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Lichenologist 35(3): 191–232 (2003)
doi:10.1016/S0024-2829(03)00027-6
The lichenicolous fungi of Great Britain and Ireland: an overview
and annotated checklist
David L. HAWKSWORTH
Abstract: Since the last compilation in 1983, the number of lichenicolous fungi (and lichenicolous
lichens) recorded in Great Britain and Ireland has increased from 218 to 403 species (a rise of 85%).
The introduction includes accounts of: the history of investigation, circumscription, phylogenetic
relationships, symbiotic interactions, dispersal and establishment, distribution, culture, host restriction,
and identification. The checklist enumerates the recorded species. Information is provided
on the systematic position of each genus, together with the authors and dates of publication of
all accepted names; synonyms used in the literature of the two countries since 1910 are
cross-indexed. References to pertinent literature with descriptions and discussions are included,
together with notes on particular species. The list of references cited includes 298 publications. The
following new scientific names are introduced: Diederichia gen. nov., and Dactylospora scapanaria
(syn. Lecidea scapanaria), Diederichia pseudeverniae (syn. Macrophomina pseudeverniae), Muellerella
ventosicola (syn. Microthelia ventosicola) and Nigromacula uniseptata (syn. Vouauxiella uniseptata)
combs. nov.
2003 The British Lichen Society. Published by Elsevier Ltd. All rights reserved.
Key words: Ascomycota, Basidiomycota, British Isles, lichens, conidial fungi, symbiosis, UK.
Introduction
In 1983, a key was presented to the 218
lichenicolous fungi and lichenicolous lichens
then known to occur in Great Britain and
Ireland (Hawksworth 1983a). The last
two decades have seen an unprecedented
increase in interest in these previously
largely ignored fungi, and so many have
been discovered in the region since that time
that a new checklist has become muchneeded.
This contribution provides an enumeration
of the recorded species, together
with systematic placements, dates of publication,
synonyms used in the literature of
the area, and references to publications with
descriptions, illustrations and keys; notes are
included where appropriate. Also provided
is an overview of the history of their study
in the region, notes on their biology and
D. L. Hawksworth: The Yellow House, Calle Aguila
12, Colonia La Maliciosa, Mataelpino, Madrid 28492,
Spain.
relationships, and an introduction to the
literature on their identification.
History of investigations
The first reports of lichenicolous fungi
from Great Britain and Ireland were unwitting
ones where infected specimens of
lichens were described, but interpreted as
lichens. Biatoropsis usnearum was illustrated
in Dillenius (1742: pl. 12 fig. 6; Diederich &
Christiansen 1994) and Smith & Sowerby
(1801a: pl. 872), and other examples in the
latter work included Sclerococcum sphaerale
(Smith & Sowerby 1801b: pl. 964) and
Unguiculariopsis lettaui (Smith & Sowerby
1804: pl. 1353). Davies (1794: 284)
described Arthonia varians, considering it a
new lichen and not appreciating it was growing
on Lecanora rupicola. The first deliberate
description of a lichenicolous fungus from
the British Isles appears to be that of Abrothallus
parmeliarum by James Edward Smith
0024-2829/03/030191+42 $30.00/0 2003 The British Lichen Society. Published by Elsevier Ltd. All rights reserved.

192 THE LICHENOLOGIST
Vol. 35
(in Smith & Sowerby 1808: pl. 1866) which
‘grows parasitically in patches or tufts, upon
fronds’ of the species now called Parmelia
omphalodes. Scattered contributions followed
in the first half of the nineteenth
century, most notably new species described
from Ireland by Taylor (1836; Hawksworth
1979c), and from Great Britain by Turner &
Borrer (1839), and Berkeley (Berkeley
1844; Berkeley & Broome 1852). Leighton
(1851) and Mudd (1861) were seemingly
more aware of their existence, covering them
routinely in their lichen surveys, but it is
W. Lauder Lindsay (1829–1880) who was
the true pioneer in the field both nationally
and internationally. As far as I know,
he was the first author to use the term
‘lichenicolous’ (Lindsay 1869a).
Lindsay first produced a monograph of
Abrothallus, in an extremely broad sense by
modern standards (Lindsay 1857 1 ), proceeded
to catalogue the lichenicolous fungi
known throughout the world (Lindsay
1869a), and then to describe meticulously
and illustrate the species he found often in
colour and with ranges of specimens to show
variability in microscopic features (Lindsay
1869b, 1871). He had a remarkable grasp
of the world literature and an ability to
interpret correctly what he observed, biologically
and systematically, always proceeding
with caution and presenting
detailed descriptions and arguments for his
decisions. He recognized that there was no
clear line between lichens, and lichenicolous
and other fungi, and also published on
lichenicolous species from Greenland and
New Zealand in particular. Most species he
described are still recognized today.
In the later half of the nineteenth century,
several British and Irish lichenologists, for
instance Crombie, Carroll and Larbalestier,
regularly sent material to Nylander in Paris
which was described as new by him, but no
1 In a set of Lindsay’s papers which he had bound for
his personal use and are now in my personal library, his
own handwritten index gives the date as 1856; the
paper was read at a meeting of the British Association in
Cheltenham in August 1856 and this part of the journal
could well have been released earlier than generally
assumed.
one took over Lindsay’s mantle in this
period.
After a lull in activity at the end of the
nineteenth century, Annie L. Smith encouraged
members of the British Mycological
Society to note these fungi on Society forays,
and produced notes (Smith 1910) on some
species encountered while working on her
monographs of lichens occurring in the
British Isles (Smith 1911). She was evidently
much impressed by Vouaux’s (1912–
14) monograph of lichenicolous fungi (see
p. 199) and included an expanded treatment
of lichenicolous taxa in the second edition of
her work, accepting some 55 species (Smith
1926). Stimulated at first by contact with
Smith, Walter Watson was the next British
person to take an interest in lichenicolous
fungi. Then, fired by enthusiasm generated
by Keissler’s (1930) voluminous treatment
of the central European species, Watson
(1948) produced a new checklist with full
literature references, and also information
on distributions as far as these were known.
Apart from passing interests developed by
T. D. V. Swinscow (e.g. Swinscow 1962)
and O. L. Gilbert (Gilbert 1966), there was
almost no interest at all in these fungi in
Great Britain and Ireland during the 1950s
and 1960s. But the situation was to change
in the 1970s, which started with the study of
Folan & Mitchell (1970) on the lichens and
lichenicolous fungi of Derryclare Wood, Co.
Galway, Ireland; Rolf Santesson, then in
Uppsala, assisted with the identifications.
I started to be fascinated by these fungi
after I discovered a new species on Alectoria
ochroleuca in the course of studies on that
genus (Hawksworth 1971). Soon after, I
started to produce a series of notes on new
and interesting lichenicolous fungi I encountered
(Hawksworth 1975a; 1978; 1980b;
1982a; 1986; 1994) as well as treatments of
di#erent groups. This was facilitated by my
being located in Kew during most of this
period with ready access to the extraordinarily
rich fungal collections, libraries, and
specialist mycologists of the Royal Botanic
Gardens and what later became the
International Mycological Institute. Brian J.
Coppins started to take a particular interest

2003 Lichenicolous fungi of Great Britain and Ireland—Hawksworth 193
Table 1. Numbers of lichenicolous fungi recorded from Great Britain and Ireland
Source Accepted species* Increase on previous list (%)
Smith (1926) 55 –
Watson (1948) 148 169
Hawksworth et al. (1980) 183 24
Hawksworth (1983a) 218 16
Present work 403 85
*Myxomycota are excluded as they have not been treated in all sources, nor in the present checklist; the numbers
used are those of the taxa accepted by authors in the works cited.
in the discomycetes on lichens, especially in
mainly lichenized genera, around this time
and passed numerous collections to me for
study, later also starting to describe new
taxa himself (e.g. Coppins 1987, 1989).
Lichenicolous fungi were included in the
checklist of Hawksworth et al. (1980a), a
practice that has since become commonplace,
and a key was subsequently provided
(Hawksworth 1983a). Following the publication
of the key, and workshops on lichenicolous
fungi, sponsored by the British
Mycological Society at Southampton in
1983 and Lyndhurst in the New Forest
in 1998, and by the British Lichen Society
at Kew in 1991, increasing numbers of
lichenologists became interested in these
fungi, many including notes of their discoveries
in the British Lichen Society’s
Bulletin (Table 1). Especially active have
been Peter M. Earland-Bennett, Oliver L.
Gilbert, Chris J. B. Hitch, Howard F. Fox,
and the late Humphrey J. M. Bowen; all
have generously sent material to me for
study.
If this new checklist and overview, despite
its short-comings, stimulates even further
interest in these fascinating fungi, it will
have served its purpose. An updated key is in
draft, also including the Spanish species,
and accounts of many genera with descriptions
have been prepared for the NERCfunded
Ascomycetes of Great Britain and
Ireland project.
Circumscription
Lichenicolous fungi in the strict sense
are fungi which obligately live on lichens,
whether as parasites, commensals, or
saprobes, at least at some stage in their
life-cycle. Species fortuitously occurring on
decayed lichens are not included, though a
few widely distributed fungal saprobes have
been retained here as exceptions as they
occurred on lichens in the field (e.g.
Aspergillus glaucus, Trichothecium roseum).
Also excluded are the numerous species that
can be isolated into culture from ground-up
lichen tissues, for example the 506 strains
obtained from 17 lichen samples by Petrini
et al. (1990), and the 117 fungi isolated from
two lichen species by Girlanda et al. (1997).
Drawing any boundary between lichenicolous
lichens and lichenicolous fungi is
problematic as many of these ‘lichens’ start
out as fungal invaders of lichen thalli and
then either take over the host’s photosynthetic
partner killing the host in the process
and becoming completely independent (e.g.
Diploschistes muscorum), or form colonies
with either an independent or the same
photobiont on (or in) the host lichen thallus
(e.g. Rimularia insularis); these methods of
establishing lichen thalli are probably much
more widespread than is generally recognized
(Hawksworth 1988; Richardson
1999). Lichens which certainly have a
lichenicolous fungal stage in their life-cycle
have been included here, and are also covered
in the latest lichen checklist (Coppins
2002).
Phylogenetic relationships
The lichenicolous habit has been
extremely successful in evolutionary terms,
with some 1559 species described (Lawrey
& Diederich 2003) and 3–4000 species estimated
world-wide (Hawksworth 2001).

