
placophyllus Anzi]: on Baeomyces;attendendum (Nyl.) Ertz &
Diederich, comb. nov. [”D. attendenda (Nyl.) Arnold]: on
Amygdalaria,Icmadophila,Pilophorus and Porpidia;australe
(Triebel & Hertel) Ertz & Diederich, comb. nov. [”D.
australis Triebel & Hertel]: on Coccotrema,Lecidea,Para-
porpidia,Poeltiaria,Poeltidea,Porpidia,Rimularia and
Tremolecia;boreale (Holien & Ihlen) Ertz & Diederich,
comb. nov. [”D. borealis Holien & Ihlen]: on Mycoblastus,
other lichens or directly on the bark; cladoniicola (Alstrup &
Olech) Ertz & Diederich, comb. nov. [”D. cladoniicola
Alstrup & Olech]: on Cladonia;crassitunicatum Zhurb.,
Diederich & U.Braun: on Cladonia;crassum (Sarri´
on &
Hafellner) Ertz & Diederich, comb. nov. [”D. crassa Sarri´
on
& Hafellner]: saprotrophic on Juniperus oxycedrus,alsoonold
Parmelia;davidii (Hafellner & H.Mayrhofer) Ertz & Die-
derich, comb. nov. [”D. davidii Hafellner & H.Mayrhofer]:
on Pseudocyphellaria;deminutum (Th.Fr.) Ertz & Diederich,
comb. nov. [”D. deminuta (Th.Fr.) Triebel;¼D. rinodinicola
Alstrup & D.Hawksw.]: on various lichens; dobrowolskii
(Olech & Alstrup) Ertz & Diederich, comb. nov. [”D.
dobrowolskii Olech & Alstrup]: on Calogaya saxicola;
epicladonia Zhurb.: on Cladonia;epiphytorum Diederich:
on Pertusaria coccodes and Varicellaria hemisphaerica;frigid-
um (Hafellner) Ertz & Diederich, comb. nov. [”D. frigida
Hafellner]: on Brigantiaea;gelidarium Etayo & F.Berger: on
Placopsis;glaucomarioides (Tuck.) Ertz & Diederich, comb.
nov. [”D. glaucomarioides (Tuck.) Hafellner]: on Ochro-
lechia;hafellnerianum (S´
erus.) Ertz & Diederich, comb. nov.
[”D. hafellneriana S´
erus.]: on Santessonia lagunebergii;
heterodermiae (Etayo) Ertz & Diederich, comb. nov. [”D.
heterodermiae Etayo]: on Heterodermia;homoclinellum
(Nyl.) Ertz & Diederich, comb. nov. [”D. homoclinella
(Nyl.) Hafellner]: on Lecanora and Tephromela;inconspic-
uum (Etayo) Ertz & Diederich, comb. nov. [”D. inconspicua
Etayo]: on Menegazzia;inquilinum (Tuck.) Ertz & Diederich,
comb. nov. [”D. inquilina (Tuck.) Hafellner]: on corticolous
Pertusaria;leuckertii Diederich & P.Scholz: on Buellia
aethalea;lobariellum (Nyl.) Ertz & Diederich, comb. nov.
[”D. lobariella (Nyl.) Hafellner]: on Lobaria and Ricasolia,
also reported on Pseudocyphellaria;microsporum (Etayo)
Ertz & Diederich, comb. nov. [”D. microspora Etayo]: on
Catinaria atropurpurea;montagnei Hafellner: on Lecanora
rupicola;olivaceum Etayo & F.Berger: on Flavoparmelia
soredians;ophthalmizae Coppins, sp. nov.: on Lepra
ophthalmiza;orygmaeum (Nyl.) Ertz & Diederich, comb.
nov. [”D. orygmaea (Nyl.) S.Y.Kondrat. & D.J.Galloway]: on
Pseudocyphellaria berberina;parasitaster (Nyl.) Ertz & Die-
derich, comb. nov. [”D. parasitaster (Nyl.) Arnold]: on
Biatora sphaeroides and Thelenella muscorum;parasiticum
(Fl¨
orke) Ertz & Diederich, comb. nov. [”D. parasitica
(Fl¨
orke) Zopf; ¼D. floerkei K¨
orb., nom. illeg.; ¼Sclerophyton
occidentale Herre]: on Lepra,Ochrolechia and Pertusaria;
parellarium (Nyl.) Ertz & Diederich, comb. nov. [”D.
parellaria (Nyl.) Arnold]: on Ochrolechia parella;pertusar-
iicola (Tuck.) Ertz & Diederich, comb. nov. [”D.
pertusariicola (Tuck.) Hafellner]: on saxicolous Pertusaria;
phaeophysciae Diederich & van den Boom: on Phaeophyscia
orbicularis;phyllobaeis Etayo: on Phyllobaeis;pleiospermum
(Triebel) Ertz & Diederich, comb. nov. [”D. pleiosperma
Triebel]: on Lecanora caesiorubella;polysporum (Triebel)
Ertz & Diederich, comb. nov. [”D. polyspora Triebel]: on
Protoparmelia;porphyreum (Hafellner & Kalb) Ertz &
Diederich, comb. nov. [”D. porphyrea Hafellner & Kalb]:
on Bacidia,Brigantiaea and Phyllopsora;protothallinum
(Anzi) Ertz & Diederich, comb. nov. [”D. protothallina
(Anzi) Hafellner]: on Fuscopannaria and Parmeliella;pur-
purascens (Triebel) Ertz & Diederich, comb. nov. [”D.
purpurascens Triebel]: on Amygdalaria,Pilophorus and
Porpidia;pyrenaicum (Etayo) Ertz & Diederich, comb. nov.
[”D. pyrenaica Etayo]: over bryophytes, perhaps associated
with Mycobilimbia pilularis;rhyparizae (Arnold) Ertz &
Diederich, comb. nov. [”D. rhyparizae Arnold]: on Bryodina
rhipariza;rimulicola (M¨
ull.Arg.) Ertz & Diederich, comb.
nov. [”D. rimulicola (M¨
ull.Arg.) Hafellner]: on Lecanora
and ?Pertusaria;rostrupii (Alstrup) Ertz & Diederich, comb.
nov. [”D. rostrupii Alstrup]: on Lepra dactylina;saxatile
(Schaer.) Ertz & Diederich, comb. nov. [”D. saxatilis
(Schaer.) Hafellner]: on Ochrolechia and Pertusaria;saxatile
var. laureri (Hafellner) Ertz & Diederich, comb. nov. [”D.
saxatilis var. laureri Hafellner]: on Pertusaria;serusiauxii
Boqueras & Diederich: on Parmelina;simplex D.Hawksw.:
on corticolous Lepra and Pertusaria;sipmanii Diederich: on
Anomomorpha cf. roseola;sphaerale (Ach.) Fr.: on saxicolous
Pertusaria [mainly P. corallina]; suburceolatum (Coppins &
Fryday) Ertz & Diederich, comb. nov. [”D. suburceolata
Coppins & Fryday]: on a whitish-yellowish muscicolous
crustose lichen; tegularum (Arnold) Ertz & Diederich, comb.
nov. [”D. tegularum (Arnold) Hafellner]: on Caloplaca s.l.,
incl. Rufoplaca;tephromelarum Etayo & Calat.: on Teph-
romela;toensbergii Diederich: on Megalaria pulverea and
Pertusaria carneopallida;urceolatum (Th.Fr.) Ertz & Die-
derich [”Dactylospora urceolata (Th.Fr.) Arnold]: on
Protothelenella sphinctrinoides, also reported from other hosts;
verrucisporum Alstrup: on Bellemerea diamarta.
Lit.: Alstrup 1993a; Alstrup & Hawksworth 1990; Alstrup & Olech
1993; Alstrup et al. 1994; Berger 2000; Boqueras & Diederich 1993;
Brackel 2014; Clauzade et al. 1989; Diederich 1990, 2015; Diederich
& Scholz 1995; Diederich & van den Boom 2017; Diederich et al.
2013; Etayo 1991, 2010c, 2017; Etayo & Berger 2002; Etayo &
Calatayud 1998; Etayo & Sancho 2008; Fryday & Coppins 2012;
Hafellner 1979, 1985b, 1996b; Hafellner & Mayrhofer 2007;
Hafellner et al. 2002; Hawksworth 1975, 1979a, 2003; Ihlen et al.
2004a; Kondratyuk & Galloway 1995c; Kondratyuk & Zelenko
2002; Olech & Alstrup 1996; Pino-Bodas et al. 2017; R´
eblova
´et al.
2017; Sarri ´
on et al. 2002; S´
erusiaux & Wessels 1984; Sparrius 2004;
368 The Bryologist 121(3): 2018
Triebel 1989; Zhurbenko & Pino-Bodas 2017; Zhurbenko et al.
2017.
Class Eurotiomycetes familia incertae sedis
Family Rhynchostomataceae Winka & O.E.Erikss.
Rhynchomeliola Speg.; type: R. pulchella Speg. Mol.: yes (fide
Wijayawardene et al. 2017).
Species: 1-0-0 (3). lichenicola Henssen & Kantvilas: on
Wawea.
Lit.: Henssen & Kantvilas 1985.
Class LECANOROMYCETES O.E.Erikss. & Winka
Subclass ACAROSPOROMYCETIDAE Reeb, Lutzo-
ni & Cl.Roux
Order Acarosporales Reeb, Lutzoni & Cl.Roux
Family Acarosporaceae Zahlbr.
Acarospora A.Massal.; type: A. schleicheri (Ach.) A.Massal.
(lichenized). Mol.: yes (T, L). Some species described by H.
Magnusson are poorly known.
Species: 0-25-0 (200). anatolica H.Magn.: lichenicolous
lichen on Dimelaena;brevilobata H.Magn.: lichenicolous
lichen on Rhizocarpon;epithallina H.Magn.: lichenicolous
lichen on Acarospora hilaris and Usnochroma carphinea;freyi
H.Magn.: facultatively lichenicolous lichen on Aspicilia;
heufleriana K¨
orb.: lichenicolous lichen on Lecanora vale-
siaca;hospitans H.Magn.: lichenicolous lichen on Aspicilia,
Bellemerea,Leanora,Lecidea and Tremolecia;imbricatula
H.Magn.: facultatively lichenicolous lichen on Aspicilia;
impressula Th.Fr.: facultatively lichenicolous lichen on
Aspicilia;insignis H.Magn.: lichenicolous lichen on Aspicilia;
insolata (B. de Lesd.) Redinger: lichenicolous lichen on
Immersaria and Rhizocarpon, becoming independent; inter-
jecta H.Magn.: lichenicolous lichen on Acarospora novomex-
icana;invadens H.Magn.: lichenicolous lichen on Aspicilia
and Dimelaena, becoming independent; jenisejensis
H.Magn.: lichenicolous lichen on Buellia,Dimelaena and
Lecanora, becoming independent; microcarpa (Nyl.) Wed-
dell: lichenicolous lichen on Diploschistes actinostomus, more
rarely Acarospora,Dimelaena and Ingvariella, becoming
independent; nashii K.Knudsen: lichenicolous lichen on
Dimelaena and Pleopsidium, becoming independent; nod-
ulosa (Dufour) Hue: facultatively lichenicolous lichen on
Diploschistes diacapsis;pulvinata H.Magn.: lichenicolous
lichen on Aspicilia;sphaerospora H.Magn.: lichenicolous
lichen on Aspicilia, becoming independent; stapfiana
(M¨
ull.Arg.) Hue: lichenicolous lichen on Caloplaca s.l. and
Rusavskia elegans;strigata (Nyl.) Jatta: lichenicolous lichen
on Aspicilia, becoming independent; succedens H.Magn.:
facultatively lichenicolous lichen on Dimelaena oreina;
superans H.Magn.: lichenicolous lichen on Aspicilia and
Dimelaena;suprasedens H.Magn.: lichenicolous lichen on
Aspicilia and Lecanora;tominiana H.Magn.: lichenicolous
lichen on Aspicilia;umbrina H.Magn.: lichenicolous lichen
on Lecanora.
Lit.: Knudsen 2011; Knudsen & Kocourkova
´2010a; Rambold &
Triebel 1992.
Myriospora N¨
ageli; type: M. rufescens (Ach.) N¨
ageli (liche-
nized). Mol.: yes (T).
Species: 0-1-0 (9). hassei (Herre) K.Knudsen & L.Arcadia:
facultatively lichenicolous lichen (juvenile obligately lichen-
icolous?) on Acarospora socialis.
Lit.: Knudsen & Bungartz 2014.
Polysporina Vˇ
ezda; type: P. simplex (Taylor) Vˇ
ezda (lichenized).
Mol.: yes (T, L).
Species: 3-0-0 (10). arenacea (H.Magn.) K.Knudsen &
Kocourk.: on crustose lichens on siliceous rocks; pusilla
(Anzi) Kantvilas: on crustose lichens on calcareous rocks,
e.g., Acarospora,Caloplaca s.l., Protoblastenia;subfusces-
cens (Nyl.) K.Knudsen & Kocourk.: on crustose lichens on
siliceous rocks, e.g., Acarospora,Buellia,Caloplaca s.l.,
Candelariella,Lecidea.
Lit.: Knudsen & Kocourkona
´2008.
Sarcogyne Flot.; type: S. corrugata Flot. (lichenized). Mol.: yes
(L).
Species: 0-1-0 (28). sphaeospora J.Steiner: lichenicolous
lichen on Candelariella.
Lit.: Lendemer et al. 2009; Pino-Bodas et al. 2017.
Subclass CANDELARIOMYCETIDAE Timdal &
M.Westb.
Order Candelariales Mia
˛dl., Lutzoni & Lumbsch
Family Candelariaceae Hakul.
Candelariella M¨
ull.Arg.; type: C. vitellina (Ehrh.) M¨
ull.Arg.
(lichenized). Mol.: yes (T).
Species: 0-2-0 (50). dispersa (R¨
as¨
anen) Hakul.: lichen-
icolous lichen on Placynthium;superdistans (Nyl.) Malme:
lichenicolous lichen on Lecanora populicola.
Lit.: Gilbert & James 2009.
Subclass LECANOROMYCETIDAE Mia
˛dl., Lutzoni
& Lumbsch
Order Caliciales Bessey
Family Caliciaceae Chevall.
Acolium (Ach.) Gray; type: A. sessile. Mol.: yes.
Species: 2-0-0 (5). marcianum (B. de Lesd.) M.Prieto &
Wedin [”Cyphelium marcianum B. de Lesd.]: on
Pertusaria, mainly P. pseudocorallina;sessile (Pers.) Trevis.
[”C. sessile (Pers.) Trevis.]: on Pertusaria coccodes and
other corticolous Lepra and Pertusaria.
Lit.: Giavarini & Purvis 2009a; Prieto & Wedin 2017; Tibell 1971.
Amandinea Scheid. & M.Mayrhofer; type: A. coniops (Wahlenb.)
Scheid. & M.Mayrhofer. Mol.: yes (T).
Diederich et al.: 2018 classification and checklist of lichenicolous fungi 369
Species: 2-0-0 (35). adjuncta (Th. Fr.) Hafellner [”Buellia
adjuncta Th. Fr.]: on Lecanora straminea (possibly also on
Caloplaca and Rinodina olivaceobrunnea); deminuta Ha-
fellner (Febr. 2004) [¼Buellia uruguayensis Etayo & Osorio
(July 2004)]: on Caloplaca felipponei.
Lit.: Etayo & Osorio 2004; Hafellner 2004c.
Buellia De Not.; type: B. disciformis (Fr.) Mudd (lichenized).
Mol.: yes (T, L).
Species: 6-8-0 (300). albulella Elix: on Buellia albula;
caloplacivora Llimona & Egea: lichenicolous lichen on
Caloplaca s.l., becoming independent; campbelliana Elix:
lichenicolous lichen on basal squamules of Cladonia;
griseosquamulata Scheidegger: lichenicolous lichen on
Buellia tirolensis;imshaugii Hafellner: lichenicolous lichen
on Dimelaena oreina;insularis Øvstedal: lichenicolous lichen
on Pannaria and Vestergrenopsis;miriquidica Scheidegger:
lichenicolous lichen on Schaereria fuscocinerea;servilosina
Elix & Kantvilas: on Xanthoparmelia mougeotina;subad-
juncta Elix & Kantvilas: on Caloplaca lateritia;teloschistis
Vˇ
ezda: on Teloschistes;thelotremicola Elix: on Thelotrema;
uberior Anzi: lichenicolous lichen on Schaereria fuscocinerea,
soon becoming independent; uberiuscula (Nyl.) Zahlbr.:
facultatively lichenicolous lichen on Acarospora fuscata;
vouauxii Calat. & Barreno: on Rhizoplaca melanophthalma.
Lit.: Calatayud & Barreno 1995; Clauzade et al. 1989; Elix 2018; Elix
et al. 2017; Hafellner 1979; Llimona & Egea 1984; Øvstedal et al.
2009; Rambold & Triebel 1992; Scheidegger 1987, 1993.
Caliciella Vain.; type: not selected. Mol.: no.
Species: 1-0-0. parasitica R¨
as¨
anen: on Chaenotheca furfuracea.
Lit.: Clauzade et al. 1989; Kaufmann & Hofmann 1998.
Calicium Pers.; type: C. viride Pers. Mol.: yes (T, L).
Species: 1-0-0. corynellum (Ach.) Ach.: on Haematomma
ochroleucum and other saxicolous lichens.
Lit.: Roux 2014.
Dimelaena Norman; type: D. oreina (Ach.) Norman (liche-
nized). Mol.: yes (T).
Species: 0-1-0 (10). lichenicola K.Knudsen., Sheard,
Kocourk. & H.Mayrhofer: lichen, juvenile lichenicolous
on D. oreina and D. thysanota.
Lit.: Knudsen et al. 2013.
Diplotomma Flot.; type: D. alboatrum (Hoffm.) Flot. (liche-
nized). Mol.: yes (T).
Species: 1-6-0 (30). murorum (A.Massal.) Coppins: lichen-
icolous lichen on Caloplaca teicholyta;nivale (Bagl. &
Carestia) Hafellner: on Rusavskia and Caloplaca s.l., soon
with its own thallus; parasiticum (B. de Lesd.) Diederich,
Cl.Roux & Van Haluwyn: on Myriolecis;renobalesiii
Cl.Roux [¼Buellia lecanoricola Renob., nom. nud.]: on
Myriolecis albescens, soon with its own thallus; scheidegger-
ianum (Bricaud & Cl.Roux) Nimis: lichenicolous lichen on
Leproplaca xantholyta;venustum K¨
orb.: juvenile lichen-
icolous lichen on Protoparmeliopsis versicolor, soon with its
own thallus; vezdanum (P.Scholz & Knoph) Coppins:
lichenicolous lichen on Polycauliona verruculifera.
Lit.: Bricaud & Roux 1991; Duke & Purvis 2009; Hafellner 1979;
Renobales 1996; Roux 2014.
Endohyalina Marbach; type: E. rappii (R.C.Harris) Marbach
(lichenized). Mol.: no.
Species: 2-1-0 (10). brandii Giralt, van den Boom & Elix:
on Aspicilia intermutans, often lichenized with an own
thallus; diederichii Giralt, van den Boom & Elix: on
Ramalina,rarelylichenizedwithareducedthallus;
insularis (Arnold) Giralt, van den Boom & Elix: on
Lecanora rupicola agg., very rarely lichenized with a reduced
thallus.
Lit.: Giralt et al. 2010.
Monerolechia Trevis.; type: M. bayrhofferi (Schaer.) Trevis.
Mol.: yes (L).
Species: 0-4-0. badia (Fr.) Kalb [”Buellia badia (Fr.)
A.Massal.]: lichen, juvenile parasite on saxicolous lichens;
californica (H.Magn.) Elix [”Dimelaena californica
(H.Magn.) Sheard]: lichen, juvenile parasite on saxicolous
Dimelaena radiata;glomerulans (M¨
ull.Arg.) Elix [”Buellia
glomerulans (M¨
ull.Arg.) Zahlbr.]: lichen, juvenile parasite
on Xanthoparmelia;norstictica Elix: lichen, juvenile
parasite on saxicolous Lecanora or Pertusaria.
Lit.: Elix 2015; Kalb 2004.
Tetramelas Norman; type: not selected. Mol.: yes (L).
Species: 2-1-0 (16). concinnus (Th.Fr.) Giralt: lichenicolous
lichen on Buellia,Melanelia,Parmelia,Physcia and
Pseudephebe, becoming independent; phaeophysciae
A.Nordin & Tibell: on Phaeophyscia and Physcia;pulver-
ulentus (Anzi) A.Nordin & Tibell [”Buellia pulverulenta
(Anzi) Jatta]: mainly on Physconia, also on Anaptychia,
Phaeophyscia and Physcia.
Lit.: Hafellner & Poelt 1980; Nordin & Tibell 2005; Scheidegger &
Ruef 1988.
Family Physciaceae Zahlbr.
Rinodina (Ach.) Gray; type: R.sophodes (Ach.) A.Massal.
(lichenized). Mol.: yes (T, L).
Species: 1-9-0 (300). afghanica Steiner & Poelt: lichen-
icolous lichen on Calogaya polycarpoides and Diplotomma
alboatrum;canariensis Matzer, H.Mayrhofer & Clerc:
lichenicolous lichen on saxicolous lichens; egedeana
(Linds.) Alstrup & D.Hawksw.: on Parmelia and Peltigera;
epimilvina H.Mayrhofer: lichenicolous lichen on Rinodina
milvina;milvina (Wahlenb.) Th.Fr.: lichen, facultatively
lichenicolous on saxicolous lichens; milvinoides
H.Mayrhofer: lichenicolous lichen on Aspicilia;obnascens
(Nyl.) H.Olivier: lichenicolous lichen on Aspicilia and
Rhizocarpon;parasitica H.Mayrhofer & Poelt: lichenicolous
lichen on Aspicilia,Lecanora and Lecidea;santorinensis
J.Steiner: lichen, facultatively lichenicolous on Diploicia,
Ochrolechia,Pertusaria, etc.; superposita H.Magn.: lichen-
icolous lichen on Lecanora and Lecidea.
Lit.: Alstrup & Hawksworth 1990; Matzer et al. 1994; Mayrhofer
1984a, 1984b; Rambold & Triebel 1992.
Order Lecanorales Nannf.
Family Catillariaceae Hafellner
Catillaria A.Massal.; type: C. chalybeia (Borrer) A.Massal.
(lichenized). Mol.: yes (T).
Species: 3-1-0 (150). lobariicola (Alstrup) Coppins &
Aptroot [”Scutula lobariicola Alstrup; ¼Phoma lobariicola
370 The Bryologist 121(3): 2018
Alstrup]: on Lobarina scrobiculata;mediterranea Hafellner:
lichenicolous lichen with a reduced thallus, on Anaptychia
ciliaris and Niebla bourgaeana;stereocaulorum (Th.Fr.)
H.Olivier [”Scutula stereocaulorum (Th.Fr.) K¨
orb.]: on
Stereocaulon;usneicola Etayo: on Usnea.
Lit.: Alstrup 1997; Etayo 2000; Hafellner 1982b.
Family Lecanoraceae K¨
orb.
Carbonea (Hertel) Hertel; type: C. atronivea [Huea C.W.Dodge
& G.E.Baker may represent an earlier synonym, fide Fryday
2011]. Mol.: yes (L).
Species: 11-6-0 (20). aggregantula (M¨
ull.Arg.) Diederich &
Triebel: on Lecanora (mainly L. polytropa), also reported
from Rhizoplaca;antarctica (C.W.Dodge & G.E.Baker)
D.Hawksw. & Iturr.: on Pseudephebe;assimilis (K¨
orb.)
Hafellner & Hertel: lichenicolous lichen on Aspicilia,
Lecanora,Lecidea,Pertusaria,Rhizocarpon and Tephromela;
atronivea (Arnold) Hertel: lichenicolous lichen on Lec-
idella, becoming independent; degelii Alstrup: on Lecanora
intricata;distans (Kremp.) Hafellner & Obermayer var.
distans: lichenicolous lichen on Lecidea obscurissima and
Orphniospora mosigii;distans var. buelliarum (Hertel)
Hertel: lichenicolous lichen on Buellia;herteliana Hafellner
& Matzer: on Rhizocarpon;Catillaria hospitans H.Magn.:
on Lecidea [a poorly known taxon, provisionally placed
here]; intrudens (H.Magn.) Hafellner: on Rhizocarpon;
invadens (H.Magn.) Andreev: lichen, juvenile lichenicolous
on Rhizocarpon geographicum;neuropogonis Etayo: on
Usnea aurantiaco-atra;Nesolechia oxysporiza J.Steiner: on
Lecanora polytropa [probably a Carbonea, fide Roux 2014];
superjecta (Nyl.) Hertel: on Lecanora disjungenda;super-
sparsa (Nyl.) Hertel: on Lecanora (mainly L. polytropa),
rarely Rhizoplaca;vitellinaria (Nyl.) Hertel: on Candelar-
iella;vorticosa (Fl¨
orke) Hertel: lichen, facultatively lichen-
icolous on Physcia dubia.
Lit.: Alstrup et al. 1994; Chambers et al. 2009; Ertz et al. 2014a;
Etayo & Sancho 2008; Hafellner 1999, 2006; Hawksworth &
Iturriaga 2006; Hertel 1967; Obermayer 1993; Rambold &Triebel
1992, Roux 2014.
Lecanora Ach.; type: L. allophana Nyl. (lichenized). Mol.: yes
(T).
Species: 3-13-0 (550). dispersa f. parasitans (Wedd.) Harm.:
lichenicolous lichen on Dermatocarpon miniatum;epithalli-
na H.Magn.: lichenicolous lichen on Tingiopsidium isidiatum;
gisleriana M¨
ull.Arg.: lichenicolous lichen on Lecanora
epanora,L. handelii and L. subaurea;gyrophorina Poelt &
Zhurb.: lichenicolous lichen on Umbilicaria hyperborea;
invadens H.Magn.: lichenicolous lichen on Aspicilia asiatica;
laatokaensis (R¨
as¨
anen) Poelt: lichenicolous lichen on Aspic-
ilia and other lichens, when young; lasalliae P´
erez-Ortega &
Etayo: on Lasallia pustulata;latro Poelt: lichenicolous lichen
on Miriquidica nigroleprosa;lecanoricola (Alstrup,
D.Hawksw. & R.Sant.) Rambold & Triebel [”Lecidella
lecanoricola Alstrup, D.Hawksw. & R.Sant.]: on Lecanora
cenisia;mosigiicola (Eitner) Hertel & Rambold var. mosigii-
cola: lichenicolous lichen on Orphniospora mosigii;mosigii-
cola var. buelliarum (Hertel) Hertel & Rambold:
lichenicolous lichen on Buellia;nohedensis Cl.Roux &
M.Barbero: lichenicolous lichen on Placopyrenium breussii;
printzenii P´
erez-Ortega, Vivas & Hafellner: lichenicolous
lichen on Umbilicaria;thallophila H.Magn.: lichenicolous
lichen on Dermatocarpon;tolypodes Poelt & Vˇ
ezda: lichen-
icolous lichen on Aspicilia cf. grisea;usneicola Etayo: on
Usnea antarctica.
