The 2016 classification of lichenized fungi in the Ascomycota and

Basidiomycota – Approaching one thousand genera

Robert Lücking1,2,6, Brendan P. Hodkinson3,4 and Steven D. Leavitt2,5

1
Botanic Garden and Botanical Museum, Königin-Luise-Str. 6–8, 14195 Berlin, Germany; 2 Research Associate,
Integrative Research Center, Science & Education, The Field Museum, 1400 South Lake Shore Drive, Chicago, IL 60605,
U.S.A.; 3 Department of Dermatology, University of Pennsylvania, BRB 1046A, 421 Curie Blvd., Philadelphia, PA
19104, U.S.A.; 4 International Plant Science Center, New York Botanical Garden, Bronx, NY 10458, U.S.A.;
5
Department of Biology, Brigham Young University, 4102 LSB, Provo, UT 84604, U.S.A.

ABSTRACT. Ninety years after Zahlbruckner, we present the most recent update to the classification of
lichen fungi in the Ascomycota and Basidiomycota to genus level, with species numbers and references to
changes compared to the 2010 Outline of Ascomycota and other recent classifications. Updated statistics on
global species richness of lichen fungi and species richness at family, order and class level are given. The
number of accepted species is 19,387 in 995 genera, 115 families, 39 orders and eight classes. Lichenized
Basidiomycota amount to 172 species (0.9% of the total), 15 genera (1.5%), five families (4.3%), five
orders (12.8%) and one class (12.5%). The most speciose genera are Xanthoparmelia, Lecanora, Arthonia,
Cladonia, Pertusaria, Ocellularia, Graphis, Caloplaca, Usnea and Buellia. The average number of species
per genus is 19.5 and 256 genera are monospecific. Using newly defined categories, two genera
(Xanthoparmelia, Lecanora) are ultradiverse (more than 500 species), 17 hyperdiverse (201–500 species)
and 12 megadiverse (101–200). The largest family is Parmeliaceae, with 2,765 species and 77 genera,
followed by Graphidaceae (2,161; 79), Verrucariaceae (943; 43), Ramalinaceae (916; 43) and Lecanoraceae
(791; 25). The largest order is Lecanorales, with 6,231 species and 234 genera, followed by Ostropales
(3,261; 138), Arthoniales (1,541, 103), Peltigerales (1,301; 67) and Caliciales (1,276; 55). The largest class is
Lecanoromycetes, with 15,131 species and 701 genera, followed by Arthoniomycetes (1,541; 103),
Eurotiomycetes (1,203; 63), Dothideomycetes (812; 39) and Lichinomycetes (390; 50). A total of 751 out
of 995 genera (75%) have molecular data. Fifty-nine genera remain in unresolved positions at the family,
order or class level. The phylogenetic position of the 39 orders containing lichenized fungi suggests 20–30
independent lichenization events during the evolution of higher Fungi, 14–23 in the Ascomycota and 6–7
in the Basidiomycota. The following names are validated: Candelariomycetidae Mia˛dl. et al. ex Timdal &
M.Westb. subcl. nov., Cystocoleaceae Locq. ex Lücking, B.P.Hodk. & S.D.Leav. fam. nov, Letrouitineae
Gaya & Lutzoni subordo nov., Rhizocarpales Mia˛dl. & Lutzoni ordo nov. and Teloschistineae Gaya &
Lutzoni subordo nov. Lectotypes are designated for Clathroporinopsis M.Choisy and Protoschistes
M.Choisy, making both synonyms of Gyalecta Ach., and Stromatothelium Trevis., making it a synonym of
Pyrenula Ach. Members of Cyphobasidiales, which are here interpreted as hyperlichenized fungi, as well as
fossil lichen fungi, are added in additional classifications in two appendices.

KEYWORDS. Global species richness, lichen fungal systematics, nomenclature.

^ ^ ^

Lichenization is one of the major lifestyles among
Fungi. The currently recognized more than 19,000
species of lichen fungi make up 17% of the known
6
Corresponding author’s e-mail: r.luecking@bgbm.org
DOI: 10.1639/0007-2745-119.4.361
110,000 fungal species and 27% of the known
Ascomycota (Feuerer & Hawksworth 2007; Jaklitsch
et al. 2016a). Lichenized Basidiomycota were con-
sidered to be species-poor (Lawrey et al. 2009;
Oberwinkler 2012), but recent findings show that
some lineages are highly diverse, containing hun-

The Bryologist 119(4), pp. 361–416 Published online: January 3, 2017 0007-2745/16/$5.75/0
Copyright Ó2016 by The American Bryological and Lichenological Society, Inc.
362 The Bryologist 119(4): 2016
dreds of taxa (L ücking et al. 2014, 2016b).
Phylogenetic evidence suggests that lichenization
evolved several times independently in both Asco-
mycota and Basidiomycota (Hodkinson et al. 2014;
Lawrey et al. 2009; Lumbsch & Rikkinen 2016;
Nelsen et al. 2009, 2011a; Prieto et al. 2013; Schoch
et al. 2009). Molecular clock studies indicate that
modern lichenized fungi first appeared in the
Permian and Triassic, about 300–200 mya, and large
macrolichens such as the Parmeliaceae in the Late
Cretaceous (Amo de Paz et al. 2011; Beimforde et al.
2014; Lücking et al. 2009a; Prieto & Wedin 2013). In
contrast, some controversial hypotheses postulate
that lichen-like organisms existed in the Precambri-
an (Hallbauer et al. 1977; Retallack 1994, 1995, 2007,
2012) and some studies suggest that so-called
protolichens may have evolved as early as 450 mya,
possibly giving rise to large lineages of non-liche-
nized fungi (Eriksson 2005; Lipnicki 2015; Lumbsch
& Rikkinen 2016). The protolichen hypothesis has
recently received support through the discovery of
415 my old lichen fossils presumably belonging in
the Pezizomycotina (Honegger et al. 2013; Taylor et
al. 2015).
Many historical and modern classifications of
lichen fungi have been published, starting with the
works of Acharius, Nylander, Zahlbruckner, and
others, which treated lichens as separate group of
organisms, and continuing with the gradual integra-
tion of lichen fungi into the fungal system over the
course of the 20th century. After the Genera of Fungi
(Clements 1909; Clements & Shear 1931), the first
comprehensive, modern attempt at classifying all
lichen fungi to genus level within the Fungi was the
Outline of the Ascomycota, started by Eriksson
(1982a) as Outline of the Ascomycetes in the journal
Mycotaxon and continued by Eriksson & Hawks-
worth (1985) and Eriksson et al. (2001) in the
journals Systema Ascomycetum and Myconet (Hawks-
worth & Eriksson 1994). The Outline of the
Ascomycota was taken over by Lumbsch & Huhndorf
(2007) and is currently published in the journal
Fieldiana, with the last edition six years ago
(Lumbsch & Huhndorf 2010). The classification
and accompanying notes are also available online
[http://archive.fieldmuseum.org/myconet].
Along with the emergence of molecular phylo-
genetics over the past two decades, the classification
of Ascomycota and Basidiomycota seemed to go
separate ways, with the Basidiomycota adopting a
trend to apply informal clade names, rather than
formal Linnean taxonomy (Moncalvo et al. 2002).
This was consolidated by Hibbett et al. (2007), who
present a formal classification to order level for all
Fungi. Since then, most papers have adopted formal
classifications, but a regular platform such as the
Outline of the Ascomycota does not yet exist for the
Basidiomycota and the lower Fungi. The most recent
works summarizing classifications of both Ascomy-
cota and Basidiomycota (the two phyla containing
lichen fungi) are the Dictionary of the Fungi (Kirk et
al. 2008), The Mycota (McLaughlin & Spatafora
2014a,b) and the Syllabus of Plant Families (Jaklitsch
et al. 2016a).
Although lichen fungi do not form a taxonomic
entity, we believe that a classification of only
lichenized taxa is useful, since many workers deal
with either lichenized, lichenicolous or non-liche-
nized fungi and a synoptic classification of taxa with
a particular lifestyle is desirable as a quick reference.
After numerous classification proposals by earlier
workers in the 18th and 19th century, the first
comprehensive classification of the lichen fungi to
genus level was compiled a century ago by
Zahlbruckner (1907, 1926) for the early editions of
A. Engler & K. Prantl’s Natürliche Pflanzenfamilien.
Further classifications were published, for instance,
by Henssen & Jahns (19737), Poelt (19747), Tehler
(1996) and Tehler & Wedin (2008), but these were
either not entirely complete or included other
groups, such as lichenicolous fungi, or focused on
other aspects, such as ascoma ontogeny. In order to
fill the gap of a dedicated classification of all lichen
fungi, we here begin a regularly updated series that
contains a combined treatement for Ascomycota and
Basidiomycota to genus level. In addition to a
hierarchic listing, the classification is accompanied
by cladograms of all higher Fungi to order level,
summarizing evidence from phylogenetic studies
and highlighting lichenized lineages. The classifica-
tion is also available as an editable spreadsheet
(Supplementary Table S1), and the cladograms
including statistics as printable poster (Supplemen-
tary Fig. S2). Besides statistics on species richness
7
Henssen & Jahn’s book ‘‘Lichenes. Eine Einführung in
die Flechtenkunde’’ was officially published in early 1974,
but Aino Henssen in December 1973 obtained a number of
copies and distributed these to various colleagues, which
can be considered effective publication according to ICN
29.1. In contrast, Ahmadjian & Hale’s book ‘‘The
Lichens’’, which contains Poelt’s classification, was sched-
uled for publication in 1973 but was effectively published
in early 1974 only.

from phylum down to genus level, other novel
functional aspects of this classification are refined
richness categories, the calculation of completeness
of molecular data at genus level, and the proportion
of genera with still unresolved positions, indices
which will serve as indicators for the progress of
future classifications. In addition, for the first time,
also exclusively fossil genera supposedly representing
lichen fungi are included in this classification.
DELIMITATION OF THE SUBJECT: WHAT CONSTITUTES A
LICHEN?
What constitutes a lichen has been controversial
and widely debated. Most often used is the definition
by Hawksworth (1988: 17): ‘‘A lichen is a stable self-
supporting association of a mycobiont and a photo-
biont in which the mycobiont is the exhabitant,’’ as
well as the one by Goward (1994: 17), stating that
lichens are ‘‘... fungi that have taken up agriculture.’’
(Will-Wolf & Lücking 2016). Both describe what
lichens are and what they do. A very broad view of
lichens was expressed by Retallack (1994, 1995, 2007,
2012), by considering Ediacaran fossils such as
Dickinsonia lichens. His reasoning was far outside
of any accepted definition of lichens; instead,
Ediacaran fossils were considered lichen-like because
of their supposedly stiff consistency and their growth
on soil, a strongly disputed view (Antcliffe & Hancy
2013; Callow et al. 2013; Fedonkin & Waggoner
1997; Lücking et al. 2009a; Taylor et al. 2015;
Waggoner 1995). The term mycobiont is phyloge-
netically defined, since mycobionts by default are
part of the kingdom Fungi. A phylogenetic definition
of the photobiont, on the other hand, is not possible,
since photobionts encompass both eukaryotic green
algae and prokaryotic cyanobacteria. In addition,
green algae are a paraphyletic assembly basal to
embryophytes (bryophytes and vascular plants), so a
phylogenetic definition of the term photobiont could
include stable, lichen-like fungal-bryophyte associa-
tions within the lichens.
To resolve this dilemma, two further aspects
should be (and often are) added to the definition of
a lichen. One is morphogenesis, since lichenization
results in a novel morphology not known from the
separate bionts. At first glance, this might exclude
some filamentous and gelatinous lichens, such as
Coenogonium and Collema species, but even in the
algal-like, filamentous Coenogonium, the cultured
alga looks different from that in the lichen thallus
Lücking et al.: 2016 lichen classification 363
(Uyenco 1965), which indicates that morphogenesis
takes place. The second aspect is the body plan of the
photobiont. Lichens are distinguished from other
fungal-plant symbioses by a photobiont with the
body plan of mostly undifferenciated, unicellular to
multicellular microalgae. In contrast, macroalgae,
bryophytes, and vascular plants involved in fungal
associations are differentiated into organs. A still
controversial case would be bryophilous fungi
associating with moss protonemata, such as in the
genus Arrhenia (Redhead et al. 2002), since proto-
nemata do have the body plan of undifferentiated
microalgae and such associations could be consid-
ered lichens. Coincidentally, Arrhenia s.l. is the
paraphyletic ancestor of the lichenized Dictyonema
clade (Dal Forno et al. 2013; Lawrey et al. 2009).
Several other cases of borderline, lichen-like
associations are known in which the mycobiont-
photobiont definitions apply, but the association is
either not stable or no morphogenetic effect is
apparent, or it is unclear whether the association
represents a bipartite lichen or a lichenicolous fungus
on an otherwise sterile lichen thallus (Grube & Wedin
2016; Lawrey & Diederich 2016). These include,
among others, Antennulariella lichenisata Coppins &
Aptroot (Coppins & Aptroot 2008), Dacampia hookeri
(Borrer) A.Massal. (Ertz et al. 2015b), Milospium
graphideorum (Nyl.) D.Hawksw. (Aptroot & Sipman
2001; Crous et al. 2015), Psammina palmata Earl.-
Benn. & D.Hawksw. (Earland-Bennett & Hawksworth
2005), Requienella seminuda (Pers.) Boise (Aptroot
1991; Jaklitsch et al. 2016b), Robergea pupula (Tuck.)
R.C.Harris (Lücking et al., unpubl. data), Schizoxylon
albescens Gilenstam, Döring & Wedin (Muggia et al.
2011a, 2016), Sclerococcum griseosporodochium Etayo
(Smith 2009; Diederich et al. 2013), ’Spiloma’
auratum Sm. (Diederich 2013) and Stigmidium
marinum (Deakin) Swinscow (Van Herk & Aptroot
2004). In most cases, these are single species in genera
and even higher lineages that otherwise contain non-
lichenized forms, so while they are few in number,
they add considerably the the phylogenetic diversity
of lichen formers.
The recent discovery of what appears to be a
regular, second fungal component present in macro-
lichens of the family Parmeliaceae and possibly other
lineages (Spribille et al. 2016) has sparked a new
discussion of what may constitute a lichen. Contrary
to the presence of a functional bacterial microbiome
in lichens (Aschenbrenner et al. 2016; Grube &
Wedin 2016; Grube et al. 2015; Hodkinson et al.
364 The Bryologist 119(4): 2016
2012), in analogy to the human microbiome, this
additional fungal component, invariably represent-
ing taxa in the newly established order Cyphobasi-
diales (genus Cyphobasidium), seems to have
structural and possibly even chemical implications
on the lichen thallus. Should we then consider this
fungus lichenized and include it in lichen fungal
classifications? By definition, a lichenized fungus
associates with a photobiont. Since Cyphobasidium
appears to be restricted to the upper cortical layer of
the lichen and is not in contact with the photobiont,
it cannot be considered lichenized in the proper
sense. Rather, in analogy to hyperparasites (Kiss
2001), this fungus could be considered hyperliche-
nized, since it associates with a fungus that is
primarily lichenized with a photobiont. Symbiotic
Cyphobasidium might more logically be placed in a
classification of lichenicolous fungi, but if the
interpretation of this phenomenon (Spribille et al.
2016) is correct, the underlying biology appears to be
quite different. To highlight the discovery of this
hyperlichenization, we have included Cyphobasi-
diales as separate component in this classification.
However, we anticipate that, with more detailed
studies elucidating the predicted high diversity of
these fungi (Spribille et al. 2016), they might
eventually deserve their own classification, as third
group along with lichenized and lichenicolous fungi.
A BRIEF RECAPITULATION OF THE CLASSIFICATION OF
LICHEN FUNGI
Long before the start of fungal nomenclature
with Linnaeus (1753) and later Persoon (1801) and
Fries (1821), the first detailed classification of lichens
was presented by Micheli (1729). Before, Tournefort
(1694, 1698) had already included lichens in two
botanical treatments, with a small number of species,
all under the genus name Lichen. Micheli continued
to use this simple nomenclature, but sorted out at
the level of ordo what he considered to be different
morphological entities that much later would
correspond to genera. Linnaeus (1753) returned to
the basic format presented by Tournefort, although
he included many more species.
Erik Acharius, who is considered the father of
lichenology (Nylander 1858; Vitikainen 1976), was
the first to consequently implement a new classifica-
tion system for lichens, with his seminal works
Lichenographiae Suecicae Prodromus (Acharius 1798),
Methodus (Acharius 1803), Lichenographia Universalis
(Acharius 1810), and Synopsis Methodica Lichenum
(Acharius 1814). His classification was largely based
on morphology of the thallus and fruiting bodies,
placing lichens with similar morphology in the same
genus and naming these genera differently. This
classification was continuously ’refined’ by several
workers, particularly based on microscopic characters
such as ascospores, including Fée (1824, 1837), De
Notatis (1846), Trevisan (1853), Nylander (1854,
1855), Massalongo (1855), Montagne (1856), Müller
(1862), Stizenberger (1862), and Tuckerman (1872).
Nylander (1858) presented an interesting synopsis
with an evolutionary hypothesis, classifying lichens in
a series from most algal-like to most fungal-like and
postulating that lichens were the evolutionary link
between algae and fungi. In doing so, he uninten-
tionally set the stage for what would become one of
the greatest discoveries in biological sciences, the
lichen symbiosis by De Bary and Schwendener (De
Bary 1866; Anonymous 18678), a fact that Nylander
himself never accepted.
The second half of the 19th century was very
productive in describing new species, and many
attempts were made to reclassify particular groups,
resulting in concepts widely used until well into the
present millenium. With the turn of the century, the
state of the art was almost nailed in stone with
Alexander Zahlbruckner’s system in two editions of
Adolf Engler and Karl Prantl’s Natürliche Pflanzen-
familien (Zahlbruckner 1907, 1926), a system that
for many groups prevailed for nearly a century, and
for some is still in existence. Zahlbruckner divided
the lichens into Asco- and Hymenolichenes (i.e.
asco- and basidiolichens) and the Ascolichenes
according to fruiting body morphology into four
large groups with either perithecia, mazaedia, lirellae
or apothecia. Within these groups, the photobiont
type played a major role in separating families. After
Zahlbruckner, Walter Watson came out with an
alternative classification at family level (Watson
1929a,b) which, however, lacked the internal coher-
ence of Zahlbruckner’s system. Nevertheless, several
of the families introduced by Watson are in use
today, albeit often in a very different sense.
Just before the turn of the century, Lindau
(1895), Reinke (1896), Clements (1897) and other
workers fiercely discussed the controversy that had
8
Schwendener’s original contribution is usually cited with
his authorship, but the corresponding publication is an
anonymous third-party report on Schwendener’s lecture.

arisen from the discovery of the lichen symbiosis,
namely whether lichens should be classified sepa-
rately (Reinke) or in the fungal system (Lindau,
Clements). Fink (1911) presented the results of a
questionnaire sent to numerous lichenologists,
mycologists and botanists, finding that 83% of those
asked favored to maintain lichens as a separate
group, whereas 17% preferred distribution among
the fungi. Lichenologists and botanists would favor a
separate classification of lichens, whereas mycolo-
gists would prefer their integration into the fungal
system. The two alternatives found their way into the
formal classifications proposed by Zahlbruckner
(1907, 1926) one one hand and Bessey’s Synopsis of
Plant Phyla (1907) and the Genera of Fungi
(Clements 1909; Clements & Shear 1931) on the
other. Both alternatives continued to be used long
into the 20th century. Nannfeldt’s (1932) work on
ascoma ontogeny and anatomical details of ascomata
and asci then founded the conceptual division of
ascohymenial versus ascolocular fungi, a classifica-
tion adopted by some mycologists and lichenologists
during the 20th century (Barr 1979, 2001; Barr &
Huhndorf 2001; Reynolds 1981).
Some of the controversy over the classification of
lichens as a separate group versus integrated with
closely related fungi was the confusion about what a
lichen actually was, not in the sense of the definition
of the symbiosis, but whether the term ’lichen’
corresponded to the association (consortium) as a
whole or just to the fungal counterpart. Fink (1913:
156) concluded that ‘‘... the lichen is a fungus pure
and simple’’ and offered the following further
definition: ‘‘The lichen is a fungus which lives during
all or part of its life in parasitic relation with the algal
host and also sustains a relation with an organic or an
inorganic substratum.’’ (Fink 1913: 157). Thus, albeit
based on an erroneous biological viewpoint, Fink
came to the correct conclusion that lichens should be
included in the fungal system, only that what he
referred to were not lichens as a whole, but the lichen
fungi. Ciferri & Tomaselli (1953a,b, 1955, 1957)
attempted to resolve this presumed dilemma by
proposing a radically new nomenclature for lichen
fungi, with the fungus in the lichenized stage (i.e.
Fink’s ’lichen’) bearing the scientific name in use and
the isolated fungal component in culture receiving a
new name with the suffix -myces. This controversy
had already been resolved in the Stockholm Code
(Lanjouw et al. 1952), but the newly proposed names
found their way into the literature and only recently
Lücking et al.: 2016 lichen classification 365
it was shown that almost all of them were illegitimate
(Lücking & Hawksworth 2007).
The second half of the 20th century saw further,
highly influential attempts at classifications of lichen
fungi that were often regarded ahead of their time
and not rarely anticipated findings confirmed much
later with molecular methods. Following Nannfeldt
(1932), Santesson (1952) adopted a classification for
foliicolous lichen fungi consequently implementing
mycological rather than lichenological terminology.
Santesson also was the first to abandon the ascospore
concept in lecidoid taxa, where genera used to be
separated based on the number and orientation of
ascospore septa. The genus Tapellaria as defined by
Santesson was the first to include species with very
differently septate ascospores, and the corresponding
plate (Santesson 1952: 495) anticipated what was
later shown to be the rule in numerous genera.
This period also saw a general dispute about the
division of the Ascomycota into ascohymenial and/
or unitunicate versus ascolocular and/or bitunicate
lineages, the so-called Lutrellian concept (Hawks-
worth 1985; Luttrell 1955; Reynolds 1981), with
some questioning the practicality of this system
(Groenhart 1965). Henssen & Thor (1994) showed
that the Arthoniales presented characters of both
ascohymenial and ascolocular lineages. In spite of
highly disparate classification attempts of the
Ascomycota (compared in Hawksworth 1985), there
was an emerging core of orders assigned to the
ascolocular fungi, namely Asterinales, Dothideales,
Hysteriales, Melanommatales, Myriangiales and
Pleosporales (Ainsworth et al. 1971, 1973; Barr
1976, 1983; Hale 1967; Henssen & Jahns 1973;
Reynolds 1981), which later would become the core
of class Dothideomycetes. Hale (1967) was the first
to adopt this concept for lichen-forming fungi,
placing arthonioid taxa, pyrenocarpous taxa with
bitunicate asci, and lirellate-pseudostromatic groups
within the ascolocular fungi. Poelt (1974) and
Hafellner (1988) also followed this approach, and
one of the remarkable components of Hafellner’s
classification was the placement of Gomphillaceae
into a separate order, Gomphillales, postulated to
have bitunicate asci.
A consequence of the early work by Nannfeldt
(1932) and Luttrell (1955) was that, even if authors
did not necessarily adopt the ascohymenial-ascoloc-
ular paradigm, increasing attention was paid to
ascoma ontogeny and ascus structure and their value
for taxonomy and systematics, particularly of lichen-
366 The Bryologist 119(4): 2016
forming fungi (Döring & Lumbsch 1998; Letrouit-
Galinou 1973; Letrouit-Galinou et al. 1994; Poelt
1974). The dominant figure in the field of ontoge-
netic studies was Aino Henssen, and perhaps the
most underrated (since published in German only)
and at the same time most elaborated and conse-
quential classification of lichen fungi in the 20th
century was presented by Henssen & Jahns (1973),
consistently basing their views on anatomical and
ontogenetic evidence and anticipating many changes
later made or confirmed in the molecular era.
Another groundbreaking work consequently using
ascus structure as systematic character was Hafell-
ner’s Studien in Richtung einer natürlicheren Glieder-
ung der Sammelfamilien Lecanoraceae und
Lecideaceae (Hafellner 1984), dividing the two
collective families into numerous, smaller families.
Although many of these families have now been
subsumed to synonymy, with Hafellner’s (1984)
work, lichenologists never looked at asci the same
way again. Indeed, taking to the (Lugol) bottle has
become basic routine when identifying lichens in
these groups, although misunderstood ontogenetic
and functional variation often makes it difficult to
assess ascus structures correctly. In parallel, second-
ary chemistry strongly influenced lichen taxonomy
during this period (Culberson 1969, 1970; Hawks-
worth 1976; Lumbsch 1998), with major changes
particularly in the classification of the family
Parmeliaceae (Elix 1993).
The 1980s marked a shift in the formal
classification of the Ascomycota (at the time
recognized as class Ascomycetes). The Outline of
the Ascomycetes, since 1999 the Outline of Ascomycota
(Eriksson 1982a,b, 1999a; Lumbsch & Huhndorf
2007a, 2010), became the widely accepted authority
for classification of the Ascomycota, including all
lichen-formers, to genus level. The appeal of this
approach was that it was simply structured and
widely available, that it consequently implemented
formal nomenclature, and that it almost worked like
an ‘‘open source wiki,’’ becoming multi-authored
(Eriksson et al. 2001), with all changes documented
in numbered notes by various authors (Eriksson
1999b). This structure allowed the Outline of
Ascomycota to seamlessly move into the molecular
area (Eriksson 1995). Unfortunately, the Outline of
Ascomycota has not been continued after 2010, but
besides the synoptic classification published by
Hodkinson (2012), Schultz & Resl (2016) have
recently started a new series summarizing changes
to the classification affecting species in Central
Europe, in a style similar to the notes accompanying
the Outline of Ascomycota.
With Armaleo & Clerc’s (1991) study on lichen
photosymbiodemes and Ahmadjian’s (1991) outlook
began the molecular era in the classification of
lichens. Molecular phylogeny of lichen fungi virtu-
ally took off in 1995 (Armaleo & Clerc 1995;
Eriksson & Strand 1995; Gargas & Taylor 1995;
Gargas et al. 1995a, b; Grube et al. 1995; Lutzoni &
Vilgalys 1995a, b; Tehler 1995). Initially, studies
focused on broad fungal phylogenies, shaping the
backbone of both the Ascomycota and Basidiomy-
cota into major lineages later recognized as subphyla
and classes (Hibbett et al. 2007; Lutzoni et al. 2004;
Schoch et al. 2009). More recently, focus shifted
towards class- and order-level studies and particu-
larly the delimitation of genera and species (Crespo
& Lumbsch 2010; Gaya et al. 2012; Gueidan et al.
2009; Lumbsch 2006; Mia˛dlikowska et al. 2006, 2014;
Lumbsch & Rikkinen 2016), gradually implementing
an increasing number of markers and new technol-
ogies such as high throughput sequencing. Apart
from their continuous representation in the Outline
of the Ascomycota and in various classifications of the
Basidiomycota, a separate, comprehensive classifica-
tion of lichen fungi was recently published by Tehler
(1996) and updated by Tehler & Wedin (2008).
Not unexpectedly, the molecular era has brought
major changes to the classification of (lichen) fungi,
much more so than in the plant and animal
kingdoms. The long-standing division of apothecial,
perithecial, hysterothecial, and mazaedial lineages
into major taxa broke down entirely (Lumbsch &
Huhndorf 2007b), with the order Ostropales being
an impressive example of ascoma variation including
almost everything imaginable even in individual
families, such as Graphidaceae (Fig. 1). Likewise, in
the Basidiomycota fruiting body morphology was
shown to have little systematic value, with the
Gasteromycetes undergoing a similar fate as the
Caliciales within the Ascomycota. In analogy to
Ostropales, the order Agaricales is now a striking
example uniting an unexpected diversity of fruiting
body morphology and lifestyles (Fig. 2). The deep
division of the Ascomycota into ascohymenial and
ascolocular fungi, as well as the grouping of
bitunicate lineages, had to be abandoned (Lumbsch
& Huhndorf 2007b). Most of the latter are now
found in the two classes Dothideomycetes and
Eurotiomycetes, which include lineages not previ-
 Lücking et al.: 2016 lichen classification 367