194 THE LICHENOLOGIST
Vol. 35
This compares with the world total of
13 500 described and 18 000 estimated
lichen-forming fungi (Sipman & Aptroot
2001). Fungi that have developed a lichenicolous
strategy are drawn from a wide range
of fungal groups, 16 of the 55 orders of
Ascomycota, and six of 31 orders of Basidiomycota
(Kirk et al. 2001; Lawrey &
Diederich 2003); in addition, many are in
families not yet placed in orders, or in
genera not referred to families, or are ascomycetous
or basidiomycetous conidial
(mitosporic) 2
fungi whose phylogenetic
relationships are uncertain.
The possibility that many fungal groups
now occurring on plants and plant products
might have been derived from lichenized
ancestors has been aired in the past (e.g.
Eriksson 1981; Hawksworth 1982c). Now,
molecular data is tending to support this
view, and Lutzoni et al. (2001) have suggested
that lichenicolous fungi might be a
‘half-way house’ facilitating transfers to
plant and other substrata. The inclusion of
both lichenized and lichenicolous species in
the same genus, and congeneric fungicolous
2 Sutton (1993) proposed the term ‘mitosporic’ for all
fungi in which no sexual stage was known, and also for
the asexual stages of pleomorphic species. This term
was broadly equivalent to the Deuteromycetes, Fungi
imperfecti, etc., of earlier workers (latinized formal
suprageneric names long considered inappropriate
as many of these fungi are unrelated and dispersed
amongst other sexual fungal groups). Although
Sutton’s term was adopted in the eighth edition of the
Dictionary of Fungi (Hawksworth et al. 1995), it was so
unpopular amongst mycologists that its use was
dropped in the ninth (Kirk et al. 2001). Instead, Kirk
et al. use ‘anamorphic fungi’, ‘anamorphic Ascomycota’,
‘anamorphic Mycosphaerellaceae’, etc. However,
the term ‘anamorph’ strictly applies only in cases
where sexual stages are known (Art. 59.1 of the Code)
and implies that the asexual taxa are never genera or
species in their own right, but states of sexual fungi yet
to be found. To even state ‘anamorphic Ascomycota’ can
be unsound in the absence of molecular or ultrastructural
information (cf. the case of Marchandiomyces
which proved to be the state of a basidiomycete). I
therefore prefer to use the informal and familiar ‘conidial
fungi’—at the same time recognizing that many
of these can be unequivocably placed even in a ‘sexual’
(teleomorphic, meiosporic) genus by molecular phylogenetic
methods. The issue of limiting future use of the
dual nomenclatural system will be debated at the next
International Botanical Congress in 2005.
and lichenicolous species, supports such a
view (see p. 193). Further, it may be pertinent
that some lichenicolous fungi are
currently placed in genera best-known as
ones of plant pathogens or saprobes (e.g.
Fusarium, Odontotrema, Phoma, Raciborskiomyces).
Additional support for this view
comes from the discovery by molecular
methods that Nesolechia and Phacopsis
belong in the Parmeliaceae (Persˇoh &
Rambold 2002).
Symbiotic interactions
The di#erent biological strategies adopted
by lichenicolous fungi were highlighted by
Santesson (1967), who pointed to single
genera which included lichenicolous and
saprobic or parasitic species. While some of
the generic concepts have changed, the
principle holds and the number of genera
which are known to include fungi that
are lichenicolous and also species which
form lichens, are increasing; examples are
Arthonia, Arthrorhaphis, Caloplaca, Carbonea,
Diplotomma, Opegrapha, Rhizocarpon,
Rimularia, Rinodina, and Toninia. Some
genera also include both lichenicolous
species and ones growing on non-lichenized
fungi (e.g. Cornutispora, Nectriopsis, Tremella),
and Chaenothecopsis encompasses
lichenicolous, fungicolous, and wood
saprobic species.
Hawksworth (1982b) classified lichenicolous
fungi into three categories: parasites
(subdivided into pathogens, restricted discolourations,
extensive discolourations, or
gall-like malformations), saprophytes, and
parasymbionts 3 . This scheme has been
developed by Lawrey & Diederich (2003)
who recognize four types of parasitic relationships:
pathogens, restricted discolourations,
galls, and commensals. I also feel it
necessary to reinstate saprophytes, but to
use the term ‘saprobes’ as lichens are not
plants, making five categories: (1) The most
pathogenic species cause extensive lesions
and can form circular colonies on mixtures
3 This term is regarded as inappropriate and its use is
no longer recommended (Hawksworth 1988).

2003 Lichenicolous fungi of Great Britain and Ireland—Hawksworth 195
of thalli (e.g. Athelia arachnoidea on Lecanora
conizaeoides, Physcia and Xanthoria stands;
Gilbert 1988; Fig. 1A), individual thalli
(e.g. Pronectria santessonii on Anaptychia
runcinata), or bleach and destroy lobes
(e.g. Weddellomyces epicallopisma on lobate
Calopaca spp.; Fig. 1B). (2) Restricted discolourations
occur when infections are confined
to discrete patches on the thallus,
often encircled by a ring of blackened lichen
tissue and recalling infection spots of many
foliar plant pathogens (e.g. Lichenoconium
lecanorae on Parmelia sulcata; Fig. 1C). (3)
Gall-formers are especially frequent in
particular lichenicolous genera, such as Polycoccum,
but there are numerous other
examples (e.g. Telogalla olivieri on Xanthoria
parietina; Fig. 1D); a survey of the thenknown
gall-forming lichens was prepared by
Grummann (1960), but it is important to
note that galls on lichens can be formed by
organisms other than fungi, particularly
nematodes (Siddiqi & Hawksworth 1982).
(4) Commensals, where the lichen thallus
appears una#ected and retains its normal
colour (e.g. Endococcus perpusillus on Rhizocarpon
geographicum; Fig. 1E); the possibility
that some commensals might be involved
in a form of mutualistic relationship
cannot be discounted. (5) Saprobes occurring
on ageing parts of the thallus (e.g.
Niesslia cladoniicola, Lettauia cladoniicola;
Fig. 1F).
The biological, physiological, and physical
(including ultrastructural) details of the
interactions between lichenicolous fungi and
their hosts have hardly been studied. In
some cases the algal partner may be the
prime target, while in others it may be the
lichenized fungal component. The algal
partner of the host can be taken over by the
invading fungus to form lichenicolous
lichens or independent lichen thalli, for
instance in Diploschistes musorum on Cladonia
species (Friedel 1987) and Rimularia
insularis on Lecanora rupicola (Ríos et al.
2002). Zwackhiomyces coepulonus attacks
and kills the algal cells in Xanthoria elegans
(Grube & Hafellner 1990). In contrast, in
four members of the Dacampiaceae, Ríos
& Grube (2000) found that the licheni-
colous fungi attacked the fungal and not
the algal partner of the lichen. Hyphae of
the invading fungus can sometimes be distinguished
microscopically within a lichen
thallus from those of the fungal component
of the lichen, the extent of invasion
varying with di#erent fungus-lichen combinations
(Fahselt et al. 2001). Damaged or
stressed lichens are more likely to become
infected by less host-restricted species,
especially when conditions are moist, as
shown experimentally by Glenn et al.
(1997).
Secondary compounds in the lichen thallus
may be important in limiting infections
by other fungi (e.g. Yamamoto et al. 2002).
Overcoming the barrier posed by these compounds
may be important in the evolution of
lichenicolous fungi. Lawrey (1997) demonstrated
experimentally that some unrelated
lichenicolous fungi grew better on lichen
thalli containing lichen compounds, than on
thalli from which such compounds had been
removed. An as yet undescribed lichenicolous
Fusarium species has been shown to
be able to degrade lichen tissue in the presence
of antibiotic compounds, while phylogenetically
allied species of the same genus
could not (Torzilli et al. 2002). However,
some lichenicolous fungi tolerate the secondary
compounds of their hosts, but not
those of other lichens (Lawrey et al. 1999).
Experimental studies on these phenomena
are summarized in Lawrey & Diederich
(2003).
One unanswered question is the extent to
which lichenicolous fungi produce secondary
compounds, and whether these have a
role in their specificity or activity. By comparing
the secondary metabolites present in
areas of lichens infected by 30 genera of
lichenicolous fungi with those in the rest of
the thallus, Hawksworth et al. (1993) found
13 di#erent compounds in five genera;
amongst these were substances tentatively
identified (by TLC characteristics) as
gyrophoric acid (in Sclerococcum sphaerale
and Skyttea nitschkei), pannarin (in Milospium
graphideorum), lipids (in Endococcus
apicicola), and a triterpenoid (in Marchandiomyces
corallinus).