Lit.: Etayo & Sancho 2006; Hafellner 1995b; Lumbsch et al. 2011;
P´
erez-Ortega & Etayo 2008; Poelt et al. 1996; Rambold & Triebel
1992; Ryan et al. 2004; Zhurbenko 2009.
Lecidella K¨
orb.; type: L. viridans (Flot.) K¨
orb. (lichenized).
Mol.: yes.
Species: 0-2-0 (80). dimelaenophila Hertel: lichen, juvenile
lichenicolous on Dimelaena oreina;vorax Leuckert & Poelt:
lichenicolous lichen on Lecanora rupicola.
Lit.: Rambold & Triebel 1992.
Miriquidica Hertel & Rambold; type: M. complanata (K¨
orb.)
Hertel & Rambold (lichenized). Mol.: yes (T).
Species: 0-6-0 (30). complanata (K¨
orb.) Hertel & Rambold:
juvenile facultatively lichenicolous lichen on Rhizocarpon gr.
geographicum;instrata (Nyl.) Hertel & Rambold: facultatively
lichenicolous lichen on Aspicilia (and on Rhizocarpon?);
intrudens (H.Magn.) Hertel & Rambold: lichenicolous lichen
on Rhizocarpon;invadens Hafellner, Obermayer & Tretiach:
lichenicolous lichen on Sporastatia, becoming independent;
nigroleprosa (Vain.) Hertel & Rambold: juvenile facultatively
lichenicolous lichen on Rhizocarpon gr. geographicum;
verrucariicola (B.D.Ryan) K.Knudsen & Kocourk. [¼Proto-
parmelia ryaniana van den Boom, Sipman & Elix, nom.
illeg.]: lichenicolous lichen on Dimelaena radiata,Tephromela
and Verrucaria subdivisa, becoming independent.
Lit.: Hafellner et al. 2005, 2014; Hertel & Rambold 1987; Knudsen
et al. 2015; Rambold & Triebel 1992.
Family Parmeliaceae Zenker
Nesolechia A.Massal; type: N. oxyspora. Mol.: yes (T, L). All
lichenicolous species, except N. falcispora, often develop
subspherical to lobate or foliose gall-like structures that
may be regarded as an own lichenized thallus. A high
morphological plasticity, sometimes correlated with host
selection, has led to the description of many poorly
characterized taxa, most of which are not accepted here.
Two species of uncertain generic position (N. cetrariicola
and N. diversispora) are provisionally kept here.
Species: 7-0-0. cetrariicola (Linds.) Arnold [”Lecidea
cetrariicola Linds.]: on Cetraria;diversispora Gr´
elet: on
Lecanora [a poorly known species]; doerfeltii (Alstrup &
P.Scholz) Diederich, comb. nov. [”Phacopsis doerfeltii
Alstrup & P.Scholz]: on Arctoparmelia centrifuga;falcispora
(Triebel & Rambold) Diederich, comb. nov. [”Ph.
falcispora Triebel & Rambold; ¼Ph. falcispora var. sipmanii
Diederich & Triebel]: on Usnea;oxyspora (Tul.) A.Massal.
var. oxyspora [”Punctelia oxyspora (Tul.) Divakar,
Crespo, D.Hawksw. & Lumbsch; ¼Ph. oxyspora var. defecta
Triebel & Rambold; ¼Ph. menegazziae Triebel & Rambold;
¼N. rufa M¨
ull.Arg.]: on Parmeliaceae [populations with
reddish to dark brown apothecia and a hyaline to pale
brown hypothecium]; oxyspora var. fusca (Triebel &
Rambold) Diederich, comb. nov. [”N. fusca (Triebel &
Rambold) P´
erez-Ortega; ”Ph. oxyspora var. fusca Triebel &
Rambold; ¼Ph. australis Aptroot & Triebel]: on Xantho-
parmelia and other Parmeliaceae [populations with dark
brown to black, immersed to superficial apothecia and a
brown to dark brown hypothecium]; thallicola (A.Massal.)
A.Massal. [¼Abrothallus curreyi Linds.]: on Flavoparmelia
caperata [type of N. thallicola], Parmotrema [type of A.
curreyi], Cetrelia,Flavopunctelia and Hypotrachyna.
Diederich et al.: 2018 classification and checklist of lichenicolous fungi 371
Lit.: Aptroot & Triebel 2002; Divakar et al. 2015, 2017; Scholtz
1998; Spribille 2010; Triebel & Rambold 1988; Triebel et al. 1995,
Zhurbenko & Hafellner 1999.
Phacopsis Tul.; type: P. vulpina. Mol.: yes (T, L).
Species: 6-0-0. cephalodioides (Nyl.) Triebel & Rambold:
on Hypogymnia physodes;lethariellae Hafellner & Ram-
bold: on Lethariella intricata;oroarcticae Zhurb.: on
Brodoa oroarctica;prolificans (M¨
ull.Arg.) Triebel & Ram-
bold: on Platismatia interrupta;usneae C.W.Dodge: on
Usnea subgen. Neuropogon;vulpina Tul.: on Letharia.
Lit.: Divakar et al. 2015; Etayo & Sancho 2006; Hafellner 1987a;
Triebel & Rambold 1988; Triebel et al. 1995; Zhurbenko 2010b.
Protoparmelia M.Choisy; type: P. badia (Hoffm.) Hafellner
(lichenized). Mol.: yes (T, L).
Species: 0-7-0 (35). atriseda (Fr.) R.Sant. & V.Wirth:
lichen, juvenile lichenicolous on Rhizocarpon;cupreobadia
(Nyl.) Poelt: lichen, juvenile lichenicolous on Rhizocarpon;
hierrensis van den Boom & Ertz: lichenicolous lichen on
Pertusaria;leproloma (R.Sant.) Rambold & Poelt: lichen-
icolous lichen on Lecidea praenubila;loricata Poelt &
Vˇ
ezda: lichenicolous lichen on Lecanora umbrosa;phaeo-
nesos Poelt: lichenicolous lichen on Aspicilia myrinii;
placentiformis (J.Steiner) Poelt: lichen, juvenile lichen-
icolous on Bellemerea cupreoatra.
Lit.: Rambold & Triebel 1992; van den Boom & Ertz 2012.
Raesaenenia D.Hawksw., C.Boluda & H.Lindgren; type: R.
husskonenii. Mol.: yes (T, L).
Species: 1-0-0. huuskonenii (R¨
as¨
anen) D.Hawksw., C.Bo-
luda & H.Lindgr. [”Phacopsis huuskonenii R¨
as¨
anen; ”
Protousnea huuskonenii (R¨
as¨
anen) Divakar, Crespo &
Lumbsch]: on Bryoria.
Lit.: Divakar et al. 2015, 2017; Lindgren et al. 2015; Triebel &
Rambold 1988.
Family Pilocarpaceae Zahlbr.
Epicladonia D.Hawksw. s.l.; type: E. sandstedei. Mol.: yes (T, L).
The generic type belongs to Leotiomycetes. Both E. simplex
and E. stenospora are not related to the type, but represent a
distinct clade in the Pilocarpaceae. More studies are needed
before a new genus can be described for them, hence they
are provisionally placed here.
Species: 2-0-0. simplex D.Hawksw.: on Cladonia;steno-
spora (Harmand) D.Hawksw.: on Cladonia.
Lit.: Hawksworth 1981; Pino-Bodas et al. 2017.
Fellhanera Vˇ
ezda; type: F. fuscatula (M¨
ull.Arg.) Vˇ
ezda (liche-
nized). Mol.: yes.
Species: 0-1-0 (100). stictae Etayo: lichenicolous lichen on
Sticta.
Lit.: Etayo 2017.
Micarea Fr.; type: M. prasina Fr. (lichenized). Mol.: yes (T, L).
Species: 4-1-0 (100). inquinans (Tul.) Coppins: on Dibaeis
baeomyces; intrusa (Th.Fr.) Coppins & H.Kilias [”
Carbonea intrusa (Th.Fr.) Rambold & Triebel; ”Scolicio-
sporum intrusum (Th.Fr.) Hafellner]: lichen, frequently
lichenicolous on Rhizocarpon,Lecidea or other saxicolous
lichens; kemmleri Brackel: on Cladonia;stereocaulorum
Etayo & van den Boom: on Stereocaulon;usneae van den
Boom & Ertz: on Usnea.
Lit.: Brackel 2016; Coppins 2009; Etayo 2017; van den Boom & Ertz
2014.
Scutula Tul.; type: S. wallrothii, i.e. S. miliaris [¼Hollosia Gyeln.;
type: H. vertescens, i.e. S. epiblastematica;¼Karsteniomyces
D.Hawksw.; type: K. peltigerae (P.Karsten) D.Hawksw.; ¼
Libertiella Speg. & Roum.; type: L. malmedyensis Speg. &
Roum., i.e. probably S. epiblastematica]. Mol.: yes (T, L).
Species on non-Peltigerales hosts probably do not belong to
Scutula s.str.
Species: 14-2-0. cladoniicola Alstrup & D.Hawksw.: on
Cladonia trassii [does not belong to Scutula]; curvispora
(D.Hawksw. & Mia
˛dl.) Diederich, comb. nov. [”Libertiella
curvispora D.Hawksw. & Mia
˛dl.]: on Peltigera;dedicata
Triebel, Wedin & Rambold: lichenicolous lichen on Peltigera;
didymospora (D.Hawksw. & Mia
˛dl.) Diederich, comb. nov.
[”Libertiella didymospora D.Hawksw. & Mia
˛dl.]: on
Peltigera;epiblastematica (Wallr.) Rehm [¼S. aggregata
(Bagl. & Carestia) Rehm; ¼Hollosia vertescens Gyeln.; ¼?
Orbilia peltigerae (Fuckel) Sacc.; ¼?Libertiella malmedyensis
Speg. & Roum.]: on cyanobacterial Peltigera species; heerii
(Hepp) Trevis.: lichenicolous lichen on Peltigera;Libertiella
fennica Alstrup: on Peltigera;Libertiella leprariae Etayo &
Diederich: on Lepraria;leptogica (Nyl.) Zopf: on Polychi-
dium muscicola;Karsteniomyces llimonae Boqueras &
Diederich: on Parmelina quercina;miliaris (Wallr.) Trevis.
[¼S. wallrothii Tul., nom. illeg.; ¼Karsteniomyces peltigerae
(P.Karst.) D.Hawksw.; ¼Diplodina peltigerae Vouaux]: on
cyanobacterial Peltigera species; nephromatis (Speg.) Etayo:
on Nephroma antarcticum;Libertiella obscurior Gerber: on
Brodoa intestiniformis [a poorly known species]; pseudocy-
phellariae Etayo & Triebel: on Pseudocyphellaria;tuber-
culosa (Th.Fr.) Rehm [¼S. krempelhuberi K¨
orb.; ¼S.
solorinaria (Nyl.) P.Karst.; ¼S. solorinicola (Vain.) Rehm]:
on Solorina;Karsteniomyces tuberculosus Alstrup &
D.Hawksw.: on green-algal Peltigera species [probably
distinct from S. epiblastematica].
Lit.: Alstrup 1997; Alstrup & Hawksworth 1990; Boqueras &
Diederich 1993; Etayo & Diederich 1995; Etayo & Sancho 2008;
Etayo & Triebel 2010; Hawksworth 1980b, 1981; Hawksworth &
Mia
˛dlikowska 1997a; Pino-Bodas et al. 2017; Triebel et al. 1997;
Wedin et al. 2007.
Family Psoraceae Zahlbr.
Glyphopeltis Brusse; type: G. eburina, i.e. G. ligustica]. Mol.: yes
(T, L).
Species: 0-1-0. ligustica (B. de Lesd.) Timdal [¼G. eburina
Brusse]: lichen, juvenile parasite on Peltula.
Lit.: Ott & Scheidegger 1992; Rambold & Triebel 1992.
Psorula Gotth.Schneid.; type: P. rufonigra. Mol.: yes (T, L).
Species: 0-1-0 (3). rufonigra (Tuck.) Gotth.Schneid.:
lichenicolous lichen on Spilonema.
Lit.: Rambold & Triebel 1992.
Family Ramalinaceae C.Agardh.
Bacidia De Not.; type: B. rosella (Pers.) De Not. (lichenized).
Mol.: yes (T).
Species: 1-0-0 (230). killiasii (Hepp) D.Hawksw.: on
Peltigera.
Lit.: Rambold & Triebel 1992.
372 The Bryologist 121(3): 2018
Myrionora R.C.Harris; type: S. albidula (Willey) R.C.Harris
(lichenized). Mol.: no.
Species: 0-1-0 (2). pseudocyphellariae (Etayo) S.Ekman &
Palice [”Scoliciosporum pseudocyphellariae Etayo]: lichen-
icolous lichen on Pseudocyphellaria.
Lit.: Etayo & Sancho 2008; Palice et al. 2013.
Toninia A.Massal.; type: T. cinereovirens (Schaer.) A.Massal.
(lichenized). Mol.: yes (T, L).
Species: 11-3-0 (85). collematicola Timdal: on Collema;
episema (Nyl.) Timdal [¼?Scutula aspiciliae (M¨
ull.Arg.)
Rehm]: on Aspicilia;leptogii Timdal: on Collema and
Leptogium;nashii Timdal: on Lecidella asema (and Lecanora
gangaleoides?); nordlandica Th.Fr.: lichenicolous lichen on
Placynthium and other lichens with cyanobacteria; plumbina
(Anzi) Hafellner & Timdal: on Degelia;poeltiana
S.Y.Kondr., L.L}
ok¨
os&J.-S.Hur:onVerrucaria fuscella;
subfuscae (Arnold) Timdal: on Lecanora,rarelyonother
lichens; subdispersa (Hasse) K.Knudsen [¼T. talparum
Timdal]: on Lecania;subtalparum van den Boom: on
Lecania;tecta C.A.Morse & Ladd: on Dermatocarpon;ualae
Etayo: lichenicolous lichen on Pseudocyphellaria and Ne-
phroma antarcticum;verrucariae (Nyl.) Timdal: on Bagliet-
toa and Encephalographa elisae;verrucarioides (Nyl.)
Timdal: lichenicolous lichen on Placynthium.
Lit.: Etayo & Sancho 2008; Knudsen & Lendemer 2007; Knudsen &
Kocourkova
´2018; Kondratyuk et al. 2016b; Ladd & Morse 2018;
Rambold & Triebel 1992; Timdal 1992; van den Boom 2004.
Family Ramboldiaceae S.Stenroos, Mia
˛dl. & Lutzoni
Ramboldia Kantvilas & Elix; type: R. stuartii (Hampe) Kantvilas
& Elix (lichenized). Mol.: yes (T, L).
Species: 0-1-0 (30). insidiosa (Th.Fr.) Hafellner: lichen-
icolous lichen on Lecanora varia.
Lit.: Hafellner & T¨
urk 1995.
Family Scoliciosporaceae Hafellner
Scoliciosporum A.Massal.; type: S. holomelaeum (Fl¨
orke)
A.Massal. (lichenized). Mol.: no.
Species: 0-1-0 (13). vouauxii (B. de Lesd.) Hafellner:
lichenicolous lichen on Ramalina.
Lit.: Hafellner 2002, 2004b.
Family Stereocaulaceae Chevall.
Paralecia Brackel, Greiner, Perˇ
soh & Rambold; type: P.
pratorum. Mol.: yes (T, L).
Species: 1-0-0. pratorum Brackel, Greiner, Perˇ
soh &
Rambold: on Protoparmeliopsis muralis.
Lit.: Liu et al. 2015.
Family Tephromelataceae Hafellner [¼Mycoblasta-
ceae Hafellner]
Tephromela M.Choisy; type: T. atra (Huds.) Hafellner (liche-
nized). Mol.: yes (T).
Species: 3-1-0 (30). campestricola (Nyl.) Rambold &
Triebel: on Lecanora pseudistera and L. sp.; cerasina
(M¨
ull.Arg.) Rambold & Triebel: on Dirinaria;koliensis
(R¨
as.) Rambold & Triebel: on Lecanora argopholis;parasitica
Øvstedal & Søchting: lichenicolous lichen on Lecidella.
Lit.: Rambold 1993; Søchting et al. 2004.
Order Lecanorales genera incertae sedis
Corticiruptor Wedin & Hafellner; type: C. abelonae. Mol.: yes.
Species: 2-0-0. abeloneae (P.M.Jørg.) Wedin & Hafellner:
on Sticta;corallinus Etayo: on Pseudocyphellaria.
Lit.: Etayo & Sancho 2008; Wedin & Hafellner 1998.
Lichenosticta Zopf; type: L. podetiicola, i.e. L. alcicorniaria. Mol.:
yes (T, L).
Species: 5-0-0. alcicorniaria (Linds.) D.Hawksw. [¼L.
podetiicola Zopf]: on Cladonia;dombrovskae Zhurb.: on
Stereocaulon;hoegnabbae Zhurb. & Pino-Bodas: on Cladia;
jurgae Kukwa & Flakus: on Lecanora;lecanorae (Vouaux)
Brackel & Zhurb.: on Lecanora.
Lit.: Flakus & Kukwa 2012; Hawksworth 1981; Pino-Bodas et al.
2017; Zhurbenko 2010a; Zhurbenko & Pino-Bodas 2015; Zhur-
benko et al. 2015a.
Nimisiostella Calat., Barreno & O.E.Erikss.; type: N. lichenicola.
Mol.: yes.
Species: 1-0-0. lichenicola Calat., Barreno & O.E.Erikss.: on
Parmelia saxatilis.
Lit.: Calatayud et al. 1997.
Umbilithecium Etayo; type: U. pseudocyphellariae. Mol.: yes.
Species: 1-0-0. pseudocyphellariae Etayo: on Pseudocyphel-
laria.
Lit.: Etayo & Sancho 2008.
Umushamyces Etayo; type: U. kuturnus. Mol.: yes.
Species: 1-0-0. kuturnus Etayo: on Coccotrema cucurbitula.
Lit.: Etayo & Sancho 2008.
Order Lecideales Vain.
Family Lecideaceae Chevall.
Bellemerea Hafellner & Cl.Roux; type: B. alpina (Sommerf.)
Clauzade & Cl.Roux (lichenized). Mol.: yes (T).
Species: 0-1-0 (9). cupreoatra (Nyl.) Clauzade & Cl.Roux:
lichenicolous lichen on Buellia, becoming independent.
Lit.: Rambold & Triebel 1992.
Cecidonia Triebel & Rambold; type: C. umbonella. Mol.: yes (T,
L).
Species: 2-0-0. umbonella (Nyl.) Triebel & Rambold: on
Lecidea;xenophana (K¨
orb.) Triebel & Rambold [¼Lecidea
hymeneliicola Alstrup & D.Hawksw.]: on Porpidia.
Lit.: Triebel & Rambold 1988.
Immersaria Rambold & Pietschm.; type: I. athroocarpa. Mol.:
yes (L).
Species: 0-3-0 (8). athroocarpa (Ach.) Rambold & Pietschm.:
lichen, juvenile facultatively lichenicolous on Aspicila and
other lichens; cupreoatra (Nyl.) Calat. & Rambold: lichen,
juvenile lichenicolous on Buellia;usbekica (Hertel) Barbero,
Diederich et al.: 2018 classification and checklist of lichenicolous fungi 373
Nav.-Ros. & Cl.Roux: lichen, juvenile facultatively lichen-
icolous on Aspicila and other lichens.
Lit.: Rambold & Triebel 1992.
Lecidea Ach. s.l.; type: L. fuscoatra (L.) Ach. (lichenized). Mol.:
yes (T).
Species: 4-6-0 (100). acervata Stiz.: on Pseudoparmelia [a
poorly known taxon]; degeliana Hertel: lichenicolous lichen
on Acarospora;diexcipula D.Hawksw. & Alstrup: on
Lecanora polytropa;grummannii Hertel: lichenicolous lichen
on Acarospora;halacsyi J.Steiner: on Rhizocarpon geographi-
cum;leprosolimbata (Arnold) Poelt: lichenicolous lichen on
Bellemerea subcandida;percutiens Poelt: lichenicolous lichen
on Diploschistes scruposus;perforans Stiz.: on Parmotrema [a
poorly known taxon]; rapax Hertel: lichenicolous lichen on
Aspicilia and Bellemerea;verruca Poelt: lichenicolous lichen
on Aspicilia.
Lit.: Hafellner 2006; Rambold & Triebel 1992.
Poeltiaria Hertel; type: P. turgescens (K¨
orb.) Hertel (lichenized).
Mol.: yes.
Species: 0-1-0 (8). coromandelica (Zahlbr.) Rambold &
Hertel: lichen, juvenile lichenicolous on Parapordia lep-
tocarpa.
Lit.: Rambold & Triebel 1992.
Order Leprocaulales Lendemer & B.P.Hodk.
Family Leprocaulaceae Lendemer & B.P.Hodk.
Halecania M.Mayrhofer; type: H. alpivaga. Mol.: yes (T, L).
Species: 1-5-0 (22). alpivaga (Th.Fr.) M.Mayrhofer: lichen-
icolous lichen on Physcia and Placynthium;fuscopannariae
Etayo & van den Boom: on Fuscopannaria;giraltiae van
den Boom & Etayo: lichenicolous lichen on saxicolous
lichens; lobulata van den Boom & Elix: lichen, possibly
lichenicolous over Hymenelia;parasitica Aptroot &
K.H.Moon: lichenicolous lichen over Schaereria fuscociner-
ea, becoming independent; santessonii Andreev: lichen-
icolous lichen on Porpidia albocaerulescens.
Lit.: Andreev 2010; Aptroot & Moon 2015; M. Mayrhofer 1987; van
den Boom 2009; van den Boom & Elix 2005; van den Boom &
Etayo 2001.
Order Rhizocarpales Mia
˛dl. & Lutzoni s.l.
Family Rhizocarpaceae Hafellner
Epilichen Clem.; type: E. scabrosus. Mol.: no.
Species: 1-2-0. glaucinigellus (Nyl.) Hafellner: lichen-
icolous lichen on Baeomyces;scabrosus (Ach.) Clements:
lichenicolous lichen on Baeomyces;stellatus Triebel: on
Lecidea cf. tessellata.
Lit.: Hafellner 1978; Triebel 1989.
Rhizocarpon DC.; type: R. geographicum (L.) DC. (lichenized).
Mol.: yes (T, L).
Species: 7-32-0 (225). advenulum (Leight.) Hafellner &
Poelt: on Pertusaria flavicans and P. rupicola;asiaticum
(Poelt) Poelt: lichenicolous lichen on Sporastatia;aspiciliae
(R¨
as¨
anen) D.Hawksw. & V.Atienza: on Hymenelia;atrovir-
ellum (Nyl.) Zahlbr.: lichenicolous lichen on Aspicilia;
bryonthae Poelt: on Pertusaria bryontha;captans Poelt:
lichenicolous lichen on Aspicilia;cleptophilum Alstrup &
E.S.Hansen: lichenicolous lichen on Acarospora picea;
dahlii Øvstedal: lichenicolous lichen on the prothallus of
Rhizocarpon geographicum;destructans Alstrup: lichen-
icolous lichen on Lecanora polytropa;dimelaena Timdal:
lichenicolous lichen on Dimelaena oreina;dinothetes Hertel
& Leuckert: lichenicolous lichen on Protoparmelia;diplo-
schistidina McCune: lichenicolous lichen on Diploschistes
muscorum;effiguratum (Anzi) Th.Fr.: facultatively lichen-
icolous lichen on Pleopsidium flavum;epispilum (Nyl.)
Zahlbr. s.l.: lichenicolous lichen on saxicolous Pertusaria;
fratricida Poelt & Nimis: lichenicolous lichen on Rhizo-
carpon;fuegianum Poelt, nom. nud.: lichenicolous lichen
on Rhizocarpon geographicum;furax Poelt & V.Wirth:
lichenicolous lichen on Lecidea;inimicum Poelt & Vˇ
ezda:
lichenicolous lichen on Lecanora rupicola s.l.; intermediel-
lum R¨
as¨
anen: lichen, juvenile lichenicolous on other
lichens; kakurgon Poelt: lichenicolous lichen on Aspicilia;
kansuense H.Magn.: lichenicolous lichen on Aspicilia;
lusitanicum (Nyl.) Arnold: lichenicolous lichen on saxic-
olous Lepra;malenconianum (Llimona & Werner) Hafell-
ner & H.Mayrhofer: on Diploschistes;mosigiae Poelt &
Obermayer: lichenicolous lichen on Rimularia gibbosa;
narssaqense Alstrup: on Lecanora atromarginata;norvegi-
cum R¨
as¨
anen: lichen, juvenile lichenicolous on other
lichens, mainly Acarospora;ochrolechiae (Poelt & Nimis)
Hafellner: on Ochrolechia parella;parvum Runem.: lichen,
juvenile lichenicolous on Tremolecia atrata;pusillum
Runem.: lichenicolous lichen on Sporastatia;rapax V.Wirth
& Poelt: lichenicolous lichen on Acarospora,Aspicilia and
Rimularia;renneri Poelt: lichenicolous lichen on Dimelae-
na oreina;santessonii Timdal: lichenicolous lichen on
Tremolecia atrata;schedomyces Hafellner & Poelt: lichen-
icolous lichen on saxicolous Pertusaria;subtile Runem.:
lichen, juvenile lichenicolous on Tremolecia;superficiale
(Schaer.) Vain.: lichen, juvenile lichenicolous on Acarospora
oxytona;tephromelae Øvstedal: lichenicolous lichen on
Tephromela atra;trapeliicola Brand: lichenicolous lichen
on Trapelia;tungurahuae Etayo & Palice: on Gyalidea;
viridiatrum (Wulf.) K¨
orb.: lichen, juvenile often lichen-
icolous on Aspicilia,Lecidea,Tremolecia;vorax Poelt &
Hafellner: lichenicolous lichen on Ochrolechia parella and
Pertusaria.
Lit.: Alstrup & Hawksworth 1990; Etayo 2017; Feuerer & Timdal
2004; Hawksworth & Atienza 1994; Holtan-Hartwig & Timdal
1987; Lumbsch et al. 2011; Øvstedal et al. 2009; Poelt & Obermayer
1995; Poelt & Vˇ
ezda 1984; Rambold & Triebel 1992.
Order Teloschistales D.Hawksw. & O.E.Erikss.
Family Teloschistaceae Zahlbr.
Subfamily Caloplacoideae Arup, Søchting & Fr¨
od´
en
Caloplaca Th.Fr. s.l.; type: C. cerina (Hedw.) Th.Fr. (lichenized).
Mol.: yes (T, L). We also include here species belonging to
Caloplaca s.l. for which no molecular data are available.