Figure 1. Representatives of Ostropales showing the morphological diversity of taxa confirmed to belong in this order. A. Stictis radiata (Stictidaceae; not
lichenized). B. Coenogonium leprieurii (Coenogoniaceae). C. Porina lucida (Porinaceae). D. Trichothelium argenteum (Porinaceae). E. Aspidothelium
cinerascens (Thelenellaceae). F. Gomphillus hyalinus (Gomphillaceae). G. Aderkomyces albostrigosus (Gomphillaceae). H. Actinoplaca strigulacea
(Gomphillaceae). I. Gyalectidium filicinum (Gomphillaceae). J. Xalocoa ocellata (Graphidaceae). K. Asteristion platycarpum (Graphidaceae). L. Graphis
chrysocarpa (Graphidaceae). M. Phaeographis rubrostroma (Graphidaceae). N. Nadvornikia hawaiiensis (Graphidaceae). O. Polistroma fernandezii
(Graphidaceae). All photographs by Robert Lücking.
368 The Bryologist 119(4): 2016

Figure 2. Representatives of Agaricales showing the morphological diversity of taxa confirmed to belong in this order. Non-lichenized: A. Amanita
muscaria (Amanitaceae). B. Marasmius siccus (Marasmiaceae). C. Coprinellus disseminatus (Psathyrellaceae). D. Termitomyces fuliginosus (Lyophyllaceae).
E. Schizophyllum commune (Schizophyllaceae). F. Fistulina hepatica (Fistulinaceae). G. Lycoperdon perlatum (Agaricaceae). H. Rhodotus palmatus
(Physalacriaceae). I. Typhula erythropus (Typhulaceae). Lichenized (all Hygrophoraceae): J. Lichenomphalina lobata. K. Semiomphalina leptoglossoides. L.
Cyphellostereum pusiolum. M. Acantholichen pannarioides. N. Dictyonema sericeum. O. Cora imi. All photographs by Robert Lücking, except F (Wikimedia
Creative Commons: Kean10), H (Wikipedia Creative Commons: Dan Molter) and I (Steve Trudell, with permission).

ously thought to be related; in addition, Euro-
tiomycetes is closer to the ascohymenial and non-
bitunicate Lecanoromycetes than to Dothideomy-
cetes. Hemisphaeriales, by several authors placed in
Loculoascomycetes, is now a synonym of Xylariales
in the Sordariomycetes. Another major change is the
abandonment of the concept of Deuteromycetes, or
imperfect fungi, with profound impact on fungal
nomenclature (Gams et al. 2003; Hawksworth et al.
2011). Within the lichen-forming fungi, Arthonia-
ceae and Roccellaceae were entirely reclassified (Ertz
et al. 2009, 2015; Ertz & Tehler 2011; Frisch et al.
2014) and some other large families, such as
Collemataceae, Graphidaceae, Lobariaceae, Pannar-
iaceae, Teloschistaceae and Verrucariaceae, have
undergone dramatic changes in generic concepts
(Arup et al. 2013; Ekman et al. 2014; Gueidan et al.
2009; Gaya et al. 2012; Moncada et al. 2013; Otálora
et al. 2014; Rivas Plata et al. 2012, 2013). Species
concepts are being revised for almost every genus
studied, not rarely with surprising results (Crespo &
Lumbsch 2010; Lücking et al. 2014; Moncada et al.
2013, 2014).
Naturally, much of the above discussion con-
centrates on the Ascomycota; after all, lichenized
Basidiomycota contain less than 1% of all lichen
formers. Indeed, basidiolichens were always consid-
ered to be generally species-poor and were often
treated as oddities, for instance by Zahlbruckner
(1907, 1926). However, their importance for the
natural classification of lichens was already recog-
nized by Fink (1913), who used the misguided study
by Möller (1893), which stated that a single species
in the genus Dictyonema was able to form both a
fungus and two different lichens, as proof that
lichens should be integrated into the fungal system.
In modern times, Oberwinkler (1984, 2001, 2012)
periodically offered treatments of basidiolichen
genera. Basidiolichens also played an important role
in the early development of molecular phylogenetic
studies (Gargas et al. 1995a; Lutzoni & Vilgalys
1995a,b) and have been identified as models to study
the evolution of the lichen thallus (Dal Forno et al.
2013). Besides the close relationship of the genera
Dictyonema s.l. and Lichenomphalia s.l. within the
single family Hygrophoraceae in Agaricales (Lawrey
et al. 2009), other substantial changes in basidio-
lichen classification are the segregation of Lepidos-
tromatales (Ertz et al. 2008; Hodkinson et al. 2014)
and the placement of Marchandiomphalina in the
Lücking et al.: 2016 lichen classification 369
corticioid fungi (Pálice et al. 2005; Lawrey et al.
2007, 2008).
Apart from the advent of a powerful new set
of tools, which continues to evolve at amazing
speed, the molecular era brought a remarkable
change to the nature of classifications. Rather than
individually characterized by conceptual approach-
es, classifications rapidly developed into synoptical
summaries of the growing body of evidence from
phylogenetic studies (Hibbett et al. 2007; Lumbsch
& Huhndorf 2007a). The present classification is
no exception. It is less an intellectual work than a
synoptic and integrated data mining exercise, with
the aim to present a snapshot at the current state-
of-the-art, with summary statistics that allow for
substantiated, precise citations of relevant num-
bers.
MATERIALS AND METHODS
We assembled both a hierarchical listing for all
lichenized fungal genera up to phylum level and
reference trees for all orders of higher Fungi
(Ascomycota and Basidiomycota). For the Asco-
mycota, we took the latest edition of the Outline of
Ascomycota (Lumbsch & Huhndorf 2010) as a
base, which was also compared to the classification
proposed by Hodkinson (2012) and the new
edition of the Syllabus of Plant Families (Ascomy-
cota) (Jaklitsch et al. 2016a), including some new
references (e.g., Réblová et al. 2016). For the
Basidiomycota, we used Hibbett et al. (2007), the
Dictionary of the Fungi (Kirk et al. 2008), and
Oberwinkler (2012) as primary references, com-
plemented by recent phylogenetic studies to assess
the position and relationships of orders (Aime et
al. 2006, 2014; Garcı́a-Sandoval et al. 2011;
Hibbett 2006; Riess et al. 2016; Wang et al. 2014,
2015a,b,c).
We then undertook a data mining exercise using
the Recent Literature on Lichens (RLL) database
[http://nhm2.uio.no/botanisk/lav/RLL/RLL.HTM],
Google Scholar [https://scholar.google.de] and Gen-
Bank [http://www.ncbi.nlm.nih.gov/genbank] to
track down all publications with changes to the
classification published since 2000, to also include
several omissions made in the 2010 Outline (for
instance, the synonymy of Dimerella with Coenogo-
nium) All data were assembled in spreadsheet format
(Supplementary Table S1). Genera were classified
within a given higher taxon if there was either
370 The Bryologist 119(4): 2016
published phylogenetic support for that placement,
if there were published (or in a few cases unpub-
lished) sequence data indicating such placement, or
in case of genera without molecular data, if the
morphology strongly suggested such placement. All
other cases were consequently placed ’incertae sedis’
within the lowest defined category. We deliberately
avoided the practice of placing taxa with question
mark within a higher category, as used in previous
versions of the Outline.
We accepted, or in some cases implemented,
changes based exclusively on systematic grounds.
The intention to stabilize nomenclature does not
mean that names long in use should continue to
be used if the topology would permit such use.
Rather, objective measures such as branch length
and support, as well as morphological disparate-
ness should be taken as only criteria to delimit
taxa, and only then should a decision be made
about what name applies to a particular taxon. For
instance, we subsumed Calycidiaceae under syn-
onymy with Sphaerophoraceae, since there is
neither a topological nor a morphological reason
to maintain the two families, even if both entities
are reciprocally monophyletic and Calycidiaceae
has been used for many decades. A similar
example is Thelotremataceae (see note below
under Graphidaceae).
Revising the data presented in the recent Syllabus
(Jaklitsch et al. 2016a), we assigned updated species
numbers to each genus, using either published
figures in recent treatments or revised estimates
based on the Dictionary (Kirk et al. 2008), with
corrections when appropriate. We then computed
species numbers for genera, families, orders and
classes. In some large genera, such as Arthonia and
Verrucaria, these numbers continue to be approxi-
mations. Data mining in GenBank was used to
calculate the number of genera for which molecular
data are available. The final spreadsheet (see
Supplementary Table S1) was edited so that the
fully formatted classification could be automatically
retrieved in text form.
In addition to the main classification of liche-
nized fungi, we present two appendices, one
including the order Cyphobasidiales, here interpret-
ed as hyperlichenized fungi, and one including fossil
lichen fungi (and those intepreted as lichen-forming
fungi) described in fossil genera in which no extant
species are known.
Table 1. Species richness of lichenized fungi in Ascomycota and
Basidiomycota.
Category Fungi Ascomycota Basidiomycota Proportion
Species 19,387 19,215 172 0.9%
Genera 995 980 15 1.5%
Families 115 110 5 4.3%
Orders 39 34 5 12.8%
Classes 8 7 1 12.5%
RESULTS AND DISCUSSION
As of this date, the total number of genera
including lichenized species is tabulated at 995 and
the total number of lichenized species is 19,387
(Table 1; Supplementary Table S1; Supplementary
Fig. S2). Overall, 115 families, 39 orders and eight
classes contain lichenized species (Table 2; Supple-
mentary Table S1; Supplementary Fig. S2). Sordar-
iomycetes has for the first time been reported to
include a species that is supposedly (at least weakly)
lichenized, in the genus Requienella (Aptroot 1991,
2009; Jaklitsch et al. 2016b). This means that now
seven of the 13 classes currently recognized in
Ascomycota include lichen-formers. Lichenized Ba-
sidiomycota amount to 172 species (0.9% of the
total), 15 genera (1.5%), five families (4.3%), five
orders (12.8%) and one class (12.5%; Table 1).
The most speciose lichenized genus continues to
be Xanthoparmelia, followed by Lecanora, Arthonia,
Cladonia, Pertusaria, Ocellularia, Graphis, Caloplaca,
Usnea and Buellia (Table 2; Supplementary Fig. S2).
However, it is likely these numbers will change,
either because of revised species concepts (e.g.,
Xanthoparmelia), genus-level splits (e.g., Graphis,
Lecanora), reassignment of species to other genera,
or all of these reasons (e.g., Arthonia, Buellia,
Caloplaca, Pertusaria). Currently, among these ten
genera, species numbers are most reliable for
Cladonia, Ocellularia and Usnea.
The average number of species per genus is 19.5,
and 256 genera are currently monospecific, 377
contain one or two species, and 582 up to five
species, which means that most genera are small to
very small. We devised logarithmically defined
categories to address species richness at the genus
level more precisely (Table 3). In these categories,
two genera (Xanthoparmelia, Lecanora) are recog-
nized as ultradiverse (more than 500 species), 17 as
hyperdiverse (201–500 species), and 13 as mega-
diverse (101–200; for listing of genus numbers, see
Supplementary Table S1). We anticipate that the
 Lücking et al.: 2016 lichen classification 371

Table 2. The 50 largest genera and families in terms of numbers of lichenized species, and all 40 orders and eight classes containing lichen-formers, in
descending order of species number, with additional listing of genus numbers.

Genera Species Families Species Genera Orders Species Genera

Xanthoparmelia 820 Parmeliaceae 2,765 77 Lecanorales 6,231 234

Lecanora 550 Graphidaceae 2,161 79 Ostropales 3,261 138
Arthonia 500 Verrucariaceae 943 43 Arthoniales 1,541 103
Cladonia 500 Ramalinaceae 916 43 Peltigerales 1,301 67
Pertusaria 500 Lecanoraceae 791 25 Caliciales 1,276 55
Ocellularia 400 Teloschistaceae 755 65 Verrucariales 946 45
Graphis 390 Arthoniaceae 717 20 Pertusariales 907 24
Caloplaca 350 Caliciaceae 675 36 Teloschistales 841 71
Usnea 350 Physciaceae 601 19 Trypetheliales 420 16
Buellia 300 Cladoniaceae 558 15 Lichinales 390 50
Opegrapha 300 Pertusariaceae 502 3 Acarosporales 259 12
Parmotrema 300 Lobariaceae 466 12 Lecideales 249 29
Rinodina 300 Pilocarpaceae 424 29 Monoblastiales 247 6
Verrucaria 300 Trypetheliaceae 420 16 Rhizocarpales 236 6
Hypotrachyna 260 Gomphillaceae 418 24 Pyrenulales 234 13
Astrothelium 242 Pannariaceae 406 28 Umbilicariales 168 14
Bacidia 230 Opegraphaceae 365 15 Trapeliales 155 15
Ramalina 230 Porinaceae 359 6 Agaricales 147 8
Rhizocarpon 225 Lichinaceae 340 46 Strigulales 79 5
Acarospora 200 Roccellaceae 311 41 Candelariales 69 5
Anisomeridium 200 Acarosporaceae 259 12 Hymeneliales 38 3
Aspicilia 200 Stereocaulaceae 252 5 Leprocaulales 35 3
Sticta 200 Monoblastiaceae 247 6 Coniocybales 31 2
Phaeographis 180 Lecideaceae 244 28 Pleosporales 31 3
Pyrenula 170 Megasporaceae 243 6 Sarrameanales 30 3
Fissurina 160 Pyrenulaceae 232 11 Collemopsidiales 26 2
Catillaria 150 Rhizocarpaceae 231 4 Thelocarpales 26 1
Porina 140 Collemataceae 192 10 Baeomycetales 21 6
Stereocaulon 140 Catillariaceae 174 5 Arctomiales 16 3
Polyblastia 120 Hygrophoraceae 147 8 Cantharellales 16 1
Thelotrema 106 Trapeliaceae 112 11 Chaetothyriales 14 3
Lecidea 100 Peltigeraceae 110 2 Vezdaeales 13 1
Micarea 100 Umbilicariaceae 91 4 Phaeomoniellales 8 1
Peltigera 100 Coenogoniaceae 90 1 Lepidostromatales 6 3
Pseudocyphellaria 100 Gyalectaceae 86 4 Capnodiales 5 5
Thelidium 100 Ochrolechiaceae 80 3 Eremithallales 4 3
Fellhanera 97 Strigulaceae 79 5 Atheliales 2 2
Gyalideopsis 95 Candelariaceae 66 5 Xylariales 1 1
Phyllopsora 95 Psoraceae 60 4 Corticiales 1 1
Cora 92 Lecanographaceae 58 7
Coenogonium 90 Icmadophilaceae 55 5 Classes Species Genera
Heterodermia 90 Stictidaceae 54 8
Hypogymnia 90 Fuscideaceae 53 13 Lecanoromycetes 15,131 701
Parmeliella 90 Tephromelataceae 52 4 Arthoniomycetes 1,541 103
Toninia 85 Malmideaceae 51 4 Eurotiomycetes 1,203 63
Lecidella 80 Haematommataceae 50 2 Dothideomycetes 812 39
Lepraria 80 Thelenellaceae 50 3 Lichinomycetes 390 50
Physcia 80 Xylographaceae 43 1 Agaricomycetes 172 15
Pseudosagedia 80 Peltulaceae 40 4 Coniocybomycetes 31 2
Pyxine 75 Coccocarpiaceae 39 3 Sordariomycetes 1 1
372 The Bryologist 119(4): 2016
Table 3. Newly defined categories to assess species richnes in genera of
lichenized fungi.
Category Range Number of genera
Monospecific 1 256
Bispecific 2 121
Nanodiverse 3 to 5
Microdiverse 6 to 10
Paucidiverse 11 to 20
Mesodiverse 21 to 50
Superdiverse 51 to 100
Megadiverse 101 to 200
Hyperdiverse 201 to 500
Ultradiverse 501 to 1,000
placement of genera within these categories will
stabilize with further assessments and hence these
categories can be used both as indices for the
progress of phylogenetic classification and for
communicating species richness patterns more
precisely in studies on ecology, biogeography,
conservation and outreach.
The largest family continues to be Parmeliaceae,
with 2,765 lichenized species and 77 genera (Table 2;
Supplementary Fig. S2), followed by Graphidaceae
(2,161; 79), Verrucariaceae (943; 43), Ramalinaceae
(916; 43) and Lecanoraceae (791; 25). Among these,
Parmeliaceae and Graphidaceae are the most well-
studied in terms of generic classification and species
delimitation, and we predict that even with further
changes, the position of these two families as by far
the two largest among lichenized lineages will not
change. The only potentially larger family among all
Ascomycota is Mycosphaerellaceae, with an estimat-
ed 3,300 species (Jaklitsch et al. 2016a); however, this
family is not well-studied phylogenetically and this
number is likely to change. Among the Basidiomy-
cota, Pucciniaceae has close to 5,000 species (Kirk et
al. 2008), but also this number has no solid
phylogenetic basis.
The largest order of lichen fungi continues to be
Lecanorales, with 6,231 species and 234 genera
(Table 2; Supplementary Fig. S2), followed by
Ostropales (3,261; 138), Arthoniales (1,541, 103),
Peltigerales (1,301; 67) and Caliciales (1,276; 55).
Also here, we predict that even with further changes
in genus and species concepts and the discovery of
additional taxa, there will be no change in the
position of the currently three largest orders. By far
the largest class in terms of genera and species of
lichenized fungi continues to be Lecanoromycetes,
with 15,131 species and 701 genera (Table 2;
Supplementary Fig. S2), followed by Arthoniomy-
cetes (1,541; 103), Eurotiomycetes (1,203; 63),
Dothideomycetes (812; 39) and Lichinomycetes
(390; 50). We predict that the positions of the first
four classes are set in stone, even with further
changes and discoveries, but that Agaricomycetes
205
132 (172; 15), currently in position six, will eventually
103 surpass Lichinomycetes, since 319 species are already
106 recognized in phylogenetic data in this class (Dal
41 Forno 2015; Lücking et al. 2016a), but not all are
12 formally published, and taking the estimate by
17 Lücking et al. (2014) as a base, Agaricomycetes is
2
expected to contain close to 700 lichenized species,
most in the genera Cora and Dictyonema. Even with
new discoveries, we predict that a similar number
will not be reached by Lichinomycetes.
According to our data, 751 out of 995 genera
(75%) currently have molecular sequence data
available, although not always for the type species
or species presumed to be congeneric with the type
(Supplementary Table S1). Filling the gaps with the
remaining 25% of the genera (Table 4) should be a
primary focus of future molecular studies. Accord-
ingly, 59 genera are currently classified as incertae
sedis: 33 at family level, 11 at order level, and 15 at
class level (Supplementary Table S1; Supplementa-
ry Fig. S2).
Compared to the latest Outline of Ascomycota
(Lumbsch & Huhndorf 2010), genus classification
underwent 14 changes at subphylum level, 36 at class
level, 43 at subclass level, 173 at order level, and 135
at family level. A total of 235 genera (24%) were
newly established or reinstated (including a few
ommitted from the 2010 Outline), and new synon-
ymy was established for 63 genera. No changes to the
classification were recorded for 415 genera. Thus,
within only six years, aspects of the classification
changed for more than half of the genera currently
recognized.
A total of 107 and 61 orders are currently
recognized in Ascomycota and Basidiomycota,
respectively (Figs. 3 & 4; Supplementary Fig. S2).
The phylogenetic positions of the 39 orders con-
taining lichenized fungi suggest that between 20 and
30 independent lichenization events took place in
the evolution of higher Fungi, 14–23 in the
Ascomycota and 6–7 in the Basidiomycota (Figs. 3
& 4; Supplementary Fig. S2). Uncertainties in these
numbers stem from the following considerations.
The exact phylogenetic position of the Thelocarpales
and Vezdaeales is unresolved and there is a
 Lücking et al.: 2016 lichen classification 373

Table 4. Genera of lichenized fungi for which currently no molecular sequence data are available.

Aciculopsora Claurouxia Heppsora Orphniospora Scleropyrenium

Adelolecia Clauzadeana Hertella Pachyascus Sculptolumina
Amazonomyces Clauzadella Heterocarpon Paraporpidia Semigyalecta
Amazonotrema Clypeopyrenis Heteromyces Parasiphula Septotrapelia
Ameliella Compsocladium Hippocrepidea Paratricharia Servitia
Amphorothecium Coniangium Homothecium Parmotremopsis Sipmania
Amylora Conotremopsis Hormosphaeria Peltularia Solorinaria
Ancistrosporella Coppinsia Hosseusia Phoebus Sphaerophoropsis
Angiactis Coronoplectrum Hueidea Phylliscidiopsis Spheconisca
Anomalographis Corynecystis Hypoflavia Phylliscidium Sphinctrinopsis
Antennulariella Crocodia Ivanpisutia Phyllisciella Sporopodiopsis
Apatoplaca Crustospathula Jamesiella Phylloblastia Sporostigma
Aplanocalenia Cryptodictyon Japewiella Phyllocratera Squamella
Argopsis Cryptothele Jarmania Phyllogyalidea Stigmidium
Arthopyrenia Culbersonia Kalbiana Physcidia Stirtoniella
Aspiciliopsis Cyanoporina Kalbographa Piccolia Streimannia
Aspilidea Davidgallowaya Kantvilasia Placolecis Sulcopyrenula
Asteroporum Dermiscellum Krogia Placothelium Synalissa
Auriculora Diaphorographis Labyrintha Placynthiopsis Synarthonia
Austrographa Diploschistella Lecidopyrenopsis Podotara Synarthothelium
Awasthiella Dirinastrum Lemmopsis Poeltidea Tania
Bacidiopsora Distopyrenis Leprantha Polistroma Tapellaria
Bactrospora Distothelia Leprocollema Psammina Tapellariopsis
Badimiella Echidnocymbium Lichinodium Psathyrophlyctis Tarbertia
Baflavia Edrudia Lignoscripta Pseudarctomia Tasmidella
Bahianora Edwardiella Lithoglypha Pseudobaeomyces Thallinocarpon
Bartlettiella Endohyalina Lithogyalideopsis Pseudocalopadia Thecographa
Barubria Enterodictyon Loflammia Pseudohepatica Thelenidia
Biatorella Eopyrenula Loflammiopsis Pseudoheppia Thelidiopsis
Blastodesmia Epilichen Logilvia Pseudolecanactis Thelignya
Bouvetiella Eremothecella Lueckingia Pseudopannaria Thelopsis
Brasilicia Eschatogonia Lyromma Pseudopaulia Thermutis
Bryodina Feigeana Magmopsis Pseudopeltula Thermutopsis
Bryogomphus Ferraroa Maronella Psorinia Tibellia
Buelliastrum Finkia Mawsonia Psoromella Toniniopsis
Byssotrema Fluctua Melanophloea Psorotheciopsis Topelia
Caeruleum Follmanniella Metamelanea Pulvinodecton Traponora
Calathaspis Frigidopyrenia Microtheliopsis Punctonora Tylophorella
Calopadiopsis Gloeoheppia Minksia Pyrenocarpon Tylophoropsis
Calotrichopsis Glomerilla Monoblastia Pyrenowilmsia Vainionora
Caprettia Glyphopsis Moriola Racoleus Vezdaea
Catarraphia Gossypiothallon Mycoporum Redonia Wadeana
Catarrhospora Graphidastra Myochroidea Rhabdopsora Xanthopsorella
Catillochroma Gudelia Myrionora Rhaphidicyrtis Xenolecia
Cenozosia Gymnographopsis Neopsoromopsis Rhizolecia Xenus
Cephalophysis Gyrocollema Notocladonia Roccellinastrum Xyleborus
Cheiromycina Halographis Oevstedalia Sagema Xylopsora
Ciposia Haplodina Oletheriostrigula Sarrameana Zahlbrucknerella
Cladidium Haploloma Orcularia Schizodiscus

possibility that they form part of an expanded morphologically and ecologically so distinct that,
Lecanoromycetes, which would reduce the number while both may share a direct, lichenized ancestor,
of separate lichenizations (maximum of 30) by two we also have to assume separate lichenization as a
(–2). Coniocybomycetes and Lichinomycetes are possibility, if both lineages represent relicts of a
374 The Bryologist 119(4): 2016

Figure 3. Synoptical cladogram of currently accepted orders in Ascomycota, shown in two parts. A. Orders excluding Eurotiomycetes and
Lecanoromycetes. B. Orders in Eurotiomycetes and Lecanoromycetes. Connection between A and B indicated. Predominantly or entirely lichenized
orders in green (dark grey with white font in print), partially lichenized orders in light blue (grey in print), and partially delichenized orders (excluding
lichenicolous lineages) indicated by white box. Numbers indicate likely range of lichenization events in the corresponding lineage.