196 THE LICHENOLOGIST
Vol. 35
Dispersal and establishment
Other aspects of dispersal and establishment
are little studied, except in the case of
certain species which become lichenized
(see p. 195). However, dispersal is clearly a
limiting factor as the richest sites for lichenicolous
fungi are ones with long histories of
ecological continuity. Indeed, lichenicolous
fungi should be more widely considered in
assessing the conservation importance of
sites. When comparing sites with old forest
indicator species (cf. Coppins & Coppins
2002), those which also have obligate
lichenicolous fungi on those indicator
lichens may be the sites with the longest
history of continuity; i.e. a site with Plectocarpon
lichenum on Lobaria, Refractohilum
galligenum on Nephroma, and Skyttea
nitschkei on Thelotrema, is likely to be of
greater conservation interest than a site with
the same host lichens but lacking such fungi.
The same principles apply to terricolous
lichens; it is the least disturbed sites that will
have the most fungi specialized on Cladonia
and Peltigera species. Dispersal in most
cases is probably primarily by discharged
ascospores and (or) rain or stem-flow. Pyatt
& Harvey (1973), however, reported that
ascospores of Muellerella lichenicola could
be dispersed along with those of the host
Caloplaca, but this cannot apply to species
which do not form fruit bodies within host
lichen ascomata. It may be significant that
many lichenicolous fungi produce large,
multi-celled, and often thick-walled spores;
such characteristics will increase longevity
and chances of germinating near potential
hosts (Hawksworth 1987). Lawrey &
Diederich (2003) discuss some of these
aspects further.
Distribution
The distribution of lichenicolous fungi is,
like that of plant-restricted fungi, intrinsicly
limited to that of their potential host lichens.
However, records of particular lichenicolous
species are often at present from only a part
of the range of a species. For example,
Homostegia hertelii occurs on Flavoparmelia
baltimorensis and F. caperata, but is currently
known only from North America despite
the wide distribution of the latter species
in Europe (Hawksworth et al. 2004). In
contrast, Lasiosphaeriopsis stereocaulicola is
known from di#erent Stereocaulon species
in Africa, Europe and South America, but
was not reported from North America
where many species of the host genus occur
(Eriksson & Santesson 1986). However, so
few detailed field surveys for these fungi
have been done, that the serendipity of
collection remains a key factor in what is
known of species distributions. For instance,
Cornutispora ciliata was described from
Tasmania, but later found in the UK only
about 100 m from the type locality of C.
lichenicola in Devon; the last species is now
known to have a wide distribution, including
North America, and occurs on a variety of
host lichen genera—something unforeseen
when the species was first described.
The keen eyes of particular collectors also
have to be considered, examples from the
British Isles being P. M. Earland-Bennett’s
eye for Psammina species (Earland-Bennett
& Hawksworth 1999a), and the late M. C.
Clark’s for Trichonectria hirta (Hawksworth
1978).
When endeavouring to identify lichenicolous
fungi, it is consequently essential to
consider the literature on species described
from all parts of the world, as well as on
di#erent hosts.
Culture
A wide range of lichenicolous fungi have
now been successfully grown in pure culture,
opening up possibilities for experimentation
as well as exploitation for novel
bioactive compounds. Conidial fungi are
most easily cultured, including species of
Acremonium (Gams 1971), Intralichen
christiansenii (Hawksworth 1979a), and
Sclerococcum sphaerale (Hawksworth & Jones
1981). Anamorphs can often be grown from
ascospores, as in Niesslia cladoniicola
(Hawksworth 1975a) and Pronectria species
(Lowen & Hawksworth 1986; Lowen
1995). Crittenden et al. (1995) reported the
successful isolation of 11 (31%) of 36

Fig. 1. Examples of kinds of relationships between lichenicolous fungi and their host lichens. A, pathogenic
(Athelia arachnoidea on Xanthoria parietina); B, pathogenic (Weddellomyces epicallopisma on Caloplaca flavescens);
C, restricted discolouration, with a blackened zone around the infection spot (Lichenoconium lecanorae on Parmelia
saxatilis); D, gall-forming (Telogalla olivieri on X. parietina); E, commensalistic (Endococcus perpusillus on
Rhizocarpon geographicum); F, saprobic (Niesslia cladoniicola on Cladonia rangiformis); G, Corticifraga fuckelii causing
a generalized infection on Peltigera hymenina; H,C. peltigerae causing a restricted and well-delimited infection on
P. didactyla. Not to scale. Photographs taken in collaboration with D. W. Fry (A, C–H) or G. Godwin (B), and
reproduced with the permission of CABI Bioscience.

198 THE LICHENOLOGIST
Vol. 35
lichenicolous species attempted (i.e. Clypeococcum
hypocenomycis, Dactylospora parasitica,
Diploschistes muscorum, Echinothecium
reticulatum, Endococcus rugulosus, Illosporiopsis
christiansenii, Lichenoconium erodens, L.
lecanorae, Muellerella pygmaea, Skyttea
nitschkei, and Vouauxiella lichenicola). It is
evident that there is the potential for much
more experimental work using pure cultures
of lichenicolous fungi. Further aspects of
culturing lichenicolous fungi are discussed
by Lawrey & Diederich (2003).
Host-restriction
Di#erent orders, families, and genera
of lichens vary in the extent to which they
serve as hosts for lichenicolous fungi. The
peltigeralean lichens are an especially rich
source of these fungi, as are Cladoniaceae
and Pertusariaceae. In contrast, very few
occur on pyrenocarpous lichens or thelotremes;
the parmelioid lichens also have
rather few in relation to the numbers of
species in the family. Hawksworth (1982c)
suggested that this phenomenon might
be related to the antiquity of the hosts,
especially where lichenicolous genera
occurred on only one family or genus.
Molecular phylogenetic data will eventually
establish whether this is really the case.
The degree to which particular lichenicolous
genera or species are restricted to
particular host lichen families, genera, or
species, varies markedly. The more pathogenic
species tend to be wide-ranging (e.g.
Athelia arachnoidea), as do some which
appear to attack often already damaged
thalli (e.g. Cornutispora lichenicola, Lichenoconium
erodens; Gilbert 1988). In gallforming
and commensalistic species,
however, the fungi tend often to be
restricted to individual genera or species, as
in the case of Abrothallus, Dactylospora, Polycoccum,
Stigmidium, and Tremella. In some
cases members of the same genus can have
di#erent e#ects on the host lichens, as in
the three Lichenoconium species able to
grow on Parmelia saxatilis (Hawksworth
1977a,b), and Corticifraga species on
Peltigera (Hawksworth & Santesson 1990;
Fig. 1G & H). In describing new lichenicolous
fungi, it is always important to study
the features of the fungi first, and consider
the links with hosts separately. Also, when
there are few collections there is a danger
of over-emphasizing small di#erences or
discontinuities in, for example, spore size,
when only part of the real variation has
actually been sampled.
Lichenicolous fungi can also have a role as
taxonomists, indicating where the relationships
between hosts merit re-examination.
Examples are the occurrences of Microcalicium
arenarium on Psilolechia lucida and
Micarea clavulifera supporting the transfer of
the latter into Psilolechia despite the di#erent
colours of the thallus (Coppins 1983). They
also support the inclusion of Cladina within
Cladonia, as species in both ‘genera’ can be
attacked by the same lichenicolous species
(e.g. Abrothallus cladoniae, Bachmanniomyces
uncialicola, Lettauia cladoniae, Lichenoconium
pyxidatae), a conclusion now supported by
molecular studies. In the case of the
parmelioid lichens, species of Abrothallus,
Lichenostigma, Nesolechia, and Stigmidium
are proving of particular assistance, supporting
some genera segregated from Parmelia
s. lat. and again tending to agree well with
molecular phylogenetic evidence (D. L.
Hawksworth, unpubl.). Other examples
of host restrictions are cited by Lawrey &
Diederich (2003).
However, caution is also required,
especially where the algal partner of the
lichen is being utilized by the fungus rather
than the fungal partner. Leightoniomyces
phillipsii, for example, occurs on Steinia
geophana (?Lecideaceae), Thrombium epigaeum
(Thrombiaceae) and probably Vezdaea
leprosa (Vezdaeaceae), all three of which have
Leptosira as photobiont.
Identification
The identification of lichenicolous fungi
requires a knowledge of the general mycological
literature as well as familiarity with
the lichen hosts. It is for this last reason that
these fungi are most commonly studied by
lichenologists; other mycologists may not