Species: 0-26-0 (350). adelphoparasitica Nimis & Poelt:
lichenicolous lichen on Caloplaca and other lichens;
anchon-phoeniceon Poelt & Clauzade: lichenicolous lichen
on Aspicilia;cacuminum Poelt: lichen, juvenile facultatively
lichenicolous on saxicolous lichens; emilii Vondra
´k,
374 The Bryologist 121(3): 2018
Khodos., Cl.Roux & V.Wirth: lichen, juvenile lichenicolous
on saxicolous lichens; epierodens M.Bertrand & Cl.Roux:
lichenicolous lichen on Pyrenodesmia erodens;epithallina
Lynge: lichenicolous lichen on Acarospora,Aspicila,Dime-
laena,Lecanora,Melanelia,Pleopsidium,Psorinia,Rhizo-
carpon,Rhizoplaca,Stereocaulon,Tephromela,Umbilicaria;
epitoninia O.Breuss: lichenicolous lichen on Toninia
aromatica;erythrocarpa (Pers.) Zwackh: lichen, juvenile
facultatively lichenicolous on Aspicilia calcarea;furax Egea
& Llimona: lichen, juvenile lichenicolous on other lichens,
especially Aspicilia;grimmiae (Nyl.) H.Olivier: lichen-
icolous lichen on Candelariella;inconnexa (Nyl.) Zahlbr.
var. inconnexa: lichen, juvenile lichenicolous on Acaro-
spora,Aspicilia,Caloplaca;inconnexa var. nesodes Poelt &
Nimis: lichenicolous lichen on Aspicilia;infestans H.Magn.:
lichenicolous lichen on Aspicilia,Caloplaca s.l., Lecanora;
insularis Poelt: lichenicolous lichen on Aspicilia;interna
Poelt & Nimis: lichenicolous lichen on Aspicilia;isidiigera
Vˇ
ezda: lichenicolous lichen on saxicolous lichens; lecanorae
F.C.Seavey & J.L.Seavey: lichenicolous lichen on Lecanora
leprosa;magni-filii Poelt: lichenicolous lichen on Miriquid-
ica nigroleprosa;megalariicola Øvstedal: lichenicolous
lichen on Megalaria grossa;necator Clauzade & Poelt:
lichenicolous lichen on Aspicilia;nubigena (Kremp.) Dalla
Torre & Sarnth.: lichen, facultatively lichenicolous on
Clauzadea immersa;peludella (Nyl.) Hasse: lichen, juvenile
facultatively lichenicolous on saxicolous lichens; placidia
(A.Massa.) J.Steiner: lichen, juvenile facultatively lichen-
icolous on Pyrenodesmia chalybaea;psoricida Hansen, Poelt
& Søchting: lichenicolous lichen on Psora rubiformis;
rinodinae-albae Poelt & Nimis: lichenicolous lichen on
Rinodina alba;tenuatula (Nyl.) Zahlbr.: lichenicolous
lichen on Bagliettoa calciseda.
Lit.: Breuss 1990; Fryday & Øvstedal 2012; Nimis et al. 1994;
Rambold & Triebel 1992; Roux 2014; Seavey & Seavey 2012;
Wetmore 1999.
Gyalolechia A.Massal.; type: G. aurea (Schaer.) A.Massal
(lichenized) [¼Fulgensia A.Massal. & De Not.; type: F.
fulgens (Sw.) Elenkin (lichenized)]. Mol.: yes (T).
Species: 0-1-0 (30). epiplacynthium (Etayo) Diederich,
comb. nov. [”Fulgensia epiplacynthium Etayo]: lichen-
icolous lichen on Placynthium filiforme.
Lit.: Etayo 2010c.
Variospora Arup, Søchting & Fr¨
od´
en; type: V. velana. Mol.: yes
(T, L).
Species: 0-1-0 (12). velana (A.Massal.) Arup, Søchting &
Fr¨
od´
en: lichen, juvenile facultatively lichenicolous on
Pyrenodesmia variabilis.
Lit.: Roux 2014.
Subfamily Teloschistoideae Arup, Søchting &
Fr¨
od´
en
Catenarina Søchting, Søgaard, Arup, Elvebakk & Elix; type: C.
desolata. Mol.: yes (T, L).
Species: 0-1-0 (3). desolata Søchting, Søgaard & Elvebakk:
lichenicolous lichen on Aspicilia.
Lit.: Søchting et al. 2014.
Sirenophila Arup, Søchting&Fr¨
od´
en;type:S. gintarasii(S.Y.Kondr.
&K
¨
arnefelt) Arup, Fr¨
od´
en & Søchting. Mol.: yes (T).
Species: 0-1-0 (8). ovis-atra Søchting, Søgaard & Sancho:
lichenicolous lichen on Hydropunctaria.
Lit.: Søchting et al. 2016.
Subfamily Xanthorioideae Arup, Søchting & Fr¨
od´
en
Flavoplaca Arup, Søchting & Fr¨
od´
en; type: F. citrina (Hoffm.)
Arup, Søchting & Fr¨
od´
en. Mol.: yes (T).
Species: 0-4-0 (27). coronata (K¨
orb.) J.Steiner: lichen, juvenile
facultativelylichenicolouson saxicolouslichens;microthallina
(Wedd.) Zahlbr.: lichen, facultatively lichenicolous on saxic-
olous lichens, especially Hydropunctaria maura;oasis (A.Mas-
sal.) Szatala: lichen, facultatively lichenicolous on saxicolous
lichens, especially Bagliettoa calciseda;polycarpa (A.Massal.)
Arup, Fr¨
od´
en & Søchting: lichenicolous lichen on Bagliettoa.
Lit.: Arup et al. 2013, Roux 2014.
Pachypeltis Søchting, Arup & Fr¨
od´
en [a hemihomonym of
Pachypeltis Signoret (Heteroptera)]; type: P. castellana
(R¨
as¨
anen) Søchting, Fr¨
od´
en & Arup (lichenized). Mol.:
yes (T, L).
Species: 0-1-0 (4). invadens (Lynge) Søchting, Fr¨
od´
en &
Arup: lichenicolous lichen on Lecanora,Physcia caesia.
Lit.: Arup et al. 2013.
Shackletonia Søchting, Fr¨
od´
en & Arup; type: S. hertelii
(Søchting, Øvstedal & Sancho) Søchting, Fr¨
od´
en & Arup
(lichenized). Mol.: yes (T, L).
Species: 1-2-0 (5). buelliae (Olech & Søchting) Søchting:
on Buellia;Caloplaca psoromatis Olech & Søchting:
lichenicolous lichen on Psoroma and Pannaria hookeri
[provisionally placed here]; sauronii (Søchting & Øvstedal)
Søchting: lichenicolous lichen on Buellia.
Lit.: Garrido-Benavent et al. 2016; Olech & Søchting 1993; Søchting
et al. 2004.
Subclass OSTROPOMYCETIDAE Reeb, Lutzoni &
Cl.Roux
Order Ostropales Nannf.
Family Gomphillaceae Walt.Watson
Corticifraga D.Hawksw. & R.Sant.; type: C. peltigerae. Mol.: yes
(T, L).
Species: 7-0-0. chugachiana Zhurb.: on Lobaria oregana;
fuckelii (Rehm) D.Hawksw. & R.Sant.: on Peltigera;
fusispora Zhurb.: on Peltigera aphthosa and P. leucophlebia;
peltigerae (Fuckel) D.Hawksw. & R.Sant.: on Peltigera,
exceptionally on Solorina and Pseudocyphellaria;pseudocy-
phellariae Etayo: on Pseudocyphellaria;santessonii Zhurb.
& Zavarzin: on Lobaria and Nephroma;scrobiculatae
P´
erez-Ortega: on Lobarina scrobiculata.
Lit.: Etayo & Sancho 2008; Hawksworth & Santesson 1990; Pino-
Bodas et al. 2017; Spribille et al. 2010; Zhurbenko 2007c;
Zhurbenko 2009b.
Gyalideopsis Vˇ
ezda; type: G. peruviana Vˇ
ezda (lichenized).
Mol.: yes.
Species: 4-0-1 (91). cochlearifer L¨
ucking & S´
erus.: on
foliicolous Gomphillaceae; epithallina R.L¨
ucking: on foliic-
Diederich et al.: 2018 classification and checklist of lichenicolous fungi 375
olous Gyalideopsis vulgaris;parvula Kalb & Vˇ
ezda: on
foliicolous Pilocarpaceae; stipitata Kalb & Vˇ
ezda: facultative-
ly lichenicolous on Dictyonema;usneicola Etayo: on Usnea.
Lit.: Etayo 2010b, 2017; L¨
ucking 1997; L¨
ucking & S´
erusiaux 1998.
Paragyalideopsis Etayo; type: P. floridae. Mol.: no.
Species: 4-0-0. breussii Etayo: on Hypotrachyna;floridae
(Etayo & Diederich) Etayo: on Parmotrema;minuta Etayo: on
Trypethelium;stereocaulicola (Etayo) Etayo: on Stereocaulon.
Lit.: Etayo 2017; Etayo & Diederich 2001.
Taitaia Suija, Kaasalainen, Kirika & Rikkinen; type: T. aurea.
Mol.: yes (T, L).
Species: 1-0-0. aurea Suija, Kaasalainen & Rikkinen: on
Crocodia cf. clathrata.
Lit.: Suija et al. 2018b.
Family Graphidaceae Dumort.
Subfamily Graphidoideae Rivas Plata, L¨
ucking &
Lumbsch
Diploschistes Norman; type: D. scruposus (Schreb.) Norman
(lichenized). Mol.: yes (T, L).
Species: 0-4-0 (31). bisporus (Bagl.) J.Steiner: lichen,
facultatively lichenicolous on Rhizocarpon geographicum s.l.;
caesioplumbeus (Nyl.) Vainio: lichen, juvenile lichenicolous
on Lecanora gangaleoides;euganeus (A.Massal.) J.Steiner:
lichen, facultatively lichenicolous on Ochrolechia parella;
muscorum (Scop.) R.Sant.: lichenicolous on Cladonia (rarely
Stereocaulon) when young, later an independent lichen.
Lit.: Lumbsch 1989.
Family Protothelenellaceae Vˇ
ezda, H.Mayrhofer &
Poelt
Protothelenella R¨
as¨
anen; type: P. reducta (Th.Fr.) R¨
as¨
anen, i.e.
P. sphinctrinoidella (Nyl.) H.Mayrhofer & Poelt (liche-
nized). Mol.: yes (T, L).
Species: 3-0-3 (11). croceae (Bagl. & Car.) Hafellner &
H.Mayrhofer: on Solorina crocea, more rarely Peltigera;
leucothelia (Nyl.) H.Mayrhofer & Poelt: facultatively
lichenicolous on moribund lichens, especially Cladonia;
pseudocyphellariae Etayo: on Pseudocyphellaria;santesso-
nii H.Mayrhofer: on Cladonia;sphinctrinoidella (Nyl.)
H.Mayrhofer & Poelt: facultatively lichenicolous on Clado-
nia,Peltigera and Stereocaulon;sphinctrinoides (Nyl.)
H.Mayrhofer & Poelt: facultatively lichenicolous on Pelti-
gera and Solorina.
Lit.: Etayo & Sancho 2008; H. Mayrhofer 1987; Pino-Bodas et al.
2017.
Family Stictidaceae Fr.
Cryptodiscus Corda; type: C. pallidus [¼Lettauia D.Hawksw. &
R.Sant.; type: L. cladoniicola]. Mol.: yes (T, L).
Species: 3-0-0 (9). cladoniicola (D.Hawksw. & R.Sant.)
Pino-Bodas, Zhurb. & S.Stenroos [”Lettauia cladoniicola
D.Hawksw. & R.Sant.]: on Cladonia;epicladonia Zhurb. &
Pino-Bodas: on Cladonia;galaninae Zhurb. & Pino-Bodas:
on Cladonia.
Lit.: Hawksworth & Santesson 1990; Pino-Bodas et al. 2017.
Lettauia D.Hawksw. & R.Sant. s.l.; type: L. cladoniicola., i.e.
Cryptodiscus cladoniicola. Mol.: no. The generic type has
recently been shown to belong to Cryptodiscus.The
phylogenetic affinities of the three other species described
in Lettauia are currently unknown.
Species: 3-0-0. hypotrachynae Etayo: on Hypotrachyna;
santessonii Ihlen & Tønsberg: on Loxospora pustulata;
usneae Etayo: on Usnea.
Lit.: Etayo 2002, 2017; Ihlen & Tønsberg 1996.
Nanostictis M.S.Christ.; type: N. peltigerae. Mol.: no.
Species: 9-0-0. caucasica Zhurb.: on Parmelia sulcata;
christiansenii Etayo: on Lobaria pulmonaria;confusa Etayo:
on Hypotrachyna;heterodermiae Etayo & Sipman: on
Heterodermia;nephromatis Etayo: on Nephroma antarcti-
cum;peltigerae M.S.Christ.: on Peltigera;pluriseptata Etayo:
on Lobariella gr. pallida;pseudocyphellariae Sherwood: on
Pseudocyphellaria;stictae Etayo: on Sticta.
Lit.: Etayo 2002a, 2017; Etayo & Diederich 1996b; Etayo & Sancho
2008; Zhurbenko 2017.
Sphaeropezia Sacc.; type: S. alpina Sacc. [¼Lethariicola
Grummann; type: L. sipei]. Mol.: yes (L).
Species: 16-0-0 (19). bryoriae (Diederich & Etayo) Baloch
& Wedin [”Odontotrema bryoriae Diederich & Etayo]: on
Bryoria;cucularis (Norman) Baloch & Wedin: on Parme-
liopsis;figulina (Norman) Baloch & Wedin: on Lecanora
polytropa;intermedia (Diederich, Zhurb. & Etayo) Baloch
& Wedin [”O. intermedium Diederich, Zhurb. & Etayo]:
on Thamnolia;japewiae (Zhurb. & Diederich) Baloch &
Wedin [”O. japewiae Zhurb. & Diederich]: on Japewia
torno¨
ensis;lecanorae (Diederich & G.Marson) Baloch &
Wedin [”O. lecanorae Diederich & G.Marson]: on
Lecanora;melaneliae (Diederich & Zhurb.) Baloch, Gilen-
stam & Wedin [”O. melaneliae Diederich & Zhurb.]: on
Melanelia hepatizon and Melanohalea olivacea;mycoblasti
Diederich, Baloch & Wedin: on Mycoblastus;navarinoi
(Etayo) Baloch & Wedin [”O. navarinoi Etayo]: on
Micarea magellanica;ochrolechiae (Diederich, Holien &
Zhurb.) Baloch & Wedin [”O. ochrolechiae Diederich,
Holien & Zhurb.]: on Ochrolechia;pertusariae (Etayo,
Diederich & Coppins) Baloch & Wedin [”O. pertusariae
Etayo, Diederich & Coppins]: on Pertusaria hymenea;
rhizocarpicola (Zhurb., Diederich & Himelbrant) Baloch
&Wedin[”O. rhizocarpicola Zhurb., Diederich &
Himelbr.]: on Rhizocarpon gr. geographicum;santessonii
(Zhurb., Etayo & Diederich) Baloch & Wedin [”O.
santessonii Zhurb., Etayo & Diederich]: on Thamnolia;sipei
(Grummann) Baloch & Wedin [”Lethariicola sipei
Grummann]: on Letharia;Odontotrema stereocaulicola
Zhurb.: on Stereocaulon [provisionally placed here; prob-
ably does not belong to Sphaeropezia]; thamnoliae (Zhurb.,
Diederich & Etayo) Baloch & Wedin [”O. thamnoliae
Zhurb., Diederich & Etayo]: on Thamnolia.
Lit.: Baloch et al. 2013; Diederich et al. 2002b; Etayo & Sancho
2008; Zhurbenko 2009b, 2010a.
Stictis Pers.; type: S. radiata (L.) Pers. Mol.: yes (T). The only
lichenicolous species probably does not belong to Stictis.
Species: 1-0-0 (4). cladoniae (Rehm) Sacc.: on Cladonia.
Lit.: Pino-Bodas et al. 2017; Sherwood 1977.
376 The Bryologist 121(3): 2018
Order Ostropales genera incertae sedis
Aabaarnia Diederich; type: A. siphulicola. Mol.: no.
Species: 1-0-0. siphulicola Diederich: on Siphula decum-
bens.
Lit.: Diederich 2014.
Biazrovia Zhurb. & Etayo; type: B. stereocaulicola. Mol.: no.
Species: 1-0-0. stereocaulicola Zhurb. & Etayo: on Stereo-
caulon.
Lit.: Zhurbenko & Etayo 2013.
Normanogalla Diederich; type: N. cribriformis. Mol.: no.
Species: 1-0-0. cribriformis (Norman) Diederich [”
Unguiculariopsis cribriformis (Norman) Alstrup &
D.Hawksw.]: on Lepra dactylina and Pertusaria oculata.
Lit.: Alstrup & Hawksworth 1990; Diederich 2014.
Paralethariicola Calat., Etayo & Diederich; type: P. aspiciliae.
Mol.: no.
Species: 1-0-0. aspiciliae Calat., Etayo & Diederich: on
Aspicilia.
Lit.: Calatayud et al. 2001a.
Spirographa Zahlbr.; type: S. fusisporella [”Graphinella Zahlbr.;
type: G. fusisporella;¼Pleospilis Clem.; type: P. vermiformis
(Leight.) Clem.; ¼Spilomela (Sacc.) Keissl.; type: S.
vermifera (Leight.) Keissl.; ¼Spirographomyces Cif. &
Tomas.; type: S. spiralis Cif. & Tomas.]. Mol.: no. No
new combinations are proposed here, as a taxonomic
revision of the genus is still missing.
Species: 5-0-0. Lecidea ascaridiella Nyl. [”Pleospilis
ascaridiella (Nyl.) D.Hawksw.]: on ?Porpidia;fusisporella
(Nyl.) Zahlbr. [”Graphinella fusisporella (Nyl.) Zahlbr.;
¼?Opegrapha spiralis M¨
ull.Arg.]: on Graphidaceae;
longispora Etayo: on Sticta;usneae Flakus, Kukwa &
Etayo: on Usnea;Melaspilea vermiformis Leight. [¼
Melaspilea vermifera Leight., nom. illeg.]: on Lepra
albescens.
Lit.: Diederich 2004; Etayo 2017; Flakus & Kukwa 2012; Sherwood-
Pike 1987.
Order Pertusariales D.Hawksw. & O.E.Erikss.
Family Megasporaceae Lumbsch, Feige & K.Schmitz
Aspicilia A.Massal.; type: A. cinerea (L.) K¨
orb. (lichenized).
Mol.: yes (T).
Species: 0-1-0 (200). parasitica B. de Lesd.: lichenicolous
lichen on saxicolous, crustose lichens.
Lit.: Rambold & Triebel 1992.
Family Microcaliciaceae Tibell
Microcalicium Vain.; type: M. disseminatum. Mol.: yes (T, L).
Species: 3-0-0 (4). arenarium (A.Massal.) Tibell: on
Psilolechia;conversum Tibell: on Calicium and Chaenothe-
ca;disseminatum (Ach.) Vain.: on Calicium and Chaeno-
theca, rarely on free living algae.
Lit.: Clauzade et al. 1989; Hawksworth 1981; Prieto et al. 2013;
Tibell 1978.
Order Sarrameanales B.P.Hodk. & Lendemer
Family Schaereriaceae Hafellner
Schaereria Th.Fr.; type: S. lugubris (Fr.) K¨
orb. [¼Hafellnera
Houmeau & Cl.Roux; type: H. parasemella]. Mol.: yes.
Species: 0-1-0 (16). parasemella (Nyl.) Lumbsch [”
Hafellnera parasemella (Nyl.) Houmeau & Cl.Roux]:
lichenicolous lichen on Biatora vernalis.
Lit.: Houmeau & Roux 1984; Rambold & Triebel 1992.
Order Trapeliales B.P.Hodk. & Lendemer
Family Xylographaceae Tuck.
Lambiella Hertel; type: L. psephota (Tuck.) Hertel (lichenized).
Mol.: yes (T, L).
Species: 0-2-0 (12). furvella (Mudd) M.Westb. & Resl:
lichenicolous lichen on various lichens; insularis (Nyl.)
T.Sprib.: lichenicolous lichen on Lecanora rupicola.
Lit.: Rambold & Triebel 1992.
Subclass Ostropomycetidae familia incertae sedis
Family Epigloeaceae Zahlbr.
Epigloea Zukal; type: E. bactrospora Zukal (algicolous). Mol.:
yes.
Species: 1-0-0 (13). urosperma D¨
obbeler: on Placynthiella.
Lit.: D¨
obbeler 1994.
Subclass Ostropomycetidae genus incertae sedis
Bachmanniomyces D.Hawksw.; type: B. uncialicola [¼Phaeo-
pyxis Rambold & Triebel; type: P. punctum]. Mol.: yes (T,
L).
Species: 6-0-0 (7). australis (Rambold & Triebel) Diederich
& Pino-Bodas, comb. nov. [”Phaeopyxis australis Rambold
&Triebel]:onParaporpidia;muscigenae (Alstrup &
E.S.Hansen) Diederich & Pino-Bodas, comb. nov. [”P.
muscigenae Alstrup & E.S.Hansen]: on Physconia;pseudo-
cyphellariae Etayo: on Pseudocyphellaria;santessonii
Etayo: on Sticta;punctum (A.Massal.) Diederich & Pino-
Bodas, comb. nov. [”P. punctum (A.Massal.) Rambold,
Triebel & Coppins; ¼B. uncialicola (Zopf) D.Hawksw.]: on
Cladonia;varius (Coppins, Rambold & Triebel) Diederich
& Pino-Bodas, comb. nov. [”P. varia Coppins, Rambold &
Triebel]: on Trapeliopsis.
Lit.: Alstrup & Hansen 2001; Etayo 2010b; Etayo & Sancho 2008;
Hawksworth 1981; Pino-Bodas et al. 2017; Rambold & Triebel
1990; Suija et al. 2015b.
Class Lecanoromycetes familia incertae sedis
Family Arthrorhaphidaceae Poelt & Hafellner
Arthrorhaphis Th.Fr.; type: A. flavovirescens, i.e. A. citrinella.
Mol.: yes (T, L).
Diederich et al.: 2018 classification and checklist of lichenicolous fungi 377
Species: 6-3-0 (13). aeruginosa R.Sant. & Tønsb.: on
Cladonia;alpina (Schaer.) R.Sant.: juvenile parasite on
Baeomyces rufus, later an independent lichen; arctoparme-
liae Kocourk. & van den Boom: on Arctoparmelia incurva;
citrinella (Ach.) Poelt [¼A. flavovirescens (A.Massal.)
Th.Fr.]: juvenile facultatively parasite on Baeomyces rufus,
later or from the beginning an independent lichen; grisea
Th.Fr.: on Baeomyces, later occasionally an independent
lichen; muddii Obermayer: on Dibaeis baeomyces;olivaceae
R.Sant. & Tønsb.: on Melanohalea olivacea;phyllobaeis
Etayo & Palice: on Phyllobaeis;vaccillans Th.Fr.: juvenile
parasite on Baeomyces, later an independent lichen.
Lit.: Etayo 2017; Kocourkova
´& van den Boom 2005; Obermayer
1994; Santesson & Tønsberg 1994.
Class Lecanoromycetes genus incertae sedis
Piccolia A.Massal.; type: P. crocea (Spreng.) A.Massal. (liche-
nized). Mol.: no.
Species: 0-1-0 (7). nannaria (Tuck.) Lendemer & Beeching:
lichen, at least facultatively lichenicolous on Pyrrhospora
varians.
Lit.: Lendemer & Harris 2014.
Class LEOTIOMYCETES O.E.Erikss. & Winka
Order Helotiales Korf & Lizon
Family Cordieritidaceae (Sacc.) Sacc.
Diplolaeviopsis Giral & D.Hawksw.; type: D. ranula. Mol.: yes
(T, L).
Species: 3-0-0. japonica Zhurb. & Diederich: on Gyalolechia
flavorubescens;ranula Giralt & D.Hawksw.: on Lecanora
strobilina and L. strobilinoides;symmictae Diederich &
Coppins: on Lecanora symmicta.
Lit.: Diederich & Coppins 2014; Etayo et al. 2015; Giralt &
Hawksworth 1991; Suija et al. 2015b; Zhurbenko et al. 2015a.
Llimoniella Hafellner & Nav.-Ros.; type: L. scabridula. Mol.: yes
(L).
Species: 19-0-0. acarosporicola (Kocourk. & K.Knudsen)
Diederich & Ertz: on Acarospora socialis;adnata Hafellner
& Nav.-Ros.: on Placidium;bergeriana Etayo: on Punctelia;
catapyrenii Zhurb., Kukwa & Flakus: on Catapyrenium;
cinnabarinae P´
erez-Ortega, Etayo & T.Sprib.: on Rambol-
dia cinnabarina;fuscatae Hafellner & Obermayer: on
Acarospora fuscata;gregorellae (Alstrup & D.Hawksw.)
Triebel & Hafellner: on Gregorella humida;groenlandiae
(Alstrup & D.Hawksw.) Triebel & Hafellner: on Athallia
and Flavoplaca, possibly on Gyalolechia and Xanthocarpia;
heppiae (Nav.-Ros., Hladun & Llimona) Diederich & Ertz:
on Heppia;parmotrematis Etayo: on Parmotrema;pertu-
sariae Diederich & Etayo: on Lepra ophthalmiza and
Pertusaria glaucomela;phaeophysciae Diederich, Ertz &
Etayo: on Phaeophyscia;placomaroneae Etayo & Diederich:
on Placomaronea candelarioides;placopsidis Diederich &
Fryday: on Placopsis;pyrenulae Diederich & Etayo: on
Pyrenula;ramalinae (M¨
ull.Arg.) Etayo & Diederich: on
Ramalina celastri and R. unilateralis;scabridula (M¨
ull.Arg.)
Nav.-Ros. & Hafellner: on Acarospora nodulosa and A.
placodiiformis;terricola (Arnold) M.Schultz, Diederich &
Ertz: on Leptogium byssinum;vinosa (Holien & Triebel)
Diederich & Etayo: on Ochrolechia and Lepra cf. borealis.
Lit.: Alstrup & Hawksworth 1990; Diederich & Etayo 2000;
Diederich et al. 2010b; Etayo 2010b, 2017; P´
erez-Ortega et al.
2011b; Suija et al. 2015b; Zhurbenko 2013a.
Macroskyttea Etayo, Flakus, Suija & Kukwa; type: M. parmo-
trematis. Mol.: yes (T, L).
Species: 1-0-0. parmotrematis Etayo, Flakus & Kukwa: on
Parmotrema.
Lit.: Etayo et al. 2015.
Rhymbocarpus Zopf; type: R. punctiformis, i.e. R. geographici.
Mol.: yes (L).
Species: 12-0-0. aggregatus Etayo & Diederich: on Buellia
griseovirens;boomii Etayo & Diederich: on Dirina cerato-
niae;cruciatus (Sherwood, D.Hawksw. & Coppins) Etayo
& Diederich: on Diploicia canescens;ericetorum (K¨
orb.)
Etayo, Diederich & Ertz: on Dibaeis baeomyces;fuscoatrae
(Hafellner) Diederich & Etayo: on Lecidea fuscoatra;
geographici (J.Steiner) Vouaux [”R. punctiformis Zopf]:
on Rhizocarpon;makarovae Diederich & Etayo: on
Porpidia;neglectus (Vain.) Diederich & Etayo: on the
Lepraria neglecta group; pertusariae Diederich, Zhurb. &
Etayo: on Lepra panyrga;pubescens (Etayo & Diederich)
Diederich & Etayo: on Lepraria;roccellae (Etayo, Paz-
Berm ´
udez & Diederich) Etayo, Diederich & Ertz: on
Roccella;stereocaulorum (Alstrup & D.Hawksw.) Etayo &
Diederich: on Stereocaulon.