Figure 3. Continued.
larger clade which became mostly extinct (–1).
Capnodiales include at least two unrelated filamen-
tous lichen-forming lineages (Cystocoleus, Racodi-
um); the third genus (Racoleus) might be related to
either one of these or not (–1). There is a possibility
that Chaetothyriomycetidae (Chaetothyriales,
Phaeomoniellales, Pyrenulales, Verrucariales), with
currently six postulated lichenization events, had a
single, lichenized ancestor (–5). This results in up to
nine reductions in the number of possible lichen-
ization events in the Ascomycota (from 23 to 14).
While the Lichenomphalina and Dictyonema clades in
Agaricales clearly represent two independent lichen-
ization events, the assessment for Atheliales is
tentative. Lichenization has been reported for Athelia
and Athelopsis and both genera are not closely
related, suggesting two independent lichenization
events, but the phylogenetic position of the liche-
nized species within these genera has not yet been
assessed. Of the two clearly lichenized species
described in Athelia, A. andina Jülich and A. poeltii
Jülich, the latter has been found to belong in
Dictyonema (Lücking, unpubl. data), so there might
only be one genuine lichenized clade in Atheliales (–
1).
Lücking et al.: 2016 lichen classification 375
While the number of independent lichenization
events within the Fungi is an important detail to
reconstruct the evolution of the lichenized lifestyle,
perhaps a more important aspect is the evolutionary
fate of these lineages. Thus, of the 30 supposedly
independent lichenization events, 11 did not lead to
subsequent radiation, consisting of a single or few
species only that are often questionably lichenized
(e.g., Arthopyrenia, Athelia, Athelopsis, Cystocoleus,
Eremithallus, Marchandiomphalina, Melaspilea, Pyr-
enothrix, Racodium, Racoleus, Requienella); to these,
one might add several genera with unresolved
placement due to lack of molecular data. Eight
lichenization events resulted in a single genus that
diversified into several species (Celothelium, Colle-
mopsidium, Lichenomphalia, Lyromma, Microtheliop-
sis, Multiclavula, Thelocarpon, Vedzaea), eight
further events diversified into several, partially
diverse genera representing a single family or part
thereof (Coniocybaceae, Hygrophoraceae tribe Ar-
rheniae, Lepidostromataceae, Monoblastiaceae, Pyr-
enulaceae, Strigulaceae, Trypetheliaceae, Verru-
cariaceae), two events resulted in orders containing
several families (Arthoniales, Lichinales), and only a
single event diversified into an entire class containing
376 The Bryologist 119(4): 2016

Figure 4. Synoptical cladogram of currently accepted orders in Basidiomycota. Predominantly or entirely lichenized orders in green (dark grey with white
font in print), partially lichenized orders in light blue (grey in print), and hyperlichenized order Cyphobasidiales indicated by white box. Numbers
indicate likely range of lichenization events in the corresponding lineage.

numerous orders and families (Lecanoromycetes).
Thus, although the number of independent lichen-
ization events is fairly high, only nine lineages (30%)
can be considered successful, namely Hygrophor-
aceae tribe Arrheniae, Monoblastiaceae, Pyrenula-
ceae, Strigulaceae, Trypetheliaceae, Verrucariaceae,
Arthoniales, Lichinales, and Lecanoromycetes, mak-
ing up more than 99% of all lichen formers within
the Fungi.
Although lichenization events during the early
evolution of Fungi outside the Dikarya (Lumbsch &
Rikkinen 2016; Taylor et al. 1997, 2015) cannot be
ruled out, the available evidence indicates that extant
lichenized lineages are not derived from a common,
lichenized ancestor (Schoch et al. 2009). The fact
that all currently recognized lichenized lineages in
the Ascomycota are found within the Leotiomyceta,
and lichenization does not occur in any of the two
basal subphyla, Taphrinomycotina and Saccharomy-
cotina, nor in the basal classes of subphylum
Pezizomycotina (Pezizomycezes, Orbiliomycetes),
precludes any such speculations. The isolated
distribution of lichenized lineages in the Basidiomy-
cota and, at least in two cases (Agaricales, Canthar-
ellales), their undisputable descendance from closely
related, non-lichenized ancestors (Lawrey et al. 2009;
Dal Forno et al. 2013), make speculations on
common ancestry of lichenized fungi mute and
support the notion of multiple independent evolu-
tion of the lichenized lifestyle in the Fungi. Notably,
although this was first shown in the phylogenetic
study by Gargas et al. (1995a), the general idea of
lichen fungi being highly polyphyletic had already
been envisioned in studies more than one hundred
years ago (Bessey 1907; Clements 1897, 1909;
Clements & Shear 1931; Fink 1911, 1913; Lindau
1895).
Cyphobasidiales, currently including the sole
genus Cyphobasidium found in the upper cortex
(epicortex) mostly of Parmeliaceae, have not been
included in the above statistics. If the assessment by
Spribille et al. (2016) is correct and these fungi are
highly species-specific, considering the size of
Parmeliaceae, this order might contain hundreds, if
not thousands, of unrecognized species and could
grow to be the third-largest fungal order involved in
lichenization (after Lecanorales and Arthoniales),
although, as stated above, these fungi should not be
considered lichenized in the proper sense, but
hyperlichenized.
Lücking et al.: 2016 lichen classification 377
THE 2016 CLASSIFICATION OF LICHENIZED FUNGI IN THE
ASCOMYCOTA AND BASIDIOMYCOTA
In the following, the 995 fungal genera containing
lichenized species are listed hierarchically by subfam-
ily (where applicable), family, suborder (where
applicable), order, subclass (where applicable), class,
subphylum and phylum. All categories are listed in
strict alphabetical order. For each genus entry, the
following information is given: number of species, any
changes in classification compared to the latest
Outline of Ascomycota (Lumbsch & Huhndorf 2010),
at any hierarchical level, with references in case of
changes, and whether molecular data are available or
not. Nomenclatural novelties are listed separately after
this classification. References are cited if they refer,
directly or indirectly, to a cited change, even if a
particular genus is not necessarily treated in a cited
reference. For instance, all genera formerly placed in
Physciaceae (Teloschistales) are now included in
either Caliciaceae or Physciaceae (Caliciales), so for
all genera in the two families the references are cited
which reflect these changes.
Phylum ASCOMYCOTA Caval.-Sm.
Subphylum PEZIZOMYCOTINA O.E.Erikss. &
Winka
Class ARTHONIOMYCETES O.E.Erikss. & Winka
Order ARTHONIALES Henssen ex D.Hawksw. &
O.E.Erikss.
Family Andreiomycetaceae B.P.Hodk. & Lendemer
Andreiomyces B.P.Hodk. & Lendemer (2 spp.). New or
reinstated (Hodkinson & Lendemer 2013). Molecular: Yes.
Family Arthoniaceae Rchb. ex Rchb.
Amazonomyces Bat. (2 spp.). No change. Molecular: No.
Arthonia Ach. [nom.cons.] (syn.: Coccopeziza Har. & P.Karst.;
Naevia Fr.; Pseudo-Arthonia Marchand; Tomaselliella Cif.;
Xerodiscus Petr.) (500 spp.). New synonymy (Sundin et al.
2012). Molecular: Yes.
Arthothelium A.Massal. (10 spp.). No change. Molecular: Yes.
Coniangium Fr. (syn.: Diarthonis Clem.) (4 spp.). New or
reinstated (Sundin et al. 2012). Molecular: No.
Coniarthonia Grube (7 spp.). No change. Molecular: Yes.
Coniocarpon DC. (syn: Conioloma Flörke; Manilaea Syd. &
P.Syd.) (5 spp.). New or reinstated and new synonymy
(Frisch et al. 2014a ; Sundin et al. 2012). Molecular: Yes.
Crypthonia Frisch & G.Thor (15 spp.). New or reinstated
(Frisch & Thor 2010). Molecular: Yes.
378 The Bryologist 119(4): 2016
Cryptophaea VandenBroeck & Ertz (1 sp.). New or reinstated
(Van den Broeck & Ertz 2016). Molecular: Yes.
Cryptothecia Stirt. [nom.cons.] (65 spp.). No change. Molec-
ular: Yes.
Eremothecella Syd. & P.Syd. (7 spp.). No change. Molecular:
No.
Glomerulophoron Frisch, Ertz & G.Thor (2 spp.). New or
reinstated (Frisch et al. 2015). Molecular: Yes.
Herpothallon Tobler (46 spp.). No change. Molecular: Yes.
Inoderma (Ach.) Gray (3 spp.). New or reinstated (Frisch et al.
2015). Molecular: Yes.
Leprantha Dufour ex Körb. (1 sp.). New or reinstated (Sundin
et al. 2012). Molecular: No.
Myriostigma Kremp. (7 spp.). New or reinstated (Frisch et al.
2014a; Jagadeesh Ram & Sinha 2016). Molecular: Yes.
Pachnolepia A.Massal. (1 sp.). New or reinstated (Frisch et al.
2014a). Molecular: Yes.
Reichlingia Diederich & Scheid. (4 spp.). New or reinstated
(Frisch et al. 2014b). Molecular: Yes.
Sporodophoron Frisch, Y.Ohmura, Ertz & G.Thor (4 spp.). New
or reinstated (Frisch et al. 2015). Molecular: Yes.
Stirtonia A.L.Sm. (25 spp.). No change. Molecular: Yes.
Tylophoron Nyl. ex Stizenb. (syn.: Sporodochiolichen Aptroot &
Sipman) (8 spp.). New synonymy (Ertz et al. 2014c).
Molecular: Yes.
Family Chrysotrichaceae Zahlbr.
Chrysothrix Mont. [nom.cons.] [syn.: Allarthonia (Nyl.)
Zahlbr.; Plearthonis Clem.] (17 spp.). New synonymy
(Sundin et al. 2012). Molecular: Yes.
Melarthonis Frisch & G.Thor (1 sp.). New or reinstated (Frisch
et al. 2014a). Molecular: Yes.
Family Lecanographaceae Ertz, Tehler, G.Thor &
Frisch
Alyxoria Ach. (12 spp.). New or reinstated (Ertz & Tehler 2011;
Diederich et al. 2012). Molecular: Yes.
Heterocyphelium Vain. (1 sp.). Changed classification (this
paper). Molecular: Yes. Notes. This genus has recently been
sequenced and a paper on its revised classification is
forthcoming (van den Broeck et al., in prep.).
Lecanographa Egea & Torrente (38 spp.). Changed classification
(Ertz & Tehler 2011). Molecular: Yes.
Plectocarpon Fée (1 sp.). Changed classification (Ertz & Tehler
2011). Molecular: Yes.
Zwackhia Körb. (6 spp.). New or reinstated (Diederich et al.
2012; Ertz & Tehler 2011). Molecular: Yes.
Family Opegraphaceae Körb. ex Stizenb.
Combea De Not. (2 spp.). Changed classification (Ertz & Tehler
2011). Molecular: Yes.
Cresponea Egea & Torrente (21 spp.). Changed classification
(Ertz & Tehler 2011). Molecular: Yes.
Dictyographa Müll.Arg. (2 spp.). New or reinstated (Ertz &
Tehler 2011). Molecular: Yes.
Dolichocarpus R.Sant. (2 spp.). Changed classification (Ertz &
Tehler 2011). Molecular: Yes.
Fouragea Trevis. (3 spp.). New or reinstated (Frisch et al.
2014a). Molecular: Yes.
Ingaderia Darb. (3 spp.). Changed classification (Ertz & Tehler
2011). Molecular: Yes.
Llimonaea Egea & Torrente (4 spp.). Changed classification
(Ertz & Tehler 2011). Molecular: Yes.
Nyungwea Sérus., Eb.Fisch. & Killmann (1 sp.). Changed
classification (Frisch et al. 2014a). Molecular: Yes.
Opegrapha Ach. [nom.cons.] (syn.: Diplogramma Müll.Arg.)
(300 spp.). Changed classification and new synonymy
(Archer 2004; Ertz & Tehler 2011). Molecular: Yes.
Paraingaderia Ertz & Tehler (1 sp.). New or reinstated (Ertz &
Tehler 2011). Molecular: Yes.
Paraschismatomma Ertz & Tehler (1 sp.). New or reinstated
(Ertz & Tehler 2011). Molecular: Yes.
Pentagenella Darb. (syn.: Camanchaca Follmann & Peine;
Darbishirella Zahlbr.) (5 spp.). Changed classification and
new synonymy (Ertz & Tehler 2011). Molecular: Yes.
Schizopelte Th.Fr. (syn.: Hubbsia W.A.Weber) (4 spp.). Changed
classification and new synonymy (Ertz & Tehler 2011).
Molecular: Yes.
Sclerophyton Eschw. (14 spp.). Changed classification (Ertz &
Tehler 2011). Molecular: Yes.
Sparria Ertz & Tehler (2 spp.). New or reinstated (Ertz & Tehler
2011). Molecular: Yes.
Family Roccellaceae Chevall.
Ancistrosporella G.Thor. (3 spp.). No change. Molecular: No.
Austrographa Sparrius, Elix & A.W.Archer (3 spp.). New or
reinstated (Sparrius et al. 2010, 2013). Molecular: No.
Austroroccella Tehler & Ertz (1 sp.). New or reinstated (Tehler
et al. 2013). Molecular: Yes.
Chiodecton Ach. (20 spp.). No change. Molecular: Yes.
Crocellina Tehler & Ertz (1 sp.). New or reinstated (Ertz et al.
2015ab). Molecular: Yes.
Dendrographa Darb. (6 spp.). No change. Molecular: Yes.
Dichosporidium Pat. (8 spp.). No change. Molecular: Yes.
Dirina Fr. (13 spp.). No change. Molecular: Yes.
Dirinastrum Müll.Arg. (3 spp.). No change. Molecular: No.
Diromma Ertz & Tehler (1 sp.). New or reinstated (Ertz et al.
2015ab). Molecular: Yes.
Enterodictyon Müll.Arg. (2 spp.). No change. Molecular: No.
Enterographa Fée (53 spp.). No change. Molecular: Yes.
Erythrodecton G.Thor (3 spp.). No change. Molecular: Yes.
Feigeana Mies, Lumbsch & Tehler (1 sp.). No change.
Molecular: No.
Follmanniella Peine & B.Werner (1 sp.). No change. Molecular:
No.
Gorgadesia Tav. (1 sp.). No change. Molecular: Yes.
Graphidastra (Redinger) G.Thor (4 spp.). No change. Molec-
ular: No.
Gyrographa Ertz & Tehler (3 spp.). New or reinstated (Ertz et al.
2015ab). Molecular: Yes.

Gyronactis Ertz & Tehler (2 spp.). New or reinstated (Ertz et al.
2015ab). Molecular: Yes.
Halographis Kohlm. & Volkm.Kohlm. (1 sp.). No change.
Molecular: No.
Haplodina Zahlbr. (3 spp.). No change. Molecular: No.
Isalonactis Ertz, Tehler, Eb.Fisch., Killmann, Razafindr. & Sérus.
(1 sp.). New or reinstated (Ertz et al. 2014b). Molecular: Yes.
Lecanactis Körb. [nom.cons.] (syn.: Sagenidium Stirt.) (25 spp.).
New synonymy (Ertz et al. 2015ab). Molecular: Yes.
Mazosia A.Massal. (syn.: Gymnographoidea Fink) (27 spp.). New
synonymy (Aptroot et al. 2014; Sparrius 2004). Molecular: Yes.
Neosergipea M.Cáceres, Ertz & Aptroot (syn.: Sergipea M.Cá-
ceres, Ertz & Aptroot [nom.illeg.]) (1 sp.). New or reinstated
(Aptroot et al. 2013; Lücking et al. 2016a). Molecular: Yes.
Ocellomma Ertz & Tehler (1 sp.). New or reinstated (Ertz et al.
2015ab). Molecular: Yes.
Pseudolecanactis Zahlbr. (1 sp.). No change. Molecular: No.
Pseudoschismatomma Ertz & Tehler (1 sp.). New or reinstated
(Ertz et al. 2015ab). Molecular: Yes.
Psoronactis Ertz & Tehler (1 sp.). New or reinstated (Ertz et al.
2015a). Molecular: Yes.
Pulvinodecton Henssen & G.Thor (2 spp.). Changed classifica-
tion (Jaklitsch et al. 2016a). Molecular: No.
Roccella DC. [nom.cons.] (40 spp.). No change. Molecular: Yes.
Roccellina Darb. (29 spp.). No change. Molecular: Yes.
Sagenidiopsis R.W.Rogers & Hafellner (4 spp.). Changed
classification (Myllys 1989). Molecular: Yes.
Schismatomma Flot. & Körb. ex A.Massal. (10 spp.). No change.
Molecular: Yes.
Sigridea Tehler (6 spp.). No change. Molecular: Yes.
Simonyella J.Steiner (1 sp.). No change. Molecular: Yes.
Sipmania Egea & Torrente (1 sp.). Changed classification
(Jaklitsch et al. 2016a). Molecular: No.
Streimannia G.Thor (1 sp.). No change. Molecular: No.
Syncesia Taylor (21 spp.). No change. Molecular: Yes.
Tania Egea, Torrente & Sipman (2 spp.). Changed classification
(Jaklitsch et al. 2016a). Molecular: No.
Vigneronia Ertz (3 spp.). New or reinstated (Ertz et al. 2015a).
Molecular: Yes.
Family Roccellographaceae Ertz & Tehler
Dimidiographa Ertz & Tehler (3 spp.). New or reinstated (Ertz
& Tehler 2011). Molecular: Yes.
Fulvophyton Ertz & Tehler (syn.: Platygraphopsis Müll.Arg.) (6
spp.). New or reinstated and new synonymy (Ertz & Tehler
2011; this paper). Molecular: Yes. Notes. Platygraphopsis
appears to be an older name for Fulvophyton.
Roccellographa J.Steiner (syn.: Peterjamesia D.Hawksw.) (4
spp.). Changed classification and new synonymy (Ertz &
Tehler 2011). Molecular: Yes.
Order Arthoniales genera incertae sedis
Angiactis Aptroot & Sparrius (3 spp.). Changed classification
(this paper). Molecular: No. Notes. Classification changed
due to lack of molecular data.
Lücking et al.: 2016 lichen classification 379
Bactrospora A.Massal. (35 spp.). Changed classification (this
paper). Molecular: No. Notes. Classification changed due to
lack of molecular data.
Bryostigma Poelt & Döbbeler (2 spp.). New or reinstated (Poelt
& Doebbeler 1979; Sundin et al. 2012; Frisch et al. 2014a).
Molecular: Yes. Notes. The genus was placed in synonymy
with Trachylia Fr. by Sundin et al. 2012 but later reinstated
(Frisch et al. 2014a).
Catarraphia A.Massal. (1 sp.). No change. Molecular: No.
Felipes Frisch & G.Thor (syn.: Catapedia Frisch & G.Thor in
Jaklitsch et al. 2016a [nom.inval.]) (1 sp.). New or
reinstated and new synonymy (Frisch et al. 2014a; Jaklitsch
et al. 2016a; this paper). Molecular: Yes.
GlyphopsisAptroot (1 sp.). New or reinstated (Aptroot 2014).
Molecular: No.
GossypiothallonAptroot (1 sp.). New or reinstated (Aptroot
2014). Molecular: No.
Helminthocarpon Fée (1 sp.). New or reinstated (Staiger 2002).
Molecular: Yes.
Hormosphaeria Lév. (1 sp.). No change. Molecular: No.
Minksia Müll.Arg. (2 spp.). Changed classification (this paper).
Molecular: No. Notes. Classification changed due to lack of
molecular data.
Phoebus R.C.Harris & Ladd (1 sp.). Changed classification (this
paper). Molecular: No. Notes. Classification changed due to
lack of molecular data.
Sporostigma Grube (1 sp.). Changed classification (this paper).
Molecular: No. Notes. Classification changed due to lack of
molecular data.
Synarthonia Müll.Arg. (3 spp.). No change. Molecular: No.
Synarthothelium Sparrius (2 spp.). Changed classification (this
paper). Molecular: No. Notes. Classification changed due to
lack of molecular data.
Tarbertia Dennis (1 sp.). No change. Molecular: No.
Tylophorella Vain. (1 sp.). No change. Molecular: No.
Class CONIOCYBOMYCETES M. Prieto & Wedin
Order CONIOCYBALES M.Prieto & Wedin
Family Coniocybaceae Rchb.
Chaenotheca (Th.Fr.) Th.Fr. (25 spp.). Changed classification
(Prieto et al. 2013). Molecular: Yes.
Sclerophora Chevall. (6 spp.). Changed classification (Prieto et
al. 2013). Molecular: Yes.
Class DOTHIDEOMYCETES O.E.Erikss. & Winka
Subclass DOTHIDEOMYCETIDAE P.M.Kirk,
P.F.Cannon, J.C.David & Stalpers ex C.L.Schoch,
Spatafora, Crous & Shoemaker
Order CAPNODIALES Woron.
Family Antennulariellaceae Woron.
Antennulariella Woron. (1 sp.). New or reinstated (Coppins &
Aptroot 2008). Molecular: No.
380 The Bryologist 119(4): 2016
Family Cystocoleaceae, Locq. ex Lücking, B.P.Hodk.
& S.D.Leav.
Cystocoleus A.Massal. (1 sp.). Changed classification (Muggia et
al. 2008; Nelsen et al. 2009; this paper). Molecular: Yes.
Family Mycosphaerellaceae Lindau [nom.cons.]
Stigmidium Trevis. (1 sp.). No change. Molecular: No.
Family Racodiaceae Link
Racodium Fr. [nom.cons.] (1 sp.). Changed classification
(Muggia et al. 2008; Nelsen et al. 2009). Molecular: Yes.
Order Capnodiales genera incertae sedis
Racoleus R.Sant. & D.Hawksw. (1 sp.). New or reinstated
(Hawksworth et al. 2011b). Molecular: No.
Subclass PLEOSPOROMYCETIAE C.L.Schoch, Spa-
tafora, Crous & Shoemaker
Order PLEOSPORALES Luttrell ex M.E.Barr
Family Arthopyreniaceae Walt.Watson
Arthopyrenia A.Massal. [nom.cons.] (syn.: Arthopyreniomyces
Cif. & Tomas.) (5 spp.). Changed classification and new
synonymy (Hyde et al. 2013; Lücking & Hawksworth 2007).
Molecular: No.
Family Dacampiaceae Körb.
Dacampia A.Massal. (1 sp.). Changed classification (Ertz et al.
2015b). Molecular: Yes.
Family Mycoporaceae Zahlbr.
Mycoporum Flot. ex Nyl. [nom.cons.] (25 spp.). Changed
classification (Jaklitsch et al. 2016a). Molecular: No.
Class Dothideomycetes orders incertae sedis
Order COLLEMOPSIDIALES Pérez-Ortega, Garrido-Be-
navent & Grube
Family Xanthopyreniaceae Zahlbr.
Collemopsidium Nyl. (26 spp.). Changed classification (Pérez-
Ortega et al. 2016). Molecular: Yes.
Order EREMITHALLALES Lücking & Lumbsch
Family Melaspileaceae Walt.Watson (syn.: Eremi-
thallaceae Lücking & Lumbsch)
Encephalographa A.Massal. (1 sp.). Changed classification (Ertz
& Diederich 2015). Molecular: Yes.
Eremithallus Lücking, Lumbsch & Umaña (2 spp.). New or
reinstated (Lücking et al. 2008; Ertz & Diederich 2015).
Molecular: Yes. Notes. Ertz & Diederich (2015) proposed to
synonymize Eremithallus with Melaspilea, but the two
genera are kept separate here due to the morpho-
anatomical differences based on the type of M. arthonioides
(Fée) Nyl.
Melaspilea Nyl. (1 sp.). Changed classification (Ertz & Diederich
2015). Molecular: Yes.
Order MONOBLASTIALES Lücking, M.P.Nelsen &
K.D.Hyde
Family Monoblastiaceae Walt.Watson
Acrocordia A.Massal. (10 spp.). Changed classification (Nelsen
et al. 2009, 2011). Molecular: Yes.
Anisomeridium (Müll.Arg.) M.Choisy [nom.cons.] (200 spp.).
Changed classification (Nelsen et al. 2009, 2011). Molec-
ular: Yes.
Caprettia Bat. & H.Maia (8 spp.). Changed classification
(Nelsen et al. 2009, 2011). Molecular: No.
Megalotremis Aptroot (12 spp.). New or reinstated (Aptroot
1991; Nelsen et al. 2009, 2011). Molecular: Yes.
Monoblastia Riddle (11 spp.). Changed classification (Nelsen et
al. 2009, 2011). Molecular: No.
Trypetheliopsis Asahina (6 spp.). Changed classification (Kashi-
wadani et al. 2009; Nelsen et al. 2009, 2011). Molecular: Yes.
Order STRIGULALES Lücking, M.P.Nelsen & K.D.Hyde
Family Strigulaceae Zahlbr. (syn.: Phyllobatheliaceae
Bitter & F.Schill.)
Flavobathelium Lücking, Aptroot & G.Thor (1 sp.). Changed
classification (Nelsen et al. 2009, 2011). Molecular: Yes.
Oletheriostrigula Huhndorf & R.C.Harris (1 sp.). Changed
classification (Jaklitsch et al. 2016a). Molecular: No.
Phyllobathelium (Müll.Arg.) Müll.Arg. (5 spp.). Changed
classification (Nelsen et al. 2009, 2011). Molecular: Yes.
Phyllocratera Sérus. & Aptroot (2 spp.). Changed classification
(Nelsen et al. 2009, 2011). Molecular: No.
Strigula Fr. (70 spp.). Changed classification (Nelsen et al. 2009,
2011). Molecular: Yes.
Order TRYPETHELIALES Lücking, Aptroot & Sipman
Family Trypetheliaceae Zenker
Aptrootia Lücking & Sipman (3 spp.). No change. Molecular:
Yes.
Architrypethelium Aptroot (7 spp.). No change. Molecular: Yes.
Astrothelium Eschw. (syn.: Campylothelium Müll.Arg., Crypto-
thelium A.Massal., Laurera Rchb.; Tremothylium Nyl.) (242
spp.). New synonymy (Aptroot & Lücking 2016; Lücking et
al. 2016c ; Nelsen et al. 2014a). Molecular: Yes.
Bathelium Ach. (16 spp.). New or reinstated (Aptroot &
Lücking 2016; Lücking et al. 2016c ; Nelsen et al. 2014a).
Molecular: Yes.