2003 Lichenicolous fungi of Great Britain and Ireland—Hawksworth 199
Table 2. Surveys, indices and keys to lichenicolous fungi occurring on particular lichen hosts
Host lichens Reference(s)
Arthrorhaphis Hafellner & Obermayer (1995)
Baeomyces Ihlen (1998)
Brigantiaea Hafellner (1995)
Cladonia Bachmann (1927–28)
Dibaeis Ihlen (1998)
Haematomma Kalb et al. (1995)
Icmadophila Ihlen (1998)
Koerberiella Rambold et al. (1990)
Lepraria Kummerling et al. (1993)
Lobaria Etayo & Diederich (1996b)
Lobariaceae Wedin & Hafellner (1998)
Omphalina Santesson (1989)
Peltigera Hawksworth (1980b), Hawksworth & Mia˛dlikowska (1997)
Kummerling & Alstrup (1992), andMartínez Moreno (1999)
Peltigerales Martínez & Hafellner (1998).
Pseudocyphellaria Kondratyuk & Galloway (1995), Kondratyuk et al. (1994)
Squamarina Navarro-Rosinés et al. (1994, 1995)
Thamnolia Ihlen (1995)
Trapelia Hafellner & Berger (2000)
Foliicolous lichens Matzer (1996)
Lecideoid lichens Triebel (1989)
Pyrenocarpous aquatic lichens Molitor & Diederich (1997)
All lichens Keissler (1930), Oudemans (1918–24), Zopf (1896), Index of Fungi (1940 on)
always recognize which ascomata or conidiomata
are those of the lichen as opposed to
an invasive fungus. There are examples in
the literature of lichen fruit bodies being
described by mycologists as lichenicolous
fungi (e.g. Pleurosticta lichenicola), and also
ones of lichenicolous fungi being described
as ‘on wood’ or ‘on bark’ with no realization
that they were actually growing on lichens
(e.g. Trichonectria hirta).
The literature on lichenicolous fungi has
grown at an amazing rate during the last two
decades, and while there is no single synthesis
of the available information, there are
several major compilations and some other
sources that are invaluable in endeavouring
to identify a lichenicolous fungus. Lindsay’s
(1869a) catalogue has already been mentioned
(see p. 192), and was followed by
Arnold’s (1874) which included 240 species.
Zopf (1896) compiled a world list by
host, and Oudemans (1918–24) compiled
another for Europe. Lichenicolous fungi
were covered with general fungal (including
lichen) literature by Lindau (1908–17) and
Ciferri (1957–60), and in the mammoth
compilation of the world’s fungi (with
descriptions) by Saccardo (1882–1931,
1972); sadly this last work misses many
names of lichenicolous fungi as the lichen
literature was poorly scanned for nonlichenized
fungal names.
However, of crucial importance for
identification is the world monograph of
Vouaux (1912–14) which endeavoured to
provide descriptions and keys and treat all
names of lichenicolous fungi known up to
that time, wherever possible basing these on
material he had examined personally.
Keissler took a keen interest in these fungi,
culminating in a major treatment of the
central European species with extensive
descriptions, synonymies, and also a host
index (Keissler 1930).
No other world compilation appeared for
46 years, when Clauzade & Roux (1976)
produced keys to all lichenicolous fungi they
could trace, covering 457 species at that
time, but mainly based on literature reports.
A second edition of those keys by Clauzade
et al. (1989) treats 686 species, and has
marginal drawings of spores. Yet while this
was a substantial increase (50%) reflecting
the resurgence of interest in the group, the

200 THE LICHENOLOGIST
Vol. 35
number now known has recently been calculated
as 1559 (Lawrey & Diederich 2003),
an increase of a staggering 127% since 1989.
In order to access those names, recourse to
the original literature or to papers cited
in national checklists (e.g. Diederich &
Sérusiaux 2000; Eriksson 1992; Esslinger
& Egan 1995; Kocourková 2000; Llimona &
Hladun 2001; Santesson 1993a; Scholz
2000) is necessary. Also helpful are treatments
of species on particular host lichens
(Table 2) and the compilation of key literature
by fungal genus of Lawrey & Diederich
(2003). Of especial value is the host index to
the twice-yearly Index of Fungi, and if a name
is already known the ‘Species Fungorum’
database (see below) can be accessed free of
charge. Bibliographic listings of papers are
included in the twice-yearly Bibliography of
Systematic Mycology (where papers have
been cross-indexed by genera treated since
1987), and in the ‘Recent literature on lichens’
series in The Bryologist (also available
on http://www.toyen.uis.no/botanisk/lav/
RRL/RRL.htm).
Checklist
This checklist is arranged alphabetically by
genus. The current systematic placement of
each genus being indicated under the
generic name; placements generally follow
Kirk et al. (2001) for basidiomycetes, and
Eriksson et al. (2003) for ascomycetes. The
phylum, order and family are listed in turn
and where one is uncertain this is indicated
by ‘incertae sedis’ (used twice if the order
and family are both uncertain). Where the
anamorphs of teleomorph genera have been
named, this is also indicated. The author
citation and date of publication of all
accepted names are provided; almost all
have been checked and verified in the
original. These names are followed by a
British Lichen Society mapping scheme
reference number printed in bold type (e.g.
2193).
Synonyms are placed in italics and crossreferenced
to the correct names; these are
given without author citations and dates
(these now being mainly of indexing
interest). I have endeavoured to include all
synonyms and variant spellings used in the
literature relating to records from Great
Britain and Ireland that have appeared since
the publication of Smith (1910). Incorrect
reports are also noted in this section, or
ones to species which are now known not to
be lichenicolous. Those wishing to locate
the original place of publication of both
accepted names and synonyms can access
most of these (in some cases indirectly by
reference to secondary sources), through the
‘Species fungorum’ database (http://www.
indexfungorum.org/). Numerous other
earlier synonyms, and those used in the
literature relating to other countries, can be
traced through the reference works listed
above under ‘Identification’. Further information
on the sources of older records can,
in the case of ascomycete genera, be located
through entries in the checklist of Cannon
et al. (1985).
Notes are included where some comment
is pertinent, and the literature cited has been
selected to cover both publications with
descriptions, and discussions of records
from Great Britain and Ireland.
New and interesting records of lichenicolous
fungi from Great Britain and Ireland
have been sporadically included in issues of
the British Lichen Society’s Bulletin since
1983; these are indicated here by an entry
after the mapping scheme number in the
form ‘B 91: 15’ where the bold number is
the issue number and the number after the
colon the page reference.
The following two abbreviations are used
after the names of some synonyms: ‘(F)’
indicates non-lichenized fungi which are not
lichenicolous, while ‘(L)’ denotes lichenforming
fungi not treated elsewhere in the
checklist; for information on the latter see
Coppins (2002). A ‘?’ is used to express
uncertainty.
Abrothallus De Not. 1845
Ascomycota: incertae sedis; incertae sedis.
Anamorph: Vouauxiomyces.
bertianus De Not. 1849 2001
caerulescens Kotte 1909 12; B87: 76

2003 Lichenicolous fungi of Great Britain and Ireland—Hawksworth 201
cetrariae Kotte 1909 2002
cladoniae R. Sant. & D. Hawksw. 1990
2003; B82: 42
microspermus Tul. 1852 4
Teleomorph: V. truncatus.
parmeliarum (Sommerf.) Arnold 1874
2004
Teleomorph: V. cf. santessonii.
prodiens (Harm.) Diederich & Hafellner
1989 2005
suecicus (Kirschst.) Nordin 1964 2006;
B 77: 37, 80: 46
Teleomorph: V. ramalinae.
usneae Rabenh. 1861 2007; B84: 46
welwitschii Mont. ex Tul. 1852 2008
glabratulae = Abrothallus bertianus
lobariellus = Dactylospora lobariella
moorei ? = Phaeopyxis punctum
Note: The genus is in need of a critical revision. The
earliest name for A. parmeliarum may be A. parasiticum
(Ach.) Nyl. ex Sacc. 1881 (syn. Endocarpon parasiticum
Ach. 1814; syn. Lichen parasiticus Sm. 1808 non
Ho#m. 1784), which merits rejection to safeguard
such a well-known name. The Vouauxiomyces anamorphs
of all species have not been accorded separate
binomials.
Lit.: Diederich (1989), Hawksworth (1990), Hawksworth
& Minter (1980), Kotte (1909), Lindsay (1857), and
Nordin (1964).
Acremonium Link 1809
Conidial fungi.
Teleomorph: Nectriopsis and Pronectria.
lichenicola W. Gams 1971 2374; B76: 47
pedatum Lowen 1989 2375
Teleomorph: Pronectria anisospora.
rhabdosporum W. Gams 1971 2210
Note: Not all anamorphs of lichenicolous species have
been formally named.
Lit.: Gams (1971), Hawksworth (1979a), andLowen
(1995).
Actinopeltis
peltigericola = Lichenopeltella peltigericola
Adelococcus Theiss. & Syd. 1918
Ascomycota: Verrucariales: Adelococcaceae.
alpestris (Zopf) Theiss. & Syd. 1918 777;
B 74: 54, 86: 42
interlatens (Arnold) Matzer & Hafellner
1990 2368; B91: 56
cladoniae = Roselliniella cladoniae
groedensis auct. angl. = ? Roselliniella atlantica p. p.
and Peridiothelia fuliguncta p. p. (F)
nephromatis = Roselliniella nephromatis
nephromicola = Roselliniella nephromatis
Lit.: Matzer & Hafellner (1990).
Aegerita
physciae = ? Marchandiomyces corallinus
Agyrium
flavescens = Skytella mulleri
Ahlesia
lichenicola = Thelocarpon lichenicola
Arrhenia Fr. 1849
Basidiomycota: Agaricales: Tricholomataceae.
peltigerina (Peck) Redhead et al. 2002
913; B86: 50
Note: This species is listed on the assumption that
Omphalina cupulatoides P. D. Orton 1977 is a synonym
(Vila 2002).
Lit.: Purvis et al. (1992), Redhead et al. (2002), and
Vila (2002).
Arthonia Ach. 1806
Ascomycota: Arthoniales: Arthoniaceae.
almquistii Vain. 1883 771; B82: 42
amylospora Almq. 1880 1930
apotheciorum (A. Massal.) Almq. 1880
1501
cohabitans Coppins 1989 27; B72: 47
diploiciae Calat. & Diederich 1995 2009;
B 83: 47
epiphyscia Nyl. 1875 122; B79: 38
fuscopurpurea (Tul.) R. Sant. 1960 775
gelidae R. Sant. 1986 1961
graphidicola Coppins 1989 735
intexta Almq. 1880 1933
invadens Coppins 1989 729; B66: 27
molendoi (Heufl. ex Frauenf.) R. Sant.
1986 1934; B88: 67
neglectula Nyl. 1874 2323; B89: 70