Lit.: Diederich & Etayo 2000; Diederich et al. 2010b; Etayo &
Diederich 2011; Suija et al. 2015b.
Skyttea Sherwood, D.Hawksw. & Coppins; type: S. nitschkei.
Mol.: yes (T, L).
Species: 30-0-0. anziae Etayo & Diederich: on Anzia;
buelliae Sherwood, D.Hawksw. & Coppins: on Amandinea
punctata;bumyoungsungii S.Y.Kondr. & J.-S.Hur: on
unidentified epilithic thallus (Gyalidea or Porpidia?); caesii
Diederich & Etayo: on Mycoblastus (incl. M. caesius);
carboneae Diederich & Etayo: on Carbonea montevidensis;
cismonicae Hafellner: on Loxospora cismonica;dacampiae
Zhurb.: on Dacampia hookeri;elachistophora (Nyl.) Sher-
wood & D.Hawksw.: on cf. Tylothallia biformigera;fusispora
Sherwood, D.Hawksw. & Coppins: on Ochrolechia;gossypi-
na Etayo: on Phlyctis argena;gregaria Sherwood, D.Hawksw.
&Coppins:onViolella fucata;hawksworthii Diederich: on
Verrucaria;heterochroae Nav.-Ros. & Mu˜
niz: on Pertusaria
heterochroa;insignis Driscoll, S.R.Clayden & R.C.Harris: on
Lecanora insignis;lecanorae Diederich & Etayo: on Lecanora;
mayrhoferi Diederich & Etayo: on Pertusaria melanospora;
megalosporae Etayo & Diederich: on Megalospora tuber-
culosa;nitschkei (K¨
orb.) Sherwood, D.Hawksw. & Coppins:
on Thelotrema lepadinum;ochrolechiae Zhurb.: on Ochro-
lechia trochophora;pertusariicola Diederich & Etayo: on
corticolous Pertusaria;pyrenulae Diederich, Etayo &
Coppins: on Pyrenula;radiatilis (Tuck.) R.Sant., Etayo &
Diederich: on cf. Loxospora pustulata;recognita Etayo &
Diederich: on epiphytic crustose lichens; richardsonii Iturr.
& D.Hawksw.: on an unidentified lichen; spinosa
D.Hawksw. & Coppins: on Verrucaria nigrescens;tavaresae
R.Sant., Etayo & Diederich: on Pyrrhospora cf. quernea;
tephromelarum Kalb & Hafellner: on Tephromela;thelo-
trematis Diederich & Etayo: on Thelotrema lepadinum;
violacea Etayo: on Tephromela;viridis D.Hawksw. &
Coppins: on Micarea cf. denigrata.
378 The Bryologist 121(3): 2018
Lit.: Diederich & Etayo 2000, 2004; Driscoll et al. 2016; Etayo
2002a, 2010c, 2017; Etayo & Sancho 2008; Hafellner 2000;
Kondratyuk et al. 2018; Navarro-Rosin`
es & Mu˜
niz 2009; Suija et
al. 2015b; Zhurbenko et al. 2015a.
Thamnogalla D.Hawksw.; type: T. crombiei. Mol.: yes (T, L).
Species: 1-0-0. crombiei (Mudd) D.Hawksw. [¼?Didymo-
cyrtis thamnoliicola Y.Joshi, R.Bajpai & Upreti]: on
Thamnolia.
Lit.: Hoffmann & Hafellner 2000; Joshi et al. 2016c; Lord et al.
2013; Suija et al. 2015b.
Unguiculariopsis Rehm; type: U. ilicincola (Berk. & Broome)
Rehm (saprotrophic). Mol.: yes (L).
Species: 15-0-0 (25). acrocordiae (Diederich) Diederich &
Etayo: on Acrocordia gemmata;ahtii D.Hawksw., D.J.Gal-
loway & S.Y.Kondr.: on Pseudocyphellaria;damingshanica
W.Y.Zhuang: on unidentified lichen on bamboo; helmutii
S.Y.Kondr., L.L}
ok¨
os&J.-S.Hur:onRinodina aff. sophor-
odes;lesdainii (Vouaux) Etayo & Diederich: on Lecanora
saligna;lettaui (Grummann) Coppins: on Evernia prunas-
tri;lobariellum S.Y.Kondr. & D.J.Galloway: on Lobaria
and Lobariella;lucaniae Brackel: on Lecidella elaeochroma;
macrocarpa Etayo: on Pseudocyphellaria;manriquei
Etayo: on Lobaria pulmonaria;Llimoniella muralicola
Halıcı:onProtoparmeliopsis muralis [belongs to Unguicu-
lariopsis s.l.]; peltigericola Etayo: on Peltigera;refractiva
(Coppins) Coppins: on Bilimbia;thallophila (P.Karst)
W.Y.Zhuang: on Lecanora;triregia S.Y.Kondr. & D.J.Gal-
loway: on Sticta.
Lit.: Brackel 2011a; Coppins 1988; Diederich & Etayo 2000; Etayo
2017; Etayo & Diederich 1996b; Etayo & Sancho 2008; Kondratyuk
et al. 1994; Kondratyuk & Galloway 1995a; Kondratyuk & Galloway
1995b; Kondratyuk et al. 2016b; Suija et al. 2015b; Zhuang 2000.
Family Helicogoniaceae Baral
Geltingia Alstrup & D.Hawksw.; type: G. associata. Mol.: yes (T, L).
Species: 1-0-0. associata (Th.Fr.) Alstrup & D.Hawksw.: on
Ochrolechia, more rarely on Lepra dactylina and Thamnolia.
Lit.: Alstrup & Hawksworth 1990; Diederich et al. 2010b; Suija et al.
2015b.
Family Helotiaceae Rehm
Bryoscyphus Spooner; type: B. conocephali (Boyd) Spooner.
Mol.: yes.
Species: 1-0-0 (19). lichenicola Alstrup & M.S.Cole: on
Cladonia cf. pyxidata.
Lit.: Alstrup & Cole 1998.
Family Hyaloscyphaceae Nannf.
Hyalopeziza Fuckel; type: H. patula (Pers.) Fuckel. Mol.: yes.
Species: 2-0-0 (15). heterodermiae Etayo: on Heterodermia;
rapax Huhtinen: on Pseudocyphellaria.
Lit.: Etayo 2017; Huhtinen et al. 2008.
Hyphodiscus Kirschst.; type: H. gregarius Kirschst. Mol.: yes (T,
L).
Species: 1-0-0 (16). ucrainicus (S.Y.Kondr.) Suija, Tsur-
ykau & Diederich [”Pezizella ucrainica S.Y.Kondr.]: on
Cladonia.
Lit.: Kondratyuk & Galloway 1995a; Oksner 2010; Suija et al. 2018;
Zhurbenko & Pino-Bodas 2017.
Polydesmia Boud.; type: P. pruinosa (Berk. & Broome) Boud.
Mol.: yes (T).
Species: 1-0-0 (7). lichenis Huhtinen & R.Sant.: on
Peltigera.
Lit.: Huhtinen & Santesson 1997.
Protounguicularia Raitv. & R.Gala
´n; type: P. brevicapitata Raitv.
& R.Gala
´n. Mol.: no.
Species: 4-0-0 (10). fasciculata (Etayo) Etayo: on Ramalina
stenospora;hispidula (Etayo) Etayo: on Psoroma hispidu-
lum;nephromatis (Zhurb. & Zavarzin) Huhtinen,
D.Hawksw. & Ihlen: on Nephroma;usneae Etayo: on
Usnea.
Lit.: Etayo 2010b, 2017; Etayo & Triebel 2010; Huhtinen et al. 2008;
Zhurbenko 2007a.
Family Lachnaceae Raitv.
Solenopezia Sacc.; type: S. solenia Sacc. Mol.: no.
Species: 1-0-0 (7). tetraspora P.Henn.: on Physcia integrata
[a poorly known species].
Lit.: Clauzade et al. 1989; Vouaux 1914.
Family Mollisiaceae Rehm
Mollisia (Fr.) P.Karst.; type: M. cinerea (Batsch) P.Karst. Mol.:
yes (T).
Species: 1-0-0 (130). collematis Boud. [”Pyrenopeziza
collematis (Boud.) Boud.]: on Collema [a poorly known
species that does not belong to Mollisia].
Lit.: Clauzade et al. 1989; Vouaux 1914.
Niptera Fr.; type: N. lacustris (Fr.) Fr. Mol.: no. Both lichenicolous
speciesare poorlyknownandpossiblydo not belong to Niptera.
Species: 2-0-0 (10). lichenicola (Speg.) Sacc.: on Flavopar-
melia caperata;microscopica (P.Crouan & H. Crouan)
Vouaux: on Placynthiella uliginosa.
Lit.: Clauzade et al. 1989; Vouaux 1914.
Trimmatostroma Corda s.l.; type: T. salicis Corda. Mol.: yes
(T). Asexual, hyphomycetous. Probably polyphyletic, the
lichenicolous taxa unlikely to be closely related to the type
species.
Species: 8-0-1 (30). arctoparmeliae Brackel & Schiefel-
bein: on Arctoparmelia incurva;cetrariae Brackel: on
Cetraria islandica;dendrographae Diederich, Ertz,
U.Braun & Heuchert: on Dendrographa;glebarum
Brackel: on Flavoparmelia caperata;hierrense Diederich
&Ertz:onArthonia endlicheri;lecanoricola Diederich,
Etayo, U.Braun & Heuchert: on Lecanora;quercicola
Diederich, U.Braun & Heuchert: facultatively lichen-
icolous on sterile, corticolous, crustose lichens; umbil-
icariicola Heuchert & U.Braun: on Umbilicaria;
varicellariae Heuchert & U.Braun: on Varicellaria
rhodocarpa.
Lit.: Brackel 2015; Diederich et al. 2010a; Heuchert & Braun 2014;
Schiefelbein et al. 2017; Zhurbenko & Brackel 2013.
Diederich et al.: 2018 classification and checklist of lichenicolous fungi 379
Family Pezizellaceae Velen.
Calycina Gray [a hemihomonym of Calycina Gregory, Echino-
zoa]; type: C. herbarum (Pers.) Gray [¼Pezizella Fuckel;
type: P. pulchella Fuckel]. Mol.: yes (T, L). Two species are
provisionally placed here pending a phylogenetic reassess-
ment of the lichenicolous taxa.
Species: 3-0-0 (30). alstrupii Suija & Motiej¯
unaite
˙:on
Lobaria pulmonaria;Pezizella epithallina (W.Phillips &
Plowr.) Sacc.: on Peltigera;Pezizella stictae Etayo: on
Sticta.
Lit.: Etayo 2002a; Hawksworth 1980b; Suija & Motiej ¯
unaite
˙2017;
Suija et al. 2015b.
Chalara (Corda) Rabenh.; type: C. fusidioides (Corda) Rabenh.
Mol.: yes (T).
Species: 3-0-1 (100). lichenicola M.S.Christ.: on Cladonia;
lobariae Etayo: on Lobaria pulmonaria; aff. microspora
(Cordes) Hughes: facultatively lichenicolous on Cladonia
coniocraea;pseudocyphellariae Etayo: on Pseudocyphellaria
faveolata.
Lit.: Brackel 2015; Christiansen 1993; Etayo & Diederich 1996b;
Etayo & Sancho 2008.
Family Sclerotiniaceae Whetzel
Pycnopeziza W.L.White & Whetzel; type: P. sympodialis
W.L.White & Whetzel [”Acarosporium Buba
´k & Vleugel;
type: A. sympodiale Buba
´k & Vleugel]. Mol.: yes (T).
Species: 1-0-0 (5). Acarosporium lichenicola Ihlen &
Tønsberg: on Icmadophila [provisionally placed here,
possibly not belong to Pycnopeziza].
Lit.: Ihlen 1998.
Order Helotiales genera incertae sedis
Brackelia Zhurb.; type: B. lunkei. Mol.: no.
Species: 1-0-0. lunkei Zhurb.: on Cladonia.
Lit.: Zhurbenko & Pino-Bodas 2017.
Cecidioskyttea Etayo; type: C. osorioi. Mol.: no.
Species: 1-0-0. osorioi Etayo: on Parmotrema fistulatum.
Lit.: Etayo & Osorio 2004.
Chloroepilichen Etayo; type: C. rolfii. Mol.: no.
Species: 1-0-0. rolfii Etayo: on Yoshimuriella cf. peltigera.
Lit.: Etayo 2010b.
Echinodiscus Etayo & Diederich [a younger synonym of
Echinodiscus Mann, diatoms, and of Echinodiscus Bentham,
Fabaceae; a hemihomonym of Echinodiscus Leske, Echino-
zoa]; type: E. lesdainii. Mol.: no.
Species: 2-0-0. kozhevnikovii Zhurb.: on Cetraria;lesdainii
(Vouaux) Etayo & Diederich: on Lecania.
Lit.: Etayo & Diederich 2000; Zhurbenko 2009a.
Pleoscutula Vouaux; type: P. arsenii. Mol.: no.
Species: 2-0-0. arsenii Vouaux: on Heterodermia;hypo-
trachynae Etayo & Diederich: on Hypotrachyna.
Lit.: Etayo 2002a; Hafellner 1982b.
Skyttella D.Hawksw. & R.Sant.; type: S. mulleri. Mol.: no.
Species: 2-0-0. mulleri (Willey) D.Hawksw. & R.Sant.: on
Peltigera;stictae Etayo: on Sticta.
Lit.: Etayo 2017; Hawksworth & Santesson 1988.
Strossmayeria Schulzer; type: S. rackii Schulzer [¼Pseudospiropes
M.B.Ellis; type: P. nodosus (Sacc.) M.B.Ellis]. Mol.: yes.
Molecular data of both facultatively lichenicolous species are
needed to determine if they belong to Strossmayeria.
Species: 0-0-2 (16). Pseudospiropes costaricensis
(E.F.Morris) de Hoog & Arx: facultatively lichenicolous
on Crocodia aurata;P. longipilus (Corda) Hol.-Jech.:
facultatively lichenicolous on Lepraria.
Lit.: Braun et al. 2008; Johnston et al. 2014.
Order Phacidiales H¨
ohn.
Order Phacidiales genus incertae sedis
Epithamnolia Zhurb.; type: E. karatyginii. Mol.: yes (L). Asexual.
Species: 7-0-0. atrolazulina (Etayo) Diederich, comb. nov.
[”Hainesia atrolazulina Etayo]: on Hypotrachyna;brevicla-
doniae (Diederich & van den Boom) Diederich & Suija [”
H. brevicladoniae Diederich & van den Boom]: on Cladonia;
karatyginii Zhurb.: on Thamnolia;longicladoniae (Die-
derich & van den Boom) Diederich & Suija [”H.
longicladoniae Diederich & van den Boom]: on Cladonia;
pertusariae (Etayo & Diederich) Diederich & Suija [”H.
pertusariae Etayo & Diederich; ¼H. bryonorae Zhurb.; ¼H.
peltigerae Zhurb. & Davydov]: on Bryonora,Cladonia,
Peltigera and Pertusaria;rangiferinae E.Zimm., Diederich
&Suija:onCladonia rangiferina;xanthoriae (Brackel)
Diederich & Suija [”H. xanthoriae Brackel; ¼H.
aeruginascens Brackel; ¼H. lecanorae Brackel]: on Cande-
laria,Hypogymnia,Lecanora,Melanohalea,Parmelia,Phaeo-
physcia,Physcia,Platismatia,Polycauliona,Protoparmeliopsis,
Pseudevernia,Punctelia,Rusavskia and Xanthoria.
Lit.: Brackel 2009, 2014; Diederich & van den Boom 2013; Etayo
2017; Etayo & Diederich 1996a; Suija et al. 2018b; Zhurbenko 2012,
2013b; Zhurbenko & Brackel 2013.
Class Leotiomycetes genus incertae sedis
Epicladonia D.Hawksw. s.str.; type: E. sandstedei. Mol.: yes (T,
L). Asexual, pycnidial. Two further species, E. simplex and
E. stenospora, are treated within Epicladonia s.l. under
Pilocarpaceae, based on phylogenetic results.
Species: 3-0-0. heterodermiae Y.Joshi, Falswal & R.Joshi:
on Heterodermia;lapponica Ihlen: on Pleopsidium chlor-
ophanum;sandstedei (Zopf) D.Hawksw.: on Cladonia.
Lit.: Hawksworth 1981; Ihlen & Wedin 2005; Joshi et al. 2017b;
Pino-Bodas et al. 2017.
Class LABOULBENIOMYCETES Engl.
Order Laboulbeniales Lindau s.l.
Order Laboulbeniales genus incertae sedis
Gliocephalis Matr.; type: G. hyalina Matr. Mol.: yes (T).
Species: 1-0-0 (2). pulchella (Penz. & Sacc.) D.Hawksw.: on
Sticta.
Lit.: Hawksworth 1979a.
380 The Bryologist 121(3): 2018
Class LICHINOMYCETES Reeb, Lutzoni & Cl.Roux
Order Lichinales Henssen & B¨
udel
Family Lichinaceae Nyl.
Lichinodium Nyl.; type: L. sirosiphoideum. Mol.: no.
Species: 0-1-0 (4). sirosiphoideum Nyl.: lichen, juvenile
parasite of Parmelia s.l.
Lit.: Rambold & Triebel 1992.
Phylliscum Nyl.; type: P. demangeonii. Mol.: yes (T, L).
Species: 0-1-0 (8). demangeonii (Moug. & Mont.) Nyl.:
lichen, juvenile parasite on Ochrolechia frigida.
Lit.: Rambold & Triebel 1992.
Class SORDARIOMYCETES O.E.Erikss. & Winka
Subclass DIAPORTHOMYCETIDAE Senan., Maha-
rachch. & K.D.Hyde
Subclass Diaporthomycetidae familiae incertae sedis
Family Sporidesmiaceae Fr.
Sporidesmium Link; type: S. atrum Link. Mol.: yes.
Species: 3-0-0 (330). bacidiicola Alstrup: on Bacidia
rubella;lichenicola Iturr., D.Hawksw. & J.L.Crane: on
Leptogium; usneae Etayo: on Usnea.
Lit.: Alstrup (1991); Etayo 2017; Iturriaga et al. (2008).
Family Trichosphaeriaceae G.Winter
Trichosphaeria Fuckel; type: T. pilosa (Pers.) Fuckel. Mol.: yes
(T).
Species: 1-0-0 (20). lichenum P.Karst. & Har.: on Peltigera
[a poorly known species].
Lit.: Clauzade et al. 1989; Hawksworth 1980b.
Subclass HYPOCREOMYCETIDAE O.E.Erikss. &
Winka
Order Coronophorales Nannf.
Family Nitschkiaceae (Fitzp.) Nannf.
Acanthonitschkea Speg.; type: A. argentinensis Speg. Mol.: yes
(T).
Species: 1-0-0 (10). peltigericola (Alstrup & Olech)
O.E.Erikss. & R.Sant. [”Hystrix peltigericola Alstrup &
Olech]: on Peltigera.
Lit.: Alstrup & Olech 1993.
LasiosphaeriopsisD.Hawksw. & Sivan.; type:L. salisburyi. Mol.:no.
Species: 7-0-0. cephalodiorum (Rostr.) Alstrup: on Pla-
copsis;christiansenii Alstrup & D.Hawksw. [¼?L. super-
sparsa]: on Porpidia tuberculosa;lecanorae P´
erez-Ortega &
Halıcı:onLecanora polytropa;pilophori Zhurb. & Triebel:
on Pilophorus;salisburyi D.Hawksw. & Sivan.: on Peltigera;
stereocaulicola (Linds.) O.E.Erikss. & R.Sant.: on Stereo-
caulon;supersparsa (Zopf) Triebel: on Porpidia.
Lit.: Alstrup et al. 1994; Alstrup & Hawksworth 1990; Eriksson &
Santesson 1986; Hawksworth 1980b; P´
erez-Ortega & Halıcı2008;
Triebel 1989; Zhurbenko & Triebel 2005.
Rhagadostoma K¨
orb.; type: R. corrugatum, i.e. R. lichenicola.
Mol.: no.
Species: 7-0-0. boleae Nav.-Ros. & Hladun: on Lecania;
brevisporum (Nav.-Ros.& Hladun) Nav.-Ros.: onNephroma,
Peltigera and Solorina;collematum Etayo & Nav.-Ros.: on
Lathagrium auriforme;lichenicola (De Not.) Keissl. [¼R.
corrugatum K¨
orb.]: on Solorina;pannariae Etayo: on Pan-
naria;rugosum Nav.-Ros. & Hladun: on Verrucaria nigrescens;
verrucariarum Nav.-Ros. & Hladun: on Verrucaria.
Lit.: Etayo & Sancho 2008; Navarro-Rosin´
es & Hladun 1994;
Navarro-Rosin´
es et al. 1999.
Rhagadostomella Etayo; type: R. gregaria. Mol.: no.
Species: 1-0-0. gregaria Etayo: on Sticta.
Lit.: Etayo 2002a.
Order Hypocreales Lindau
Family Bionectriaceae Samuels & Rossman
Acremonium Link; type: A. alternatum Link. Mol.: yes (T, L).
Asexual, hyphomycetous.
Species: 4-0-6 (150). antarcticum (Speg.) D.Hawksw.: on
various lichens, facultatively lichenicolous; bavaricum
Brackel: on Melanelixia glabratula;hypholomatis (Boedijn)
D.Hawksw.: on Physcia stellaris, facultatively lichenicolous
[the lichenicolous populations are possible the asexual stage
of Pronectria echinulata Lowen]; lichenicola W.Gams: on
lichens and other substrata; persicinum (Nicot) W.Gams:
isolated from lichens, facultatively lichenicolous; pertusar-
iae Brackel & Etayo: on Lepra and Pertusaria;psychrophi-
lum M¨
oller & Gams: isolated from lichens, probably
lichenicolous; spegazzinii D.Hawksw.: on Leptogium;
strictum W.Gams: facultatively lichenicolous; subeffusum
(Ellis & Galw.) Etayo & Brackel: on Physcia millegrana and
possibly other lichens.
Lit.: Brackel et al. 2012; Diederich 1989; Diederich & Braun 2009;
Hawksworth 1979a; Lowen 1995.
Globonectria Etayo; type: G. cochensis. Mol.: no.
Species: 1-0-0. cochensis Etayo: on Lobaria pallida.
Lit.: Etayo 2002a.
Nectriopsis Maire; type: N. violacea (Fr.) Maire. Mol.: yes (T, L).
Species: 24-0-1 (70). albida Etayo: on Sticta;cariosae
Brackel & D.G.Zimmermann: on Cladonia;cladoniicola
M.S.Cole & D.Hawksw.: on Cladonia;collematis Diederich:
on Collema;curtiseta Etayo: on Sticta;frangospora
Brackel: on Lepra albescens and Pertusaria rubefacta;
gangwondoensis S.Y.Kondr., L.L}
ok¨
os & J.-S.Hur: on
Phaeophyscia adiastola;guamuesii Etayo: on Sticta;hetero-
dermiae Etayo: on Heterodermia;hirta Etayo: on various
lichens; indigens (Arnold) Diederich & Schroers: on
perithecioid lichens, often near water; lecanodes (Ces.)
Diederich & Schroers: on cyanobacterial macrolichens,
Diederich et al.: 2018 classification and checklist of lichenicolous fungi 381
mainly Lobaria,Peltigera and Pseudocyphellaria, also on
Collema and Leptogium;leptogii Diederich: on Leptogium;
lichenophila (Speg.) Etayo: on Heterodermia;melongenoi-
dea Etayo: on Gomphillus;micareae Diederich, van den
Boom & Ernst: on Micarea viridileprosa;mindoensis
(Petrak) (Petr.) Samuels: facultatively lichenicolous on
Coenogonium;peruviana Etayo & R.Sant.: on Cora and
Coccocarpia;physciicola D.Hawksw. & Earl.-Benn.: on
Physcia stellaris;porinicola Samuels: on foliicolous Porina;
silvaustralis Etayo: on Pseudocyphellaria;tatrensis (Alstr-
up) Lisicka
´& Alstrup: on Thelidium;verseghyklarae
S.Y.Kondr., L.L}
ok¨
os & J.-S.Hur: on Verseghya klarae;
vinosa Etayo: on Usnea;vivida Etayo & Sipman: on
unidentified crustose lichen.
Lit.: Alstrup 1996; Aptroot et al. 1997; Brackel 2010a; Brackel &
Zimmermann 2012; Cole & Hawksworth 2001; Earland-Bennett et
al. 2006; Etayo 2002a, 2010a, 2017; Etayo & Sancho 2008;
Kondratyuk et al. 2016a, 2018; Matzer 1996; Samuels 1988;
S´
erusiaux et al. 1999.
Ovicuculispora Etayo; type: O. parmeliae. Mol.: yes (T, L).
Species: 2-0-0. macrospora Etayo: on sterile saxicolous
crust (Gomphillaceae?); parmeliae (Berk. & Curt.) Etayo:
on various lichens.
Lit.: Etayo 2010b.
Paranectria Sacc.; type: P. affinis. Mol.: no.
Species: 4-0-0. affinis (Grev.) Sacc.: on Ephebe pubescens;
alstrupii Zhurb.: on various lichens; oropensis (Rabenh.)
D.Hawksw. & Piroz.: on various lichens; superba
D.Hawksw.: on Peltigera.
Lit.: Clauzade et al. 1989; Hawksworth 1982a; Rossman et al. 1999;
Zhurbenko 2009d.
Pronectria Clem.; type: P. lichenicola, i.e. P. robergei. Mol.: yes
(T, L).
Species: 43-0-1. algicola Vondra
´k, Naumovich & Khodos.:
on terricolous microbial crusts possibly associated with
Sarcosagium or Steinia;angulospora Etayo: on Hetero-
dermia;biglobosa Etayo: on Hypotrachyna;caloplacae
Khodos., Vondra
´k & Naumovich: on Flavoplaca austro-
citrina;casaresii Etayo: on Evernia prunastri;collematis
Etayo & Brackel: on Lathagrium;dealbans (M¨
ull.Arg.)
Etayo & Breuss: on Endocarpon;dillmaniae Zhurb.: on
Catapyrenium cinereum;diplococca Kocourk., Khodos.,
Naumovich, Vondra
´k & Motiej.: on Collema;echinulata
Lowen: on Phaeophyscia,Physcia and Physconia;erythri-
nella (Nyl.) Lowen: on Peltigera;fragmospora Etayo: on
Usnea;hymeniicola Etayo: on Collema;invisibilis Etayo:
on Nephroma antarcticum;japonica Zhurb., Tadome & Y.