Bogoriella Zahlbr. (29 spp.). New or reinstated (Aptroot &
Lücking 2016; Lücking et al. 2016c ; Nelsen et al. 2014a).
Molecular: Yes.
Constrictolumina Lücking, M.P.Nelsen & Aptroot (9 spp.). New
or reinstated (Aptroot & Lücking 2016; Lücking et al.
2016c; Nelsen et al. 2014a). Molecular: Yes.
Dictyomeridium Aptroot, M.P.Nelsen & Lücking (7 spp.). New
or reinstated (Aptroot & Lücking 2016; Lücking et al.
2016c; Nelsen et al. 2014a). Molecular: Yes.
Distothelia Aptroot (3 spp.). New or reinstated (Aptroot &
Lücking 2016). Molecular: No.
Marcelaria Aptroot, M.P.Nelsen & Parnmen (3 spp.). New
or reinstated (Aptroot & Lücking 2016; Lücking et al.
2016c ; Nelsen et al. 2013; Nelsen et al. 2014a).
Molecular: Yes.
Nigrovothelium Lücking, M.P.Nelsen & Aptroot (2 spp.). New
or reinstated (Aptroot & Lücking 2016; Lücking et al.
2016c; Nelsen et al. 2014a). Molecular: Yes.
Novomicrothelia Aptroot, M.P.Nelsen & L ücking (1 sp.).
New or reinstated (Aptroot & L ücking 2016; L ücking
et al. 2016c; Nelsen et al. 2014a). Molecular: Yes.
Polymeridium (M üll.Arg.) R.C.Harris (syn.: Exiliseptum
R.C.Harris) (50 spp.). New synonymy (Aptroot & Lücking
2016). Molecular: Yes.
Polypyrenula D.Hawksw. (2 spp.). Changed classification (this
paper). Molecular: Yes. Notes. A forthcoming phylogenetic
analysis places this genus in Trypetheliaceae (Miranda-
Gonzáles et al., in prep.).
Pseudopyrenula Müll.Arg. (20 spp.). No change. Molecular: Yes.
Trypethelium Spreng. [nom.cons.] (16 spp.). No change.
Molecular: Yes.
Viridothelium Lücking, M.P.Nelsen & Aptroot (10 spp.). New
or reinstated (Aptroot & Lücking 2016; Lücking et al.
2016c). Molecular: Yes.
Class EUROTIOMYCETES O.E.Erikss. & Winka
Subclass CHAETOTHYRIOMYCETIDAE Dowell
Order CHAETOTHYRIALES M.E.Barr
Family Lyrommataceae Lücking
Lyromma Bat. (8 spp.). Changed classification (Jaklitsch et al.
2016a). Molecular: No.
Family Microtheliopsidaceae O.E.Erikss.
Microtheliopsis Müll.Arg. (4 spp.). Changed classification
(Jaklitsch et al. 2016a). Molecular: No.
Family Pyrenotrichaceae Zahlbr.
Pyrenothrix Riddle (2 spp.). Changed classification (Jaklitsch et
al. 2016a). Molecular: Yes.
Order PHAEOMONIELLALES K.-H.Chen, A.E.Arnold,
Gueidan & Lutzoni
Lücking et al.: 2016 lichen classification 381
Family Celotheliaceae Lücking, Aptroot & Sipman
Celothelium A.Massal. (8 spp.). Changed classification (Chen et
al. 2015; Gueidan et al. 2014b). Molecular: Yes.
Order PYRENULALES Fink ex D.Hawksw. & O.E.Erikss.
Family Pyrenulaceae Rabenh.
Anthracothecium Hampe ex A.Massal. (5 spp.). No change.
Molecular: Yes.
Clypeopyrenis Aptroot (2 spp.). No change. Molecular: No.
Distopyrenis Aptroot (8 spp.). No change. Molecular: No.
Lithothelium Müll.Arg. (28 spp.). No change. Molecular: Yes.
Mazaediothecium Aptroot (3 spp.). No change. Molecular: Yes.
Pyrenowilmsia R.C.Harris & Aptroot (1 sp.). No change.
Molecular: No.
Pyrenula Ach. [nom.cons.] (syn.: Heufleridium Müll.Arg.,
Micromma Zahlbr., Porothelium Eschw., Stromatothelium
Trevis.) (170 spp.). New synonymy (Aptroot 2012).
Molecular: Yes.
Pyrgillus Nyl. (4 spp.). No change. Molecular: Yes.
Sulcopyrenula H.Harada (4 spp.). No change. Molecular: No.
Granulopyrenis Aptroot (6 spp.). No change. Molecular: Yes.
Blastodesmia A.Massal. (1 sp.). No change. Molecular: No.
Order Pyrenulales genera incertae sedis
Rhaphidicyrtis Vain. (1 sp.). No change. Molecular: No.
Xenus Kohlm. & Volkm.Kohlm. (1 sp.). No change. Molecular:
No.
Order VERRUCARIALES Mattick ex D.Hawksw. &
O.E.Erikss.
Family Verrucariaceae Zenker
Agonimia Zahlbr. (17 spp.). No change. Molecular: Yes.
Anthracocarpon Breuss (3 spp.). No change. Molecular: Yes.
Atla Savić & Tibell (9 spp.). No change. Molecular: Yes.
Awasthiella Kr.P.Singh (1 sp.). No change. Molecular: No.
Bagliettoa A.Massal. (20 spp.). No change. Molecular: Yes.
Catapyrenium Flot. (6 spp.). No change. Molecular: Yes.
Clauzadella Nav.Ros. & Roux (1 sp.). No change. Molecular:
No.
Clavascidium Breuss (9 spp.). No change. Molecular: Yes.
Dermatocarpon Eschw. (20 spp.). No change. Molecular: Yes.
Endocarpon Hedw. (70 spp.). No change. Molecular: Yes.
Flakea O.E.Erikss. (1 sp.). No change. Molecular: Yes.
Glomerilla Norman (1 sp.). No change. Molecular: No.
Henrica B.de Lesd. (4 spp.). No change. Molecular: Yes.
Heterocarpon Müll.Arg. (1 sp.). No change. Molecular: No.
Heteroplacidium Breuss (12 spp.). No change. Molecular:
Yes.
382 The Bryologist 119(4): 2016
Hydropunctaria C.Keller, Gueidan & Thüs (8 spp.). No change.
Molecular: Yes.
Involucropyrenium Breuss (9 spp.). No change. Molecular:
Yes.
Mastodia Hook.f. & Harv. (5 spp.). Changed classification
(Pérez-Ortega et al. 2010). Molecular: Yes.
Moriola Norman (15 spp.). Changed classification (Grube
2005). Molecular: No.
Neocatapyrenium H.Harada (5 spp.). No change. Molecular:
Yes.
Normandina Nyl. (3 spp.). Changed classification (Frisch &
Ohmura 2015; Muggia et al. 2010). Molecular: Yes.
Parabagliettoa Gueidan & Cl.Roux (3 spp.). No change.
Molecular: Yes.
Phylloblastia Vain. (12 spp.). No change. Molecular: No.
Placidiopsis Beltr. (20 spp.). No change. Molecular: Yes.
Placidium A.Massal. (syn: Dermatocarpella H.Harada) (28 spp.).
New synonymy (Breuss 1996) Molecular: Yes.
Placocarpus Trevis. (5 spp.). No change. Molecular: Yes.
Placopyrenium Breuss (22 spp.). No change. Molecular: Yes.
Placothelium Müll.Arg. (1 sp.). No change. Molecular: No.
Polyblastia A.Massal. [nom.cons.] (120 spp.). No change.
Molecular: Yes.
Psoroglaena Müll.Arg. (syn.: Leucocarpia Vězda; Macentina
Vězda) (17 spp.). New synonymy (Lücking 2008). Molec-
ular: Yes.
Rhabdopsora Müll.Arg. (2 spp.). No change. Molecular: No.
Scleropyrenium H.Harada (2 spp.). No change. Molecular: No.
Servitia M.S.Christ. & Alstrup (1 sp.). No change. Molecular:
No.
Spheconisca (Norman) Norman (20 spp.). No change. Molec-
ular: No.
Sporodictyon A.Massal. (5 spp.). No change. Molecular: Yes.
Staurothele Norman [nom.cons.] (40 spp.). No change.
Molecular: Yes.
Thelidiopsis Vain. (4 spp.). No change. Molecular: No.
Thelidium A.Massal. (100 spp.). No change. Molecular: Yes.
Trimmatothele Norman ex Zahlbr. (3 spp.). No change.
Molecular: Yes.
Verrucaria Schrad. [nom.cons.] (300 spp.). No change.
Molecular: Yes.
Verruculopsis Gueidan, Nav.Ros. & Cl.Roux (4 spp.). No
change. Molecular: Yes.
Wahlenbergiella Gueidan & Thüs (3 spp.). No change.
Molecular: Yes.
Willeya Müll.Arg. (11 spp.). New or reinstated (Gueidan et al.
2014a; Gueidan & Lendemer 2015). Molecular: Yes.
Order Verrucariales genera incertae sedis
Kalbiana Henssen (1 sp.). No change. Molecular: No.
Botryolepraria Canals, Hern.-Mar., Gómez-Bolea & Llimona (2
spp.). Changed classification (Kukwa & Pérez-Ortega
2010). Molecular: Yes.
Class Eurotiomycetes families incertae sedis
Family Dactylosporaceae Bellem. & Hafellner
Sclerococcum Fr. (1 sp.). New or reinstated (Etayo 1995; Smith
2009; Diederich et al. 2013). Molecular: Yes.
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. (200 spp.). No change. Molecular: Yes.
Caeruleum Arcadia (2 spp.). New or reinstated (Arcadia &
Knudsen 2012). Molecular: No.
Eiglera Hafellner (2 spp.). Changed classification (Mia˛dlikowska
et al. 2014). Molecular: Yes.
Glypholecia Nyl. (1 sp.). No change. Molecular: Yes.
Lithoglypha Brusse (1 sp.). No change. Molecular: No.
Myriospora Nägeli ex Uloth (syn.: Silobia M.Westb. & Wedin)
(9 spp.). New or reinstated and new synonymy (Arcadia &
Knudsen 2012; Gueidan et al. 2014c; Roux et al. 2016;
Westberg et al. 2011, 2015). Molecular: Yes.
Pleopsidium Körb. (3 spp.). No change. Molecular: Yes.
Polysporina Vězda [nom.cons.] (10 spp.). No change. Molec-
ular: Yes.
Sarcogyne Flot. [nom.cons.] (28 spp.). No change. Molecular:
Yes.
Thelocarpella Nav.Ros. & Cl.Roux (1 sp.). New synonymy
(Gueidan et al. 2014c, Knudsen & Lendemer 2016; Roux et
al. 2016). Molecular: Yes.
Trimmatothelopsis Zschacke (1 sp.). Changed classification
(Gueidan et al. 2014c, Knudsen & Lendemer 2016; Roux et
al. 2016). Molecular: Yes.
Timdalia Hafellner (1 sp.). Changed classification (Westberg et
al. 2015). Molecular: Yes.
Subclass CANDELARIOMYCETIDAE Mia˛dl. et al.
ex Timdal & M.Westb.
Order CANDELARIALES Mia˛dl., Lutzoni & Lumbsch
Family Candelariaceae Hakul.
Candelaria A.Massal. (7 spp.). Changed classification (Bend-
iksby & Timdal 2013). Molecular: Yes.
Candelariella Müll.Arg. (50 spp.). Changed classification
(Bendiksby & Timdal 2013). Molecular: Yes.
Candelina Poelt (3 spp.). Changed classification (Bendiksby &
Timdal 2013). Molecular: Yes.

Placomaronea Räsänen (6 spp.). Changed classification (Bend-
iksby & Timdal 2013). Molecular: Yes.
Family Pycnoraceae Bendiksby & Timdal
Pycnora Hafellner (3 spp.). Changed classification (Bendiksby &
Timdal 2013). Molecular: Yes.
Subclass LECANOROMYCETIDAE P.M.Kirk,
P.F.Cannon, J.C.David & Stalpers ex Mia˛dl.,
Lutzoni & Lumbsch
Order CALICIALES Bessey
Family Caliciaceae Chevall.
Acolium (Ach.) Grev. (5 spp.). Changed classification (Gaya et
al. 2012; Prieto et al. 2013; Prieto & Wedin 2016).
Molecular: Yes.
Acroscyphus Lév. (1 sp.). Changed classification (Gaya et al.
2012; Prieto et al. 2013; Prieto & Wedin 2016). Molecular:
Yes.
Allocalicium M.Prieto & Wedin (1 sp.). New or reinstated
(Prieto & Wedin 2016). Molecular: Yes.
Amandinea M.Choisy ex Scheid. & M.Mayrhofer (35 spp.).
Changed classification (Gaya et al. 2012; Prieto et al. 2013;
Prieto & Wedin 2016). Molecular: Yes.
Australiaena Matzer, H.Mayrhofer & Elix (1 sp.). Changed
classification (Gaya et al. 2012; Prieto et al. 2013; Prieto &
Wedin 2016). Molecular: Yes.
Baculifera Marbach (14 spp.). New or reinstated (Marbach
2000). Molecular: Yes.
Buellia De Not. [nom.cons.] (syn.: Hafellia Kalb, H.Mayrhofer
& Scheid.) (300 spp.). Changed classification and new
synonymy (Bungartz et al. 2007; Gaya et al. 2012; Prieto &
Wedin 2016). Molecular: Yes. Notes. The genera Buellia,
typified by B. disciformis (Fr.) Mudd (conserved type) and
Hafellia, typified by H. leptoclinoides (Nyl.) Scheid. & H.
Mayrhofer, are congeneric, since the type species are closely
related (Bungartz et al. 2007; Marbach 2000). The
conservation of Buellia with B. disciformis as type was
proposed by Aptroot (1987) and accepted, although based
on an erroneous lectotypification, since the name B.
disciformis was not part of the protologue of Buellia. For
unknown reasons, an attempt to correct this error and
make the name Buellia available for a widespread and
common group of saxicolous species centered around B.
aethalea (Ach.) Th. Fr., thus maintaining Hafellia for the
corticolous species related to B. disciformis (Mobert et al.
1999) was not accepted (Bungartz et al. 2007). The problem
continues to linger around in the literature, with both
Buellia and Hafellia continuing to be used.
Calicium Pers. (30 spp.). Changed classification (Gaya et al.
2012; Prieto et al. 2013; Prieto & Wedin 2016). Molecular:
Yes.
Chrismofulvea Marbach (4 spp.). New or reinstated (Marbach
2000). Molecular: Yes.
Ciposia Marbach (1 sp.). New or reinstated (Marbach 2000).
Molecular: No.
Lücking et al.: 2016 lichen classification 383
Cratiria Marbach (20 spp.). Changed classification (Gaya et al.
2012; Prieto et al. 2013; Prieto & Wedin 2016). Molecular:
Yes.
Cyphelium Ach. (7 spp.). Changed classification (Gaya et al.
2012; Prieto et al. 2013; Prieto & Wedin 2016). Molecular:
Yes.
Dermatiscum Nyl. (3 spp.). Changed classification (Gaya et al.
2012; Prieto et al. 2013; Prieto & Wedin 2016). Molecular:
Yes.
Dermiscellum Hafellner, H.Mayrhofer & Poelt (1 sp.). Changed
classification (Gaya et al. 2012; Prieto et al. 2013; Prieto &
Wedin 2016). Molecular: No.
Dimelaena Norman (8 spp.). Changed classification (Gaya et al.
2012; Prieto et al. 2013; Prieto & Wedin 2016). Molecular:
Yes.
Diploicia A.Massal. (4 spp.). Changed classification (Gaya et al.
2012; Prieto et al. 2013; Prieto & Wedin 2016). Molecular:
Yes.
Diplotomma Flot. (syn.: Mattickiolichen Tomas. & Cif.) (30
spp.). Changed classification and new synonymy (Gaya et
al. 2012; Lamb 1968; Prieto & Wedin 2016). Molecular: Yes.
Dirinaria (Tuck.) Clem. (35 spp.). Changed classification (Gaya
et al. 2012; Prieto & Wedin 2016). Molecular: Yes.
Endohyalina Marbach (10 spp.). New or reinstated (Marbach
2000). Molecular: No.
Fluctua Marbach (1 sp.). New or reinstated (Marbach 2000).
Molecular: No.
Gassicurtia Feé (27 spp.). Changed classification (Gaya et al.
2012; Prieto et al. 2013; Prieto & Wedin 2016). Molecular:
Yes.
Hypoflavia Marbach (3 spp.). Changed classification (Gaya et al.
2012; Prieto et al. 2013; Prieto & Wedin 2016). Molecular:
No.
Monerolechia Trevis. (5 spp.). Changed classification (Gaya et
al. 2012; Prieto et al. 2013; Prieto & Wedin 2016).
Molecular: Yes.
Orcularia (Malme) Kalb & Giralt (4 spp.). New or reinstated
(Kalb & Giralt 2011). Molecular: No.
Pseudothelomma M.Prieto & Wedin (2 spp.). New or reinstated
(Prieto & Wedin 2016). Molecular: Yes.
Pyxine Fr. (75 spp.). Changed classification (Gaya et al. 2012;
Prieto et al. 2013; Prieto & Wedin 2016). Molecular: Yes.
Redonia C.W.Dodge (2 spp.). Changed classification (Gaya et al.
2012; Prieto et al. 2013; Prieto & Wedin 2016). Molecular:
No.
Santessonia Hale & Vobis (7 spp.). Changed classification (Gaya
et al. 2012; Prieto et al. 2013; Prieto & Wedin 2016).
Molecular: Yes.
Sculptolumina Marbach (3 spp.). Changed classification (Gaya
et al. 2012; Prieto et al. 2013; Prieto & Wedin 2016).
Molecular: No.
Sphinctrinopsis Woron. (1 sp.). Changed classification (Gaya et
al. 2012; Prieto et al. 2013; Prieto & Wedin 2016).
Molecular: No.
Stigmatochroma Marbach (9 spp.). Changed classification
(Gaya et al. 2012; Prieto et al. 2013; Prieto & Wedin
2016). Molecular: Yes.
384 The Bryologist 119(4): 2016
Tetramelas Norman (16 spp.). Changed classification (Gaya et
al. 2012; Prieto et al. 2013; Prieto & Wedin 2016).
Molecular: Yes.
Texosporium Nádv. ex Tibell & Hofsten (1 sp.). Changed
classification (Gaya et al. 2012; Prieto et al. 2013; Prieto &
Wedin 2016). Molecular: Yes.
Thelomma A.Massal. (7 spp.). Changed classification (Gaya et
al. 2012; Prieto et al. 2013; Prieto & Wedin 2016).
Molecular: Yes.
Tholurna Norman [nom.cons.] (1 sp.). Changed classification
(Gaya et al. 2012; Prieto et al. 2013; Prieto & Wedin 2016).
Molecular: Yes.
Tylophoropsis Sambo (1 sp.). Changed classification (Gaya et al.
2012; Prieto et al. 2013; Prieto & Wedin 2016). Molecular:
No.
Family Physciaceae Zahlbr. [nom.cons.]
Anaptychia Körb. (15 spp.). Changed classification (Gaya et al.
2012; Prieto et al. 2013; Prieto & Wedin 2016). Molecular:
Yes.
Coscinocladium Kunze (2 spp.). Changed classification (Gaya et
al. 2012; Prieto et al. 2013; Prieto & Wedin 2016).
Molecular: Yes. Notes. The genus was listed twice in
Lumbsch & Huhndorf (2010).
Culbersonia Essl. (2 spp.). Changed classification (Gaya et al.
2012; Prieto et al. 2013; Prieto & Wedin 2016). Molecular:
No.
Heterodermia Trevis. (90 spp.). Changed classification (Gaya et
al. 2012; Prieto et al. 2013; Prieto & Wedin 2016).
Molecular: Yes.
Hyperphyscia Müll.Arg. (9 spp.). Changed classification (Gaya
et al. 2012; Prieto et al. 2013; Prieto & Wedin 2016).
Molecular: Yes.
Kashiwadia S.Y.Kondr., L.L} okös & Hur (1 sp.). New or
reinstated (Kondratyuk et al. 2014c). Molecular: Yes.
Leucodermia Kalb (10 spp.). New or reinstated (Mongkolsuk et
al. 2015). Molecular: Yes.
Mischoblastia A.Massal. (3 spp.). Changed classification (Gaya
et al. 2012; Prieto et al. 2013; Prieto & Wedin 2016).
Molecular: Yes.
Mobergia H.Mayrhofer, Sheard & Matzer (2 spp.). Changed
classification (Gaya et al. 2012; Prieto et al. 2013; Prieto &
Wedin 2016). Molecular: Yes.
Oxnerella S.Y.Kondr., L}
okös & Hur (1 sp.). New or reinstated
(Kondratyuk et al. 2014d; Resl et al. 2016). Molecular: Yes.
Phaeophyscia Mob. (30 spp.). Changed classification (Gaya et al.
2012; Prieto et al. 2013; Prieto & Wedin 2016). Molecular:
Yes.
Phaeorrhiza H.Mayrhofer & Poelt (2 spp.). Changed classifica-
tion (Gaya et al. 2012; Prieto et al. 2013; Prieto & Wedin
2016). Molecular: Yes.
Physcia (Schreb.) Michaux (80 spp.). Changed classification
(Gaya et al. 2012; Prieto et al. 2013; Prieto & Wedin 2016).
Molecular: Yes.
Physciella Essl. (4 spp.). Changed classification (Gaya et al.
2012; Prieto et al. 2013; Prieto & Wedin 2016). Molecular:
Yes.
Physconia Poelt [nom.cons.] (25 spp.). Changed classification
(Gaya et al. 2012; Prieto et al. 2013; Prieto & Wedin 2016).
Molecular: Yes.
Polyblastidium Kalb (18 spp.). New or reinstated (Mongkolsuk
et al. 2015). Molecular: Yes.
Rinodina (Ach.) Gray (300 spp.). Changed classification (Gaya
et al. 2012; Prieto et al. 2013; Prieto & Wedin 2016).
Molecular: Yes.
Rinodinella H.Mayrhofer & Poelt (6 spp.). Changed classifica-
tion (Gaya et al. 2012; Prieto et al. 2013; Prieto & Wedin
2016). Molecular: Yes.
Tornabea Oesth. (1 sp.). Changed classification (Gaya et al. 2012;
Prieto et al. 2013; Prieto & Wedin 2016). Molecular: Yes.
Order LECANORALES Nannf.
Family Biatorellaceae M.Choisy ex Hafellner &
Casares-Porcel
Biatorella De Not. (30 spp.). No change. Molecular: No.
Family Carbonicolaceae Bendiksby & Timdal
Carbonicola Bendiksby & Timdal (3 spp.). New or reinstated
(Bendiksby & Timdal 2013). Molecular: Yes.
Family Catillariaceae Hafellner
Austrolecia Hertel (1 sp.). No change. Molecular: Yes.
Catillaria A.Massal. (150 spp.). No change. Molecular: Yes.
Placolecis Trevis. (1 sp.). No change. Molecular: No.
Solenopsora A.Massal. (20 spp.). No change. Molecular: Yes.
Xanthopsorella Kalb & Hafellner (2 spp.). No change.
Molecular: No.
Family Cladoniaceae Zenker
Calathaspis I.M.Lamb & W.A.Weber (1 sp.). No change.
Molecular: No.
Carassea S.Stenroos (1 sp.). No change. Molecular: Yes.
Cetradonia J.C.Wei & Ahti (1 sp.). No change. Molecular: Yes.
Cladia Nyl. (syn.: Heterodea Nyl.; Ramalinora Lumbsch,
Rambold & Elix) (25 spp.). New synonymy (Lumbsch et
al. 2010). Molecular: Yes.
Cladonia Hill ex P.Browne [nom.cons.] (500 spp.). No change.
Molecular: Yes.
Gymnoderma Nyl. (3 spp.). No change. Molecular: Yes.
Heteromyces Müll.Arg. (1 sp.). No change. Molecular: No.
Metus D.J.Galloway & P.James (3 spp.). No change. Molecular:
Yes.
Myelorrhiza Verdon & Elix (2 spp.). No change. Molecular: Yes.
Notocladonia S.Hammer (2 spp.). No change. Molecular: No.
Pilophorus Th.Fr. (11 spp.). No change. Molecular: Yes.
Pycnothelia Duf. (2 spp.). No change. Molecular: Yes.
Sphaerophoropsis Vain. (2 spp.). No change. Molecular: No.
Squamella S.Hammer (1 sp.). No change. Molecular: No.