202 THE LICHENOLOGIST
Vol. 35
peltigerae Th. Fr. 1866 1935; B84: 46
phaeophysciae Grube & Matzer 1997
1982; B85: 46
punctella Nyl. 1859 1929; B86: 43
subfuscicola (Linds.) Triebel 1991 1936;
B 89: 71
thelotrematis Coppins 1989 1937
varians (Davies) Nyl. 1861 714
abelonae = Corticiruptor abeloneae
clemens auct. angl. = Arthonia apotheciorum
dispersa auct. angl. p. p. = Arthonia excipienda (F)
glaucomaria = Arthonia varians
hibernica = Arthonia excipienda (F)
melaspermella = Rhizodiscina lignyota (F)
stictarum = Plectocarpon lichenum
Lit.: Coppins (1989), Grube et al. (1995), Hawksworth
(1980b), Purvis et al. (1992), and Wedin & Hafellner
(1998).
Arthophacopsis Hafellner 1998
Ascomycota: Arthoniales: incertae sedis.
parmeliarum Hafellner 1998 1983; B84:
47
Lit.: Hafellner (1998).
Arthopyrenia A. Massal. 1852
Ascomycota: incertae sedis: Arthopyreniaceae.
allogena (Nyl) Arnold 1870 2011
desistens (Nyl.) A. L. Sm. 1911 2012
chlorococca = Lauderlindsaya acroglypta (L)
leptotera = Stigmidium marinum
lomnitzensis = Sagediopsis lomnitzensis
marina = Stigmidium marinum
spilobola = Dubiously recorded
strontianensis = Collemopsidium angermannicum (L)
Note: Neither of the two ‘accepted’ species belong in
Arthopyrenia s. str. and a new generic name may be
required for A. allogena.
Arthrorhaphis Th. Fr. 1860
Ascomycota: ? Lecanorales: Arthrorhaphidaceae.
aeruginosa R. Sant. & Tønsberg 1994
1916; B75: 34
citrinella (Ach.) Poelt 1990 100
grisea Th. Fr. 1860 313; B67: 38
muddii Obermayer 1994 1923
fuscireagens auct. = Arthrorhaphis muddii
Note: A. citrinella has not certainly been seen acting as a
lichenicolous species in Great Britain and Ireland.
Lit.: Hansen & Obermayer (1999), Ilhen (1998), Poelt
&Vězda (1977), Purvis et al. (1992), andSantesson &
Tønsberg (1994).
Ascochyta
lichenoides = Crustodiplodia lichenoides (F)
Lit.: Punithalingam (1988).
Ascochytula
lecanorae = Lichenodiplis lecanorae
Aspergillus Link 1809
Conidial fungi.
Teleomorph: Eurotium (also Neosartorya
and Emericella in non-lichenicolous
species).
glaucus Link 1809 2014
Teleomorph: E. herbariorum Link
1809.
Note: Perhaps fortuitously lichenicolous; the teleomorph
has not been found on lichens in the British Isles.
Lit.: Klich (2002), Kozakiewicz (1989), andPitt et al.
(2000).
Asterophoma
See Chaenothecopsis.
Athelia Pers. 1822
Basidiomycota: Polyporales: Atheliaceae.
arachnoidea (Berk.) Jülich 1972 2015
Anamorph: Rhizoctonia carotae Arder
1948
bispora = Athelia arachnoidea
epiphylla auct. lich. pro. max. p. = Athelia
arachnoidea
Note: A. epiphylla Pers. 1822 is essentially a bark
saprobe that sometimes spreads onto adjacent lichens,
but has often been confused with A. arachnoidea by
lichenologists in the past.
Lit.: Adams & Kropp (1996), Arvidsson (1979), and
Jülich (1972, 1978).
Atractium
flammeum = Microcera coccophila (F)

2003 Lichenicolous fungi of Great Britain and Ireland—Hawksworth 203
Lit.: Hawksworth (1979a).
Bachmanniomyces D. Hawksw. 1981
Conidial fungi.
uncialicola (Zopf) D. Hawksw. 1981
2016; B86: 43
Lit.: Hawksworth (1981, 1990).
Bacidia De Not. 1846
Ascomycota: Lecanorales: Ramalinaceae.
killiasii (Hepp) D. Hawksw. 1983 2017
ascaridiella = Spirographa fusisporella
beckhausii = Bacidia beckhausii (L)
plumbina = Toninia plumbina
Note: The generic position of B. killiasii merits
re-investigation; reports of B. beckhausii Körb. 1860 as
lichenicolous are erroneous (Hawksworth 1982a).
Beloniella
nitschkei = Skyttea nitschkei
Biatoropsis Räsänen 1934
Basidiomycota: Platygloeales: incertae
sedis.
usnearum Räsänen 1934 2018
Lit.: Diederich & Christiansen (1994).
Bispora
christiansenii = Intralichen christiansenii
lichenum = Intralichen lichenum
Blarneya D. Hawksw., Coppins & P. James
1980
Conidial fungi.
hibernica D. Hawksw., Coppins & P.
James 1980 185; B69: 33, 71: 42, 72:
47, 74: 55, 75: 35
Note: Becomes established as an independent lichen,
but evidently parasitic at first.
Lit.: Hawksworth et al. (1980b).
Buellia De Not. 1846
Ascomycota: Lecanorales: Physciaceae.
badia (Fr.) A. Massal. 1853 1546; B72:
47, 70: 38, 79: 39, 87: 78
adjuncta auct. brit. = Buellia pulverea (L)
advenula = Rhizocarpon advenulum
athallina = Dactylospora athallina
bloxamii = Dactylospora bloxamii (F)
epipolia var. murorum = Diplotomma murorum
lignyota = Rhizodiscina lignyota (F)
pulverulenta = Diplotomma pulverulentum
saxatilis = Dactylospora saxatilis
vezdana = Diplotomma vezdanum
Note: British material of B. badia generally has a welldeveloped
autonomous thallus.
Lit.: Hafellner (1979a), andHawksworth (1982a).
Buelliella Fink 1935
Ascomycota: incertae sedis: incertae sedis.
physciicola Poelt & Hafellner 1979 2022;
B 67: 38
Lit.: Hafellner (1979a), and Hawksworth (1990).
Calicium
arenarium = Microcalicium arenarium
corynellum = Calicium corynellum (L)
parietinum = Mycocalicium subtile (F)
retinens = Chaenothecopsis retinens
subtile = Mycocalicium subtile (F)
Calycina
See Pezizella.
Capronia Sacc. 1883
Ascomycota: Chaetothyriales: Herpotrichiellaceae.
normandinae R. Sant. & D. Hawksw.
1990 2023
Lit.: Hawksworth (1990).
Carbonea (Hertel) Hertel 1983
Ascomycota: Lecanorales: Lecanoraceae.
supersparsa (Nyl.) Hertel 1983 706;B82:
44
vitellinaria (Nyl.) Hertel 1983 1880;B82:
44
Lit.: Hertel (1967, 1983), andTriebel (1989).
Catenomycopsis
See Chaenothecopsis.
Catillaria
epiblastematica = Scutula epiblastematica
episema = Toninia episema