Ohmura: on Ochrolechia;lecideicola Zhurb.: on Lecidea
ramulosa;leptogii Etayo: on Leptogium;microspora Etayo:
on Lobaria pallida;minuta Motiej. & Kukwa: on Cladonia
subg. Cladina;neofissuriprodiens F.Berger & E.Zimm.: on
Lobaria;occulta Etayo: on Usnea;oligospora Lowen &
Rogersen var. oligospora:onPunctelia;oligospora var.
octospora Etayo: on Punctelia;parmotrematis Etayo: on
Parmotrema;pedemontana Brackel: on Lathagrium auri-
forme;pertusariicola Lowen: on Lepra and Pertusaria;
pilosa Etayo & L ´
opez Silanes: on Collema;pycnidioidea
Etayo: on Heterodermia;rhizocarpicola Brackel: on Rhizo-
carpon geographicum;robergei (Mont. & Desm.) Lowen [¼
P. lichenicola (Ces.) Clem.; ¼Illosporium carneum Fr.]: on
Peltigera and Solorina;rolfiana Etayo: on Solorina;rose-
opunctata Etayo: on Hypotrachyna;santessonii (Lowen &
D.Hawksw.) Lowen: on Anaptychia runcinata;sticticola
Etayo: on Sticta;tenacis (Vouaux) Lowen: on Collema;
tenuispora (D.Hawksw.) Lowen: on Peltigera;terrestris
Lowen & Diederich: on Thrombium epigaeum;tibellii
Zhurb.: on Cladonia;tincta (Fuckel) Lowen: on Anaptychia
ciliaris;toniniae Etayo: on Toninia sedifolia;verrucariae
(Vouaux) Lowen: on Verrucaria;walkerorum Zhurb.: on
Ochrolechia;xanthoriae Lowen & Diederich: on Xanthoria;
zhurbenkoi Brackel: on Peltigera praetextata.
Lit.: Berger & Zimmermann 2016; Brackel 2013a, 2013b, 2015;
Brackel & Etayo 2010; Etayo 1998, 2001, 2002a, 2010b, 2010c,
2017; Etayo & Breuss 1996; Etayo & Diederich 1996b; Etayo &
Sancho 2008; Etayo & Triebel 2010; Hawksworth 1979a;
Khodosovts ev 201 2; L ´
opez de Silanes et al. 2009; Lowen 1995;
Lowen & Diederich 1990; Lowen & Hawksworth 1986; Mo-
tiej¯
unaite
˙& Kukwa 2008; Rossman et al. 1999; Sikaroodi et al.
2001; Zhurbenko 2007b, 2009d; Zhurbenko & Alstrup 2004;
Zhurbenko et al. 2005, 2018.
Trichonectria Kirschst.; type: T. aculeata Kirschst. Mol.: yes.
Species: 15-0-0 (19). anisospora (Lowen) van den Boom &
Diederich [¼Acremonium pedatum Lowen]: on Hypogymnia;
apiculata Etayo: on Usnea;australis Etayo: on Usnea;
calopadiicola Farkas & Flakus: on foliicolous Calopadia cf.
puiggari;cladoniicola (Alstrup & Svane) Alstrup: on
Cladonia;furcatosetosa Brackel: on necrosed Melanelixia,
Parmelia,Physcia and Candelariella;hirta (A.Bolxam) Petch:
on corticolous lichens; hypotrachynae Etayo: on Hypotrachy-
na;intermedia Etayo: on Parmotrema;leptogiicola Etayo: on
Leptogium;pertusariae Etayo & van den Boom: on Pertusaria;
pyrenaica Gardiennet & Lechat: on Usnea;rubefaciens (Ellis
& Everh.) Diederich & Schroers [”Nectriopsis rubefaciens
(Ellis & Everh.) M.S.Cole & D.Hawksw.; ¼Acremonium
rhabdosporum W.Gams]: on Parmeliaceae; setadpressa Etayo:
on Lobaria;usneicola Etayo: on Usnea.
Lit.: Alstrup & Svane 1998; Brackel 2014; Etayo 2001, 2002a, 2017;
Etayo & Sancho 2008; Etayo & van den Boom 2005; Farkas &
Flakus 2016; Gardiennet & Lechat 2012; Hawksworth 1978b;
Rossman et al. 1999; S´
erusiaux et al. 1999.
Family Clavicipitaceae (Lindau) O.E.Erikss.
Neobarya Lowen; type: N. parasitica (fungicolous). Mol.: yes (T).
Species: 4-0-0 (10). ciliaris Etayo: on Heterodermia;
darwiniana Etayo: on Nephroma antarcticum;lichenophila
(Ferd. & Winge) Lowen & Samuels: on Cladonia;peltigerae
Lowen, Boqueras & G ´
omez-Bolea: on Peltigera.
Lit.: Candoussau et al. 2007; Etayo 2002a; Etayo & Sancho 2008.
Family Cordycipitaceae G.H.Sung, J.M.Sung, Hy-
wel-Jones & Spatafora
Syspastospora P.F.Cannon & D.Hawksw.; type: S. parasitica
(Tul.) P.F.Cannon & D.Hawksw. Mol.: yes (T).
Species: 1-0-0 (4). cladoniae Etayo: on Cladonia foliacea.
Lit.: Etayo 2008.
Family Hypocreaceae De Not.
Hypomyces (Fr.) Tul. & C.Tul.; type: H. lactifluorum (Schwein.)
Tul. & C.Tul. Mol.: yes (T, L).
Species: 1-0-0 (150). peltigericola Lechat & Gardiennet: on
Peltigera canina.
Lit.: Lechat et al. 2017.
382 The Bryologist 121(3): 2018
Lichenobarya Etayo, Diederich & Lawrey; type: L. usneae. Mol.:
yes (T, L).
Species: 1-0-0. usneae (Etayo) Etayo, Diederich & Lawrey:
on Usnea.
Lit.: Etayo 2000a; Lawrey et al. 2015.
Family Nectriaceae Tul. & C.Tul.
Fusarium Link; type: F. roseum Link. Mol.: yes. Asexual,
hyphomycetous.
Species: 2-0-0 (120). pallens Link: on Myriolecis hagenii;
peltigerae Westend.: on Peltigera.
Lit.: Hawksworth 1979a.
Nectria (Fr.) Fr. s.l. [a hemihomonym of Nectria Gray,
Asterozoa]; type: N. cinnabarina (Tode) Fr. Mol.: yes (T).
Species: 2-0-1 (29). brutia Diederich & Puntillo: on
Collema [does not belong to Nectria s.str.]; byssophila
Rossman [possibly related to Lichenobarya or Neobarya]:
on macrolichens; paraguayensis Speg.: facultatively lichen-
icolous on lichens.
Lit.: Diederich & Puntillo 1995; Etayo & Sancho 2008; Hirooka et
al. 2012.
Xenonectriella Weese; type: X. lutescens. Mol.: no.
Species: 18-0-0. aurantiaca Etayo: on Pseudocyphellaria;
calabrica Brackel & Puntillo: on Pertusaria pertusa;
coppinsiana Etayo: on Yoshimuriella subdissecta;dirinar-
iae Etayo & van den Boom: on Dirinaria;fissuriprodiens
(Etayo) Etayo: on Lobaria;humilis Etayo: on Pseudocy-
phellaria;leptaleae (J.Steiner) Rossman & Lowen: on
Phaeophyscia,Physcia and Physconia;leptaleoides (Etayo)
Etayo: on Leptogium;lutescens (Rehm) Weese: on
Peltigera and Solorina;ornamentata (D.Hawksw.) Ross-
man: on Peltigera;protopannariae (Zhurb.) Brackel: on
Protopannaria pezizoides;rosea Etayo: on Pannaria;
rugulatispora Etayo: on Lobariella;septemseptata (Etayo)
Etayo & van den Boom: on Melanelixia and Melanohalea;
streimannii (S.Y.Kondr., Coppins & D.J.Galloway) Ross-
man: on Lobaria and Sticta;subimperspicua (Speg.) Etayo
var. subimperspicua:onParmelia and Punctelia,also
reported from Physcia;subimperspicua var. degenerans
Etayo: on Parmotrema;vertebrata Etayo: on Heteroder-
mia.
Lit.: Brackel 2014; Brackel & Puntillo 2016; Etayo 2002a, 2017;
Etayo & Sancho 2008; Etayo & Triebel 2010; Etayo & van den
Boom 2013; Rossman et al. 1999; van den Boom & Etayo
2014.
Family Niessliaceae Kirschst.
Atronectria Etayo; type: A. magellanica. Mol.: no.
Species: 1-0-0. magellanica Etayo: on Pseudocyphellaria and
Nephroma.
Lit.: Etayo & Sancho 2008.
Melanopsamma Niessl.; type: M. pomiformis (Pers.) Sacc. Mol.:
yes (T).
Species: 1-0-0 (40). lettauiana (Keissl.) Vouaux: on
saxicolous, crustose lichens [a poorly known species].
Lit.: Clauzade et al. 1989.
Niesslia Auersw.; type: N. chaetomium (Corda) Auersw. [¼
Nitschkiopsis Nannf. & R.Sant.; type: N. stictarum]. Mol.:
yes (L).
Species: 17-0-0 (43). cladoniicola D.Hawksw. & W.Gams:
on Cladia and Cladonia [anamorph: Monocillium sp.];
coenogonii van den Boom: on Coenogonium;echinoides
Etayo, Flakus & Kukwa: on Erioderma;evae Etayo: on
Erioderma;globospora Etayo: on Cora;keissleri Zhurb. [¼
Echinothecium cladoniae Keissl., nom. nud.]: on Cladia and
Cladonia;kununguakii Alstrup & E.S.Hansen: on Arthro-
rhaphis alpina;lobariae Etayo & Diederich: on Lobaria;
peltigericola (D.Hawksw.) Etayo [”Wentiomyces peltiger-
icola D.Hawksw.; ”Raciborskiomyces peltigericola
(D.Hawksw.) M.E.Barr]: on Peltigera;pseudocyphellariae
Etayo & Diederich: on Pseudocyphellaria;robusta Tretiach:
on Tephromela grumosa;schizospora Etayo: on Hypotra-
chyna;stictarum (Nannf. & R.Sant.) R.Sant. & Tretiach
subsp. stictarum [”Nitschkiopsis stictarum Nannf. &
R.Sant.]: on Sticta;stictarum subsp. nuda Etayo: on
Lobariella;tatjanae (S.Y.Kondr.) Etayo: on Pseudocyphel-
laria;tetrahedrospora Etayo: on Dichosporidium [possibly
belongs to Trichosphaerella, fide Lendemer et al. 2016];
yaganae Etayo: on Nephroma antarcticum.
Lit.: Alstrup & Hansen 2001; Etayo 2000, 2002a, 2017; Etayo &
Diederich 1996b; Etayo & Sancho 2008; Etayo et al. 2013a;
Hawksworth 1975, 1979a; Tretiach 2002; van den Boom et al. 2017;
Zhurbenko & Pino-Bodas 2017.
Trichosphaerella E.Bommer, M.Rousseau & Sacc.; type: T.
decipiens E.Bommer, M.Rousseau & Sacc. Mol.: yes.
Species: 1-0-0 (4). buckii R.C.Harris & Lendemer: on
Punctelia rudecta.
Lit.: Lendemer et al. 2016.
Order Hypocreales genera incertae sedis
Cephalosporiopsis Peyronel; type: none. Mol.: no. Asexual,
hyphomycetous. A genus of uncertain application.
Species: 0-0-1 (10). epiparasitaster Alstrup: on the lichen-
icolous fungus Chaenothecopsis parasitaster.
Lit.: Alstrup 1997.
Chondronectria Etayo, Flakus & Kukwa; type: C. eriodermati-
cola. Mol.: no.
Species: 1-0-0. eriodermaticola Etayo, Flakus & Kukwa: on
Erioderma.
Lit.: Etayo 2017.
Cylindronectria Etayo; type: C. cyanobactericola. Mol.: no.
Species: 0-0-1. cyanobactericola Etayo: on epilichenic
cyanobacteria, doubtfully lichenicolous.
Lit.: Etayo 2017.
Illosporiopsis D.Hawksw.; type: I. christiansenii. Mol.: yes (T,
L). Asexual, sporodochial.
Species: 1-0-0. christiansenii (B.L.Brady & D.Hawksw.)
D.Hawksw.: on various corticolous lichens.
Lit.: Lowen et al. 1986; Sikaroodi et al. 2001.
Leptobarya Etayo; type: L. nigra. Mol.: no.
Species: 2-0-0. auranticarpa Etayo: on Leptogium;nigra
Etayo: on Leptogium.
Lit.: Etayo 2017.
Diederich et al.: 2018 classification and checklist of lichenicolous fungi 383
Roselliniella Vain.; type: R. nephromatis. Mol.: yes (L).
Species: 19-0-0. africana Diederich: on Usnea;atlantica
Matzer & Hafellner: on Xanthoparmelia,Hypotrachyna
and Platismatia;cladoniae (Anzi) Matzer & Hafellner: on
Cladonia;coccocarpiae (Pat.) Matzer & R.Sant.: on
Degelia;epiphylla Matzer & Hafellner: on foliicolous
Sporopodium;eriodermicola Matzer & R.Sant.: on Erio-
derma tomentosum;euparmeliicola Millanes &
D.Hawksw.: on Parmelia s.str.; heterodeae Matzer &
Hafellner: on Cladia muelleri;lecideae Darmostuk,
Khodos. & Naumovich: on Lecidea fuscoatra;microthelia
(Wallr.) Nik.Hoffm. & Hafellner: on Trapelia;muralis van
den Boom: on Protoparmeliopsis muralis;nephromatis
(Crouan) Matzer & Hafellner: on Nephroma;oxyspora
Matzer & R.Sant.: on Coccocarpia;pannariae Matzer &
Hafellner: on Protopannaria pezizoides;papuana Die-
derich: on Usnea;peltigericola D.Hawksw. & Mia
˛dl.: on
Peltigera;ramirezii Etayo: on Lobaria;stereocaulorum
Zhurb., Kukwa & Oset: on Stereocaulon;stictae Etayo: on
Sticta.
Lit.: Aptroot et al. 1997; Darmostuk et al. 2018; Etayo 2002a;
Hawksworth & Mia
˛dlikowska 1997a; Hawksworth et al. 2010b;
Hoffmann & Hafellner 2000; Matzer & Hafellner 1990; van den
Boom 2015; Zhurbenko et al. 2009.
Trichothecium Link; type: T. roseum (Pers.) Link. Mol.: yes (T,
L). Asexual, hyphomycetous.
Species: 0-0-1 (9). roseum (Pers.) Link: facultatively
lichenicolous.
Lit.: Hawksworth 1979a.
Order Microascales Benny & Kimbr.
Family Graphiaceae De Beer
Graphium Corda [a hemihomonym of Graphium Scopoli,
Lepidoptera]; type: G. penicillioides Corda. Mol.: yes (T).
Asexual, synnematous.
Species: 2-0-0 (20). aphthosae Alstrup & D.Hawksw.: on
Peltigera and other terricolous lichens, possibly algicolous;
samogiticum Motiej. & Alstrup: over Sarcosagium and
Verrucaria, possibly algicolous.
Lit.: Alstrup & Hawksworth 1990; Motiej¯
unaite
˙& Alstrup 2006.
Subclass SORDARIOMYCETIDAE O.E.Erikss. &
Winka
Order Chaetosphaeriales Huhndorf, A.N.Mill. &
F.A.Ferna
´ndez
Family Chaetosphaeriaceae R´
eblova
´, M.E.Barr &
Samuels
Dinemasporium L´
ev.; type: D. graminum, i.e. D. strigosum.
Mol.: yes (T, L). Asexual, pycnidial.
Species: 0-0-1 (35). strigosum (Fr.) Sacc. [¼D. gramineum
(Lib.) L´
ev.]: facultatively lichenicolous on Peltigera, more
rarely other lichens.
Lit.: Etayo 2017; Mia
˛dlikowska & Alstrup 1995.
Order Phyllachorales M.E.Barr.
Family Phyllachoraceae Theiss. & P.Syd.
Lichenochora Hafellner; type: L. thallina, i.e. L. obscuroides.
Mol.: no.
Species: 44-0-0. acutispora Etayo: on Fuscopannaria
magellanica;aipoliae Etayo, Nav.-Ros. & Coppins: on
Physcia aipolia;aprica Hafellner & Nik.Hoffm.: on
Aspicilia;arctica Zhurb.: on Candelariella canadensis
and neighbouring Rinodina terrestris;atrans Halıcı,
K.Knudsen & Candan: on Psora decipiens;bacidiispora
Etayo: on Parmotrema;bellemerei Nav.-Ros., Cl.Roux &
Diederich: on Menegazzia;caloplacae Zhurb.: on Athal-
lia,Blastenia,Bryoplaca and Caloplaca s.str.; chimaerica
Etayo: on Pertusaria [close to Parapyrenis,fideEtayo,
pers. comm.]; clauzadei Nav.-Ros., Cl.Roux & Llimona:
on Squamarina;coarctatae (B. de Lesd.) Hafellner &
F.Berger: on Trapelia;collematum Nik.Hoffm. & Ha-
fellner: on Rostania ceranisca;constrictella (M ¨
ull.Arg.)
Hafellner: on Gyalolechia bracteata;coppinsii Etayo &
Nav.-Ros.: on Protopannaria pezizoides;elegantis Hafell-
ner: on Rusavskia elegans;epidesertorum Nav.-Ros.: on
Gyalolechia desertorum;epifulgens Nav.-Ros. & Cl.Roux:
on Gyalolechia fulgens;epimarmorata Nav.-Ros.: on
Xanthocarpia marmorata;epinashii Nav.-Ros. & Etayo:
on Polycauliona nashii;gahavisukae Diederich: on
?Graphis;galligena R.Sant. & Hafellner: on Physcia;
haematommatum R.C.Harris & Lendemer: on Haema-
tomma personii;heppiae Cl.Roux: on Heppia lutosa;
hypanica S.Y.Kondr., L.L˜
ok¨
os&J.-S.Hur:onEndocar-
pon;hyperphysciae Etayo: on Hyperphyscia adglutinata;
inconspicua Hafellner: on Lecidea berengeriana;lecidel-
lae Boqueras & Nav.-Ros.: on Lecidella elaeochroma;
lepidiotae (Anzi) Etayo & Nav.-Ros.: on Fuscopannaria
praetermissa;makareviczae S.Y.Kondr.: on Lecanora aff.
campestris;mediterraneae Calat., Nav.-Ros. & E.Calvo:
on Fuscopannaria mediterranea;monegrina Etayo: on
terricolous Polyblastia;obscuroides (Linds.) Triebel &
Rambold [¼L. thallina (Cooke) Hafellner]: on Phaeo-
physcia,morerarelyPhyscia;paucispora Etayo & Nav.-
Ros.: on Bilimbia sabuletorum;physciicola (Ihlen &
R.Sant.) Hafellner: on Physcia dubia;polycoccoides
Hafellner & R.Sant.: on Physcia tribacia;pyrenodesmiae
Nav.-Ros. & Cl.Roux: on Pyrenodesmia;rinodinae
Zhurb.: on Rinodina;sinapispermae Etayo & Nav.-
Ros.: on Bryoplaca sinapisperma;tertia Etayo, Flakus &
Rodr.Flakus: on Rusavskia elegans;thorii Zhurb.: on
Aspicilia moenium;verrucicola (Wedd.) Nik.Hoffm. &
Hafellner: on Aspicilia calcarea,A. cinerea and A.
contorta;wasseri S.Y.Kondr.: on Flavoplaca;weillii
(Werner) Hafellner & R.Sant.: on Physcia and Physconia;
xanthoriae Triebel & Rambold: on Leproplaca proteus
and Rusavskia elegans.
Lit.: Aptroot et al. 1997; Calatayud et al. 2000; Etayo 2010c, 2011,
2017; Etayo & Navarro-Rosin´
es 2008; Etayo & Sancho 2008; Etayo
et al. 2013c; Hafellner 1989; Hafellner & Berger 2000; Hafellner &
Zimmermann 2012; Hoffmann & Hafellner 2000; Ihlen & Wedin
2005; Kondratyuk et al. 2014, 2015; Lendemer et al. 2016;
Navarro-Rosin´
es & Etayo 2001; Navarro-Rosin´
es et al. 1994,
1998a, 1998c; Navrotskaya et al. 1996; Roux & Triebel 1994;
Triebel et al. 1991; Zhurbenko 2008b, 2013a; Zhurbenko &
Brackel 2013.
384 The Bryologist 121(3): 2018
Order Sordariales D.Hawksw. & O.E.Erikss.
Family Helminthosphaeriaceae Samuels, Cand. &
Magni
Endophragmiella B.Sutton; type: E. pallescens B.Sutton. Mol.:
yes. Asexual, hyphomycetous.
Species: 3-0-0 (70). franconica Brackel & Markovskaja: on
Cladonia,Platismatia glauca and Hypogymnia physodes;
hughesii D.Hawksw.: on Lobaria pulmonaria;stordeuriana
U.Braun, Zhurb., Diederich, Tsurykau & Heuchert: on
Cladonia.
Lit.: Brackel & Markovskaja 2009; Hawksworth 1979a; Zhurbenko
et al. 2015d.
Order Sordariales genera incertae sedis
Globosphaeria D.Hawksw.; type: G. jamesii. Mol.: no.
Species: 1-0-0. jamesii D.Hawksw.: on Normandina
pulchella.
Lit.: Hawksworth 1990b.
Reconditella Matzer & Hafellner; type: R. physconiarum. Mol.:
no.
Species: 1-0-0. physconiarum Hafellner & Matzer: on
Physconia distorta and P. venusta.
Lit.: Matzer & Hafellner 1990.
Roselliniomyces Matzer & Hafellner; type: R. trichotheliorum.
Mol.: no.
Species: 2-0-0. erinaceus Etayo: on Sticta;trichotheliorum
Matzer & Hafellner: on foliicolous Trichothelium.
Lit.: Etayo 2017; Matzer & Hafellner 1990.
Roselliniopsis Matzer & Hafellner; type: R. groedensis.Mol.:
no.
Species: 7-0-0. gelidaria (Mudd) Matzer: on Placopsis gelida;
groedensis (Zopf) Matzer & Hafellner: on Varicellaria lactea;
palicei Etayo: on Placopsis gelida;phaeophysciae S.Y.Kondr.,
L.L}
ok¨
os & J.-S.Hur: on Phaeophyscia;tartaricola (Leight.)
Matzer: on Lepra,Ochrolechia,Pertusaria pupillaris and
Varicellaria hemisphaerica;tropica Matzer & R.Sant.: on
?Ochrolechia;ventosa (Rostr.) Alstrup: on Placopsis gelida.
Lit.: Alstrup et al. 1994; Etayo 2003; Kondratyuk et al. 2013; Matzer
1993; Matzer & Hafellner 1990.
Subclass XYLARIOMYCETIDAE O.E.Erikss.
& Winka
Order Xylariales Nannf.
Family Hyponectriaceae Petr.
Charonectria Sacc.; type: C. consolationis Sacc. Mol.: no.
Species: 1-0-0 (3). amabilis Lowen & D.Hawksw.: on
Phaeorrhiza.
Lit.: Rossman et al. 1999.
Physalospora Niessl; type: P. alpestris Niessl. Mol.: yes.
Species: 2-0-0 (37). collematis (Stein) G.Winter: on
Collema [a poorly known species]; jonaspidis (Stein)
Vouaux: on Ionaspis chrysophana [a poorly known
species].
Lit.: Hoffmann & Hafellner 2000.
Family Hypoxylaceae DC.
Hypoxylon Bull.; type: H. fragiforme (Pers.) J.Kickx f. [¼
Sphaeria Haller, nom. rejic.]. Mol.: yes (T).
Species: 1-0-0 (140). Sphaeria sociella Nyl.: on Trapelia
coarctata [phylogenetic position uncertain].
Lit.: Triebel 1989.
Family Xylariaceae Tul. & C.Tul.
Anthostomella Sacc.; type: A. limitata Sacc. Mol.: yes. Currently
heterogeneous.
Species: 1-0-0 (85). apogyra (Nyl.) Saccardo & D.Saccardo:
on Umbilicaria [a poorly known species].
Lit.: Clauzade et al. 1989; Matzer & Hafellner 1990.
Class Sordariomycetes ordo incertae sedis
Order Coniochaetales Huhndorf, A.N.Mill. &
F.A.Ferna
´ndez
Family Coniochaetaceae Malloch & Cain
Coniochaeta (Sacc.) Cooke; type: C. ligniaria. Mol.: yes (T, L).
Species: 0-0-1 (80). ligniaria (Grev.) Massee: facultatively
lichenicolous on Diploschistes diacapsis.
Lit.: Etayo 2008.
Subphylum Pezizomycotina ordo incertae sedis
Order Thelocarpales L¨
ucking & Lumbsch
Family Thelocarpaceae Zukal
Thelocarpon Nyl.; type: T. laureri (Flot.) Nyl. (lichenized). Mol.:
yes (T, L).
Species: 0-0-4 (25). cyaneum Olech & Alstrup: on
‘Polyblastia cf. gothica’, probably facultatively lichenicolous;
epibolum Nyl. var. epibolum: facultatively lichenicolous
on Arthrorhaphis,Baeomyces,Lichenomphalia hudsoniana,
Peltigera and Solorina;epibolum var. epithallinum
(Leight.) G.Salisb.: facultatively lichenicolous on Arthro-
rhaphis,Baeomyces,Catapyrenium,Peltigera,Protopannaria
and Solorina;lichenicola (Fuckel) Poelt & Hafellner:
facultatively lichenicolous on Baeomyces,Peltigera,Solorina
and other terricolous lichens.
Lit.: Olech & Alstrup 1990; Rambold & Triebel 1992.
Diederich et al.: 2018 classification and checklist of lichenicolous fungi 385
Subphylum Pezizomycotina familiae incertae sedis
Family Myxotrichaceae Currah
Myxotrichum Kunze; type: M. chartarum Kunze. Mol.: yes (T).
Species: 0-0-1 (12). bicolor (Ehrenb.) Fr.: on dying thalli of
Usnea, possibly only fortuitously lichenicolous.
Lit.: Hawksworth 1979a.
Family Obryzaceae O.E.Erikss.
Obryzum Wallr.; type: O. corniculatum. Mol.: no.
Species: 3-0-0. corniculatum Wallr.: on Scytinium;friesii
(Keissl.) Nik.Hoffm. & Hafellner: on Leptogium;strigu-
loides Aptroot & Sipman: on Agonimia vouauxii.
Lit.: Aptroot & Sipman 2001; Hoffmann & Hafellner 2000; L¨
ucking
et al. 2017.
Subphylum Pezizomycotina genera incertae sedis
Acaroconium Kocourk. & D.Hawksw.; type: A. punctiforme.
Mol.: no. Asexual, pycnidial.
Species: 1-0-0. punctiforme Kocourk. & D.Hawksw.: on
Acarospora and Sarcogyne.
Lit.: Kocourkova
´& Hawksworth 2008.
Actinocladium Ehrenb.; type: A. rhodosporum.Mol.:no.
Asexual, hyphomycetous.
Species: 0-0-1 (8). rhodosporum Ehrenb.: facultatively
lichenicolous on Flavoparmelia,Hypotrachyna,Lecanora,
Lepra and Lepraria.
Lit.: Brackel 2014.
Arborillus Munt.-Cvetk. & G ´
omez-Bolea; type: A. llimonae.
Mol.: no. Asexual, synnematous.
Species: 1-0-0. llimonae Munt.-Cvetk. & G ´
omez-Bolea: on
Diploschistes.
Lit.: Munta˜
nola-Cvetkovic & G´
omez-Bolea 1998.
Asteroglobulus Brackel; type: A. giselae. Mol.: no. Asexual,
pycnidial.
Species: 2-0-0. giselae Brackel: on Ramalina farinacea;
pyramidalis (Etayo) Diederich, comb. nov.: on Hypotra-
chyna and Parmotrema.
Lit.: Brackel 2011a, Etayo 2010a.