Thysanothecium Mont. & Berk. (3 spp.). No change. Molecular:
Yes.
Family Gypsoplacaceae Timdal
Gypsoplaca Timdal (1 sp.). No change. Molecular: Yes.
Family Haematommataceae Hafellner
Haematomma A.Massal. [nom.cons.] (50 spp.). No change.
Molecular: Yes.
Family Lecanoraceae Körb.
Ameliella Fryday & Coppins (2 spp.). New or reinstated (Fryday
& Coppins 2008). Molecular: No.
Bryodina Hafellner (2 spp.). No change. Molecular: No.
Bryonora Poelt (11 spp.). No change. Molecular: Yes.
Cladidium Hafellner (2 spp.). No change. Molecular: No.
Claurouxia D.Hawksw. (1 sp.). No change. Molecular: No.
Clauzadeana Cl.Roux (1 sp.). No change. Molecular: No.
Edrudia W.P.Jordan (1 sp.). No change. Molecular: No.
Huea C.W.Dodge & G.E.Baker [syn.: Carbonea (Hertel) Hertel]
(20 spp.). Changed classification and new synonymy
(Fryday 2011). Molecular: Yes.
Japewiella Printzen (6 spp.). No change. Molecular: No.
Lecanora Ach. (550 spp.). No change. Molecular: Yes.
Lecidella Körb. (80 spp.). No change. Molecular: Yes.
Miriquidica Hertel & Rambold (25 spp.). No change. Molecular:
Yes.
Myriolecis Clements (syn.: Arctopeltis Poelt) (31 spp.). New or
reinstated and new synonymy (Śliwa et al. 2012; Zhao et al.
2016). Molecular: Yes.
Myrionora R.C.Harris (2 spp.). No change. Molecular: No.
Palicella Rodr.Flakus & Printzen (3 spp.). New or reinstated
(Rodriguez-Flakus & Printzen 2014). Molecular: Yes.
Protoparmeliopsis Choisy (20 spp.). New or reinstated (Zhao et
al. 2016). Molecular: Yes.
Psorinia Gotth.Schneid. (2 spp.). No change. Molecular: No.
Punctonora Aptroot (1 sp.). No change. Molecular: No.
Pyrrhospora Körb. (1 sp.). No change. Molecular: Yes.
Rhizoplaca Zopf (11 spp.). New or reinstated (Zhao et al. 2016).
Molecular: Yes.
Sagema Poelt & Grube (1 sp.). No change. Molecular: No.
Sedelnikovaea S.Y.Kondr., M.H.Jeong & Hur (1 sp.). New or
reinstated (Kondratyuk et al. 2014e). Molecular: Yes.
Traponora Aptroot (5 spp.). No change. Molecular: No.
Tylothallia P.James & R.Kilias (3 spp.). No change. Molecular:
Yes.
Vainionora Kalb (9 spp.). No change. Molecular: No.
Family Malmideaceae Kalb, RivasPlata & Lumbsch
Malmidea Kalb, Rivas Plata & Lumbsch (50 spp.). New or
reinstated (Kalb et al. 2011). Molecular: Yes.
Lücking et al.: 2016 lichen classification 385
Savoronala Ertz, Eb.Fisch., Killmann, Razafindr. & Sérus. (1
sp.). New or reinstated (Ertz et al. 2014c). Molecular: Yes.
Family Pachyascaceae Poelt ex P.M.Kirk, P.F.Can-
non & J.C.David
Pachyascus Poelt & Hertel (1 sp.). No change. Molecular: No.
Family Parmeliaceae Zenker
Ahtiana Goward (3 spp.). No change. Molecular: Yes.
Alectoria Ach. (7 spp.). No change. Molecular: Yes.
Allantoparmelia (Vain.) Essl. (3 spp.). No change. Molecular:
Yes.
Allocetraria Kurok. & M.Y.Lai (9 spp.). No change. Molecular:
Yes.
Anzia Stizenb. [nom.cons.] (34 spp.). No change. Molecular:
Yes.
Arctocetraria Kärnefelt & Thell (3 spp.). No change. Molecular:
Yes.
Arctoparmelia Hale (5 spp.). No change. Molecular: Yes.
Asahinea W.L.Culb. & C.F.Culb. (4 spp.). No change.
Molecular: Yes.
Austroparmelina A.Crespo, Divakar & Elix (15 spp.). New or
reinstated (Crespo et al. 2010b). Molecular: Yes.
Brodoa Goward (3 spp.). No change. Molecular: Yes.
Bryocaulon Kärnefelt (4 spp.). No change. Molecular: Yes.
Bryoria Brodo & D.Hawksw. (50 spp.). No change. Molecular:
Yes.
Bulbothrix Hale (syn.: Bulborrhizina Kurok.) (60 spp.). New
synonymy (Kirika et al. 2015) Molecular: Yes.
Canoparmelia Elix & Hale (40 spp.). No change. Molecular: Yes.
Cetraria Ach. [nom.cons.] (syn.: Coelocaulon Link) (15 spp.).
New synonymy (Thell et al. 2002, 2009). Molecular: Yes.
Cetrariella Kärnefelt & Thell (4 spp.). No change. Molecular:
Yes.
Cetrelia W.L.Culb. & C.F.Culb. (18 spp.). No change.
Molecular: Yes.
Cetreliopsis Kurok. (7 spp.). No change. Molecular: Yes.
Coelopogon Brusse & Kärnefelt (2 spp.). No change. Molecular:
Yes.
Cornicularia (Schreb.) Ach. (1 sp.). No change. Molecular: Yes.
Crespoa (D.Hawksw.) Lendemer & B.P.Hodk. (5 spp.). New or
reinstated (Lendemer & Hodkinson 2012b). Molecular: Yes.
Dactylina Nyl. (2 spp.). No change. Molecular: Yes.
Davidgallowaya Aptroot (1 sp.). No change. Molecular: No.
Emodomelanelia Divakar & A.Crespo (1 sp.). New or reinstated
(Crespo et al. 2010a). Molecular: Yes.
Esslingeriana Hale & M.J.Lai (1 sp.). No change. Molecular: Yes.
Evernia Ach. (10 spp.). No change. Molecular: Yes.
Everniopsis Nyl. (1 sp.). No change. Molecular: Yes.
Flavocetraria Kärnefelt & Thell (3 spp.). No change. Molecular:
Yes.
Flavoparmelia Hale (40 spp.). No change. Molecular: Yes.
Flavopunctelia Hale (7 spp.). No change. Molecular: Yes.
386 The Bryologist 119(4): 2016
Gowardia Halonen, Myllys, Velmala & Hyvärinen (2 spp.).
Reinstated genus (Myllys et al. 2014). Molecular: Yes.
Himantormia I.M.Lamb (2 spp.). No change. Molecular: Yes.
Hypogymnia (Nyl.) Nyl. (syn.: Cavernularia Degel.) (90 spp.).
New synonymy (Mia˛dlikowska et al. 2011). Molecular: Yes.
Hypotrachyna (Vain.) Hale (syn.: Cetrariastrum Sipman, Ever-
niastrum Hale ex Sipman, Parmelinopsis Elix & Hale) (260
spp.). New synonymy (Divakar et al. 2013b). Molecular:
Yes.
Imshaugia F.C.Mey. (7 spp.). No change. Molecular: Yes.
Kaernefeltia Thell & Goward (2 spp.). No change. Molecular:
Yes.
Letharia (Th.Fr.) Zahlbr. [nom.cons.] (6 spp.). No change.
Molecular: Yes.
Lethariella (Motyka) Krog (11 spp.). No change. Molecular:
Yes.
Masonhalea Kärnefelt (2 spp.). No change. Molecular: Yes.
Melanelia Essl. (6 spp.). No change. Molecular: Yes.
Melanelixia O.Blanco, A.Crespo, Divakar, Essl., D.Hawksw. &
Lumbsch (16 spp.). No change. Molecular: Yes.
Melanohalea O.Blanco, A.Crespo, Divakar, Essl., D.Hawksw. &
Lumbsch (24 spp.). New or reinstated (Blanco et al. 2004).
Molecular: Yes.
Menegazzia A.Massal. (70 spp.). No change. Molecular: Yes.
Montanelia Divakar, A.Crespo, Wedin & Essl. (5 spp.). New or
reinstated (Divakar et al. 2012). Molecular: Yes.
Myelochroa (Asahina) Elix & Hale (30 spp.). No change.
Molecular: Yes.
Nephromopsis Müll.Arg. (syn.: Cetrariopsis Kurok.; Flavocetrar-
iella D.D.Awasthi) (24 spp.). New synonymy (Nelsen et al.
2011b ; Randlane & Saag 1998). Molecular: Yes.
Nipponoparmelia (Kurok.) K.H.Moon, Y.Ohmura & Kashiw. (5
spp.). New or reinstated (Crespo et al. 2010a). Molecular:
Yes.
Nodobryoria Common & Brodo (3 spp.). No change. Molecular:
Yes.
Notoparmelia A.Crespo, Ferencova & Divakar (16 spp.). New or
reinstated (Ferencova et al. 2014). Molecular: Yes.
Omphalodium Meyen & Flot. (4 spp.). No change. Molecular:
Yes.
Omphalora T.H.Nash & Hafellner (1 sp.). No change.
Molecular: Yes.
Oropogon Th.Fr. (40 spp.). No change. Molecular: Yes.
Pannoparmelia (Müll.Arg.) Darb. (6 spp.). No change.
Molecular: Yes.
Parmelia Ach. [nom.cons.] (50 spp.). No change. Molecular:
Yes.
Parmelina Hale (10 spp.). No change. Molecular: Yes.
Parmelinella Elix & Hale (7 spp.). No change. Molecular: Yes.
Parmeliopsis (Nyl.) Nyl. [nom.cons.] (3 spp.). No change.
Molecular: Yes.
Parmotrema A.Massal. (syn.: Parmelaria D.D.Awasthi) (300
spp.). New synonymy (Crespo et al. 2010a). Molecular: Yes.
Parmotremopsis Elix & Hale (2 spp.). No change. Molecular:
No.
Platismatia W.L.Culb. & C.F.Culb. (2 spp.). No change.
Molecular: Yes.
Pleurosticta Petr. (2 spp.). No change. Molecular: Yes.
Protoparmelia M.Choisy (syn.: Maronina Hafellner & R.W.Ro-
gers) (35 spp.). New synonymy (Papong et al. 2011).
Molecular: Yes.
Protousnea (Motyka) Krog (8 spp.). No change. Molecular: Yes.
Pseudephebe M.Choisy (2 spp.). No change. Molecular: Yes.
Pseudevernia Zopf (4 spp.). No change. Molecular: Yes.
Pseudoparmelia Lynge (15 spp.). No change. Molecular: Yes.
Psiloparmelia Hale (13 spp.). No change. Molecular: Yes.
Punctelia Krog (45 spp.). No change. Molecular: Yes.
Relicina (Hale & Kurok.) Hale (syn.: Relicinopsis Elix & Verdon)
(60 spp.). New synonymy (Kirika et al. 2017) Molecular:
Yes.
Remototrachyna Divakar & A.Crespo (20 spp.). New or
reinstated (Divakar et al. 2010). Molecular: Yes.
Sulcaria Bystr. (5 spp.). No change. Molecular: Yes.
Tuckermanella Essl. (6 spp.). No change. Molecular: Yes.
Tuckermanopsis Gyeln. (9 spp.). No change. Molecular: Yes.
Usnea Dill. ex Adans. (350 spp.). No change. Molecular: Yes.
Usnocetraria M.J.Lai & J.C.Wei (1 sp.). New or reinstated (Thell
et al. 2009, 2012). Molecular: Yes.
Vulpicida Mattson & M.J.Lai (6 spp.). No change. Molecular:
Yes.
Xanthoparmelia (Vain.) Hale [nom.cons.] (syn.: Karoowia Hale;
Omphalodiella Henssen; Placoparmelia Henssen) (820
spp.). New synonymy (Amo de Paz 2010a,b). Molecular:
Yes.
Family Pilocarpaceae Zahlbr.
Badimiella Malcolm & Vězda (1 sp.). No change. Molecular:
No.
Baflavia Lücking (1 sp.). No change. Molecular: No.
Bapalmuia Sérus. (23 spp.). No change. Molecular: Yes.
Barubria Vězda (2 spp.). No change. Molecular: No.
Brasilicia Lücking, Kalb & Sérus. (6 spp.). No change.
Molecular: No.
Bryogomphus Lücking, W.R.Buck, Sérus. & L.I.Ferraro (1 sp.).
No change. Molecular: No.
Byssolecania Vain. (7 spp.). No change. Molecular: Yes.
Byssoloma Trevis. (58 spp.). No change. Molecular: Yes.
Calopadia Vězda (27 spp.). No change. Molecular: Yes.
Calopadiopsis Lücking & R.Sant. (2 spp.). No change.
Molecular: No.
Eugeniella Lücking, Sérus. & Kalb (9 spp.). New or reinstated
(Lücking 2008). Molecular: Yes.
Fellhanera Vězda (97 spp.). No change. Molecular: Yes.
Fellhaneropsis Sérus. & Coppins (7 spp.). No change. Molec-
ular: Yes.
Kantvilasia P.M.McCarthy, Elix & Sérus. (1 sp.). No change.
Molecular: No.
Lasioloma R.Sant. (9 spp.). No change. Molecular: Yes.

Leimonis R.C.Harris (1 sp.). No change. Molecular: Yes.
Loflammia Vězda (5 spp.). No change. Molecular: No.
Loflammiopsis Lücking & Kalb (1 sp.). No change. Molecular:
No.
Logilvia Vězda (1 sp.). No change. Molecular: No.
Micarea Fr. [nom.cons.] (100 spp.). No change. Molecular: Yes.
Podotara Malcolm & Vězda (1 sp.). Changed classification
(Jaklitsch et al. 2016a). Molecular: No.
Pseudocalopadia Lücking (1 sp.). No change. Molecular: No.
Roccellinastrum Follmann (6 spp.). Changed classification
(Jaklitsch et al. 2016a). Molecular: No.
Septotrapelia Aptroot & Chaves (syn.: Nelsenium Lendemer &
B.P.Hodk.) (4 spp.). New synonymy (Bungartz et al. 2013).
Molecular: No.
Sporopodiopsis Sérusiaux (2 spp.). No change. Molecular: No.
Sporopodium Mont. (24 spp.). No change. Molecular: Yes.
Szczawinskia A.Funk (4 spp.). No change. Molecular: Yes.
Tapellaria Müll.Arg. (22 spp.). No change. Molecular: No.
Tapellariopsis Lücking (1 sp.). No change. Molecular: No.
Family Psilolechiaceae S.Stenroos, Mia˛dl. & Lutzoni
Psilolechia A.Massal. (4 spp.). Changed classification (Mia˛dli-
kowska et al. 2014). Molecular: Yes.
Family Psoraceae Zahlbr.
Brianaria S.Ekman & M.Svensson (4 spp.). New or reinstated
(Ekman & Svensson 2014). Molecular: Yes.
Eremastrella Vogel (2 spp.). No change. Molecular: Yes.
Glyphopeltis Brusse (1 sp.). No change. Molecular: Yes.
Protoblastenia (Zahlbr.) J.Steiner (14 spp.). No change.
Molecular: Yes.
Protomicarea Hafellner (1 sp.). No change. Molecular: Yes.
Psora Hoffm. [nom.cons.] (35 spp.). No change. Molecular: Yes.
Psorula Gotth.Schneid. (3 spp.). No change. Molecular: Yes.
Family Ramalinaceae C.Agardh (syn.: Crocyniaceae
M. Choisy ex Hafellner, Megalariaceae Hafell-
ner)
Aciculopsora Aptroot & Trest (2 spp.). No change. Molecular:
No.
Adelolecia Hertel & Hafellner (4 spp.). No change. Molecular:
No.
Arthrosporum A.Massal. (1 sp.). No change. Molecular: Yes.
Auriculora Kalb (1 sp.). Changed classification (Jaklitsch et al.
2016a). Molecular: No.
Bacidia De Not. (syn.: Lopacidia Kalb) (230 spp.). New
synonymy (this paper). Molecular: Yes. Notes. Lopacidia
was listed as separate genus by Lumbsch & Huhndorf
(2010) but the type is congeneric with Bacidia.
Bacidina Vězda [nom.cons.] (12 spp.). No change. Molecular:
Yes.
Bacidiopsora Kalb (6 spp.). No change. Molecular: No.
Lücking et al.: 2016 lichen classification 387
Badimia Vězda [nom.cons.] (18 spp.). Changed classification
(Andersen et al. 2005). Molecular: Yes.
Bilimbia De Not. (6 spp.). Changed classification (Ekman et al.
2008; Printzen 2014). Molecular: Yes.
Biatora Fr. (42 spp.). No change. Molecular: Yes.
Catillochroma Kalb (7 spp.). Changed classification (Ekman et
al. 2008; Mia˛dlikowska et al. 2014; Schmull et al. 2011).
Molecular: No.
Catinaria Vain. [nom.cons.] (2 spp.). No change. Molecular:
Yes.
Cenozosia A.Massal. (1 sp.). No change. Molecular: No.
Cliostomum Fr. (7 spp.). No change. Molecular: Yes.
Compsocladium I.M.Lamb (2 spp.). No change. Molecular: No.
Crocynia (Ach.) A.Massal. [nom.cons.] (3 spp.). Changed
classification (Andersen et al. 2005; Ekman et al. 2008).
Molecular: Yes.
Crustospathula Aptroot (3 spp.). No change. Molecular: No.
Echidnocymbium Brusse (1 sp.). No change. Molecular: No.
Eschatogonia Trevis. (7 spp.). Changed classification (Jaklitsch
et al. 2016a). Molecular: No.
Frutidella Kalb (2 spp.). No change. Molecular: Yes.
Heppsora D.D.Awasthi & K.Singh (2 spp.). No change.
Molecular: No.
Herteliana P.James (3 spp.). No change. Molecular: Yes.
Japewia Tønsberg (3 spp.). No change. Molecular: Yes.
Jarmania Kantvilas (2 spp.). No change. Molecular: No.
Krogia Timdal (3 spp.). No change. Molecular: No.
Lecania A.Massal. (50 spp.). No change. Molecular: Yes.
Lopezaria Kalb & Hafellner (2 spp.). No change. Molecular: Yes.
Lueckingia Aptroot & Umaña (1 sp.). No change. Molecular:
No.
Megalaria Hafellner (29 spp.). Changed classification (Ekman
2001; Ekman et al. 2008; Mia˛dlikowska et al. 2014; Schmull
et al. 2011). Molecular: Yes.
Niebla Rundel & Bowler (22 spp.). No change. Molecular: Yes.
Notes. The over 40 species of Niebla described by Spjut
(1996) are not generally accepted (Bowler & Marsh 2004).
Phyllopsora Müll.Arg. (syn.: Triclinum Fée) (95 spp.). New
synonymy (Timdal 2008). Molecular: Yes. Notes. The
synonymy was already proposed by Timdal (2008) but the
name Triclinum was still used in later publications,
including Aptroot & Cáceres (2014).
Physcidia Tuck. (7 spp.). No change. Molecular: No.
Pseudohepatica P.M.Jørg. (2 spp.). Changed classification
(Jaklitsch et al. 2016a). Molecular: No.
Ramalina Ach. [nom.cons.] [syn.: Ramalinopsis (Zahlbr.)
Follmann & Huneck; Vermilacinia Spjut & Hale] (230
spp.). New synonymy (Moncada et al. 2014). Molecular: Yes.
Rolfidium Moberg (2 spp.). No change. Molecular: Yes.
Schadonia Körb. (3 spp.). No change. Molecular: Yes.
Stirtoniella D.J.Galloway, Hafellner & Elix (1 sp.). No change.
Molecular: No.
Tasmidella Kantvilas, Hafellner & Elix (1 sp.). Changed
classification (Ekman et al. 2008; Mia˛dlikowska et al.
2014; Schmull et al. 2011). Molecular: No.
388 The Bryologist 119(4): 2016
Thamnolecania (Vain.) Gyeln. (7 spp.). No change. Molecular:
Yes.
Tibellia Vězda & Hafellner (1 sp.). No change. Molecular: No.
Toninia A.Massal. [nom.cons.] (85 spp.). No change. Molecular:
Yes.
Toniniopsis Frey (1 sp.). No change. Molecular: No.
Waynea Moberg (7 spp.). No change. Molecular: Yes.
Family Ramboldiaceae S.Stenroos, Mia˛dl. & Lutzoni
Ramboldia Kantvilas & Elix (30 spp.). Changed classification
(Mia˛dlikowska et al. 2014). Molecular: Yes.
Family Scoliciosporaceae Hafellner
Scoliciosporum A.Massal. (15 spp.). No change. Molecular: Yes.
Family Sphaerophoraceae Fr. (syn.: Calycidiaceae
Elenkin)
Austropeltum Henssen, Döring & Kantvilas (1 sp.). No change.
Molecular: Yes.
Bunodophoron A.Massal. (24 spp.). No change. Molecular: Yes.
Calycidium Stirt. (2 spp.). Changed classification (Jaklitsch et al.
2016a). Molecular: Yes.
Leifidium Wedin (1 sp.). No change. Molecular: Yes.
Neophyllis F.Wilson (2 spp.). No change. Molecular: Yes.
Sphaerophorus Pers. [nom.cons.] (8 spp.). No change. Molec-
ular: Yes.
Family Stereocaulaceae Chevall.
HertelideaPrintzen& Kantvilas (6 spp.). No change.Molecular:Yes.
Lepraria Ach. [nom.cons.] (80 spp.). No change. Molecular:
Yes.
Stereocaulon Hoffm. [nom.cons.] (140 spp.). No change.
Molecular: Yes.
Squamarina Poelt (25 spp.). No change. Molecular: Yes.
Xyleborus R.C.Harris & Ladd (1 sp.). No change. Molecular: No.
Family Tephromelataceae Hafellner (syn.: Mycoblas-
taceae Hafellner)
Calvitimela Hafellner (10 spp.). Changed classification (Bend-
iksby et al. 2015). Molecular: Yes.
Mycoblastus Norman [nom.cons.] (10 spp.). Changed classifi-
cation (Spribille et al. 2011). Molecular: Yes.
Tephromela M.Choisy (30 spp.). Changed classification (Spri-
bille et al. 2011). Molecular: Yes.
Violella T.Sprib. (2 spp.). New or reinstated (Spribille et al.
2011). Molecular: Yes.
Order Lecanorales genera incertae sedis
Coronoplectrum Brusse (1 sp.). Changed classification (Thell et
al. 2012). Molecular: No.
Ivanpisutia S.Y.Kondr., L.Lokös & Hur (1 sp.). New or
reinstated (Kondratyuk et al. 2015a). Molecular: No.
Joergensenia Passo, Stenroos & Calvelo (1 sp.). No change.
Molecular: Yes.
Myochroidea Printzen, T.Sprib. & Tønsberg (4 spp.). No
change. Molecular: No.
Neopsoromopsis Gyeln. (1 sp.). Changed classification (Thell et
al. 2012). Molecular: No.
Psoromella Gyeln. (1 sp.). Changed classification (Thell et al.
2012). Molecular: No.
Puttea S.Stenroos & Huhtinen (3 spp.). No change. Molecular:
Yes.
Ramalea Nyl. (4 spp.). No change. Molecular: Yes.
Order LECIDEALES Vain.
Family Lecideaceae Chevall.
Amygdalaria Norman (14 spp.). Changed classification
(Schmull et al. 2011). Molecular: Yes.
Bahianora Kalb (1 sp.). Changed classification (Schmull et al.
2011). Molecular: No.
Bellemerea Hafellner & Cl.Roux (8 spp.). Changed classification
(Schmull et al. 2011). Molecular: Yes.
Bryobilimbia Fryday, Printzen & S.Ekman (6 spp.). New or
reinstated (Fryday et al. 2014). Molecular: Yes.
Catarrhospora Brusse (2 spp.). Changed classification (Schmull
et al. 2011). Molecular: No.
Cecidonia Triebel & Rambold (2 spp.). Changed classification
(Schmull et al. 2011). Molecular: Yes.
Clauzadea Hafellner & Bellem. (7 spp.). Changed classification
(Schmull et al. 2011). Molecular: Yes.
Cryptodictyon A.Massal. (2 spp.). Changed classification
(Schmull et al. 2011). Molecular: No.
Farnoldia Hertel (6 spp.). Changed classification (Schmull et al.
2011). Molecular: Yes.
Immersaria Rambold & Pietschm. (8 spp.). Changed classifica-
tion (Schmull et al. 2011). Molecular: Yes.
Koerberiella Stein (2 spp.). Changed classification (Schmull et
al. 2011). Molecular: Yes.
Labyrintha Malcolm, Elix & OweLarsson (1 sp.). Changed
classification (Schmull et al. 2011). Molecular: No.
Lecidea Ach. (100 spp.). Changed classification (Schmull et al.
2011). Molecular: Yes.
Lecidoma Gotth.Schneid. & Hertel (1 sp.). Changed classifica-
tion (Schmull et al. 2011). Molecular: Yes.
Melanolecia Hertel (7 spp.). Changed classification (Buschbom
& Müller 2004). Molecular: Yes. Notes. Published phylog-
enies are inconclusive about the position of this genus, but
revision of sequence data suggest placement in Lecideaceae.
Mycobilimbia Rehm (5 spp.). Changed classification (Schmull et
al. 2011). Molecular: Yes.
Pachyphysis R.C.Harris & Ladd (1 sp.). Changed classification
(Schmull et al. 2011). Molecular: Yes.
Paraporpidia Rambold & Pietschm. (3 spp.). Changed classi-
fication (Schmull et al. 2011). Molecular: No.

Poeltiaria Hertel (8 spp.). Changed classification (Schmull et al.
2011). Molecular: Yes.
Poeltidea Hertel & Hafellner (3 spp.). Changed classification
(Schmull et al. 2011). Molecular: No.
Porpidia Körb. (50 spp.). Changed classification (Schmull et al.
2011). Molecular: Yes.
Pseudopannaria (B.de Lesd.) Zahlbr. (1 sp.). Changed classifi-
cation (Schmull et al. 2011). Molecular: No.
Rhizolecia Hertel (1 sp.). No change. Molecular: No.
Romjularia Timdal (1 sp.). Changed classification (Schmull et
al. 2011). Molecular: Yes.
Schizodiscus Brusse (1 sp.). Changed classification (Schmull et
al. 2011). Molecular: No.
Stenhammarella Hertel (1 sp.). Changed classification (Busch-
bom & Müller 2004). Molecular: Yes. Notes. Published
phylogenies are inconclusive about the position of this
genus, but revision of sequence data suggest placement in
Lecideaceae.
Stephanocyclos Hertel (1 sp.). Changed classification (Schmull
et al. 2011). Molecular: Yes.
Xenolecia Hertel (1 sp.). Changed classification (Schmull et al.
2011). Molecular: No.
Family Lopadiaceae Hafellner
Lopadium Körb. [nom.cons.] (5 spp.). Changed classification
(Ekman et al. 2008). Molecular: Yes.
Order LEPROCAULALES Lendemer & B.P.Hodk.
Family Leprocaulaceae Lendemer & B.P.Hodk.
Halecania M.Mayrhofer (22 spp.). Changed classification
(Lendemer & Hodkinson 2013). Molecular: Yes.
Leprocaulon Nyl. (syn.: Byssocaulon Mont.) (12 spp.). Changed
classification and new synonymy (Lendemer & Hodkinson
2013; this paper). Molecular: Yes. Notes. Byssocaulon was
listed under Chrysotrichaceae (Lumbsch & Huhndorf
2010), but the type appears to be congeneric with
Leprocaulon.
Speerschneidera Trevis. (1 sp.). Changed classification (Lend-
emer & Hodkinson 2013). Molecular: Yes.
Order PELTIGERALES Walt.Watson
Suborder Collematineae Mia˛dl. & Lutzoni
Family Coccocarpiaceae (Mont. ex Müll.Stuttg.)
Henssen
Coccocarpia Pers. (30 spp.). No change. Molecular: Yes.
Peltularia R.Sant. (4 spp.). No change. Molecular: No.
Spilonema Bornet (syn.: Spilonemella Henssen & Tønsberg) (5
spp.). New synonymy (Spribille et al. 2014). Molecular: Yes.
Family Collemataceae Zenker
Blennothallia Trevis. (4 spp.). New or reinstated (Otálora et al.
2013a, 2014). Molecular: Yes.
Lücking et al.: 2016 lichen classification 389
Callome Otálora & Wedin (1 sp.). New or reinstated (Otálora et
al. 2013a, 2014). Molecular: Yes.
Collema F.H.Wigg. [nom.cons.] (40 spp.). No change. Molec-
ular: Yes.
Enchylium (Ach.) Gray (9 spp.). New or reinstated (Otálora et
al. 2013a, 2014). Molecular: Yes.
Lathagrium (Ach.) Gray (10 spp.). New or reinstated (Otálora et
al. 2013a, 2014). Molecular: Yes.
Leptogium (Ach.) Gray(syn.: Epiphloea Trevis.) (70 spp.). New
synonymy (Schultz et al. 2015). Molecular: Yes.
Paracollema Otálora & Wedin (2 spp.). New or reinstated
(Otálora et al. 2013a, 2014). Molecular: Yes.
Pseudoleptogium Müll.Arg. (1 sp.). New or reinstated (Otálora
et al. 2013a, 2014). Molecular: Yes.
Rostania Trevis. (7 spp.). New or reinstated (Otálora et al.
2013a, 2014). Molecular: Yes.
Scytinium (Ach.) Gray (48 spp.). New or reinstated (Otálora et
al. 2013a, 2014). Molecular: Yes.
Family Pannariaceae Tuck.
Austrella P.M.Jørg. (2 spp.). No change. Molecular: Yes.
Degelia Arv. & D.J.Galloway (22 spp.). No change. Molecular:
Yes.
Degeliella P.M.Jørg. (2 spp.). No change. Molecular: Yes.
Erioderma Feé (40 spp.). No change. Molecular: Yes.
Fuscoderma (D.J.Galloway & P.M.Jørg.) P.M.Jørg. & D.J.Gallo-
way (5 spp.). No change. Molecular: Yes.
Fuscopannaria P.M.Jørg. [nom.cons.prop.] (syn.: Moelleropsis
Gyeln.) (58 spp.). New synonymy (Ekman et al. 2014).
Molecular: Yes.
Gibbosporina Elvebakk, Hong & P.M.Jørg. (13 spp.). New or
reinstated (Elvebakk et al. 2016). Molecular: Yes.
Homothecium A.Massal. (5 spp.). Changed classification (Ek-
man et al. 2014). Molecular: No.
Hosseusia Gyeln. (1 sp.). No change. Molecular: No.
Kroswia P.M.Jørg. (4 spp.). No change. Molecular: Yes.
Leciophysma Th.Fr. (2 spp.). Changed classification (Ekman et
al. 2014; Wedin et al. 2009). Molecular: Yes.
Leightoniella Henssen (1 sp.). Changed classification (Weer-
akoon et al. 2016). Molecular: Yes.
Leioderma Nyl. (7 spp.). No change. Molecular: Yes.
Lepidocollema Vain. (2 spp.). No change. Molecular: Yes.
Leptogidium Nyl. (3 spp.). New or reinstated (Muggia et al.
2011). Molecular: Yes.
Nebularia P.M.Jørg. (2 spp.). New or reinstated (Ekman et al.
2014). Molecular: Yes.
Nevesia P.M.Jørg, L.Lindblom, Wedin & S.Ekman (1 sp.). New
or reinstated (Ekman et al. 2014). Molecular: Yes.
Pannaria Del. ex Bory (40 spp.). No change. Molecular: Yes.
Parmeliella Müll.Arg. (90 spp.). No change. Molecular: Yes.
Pectenia P.M.Jørg., L.Lindblom, Wedin & S.Ekman (4 spp.).
New or reinstated (Ekman et al. 2014). Molecular: Yes.
Physma A.Massal. (12 spp.). Changed classification (Ekman et
al. 2014; Wedin et al. 2009). Molecular: Yes.
390 The Bryologist 119(4): 2016
Protopannaria (Gyeln.) P.M.Jørg. & S.Ekman (5 spp.). No
change. Molecular: Yes.
Psoroma Michaux (syn.: Santessoniella Henssen) (70 spp.). New
synonymy (Ekman et al. 2014). Molecular: Yes.
Psoromidium Stirt. (2 spp.). No change. Molecular: Yes.
Ramalodium Nyl. (5 spp.). Changed classification (Ekman et al.
2014; Wedin et al. 2009). Molecular: Yes.
Siphulastrum Müll.Arg. (4 spp.). No change. Molecular: Yes.
Staurolemma Körb. (3 spp.). Changed classification (Ekman et
al. 2014; Wedin et al. 2009). Molecular: Yes.
Steineropsis T.Sprib. & Muggia (1 sp.). New or reinstated (Spribille
et al. 2010; Spribille & Muggia 2013). Molecular: Yes.
Family Placynthiaceae E.Dahl [nom.cons.]
Hertella Henssen (3 spp.). No change. Molecular: No.
Placynthiopsis Zahlbr. (1 sp.). No change. Molecular: No.
Placynthium (Ach.) Gray (syn.: Collolechia A.Massal.; Psoroti-
chiella Werner) (25 spp.). New synonymy (Košuthová et al.
2016; Moreno & Egea 1990). Molecular: Yes.
Suborder Peltigerineae Mia˛dl. & Lutzoni
Family Koerberiaceae T.Sprib. & Muggia
Koerberia A.Massal. (2 spp.). Changed classification (Spribille &
Muggia 2013). Molecular: Yes.
Steinera Zahlbr. (4 spp.). Changed classification (Spribille &
Muggia 2013). Molecular: Yes.
Vestergrenopsis Gyeln. (2 spp.). Changed classification (Spribille
& Muggia 2013). Molecular: Yes.
Family Lobariaceae Chevall.
Crocodia Link (5 spp.). New or reinstated (Galloway & Elix
2013; Moncada et al. 2013). Molecular: No.
Dendriscosticta Moncada & Lücking (5 spp.). New or reinstated
(Moncada et al. 2013a). Molecular: Yes.
Lobaria (Schreb.) Hoffm. (syn.: Anomalobaria Moncada &
Lücking) (60 spp.). New synonymy (Cornejo & Scheidegger
2015; Mia˛dlikowska et al. 2014; Moncada et al. 2013a).
Molecular: Yes.
Lobariella Yoshim. (35 spp.). New or reinstated (Moncada et al.
2013a). Molecular: Yes.
Lobarina Nyl. ex Cromb. (15 spp.). New or reinstated (Moncada
et al. 2013a). Molecular: Yes.
Parmostictina Nyl. (15 spp.). New or reinstated (Moncada et al.
2013a). Molecular: Yes.
Podostictina Clem. (6 spp.). New or reinstated (Moncada et al.
2013a). Molecular: Yes.
Pseudocyphellaria Vain. [nom.cons.] (100 spp.). No change.
Molecular: Yes.
Ricasolia De Not. (syn.: Dendriscocaulon Nyl.) (15 spp.). New or
reinstated and new synonymy (Moncada et al. 2013a).
Molecular: Yes.
Sticta (Schreb.) Ach. (200 spp.). No change. Molecular: Yes.
Yarrumia D.J.Galloway (2 spp.). New or reinstated (Galloway
2015). Molecular: Yes.
Yoshimuriella Moncada & Lücking (8 spp.). New or reinstated
(Moncada et al. 2013a). Molecular: Yes.
Family Massalongiaceae Wedin, P.M.Jørg. & Wiklund
Leptochidium M.Choisy (2 spp.). Changed classification (Wedin
et al. 2007). Molecular: Yes.
Massalongia Körb. (2 spp.). Changed classification (Wedin et al.
2007). Molecular: Yes.
Polychidium (Ach.) Gray (1 sp.). Changed classification (Wedin
et al. 2007). Molecular: Yes.
Family Nephromataceae Wetm. ex J.C.David &
D.Hawksw.
Nephroma Ach. (36 spp.). No change. Molecular: Yes.
Family Peltigeraceae Dumort.
Peltigera Willd. [nom.cons.] (100 spp.). No change. Molecular: Yes.
Solorina Ach. (10 spp.). No change. Molecular: Yes.
Family Vahliellaceae Wedin, P.M.Jørg. & S.Ekman
Vahliella P.M.Jørg. (8 spp.). Changed classification (Wedin et
al. 2011). Molecular: Yes.
Order Peltigerales genera incertae sedis
Erinacellus T.Sprib., Muggia & Tønsberg (2 spp.). New or
reinstated (Spribille et al. 2014). Molecular: Yes.
Order RHIZOCARPALES Mia˛dl. et al. ex Mia˛dl. &
Lutzoni
Family Rhizocarpaceae M.Choisy & Hafellner
Catolechia Flot. (1 sp.). Changed classification (Mia˛dlikowska et
al. 2006). Molecular: Yes.
Epilichen Clem. (3 spp.). Changed classification (Mia˛dlikowska
et al. 2006). Molecular: No.
Poeltinula Hafellner (2 spp.). Changed classification (Mia˛dli-
kowska et al. 2006). Molecular: Yes.
Rhizocarpon Ramond ex DC. (225 spp.). Changed classification
(Mia˛dlikowska et al. 2006). Molecular: Yes.
Family Sporastatiaceae Bendiksby & Timdal
Sporastatia A.Massal. (4 spp.). Changed classification (Bend-
iksby & Timdal 2013; Mia˛dlikowska et al. 2006). Molecular:
Yes.
Toensbergia Bendiksby & Timdal (1 sp.). New or reinstated
(Bendiksby & Timdal 2013). Molecular: Yes.
Order TELOSCHISTALES D.Hawksw. & O.E.Erikss.
Suborder Letrouitineae Gaya & Lutzoni