204 THE LICHENOLOGIST
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Celidium
dubium = Plectocarpon lichenum
lichenum = Plectocarpon lichenum
squamaricola = uncertain application
varians = Arthonia varians
varians f. parasemoides = Arthonia intexta
varium = Arthonia varians
Cecidonia Triebel & Rambold 1988
Ascomycota: Lecanorales: Lecideaceae.
umbonella (Nyl.) Triebel & Rambold
1988 789
xenophana (Körb.) Triebel & Rambold
1988 1838; B81: 33, 90: 79
Lit.: Hawksworth (1990), Triebel (1989), andTriebel
& Rambold (1988).
Cercidospora Körb. 1865
Ascomycota: incertae sedis: incertae sedis.
cladoniicola Alstrup 1997 2024; B82: 44
decolorella (Nyl.) O. E. Erikss. & J. Z.
Yue 1992 2346
epipolytropa (Mudd) Arnold 1874 2025
lichenicola (Zopf) Hafellner 1987 2026
parva Hafellner & Ihlen 1998 2027
stereocaulorum (Arnold) Hafellner 1987
2028
ulothii Körb. 1865 2029
verrucosaria (Linds.) Arnold 1874 2030
Lit.: Alstrup (1997), Grube & Hafellner (1990),
Hafellner (1987), Hawksworth (1982a, 1990), Ihlen
(1998), andZhurbenko & Triebel (2003).
Chaenothecopsis Vain. 1927
Ascomycota: Mycocaliciales: Mycocaliciaceae.
Anamorphs: Asterophoma, Catenomycopsis,
orPhialophora-like.
epithallina Tibell 1975 1939
parasitaster (Bagl. & Carestía) D.
Hawksw. 1978 1515
pusilla (Flörke) A. F. W. Schmidt 1970
1931
pusiola (Ach.) Vain. 1927 351
retinens (Nyl.) Tibell 1991 1396
rubescens Vain. 1927 1940
subparoica (Nyl.) Tibell 1995 2313;B88:
68
vainioana (Nádv.) Tibell 1979 1833
viridireagens (Nádv.) A. F. W. Schmidt
1970 1942
Note: Asterophoma (Hawksworth 1981; Tibell 1991,
1993) and Catenomycopsis (Tibell & Constantinescu
1991) anamorphs have not been specifically recorded in
Great Britain and Ireland; some species also produce
Phialophora-like anamorphs in culture
Lit.: Hawksworth (1978), Schmidt (1970), Tibell
(1975, 1987, 1991, 1993, 1999), Tibell & Ryman
(1995), Titov & Tibell (1993), and Vainio (1927).
Chionosphaera D. E. Cox 1976
Basidiomycota: Atractiellales: Chionosphaeraceae.
coppinsii P. Roberts 1997 2031
Lit.: Diederich (1996), andRoberts (1997).
Ciliomyces
oropensis = Paranectria oropensis
Clypeococcum D. Hawksw. 1977
Ascomycota: incertae sedis: Dacampiaceae.
cladonema (Wedd.) D. Hawksw. 1977
2032
epicrassum (H. Olivier) Nav.-Ros. & Cl.
Roux 1995 2033
hypocenomycis D. Hawksw. 1980 2034
hypocenomyceae = Clypeococcum hypocenomycis
Lit.: Hawksworth (1977a, 1980b, 1986), Hafellner
(1995), and Hafellner & Navarro-Rosinés (1994).
Conida
abrothallus = uncertain application
clemens auct. brit. = Arthonia apotheciorum
epicladonia = Scutula epicladonia
epiphorbia = Arthonia fuscopurpuea
fuscopurpurea = Arthonia fuscopurpurea
Coniocybe
citrina = Microcalicium arenarium
Coniocybopsis
arenaria = Microcalicium arenarium
Coniosporium
physciae = Xanthoriicola physciae
Coniothecium
graphideorum = Milospium graphideorum
lichenicola = Sclerococcum sphaerale
silaceum = not correctly reported
sphaerale = Sclerococcum sphaerale
Cornutispora Piroz. 1973
Conidial fungi.

2003 Lichenicolous fungi of Great Britain and Ireland—Hawksworth 205
ciliata Kalb 1993 2035; B90: 70
lichenicola D. Hawksw. & B. Sutton
1976 2036
triangularis Diederich & Etayo 1995
2324; B89: 73, 91: 58
Lit.: Etayo & Diederich (1995), Hawksworth (1976,
1981), and Punithalingam (2003).
Corticifraga D. Hawksw. & R. Sant. 1990
Ascomycota: ? Lecanorales: incertae
sedis.
fuckelii (Rehm) D. Hawksw. & R. Sant.
1990 2037; B89: 73
peltigerae (Fuckel) D. Hawksw. & R.
Sant. 1990 2038
Lit.: Hawksworth & Santesson (1990).
Corticiruptor Wedin & Hafellner 1998
Ascomycota: Lecanorales: incertae sedis.
abeloneae (P. M. Jørg.) Wedin &
Hafellner 1998 1915
Lit.: Jørgensen (1969), andWedin & Hafellner (1998).
Corticium
arachnoideum = Athelia arachnoidea
centrifugum = Athelia arachnoidea
Cryptomyces
peltigerae = Corticifraga fuckelii
Cyphelium Ach., 1815
Ascomycota: Lecanorales: Physciaceae
marcianum de Lesd. 1908 1865; B89: 73
sessile (Pers.) Trevis. 1862 1545
stigonellum auct. brit. = Cyphelium sessile
Lit.: Tibell (1971, 1999).
Dacampia A. Massal. 1853
Ascomycota: incertae sedis: Dacampiaceae.
hookeri (Borrer) A. Massal., 1853 2039
rufescentis (Vouaux) D. Hawksw. 1986
2040
Lit.: Hawksworth (1986), and Henssen (1995).
Dacampiosphaeria
rivana = Pyrenidium actinellum
Dactylium
lichenicola = uncertain application
Dactylospora Körb. 1855
Ascomycota: Lecanorales: Dactylosporaceae.
amygdalariae Triebel 1989 2041; B 73:
58, 85: 50
athallina (Müll. Arg.) Hafellner 1979
2042; B79: 40
attendenda (Nyl.) Arnold 1874 2043;
B 73: 58
australis Triebel & Hertel 1989 2044;
B 73: 58
frigida Hafellner 1985 2045
lobariella (Nyl.) Hafellner 1979 2046
microspora Etayo 1991 2047
ophthalamizae Coppins ined. 2000 4
parasitica (Flörke ex Spreng.) Zopf 1896
1973
parellaria (Nyl.) Arnold 1877 2048; B81:
33
purpurascens Triebel 1989 2049; B81:
34
saxatilis (Schaer.) Hafellner 1979 2050;
B 80: 49
lamyi = Opegrapha lamyi
scapanaria 5 (F)
stygia = Dactylospora stygia (F)
urceolata auct. angl. = Dactylospora frigida
Lit.: Etayo (1991), Hafellner (1979a), Hawksworth
(1990, 1994), andTriebel (1989).
Dendrophoma
alcicornaria = Lichenosticta alcicornaria
Didymella
collemata = Didymellopsis collematum
epipolytropa = Cercidospora epipolytropa
pulposi = Didymellopsis pulposi
sphinctrinoides = Zwackhiomyces sphinctrinoides
Didymellopsis (Sacc.) Clem. & Shear
1931
Ascomycota: incertae sedis: Xanthopyreniaceae.
4 Although not yet validly published, this name is
listed as it is in the Society’s mapping scheme database
and has been accorded a reference number.
5 Dactylospora scapanaria (Carrington) D.
Hawksw., comb. nov. (basionym: Lecidea scapanaria
Carrington, Trans. Bot. Soc. Edinb. 7: 382, 1863).

206 THE LICHENOLOGIST
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collematum ( J. Steiner) Grube &
Hafellner 1990 2052
pulposi (Zopf) Grube & Hafellner 1990
2053
gelidaria = Roselliniopsis gelidaria
Lit.: Grube & Hafellner (1990).
Didymosphaeria
epipolytropa = Cercidospora epipolytropa
microstictica = Polycoccum microsticticum
micula auct. angl. = Mycomicrothelia confusa (F)
neottizans = Pyrenidium actinellum
peltigerae = Polycoccum peltigerae
Diederichia D. Hawksw. 2003 6
Conidial fungi.
pseudeverniae (Etayo & Diederich) D.
Hawksw. 2003 2107; B80: 52
Note: The type species of Macrophomina Petr. 1923, M.
phaseolina (Tassi) Goid. 1947, is a plurivorous plant
pathogen producing sclerotia and with thick and
heavily pigmented pycnidial walls. In addition, the
conidiogenous cells are much more elongated and
the conidia more regular in shape (Holliday &
Punithalingam 1970). It is clear that M. pseudeverniae
Etayo & Diederich 1996 is fundamentally di#erent and
a new generic name was therefore necessary for the
lichenicolous species.
Lit.: Etayo & Diederich (1996a).
Diederimyces
See Phaeosphaerobolus.
Diplodina
lichenoides = Crustodiplodia lichenoides (F)
sandstedei = Epicladonia sandstedei
solorinaria = Rhagadostoma lichenicola
vouauxii ? = Lichenodiplis lecanorae
Lit.: Punithalingam (1988).
Diplolaeviopsis Giralt & D. Hawksw.
1991
Conidial fungi.
6 Diederichia D. Hawksw., gen. nov.
Similis generis Phoma Sacc. 1880 sed di#ert in muris
pycnidiis crassis et valde pigmentis, et in conidiis
majiusculis et plerumque irregularis.
Typus: Diederichia pseudeverniae (Etayo &
Diederich) D. Hawksw., comb. nov. (basionym:
Macrophomina pseudeverniae Etayo & Diederich,
Mycotaxon 60: 419, 1996).
ranula Giralt & D. Hawksw. 1991 2283;
B 84: 48
Lit.: Giralt & Hawksworth (1991).
Diploschistes Norman 1853
Ascomycota: Ostropales: Thelotremataceae.
muscorum (Hepp) R. Sant. 1980 494;
B 84: 49
lichenicola = Diploschistes muscorum
Lit.: Friedel (1987), andLumbsch (1989).
Diplotomma Flot. 1850
Ascomycota: Lecanorales: Physciaceae.
murorum (A. Massal.) Coppins 1980 317
pulverulentum (Anzi) D. Hawksw. 2002
1855; B84: 47
vezdanum (P. Scholz & Knoph) Coppins
2002 2339
Lit.: Hafellner & Poelt (1980), Molina et al. (2002),
and Nordin (2000).
Discocera
lichenicola = Lecidea lichenicola (L)
Lit.: Hawksworth (1978).
Discothecium
acervatum = Polycoccum trypethelioides
gemmiferum auct. = Endococcus propinquus
gemmiferum var. brachysporum auct. angl. = uncertain
application
gemmiferum var. calcaricola ? = Endococcus rugulosus
gemmiferum var. physciicola ? = Endococcus parietinarius
physciicola ? = Endococcus parietinarius
squamarioides = Polycoccum squamarioides
stereocaulicola = Polycoccum trypethelioides
stigma auct. = Endococcus perpusillus
vermicularium = Polycoccum vermicularium (not
correctly reported)
Echinodiscus Etayo & Diederich 2000
Ascomycota: ? Leotiales: incertae sedis.
lesdainii (Vouaux) Etayo & Diederich
2000 2294; B86: 46
Lit.: Etayo & Diederich (2000).
Echinothecium
glabrum = Sphaerellothecium araneosum