Asterophoma D.Hawksw.; type: A. mazaediicola. Mol.: no.
Asexual, pycnidial.
Species: 1-0-0. mazaediicola D.Hawksw.: on Calicium and
Chaenotheca.
Lit.: Hawksworth 1981; Tibell 1991a.
Biciliopsis Diederich; type: B. leptogiicola. Mol.: no.
Species: 2-0-0. cladoniae Zhurb., Pino-Bodas & Diederich:
on Cladonia; leptogiicola Diederich: on Leptogium.
Lit.: Aptroot el al.1997; Hoffmann & Hafellner 2000; Zhurbenko &
Pino-Bodas 2017.
Bicoloromyces Heuchert, U.Braun & D.Hawksw.; type; B.
kyffinensis. Mol.: no. Asexual, hyphomycetous.
Species: 1-0-0. kyffinensis Heuchert, U.Braun &
D.Hawksw.: on Lecanora fuscobrunnea or Lecidella.
Lit.: Heuchert et al. 2015.
Calongeomyces D.Hawksw. & Etayo; type: C. gibelluloides [”
Calongia D.Hawksw. & Etayo, nom. illeg.]. Mol.: no.
Asexual, hyphomycetous.
Species: 1-0-0. gibelluloides (D.Hawksw. & Etayo)
D.Hawksw. & Etayo: on Cladonia,Flavoparmelia caperata
and Lepra amara.
Lit.: Hawksworth & Etayo 2010, 2011.
Caeruleoconidia Zhurb. & Diederich; type: C. ochrolechiae.
Mol.: no. Asexual, pycnidial.
Species: 2-0-0. biazrovii Zhurb.: on Cladonia;ochrolechiae
Zhurb. & Diederich: on Ochrolechia.
Lit.: Zhurbenko & Pino-Bodas 2017; Zhurbenko et al. 2015a.
Carnegieispora Etayo & F.Berger; type: C. rimeliae. Mol.: no.
Asexual, pycnidial.
Species: 1-0-0. rimeliae Etayo & F.Berger: on Parmotrema.
Lit.: Etayo & Berger 2009.
Cecidiomyces U.Braun & Zhurb.; type: C. santessonii. Mol.: no.
Asexual, hyphomycetous.
Species: 1-0-0. santessonii U.Braun & Zhurb.: on Stereo-
caulon.
Lit.: Zhurbenko & Braun 2012.
Choreospora Constant. & R.Sant.; type: C. tibellii. Mol.: no.
Asexual, sporodochial.
Species: 1-0-0. tibellii Constant. & R.Sant.: on foliicolous
?Echinoplaca.
Lit.: Constantinescu & Santesson 1987.
Cladoniicola Diederich, van den Boom & Aptroot; type: C.
staurospora. Mol.: no. Asexual, pycnidial.
Species: 2-0-0. irregularis Etayo: on Diploschistes diacapsis;
staurospora Diederich, van den Boom & Aptroot: on
Cladonia.
Lit.: Diederich et al. 2002a; Etayo 2010c.
Clauzadeomyces Diederich; type: C. verrucosus.Mol.:no.
Asexual, sporodochial.
Species: 1-0-0. verrucosus Diederich: on Placopsis.
Lit.: Diederich 1994.
Codonmyces Calat. & Etayo; type: C. lecanorae. Mol.: no.
Asexual, hyphomycetous.
Species: 1-0-0. lecanorae Calat. & Etayo: on Lecanora
valesiaca.
Lit.: Calatayud & Etayo 1999a.
Coniambigua Etayo & Diederich; type: C. phaeographidis. Mol.:
no. Asexual, pycnidial.
Species: 1-0-0. phaeographidis Etayo & Diederich: on
Phaeographis s.l. (incl. Leiorreuma).
Lit.: Etayo & Diederich 1995.
Cornutispora Piroz.; type: C. limaciformis. Mol.: no. Asexual,
pycnidial.
Species: 8-0-1. ciliata Kalb: on various lichens; herteliana
Knoph: on Lecidella;intermedia Punith. & D.Hawksw.: on
various lichens; lichenicola D.Hawksw. & B.Sutton: on
386 The Bryologist 121(3): 2018
various lichens; limaciformis Piroz.: facultatively lichen-
icolous on Protoparmeliopsis muralis;ophiurospora Etayo:
on Lobariella;pittii D.Hawksw. & Punith.: on Hypotrachyna;
triangularis Diederich & Etayo: on Pertusaria pertusa;
tricupulata F.Berger & E.Zimm.: on Physcia stellaris.
Lit.: Etayo 2017; Etayo & Diederich 1995; Gierl & Kalb 1993;
Hawksworth 1976, 1981; Knoph 2004; Punithalingam 2003;
Sutton 1980; van den Boom & Etayo 2000; Zimmermann &
Berger 2018.
Deichmannia Alstrup & D.Hawksw.; type: D. verrucosa. Mol.:
no. Asexual, sporodochial.
Species: 1-0-0. verrucosa Alstrup & D.Hawksw.: on
Diploschistes.
Lit.: Alstrup & Hawksworth 1990.
Dictyophrynella Bat. & Cavalc.; type: D. bignoniacearum. Mol.:
no. Asexual, hyphomycetous.
Species: 1-0-0. bignoniacearum Bat. & Caval.: on foliico-
lous lichens.
Lit.: Hawksworth 1979a.
Epaphroconidia Calat. & V.Atienza; type: E. hawksworthii. Mol.:
no. Asexual, pycnidial.
Species: 1-0-0. hawksworthii Calat. & V.Atienza: on
Pertusaria pertusa.
Lit.: Calatayud & Atienza 1995.
Everniicola D.Hawksw.; type: E. flexispora. Mol.: no. Asexual,
pycnidial.
Species: 1-0-0. flexispora D.Hawksw.: on Evernia and
Nephroma arcticum.
Lit.: Hawksworth 1982a.
Feltgeniomyces Diederich; type: F. luxemburgensis. Mol.: no.
Asexual, sporodochial.
Species: 4-0-0. dichotomus Etayo & Calat.: on Physcia;
luxemburgensis Diederich: on Lecidella;physciae Etayo &
Breuss: on Physcia;uniseptatus Diederich: on Hypotrachy-
na.
Lit.: Aptroot et al. 1997; Calatayud & Etayo 2001; Diederich 1990;
Etayo & Breuss 1998.
Fenestroconidia Calat. & Etayo; type: C. caloplacae. Mol.: no.
Asexual, sporodochial.
Species: 1-0-0. caloplacae Calat. & Etayo: on Calogaya.
Lit.: Calatayud & Etayo 1999b.
Gyrophthorus Hafellner & Sancho; type: G. perforans. Mol.: no.
Species: 3-0-0. crustulosae (Creveld) Hafellner & Sancho:
on Umbilicaria;gracilis Nik.Hoffm. & Hafellner: on
Umbilicaria;perforans Hafellner & Sancho: on Umbilicaria.
Lit.: Hafellner & Sancho 1990; Hoffman & Hafellner 2000.
Hawksworthiana U.Braun; type: H. peltigericola. Mol.: no.
Asexual, hyphomycetous.
Species: 1-0-0. peltigericola (D.Hawksw.) U.Braun: on
Peltigera.
Lit.: Braun 1988; Hawksworth 1980a.
Henfellra Halıcı, D.Hawksw., Z.Kocak. & M.Kocak.; type: H.
muriformis. Mol.: no. Asexual, pycnidial.
Species: 1-0-0. muriformis Halıcı, D.Hawksw., Z.Kocak. &
M.Kocak.: on Candelariella antennaria.
Lit.: Hawksworth et al. 2016.
Hymenobia Nyl.; type: H. insidiosa [¼Hymenobiella Triebel;
type: H. aporea]. Mol.: no. A poorly known genus.
Species: 1-0-0. insidiosa Nyl. [¼Hymenobiella aporea (Nyl.)
Triebel]: on Lecidea.
Lit.: Eriksson & Hawksworth 1991; Triebel 1989.
Hypotrachynicola Etayo; type: H. rubra. Mol.: no. Sexual.
Species: 1-0-0. rubra Etayo: on Hypotrachyna.
Lit.: Etayo 2002.
Intralichen D.Hawksw. & M.S.Cole; type: I. christiansenii.
Mol.: yes (T, L). Asexual, hyphomycetous. Probably
polyphyletic.
Species: 4-0-0. baccisporus D.Hawksw. & M.S.Cole: on
Calogaya,Flavoplaca,Gyalolechia,Xanthocarpia,Xantho-
mendoza and other lichens; christiansenii (D.Hawksw.)
D.Hawksw. & M.S.Cole [”Bispora christiansenii
D.Hawksw.]: on Candelariella and other lichens; lichen-
icola (M.S.Christ. & D.Hawksw.) D.Hawksw. & M.S.Cole
[”Trimmatostroma lichenicola M.S.Christ. & D.Hawksw.]:
on Candelariella and other lichens; lichenum (Diederich)
D.Hawksw. & M.S.Cole [”Bispora lichenum Diederich]: on
various lichens.
Lit.: Diederich 1990; Hawksworth 1979a; Hawksworth & Cole
2002.
Isthmoconidium Etayo & F.Berger; type: I. bacidiicola. Mol.: no.
Asexual, pycnidial.
Species: 1-0-0. bacidiicola Etayo & F.Berger: on Bacidia.
Lit.: Etayo & Berger 2013.
Kalchbrenneriella Diederich & M.S.Christ.; type: K. cyanescens.
Mol.: no. Asexual, hyphomycetous.
Species: 1-0-0. cyanescens (Kalchbr.) Diederich &
M.S.Christ.: on Usnea.
Lit.: Diederich 2002.
Katherinomyces Khodos.; type: K. cetrariae. Mol.: no. Asexual,
coelomycetous, stromatic.
Species: 1-0-0. cetrariae Khodos: on Cetraria aculeata.
Lit.: Khodosovtsev et al. 2016, 2018.
Keissleriomyces D.Hawksw.; type: K. sandstedeanus. Mol.: no.
Asexual, pycnidial.
Species: 1-0-0. sandstedeanus (Keissler) D.Hawksw.: on
Cladonia.
Lit.: Hawksworth 1981.
Lambinonia S´
erus. & Diederich; type: L. strigulae. Mol.: no.
Asexual, sporodochial.
Species: 1-0-0. strigulae (Elenkin & Woron.) S´
erus. &
Diederich: on foliicolous Strigula.
Lit.: S´
erusiaux & Diederich 2005.
Lawalreea Diederich; type: L. lecanorae. Mol.: no. Asexual,
pycnidial. Resembles pycnidia of some Lecanora s.l.
Species: 1-0-0. lecanorae Diederich: on Myriolecis persimilis
and other Myriolecis species.
Lit.: Diederich 1990.
Leightoniomyces D.Hawksw. & B.Sutton; type: L. phillipsii.
Mol.: no. Asexual, hyphomycetous.
Diederich et al.: 2018 classification and checklist of lichenicolous fungi 387
Species: 2-0-0. canariensis Etayo & P ´
erez-Vargas: on
crustose saxicolous lichens; phillipsii (Berk. & Leight.)
D.Hawksw. & B.Sutton: on Steinia geophana and
Thrombium epigaeum, possibly algicolous.
Lit.: Hawksworth 1977a, 1979a; P´
erez-Vargas et al. 2013.
Lichenobactridium Diederich & Etayo; type: L. pertusariae.
Mol.: no. Asexual, sporodochial.
Species: 1-0-0. pertusariae Diederich & Etayo: on Pertu-
saria pertusa.
Lit.: Etayo & Diederich 1995.
Lichenohendersonia Calat. & Etayo; type: L. squamarinae. Mol.:
no. Asexual, pycnidial.
Species: 3-0-0. squamarinae Calat. & Etayo: on Squamar-
ina;uniseptata Etayo & Calat.: on Parmelia and Parmelina;
varians Calat. & Etayo: on Rhizoplaca.
Lit.: Calatayud & Etayo 2001.
Lichenopenicillus Etayo; type: L. versicolor. Mol.: no. Asexual,
synnematous.
Species: 1-0-0. versicolor Etayo: on Sticta and Leptogium.
Lit.: Etayo 2017.
Lichenophoma Keissl.; type: L. haematommatis. Mol.: no.
Asexual, pycnidial. A poorly known genus.
Species: 1-0-0 (2). haematommatis Keissl.: on Haematomma.
Lit.: Hawksworth 1981.
Lichenopuccinia D.Hawksw. & Hafellner; type: L. poeltii. Mol.:
no. Asexual, sporodochial.
Species: 1-0-0. poeltii D.Hawksw.&Hafellner:onParmelia.
Lit.: Hawksworth 1984.
Lichenostella Calat. & Etayo; type: L. hypotrachynae. Mol.: no.
Asexual, sporodochial.
Species: 1-0-0. hypotrachynae Etayo & Calat.: on Hypo-
trachyna.
Lit.: Calatayud & Etayo 1999a.
Milospium D.Hawksw.; type: M. graphideorum. Mol.: yes (T).
Asexual, hyphomycetous.
Species: 2-0-0 (4). graphideorum (Nyl.) D.Hawksw.: on
Dirina massiliensis, sometimes lichenized; lacoizquetae
Etayo & Diederich: on Cladonia [often co-occurring with
Arthonia digitatae].
Lit.: Crous et al. 2015; Etayo & Diederich 1996a; Hawksworth 1975,
1979a.
Minutophoma D.Hawksw.; type: M. chrysophthalmae. Mol.: no.
Asexual, pycnidial.
Species: 1-0-0. chrysophthalmae D.Hawksw.: on Chryso-
thrix chrysophthalmae.
Lit.: Hawksworth 1981.
Neolamya Theiss. & Syd.; type: N. peltigerae. Mol.: no. Sexual.
Species: 3-0-0. ahtii Zhurb.: on Cladonia;peltigerae
(Mont.) Theiss. & Syd.: on Peltigera (mainly P. didactyla);
xanthoparmeliae Kocourk.: on Xanthoparmelia conspersa.
Lit.: Keissler 1930; Kocourkova
´2009; Zhurbenko & Pino-Bodas
2017.
Nigromacula Etayo; type: N. hypotrachynae. Mol.: no. Asexual,
sporodochial.
Species: 1-0-0. uniseptata (D.Hawksw.) D.Hawksw. [”
Vouauxiella u. D.Hawksw.; ¼N. hypotrachynae Etayo]: on
Hypotrachyna.
Lit.: Etayo 2002.
Nigropuncta D.Hawksw.; type: N. rugulosa. Mol.: no. Asexual,
pycnidial.
Species: 2-0-0. groenlandica Alstrup: on crustose lichen;
rugulosa D.Hawksw.: on Bellemerea.
Lit.: Alstrup 1993a; Hawksworth 1981; Hawksworth & Poelt 1986.
Patriciomyces D.Hawksw.; type: P. valentinianus. Mol.: no.
Asexual, hyphomycetous.
Species: 1-0-0. valentinianus D.Hawksw.: on Physcia.
Lit.: Cole & Hawksworth 2001.
Psammina Sacc. & M.Rousseau; type: P. bommeriae M.Rousseau
& Sacc. Mol.: no. Asexual, hyphomycetous.
Species: 3-0-0 (8). inflata Earl.-Benn. & D.Hawksw.: on
Lepraria and corticolous algae; simplex Earl.-Benn. &
D.Hawksw.:on various corticolous lichens and algae; stipitata
D.Hawksw.: on various corticolous lichens and algae.
Lit.: Earland-Bennett & Hawksworth 1999a; Hawksworth 1979a.
Pycnopsammina Diederich & Etayo; type: P. lobariae. Mol.: no.
Asexual, pycnidial.
Species: 1-0-0. lobariae Diederich & Etayo [”Psammina
lobariae (Diederich & Etayo) Earl.-Benn. & D.Hawksw.]: on
Lobaria pulmonaria.
Lit.: Etayo & Diederich 1995.
Pygmaeosphaera Etayo & Diederich; type: P. coccocarpiae. Mol.:
no. Possibly related to Tubeufiales.
Species: 3-0-0. coccocarpiae (Diederich) Etayo & Diederich
[”Nectriopsis coccocarpiae Diederich]: on Coccocarpia;
epigraphis Etayo: on Graphis;sipmaniana Etayo: on
Parmeliella.
Lit.: Etayo 2017.
Refractohilum D.Hawksw.; type: R. galligenum. Mol.: no.
Asexual, hyphomycetous.
Species: 5-0-0. achromaticum (B.Sutton) D.Hawksw.: on
Gyalecta;galligenum D.Hawksw.: on Nephroma laeviga-
tum;intermedium Cl.Roux & Etayo: on Gyalecta;peltiger-
ae (Keissl.) D.Hawksw.: on Peltigera;pluriseptatum Etayo
&Cl.Roux:onGyalecta,possiblyalsoonNephroma
antarcticum.
Lit.: Etayo & Sancho 2008; Hawksworth 1977a, 1979a; Roux et al.
1997.
Sarcoexcipula Etayo; type: S. pannariae. Mol.: no. Sexual.
Species: 1-0-0. pannariae Etayo: on Pannaria pallida.
Lit.: Etayo & Sancho 2008.
Sessiliospora D.Hawksw.; type: S. bicolor. Mol.: no. Asexual,
hyphomycetous.
Species: 1-0-0. bicolor D.Hawksw.: foliicolous lichens.
Lit.: Hawksworth 1979a.
Spilomium Nyl.; typification not clear. Mol.: no. Asexual,
sporodochial.
Species: 0-1-0. auratum Sm.: lichenized, facultatively
lichenicolous on Dirina spp.
Lit.: Giavarini & Purvis 2009b.
388 The Bryologist 121(3): 2018
Stellifraga Alstrup & Olech; type: S. cladoniicola. Mol.: no.
Sexual.
Species: 1-0-0. cladoniicola Alstrup & Olech: on Cladonia.
Lit.: Alstrup & Olech 1993.
Stygiomyces Coppins & S.Y.Kondr.; type: S. galliformis. Mol.:
no. Asexual, pycnidial.
Species: 1-0-0. galliformis Coppins & S.Y.Kondr.: on
Pseudocyphellaria.
Lit.: Coppins & Kondratyuk 1995.
Taeniolina M.B.Ellis; type: T. centaurii (Fuckel) M.B.Ellis. Mol.:
no. Asexual, hyphomycetous.
Species: 0-0-1 (4). scripta (P.Karst) P.M.Kirk: facultatively
lichenicolous on corticolous lichens.
Lit.: Diederich 1989; Hawksworth 1979a; Heuchert et al. 2018.
Talpapellis Alstrup & M.Cole; type: T. peltigerae. Mol.: no.
Asexual, hyphomycetous.
Species: 5-0-0. beschiana (Diederich) Zhurb., U.Braun,
Diederich & Heuchert [”Taeniolella beschiana Diederich;
¼Ameroconium cladoniae U.Braun & Zhurb.]: on Cladonia;
mahensis Diederich, U.Braun & Ertz: on crustose lichen
with Trentepohlia;peltigerae Alstrup & M.S.Cole var.
peltigerae:onPeltigera venosa;peltigerae Alstrup &
M.S.Cole var. rossica Heuchert, Zhurb. & U.Braun: on
Peltigera spp.; solorinae Zhurb., Heuchert & U.Braun: on
Solorina crocea.
Lit.: Alstrup & Cole 1998; Diederich 1992; Diederich et al. 2017;
Heuchert et al. 2014, 2018; Zhurbenko & Braun 2013; Zhurbenko
et al. 2015c.
Teratosperma Syd. & P.Syd.; type: T. singulare Syd. & P.Syd.
Mol.: no. Asexual, hyphomycetous.
Species: 2-0-0 (14). anacardii Hansf.: on foliicolous lichens;
lichenicola D.Hawksw.: on foliicolous lichens.
Lit.: Hawksworth 1979a.
Trichoconis Clem.; type: T. caudata (Appel & Strunk) Clem.
Mol.: yes. Asexual, hyphomycetous.
Species: 3-0-0 (21). hafellneri U.Braun, Khodos., Darmos-
tuk & Diederich: on Athallia pyracea and Xanthoria
parietina;lichenicola D.Hawksw.: on Peltigera collina;
physciicola Brackel: on Physcia.
Lit.: Brackel 2014; Braun et al. 2016; Hawskworth 1980b.
Vagnia D.Hawksw. & Mia
˛dl.; type: V. cirriformis. Mol.: no.
Asexual, coelomycetous.
Species: 1-0-0. cirriformis D.Hawksw. & Mia
˛dl. [as
‘cirriformia’]: on Peltigera.
Lit.: Hawksworth & Mia
˛dlikowska 1997b.
Verrucaster Tobler; type: V. lichenicola. Mol.: no. Asexual,
pycnidial. A poorly known taxon.
Species: 1-0-0. lichenicola Tobler: on Cladonia.
Lit.: Hawksworth 1981.
Viridiannula Etayo; type: V. pertusariae. Mol.: no. Asexual,
pycnidial.
Species: 1-0-0. pertusariae Etayo: on Pertusaria pertusa.
Lit.: Etayo 2002b.
Vouauxiella Petr. & Syd.; type: V. verrucosa. Mol.: no. Asexual,
pycnidial.
Species: 3-0-0. lichenicola (Linds.) Petrak & H.Sydow: on
Lecanora;pithospora (Cavalc. & Silva) B.Sutton: on
foliicolous lichens; verrucosa (Vouaux) Petrak & H.Sydow:
on Lecanora.
Lit.: Hawksworth 1981.
Zevadia J.C.David & D.Hawksw.; type: Z. peroccidentalis. Mol.:
no. Asexual, hyphomycetous.
Species: 1-0-0. peroccidentalis J.C.David & D.Hawksw.: on
Usnea.
Lit.: David & Hawksworth 1995.
Zwackhiomacromyces Etayo & van den Boom; type: Z.
constrictocarpus. Mol.: no. Sexual.
Species: 2-0-0. constrictocarpus Etayo & van den Boom: on
Megalospora verrucosa;hyalosporus (Alstrup, D.Hawksw. &
R.Sant.) Etayo & F.Berger [”Pyrenidium hyalosporum
Alstrup, D.Hawksw. & R.Sant.]: on Placopsis.
Lit.: Alstrup & Hawksworth 1990; van den Boom & Etayo
2014.
Phylum BASIDIOMYCOTA R.T.Moore
Subphylum PUCCINIOMYCOTINA R.Bauer, Be-
gerow, J.P.Samp., M.Weiss & Oberw.
Order Agaricostilbales Oberw. & R.Bauer
Family Chionosphaeraceae Oberw. & Bandoni
Chionosphaera D.E.Cox; type: C. apobasidialis D.E.Cox (sapro-
trophic). Mol.: yes (T).
Species: 2-0-0 (6). coppinsii P.Roberts: on various lichens;
lichenicola Alstrup, Sutton & Tønsberg: on various lichens.
Lit.: Diederich 1996; Kirschner et al. 2001; Roberts 1997.
Order Cyphobasidiales T.Sprib. & H.Mayrhofer
Family Cyphobasidiaceae T.Sprib. & H.Mayrhofer
Cyphobasidium Millanes, Diederich & Wedin; type: C.
hypogymniicola. Mol.: yes (T, L). Sexual stage in galls, yeast
stage asymptomatic in lichen cortex; DNA sequences of
many unnamed species obtained from the cortex of
macrolichens, especially of Parmeliaceae.
Species: 2-0-0. hypogymniicola (Diederich & Ahti) Mill-
anes, Diederich & Wedin [”Cystobasidium hypogymniicola
Diederich & Ahti]: on Hypogymnia;usneicola (Diederich &
Alstrup) Millanes, Diederich & Wedin [”Cystobasidium
usneicola Diederich & Alstrup]: on Usnea [consists of two
genetically distinct species].
Lit.: Diederich 1996; Diederich & Christiansen 1994; Millanes et al.
2016a; Spribille et al. 2016.
Subphylum AGARICOMYCOTINA Doweld [¼Ba-
sidiomycotina Ainsw.]
Class TREMELLOMYCETES Doweld
Diederich et al.: 2018 classification and checklist of lichenicolous fungi 389
Order Filobasidiales J¨
ulich
Family Filobasidiaceae L.S.Olive
Heterocephalacria Berthier; type: H. solida Berthier. Mol.: yes
(T, L).
Species: 2-0-0 (4). bachmannii (Diederich & M.S.Christ.)
Millanes & Wedin [”Syzygospora bachmannii Diederich &
M.S.Christ.]: on Cladonia;physciacearum (Diederich)
Millanes & Wedin [”S. physciacearum Diederich]: on
Leucodermia,Physcia,Physconia and Polyblastidium, also
reported from Phaeophyscia.
Lit.: Diederich 1996; Liu et al. 2016.
Syzygospora G.W.Martin; type: S. alba G.W.Martin. Mol.: yes
(T).
Species: 1-0-0 (3). parmeliicola Diederich: on Hypotrachy-
na [phylogenetic position unknown].
Lit.: Diederich 1996.
Order Tremellales Fr.
Family Tremellaceae Fr.
Biatoropsis R¨
as¨
anen; type: B. usnearum. Mol.: yes (T, L).
Species: 4-0-0. hafellneri Millanes, Diederich, M.Westb. &
Wedin: on Usnea fragilescens aggr., mainly U. cornuta;
minuta Millanes, Diederich, M.Westb. & Wedin: on Usnea
barbata and U. lapponica;protousneae Millanes, Diederich,
M.Westb. & Wedin: on Protousnea dusenii;usnearum
R¨
as¨
anen s.str.: on Usnea (incl. U. flavocardia,U. florida/
subfloridana,U. fragilescens,U. glabrescens,U. cf. krogiana
and U. macaronesica); in continental Europe so far only on
U. florida/subfloridana.
Lit.: Diederich 1996; Diederich & Christiansen 1994; Millanes et al.
2014b, 2016b.
Tremella Pers. s.l.; type: T. mesenterica (Schaeff.) Retz. Mol.: yes
(T, L). All lichenicolous species sexual, often with asexual
stage.