Family Brigantieaceae Hafellner & Bellem.
Brigantiaea Trevis. (29 spp.). Changed classification (Gaya et al.
2012). Molecular: Yes.
Family Letrouitiaceae Bellem. & Hafellner
Letrouitia Hafellner & Bellem. (18 spp.). No change. Molecular:
Yes.
Suborder Teloschistineae Gaya & Lutzoni
Family Megalosporaceae Vězda ex Hafellner &
Bellem.
Megaloblastenia Sipman (2 spp.). No change. Molecular: Yes.
Megalospora Meyen (syn.: Austroblastenia Sipman) (35 spp.).
New synonymy (Kantvilas & Lumbsch 2012). Molecular:
Yes.
Sipmaniella Kalb (1 sp.). Changed classification (Gaya et al.
2012). Molecular: Yes.
Family Teloschistaceae Zahlbr.
Subfamily Brownlielloideae S.Y.Kondr., Kärnefelt,
Elix & A.Thell
Brownliella S.Y.Kondr., Kärnefelt, Elix, A.Thell & Hur (4 spp.).
New or reinstated (Kondratyuk et al. 2013, 2015b).
Molecular: Yes.
Lazarenkoella S.Y.Kondr., Kärnefelt, A.Thell, Elix & Hur (1 sp.).
New or reinstated (Kondratyuk et al. 2015b). Molecular:
Yes.
Marchantiana S.Y.Kondr., Kärnefelt, Elix, A.Thell & Hur (6
spp.). New or reinstated (Kondratyuk et al. 2014a, 2015b).
Molecular: Yes.
Raesaeneniana S.Y.Kondr., Kärnefelt, A.Thell, Elix & Hur (1
sp.). New or reinstated (Kondratyuk et al. 2015b).
Molecular: Yes.
Streimanniella S.Y.Kondr., Kärnefelt, A.Thell, Elix & Hur (4
spp.). New or reinstated (Kondratyuk et al. 2015b).
Molecular: Yes.
Tarasginia S.Y.Kondr., Kärnefelt, A.Thell, Elix & Hur (2 spp.).
New or reinstated (Kondratyuk et al. 2015b). Molecular:
Yes.
Tayloriellina S.Y.Kondr., Kärnefelt., A.Thell, Elix & Hur [syn.:
Tayloriella S.Y.Kondr., Kärnefelt, A.Thell, Elix & Hur
[nom.illeg.], non Tayloriella Kylin (Rhodophyta)] (1 sp.).
New or reinstated and new synonymy (Kondratyuk et al.
2015b, 2016). Molecular: Yes.
Thelliana S.Y.Kondr., Kärnefelt, Elix & Hur (1 sp.). New or
reinstated (Kondratyuk et al. 2015b). Molecular: Yes.
Subfamily Caloplacoideae Arup, Søchting & Frödén
Apatoplaca Poelt & Hafellner (1 sp.). New or reinstated
(Vondrák et al. 2012). Molecular: No.
Blastenia A.Massal. (11 spp.). New or reinstated (Arup et al.
2013). Molecular: Yes.
Bryoplaca Søchting, Frödén & Arup (3 spp.). New or reinstated
(Arup et al. 2013). Molecular: Yes.
Lücking et al.: 2016 lichen classification 391
Caloplaca Th.Fr. [nom.cons.] (350 spp.). Changed classification
(Arup et al. 2013). Molecular: Yes.
Cephalophysis (Hertel) H.Kilias (1 sp.). Changed classification
(Arup et al. 2013). Molecular: No.
Eilifdahlia S.Y.Kondr., Kärnefelt, Elix, A.Thell & Hur (2 spp.).
New or reinstated (Kondratyuk et al. 2014a). Molecular:
Yes.
Fauriea S.Y.Kondr., L.L}okös & J.-S.Hur (2 spp.). New or
reinstated (Kondratyuk et al. 2016). Molecular: Yes.
Franwilsia S.Y.Kondr., Kärnefelt, Elix, A.Thell & Hur (3 spp.).
New or reinstated (Kondratyuk et al. 2014a). Molecular:
Yes.
Gyalolechia A.Massal. (syn.: Elenkiniana S.Y.Kondr., Kärnefelt,
Elix & Hur; Fulgensia A.Massal. & De Not.; Mikhtomia
S.Y.Kondr., Kärnefelt, Elix & Hur) (30 spp.). New or
reinstated and new synonymy (Arup et al. 2013; Jaklitsch et
al. 2016a ; Kondratyuk et al. 2014a). Molecular: Yes.
Huneckia S.Y.Kondr., Elix, Kärnefelt, A.Thell & Hur (2 spp.).
New or reinstated (Kondratyuk et al. 2014a). Molecular:
Yes.
Ioplaca Poelt (2 spp.). Changed classification (Arup et al. 2013).
Molecular: Yes.
Jasonhuria S.Y.Kondr., L.L}okös & S.O.Oh (1 sp.). New or
reinstated (Kondratyuk et al. 2015d). Molecular: Yes.
Leproplaca (Nyl.) Nyl. (6 spp.). New or reinstated (Arup et al.
2013). Molecular: Yes.
Loekoesia S.Y.Kondr., S.O.Oh & Hur (1 sp.). New or reinstated
(Kondratyuk et al. 2015d). Molecular: Yes.
Olegblumia S.Y.Kondr., L.L} okös & Hur (1 sp.). New or
reinstated (Kondratyuk et al. 2015d). Molecular: Yes.
Pyrenodesmia A.Massal. (6 spp.). New or reinstated (Arup et al.
2013). Molecular: Yes.
Rufoplaca Arup, Søchting & Frödén (6 spp.). New or reinstated
(Arup et al. 2013). Molecular: Yes.
Seirophora Poelt (11 spp.). Changed classification (Arup et al.
2013). Molecular: Yes.
Usnochroma Søchting, Arup & Frödén (2 spp.). New or
reinstated (Arup et al. 2013). Molecular: Yes.
Variospora Arup, Søchting & Frödén (12 spp.). New or
reinstated (Arup et al. 2013). Molecular: Yes.
Yoshimuria S.Y.Kondr., Kärnefelt, Elix, A.Thell & Hur (3 spp.).
New or reinstated (Kondratyuk et al. 2014a). Molecular:
Yes.
Subfamily Teloschistoideae Arup, Søchting & Frödén
Catenarina Søchting, Søgaard, Arup, Elvebakk & Elix (3 spp.).
New or reinstated (Søchting et al. 2014a). Molecular: Yes.
Filsoniana S.Y.Kondr., Kärnefelt, Elix, A.Thell & Hur (5 spp.).
New or reinstated (Kondratyuk et al. 2013). Molecular: Yes.
Follmannia C.W.Dodge (2 spp.). New or reinstated (Arup et al.
2013). Molecular: Yes.
Haloplaca Arup, Søchting & Frödén (3 spp.). New or reinstated
(Arup et al. 2013). Molecular: Yes.
Josefpoeltia S.Y.Kondr. & Kärnefelt (3 spp.). Changed classifi-
cation (Arup et al. 2013). Molecular: Yes.
Kaernefia S.Y.Kondr., Elix, A.Thell & Hur (3 spp.). New or
reinstated (Kondratyuk et al. 2013). Molecular: Yes.
392 The Bryologist 119(4): 2016
Scutaria Søchting, Arup & Frödén (1 sp.). New or reinstated
(Arup et al. 2013). Molecular: Yes.
Sirenophila Søchting, Arup & Frödén (8 spp.). New or
reinstated (Arup et al. 2013). Molecular: Yes.
Stellarangia Frödén, Arup & Søchting (3 spp.). New or
reinstated (Arup et al. 2013). Molecular: Yes.
Tassiloa S.Y.Kondr., Kärnefelt, A.Thell, Elix & Hur (2 spp.).
New or reinstated (Kondratyuk et al. 2015e). Molecular:
Yes.
Teloschistes Norman (syn.: Niorma A.Massal.) (20 spp.). New
synonymy (Kondratyuk et al. 2013). Molecular: Yes.
Teloschistopsis Frödén, Søchting & Arup (3 spp.). New or
reinstated (Arup et al. 2013). Molecular: Yes.
Villophora Søchting, Arup & Frödén (2 spp.). New or reinstated
(Arup et al. 2013). Molecular: Yes.
Wetmoreana Arup, Søchting & Frödén (syn.: Fulgogasparrea
S.Y.Kondr., N.-H.Jeong, Kärnefelt, Elix, A.Thell & Hur) (3
spp.). New or reinstated and new synonymy (Arup et al.
2013; Kondratyuk et al. 2013; Jaklitsch et al. 2016a).
Molecular: Yes.
Subfamily Xanthorioideae Arup, Søchting & Frödén
Amundsenia Søchting, Garrido-Benavent, Arup & Frödén (2
spp.). New or reinstated (Søchting et al. 2014b). Molecular:
Yes.
Athallia Arup, Frödén & Søchting (13 spp.). New or reinstated
(Arup et al. 2013). Molecular: Yes.
Austroplaca Søchting, Frödén & Arup (10 spp.). New or
reinstated (Arup et al. 2013). Molecular: Yes.
Calogaya Arup, Frödén & Søchting (14 spp.). New or reinstated
(Arup et al. 2013). Molecular: Yes.
Cerothallia Arup, Frödén & Søchting (4 spp.). New or reinstated
(Arup et al. 2013). Molecular: Yes.
Charcotiana Søchting, Garrido-Benavent & Arup (1 sp.). New
or reinstated (Søchting et al. 2014b). Molecular: Yes.
Dufourea Ach. (syn.: Jackelixia S.Y.Kondr., Fedorenko, S.Sten-
roos, Kärnefelt & A.Thell; Langeottia S.Y.Kondr., Kärnefelt,
Elix, A.Thell & Hur; Ovealmbornia S.Y.Kondr., Fedorenko,
S.Stenroos, Kärnefelt, Elix & A.Thell; Xanthodactylon
P.A.Duvign.; Xanthokarrooa S.Y.Kondr., Fedorenko,
S.Stenroos, Kärnefelt, Elix & A.Thell) (23 spp.). New or
reinstated and new synonymy (Arup et al. 2013; Arup et al.
2013; Jaklitsch et al. 2016a ; Kondratyuk et al. 2014b).
Molecular: Yes.
Flavoplaca Arup, Søchting & Frödén (27 spp.). New or
reinstated (Arup et al. 2013). Molecular: Yes.
Gondwania Søchting, Frödén & Arup (4 spp.). New or
reinstated (Arup et al. 2013). Molecular: Yes.
Orientophila Arup, Søchting & Frödén (5 spp.). New or
reinstated (Arup et al. 2013). Molecular: Yes.
Pachypeltis Søchting, Arup & Frödén (4 spp.). New or reinstated
(Arup et al. 2013). Molecular: Yes.
Parvoplaca Arup, Søchting & Frödén (4 spp.). New or reinstated
(Arup et al. 2013). Molecular: Yes.
Polycauliona Hue (syn.: Igneoplaca S.Y.Kondr., Kärnefelt, Elix,
A.Thell & Hur; Massjukiella S.Y.Kondr., Fedorenko,
S.Stenroos, Kärnefelt, Elix, J.S.Hur & A.Thell [nom. illeg.],
Scythioria S.Y.Kondr., Kärnefelt, Elix, A.Thell & Hur;
Verrucoplaca S.Y.Kondr., Kärnefelt, Elix, A.Thell & Hur)
(25 spp.). New or reinstated and new synonymy (Arup et al.
2013; Jaklitsch et al. 2016a ; Kondratyuk et al. 2014b).
Molecular: Yes. Notes. The genus Diblastia Trevis. was
originally based on Lichen candelarius L. and hence
provides an earlier name for Polycauliona; however, Arup
et al. (2013) changed the type to Lichen vitellinus Ehrh.,
presumably through lectotypification, thus making Diblas-
tia an earlier name threatening Candelariella. We believe
that this lectotypification is incorrect, since L. vitellinus was
listed as synonym of L. candelarius in the protologue and
hence a single species was accepted in the genus Diblastia,
which consequently must be considered the holotype;
therefore, the type can only be changed by conservation.
This issue needs further clarification, as it affects the
current use of the names Polycauliona and Candelariella.
Rusavskia S.Y.Kondr. & Kärnefelt (18 spp.). New or reinstated
(Arup et al. 2013; Kondratyuk & Kärnefelt 2003).
Molecular: Yes.
Shackletonia Søchting, Frödén & Arup (5 spp.). New or
reinstated (Arup et al. 2013). Molecular: Yes.
Solitaria Arup, Søchting & Frödén (1 sp.). New or reinstated
(Arup et al. 2013). Molecular: Yes.
Squamulea Arup, Søchting & Frödén (6 spp.). New or reinstated
(Arup et al. 2013). Molecular: Yes.
Xanthocarpia A.Massal. & De Not. (13 spp.). New or reinstated
(Arup et al. 2013). Molecular: Yes.
Xanthomendoza S.Y.Kondr. & Kärnefelt (syn.: Gallowayella
S.Y.Kondr., Fedorenko, S.Stenroos, Kärnefelt, Elix &
A.Thell; Golubkovia S.Y.Kondr., Kärnefelt, Elix, A.Thell &
Hur; Honeggeria S.Y.Kondr., Fedorenko, S.Stenroos,
Kärnefelt, Elix, Hur & A.Thell; Jesmurraya S.Y.Kondr.,
Fedorenko, S.Stenroos, Kärnefelt, Elix, Hur & A.Thell;
Oxneria S.Y.Kondr. & Kärnefelt) (20 spp.). New synonymy
(Arup et al. 2013; Jaklitsch et al. 2016a ; Kondratyuk et al.
2014b). Molecular: Yes.
Xanthopeltis R.Sant. (1 sp.). Changed classification (Arup et al.
2013). Molecular: Yes.
Xanthoria (Fr.) Th.Fr. [nom.cons.] (10 spp.). Changed
classification (Arup et al. 2013). Molecular: Yes.
Zeroviella S.Y.Kondr. & J.-S.Hur (8 spp.). New or reinstated
(Kondratyuk et al. 2015f). Molecular: Yes.
Order Teloschistales genera incertae sedis
Malcolmiella Vězda (1 sp.). Changed classification (Kalb et al.
2011). Molecular: Yes.
Subclass Lecanoromycetidae families incertae sedis
Family Helocarpaceae Hafellner
Helocarpon Fr. (3 spp.). Changed classification (Mia˛dlikowska
et al. 2014). Molecular: Yes.
Subclass OSTROPOMYCETIDAE Reeb, Lutzoni &
Cl.Roux
Order ARCTOMIALES S.Stenroos, Mia˛dl. & Lutzoni

Family Arctomiaceae Th.Fr.
Arctomia Th.Fr. (14 spp.). Changed classification (Mia˛dlikow-
ska et al. 2014; Otálora et al. 2013b). Molecular: Yes.
Gregorella Lumbsch (1 sp.). Changed classification (Mia˛dli-
kowska et al. 2014). Molecular: Yes.
Wawea Henssen & Kantvilas (1 sp.). Changed classification
(Mia˛dlikowska et al. 2014). Molecular: Yes.
Order BAEOMYCETALES Lumbsch, Huhndorf & Lutzo-
ni
Family Baeomycetaceae Dumort. (syn.: Anamylop-
soraceae Lumbsch & Lunke)
Ainoa Lumbsch & I.Schmitt (3 spp.). No change. Molecular:
Yes.
Anamylopsora Timdal (1 sp.). Changed classification (Resl et al.
2015). Molecular: Yes.
Baeomyces Pers. (9 spp.). No change. Molecular: Yes.
Parainoa Resl & T.Sprib. (1 sp.). New or reinstated (Resl et al.
2015). Molecular: Yes.
Phyllobaeis Kalb & Gierl (5 spp.). No change. Molecular: Yes.
Family Cameroniaceae Kantvilas & Lumbsch
Cameronia Kantvilas (2 spp.). New or reinstated (Kantvilas
2012; Lumbsch et al. 2012). Molecular: Yes.
Order HYMENELIALES S.Stenroos, Mia˛dl. & Lutzoni
Family Hymeneliaceae Körb.
Hymenelia Kremp. (25 spp.). Changed classification (Mia˛dli-
kowska et al. 2014). Molecular: Yes.
Ionaspis Th.Fr. (7 spp.). Changed classification (Mia˛dlikowska
et al. 2014). Molecular: Yes.
Tremolecia M.Choisy (6 spp.). Changed classification (Mia˛dli-
kowska et al. 2014). Molecular: Yes.
Order OSTROPALES Nannf.
Family Coenogoniaceae (Fr.)Stizenb.
Coenogonium Ehrenb. ex Nees (syn.: Dimerella Trevis.) (90
spp.). New synonymy (Baloch et al. 2010; Kauff & Lutzoni
2002; Lücking & Kalb 2000; Rivas Plata et al. 2006).
Molecular: Yes.
Family Gomphillaceae Walt.Watson
Actinoplaca Müll.Arg. (2 spp.). No change. Molecular: Yes.
Aderkomyces Bat. (30 spp.). No change. Molecular: Yes.
Aplanocalenia Lücking, Sérus. & Vězda (1 sp.). No change.
Molecular: No.
Arthotheliopsis Vain. (5 spp.). No change. Molecular: Yes.
Asterothyrium Müll.Arg. (33 spp.). No change. Molecular: Yes.
Aulaxina Fée (14 spp.). No change. Molecular: Yes.
Lücking et al.: 2016 lichen classification 393
Calenia Müll.Arg. (30 spp.). No change. Molecular: Yes.
Caleniopsis Vězda & Poelt (2 spp.). No change. Molecular:
Yes.
Diploschistella Vain. (4 spp.). No change. Molecular: No.
Echinoplaca Fée (40 spp.). No change. Molecular: Yes.
Ferraroa Lücking, Sérus. & Vězda (1 sp.). No change. Molecular:
No.
Gomphillus Nyl. (6 spp.). No change. Molecular: Yes.
Gyalectidium Müll.Arg. (52 spp.). No change. Molecular: Yes.
Gyalidea Lettau ex Vězda [nom.cons.] (50 spp.). No change.
Molecular: Yes.
Gyalideopsis Vězda [nom.cons.] (95 spp.). No change. Molec-
ular: Yes.
Hippocrepidea Sérus. (1 sp.). No change. Molecular: No.
Jamesiella Lücking, Sérus. & Vězda (4 spp.). No change.
Molecular: No.
Lithogyalideopsis Lücking, Sérus. & Vězda (4 spp.). No change.
Molecular: No.
Paratricharia Lücking (1 sp.). No change. Molecular: No.
Phyllogyalidea Lücking (2 spp.). No change. Molecular: No.
Psorotheciopsis Rehm (7 spp.). No change. Molecular: No.
Rolueckia Papong, Thammath. & Boonpr. (2 spp.). No change.
Molecular: Yes.
Rubrotricha Lücking, Sérus. & Vězda (2 spp.). No change.
Molecular: Yes.
Tricharia Fée (30 spp.). No change. Molecular: Yes.
Family Graphidaceae Dumort. [syn.: Fissurinaceae (Rivas
Plata, Lücking & Lumbsch) B.P. Hodk.; Thelotremataceae
Stizenb.]. Notes. The division of Graphidaceae into three
families as proposed by Hodkinson (2012) is not followed
here. While Fissurinaceae sensu Hodkinson is monophy-
letic, it does not exhibit consistent phenotypic differences
towards the remainder of the family; Graphidaceae sensu
Hodkinson (2012) is not monophyletic and both Graph-
idaceae and Thelotremataceae sensu Hodkinson (2012)
include graphidoid and thelotremoid lineages, making their
phenotypic distinction arbitrary (Rivas Plata et al. 2013;
Lumbsch et al. 2014).
Subfamily Fissurinoideae Rivas Plata, Lücking &
Lumbsch
Clandestinotrema Rivas Plata, Lücking & Lumbsch (17 spp.).
New or reinstated (Rivas Plata et al. 2012a). Molecular:
Yes.
Cruentotrema Rivas Plata, Papong, Lumbsch & Lücking (5
spp.). New or reinstated (Rivas Plata et al. 2012a).
Molecular: Yes.
Dyplolabia A.Massal. (4 spp.). Changed classification (Rivas
Plata et al. 2012a). Molecular: Yes.
Enigmotrema Lücking (1 sp.). New or reinstated (Sipman et al.
2012). Molecular: Yes.
Fissurina Fée (syn.: Leucogymnospora Fink) (160 spp.). New
synonymy (this paper). Molecular: Yes. Notes. The name
Leucogymnospora was found to be based on a species
congeneric with Fissurina.
394 The Bryologist 119(4): 2016
Pycnotrema Rivas Plata & Lücking (3 spp.). New or reinstated
(Rivas Plata et al. 2012a). Molecular: Yes.
Subfamily Graphidoideae Rivas Plata, Lücking &
Lumbsch
Acanthothecis Clem. (56 spp.). Changed classification (Rivas
Plata et al. 2012a). Molecular: Yes.
Acanthotrema Frisch (5 spp.). Changed classification (Rivas
Plata et al. 2012a). Molecular: Yes.
Aggregatorygma M.Cáceres, Aptroot & Lücking (1 sp.). New or
reinstated (Cáceres et al. 2014). Molecular: Yes.
Amazonotrema Kalb & Lücking (1 sp.). Changed classification
(Rivas Plata et al. 2012a). Molecular: No.
Ampliotrema Kalb ex Kalb (18 spp.). New or reinstated (Rivas
Plata et al. 2012b). Molecular: Yes.
Anomalographis Kalb (2 spp.). Changed classification (Rivas
Plata et al. 2012a). Molecular: No.
Anomomorpha Nyl. ex Hue (8 spp.). Changed classification
(Rivas Plata et al. 2012a). Molecular: Yes.
Asteristion Leight. (5 spp.). New or reinstated (Medeiros et al.
2017). Molecular: Yes.
Astrochapsa Parnmen, Lücking & Lumbsch (26 spp.). New or
reinstated (Parnmen et al. 2012). Molecular: Yes.
Austrotrema I.Medeiros, Lücking & Lumbsch (3 spp.). New or
reinstated (Medeiros et al. 2017). Molecular: Yes.
Borinquenotrema Mercado-Dı́az, Lücking & Parnmen (1
sp.). New or reinstated (Lumbsch et al. 2014).
Molecular: Yes.
Byssotrema M.Cáceres, Aptroot & Lücking (1 sp.). New or
reinstated (Cáceres et al. 2014). Molecular: No.
Carbacanthographis Staiger & Kalb (28 spp.). Changed
classification (Rivas Plata et al. 2012a). Molecular: Yes.
Chapsa A.Massal. (62 spp.). Changed classification (Rivas Plata
et al. 2012a). Molecular: Yes.
Chroodiscus (Müll.Arg.) Müll.Arg. (14 spp.). Changed classifi-
cation (Rivas Plata et al. 2012a). Molecular: Yes.
Compositrema Rivas Plata, Lücking & Lumbsch (4 spp.). New
or reinstated (Rivas Plata et al. 2012b). Molecular: Yes.
Corticorygma M.Cáceres, Feuerstein, Aptroot & Lücking (1 sp.).
New or reinstated (Cáceres et al. 2014). Molecular: Yes.
Creographa A.Massal. (30 spp.). New or reinstated (Rivas Plata
et al. 2013). Molecular: Yes.
Crutarndina Parnmen, Lücking & Lumbsch (1 sp.). New or
reinstated (Parnmen et al. 2012). Molecular: Yes.
Diaphorographis A.W.Archer (2 spp.). Changed classification
(Rivas Plata et al. 2012a). Molecular: No.
Diorygma Eschw. (71 spp.). Changed classification (Rivas Plata
et al. 2012a). Molecular: Yes.
Diploschistes Norman (31 spp.). Changed classification (Rivas
Plata et al. 2012a). Molecular: Yes.
Fibrillithecis Frisch (16 spp.). Changed classification (Rivas
Plata et al. 2012a). Molecular: Yes.
Flegographa A.Massal. (1 sp.). New or reinstated (Rivas Plata et
al. 2013). Molecular: Yes.
Gintarasia Kraichak, Lücking & Lumbsch (4 spp.). New or
reinstated (Kraichak et al. 2013). Molecular: Yes.
Glaucotrema Rivas Plata & Lumbsch (4 spp.). New or reinstated
(Rivas Plata et al. 2012b). Molecular: Yes.
Glyphis Ach. (7 spp.). Changed classification (Rivas Plata et al.
2012a). Molecular: Yes.
Graphis Adans. (390 spp.). Changed classification (Rivas Plata et
al. 2012a). Molecular: Yes.
Gyrotrema Frisch (6 spp.). Changed classification (Rivas Plata et
al. 2012a). Molecular: Yes.
Halegrapha Rivas Plata & Lücking (8 spp.). New or reinstated
(Lücking et al. 2011a). Molecular: Yes.
Heiomasia Nelsen, Lücking & Rivas Plata (3 spp.). New or
reinstated (Nelsen et al. 2010). Molecular: Yes.
Hemithecium Trevis. (51 spp.). Changed classification (Rivas
Plata et al. 2012a). Molecular: Yes.
Kalbographa Lücking (5 spp.). Changed classification (Rivas
Plata et al. 2012a). Molecular: No.
Leiorreuma Eschw. (15 spp.). New or reinstated (Staiger 2002).
Molecular: Yes.
Leucodecton A.Massal. (28 spp.). Changed classification (Rivas
Plata et al. 2012a). Molecular: Yes.
Malmographina M.Cáceres, Rivas Plata & Lücking (1 sp.). New
or reinstated (Cáceres et al. 2012). Molecular: Yes.
Mangoldia Lücking, Parnmen & Lumbsch (2 spp.). New or
reinstated (Lücking et al. 2012). Molecular: Yes.
Melanotopelia Lumbsch & Mangold (4 spp.). Changed
classification (Rivas Plata et al. 2012a). Molecular: Yes.
Melanotrema Frisch (18 spp.). Changed classification (Rivas
Plata et al. 2012a). Molecular: Yes.
Myriochapsa M.Cáceres, Lücking & Lumbsch (3 spp.). New or
reinstated (Parnmen et al. 2013). Molecular: Yes.
Myriotrema Fée (syn.: Leptotrema Mont. & Bosch) (54 spp.).
Changed classification and new synonymy (Lücking et al.
2015, 2016d ; Rivas Plata et al. 2012a). Molecular: Yes.
Nadvornikia Tibell (4 spp.). Changed classification (Rivas Plata
et al. 2012a). Molecular: Yes.
Nitidochapsa Parnmen, Lücking & Lumbsch (5 spp.). New or
reinstated (Parnmen et al. 2013). Molecular: Yes.
Ocellularia G.Mey. [nom.cons.] (400 spp.). Changed classifica-
tion (Rivas Plata et al. 2012a). Molecular: Yes.
Pallidogramme Staiger, Kalb & Lücking (12 spp.). Changed
classification (Rivas Plata et al. 2012a). Molecular: Yes.
Paratopeliopsis Mercado-Dı́az, Lücking & Parnmen (1 sp.).
New or reinstated (Lumbsch et al. 2014). Molecular: Yes.
Phaeographis Müll.Arg. (180 spp.). Changed classification
(Rivas Plata et al. 2012a). Molecular: Yes.
Phaeographopsis Sipman (3 spp.). Changed classification (Rivas
Plata et al. 2013). Molecular: Yes.
Platygramme Fée (30 spp.). Changed classification (Rivas Plata
et al. 2012a). Molecular: Yes.
Platythecium Staiger (26 spp.). Changed classification (Rivas
Plata et al. 2012a). Molecular: Yes.
Pliariona A.Massal. (1 sp.). New or reinstated (Rivas Plata et al.
2013). Molecular: Yes.
Polistroma Clemente (1 sp.). Changed classification (Rivas Plata
et al. 2012a). Molecular: No.