2003 Lichenicolous fungi of Great Britain and Ireland—Hawksworth 207
Endocarpon
crombiei = Thamnogalla crombiei
Endococcus Nyl. 1855
Ascomycota: ? Dothideales: incertae sedis.
apiciicola Nyl. 1855 2054
brachysporus (Zopf) M. Brand & Diederich
1999 2295; B86: 46
caudisporus J. C. David & Etayo 1995
2055
exerrans Nyl. 1879 2056
macrosporus (Arnold) Nyl. 1878 2296;
B 86: 46
parietinarius (Linds.) Clauzade & Cl.
Roux 1976 2058; B77: 38, 83: 57
perpusillus Nyl. 1857 2059
propinquus (Körb.) D. Hawksw. 1979
2060; B86: 47
rugulosus Nyl., 1855 2061
verrucosporus Alstrup 1994 2297; B86:
47
alpestris = Endococcus apiciicola
alpicola = Endococcus apiciicola
gyrophorarum = Stigmidium gyrophorarum
parietinus = Endococccus parietinarius
stigma auct. = Endococcus perpusillus
triphractoides ? = Phaeospora parasitica
vermicularius = Not correctly reported
Lit.: Alstrup et al. (1994), David & Etayo (1995),
Hawksworth (1979b, 1982a), and Sérusiaux et al.
(1999).
Endophragmiella B. Sutton 1973
Conidial fungi.
hughesii D. Hawksw. 1979 2062
Lit.: Hawksworth (1979a).
Epicladonia D. Hawksw. 1981
Conidial fungi.
sandstedei (Zopf) D. Hawksw. 1981
2063; B77: 38
stenospora (Harm.) D. Hawksw. 1981
2064
Lit.: Hawksworth (1981, 1986).
Epicymatica
thallophila = Anisomeridium biforme (L)
Lit: Hawksworth (1986).
Epilichen Clem. 1909
Ascomycota: Lecanorales: ? Rhizocarpaceae.
scabrosus (Ach.) Clem. 1909 510; B61:
27
Lit: Hafellner (1979a, b).
Eurotium
See Aspergillus.
Everniicola D. Hawksw. 1982
Conidial fungi.
flexispora D. Hawksw. 1982 2065; B79:
40
Lit.: Alstrup & Hawksworth (1990), andHawksworth
(1982a).
Fusarium Link 1809
Conidial fungi.
peltigerae Westend. 1849 2288; B85: 50
kuhnii auct. = ? Athelia arachnoidea
Lit.: Hawksworth (1979a).
Gelatinopsis Rambold & Triebel 1990
Ascomycota: Helotiales: Helotiaceae.
ericetorum (Körb.) Rambold & Triebel
1990 2066; B77: 38, 79: 40
Lit.: Rambold & Triebel (1990).
Geltingia Alstrup & D. Hawksw. 1990
Ascomycota: Ostropales: Odontotremataceae.
associata (Th. Fr.) Alstrup & D.
Hawksw. 1990 1943
groenlandiae = Unguiculariopsis groenlandiae
Lit.: Alstrup & Hawksworth (1990), and Diederich &
Etayo (2000).
Gloeopyrenia
gelatinosa ? = Protothelenella sphinctrinoides (L)
Gongylia
sabuletorum pro parte ? = Arthrorhaphis grisea
viridis = Arthrorhaphis grisea
Guignardia
corniculata = Obryzum corniculatum
fimbriata auct. brit? = Phaeopyxis punctum

208 THE LICHENOLOGIST
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insularis auct. angl. = Verrucaria latericola
olivieri = Telogalla olivieri
psoromoides = Catapyrenium psoromoides (L)
verrucicola f. olivieri = Telogalla olivieri
Lit.: Ho#mann & Hafellner (2000).
Gyalecta
ascaridiella = Spirographa fusisporella
Hainesia Ellis & Sacc. 1884
Conidial fungi.
pertusariae Etayo & Diederich 1996 2292
Lit.: Etayo & Diederich (1996a).
Hawksworthiana U. Braun 1988
Conidial fungi.
peltigericola (D. Hawksw.) U. Braun
1988 2069
Lit.: Braun (1988), andHawksworth (1980a).
Hemigrapha (Müll. Arg.) R. Sant. ex D.
Hawksw. 1975
Ascomycota: incertae sedis: ? Parmulariaceae.
atlantica Diederich & Wedin 2000 2307;
B 87: 80
astericus auct. mult. = Hemigrapha atlantica
Lit.: Diederich & Wedin (2000).
Hobsonia
christiansenii = Illosporiopsis christiansenii
Homostegia Fuckel 1870
Ascomycota: incertae sedis: ? Dacampiaceae.
piggotii (Berk. & Broome) P. Karst.
1873 2072
pelvetii ? = Arthonia fuscopurpurea
Lit.: Hawksworth et al. (2004), and Schæchtelin &
Werner (1928).
Hymenobia Nyl. 1854
Ascomycota: incertae sedis: incertae
sedis.
aporea (Nyl.) D. Hawksw. & O. E.
Erikss. 1992 2073
Lit.: Eriksson & Hawksworth (1991), and Triebel
(1989).
Hyphoderma
roseum = Trichothecium roseum
Illosporiopsis D. Hawksw. 2001
Conidial fungi.
christiansenii (B. L. Brady & D.
Hawksw.) D. Hawksw. 2001 2071; B
88: 70, 89: 75
Lit.: Lowen et al. (1986), andSikaroodi et al. (2001).
Illosporium Mart. 1817
Conidial fungi.
carneum Fr. 1829 2074
coccinum = Phlyctis argena (L)
corallinum = Marchandiomyces corallinus
roseum auct. = Marchandiomyces corallinus
roseum var. corallinum = Marchandiomyces corallinus
Note: I. carneum has been presumed to be the teleomorph
of Pronectria robergei.
Lit.: Hawksworth (1979a), Killian & Werner (1924),
and Sikaroodi et al. (2001).
Intralichen D. Hawksw. & M. S. Cole
2002
Conidial fungi.
christiansenii (D. Hawksw.) D. Hawksw.
& M. S. Cole 2002 2019
lichenum (Diederich) D. Hawksw. &
M. S. Cole 2002 2020; B68: 34
Lit.: Diederich (1990), Hawksworth (1979a, 1994),
and Hawksworth & Cole (2002).
Kalaallia Alstrup & D. Hawksw. 1990
Ascomycota: incertae sedis: Dacampiaceae.
reactiva Alstrup & D. Hawksw. 1990
2075
Lit.: Alstrup & Hawksworth (1990).
Karschia
adjuncta auct. angl. = Buellia pulverea (L)
advenula = Rhizocarpon advenulum
athallina = Dactylospora athallina
bloxamii = Dactylospora bloxamii (F)
destructans = Buellia schaereri (L)
ligynota = Rhizodiscina ligynota (F)
pulverulenta = Diplotomma pulverulentum
saxatilis = Dactylospora saxatilis