Species: 60-0-0 (250). anaptychiae J.C.Zamora & Diederich:
on Anaptychia ciliaris;caloplacae (Zahlbr.) Diederich s.l.: on
Caloplaca s.l. and Xanthoria;candelariellae Diederich &
Etayo: on Candelariella;celata J.C.Zamora, Millanes, V.J.Rico
&P
´
erez-Ortega: on Ramalina fraxinea;cephalodiicola Die-
derich: on Psoroma;cetrariellae Millanes, Diederich,
M.Westb., Pippola & Wedin: on Cetraria delisei;cetrariicola
Diederich & Coppins: on Nephromopsis;christiansenii
Diederich: on Physcia;cladoniae Diederich & M.S.Christ.:
on Cladonia;coccocarpiae Diederich: on Coccocarpia;
coppinsii Diederich & G.Marson: on Platismatia;dendrog-
raphae Diederich & Tehler: on Dendrographa;diederichiana
P´
erez-Ortega, Millanes, Wedin, V.J.Rico & J.C.Zamora: on
Lecidea aff. erythrophaea;diploschistina Millanes, M.Westb.,
Wedin & Diederich: on Diploschistes;dirinariae Diederich,
Millanes & Wedin: on Dirinaria aegialita;endosporogena
J.C.Zamora, Millanes, V.J.Rico & P ´
erez-Ortega: on Lecanora
carpinea (apothecia); everniae Diederich: on Evernia meso-
morpha;graphidastrae Diederich: on Graphidastra;graph-
idis Diederich, Millanes & Wedin: on Graphis;
haematommatis Diederich: on Haematomma;harrisii Die-
derich: on Polymeridium and Trypethelium;huuskonenii
Diederich, Myllys, Goward & Lindgren: hyperparasitic on
Raesaenenia huuskonenii over Bryoria;hypocenomycis Die-
derich: on Hypocenomyce;hypogymniae Diederich &
M.S.Christ.: on Hypogymnia;leptogii Diederich: on Lepto-
gium;lethariae:onLetharia vulpina;lichenicola Diederich:
on Violella fucata;lobariacearum Diederich & M.S.Christ.:
on Lobaria and Pseudocyphellaria;macrobasidiata J.C.Za-
mora, P´
erez-Ortega & V.J.Rico: on Lecanora;macroceratis
Diederich & Hafellner: on Cladonia;mayrhoferi J.C.Zamora,
Millanes, Etayo & Wedin: on Lecanora allophana;microcarpa
Diederich: on cf. Megalospora;monospora Diederich: on
Leptogium;montis-wilhelmii Diederich: on Normandina
simodense;nashii Diederich: on Usnea;nephromatis Die-
derich: on Nephroma;normandinae Diederich: on Norman-
dina pulchella;nieblae Diederich: on Niebla;papuana
Diederich: on Hypogymnia;parmeliarum Diederich: on
Parmotrema;parmeliellae Diederich & S´
erusiaux: on Parme-
liella;pertusariae Diederich: on Pertusaria;phaeographidis
Diederich, Coppins & Bandoni: on Phaeographis;phaeogra-
phinae Diederich & Aptroot: on ‘Phaeographina’; phaeophy-
sciae Diederich & M.S.Christ.: on Phaeophyscia;
protoparmeliae Diederich & Coppins: on Protoparmelia;
psoroglaenae Diederich: on Psoroglaena;psoromicola Die-
derich: on Psoroma;pyrenulae Diederich, Millanes, Wedin &
Common: on Pyrenula;ramalinae Diederich: on Ramalina;
rhizocarpicola Diederich, Millanes & Wedin: on Rhizocar-
pon;rinodinae Diederich & M.S.Christ.: on Rinodina;
santessonii Diederich: on Usnea;stevensiana Diederich: on
Usnea;stictae Diederich: on Dendriscocaulon and Sticta;
sulcariae Diederich & M.S.Christ.: on Sulcaria;umbilicariae
Diederich & Millanes: on Umbilicaria;tuckerae Diederich:
on Ramalina;variae P´
erez-Ortega, Millanes, V.J.Rico &
J.C.Zamora: on Lecanora varia;wirthii Diederich: on
Protoparmelia.
Lit.: Ariyawansa et al. 2015; Diederich 1996, 2003, 2004d, 2007a,
2007b; Diederich et al. 2014b; Etayo 2018a; Lindgren et al. 2015;
S´
erusiaux et al. 2003; Millanes et al. 2011, 2012, 2014a, 2015; Werth
et al. 2013; Zamora et al. 2016, 2017.
Class Tremellomycetes familia incertae sedis
Family Heteroacanthellaceae P.Roberts
Heteroacanthella Oberw.; type: H. variabilis Oberw. & Langer.
Mol.: yes (T).
Species: 1-0-0 (3). ellipsospora J.C.Zamora, P´
erez-Ortega
& V.J.Rico: on Lecanora carpinea.
Lit.: Zamora et al. 2014.
Class AGARICOMYCETES Doweld
Order Agaricales Underw.
Family Hygrophoraceae Lotsy
Subfamily Lichenomphaloideae L¨
ucking & Redhead
Arrhenia Fr.; type: A. auriscalpia (Fr.) Fr. (bryophylous). Mol.:
yes (T, L). Most species bryophylous or phycophilous.
390 The Bryologist 121(3): 2018
Species: 1-0-0 (40). peltigerina (Peck) Redhead, Lutzoni,
Moncalvo & Vilgalys [”Omphalina p. (Peck) P.Collin]: on
Peltigera.
Lit.: Barrasa & Rico 2010; Collin & Lauron 1994; Garnier-Delcourt
2008; Orton 1977; Redhead et al. 2002; Santesson 1993a.
Eonema Redhead, L¨
ucking & Lawrey; type: E. pyriforme. Mol.:
yes (T, L).
Species: 0-0-1. pyriforme (M.P.Christ.) Redhead, L¨
ucking
& Lawrey [”Athelia pyriformis (M.P.Christ.) J ¨
ulich]:
facultatively lichenicolous on Cetraria aculeata.
Lit.: Khodosovtsev et al. 2018.
Family Tricholomataceae Lotsy
Gamundia Raitelh.; type: G. pseudoclusilis (Joss. & Konrad)
Raitelh. (saprotrophic) [possibly a synomym of G. striatula
(K¨
uhner) Raitelh.]. Mol.: yes. G. leucophylla (Fr.) H.E.Bi-
gelow and G. xerophila (L¨
uthi & R¨
ollin) Raithelh. are only
doubtfully lichenicolous on moribund Peltigera.
Species: 0-0-1 (10). striatula (K¨
uhner) Raithelh.: faculta-
tively lichenicolous on Cladonia and Peltigera.
Lit.: Alstrup & Hawksworth 1990; Barrasa & Rico 2010; Moreau
2005.
Order Atheliales J¨
ulich
Family Atheliaceae J¨
ulich
Athelia Pers.; type: A. epiphylla. Mol.: yes (T, L). Sexual and
asexual (sclerotia-forming).
Species: 0-0-4 (100). alnicola (Bourd. & Galz.) J¨
ulich:
facultatively lichenicolous; arachnoidea (Berk.) J ¨
ulich:
frequently lichenicolous, on various lichens, also over
bryophytes or bark; epiphylla Pers.: facultatively lichen-
icolous; salicum Pers.: facultatively lichenicolous.
Lit.: Arvidsson 1976, 1978; J¨
ulich 1972; Lawrey et al. 2007;
Parmasto 1998.
Order Boletales E.-J.Gilbert
Family Coniophoraceae Ulbr.
Leucogyrophana Pouzar; type: L. mollusca (Fr.) Pouzar
(saprotrophic). Mol.: yes (T, L). Sexual and asexual
(sclerotia-forming).
Species: 1-0-0 (10). lichenicola Thorn, Malloch & Ginns:
on Cladonia and Stereocaulon.
Lit.: Lawrey et al. 2007; Thorn et al. 1998.
Order Cantharellales G¨
aum.
Order Cantharellales genera incertae sedis
Adamflakia Diederich & Lawrey; type: A. applanata [”Bulbilla
Diederich, Flakus & Etayo, nom. inval.; type: B. applanata].
Mol.: yes (T, L). Asexual (bulbil-forming).
Species: 1-0-0. applanata Diederich & Lawrey [”Bulbilla
applanata Diederich, Flakus & Etayo, nom. inval.]: on
Peltigerales.
Lit.: Diederich et al. 2014a; Lawrey et al. 2016.
Burgella Diederich & Lawrey; type: B. flavoparmeliae. Mol.: yes
(T, L). Asexual (bulbil-forming).
Species: 1-0-1. flavoparmeliae Diederich & Lawrey: on
Flavoparmelia baltimorensis;lutea Diederich, Capdet,
A.I.Romero & Etayo: facultatively lichenicolous over
various lichens.
Lit.:Diederich& Lawrey2007; Diederichetal.2014a;Lawrey et al. 2007.
Burgellopsis Diederich & Lawrey; type: B. nivea. Mol.: yes (T,
L). Asexual (bulbil-forming).
Species: 1-0-0. nivea Diederich & Lawrey: on sterile,
crustose lichen.
Lit.: Diederich et al. 2014a.
Burgoa Goid.; type: B. verzuoliana Goid. (saprotrophic). Mol.:
yes (T, L). Asexual (bulbil-forming). Related to Sistotrema
Fr. s.l. (sexual)
Species: 1-0-0 (7). angulosa Diederich, Lawrey & Etayo: on
various lichens.
Lit.: Diederich & Lawrey 2007; Lawrey et al. 2007.
Ceratobasidium D.P.Rogers; type: C. calosporum S.P.Rogers
(saprotrophic). Mol.: yes (L). Sexual and asexual (sclerotia/
bulbil-forming).
Species: 1-0-0 (10). bulbillifaciens Diederich & Lawrey: on
various lichens.
Lit.: Diederich et al. 2014a.
Minimedusa Weresub & P.M.LeClair; type: M. polyspora
(Hotson) Weresub & P.M.LeClair (saprotrophic). Mol.:
yes (T, L). Asexual (bulbil-forming).
Species: 1-0-0 (3). pubescens Diederich, Lawrey & Heylen:
on various lichens.
Lit.: Diederich & Lawrey 2007; Lawrey et al. 2007.
Neoburgoa Diederich, E.Zimm. & Lawrey; type: N. freyi. Mol.:
yes (T, L). Asexual (bulbil-forming).
Species: 1-0-0. freyi Diederich, E.Zimm. & Lawrey: on
Cladonia rangiferina.
Lit.: Lawrey et al. 2016.
Order Corticiales K.-H.Larss.
Family Corticiaceae Herter
Corticium Pers.; type: C. roseum Pers. Mol.: yes (T, L). Sexual.
Species: 1-0-0 (20). silviae Diederich, E.Zimm. & Lawrey:
on Thamnolia.
Lit.: Diederich et al. 2018.
Erythricium J.Erikss. & Hjortstam; type: E. laetum (P.Karst.)
J.Erikss. & Hjortstam (saprotrophic) [¼Marchandiobasi-
dium Diederich & Schultheis; type: M. aurantiacum]. Mol.:
yes (T, L). Sexual and asexual (bulbil-forming).
Species: 1-0-0 (7). aurantiacum (Lasch) D.Hawksw. &
A.Henrici [”Marchandiomyces aurantiacus (Lasch) Die-
derich & Etayo; ¼Marchandiobasidium aurantiacum
Diederich et al.: 2018 classification and checklist of lichenicolous fungi 391
Diederich & Schultheis]: on Physcia and rarely other
lichens.
Lit.: Diederich 1990; Diederich & Lawrey 2007; Diederich et al.
2003; Hawksworth & Henrici 2015; Lawrey et al. 2008.
Laetisaria Burds.; type: L. fuciformis (McAlpine) Burds.
(graminicolous). Mol.: yes (T, L). Sexual or asexual
(bulbil-forming).
Species: 2-0-0 (7). buckii (Diederich & Lawrey) Diederich,
Lawrey & Ghobad-Mejhad [”Marchandiomyces buckii
Diederich & Lawrey]: on various lichens; lichenicola
Diederich, Lawrey & Van den Broeck: on Physcia.
Lit.: Diederich & Lawrey 2007; Diederich et al. 2011a.
Lawreymyces L¨
ucking & Moncada; type: L. palicei. Mol.: yes (T,
L). Sexual, asexual, hyphal and yeast stages unknown.
Endolichenic. Cannot be detected or identified without
DNA sequences. Not validly published (Hawksworth et al.
2017). DNA sequences obtained from the host of the
generic type (Agonimia foliacea) led previous authors to
erroneously conclude that the host was a lichenized
member of Corticiaceae.
Species: 7-0-0. bogotensis L¨
ucking & Moncada: endoli-
chenic in Normandina sp.; columbiensis L¨
ucking &
Moncada: endolichenic in Normandina columbiensis;con-
fusus L¨
ucking & Moncada: endolichenic in Normandina
aff. columbiensis;foliaceae L¨
ucking & Moncada: endoli-
chenic in Agonimia foliacea;palicei L¨
ucking & Moncada:
endolichenic in Agonimia foliacea;pulchellae L¨
ucking &
Moncada: endolichenic in Normandina pulchella;spribillei
L¨
ucking & Moncada: endolichenic in Agonimia foliacea.
Lit.: L¨
ucking & Moncada 2017.
Marchandiomyces Diederich & D.Hawksw.; type: M. corallinus.
Mol.: yes (T, L). Asexual (bulbil-forming).
Species: 1-0-0 (3). corallinus (Roberge) Diederich &
D.Hawksw.: on various lichens.
Lit.: Diederich & Lawrey 2007; Etayo & Diederich 1996a; Hawks-
worth 1979a; Lawrey et al. 2007; Sikaroodi et al. 2001.
NOMENCLATURAL AND TAXONOMIC NOVELTIES
After the revision of the genus Didymocyrtis,
including now many species that have a former
Phoma species as the asexual stage (Ertz et al. 2015),
it became obvious that both Phoma lobariae and P.
psoromatis do not belong there. Instead, the
morphological characters of both perfectly fit the
current circumscription of Abrothallus, including
species mainly known from the asexual stage that
were formerly described in Vouauxiomyces. The two
following new combinations become therefore
necessary.
Abrothallus lobariae (Diederich & Etayo) Diederich
& Ertz, comb. nov.
BASIONYM:Phoma lobariae Diederich & Etayo, in
Etayo & Diederich, Flechten Follmann, Contri-
butions to Lichenology in Honour of Gerhard
Follmann: 213 (1995)
MYCOBANK MB 827766
Abrothallus psoromatis (Zhurb. & U.Braun) Die-
derich & Zhurb., comb. nov.
BASIONYM:Phoma psoromatis Zhurb. & U.Braun,
Lichenologist 45: 587 (2013)
MYCOBANK MB 827767
Arthonia polydactylonis Diederich & Ertz, nom.
nov.
REPLACED SYNONYM:Arthonia ceracea Etayo & Breuss,
¨
Osterr. Z. Pilzk. 7: 206 (1998) nom. illeg. non
Arthonia ceracea F´
ee, Essai Crypt. Ecorces Exot.
Offic., Suppl. R´
evis. (Paris): 42 (1837)
MYCOBANK MB 827768
TYPE: CANADA. BRITISH COLUMBIA:WellsGray
Provincial Park, Trophy Mountain Recreation
Area, 518460N, 1198540W, 1800-1900 m, on
Peltigera polydactylon, 24 Aug. 1994. O. Breuss
10.658 (LI–holotype, UPS–isotype, non vid.).
Asteroglobulus pyramidalis (Etayo) Diederich,
comb. nov.
BASIONYM:Cornutispora pyramidalis Etayo, Opuscula
Philolichenum 8: 134 (2010)
MYCOBANK MB 827769
Based on the original description and a further
specimen studied by us, this species does not belong
to Cornutispora, but is a typical member of
Asteroglobulus.
Specimen examined: AZORES. SAO MIGUE:
Furnas, on the shore of Lagoa das Furnas, on bark
of Cryptomeria japonica,onParmotrema robustum,
30 July 2010, P. Diederich 16968 (herb. Diederich).
Bachmanniomyces D.Hawksw., Bull. Brit. Mus.
(Nat. Hist.), Bot. 9: 10 (1981); type: B.
uncialicola (Zopf) D.Hawksw.
SYN.NOV.: Phaeopyxis Rambold & Triebel, Notes Roy.
Bot. Gard. Edinburgh 46: 380 (1990); type P.
punctum (A.Massal.) Rambold, Triebel & Cop-
pins
Pino-Bodas et al. (2017) included the type
species of both genera in a molecular phylogenetic
analysis and found that both cluster together in a
well-supported clade within the Ostropomycetidae,
and their results suggested that Bachmanniomyces
uncialicola represents the asexual stage of Phaeo-
pyxis punctum. Consequently, we consider both
392 The Bryologist 121(3): 2018
species and both genera to be synonymous. As
Bachmanniomyces (1981) predates Phaeopyxis
(1990), and after the deletion of Art. 57.2 in the
new Shenzhen Code (ICN), the name Bachmannio-
myces has to be used for this genus in the absence of
a formal conservation proposal. The following new
combinations and synonymy become therefore
necessary:
Bachmanniomyces australis (Rambold & Triebel)
Diederich & Pino-Bodas, comb. nov.
BASIONYM:Phaeopyxis australis Rambold & Triebel,
Notes Roy. Bot. Gard. Edinburgh 46: 381 (1990)
MYCOBANK MB 827770
Bachmanniomyces carniolicus (Arnold) Diederich
& Pino-Bodas, comb. nov.
BASIONYM:Biatora carniolica Arnold, in Głowacki,
Verh. K. K. Zool.-Bot. Ges. Wien 20: 453 (1870);
”Phaeopyxis carniolica (Arnold) Rambold &
Triebel, Notes Roy. Bot. Gard. Edinburgh 46:
382 (1990)
MYCOBANK MB 827771
Bachmanniomyces muscigenae (Alstrup & E.S.Han-
sen) Diederich & Pino-Bodas, comb. nov.
BASIONYM:Phaeopyxis muscigenae Alstrup & E.S.Han-
sen, Graphis Scripta 12: 44 (2001)
MYCOBANK MB 827772
Bachmanniomyces punctum (A.Massal.) Diederich
& Pino-Bodas, comb. nov.
BASIONYM:Nesolechia punctum A.Massal., Schedulae
criticae in lichenes exsiccatos Italiae: 96 (1856);
”Phaeopyxis punctum (A.Massal.) Rambold,
Triebel & Coppins, Notes Roy. Bot. Gard.
Edinburgh 46: 384 (1990)
SYN.NOV.: Phyllosticta uncialicola Zopf, Hedwigia 35:
324 (1896); ”Bachmanniomyces uncialicola
(Zopf) D.Hawksw., Bull. Brit. Mus. (Nat. Hist.),
Bot. 9: 11 (1981)
MYCOBANK MB 827773
Bachmanniomyces varius (Coppins, Rambold &
Triebel) Diederich & Pino-Bodas, comb. nov.
BASIONYM:Phaeopyxis varia Coppins, Rambold &
Triebel, Notes Roy. Bot. Gard. Edinburgh 46:
385 (1990)
MYCOBANK MB 827774
Didymocyrtis grumantiana (Zhurb. & Diederich)
Zhurb. & Diederich, comb. nov.
BASIONYM:Phoma grumantiana Zhurb. & Diederich,
in Diederich et al., Lichenologist 39: 160 (2007)
MYCOBANK MB 827775
Although no sequences are available from this
species, it strongly resembles other Phoma species
recently included in Didymocyrtis and is most
probably closely related to them.
Epithamnolia atrolazulina (Etayo) Diederich,
comb. nov.
BASIONYM:Hainesia atrolazulina Etayo, Opera Lilloa-
na 50: 205 (2017)
MYCOBANK MB 827776
Suija et al. (2018) revised the lichenicolous
Hainesia Ellis & Sacc. species and suggested includ-
ing them in an enlarged concept of the genus
Epithamnolia Zhurb. Hainesia atrolazulina obviously
belongs to the same genus, but is distinguished from
all other species by the dark blue, instead of
brownish conidiomatal wall (Etayo 2017).
Gyalolechia epiplacynthium (Etayo) Diederich,
comb. nov.
BASIONYM:Fulgensia epiplacynthium Etayo, Guineana
16: 170 (2010)
MYCOBANK MB 827777
This species was described for a lichenized taxon
with a reduced or sometimes almost absent thallus,
growing parasitically over Placynthium, otherwise
strongly resembling Fulgensia bracteata. As the genus
Fulgensia has recently been shown to be a younger
synonym of Gyalolechia, and as molecular data
placed F. bracteata close to the generic types of both
Gyalolechia and Fulgensia (Arup et al. 2013), this
new combination becomes necessary.
Divakar et al. (2015) showed that Phacopsis s.l.
(see Triebel et al. 1995) represents three phylogenet-
ically distinct genera, Phacopsis s.str., Nesolechia and
the new Raesaenenia. Divakar et al. (2017) used a
temporal phylogenetic method to harmonize family-
and genus-level classification and concluded that
Nesolechia should be considered as a synonym of
Punctelia,andRaesaenenia of Protousnea. Molecular
data obtained from Perˇ
soh & Rambold (2002) are still
unsufficient for Phacopsis s.str., although Divakar et
al. (2015) found, based on a single ITS sequence, that
the generic type P. vulpina formed a sister relationship
with the Relicina þPseudoparmelia clade.
Diederich et al.: 2018 classification and checklist of lichenicolous fungi 393
As Nesolechia was the sister clade of Punctelia,and
Raesaenenia of Protousnea, in the phylogenetic tree
obtained by Divakar et al. (2017), and as both lichen-
icolous genera are morphologically and anatomically
rather different from the lichenized genera, we continue
accepting Nesolechia and Raesaenenia as distinct genera.
The three taxa combined below were described
within Phacopsis s.l., but are oviously congeneric
with the type of Nesolechia, and consequently are
combined there.
Nesolechia doerfeltii (Alstrup & P.Scholz) Dieder-
ich, comb. nov.
BASIONYM:Phacopsis doerfeltii Alstrup & P.Scholz, in
Scholz, Sauteria 9: 38 (1998)
MYCOBANK MB 827779
Nesolechia falcispora (Triebel & Rambold) Die-
derich, comb. nov.
BASIONYM:Phacopsis falcispora Triebel & Rambold, in
Triebel et al., Bryologist 98: 76 (1995)
MYCOBANK MB 827780
Nesolechia oxyspora var. fusca (Triebel & Rambold)
Diederich, comb. nov.
BASIONYM:Phacopsis oxyspora var. fusca Triebel &
Rambold, in Triebel et al., Bryologist 98: 80 (1995)
MYCOBANK MB 827781
Phaeoseptoria Speg.
Quaedvlieg et al. (2013) and Phookamsak et al.
(2014) showed in multigene phylogenetic analyses
that sequences of the type of Phaeosphaeria (sexual)
and Phaeoseptoria (asexual) group together and
consequently synonymized Phaeoseptoria under
Phaeosphaeria. However, after the deletion of Art.
57.2 in the new Shenzhen Code (ICN), the earlier
name Phaeoseptoria (1908) must be used for these
species, and not Phaeosphaeria (1909).
Although no sequences are available from
Phaeoseptoria peltigerae, the only known lichenicolous
species of the genus, we nevertheless regard it here as
a typical member of Phaeoseptoria,basedonmor-
phological resemblances with the generic type.
Preussia peltigerae (Brackel) Diederich, comb. nov.
BASIONYM:Sporormiella peltigerae Brackel, Ber. Bayer.
Bot. Ges. 79: 36 (2009)
MYCOBANK MB 827782
Kruys & Wedin (2009) proposed to treat
Sporormiella Ellis & Everh. as a synonym of Preussia
Fuckel, based on multigene phylogenetic analyses.
Consequently the only known lichenicolous species
of Sporormiella needs to be combined in Preussia.
Sclerococcum Fr. : Fr., Syst. mycol. 1: XL (1821);
type: S. sphaerale (Ach. : Fr.) Fr.
SYN.NOV.: Dactylospora K¨
orb., Syst. Lich. Germ.: 271
(1855); type: D. floerkei K¨
orb. [¼D. parasitica
(Fl¨
orke) Zopf]
Dactylosporaceae Bellem. & Hafellner, Cryptog.
Mycol. 3: 79 (1982); type: Dactylospora K¨
orb.
SYN.NOV.: Sclerococcaceae R´
eblova
´,Unter.&
W.Gams, Mycol. Progr. 16: 35 (2017); type:
Sclerococcum Fr. : Fr.
Diederich et al. (2013) presented a first
phylogenetic study of Sclerococcum in which five
species were transferred to Cladophialophora.The
generic type, S. sphaerale, was found to be a
member of a strongly supported clade that also
included the two non-lichenicolous marine Dacty-
lospora,D. haliotropha and D. mangrovei,andan
unidentified rock isolate. Schoch et al. (2009) found
that two further sequences of the lichenicolous D.
lobariella and the saprotrophic D. imperfecta
belonged to another lineage, concluding that the
genus Dactylospora was polyphyletic. R´
eblova
´et al.
(2017) erroneously concluded that the generic type
of Dactylospora, the lichenicolous D. parasitica,
must belong to the clade also containing the
lichenicolous D. lobariella,andthatthefamily
name Dactylosporaceae consequently refers to that
lineage. Subsequently, they described the new
family Sclerococcaceae for the lineage containing
thetypeofSclerococcum and the two marine
Dactylospora species. However, Pino-Bodas et al.
(2017) have shown that the generic type D.
parasitica clusters with S. sphaerale,bothspecies
being included in a lineage with the lichenicolous
D. ahtii,D. deminuta,D. glaucomarioides,D.sp.
RP391 and the saprotrophic D. haliotrepha,D.
mangrovei and D. vrijmoediae.Ourownanalyses
including newly generated sequences of D. para-
sitica confirm its close relationship with Sclerococ-
cum sphaerale (Fig. 4). Moreover, our newly
generated sequences of D. lobariella cluster with
D. parasitica suggesting that the published nuLSU
sequence of D. lobariella by Schoch et al. (2009) is a
394 The Bryologist 121(3): 2018
contaminant. In our phylogenetic tree (Fig. 4), this
sequence (‘Dactylospora lobariella 40)isplacedin
the outgroup, thus outside of the Sclerococcales
because it belongs to the Chaethotyriales. The
presumed polyphyly of Dactylospora (Schoch et al.
2009) seems thus to be based on contaminant
sequences. Two species were reported by Schoch et
al. (2009) as belonging to another lineage: D.
lobariella, for which we can prove, based on newly
obtained sequences, that the sequence is a contam-
inant, and the saprotrophic D. imperfecta,forwhich
we had no way to check the quality of the sequence,
but we suspect a similar explanation. Moreover, the
published mtSSU sequence of D. deminuta
(KY661681) was not included in our phylogenetic
analysis (Fig. 4) because it is obviously also a
contaminant with a Chaetothyriales species, while
the ITS sequence places the species close to other
Dactylospora s.str. As a result, in Pino-Bodas et al.
(2017), the single specimen of D. deminuta is placed
in the Dactylosporaceae but as the earliest diverging
taxon of the family because the mtSSU pulls it out
of the core group, giving the impression that it
mightrepresentadistinctgenus.
In Pino-Bodas et al. (2017) and in our
phylogenetic tree (Fig. 4), D. parasitica is heteroge-
neous, rending the correct phylogenetic position of
the type of Dactylospora within the Dactylosporaceae
uncertain. One specimen (‘D. parasitica 20in Fig. 4)
appears to be so close to Sclerococcum sphaerale that
it might represent its sexual stage.
As a conclusion, Dactylospora þSclerococcum
appears to represent a monophyletic group, for
which the family name Dactylosporaceae has to be
used, Sclerococcaceae becoming a new synonym.
Further, the two genera Sclerococcum and Dactylo-
spora cannot be kept as separate. As Sclerococcum
(1821) predates Dactylospora (1855), and after the
deletion of Art. 57.2 in the new Shenzhen Code
(ICN), the name Sclerococcum has to be used for
this genus. A proposal to conserve the more
species-rich genus Dactylospora would not likely
be accepted, as Sclerococcum currently includes 20
species, as many recent papers dealt with the
taxonomy of Sclerococcum species, as several recent
papers dealt with the molecular phylogeny of
Sclerococcum,andasthegenusisthetypeofthe
order Sclerococcales. For the moment, we recognize
a single genus in Dactylosporaceae because no
morphological characters support the recognition
of more than one genus. Indeed, Hafellner (1979)
studied many Dactylospora species in his Karschia
monograph, including lichenicolous and saprotro-
phic species, and the genus appeared as morpho-
logically homogeneous. Now, almost 40 years later,
no attempt of splitting the genus or of excluding
some species has been done, and many more species
have been described that fit the genus as circum-
scribed by Hafellner (1979). Molecular phylogenetic
results (except one sequence that we prefer to
consider as doubtful) also support the hypothesis of
the monophyly of the genus.