Pseudochapsa Parnmen, Lücking & Lumbsch (18 spp.).
New or reinstated (Parnmen et al. 2012). Molecular:
Yes.
Pseudoramonia Kantvilas & Vězda (3 spp.). Changed classifi-
cation (Rivas Plata et al. 2012a). Molecular: Yes.
Pseudotopeliopsis Parnmen, Lücking & Lumbsch (4 spp.).
New or reinstated (Parnmen et al. 2012). Molecular:
Yes.
Redingeria Frisch (12 spp.). Changed classification (Rivas Plata
et al. 2012a). Molecular: Yes.
Reimnitzia Kalb (1 sp.). Changed classification (Rivas Plata et al.
2012a). Molecular: Yes.
Rhabdodiscus Vain. (42 spp.). New or reinstated (Rivas Plata et
al. 2012b). Molecular: Yes.
Sanguinotrema Lücking (1 sp.). New or reinstated (Lücking et
al. 2015). Molecular: Yes.
Sarcographa Fée (37 spp.). Changed classification (Rivas Plata et
al. 2012a). Molecular: Yes.
Sarcographina Müll.Arg. (1 sp.). New or reinstated (Kraichak et
al. 2014). Molecular: Yes.
Schistophoron Stirt. (4 spp.). Changed classification (Rivas Plata
et al. 2012a). Molecular: Yes.
Schizotrema Mangold & Lumbsch (13 spp.). Changed classifi-
cation (Rivas Plata et al. 2012a). Molecular: Yes.
Stegobolus Mont. (15 spp.). Changed classification (Rivas Plata
et al. 2012a). Molecular: Yes.
Thalloloma Trevis. (19 spp.). Changed classification (Rivas Plata
et al. 2012a). Molecular: Yes.
Thecaria Fée (3 spp.). Changed classification (Rivas Plata et al.
2012a). Molecular: Yes.
Thecographa A.Massal. (3 spp.). New or reinstated (Rivas Plata
et al. 2012a). Molecular: No.
Thelotrema Ach. (106 spp.). Changed classification (Rivas Plata
et al. 2012a). Molecular: Yes.
Topeliopsis Kantvilas & Vězda (20 spp.). Changed classification
(Rivas Plata et al. 2012a). Molecular: Yes.
Wirthiotrema Rivas Plata, Kalb, Frisch & Lumbsch (7 spp.).
New or reinstated (Rivas Plata et al. 2010). Molecular:
Yes.
Xalocoa Kraichak, Lücking & Lumbsch (1 sp.). New or
reinstated (Kraichak et al. 2013). Molecular: Yes.
Subfamily Redonographoideae Lücking, Tehler &
Lumbsch
Gymnographopsis C.W.Dodge (2 spp.). Changed classification
(Rivas Plata et al. 2012a). Molecular: No.
Redonographa Lücking, Tehler & Lumbsch (5 spp.). New or
reinstated (Lücking et al. 2013). Molecular: Yes.
Family Gyalectaceae (A.Massal.)Stizenb.
Cryptolechia A.Massal. (11 spp.). No change. Molecular: Yes.
Gyalecta Ach. (syn.: Belonia K örb.; Clathroporinopsis
M.Choisy; Pachyphiale Lönnr.; Protoschistes M.Choisy)
(50 spp.). New synonymy (Baloch et al. 2010, 2013).
Molecular: Yes.
Ramonia Stizenb. (24 spp.). No change. Molecular: Yes.
Lücking et al.: 2016 lichen classification 395
Semigyalecta Vain. (1 sp.). No change. Molecular: No.
Family Phlyctidaceae Poelt & Vězda ex J.C.David &
D.Hawksw.
Phlyctis (Wallr.) Flot. [nom.cons.] (20 spp.). No change.
Molecular: Yes.
Psathyrophlyctis Brusse (1 sp.). No change. Molecular: No.
Family Porinaceae Rchb. (syn.: Myeloconidaceae
P.M.McCarthy)
Clathroporina Müll.Arg. (25 spp.). No change. Molecular: Yes.
Myeloconis P.M.McCarthy & Elix (4 spp.). Changed classifica-
tion (Nelsen et al. 2014b). Molecular: Yes.
Porina Müll.Arg. [nom.cons.] (140 spp.). No change. Molecu-
lar: Yes.
Pseudosagedia (Müll.Arg.) Choisy (80 spp.). New or reinstated
(Brodo et al. 2013; Hafellner & Kalb 2005; Harris 2005).
Molecular: Yes.
Segestria Fr. (syn.: Polycornum Malcolm & Vězda) (70 spp.).
New synonymy (Lücking 2008). Molecular: Yes.
Trichothelium Müll.Arg. (40 spp.). No change. Molecular: Yes.
Family Protothelenellaceae Vězda, H.Mayrhofer &
Poelt
Protothelenella Räsänen (11 spp.). Changed classification (Resl
et al. 2015). Molecular: Yes.
Family Sagiolechiaceae Baloch, Lücking, Lumbsch &
Wedin
Rhexophiale Th.Fr. (1 sp.). New or reinstated (Baloch et al.
2010). Molecular: Yes.
Sagiolechia A.Massal. (3 spp.). Changed classification (Baloch et
al. 2010). Molecular: Yes.
Family Stictidaceae Fr.
Absconditella Vězda (12 spp.). No change. Molecular: Yes.
Acarosporina Sherwood (5 spp.). No change. Molecular: Yes.
Conotremopsis Vězda (1 sp.). No change. Molecular: No.
Cryptodiscus Corda (syn.: Bryophagus Nitschke ex Arnold) (9
spp.). New synonymy (Baloch et al. 2009). Molecular: Yes.
Geisleria Nitschke (1 sp.). New or reinstated (Aptroot et al.
2015). Molecular: Yes.
Ingvariella Guderley & Lumbsch (1 sp.). Changed classification
(Fernández-Brime et al. 2011). Molecular: Yes.
Robergea Desm. (1 sp.). No change. Molecular: Yes.
Schizoxylon Pers. (1 sp.). No change. Molecular: Yes.
Stictis Pers. (4 spp.). No change. Molecular: Yes.
Thelopsis Nyl. [nom.cons.] (9 spp.). No change. Molecular: No.
Topelia P.M.Jørg. & Vězda (6 spp.). No change. Molecular: No.
Trinathotrema Lücking, Rivas Plata & Mangold (3 spp.). New
or reinstated (Lücking et al. 2011b). Molecular: Yes.
396 The Bryologist 119(4): 2016
Xyloschistes Vain. ex Zahlbr. (1 sp.). Changed classification
(Mia˛dlikowska et al. 2014). Molecular: Yes.
Family Thelenellaceae O.E.Erikss. ex H.Mayrhofer
(syn.: Aspidotheliaceae Räsänen ex J.C. David &
D.Hawksw.)
Aspidothelium Vain. (17 spp.). Changed classification (Nelsen et
al. 2016). Molecular: Yes.
Chromatochlamys Trevis. (3 spp.). Changed classification
(Nelsen et al. 2016). Molecular: Yes.
Thelenella Nyl. (30 spp.). Changed classification (Nelsen et al.
2016). Molecular: Yes.
Family Thrombiaceae Poelt & Vězda ex J.C.David &
D.Hawksw.
Thrombium Wallr. (5 spp.). Changed classification (Resl et al.
2015). Molecular: Yes.
Order Ostropales genera incertae sedis
Amphorothecium P.M.McCarthy, Kantvilas & Elix (1 sp.). No
change. Molecular: No.
Anzina Scheid. (1 sp.). Changed classification (Resl et al. 2015).
Molecular: Yes.
Order PERTUSARIALES M.Choisy ex D.Hawksw. &
O.E.Erikss.
Family Coccotremataceae Henssen ex J.C.David &
D.Hawksw.
Coccotrema Müll.Arg. (16 spp.). No change. Molecular: Yes.
Gyalectaria Lumbsch, Kalb & Schmitt (3 spp.). New or
reinstated (Schmitt et al. 2010). Molecular: Yes.
Parasiphula Kantvilas & Grube (7 spp.). No change. Molecular: No.
Family Icmadophilaceae Triebel
Chirleja Lendemer & B.P.Hodk. (1 sp.). New or reinstated
(Lendemer & Hodkinson 2012a). Molecular: Yes.
Dibaeis Clem. (13 spp.). No change. Molecular: Yes.
Endocena Cromb. (1 sp.). No change. Molecular: Yes.
Icmadophila Trevis. [nom.cons.] (8 spp.). No change. Molec-
ular: Yes.
Pseudobaeomyces M.Satı̀ (1 sp.). No change. Molecular: No.
Siphula Fr. [nom.cons.] (26 spp.). No change. Molecular: Yes.
Siphulella Kantvilas, Elix & P.James (1 sp.). No change.
Molecular: Yes.
Thamnolia Ach. ex Schaerer [nom.cons.] (4 spp.). No change.
Molecular: Yes.
Family Megasporaceae Lumbsch, Feige & K.Schmitz
Aspicilia A.Massal. [nom.cons.] (200 spp.). No change.
Molecular: Yes.
Circinaria Link (25 spp.). New or reinstated (Nordin et al. 2010;
Sohrabi et al. 2013a). Molecular: Yes.
Lobothallia (Clauzade & Cl.Roux) Hafellner (9 spp.). No
change. Molecular: Yes.
Megaspora (Clauzade & Cl.Roux) Hafellner & V.Wirth (3 spp.).
No change. Molecular: Yes.
Sagedia Ach. (3 spp.). New or reinstated (Nordin et al. 2010).
Molecular: Yes.
Teuvoa Sohrabi & S.Leavitt (3 spp.). New or reinstated (Sohrabi
et al. 2013b). Molecular: Yes.
Family Miltideaceae Hafellner
Miltidea Stirt. (1 sp.). Changed classification (Widhelm &
Lumbsch 2011). Molecular: Yes.
Family Ochrolechiaceae R.C.Harris ex Lumbsch &
I.Schmitt
Marfloraea S.Y.Kondr., L.L}
okös & Hur (syn.: Variolaria Pers.
[nom. illeg.]) (13 spp.). New or reinstated and new
synonymy (Kondratyuk et al. 2015c). Molecular: Yes.
Ochrolechia A.Massal. (60 spp.). No change. Molecular: Yes.
Varicellaria Nyl. (7 spp.). No change. Molecular: Yes.
Family Pertusariaceae Körb. ex Körb.
Loxosporopsis Brodo, Henssen & Imshaug (1 sp.). No change.
Molecular: Yes.
Pertusaria DC. [nom.cons.] (500 spp.). No change. Molecular: Yes.
Thamnochrolechia Aptroot & Sipman (1 sp.). No change.
Molecular: Yes.
Order SARRAMEANALES B.P.Hodk. & Lendemer
Family Sarrameanaceae Hafellner
Loxospora A.Massal. (13 spp.). Changed classification (Hodkin-
son & Lendemer 2011). Molecular: Yes.
Sarrameana Vězda & P.James (1 sp.). Changed classification
(Hodkinson & Lendemer 2011). Molecular: No.
Family Schaereriaceae Hafellner
Schaereria Th.Fr. [nom.cons.] (16 spp.). Changed classification
(Hodkinson & Lendemer 2011). Molecular: Yes.
Order TRAPELIALES B.P.Hodk. & Lendemer
Family Trapeliaceae M.Choisy ex Hertel [nom.cons.]
Amylora Rambold (1 sp.). Changed classification (Hodkinson &
Lendemer 2011). Molecular: No.
Aspiciliopsis (Müll.Arg.) M.Choisy (1 sp.). Changed classifica-
tion (Hodkinson & Lendemer 2011). Molecular: No.
Coppinsia Lumbsch & Heibel (1 sp.). Changed classification
(Hodkinson & Lendemer 2011). Molecular: No.
Lignoscripta B.D.Ryan (1 sp.). Changed classification (Hodkin-
son & Lendemer 2011). Molecular: No.

Orceolina Hertel (2 spp.). Changed classification (Hodkinson &
Lendemer 2011). Molecular: Yes.
Placopsis (Nyl.) Linds. (60 spp.). Changed classification
(Hodkinson & Lendemer 2011). Molecular: Yes.
Placynthiella Elenkin (7 spp.). Changed classification (Hodkin-
son & Lendemer 2011). Molecular: Yes.
Rimularia Nyl. (4 spp.). Changed classification (Hodkinson &
Lendemer 2011). Molecular: Yes.
Sarea Fr. (2 spp.). Changed classification (Hodkinson &
Lendemer 2011). Molecular: Yes.
Trapelia M.Choisy [nom.cons.] (13 spp.). Changed classifica-
tion (Hodkinson & Lendemer 2011). Molecular: Yes.
Trapeliopsis Hertel & Gotth.Schneid. (20 spp.). Changed
classification (Hodkinson & Lendemer 2011). Molecular:
Yes.
Family Xylographaceae Tuck. (syn.: Lithographaceae
Poelt [nom. inval.])
Lambiella Hertel (12 spp.). New or reinstated (Resl et al. 2015;
Spribille et al. 2014). Molecular: Yes.
Lithographa Nyl. (10 spp.). Changed classification (Resl et al.
2015). Molecular: Yes.
Ptychographa Nyl. (1 sp.). Changed classification (Resl et al.
2015). Molecular: Yes.
Xylographa (Fr.) Fr. (20 spp.). Changed classification (Resl et al.
2015). Molecular: Yes.
Subclass Ostropomycetidae genera incertae sedis
Aspilidea Hafellner (1 sp.). No change. Molecular: No.
Dictyocatenulata Finley & E.F.Morris (1 sp.). New or reinstated
(Lendemer & Harris 2004; Diederich et al. 2008; An et al.
2012). Molecular: Yes.
Subclass UMBILICARIOMYCETIDAE Bendiksby,
Hestmark & Timdal
Order UMBILICARIALES Lumbsch, Hestmark & Lutzoni
Family Elixiaceae Lumbsch
Elixia Lumbsch (2 spp.). Changed classification (Bendiksby &
Timdal 2013; Mia˛dlikowska et al. 2014). Molecular: Yes.
Meridianelia Kantvilas & Lumbsch (1 sp.). Changed classifica-
tion (Bendiksby & Timdal 2013; Mia˛dlikowska et al. 2014).
Molecular: Yes.
Family Fuscideaceae Hafellner (syn.: Orphniospor-
aceae Hafellner)
Fuscidea V.Wirth & Vězda (35 spp.). Changed classification
(Bendiksby & Timdal 2013; Mia˛dlikowska et al. 2014).
Molecular: Yes.
Hueidea Kantvilas & P.M.McCarthy (1 sp.). New or reinstated
(Kantvilas & McCarthy 2003). Molecular: No.
Lücking et al.: 2016 lichen classification 397
Maronea A.Massal. (13 spp.). Changed classification (Bendiksby
& Timdal 2013; Mia˛dlikowska et al. 2014). Molecular: Yes.
Orphniospora Körb. (4 spp.). Changed classification (Bendiksby
& Timdal 2013; Mia˛dlikowska et al. 2014). Molecular: No.
Family Ophioparmaceae R.W.Rogers & Hafellner
(syn.: Rhizoplacopsidaceae J.C.Wei & Q.M.
Zhou)
Boreoplaca Timdal (1 sp.). Changed classification (Bendiksby &
Timdal 2013; Mia˛dlikowska et al. 2014). Molecular: Yes.
Hypocenomyce M.Choisy (3 spp.). Changed classification
(Bendiksby & Timdal 2013; Mia˛dlikowska et al. 2014).
Molecular: Yes.
Ophioparma Norman (8 spp.). Changed classification (Bend-
iksby & Timdal 2013; Mia˛dlikowska et al. 2014). Molecular:
Yes.
Family Ropalosporaceae Hafellner
Ropalospora A.Massal. (9 spp.). Changed classification (Bend-
iksby & Timdal 2013; Mia˛dlikowska et al. 2014). Molecular:
Yes.
Family Umbilicariaceae Chevall.
Fulgidea Bendiksby & Timdal (2 spp.). New or reinstated
(Bendiksby & Timdal 2013). Molecular: Yes.
Lasallia Mérat (17 spp.). Changed classification (Bendiksby &
Timdal 2013). Molecular: Yes.
Umbilicaria Hoffm. [nom.cons.] (70 spp.). Changed classifica-
tion (Bendiksby & Timdal 2013). Molecular: Yes.
Xylopsora Bendiksby & Timdal (2 spp.). New or reinstated
(Bendiksby & Timdal 2013). Molecular: No.
Class Lecanoromycetes families incertae sedis
Family Arthrorhaphidaceae Poelt & Hafellner
Arthrorhaphis Th.Fr. [nom.cons.] (13 spp.). Changed classifi-
cation (Mia˛dlikowska et al. 2014). Molecular: Yes.
Class Lecanoromycetes genera incertae sedis
Argopsis Th.Fr. (3 spp.). Changed classification (this paper).
Molecular: No. Notes. Argopsis was long placed with
questionmark in Brigantiaeaceae, based on Hafellner
(1984), but in our view is unrelated to that genus and its
placement must be considered unresolved.
Bartlettiella D.J.Galloway & P.M.Jørg. (1 sp.). No change.
Molecular: No.
Bouvetiella Øvstedal (1 sp.). No change. Molecular: No.
Buelliastrum Zahlbr. (2 spp.). No change. Molecular: No.
Haploloma Trevis. (1 sp.). No change. Molecular: No.
Maronella M.Steiger (1 sp.). No change. Molecular: No.
Notolecidea Hertel (1 sp.). No change. Molecular: Yes.
Petractis Fr. (3 spp.). Changed classification (this paper).
Molecular: Yes. Notes. The genus was listed in Stictidaceae
398 The Bryologist 119(4): 2016
(Lumbsch & Huhndorf 2010) but suggested to form part of
Phlyctidaceae by Mia˛dlikowska et al. (2014). However, the
two available sequences give conflicting results, with no
close relative within the Lecanoromycetes. This taxon was
therefore reclassified and needs further study.
Piccolia A.Massal. (7 spp.). No change. Molecular: No.
Class LICHINOMYCETES Reeb, Lutzoni & Cl.Roux
Order LICHINALES Henssen & Büdel
Family Gloeoheppiaceae Henssen
Gloeoheppia Gyeln. (5 spp.). No change. Molecular: No.
Gudelia Henssen (1 sp.). No change. Molecular: No.
Pseudopeltula Henssen (4 spp.). No change. Molecular: No.
Family Lichinaceae Nyl. (syn.: Heppiaceae Zahlbr.)
Anema Nyl. (5 spp.). No change. Molecular: Yes.
Calotrichopsis Vain. (3 spp.). No change. Molecular: No.
Corynecystis Brusse (1 sp.). New or reinstated (Tehler & Wedin
2008). Molecular: No.
Cryptothele Th.Fr. (6 spp.). No change. Molecular: No.
Digitothyrea P.Moreno & Egea (3 spp.). No change. Molecular:
Yes.
Edwardiella Henssen (1 sp.). No change. Molecular: No.
Ephebe Fr. (13 spp.). No change. Molecular: Yes.
Euopsis Nyl. (2 spp.). No change. Molecular: Yes.
Finkia Vain. (1 sp.). No change. Molecular: No.
Gyrocollema Vain. (2 spp.). No change. Molecular: No.
Harpidium Körb. (3 spp.). No change. Molecular: Yes.
Heppia Nägeli (7 spp.). Changed classification (Schultz & Büdel
2003). Molecular: Yes.
Jenmania W.Wächt. (2 spp.). No change. Molecular: Yes.
Lecidopyrenopsis Vain. (1 sp.). No change. Molecular: No.
Lemmopsis (Vain.) Zahlbr. (3 spp.). No change. Molecular: No.
Lempholemma Körb. (35 spp.). No change. Molecular: Yes.
Leprocollema Vain. (3 spp.). No change. Molecular: No.
Lichina C.Agardh [nom.cons.] (9 spp.). No change. Molecular:
Yes.
Lichinella Nyl. (syn.: Gonohymenia J.Steiner) (30 spp.). New
synonymy (Moreno & Egea 1992). Molecular: Yes.
Lichinodium Nyl. (4 spp.). No change. Molecular: No.
Mawsonia C.W.Dodge (1 sp.). No change. Molecular: No.
Metamelanea Henssen (3 spp.). No change. Molecular: No.
Paulia Feé (14 spp.). No change. Molecular: Yes.
Peccania A.Massal. ex Arnold [nom.cons.] (15 spp.). No change.
Molecular: Yes.
Phloeopeccania J.Steiner (4 spp.). No change. Molecular: Yes.
Phylliscidium Forssell (1 sp.). No change. Molecular: No.
Phyllisciella Henssen & Büdel (3 spp.). No change. Molecular:
No.
Phylliscidiopsis Sambo (1 sp.). No change. Molecular: No.
Phylliscum Nyl. (8 spp.). No change. Molecular: Yes.
Porocyphus Körb. (8 spp.). No change. Molecular: Yes.
Pseudarctomia Gyeln. (1 sp.). No change. Molecular: No.
Pseudoheppia Zahlbr. (1 sp.). Changed classification (Schultz &
Büdel 2003). Molecular: No.
Pseudopaulia M.Schultz (1 sp.). No change. Molecular: No.
Psorotichia A.Massal. (50 spp.). No change. Molecular: Yes.
Pterygiopsis Vain. (17 spp.). No change. Molecular: Yes.
Pyrenocarpon Trevis. (1 sp.). No change. Molecular: No.
Pyrenopsis Nyl. [nom.cons.] (40 spp.). No change. Molecular: Yes.
Solorinaria (Vain.) Gyeln. (1 sp.). Changed classification
(Schultz & Büdel 2003). Molecular: No.
Stromatella Henssen (1 sp.). No change. Molecular: Yes.
Synalissa Fr. (5 spp.). No change. Molecular: No.
ThallinocarponÅ.E.Dahl (2 spp.). New or reinstated (Jørgensen
2007). Molecular: No.
Thelignya A.Massal. (2 spp.). No change. Molecular: No.
Thermutis Fr. (2 spp.). No change. Molecular: No.
Thermutopsis Henssen (1 sp.). No change. Molecular: No.
Thyrea A.Massal. (13 spp.). No change. Molecular: Yes.
Zahlbrucknerella Herre (10 spp.). No change. Molecular: No.
Family Peltulaceae Büdel
Peltula Nyl. (syn.: Neoheppia Zahlbr.; Phyllopeltula Kalb) (40
spp.). New synonymy (Rauhut 2007). Molecular: Yes.
Class SORDARIOMYCETES O.E.Erikss. & Winka
Subclass XYLARIOMYCETIDAE O.E.Erikss. & Winka
Order XYLARIALES Nannf.
Family Requienellaceae Boise
Requienella Fabre (1 sp.). Changed classification (Jaklitsch et al.
2016b). Molecular: Yes.
Subphylum Pezizomycotina orders incertae sedis
Order THELOCARPALES Lücking & Lumbsch
Family Thelocarpaceae Zukal
Sarcosagium A.Massal. (1 sp.). Changed classification (Jaklitsch
et al. 2016a). Molecular: Yes.
Thelocarpon Nyl. (25 spp.). Changed classification (Jaklitsch et
al. 2016a). Molecular: Yes.
Order VEZDAEALES Lumbsch & Lücking
Family Vezdaeaceae Poelt & Vězda ex J.C.David &
D.Hawksw.
Vezdaea Tsch.-Woess & Poelt (13 spp.). Changed classification
(Jaklitsch et al. 2016a). Molecular: No.

Subphylum Pezizomycotina families incertae sedis
Family Aphanopsidaceae Printzen & Rambold
Aphanopsis Nyl. ex Sydow (2 spp.). Changed classification
(Printzen et al. 2012). Molecular: Yes.
Steinia Körb. (3 spp.). New or reinstated (Printzen et al. 2012).
Molecular: Yes.
Family Strangosporaceae S.Stenroos, Mia˛dl. & Lut-
zoni
Strangospora Körb. (10 spp.). Changed classification (Mia˛dli-
kowska et al. 2014). Molecular: Yes.
Subphylum Pezizomycotina genera incertae sedis
Asteroporum Müll.Arg. (7 spp.). Changed classification (this
paper). Molecular: No. Notes. No molecular data are
available for this little known genus and it therefore has
been moved from Dothideomycetes to Pezizomycotina
incertae sedis.
Biatoridium J.Lahm ex Körb. (3 spp.). Changed classification
(Reeb et al. 2004). Molecular: Yes.
Cheiromycina B.Sutton (4 spp.). New or reinstated (Aptroot &
Schiefelbein 2003; Printzen 2007; Diederich et al. 2008; An
et al. 2012). Molecular: No.
Cyanoporina Groenh. (1 sp.). Changed classification (this
paper). Molecular: No. Notes. No molecular data are
available for this little known genus and it therefore has
been moved from Dothideomycetes to Pezizomycotina
incertae sedis.
Eopyrenula R.C.Harris (6 spp.). Changed classification (this
paper). Molecular: No. Notes. No molecular data are
available for this little known genus and it therefore has
been moved from Dothideomycetes to Pezizomycotina
incertae sedis.
Frigidopyrenia Grube (1 sp.). Changed classification (this
paper). Molecular: No. Notes. No molecular data are
available for this little known genus and it therefore has
been moved from Dothideomycetes to Pezizomycotina
incertae sedis.
Magmopsis Nyl. (1 sp.). New or reinstated (Grube 2005).
Molecular: No.
Melanophloea P.James & Vězda (1 sp.). Changed classification (this
paper). Molecular: No. Notes. No molecular data are available
for this little known genus and it therefore has been moved
from Dothideomycetes to Pezizomycotina incertae sedis.
Milospium D.Hawksw. (3 spp.). New or reinstated (Aptroot &
Sipman 2001; Crous et al. 2015). Molecular: Yes.
Oevstedalia Ertz & Diederich (1 sp.). Changed classification
(this paper). Molecular: No. Notes. No molecular data are
available for this little known genus and it therefore has
been moved from Dothideomycetes to Pezizomycotina
incertae sedis.
Psammina Sacc. & M.Rousseau ex E.Bommer & M.Rousseau (1
sp.). New or reinstated (Earland-Bennett & Hawksworth
2005). Molecular: No.
Pyrenocollema Reinke (19 spp.). Changed classification (Pérez-
Ortega et al. 2016). Molecular: Yes.
Lücking et al.: 2016 lichen classification 399
Thelenidia Nyl. (1 sp.). Changed classification (this paper).
Molecular: No. Notes. No molecular data are available for
this little known genus and it therefore has been moved
from Dothideomycetes to Pezizomycotina incertae sedis.
Trizodia Laukka (1 sp.). New or reinstated (Stenroos et al.
2010). Molecular: Yes.
Wadeana Coppins & P.James (2 spp.). Changed classification
(this paper). Molecular: No. Notes. The genus was listed
under Ascomycota incertae sedis, but since all lichenized
forms are found in subphylum Pezizomycotina, the listing
was changed.
Phylum BASIDIOMYCOTA R.T.Moore
Subphylum AGARICOMYCOTINA Doweld
Class AGARICOMYCETES Doweld
Subclass AGARICOMYCETIDAE Loqu. ex Parmasto
Order AGARICALES Underw.
Family Hygrophoraceae Lotsy
Subfamily Lichenomphalioideae Lücking & Redhead
Acantholichen P.M.Jørg. (6 spp.). Changed classification (Law-
rey et al. 2009; Lodge et al. 2014). Molecular: Yes.
Arrhenia Fr. (1 sp.). Changed classification (Lawrey et al. 2009;
Lodge et al. 2014). Molecular: Yes.
Cora Fr. (92 spp.). Reinstated genus (Dal Forno et al. 2013;
Lawrey et al. 2009; Lodge et al. 2014; Lücking et al. 2016b).
Molecular: Yes.
Corella Vain. (4 spp.). Reinstated genus (Dal Forno et al. 2013;
Lawrey et al. 2009; Lodge et al. 2014). Molecular: Yes.
Cyphellostereum D.A.Reid (5 spp.). Changed classification
(Lawrey et al. 2009; Lodge et al. 2014). Molecular: Yes.
Dictyonema C.Agardh ex Kunth (22 spp.). Changed classifica-
tion (Lawrey et al. 2009; Lodge et al. 2014). Molecular: Yes.
Lichenomphalia Redhead, Lutzoni, Moncalvo & Vilgalys (16
spp.). Changed classification (Lodge et al. 2014). Molecular:
Yes.
Semiomphalina Redhead (1 sp.). Changed classification (Lodge
et al. 2014). Molecular: Yes.
Order ATHELIALES Jülich
Family Atheliaceae Jülich
Athelia Pers. (1 sp.). No change. Molecular: Yes.
Athelopsis Oberw. ex Parmasto (1 sp.). No change. Molecular:
Yes.
Order LEPIDOSTROMATALES B.P.Hodk. & Lücking
Family Lepidostromataceae Ertz, Eb.Fisch., Kill-
mann, Sérus. & Lawrey
Ertzia B.P.Hodk. & Lücking (1 sp.). New or reinstated
(Hodkinson et al. 2014). Molecular: Yes.
400 The Bryologist 119(4): 2016
Lepidostroma Mägd. & S.Winkl. (3 spp.). Changed classification
(Ertz et al. 2008; Hodkinson et al. 2014). Molecular: Yes.
Sulzbacheromyces B.P.Hodk. & Lücking (2 spp.). New or
reinstated (Hodkinson et al. 2014). Molecular: Yes.
Class Agaricomycetes orders incertae sedis
Order CANTHARELLALES Gäum.
Family Clavulinaceae Donk
Multiclavula R.H.Petersen (16 spp.). No change. Molecular:
Yes.
Order CORTICIALES K.-H.Larsson
Family Corticiaceae Herter [nom.cons.]
Marchandiomphalina Diederich, Manfr.Binder & Lawrey (1
sp.). No change. Molecular: Yes.
APPENDIX TO THE CLASSIFICATION: HYPERLICHENIZED
FUNGI
Phylum BASIDIOMYCOTA R.T.Moore
Subphylum PUCCINIOMYCOTINA Bauer, Beger-
ow, J.P.Samp., M.Weiß & Oberw.
Order CYPHOBASIDIALES T.Sprib. & H.Mayrhofer
Family Cyphobasidiaceae T.Sprib. & H.Mayrhofer
Cyphobasidium Millanes, Diederich & Wedin (2 spp.). New or
reinstated (Millanes et al. 2016; Spribille et al. 2016).
Molecular: Yes. Notes. The genus was not effectively (and
validly) published at the time of publication of the family
and order name (Spribille et al. 2016), since the original
publication (Millanes et al. 2016) at that time and until 11
October 2016 was listed as ‘‘Article in press’’ (ICN. Art.
30.2), with a posting date of 31 December 2015. While the
genus and species names are valid with the subsequent
effective publication of the final version of that article on 11
October 2016 (Millanes et al. 2016), the order and family
names were subsequently validated (Spribille & Mayrhofer
2016). Unfortunately, the online publication date of the
final article (Millanes et al. 2016) is still given as 31
December 2015, which creates a confusing situation and is
in violation of the Code, since the journal erroneously states
that the ‘‘Article in press’’ version was the final version.
APPENDIX TO THE CLASSIFICATION: FOSSIL LICHEN FUNGI
Chlorolichenomycites Honegger, D.Edwards & L.Axe (1 sp.).
415 mya, Early Devonian (Honegger et al. 2013a, b).
Cyanolichenomycites Honegger, D.Edwards & L.Axe (1 sp.). 415
mya, Early Devonian (Honegger et al. 2013a).
Daohugouthallus X.Wang, M.Krings & T.N.Taylor (1 sp.). 168–
152 mya, Middle Jurassic (Wang et al. 2010).
Flabellitha Krassilov & Jurina (1 sp.). 388–383 mya, Middle
Devonian (Jurina & Krassilov 2002).
Honeggeriella Matsunaga, Stockey & Tomescu [nom. inval.;
ICN Art. 42.1] (1 sp.). 133 mya, Early Cretaceous
(Matsunaga et al. 2013).
Pelicothallos Dilcher (1 sp.). 40 mya, Eocene (Dilcher 1965;
Reynolds & Dilcher 1984; Sherwood-Pike 1985). Notes.
This fossil was first reinterpreted as non-lichenized
Cephaleuros (Reynolds & Dilcher 1984) and then as
lichenized (in an early stage) Strigula (Sherwood-Pike
1985). If restudy of the fossil supports this view, it would
have to be listed as synonym of the lichen fungus Strigula,
not as synonym of the photobiont genus Cephaleuros, even
if the characters in the protologue mostly refer to the algal
features, since Pelicothallus was originally described as
ascomycete and then ICN Art. 13.2. comes into play: ‘‘The
group to which a name is assigned for the purposes of this
Article is determined by the accepted taxonomic position of
the type of the name."
Spongiophyton Kräusel (1 sp.). 409–386 mya, Early Devonian
(Jahren et al. 2003; Matsumara et al. 2016; Taylor et al.
2004b). Notes. Spongiophyton minutissimum Kräusel was
interpreted as a lichen based on physiological (isotope data)
and morphological evidence (Jahren et al. 2003; Matsumara
et al. 2016; Taylor et al. 2004b), but according to a
reevaluation by Fletcher et al. (2004), the isotope data do
not provide conclusive evidence about the placement of this
fossil.
Thuchomyces Hallbauer & Jahns (1 sp.). 2,500 mya, Precam-
brian Witwatersrand (Hallbauer et al. 1977). Notes. This is
by far the oldest fossil morphologically and anatomically
intepreted as a lichen, but no evidence exists for the
presence of a photobiont. The age of the material makes its
interpretation as a lichen very unlikely, since it antedates
the estimated origin of the Eukaryota. Some workers
intepret these structures as abiotic (Cloud 1976).
Winfrenatia T.N.Taylor, Hass & Kerp (1 sp.). 415 mya, Early
Devonian, Rhynie chert (Karatygin et al. 2009, 2010; Taylor
et al. 1995, 1997, 2004a).
NOMENCLATURAL NOTES
Candelariomycetidae Mia˛dl. et al. ex Timdal &
M.Westb. subcl. nov. [MYCOBANK 819260]. Cande-
lariomycetidae Mia˛dl. et al., Mycologia 98: 1091.
2006; nom. inval. [ICN Art. 32.1(d)]
A new subclass within class Lecanoromycetes
containing the single order Candelariales with the
two families Candelariaceae and Pycnoraceae. Thal-
lus crustose to squamulose, peltate-subumbilicate, or
microfoliose, rarely lichenicolous and without thal-
lus, often bright yellow; photobiont chlorococcoid;
cephalodia absent. Ascomata apotheciate, lecanorine
or rarely biatorine (Candelariaceae), or lecideine
(Pycnoraceae). Hamathecium consisting of un-
branched to slightly branched paraphyses, amyloid.
Asci with apical tholus, clavate; outer wall amyloid,
tholus weakly amyloid except for darker amyloid,
ring-shaped structure in the lower part. Ascospores 8

to many (64) per ascus (Candelariaceae), non-
septate to (indistinctly) 1–3-septate, ellipsoid to
citriform, hyaline, non-amyloid. Conidiomata pyc-
nidia; conidia non-septate, ellipsoid to bacillar,
sometimes curved, hyaline. Secondary chemistry:
pulvinic acid derivates (Candelariaceae) or depsides
(alectorialic acid; Pycnoraceae). On rock (many
Candelariaceae), rarely on bryophytes or soil or
epiphytic on bark, or typically on wood (Pycnor-
aceae). HOLOTYPE order: Candelariales Mia˛dl., Lut-
zoni & Lumbsch. HOLOTYPE genus: Candelaria
A.Massal.
Diagnostic characters of this new subclass are the
thallus containing green algae but lacking cepha-
lodia, the clavate, octo- to polysporous asci with a
broad, amyloid ring structure in the lower part of the
tholus (Candelaria-type), and the hyaline, simple or
1-septate ascospores. Except for the Candelaria-type
ascus (see Fig. 4 in Westberg 2005), there are no
known morphological characters separating this
subclass from Lecanoromycetidae and the subclass
is based on molecular phylogenetic analysis by
Bendiksby & Timdal (2013) and Mia˛dlikowska et
al. (2014).
Clathroporinopsis M.Choisy
LECTOTYPE: Clathroporina calcarea Walt.Watson ¼
Gyalecta nidarosiensis (Kindt) Baloch & Lücking;
here designated.
This name was introduced by Choisy (1929), in a
handwritten scheme, with a question mark, without
listing a type and with the only description ‘‘in
Clathroporina inmixtis, dictyospores.’’ The genus
name was later listed as synonym of Clathroporina
(McCarthy 1995) or suggested to possibly represent
Topelia (Hafellner & Kalb 1995; Lumbsch &
Huhndort 2010). There is no way to elucidate the
exact meaning of this name from the protologue,
and in spite of the unusual form of publication, the
name is valid. Handwritten publication (‘‘indelible
autograph’’) is permitted before 1953 [ICN Art.
30.4], the question mark does not render the name
invalid [ICN Art. 36.1, Ex. 10], and the description
in French (‘‘dictyospores’’) was permitted before
1936 [ICN Art. 39.1]. The phrase ‘‘in Clathroporina
inmixtis’’ dictates that a lectotype is to be selected
among species at the time of publication of
Clathroporinopsis listed in Clathroporina but, accord-
ing to Choisy (1929), not belonging there. There are
37 names combined in Clathroporina before 1929
Lücking et al.: 2016 lichen classification 401
and of these, 22 belong to Porina s.l. including
Clathroporina, which leaves 15 names providing a
potential lectotype for Clathroporinopsis. Since
according to the scheme presented by Choisy
(1929), the ascomata should be urceolate to
perithecioid but with an apothecioid (coherent)
hymenium, an interpretation also followed by
Hafellner & Kalb (1995), all names referring to taxa
with genuine perithecia (loose hymenium) need also
to be disregarded, including synonyms of Astrothe-
lium, Polymeridium and Thelenella. Coincidentally,
the only name referring to a species now included in
Topelia, Clathroporina heterospora Zahlbr., is also
based on a specimen producing genuine perithecia,
which probably does not belong in Topelia s.str. This
leaves but a single name available as suitable
lectotype for Clathroporinopsis, namely Clathropor-
ina calcarea, which is a synonym of Gyalecta
nidarosiensis, until recently known as Belonia
nidarosiensis (Kindt) P.M.Jørg. & Vězda. This
lectotypification makes Clathroporinopsis a synonym
of Gyalecta, which not only fits Choisy’s scheme well
but also avoids any nomenclatural disruption.
Cystocoleaceae Locq. ex Lücking, B.P.Hodk. &
S.D.Leav., fam. nov. [MYCOBANK 819262] Cystoco-
leaceae Locq., Mycol. Gén. Struct. (Paris): 213
(1984), nom. inval. [ICN Art. 39.1].
A new family within order Capnodiales. Liche-
nized; thallus filamentose; photobiont Trentepohlia.
Hyphal sheath around the algal filaments composed
of somewhat contorted, jigsaw puzzle-shaped cells
forming a closed layer. Ascomata and conidiomata
unknown. Secondary chemistry: no substances. On
rocks or bark near water. HOLOTYPE genus: Cystoco-
leus A.Massal.
Letrouitineae Gaya & Lutzoni, subordo nov. [MYCO-
BANK 819265]
A new suborder within order Teloschistales
containing the two families Brigantiaeaceae and
Letrouitiaceae. Lichenized; thallus crustose; photo-
biont chlorococcoid. Ascomata apotheciate, bia-
torine, rarely lecideine or lecanorine, usually
yellow-orange to purple-brown. Hamathecium
consisting of mostly unbranched paraphyses, am-
yloid. Asci semifissitunicate, with amyloid tholus
and wall layers but no further apical structure
discernable except small ocular chamber in Le-
trouitiaceae, clavate. Ascospores 1–8 per ascus,
402 The Bryologist 119(4): 2016
transversely septate to muriform, oblong-ellipsoid,
euseptate with rectangular lumina (Brigantiaea-
ceae) or distoseptate with rounded to rectangular
lumina (Letrouitiaceae), hyaline, non-amyloid.
Conidiomata pycnidia; conidia non-septate, bacil-
lar to oblong, hyaline. Secondary chemistry:
variable, with depsides (atranorin), dibenzofuranes
(usnic acid), terpenes (zeorin), and anthraquinones
(usually in the ascomata). Usually epiphytic on
bark. HOLOTYPE family: Letrouitiaceae Bellem. &
Hafellner. HOLOTYPE genus: Letrouitia Hafellner &
Bellem.
Protoschistes M.Choisy
LECTOTYPE: Porina herculana (Rehm) Suza ¼ Gyalecta
herculana (Rehm) Baloch, Lumbsch & Wedin;
here designated.
Like Clathroporinopsis (see above), this name
was introduced by Choisy (1929), in a handwritten
scheme, without listing a type and with the only
description ‘‘in Porina inmixtis, hyalophragmo-
spores a loges cylindriques, paraphyses simples,
libres.’’ MycoBank lists Protoschistes [MYCOBANK
4393] as synonym of Diploschistes, but this is in
clear conflict with the protologue, since Diplo-
schistes has brown, muriform ascospores, not
hyaline and transseptate. Following the logic used
for lectotypification of Clathroporinopsis (see
above), Protoschistes needs to be lectotypified with
a species that is in accordance with the protologue,
i.e. having been listed in Porina at the time of
publication but not belonging in that genus, having
hyaline, transseptate ascospores, and not causing
any nomenclatural disruption. The logical choice is
Porina herculana, now known as Gyalecta herculana
and until recently as Belonia herculana (Rehm)
Keissl., which then makes Protoschistes another
synonym of Gyalecta Ach. This also fits Choisy’s
concept well, since according to this concept,
Protoschistes and Clathroporinopsis form an evolu-
tionary series of shared morphology but trans-
septate versus muriform ascospores, parallel to
Porina versus Clathroporina. In addition, neither
Porina herculana nor Clathroporina calcarea were
recognized as part of Belonia, a genus known to
Choisy at the time.
Rhizocarpales Mia˛dl. et al. ex Mia˛dl. & Lutzoni, ordo
nov. [MYCOBANK 819267] Rhizocarpales Mia˛dl. et al.,
Mycologia 98: 1097. 2006; nom. inval. [ICN Art.
32.1(d)]
A new order within subclass Lecanoromycetidae
containing the two families Rhizocarpaceae and
Sporastatiaceae. Lichenized or rarely lichenicolous;
thallus crustose, black prothallus often present;
photobiont chlorococcoid. Ascomata apotheciate,
immersed to sessile, lecideine or emarginate. Hama-
thecium of unbranched (Sporastatiaceae) to
branched and anastomosing (Rhizocarpaceae) pa-
raphyses, amyloid. Asci semifissitunicate, clavate,
with amyloid tholus lacking other structures (Spor-
astatiaceae) or with weakly amyloid tholus with
stronger amyloid cap (Rhizocarpaceae). Ascospores
(1–)8 (Rhizocarpaceae) to many (ca. 100; Sporasta-
tiaceae) per ascus, non-septate (Sporastatiaceae) or
transversally septate to muriform (Rhizocarpaceae),
ellipsoid, hyaline to dark brown, non-amyloid.
Conidiomata pycnidia; conidia non-septate, bacillar
to acicular or filiform, hyaline. Secondary chemistry:
depsides, depsidones, pulvinic acid derivates. On
rock or lichenicolous on saxicolous lichens, more
rarely epiphytic on bark. HOLOTYPE family: Rhizo-
carpaceae M.Choisy & Hafellner. HOLOTYPE genus:
Rhizocarpon Ramond ex DC.
Stromatothelium Trevis.
LECTOTYPE: Stromathothelium cruentum (Mont.) Tre-
vis. ” Pyrenula cruenta (Mont.) Vain.; here
designated.
This genus was introduced by Trevisan (1861)
for four species, without naming a type species. Two
of these, Stromatothelium arctecinctum (Fée) Trevis.
and S. uberinum (Fée) Trevis. are conspecific and
represent Architrypethelium uberinum (Fée) Aptroot.
The other two, S. cruentum and S. melanophthalmum
(Mont.) Trevis., are species accepted in Pyrenula
(Aptroot 2012). Notably, the four species differ
markedly in ascospore size, but the protologue does
not mention this, so there is no possibility to
determine a lectotype based on features listed as part
of the genus description. Lectotypification of Stro-
mathelium with either of the first two species would
give this name priority over Architrypethelium Apt-
root, whereas lectotypification with either of the
remaining two species would make it a synonym of
Pyrenula, which is the less disruptive solution and
therefore preferred. We selected S. cruentum as
lectotype to make this name potentially available

should Pyrenula be split into several genera following
a recent phylogenetic study (Weerakoon et al. 2012).
In that study, the type of Pyrenula, P. nitida (Weigel)
Ach. and P. cruenta are separated in two large clades
and P. cruenta also forms a smaller clade with several
other species, sister to a clade including the genus
Pyrgillus. The fourth species listed under Stromatho-
thelium, now named P. melanophthalma (Mont.)
Trevis., has not yet been sequenced.
Teloschistineae Gaya & Lutzoni, subordo nov.
[MYCOBANK 819268]
A new suborder within order Teloschistales
containing the families Megalosporaceae and Telo-
schistaceae. Lichenized, very rarely lichenicolous;
thallus crustose to squamulose, foliose, or fruticose,
often yellow-orange to orange-red (Teloschistaceae);
photobiont chlorococcoid. Ascomata apotheciate,
mostly lecanorine or pseudolecanorine, more rarely
biatorine or lecideine. Hamathecium consisting of
unbranched to slightly branched and sometimes
anastomosing, often slightly capitate paraphyses,
amyloid. Asci semifissitunicate, with apical tholus
and (in Teloschistaceae) distinct ocular chamber,
upper part of tholus (cap) and lower part lining the
ocular chamber darker amyloid, cylindrical to
clavate. Ascospores (1–)8(–16) per ascus, transverse-
ly septate to muriform, oblong-ellipsoid, either
euseptate with rectangular lumina and usually
megalosporous with large lumina and thickened
walls (Megalosporaceae) or polarilocular with very
thick septa and central canal connecting the lumina
(Teloschistaceae), hyaline, non-amyloid. Conidio-
mata pycnidia; conidia non-septate, bacillar to
clavate, ellipsoid or bifusiform, hyaline. Secondary
chemistry: variable, commonly with anthraquinones,
depsides (atranorin), depsidones (pannarin), diben-
zofuranes (usnic acid), terpenes (zeorin), and
sometimes lichexanthone. Mostly on rock and
epiphytic on bark. HOLOTYPE family: Teloschistaceae
Zahlbr. HOLOTYPE genus: Teloschistes Norman.
OUTLOOK
The classification of lichen fungi, and the
classification of Fungi in general, has made consid-
erable progress in the past six to eight years after
publication of the latest Outline of the Ascomycota
(Lumbsch et al. 2010) or the latest Dictionary of the
Fungi (Kirk et al. 2008). Yet, much remains to be
done, even in lichenized lineages, which are arguably
Lücking et al.: 2016 lichen classification 403
much better studied that other fungi. In our view,
there should be several priorities for research into
lichen fungal systematics for the coming years:
(1) Molecular data should be obtained for as many
genera as possible among the 244 currently listed
without such data (Table 4), to clarify their
phylogenetic relationships.
(2) Large, megadiverse to ultradiverse genera that
contain more than 100 species (Tables 2 & 3)
but have comparatively few molecular data
available should be broadly sampled to reassess
generic delimitation, as these are likely to
contain unrecognized genus-level lineages. In
addition, the status of names described in these
genera should be assessed in order to obtain
more reliable estimates of their species richness.
(3) Studies on species delimitation should be
extended to include a broad sampling across
all major lineages and obtain a statistically based
mean on how much species richness in currently
accepted species is under- or overestimated.
(4) Families such as Teloschistaceae, in which a
large number of changes to the genus concept
have been introduced in widely dispersed studies
by different authors, need to be consolidated by
a broad, multi-locus analysis including all
available data, and with rigorous analytical
methods, as has been done for Collemataceae,
Graphidaceae, Pannariaceae and Parmeliaceae,
among others.
(5) Basal orphaned lineages within the Leotiomyceta
for which molecular data are available but which
have an unresolved position in the vicinity of
Lecanoromycetes, such as Thelocarpales and
Vezdaeales, need to be reanalyzed using a broad
data set, in order to resolve such phylogenetic
uncertainties. For such cases, new approaches
using high throughput sequencing of a broad
selection of markers might be an appropriate
approach.
(6) Borderline lichens need to be critically assessed
using sophisticated microscopical and molecular
as well as culture methods, following the model
study of Schizoxylon albescens (Grube & Wedin
2016; Muggia et al. 2016), to test whether these
associations actually do represent lichens and to
elucidate their biological status.
404 The Bryologist 119(4): 2016
ACKNOWLEDGMENTS
Mats Wedin provided a thorough revision of an earlier version of
this work. Numerous colleagues helped in checking details of the
classification or nomenclatural matters and are warmly thanked:
André Aptroot, Mika Bendiksby, Othmar Breuss, Paul Diederich,
Pradeep Divakar, Damien Ertz, Ester Gaya, Cecile Gueidan, Martin
Grube, Rosmarie Honegger, Klaus Kalb, Thorsten Lumbsch,
Francois Lutzoni, Jolanta Mia˛dlikowska, Claude Roux, Matthias
Schulz, Harrie Sipman, Toby Spribille, Einar Timdal, Nicholas
Turland and Martin Westberg. James Lawrey assisted in developing
the general format and layout. Teppo Rämä and Steve Trudell kindly
placed images of Requienella seminuda and Typhula erythropus at
our disposal, and Dan Molter and Kean10 are thanked for providing
images through Wikimedia Commons.
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manuscript received October 13, 2016; accepted November 22, 2016.
Supplementary Documents Online:
Supplementary Table S1. The 2016 classification of lichenized fungi
in the Ascomycota and Basidiomycota in spreadsheet format.
Supplementary Figure S2. The 2016 classification of lichenized
fungi in the Ascomycota and Basidiomycota in poster format.