The following new name and new combinations
become therefore necessary for the lichenicolous
taxa:
Sclerococcum acarosporicola Ertz & Diederich,
nom. nov.
REPLACED SYNONYM:Karschia acarosporae H.Magn.,
Svensk Bot. Tidskr. 37: 285 (1943) non Scle-
rococcum acarosporae S.Y.Kondr., in Kondratyuk
& Zelenko, Ukr. Bot. Journ. 59: 600 (2002); ”
Dactylospora acarosporae (H.Magn.) Hafellner,
Nova Hedw., Beih. 62: 96 (1979)
MYCOBANK MB 827701
Sclerococcum aeruginosum (Holien & Ihlen) Ertz &
Diederich, comb. nov.
BASIONYM:Dactylospora aeruginosa Holien & Ihlen, in
Ihlen et al., Bryologist 107: 358 (2004)
MYCOBANK MB 827702
Sclerococcum ahtii (Zhurb. & Pino-Bodas) Ertz &
Diederich, comb. nov.
BASIONYM:Dactylospora ahtii Zhurb. & Pino-Bodas,
in Pino-Bodas et al., Persoonia 39: 102 (2017)
MYCOBANK MB 827703
Sclerococcum allantoideum (Alstrup & D.Hawksw.)
Ertz & Diederich, comb. nov.
BASIONYM:Dactylospora allantoidea Alstrup &
D.Hawksw., Meddel. Grønland Biosci. 31: 23 (1990)
MYCOBANK MB 827704
Sclerococcum amygdalariae (Triebel) Ertz & Die-
derich, comb. nov.
BASIONYM:Dactylospora amygdalariae Triebel, Bib-
lioth. Lichenol. 35: 197 (1989)
MYCOBANK MB 827705
Diederich et al.: 2018 classification and checklist of lichenicolous fungi 395
Figure 4. Phylogenetic relationships among 24 samples of Sclerococcales resulting from a Bayesian analysis based on a data set of nuLSU and mtSSU
sequences. Penicillium javanicum, Pyrgillus javanicus and Sphinctrina turbinata were chosen as outgroup. Posterior probabilities are shown near internal
branches. Internal branches, considered strongly supported (Posterior probabilities 0.95 ) are represented by thicker lines. The newly sequenced samples
are in bold. The genus Sclerococcum (including Dactylospora, considered here as a synonym of Sclerococcum) is highlighted with a colored box.
396 The Bryologist 121(3): 2018
Sclerococcum anziae (Zhurb., Ezhkin, Skirina &
Y.Ohmura) Ertz & Diederich, comb. nov.
BASIONYM:Dactylospora anziae Zhurb., Ezhkin, Skir-
ina & Y.Ohmura, Folia Cryptog. Estonica 54: 13
(2017)
MYCOBANK MB 827706
Sclerococcum aspiciliicola (Alstrup & D.Hawksw.)
Ertz & Diederich, comb. nov.
BASIONYM:Dactylospora aspiciliicola Alstrup &
D.Hawksw., Meddel. Grønland Biosci. 31: 24 (1990)
MYCOBANK MB 827707
Sclerococcum athallinum (M¨
ull.Arg.) Ertz & Die-
derich, comb. nov.
BASIONYM:Buellia athallina M¨
ull.Arg., Mem. Soc.
Phys. Gen`
eve 16: 64 (1862); ”Dactylospora
athallina (M¨
ull.Arg.) Hafellner, Beih. Nova
Hedwigia 62: 99 (1979)
MYCOBANK MB 827708
Sclerococcum attendendum (Nyl.) Ertz & Diederich,
comb. nov.
BASIONYM:Lecidea attendenda Nyl., Flora 49: 419
(1866); ”Dactylospora attendenda (Nyl.) Ar-
nold, Flora 57: 108 (1874)
MYCOBANK MB 827709
Sclerococcum australe (Triebel & Hertel) Ertz &
Diederich, comb. nov.
BASIONYM:Dactylospora australis Triebel & Hertel, in
Triebel, Biblioth. Lichenol. 35: 205 (1989)
MYCOBANK MB 827710
Table 6. Species names, voucher specimens and GenBank accession numbers for the specimens included in the phylogenetic inference depicted in Fig. 4.
GenBank accession numbers in bold refer to sequences generated by this project.
Name Voucher nuLSU mtSSU Host for lichenicolous fungi
Pyrgillus javanicus AFTOL-ID 342 DQ823103 FJ225774
Penicillium javanicum AFTOL-ID 429 EF413621 FJ225778
Sphinctrina turbinata AFTOL-ID 1721 EF413632 FJ713611
Fusichalara minuta CBS 709.88 KX537758 KX537762
Rhopalophora clavispora 1 CBS 129.74 KX537755 KX537759
Rhopalophora clavispora 2 CBS 281.75 KX537756 KX537760
Dactylospora ahtii 1 USA, Talbot CHI17-37a (H), RP182 NA KY661687 Cladonia rangiferina
Dactylospora ahtii 2 Iceland, Hognabba 1325a (H), RP23 KY661659 KY661686 Cladonia uncialis subsp.
biuncialis
Dactylospora glaucomarioides Russia, Zhurbenko 13107 (LE), RP275 KY661660 KY661683 Ochrolechia akagiensis
Dactylospora haliotrepha 1 AFTOL-ID 798 FJ713617 NA
Dactylospora haliotrepha 2 AFTOL-ID 758 FJ176855 KJ766382
Dactylospora imperfecta AFTOL-ID 5006 FJ176896 NA
Dactylospora lobariella 1 Canary Islands, Ertz 13794 (BR)MH698497 NA Lobaria pulmonaria
Dactylospora lobariella 2 Austria, Diederich 17708 (hb Diederich) MH698498 MH698502 Lobaria pulmonaria
Dactylospora lobariella 3 France, Diederich 18109 (hb Diederich) MH698499 MH698503 Lobaria pulmonaria
Dactylospora lobariella 4 AFTOL-ID 2137 FJ176891 NA ?
Dactylospora mangrovei AFTOL-ID 2108; CBS 110444 FJ176890 KJ766383
Dactylospora parasitica 1 Russia, Tugi (LE), RP422 KY661666 KY661690 Ochrolechia sp.
Dactylospora parasitica 2 Russia, Zhurbenko 12135 (LE), RP423 NA KY661691 Ochrolechia sp.
Dactylospora parasitica 3 Finland, Pykala 39145 (H), RP424 KY661667 KY661692 Ochrolechia sp.
Dactylospora parasitica 4 Belgium, Ertz 16080 (BR)MH698500 MH698504 Ochrolechia turneri
Dactylospora parasitica 5 Canary Islands, van den Boom 46349 (hb v.d.Boom) MH698501 MH698505 on sorediate crust cf.
Pertusaria flavicans
Dactylospora sp RP391 Chile, Buck 60495a (H) KY661664 KY661689 Cladonia rangiferina
Dactylospora sp KLP-2013 isolate NTOU4002 KC692153 NA
Ascomycete ALr-1 isolate ALr-1 EF210108 NA Lecanora rupicola
Sclerococcum sphaerale 1 Luxembourg, Diederich 17279 (hb Diederich) JX081672 JX081677 Pertusaria corallina
Sclerococcum sphaerale 2 Belgium, Ertz 17425 (BR) JX081674 JX081676 Pertusaria corallina
Sclerococcum A1016 Austria, Muggia & Fleischhacker (GZU) KT263077 KT263115 lichen thalli
Sclerococcum A1153 Austria, Muggia MF071425 MF085485 Pertusaria corallina
Diederich et al.: 2018 classification and checklist of lichenicolous fungi 397
Sclerococcum boreale (Holien & Ihlen) Ertz &
Diederich, comb. nov.
BASIONYM:Dactylospora borealis Holien & Ihlen, in
Ihlen et al., Bryologist 107: 359 (2004)
MYCOBANK MB 827711
Sclerococcum cladoniicola (Alstrup & Olech) Ertz &
Diederich, comb. nov.
BASIONYM:Dactylospora cladoniicola Alstrup & Olech,
Polish Polar Res. 14: 35 (1993)
MYCOBANK MB 827712
Sclerococcum crassum (Sarri ´
on & Hafellner) Ertz &
Diederich, comb. nov.
BASIONYM:Dactylospora crassa Sarri ´
on & Hafellner,
Lichenologist 34: 361 (2002)
MYCOBANK MB 827713
Sclerococcum davidii (Hafellner & H.Mayrhofer)
Ertz & Diederich, comb. nov.
BASIONYM:Dactylospora davidii Hafellner &
H.Mayrhofer, Biblioth. Lichenol. 95: 233 (2007)
MYCOBANK MB 827714
Sclerococcum deminutum (Th.Fr.) Ertz & Diederich,
comb. nov.
BASIONYM:Lecidea parasitica var. deminuta Th.Fr.,
¨
Ofvers. Kongl. Vetensk.-Akad. F¨
orh. 13: 127
(1856); ”Dactylospora deminuta (Th.Fr.) Trie-
bel, Biblioth. Lichenol. 35: 209 (1989)
MYCOBANK MB 827715
Sclerococcum dobrowolskii (Olech & Alstrup) Ertz
& Diederich, comb. nov.
BASIONYM:Dactylospora dobrowolskii Olech & Alstr-
up, Polish Polar Res. 17: 166 (1996)
MYCOBANK MB 827716
Sclerococcum frigidum (Hafellner) Ertz & Diederich,
comb. nov.
BASIONYM:Dactylospora frigida Hafellner, Herzogia 7:
166 (1985)
MYCOBANK MB 827717
Sclerococcum glaucomarioides (Tuck.) Ertz & Die-
derich, comb. nov.
BASIONYM:Buellia glaucomarioides Tuck., Synops.
North Americ. Lich. 2: 108 (1888); ”Dactylo-
spora glaucomarioides (Tuck.) Hafellner, Beih.
Nova Hedwigia 62: 109 (1979)
MYCOBANK MB 827718
Sclerococcum hafellnerianum (S´
erus.) Ertz & Die-
derich, comb. nov.
BASIONYM:Dactylospora hafellneriana S´
erus., in
S´
erusiaux & Wessels, Mycotaxon 19: 499 (1984)
MYCOBANK MB 827719
Sclerococcum heterodermiae (Etayo) Ertz & Die-
derich, comb. nov.
BASIONYM:Dactylospora heterodermiae Etayo, Opera
Lilloana 50: 165 (2017)
MYCOBANK MB 827720
Sclerococcum homoclinellum (Nyl.) Ertz & Die-
derich, comb. nov.
BASIONYM:Lecidea homoclinella Nyl., Flora 55: 361
(1872); ”Dactylospora homoclinella (Nyl.)
Hafellner,Beih.NovaHedwigia62:112
(1979)
MYCOBANK MB 827721
Sclerococcum inconspicuum (Etayo) Ertz & Die-
derich, comb. nov.
BASIONYM:Dactylospora inconspicua Etayo, Biblioth.
Lichenol. 98: 79 (2008)
MYCOBANK MB 827722
Sclerococcum inquilinum (Tuck.) Ertz & Diederich,
comb. nov.
BASIONYM:Buellia inquilina Tuck., Lich. Calif. 32
(1866); ”Dactylospora inquilina (Tuck.) Hafell-
ner, Beih. Nova Hedwigia 62: 116 (1979)
MYCOBANK MB 827723
Sclerococcum lobariellum (Nyl.) Ertz & Diederich,
comb. nov.
BASIONYM:Lecidea lobariella Nyl., Flora 52: 296
(1869); ”Dactylospora lobariella (Nyl.) Hafell-
ner, Beih. Nova Hedwigia 62: 118 (1979)
MYCOBANK MB 827724
Sclerococcum microsporum (Etayo) Ertz & Die-
derich, comb. nov.
BASIONYM:Dactylospora microspora Etayo, Candollea
46: 391 (1991)
MYCOBANK MB 827725
398 The Bryologist 121(3): 2018
Sclerococcum orygmaeum (Nyl.) Ertz & Diederich,
comb. nov.
BASIONYM:Lecidea orygmaea Nyl., Flora 52: 296
(1869); ”Dactylospora orygmaea (Nyl.)
S.Y.Kondrat. & D.J.Galloway, Biblioth. Lichenol.
57: 336 (1995)
MYCOBANK MB 827726
Sclerococcum parasitaster (Nyl.) Ertz & Diederich,
comb. nov.
BASIONYM:Lecidea parasitaster Nyl., Flora 58: 105
(1875); ”Dactylospora parasitaster (Nyl.) Ar-
nold, Flora 58: 346 (1875)
MYCOBANK MB 827727
Sclerococcum parasiticum (Fl ¨
orke) Ertz & Die-
derich, comb. nov.
BASIONYM:Lecidea parasitica Fl¨
orke, Deutsche Li-
chenen 6: 3 (1819); ”Dactylospora parasitica
(Fl¨
orke) Zopf, Hedwigia 35: 159 (1896)
MYCOBANK MB 827728
¼Dactylospora floerkei K¨
orb., Systema lichenum
Germaniae: 271 (1855), nom. illeg. superfl.
SYN.NOV.: Sclerophyton occidentale Herre [as occi-
dentalum], Bryologist 43: 27 (1940). TYPE: U.S.A.
CALIFORNIA: Santa Clara Co., Los Gatos, on
Quercus californica,onOchrolechia turneri
(identification kindly confirmed by M. Kukwa
based on photographs), 30.ix.1907, A. Herre
1183 (F– lichenicolous apothecia, lectotype here
designated; M! – isolectotype).
Notes: The holotype is now kept in F(kindly
examined by T. Lumbsch and compared with
our photos from the M specimen) and entirely
agrees with an isotype kept in Mexamined by us.
The type material represents a thallus of Ochro-
lechia turneri parasitized by apothecia of Sclero-
coccum parasiticum. As Herre’s name is based on
a mixture of a lichen and a lichenicolous fungus,
we herewith lectotypify it on the lichenicolous
apothecia present in the type specimen kept in F.
Sclerococcum parellarium (Nyl.) Ertz & Diederich,
comb. nov.
BASIONYM:Lecidea parellaria Nyl., Flora 59: 239
(1876); ”Dactylospora parellaria (Nyl.) Arnold,
Flora 60: 300 (1877)
MYCOBANK MB 827729
Sclerococcum pertusariicola (Tuck.) Ertz & Die-
derich, comb. nov.
BASIONYM:Buellia pertusariicola Tuck. [as pertusar-
icola], Synops. North Americ. Lich. 2: 108
(1888); ”Dactylospora pertusariicola (Tuck.)
Hafellner, Beih. Nova Hedwigia 62: 122 (1979)
MYCOBANK MB 827730
Sclerococcum pleiospermum (Triebel) Ertz & Die-
derich, comb. nov.
BASIONYM:Dactylospora pleiosperma Triebel, in Ha-
fellner, Mycotaxon 84: 300 (2002)
MYCOBANK MB 827731
Sclerococcum polysporum (Triebel) Ertz & Die-
derich, comb. nov.
BASIONYM:Dactylospora polyspora Triebel, Biblioth.
Lichenol. 35: 212 (1989)
MYCOBANK MB 827732
Sclerococcum porphyreum (Hafellner & Kalb) Ertz
& Diederich, comb. nov.
BASIONYM:Dactylospora porphyrea Hafellner & Kalb,
in Hafellner, Herzogia 7: 169 (1985)
MYCOBANK MB 827733
Sclerococcum protothallinum (Anzi) Ertz & Die-
derich, comb. nov.
BASIONYM:Abrothallus prothallinus Anzi, Catal. Lich.
Sond.: 116 (1860); ”Dactylospora protothallina
(Anzi) Hafellner, Beih. Nova Hedwigia 62: 124
(1979)
MYCOBANK MB 827734
Sclerococcum purpurascens (Triebel) Ertz & Die-
derich, comb. nov.
BASIONYM:Dactylospora purpurascens Triebel, Bib-
lioth. Lichenol. 35: 214 (1989)
MYCOBANK MB 827735
Sclerococcum pyrenaicum (Etayo) Ertz & Diederich,
comb. nov.
BASIONYM:Dactylospora pyrenaica Etayo, Guineana
16: 147 (2010)
MYCOBANK MB 827736
Sclerococcum rhyparizae (Arnold) Ertz & Diederich,
comb. nov.
Diederich et al.: 2018 classification and checklist of lichenicolous fungi 399
BASIONYM:Dactylospora rhyparizae Arnold, Flora 57:
108 (1874)
MYCOBANK MB 827737
Sclerococcum rimulicola (M¨
ull.Arg.) Ertz & Die-
derich, comb. nov.
BASIONYM:Buellia rimulicola M¨
ull.Arg., Flora 55: 500
(1872); ”Dactylospora rimulicola (M¨
ull.Arg.)
Hafellner, Beih. Nova Hedwigia 62: 126 (1929)
MYCOBANK MB 827738
Sclerococcum rostrupii (Alstrup) Ertz & Diederich,
comb. nov.
BASIONYM:Dactylospora rostrupii Alstrup, in Alstrup
et al., Frodskaparrit 40: 89 (1994)
MYCOBANK MB 827739
Sclerococcum saxatile (Schaer.) Ertz & Diederich,
comb. nov.
BASIONYM:Calicium saxatile Schaer., Naturwiss. Anz.
Allg. Schweiz. Ges. Gesammten Naturwiss. 5: 35
(1821); ”Dactylospora saxatilis (Schaer.) Ha-
fellner, Beih. Nova Hedwigia 62: 129 (1979)
MYCOBANK MB 827740
Sclerococcum saxatile var. laureri (Hafellner) Ertz &
Diederich, comb. nov.
BASIONYM:Dactylospora saxatilis var. laureri Hafell-
ner, Beih. Nova Hedwigia 62: 135 (1979)
MYCOBANK MB 827741
Sclerococcum suburceolatum (Coppins & Fryday)
Ertz & Diederich, comb. nov.
BASIONYM:Dactylospora suburceolata Coppins & Fry-
day, Lichenologist 44: 725 (2012)
MYCOBANK MB 827742
Sclerococcum tegularum (Arnold) Ertz & Diederich,
comb. nov.
BASIONYM:Buellia tegularum Arnold, Ber. Bayer. Bot.
Ges. 1 (suppl.): 130 (1891); ”Dactylospora
tegularum (Arnold) Hafellner, Beih. Nova Hed-
wigia 62: 149 (1979)
MYCOBANK MB 827743
Sclerococcum urceolatum (Th.Fr.) Ertz & Diederich,
comb. nov.
BASIONYM:Buellia urceolata Th.Fr., Lich. Arctoi 3: 333
(1860)
MYCOBANK MB 827744
The following new species has been contributed
by Dr B. Coppins:
Sclerococcum ophthalmizae Coppins, sp. nov.
MYCOBANK MB 827745
Differs from S. parasitica in having 16-spored asci,
ascospores to 7-septate, and a different internal
pigmentation.
TYPE: BRITISH ISLES. SCOTLAND:WESTERNESS (VC
97): N side of Loch Sunart, c. 10 km WSW of
Salen, Rubha Aird Druimnich, NM597610, 0–30
m elev., on thallus of Lepra ophthalmiza on
Betula trunk, 8.iii.1983, B.J. Coppins 9274 &P.M.
Jørgensen (E– holotype; GZU – isotype).
Ascomata 0.12-0.4 mm diam., scattered, black;
disc concave to flat; margin smooth. Exciple 25-30
lm wide, reddish brown (K–) or green tinged in
upper part; cells to 10 lm diam. Hymenium 50-60
lm tall, colourless; epihymenium brown, K–.
Hypothecium pale grey- or reddish-brown, K–.
Paraphyses 1.5–1.7 lm thick; apices to 3 lm. Asci
c. 50 312-17 lm, 16-spored. Ascospores (10.5–)12–
18 33.5–4.5 lm, narrowly ellipsoid, oblong or
fusiform, sometimes curved, brown, smooth-walled
or slightly roughened when old, 3(–7)-septate.
Distribution and host: Known from Scotland
(Isle of Skye, Westerness: Loch Sunart, West Ross),
always on the thallus of Lepra ophthalmiza.
Additional specimens examined: BRITISH
ISLES. SCOTLAND. WESTERNESS (VC 97): N side of
Loch Sunart, Glenborrodale, RSPB Nature Reserve,
NM6060, on Quercus, 15.vi.1992, B.J. Coppins 15293
(E). KINTYRE (VC 101): Knapdale, Loch Coill-Bharr,
NR780897, birch-oak wood, on Quercus, 22.x.2001,
B.J.&A.M. Coppins 20460 (E). WEST ROSS (VC 105):
Inverpolly NNR, N side of Allt Geann an t-Strathain,
NC090163, 30–50 m elev., SE-facing hazel pasture
woodland, on large boulder, 8.vi.1999, B.J.&A.M.
Coppins 18765 (E, sub Lepra ophthalmiza); Beinn
Eighe NNR, Coille na Glas-leitir, NE of Cnoc na
Gaoithe, NG9964, c. 150 m elev., 14.iv.2001, B.J.&
A.M. Coppins 19731 (E); Loch Maree, Eilean Dubh
na Sr `
oine. NG910720. 15 m elev., on Pinus,
31.v.1999, B.J. Coppins 18555 & V.J. Giavarini (E,
sub Lepra ophthalmiza).
400 The Bryologist 121(3): 2018
Although this species has never been formally
described, the name Dactylospora ophthalmizae
Coppins has been used for many years by British
lichenologists, as it is included in the keys to British,
Iberian and Canarian lichenicolous fungi (Hawks-
worth et al. 2010a).
We further combine in Sclerococcum three non-
lichenicolous Dactylospora species that have been
included in recent phylogenetic analyses (Diederich
et al. 2013; Pino-Bodas et al. 2017; R´
eblova
´et al.
2017) and that belonged to the same clade as the
type:
Sclerococcum haliotrephum (Kohlm. & E.Hohlm.)
Ertz & Diederich, comb. nov.
BASIONYM:Buellia haliotrepha Kohlm. & E.Kohlm.,
Nova Hedwigia 9: 90 (1965); ”Dactylospora
haliotrepha (Kohlm. & E.Kohlm.) Hafellner,
Beih. Nova Hedwigia 62: 111 (1979)
MYCOBANK MB 827746
Sclerococcum mangrovei (E.B.G.Jones, Alias, Abdel-
Wahab & S.Y.Hsieh) Ertz & Diederich, comb.
nov.
BASIONYM:Dactylospora mangrovei E.B.G.Jones, Alias,
Abdel-Wahab & S.-Y.Hsieh, in Jones et al.,
Mycoscience 40: 317 (1999)
MYCOBANK MB 827747
Sclerococcum vrijmoediae (K.L.Pang, S.Y.Guo,
Alias, Hafellner & E.B.G.Jones) Ertz & Diederich,
comb. nov.
BASIONYM:Dactylospora vrijmoediae K.L.Pang,
S.Y.Guo, Alias, Hafellner & E.B.G.Jones, Botan-
ica Marina 57: 317 (2014)
MYCOBANK MB 827748
The genus Libertiella Speg. & Roum. with the
type L. malmedyensis Speg. & Roum. resembles
mesoconidial stages of Scutula so much that there is
no doubt that both genera are synonyms (Triebel et
al. 1997). As all currently accepted species of Scutula
are confined to hosts belonging to Peltigerales, we
combine here in Scutula two species of Libertiella
described from Peltigera. We refrain from combining
a further species, L. fennica Alstrup, also growing on
Peltigera, as it differs by the ovoid or pyriform to
subglobose conidia, distinct from conidia of all other
known Scutula species, as well as L. leprariae Etayo &
Diederich (on Lepraria) and the poorly known L.
obscurior Gerber (on Brodoa) known from hosts not
belonging to the Peltigerales.
Scutula curvispora (D.Hawksw. & Mia
˛dl.) Die-
derich, comb. nov.
BASIONYM:Libertiella curvispora D.Hawksw. & Mia
˛dl.,
Mycol. Res. 101: 1127 (1997)
MYCOBANK MB 827783
Scutula didymospora (D.Hawksw. & Mia
˛dl.) Die-
derich, comb. nov.
BASIONYM:Libertiella didymospora D.Hawksw. &
Mia
˛dl., Mycol. Res. 101: 1128 (1997)
MYCOBANK MB 827784
Stigmidium haesitans (Nyl.) Diederich, comb. nov.
BASIONYM:Verrucaria haesitans Nyl., in Kihlman,
Meddeland. Soc. Fauna Fl. Fenn. 18: 59 (1891);
”Pharcidia haesitans (Nyl.) Vouaux, Bull. Soc.
Mycol. France 28: 249 (1912)
MYCOBANK MB 827785
Mayrhofer (1987: 336) reported Pharcidia hae-
sitans from Sweden and wrote that it belonged to
Stigmidium, fide Hafellner (pers. comm.). Later, the
species was incorrectly referred to as S. haesitans
(Nyl.) Hafellner (Santesson 1993a), although such a
combination was never published. This error was
later addressed by Santesson et al. (2004) who used
the same name and authorities, by adding ‘‘comb.
not published’’. The name is hereby validly com-
bined in Stigmidium.
Stigmidium parvum (Henssen) Diederich, comb.
nov.
BASIONYM:Pharcidia parva Henssen, Symb. Bot.
Upsal. 18: 112 (1963)
MYCOBANK MB 827786
Santesson (1993a) named this taxon ‘‘Stigmi-
dium parvum (Henssen) (Provisionally placed here,
Art. 34.1b)’’, followed by the basionym, but without
a full reference to that basionym. As this author
clearly did not intend to publish a new combination,
and according to Art. 41.1 of ICN, the new
combination was not validly published. We herewith
validly combine the species in Stigmidium.
ACKNOWLEDGMENTS
We wish to thank all the colleagues who over years drew our
attention to errors or missing data in our draft online catalogue
Diederich et al.: 2018 classification and checklist of lichenicolous fungi 401
(www.lichenicolous.net), and especially Drs. Franz Berger (Kopfing,
Austria), Wolfgang von Brackel (R¨
ottenbach, Germany), Adam
Flakus (Krak ´
ow, Poland), Martin Kukwa (Gdansk, Poland), Robert
L¨
ucking (Berlin, Germany), Mikhail Zhurbenko (St. Petersburg,
Russia) and two anonymous referees for carefully checking the
present manuscript. Brian Coppins (Edinburgh, UK), Adam Flakus,
Raquel Pino-Bodas (Madrid, Spain) and Mikhail Zhurbenko kindly
contributed to the nomenclatural and taxonomic novelties proposed
here. The curator of the herbarium Mis thanked for the loan of a
type of Sclerophyton occidentale and T. Lumbsch (Chicago, U.S.A.)
for examining the type of the same species kept in F.
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manuscript received May 16, 2018; accepted August 17, 2018.
Supplementary document online:
Supplementary Table S1. Summary statistics for
all taxa listed in the 2018 classification and checklist
of lichenicolous fungi in spreadsheet format.
Diederich et al.: 2018 classification and checklist of lichenicolous fungi 425