2003 Lichenicolous fungi of Great Britain and Ireland—Hawksworth 209
scabrosa = Epilichen scabrosus
stygia = Dactylospora stygia (F)
talcophila = Not correctly reported
thalloides auct. angl. = Uncertain application
thallophila = Amandinea punctata (L)
Lit.: Hafellner (1979a).
Kalchbrenneriella Diederich & M. S.
Christ. 2002
Conidial fungi.
cyanescens (Kalchbr.) Diederich & M. S.
Christ. 2002 2329
Lit.: Diederich (2002).
Karsteniomyces
See Scutula.
Laestaedia
olivieri = Telogalla olivieri
Laeviomyces D. Hawksw. 1981
Conidial fungi.
opegraphae D. Hawksw. 1981 2076
pertusariicola (Nyl.) D. Hawksw. 1981
2077
Lit.: Hawksworth (1981).
Lahmia
fueistingii ? = Arthrorhaphis grisea
plumbina = Toninia plumbina
Lasiosphaeriopsis D. Hawksw. & Sivan.
1980
Ascomycota: Sordariales: Nitschkiaceae.
salisburyi D. Hawksw. & Sivan. 1980
2078; B83: 49
supersparsa (Arnold ex Zopf) Triebel
1989 2079
Lit.: Eriksson & Santesson (1986), Hawksworth
(1980a), andTriebel (1989).
Lauderlindsaya J. C. David & D. Hawksw.
1989
Ascomycota: Verrucariales: Verrucariaceae.
borreri (Tul.) J. C. David & D. Hawksw.
1989 1946
Note: Possibly fertile Normandina pulchella (L); robust
molecular data to establish this may be forthcoming.
Lit.: Aptroot (1998), Diederich & Sérusiaux (1993),
and David & Hawksworth (1989).
Lecidea
alumnula = Cecidonia xenophana
associata = Geltingia associata
cetrariicola ? = Phaeopyxis punctum
citrinella f. arenicola = Arthrorhaphis grisea
cladoniaria ? = Phaeopyxis punctum
dealbatula = Cecidonia xenophana
endocarpicola ? = Toninia verrucarioides
furvella = Rimularia furvella (L)
imponens = Carbonea vitellinaria
inquinans = Micarea inquinans
insidiosa = Ramboldia insidiosa
insularis = Rimularia insularis
leptostigma = Geltingia associata
lichenicola (L)
oxyspora = Nesolechia oxyspora
puncta = Phaeopyxis punctum
scapanaria = Dactylospora scapanaria (F)
supersparsa = Carbonea supersparsa
umbonata auct. angl. = Cecidonia umbonella
umbonella = Cecidonia umbonella
umbonella var. alumnula = Cecidonia xenophana
vitellinaria = Carbonea vitellinaria
watsonii = Lecidea lichenicola (L)
Leciographa
centrífuga = Opegrapha rupestris
inspersa auct. = Dactylospora parasitica
lamyí = Opegrapha lamyi
muscigena = Diplotomma pulverulentum
parasitica = Opegrapha parasitica
parasitica auct. brit. p. max. p. = Opegrapha rupestris
parellaria = Dactylospora parellaria
persimilis ? = Dactylospora scapanaria (F)
scapanaria = Dactylospora scapanaria (F)
urceolata = Dactylospora urceolata
zwackhii = Opegrapha zwackhii (L)
Leightoniomyces D. Hawksw. & B. Sutton
1977
Conidial fungi.
phillipsii (Berk. & Leight.) D. Hawksw. &
B. Sutton 1977 2080
Lit.: Hawksworth (1977a).
Lepraria
nigra = Monodictys cellulosa
Leptosphaeria Ces. & De Not. 1863
Ascomycota: Pleosporales: Leptosphaeriaceae.
clarkii D. Hawksw. 1980 2081
leucomelaria (Mudd) Vouaux 1913 2082
baeomycearia ? = Pyrenidium actinellum
caninae = Pyrenidium actinellum
crozalsii = Not correctly reported

210 THE LICHENOLOGIST
Vol. 35
neottizans = Pyrenidium actinellum
oligospora = Pyrenidium actinellum
pycnostigma auct. angl. = Pyrenidium actinellum
rivana = Pyrenidium actinellum
Note: L. leucomelaria may also be a synonym of Pyrenidium
actinellum.
Lit.: Crane & Shearer (1991).
Leptosphaerulina McAlpine 1902
Ascomycota: Pleosporales: Pleosporaceae.
peltigerae (Fuckel) Riedl 1969 2083
Lit.: Hawksworth (1980a, 1990).
Lethariicola
See Odontotrema.
Lettauia D. Hawksw. & R. Sant. 1990
Ascomycota: Lecanorales: ? Fuscideaceae.
cladoniicola D. Hawksw. & R. Sant.
1990 2084; B82: 47, 86: 49
Lit.: Hawksworth & Santesson (1990), and Ihlen &
Tønsberg (1996).
Libertiella Speg. & Roum. 1880
Conidial fungi.
Teleomorph: Scutula.
malmedyensis Speg. & Roum. 1880 2085
peltigerae = Libertiella malmedyensis
Note: The microconidial state of Scutula epiblastematica
was regarded as very similar to L. malymedyensis by
Triebel et al. (1997).
Lit.: Hawksworth (1981, 1982a), andHawksworth &
Mia˛dlikowska (1997).
Lichenochora Hafellner 1989
Ascomycota: Phyllachorales: Phyllachoraceae.
aprica Hafellner & Nik. Ho#m. 2000
coarctatae (de Lesd.) Hafellner & F.
Berger 2000 2328; B88: 72
inconspicua Hafellner 1989 2086; B73:
60
lecidellae Boqueras & Nav.-Ros. 1998
2284; B84: 50
mediterraneae Calat., Nav.-Ros. & E.
Calvo 2000 2319; B88: 72, 89: 76
obscuroides (Linds.) Triebel & Rambold
1992 2087
weillii (Werner) Hafellner & R. Sant.
1989 2089; B74: 59
thallina = Lichenochora obscuroides
Lit.: Calatyud et al. (2000), Hafellner (1989), Navarro-
Rosinés et al. (1998), andRambold & Triebel (1992).
Lichenoconium Petr. & Syd. 1927
Conidial fungi.
cargillianum (Linds.) D. Hawksw. 1977
2090
erodens M.S. Christ. & D. Hawksw.
1977 2091
lecanorae (Jaap) D. Hawskw. 1979 2092;
B 67: 39
pyxidatae (Oudem.) Petr. & Syd. 1927
2093; B74: 59
usneae (Anzi) D. Hawksw. 1977 2094
xanthoriae M.S. Christ. 1956 2095; B74:
59
imbricariae = Lichenoconium usneae
jaapii = Lichenoconium usneae
lecanoracearum auct. angl. = Lichenoconium lecanorae
parasiticum = Lichenoconium lecanorae
pertusariicola = Laeviomyces pertusariicola
Lit.: Christiansen (1980), and Hawksworth (1977b,
1979a, 1981).
Lichenodiplis Dyko & D. Hawksw. 1979
Conidial fungi.
lecanorae (Vouaux) Dyko & D. Hawksw.
1979 2096; B67: 39
lichenicola Dyko & D. Hawksw. 1979
2097; B78: 60
Lit.: Hawksworth (1981), and Hawksworth & Dyko
(1979).
Lichenopeltella Höhn. 1918
Ascomycota: Microthyriales: Microthyriaceae.
cetrariicola (Nyl.) R. Sant. 1989 2098
coppinsii Earl.-Benn. & D. Hawksw.
1999 2289
peltigericola (D. Hawksw.) R. Sant.
1993 2099; B88: 72

2003 Lichenicolous fungi of Great Britain and Ireland—Hawksworth 211
ramalinae Etayo & Diederich 1997 2181;
B 85: 52
Lit.: Aptroot et al. (1997), Earland-Bennett &
Hawksworth (1999a), Hawkworth (1980b, 1982),
Matzer (1996), andSpooner & Kirk (1990).
Lichenopuccinia D. Hawksw. & Hafellner
1984
Conidial fungi.
poeltii D. Hawksw. & Hafellner 1984
2100
Lit.: Hawksworth (1984, 1990).
Lichenomyces
lichenum = Plectocarpon lichenum
Lichenosticta Zopf 1898
Conidial fungi.
alcicornaria (Linds.) D. Hawksw. 1980
2101
podetiicola = Lichenosticta alcicornaria
Lit.: Hawksworth (1981).
Lichenostigma Hafellner 1983
Ascomycota: ? Dothideales: Lichenotheliaceae.
elongata Nav.-Ros. & Hafellner 1996
2102; B81: 36
maureri Hafellner 1983 2103
rugosa G. Thor 1985 2104
Lit.: Calatayud et al. (2002), Hafellner (1983),
Hawksworth (1986, 1990), Navarro-Rosinés &
Hafellner (1996), and Thor (1985).
Lithographa
andrewii = Geltingia associata
Llimoniella
groenlandiae = Unguiculariopsis groenlandiae
neglecta = Rhymbocarpus neglectus
pubescens = Rhymbocarpus pubescens
Lit.: Diederich & Etayo (2000).
Lophothelium
acervatum = Polycoccum trypethelioides
Macrophomina
pseudeverniae = Diederichia pseudeverniae
Marchandiobasidium
See Marchandiomyces.
Marchandiomyces Diederich & D.
Hawksw. 1990
Conidial fungi.
Teleomorph: Marchandiobasidium.
aurantiacus (Lasch) Diederich & Etayo
1996 2108
corallinus (Roberge) Diederich & D.
Hawksw. 1990 2109
Note: The teleomorph of M. aurantiacus, Marchandiobasidium
aurantiacum Diederich et al. 2003
(Basidiomycota: Ceratobasidiales) has not yet been
recorded in the British Isles but may have been
overlooked.
Lit.: Diederich (1990), Diederich et al. (2003), Etayo &
Diederich (1996a), andHawksworth (1979a).
Massaria
scoriadea = uncertain application
Melaspilea Nyl. 1857
Ascomycota: ? Arthoniales: Melaspileaceae.
diplasiospora auct. brit. 866
leciographiodes Vouaux 1913 1948
lentiginosa (Lyell ex Leight.) Müll. Arg.
1887 1554; B72: 49
lentiginosula (F)
proximella (F)
Note: The British fungus named as M. diplasiospora
(Nyl.) Müll. Arg. 1887 occurs in apothecia of Graphis
elegans and requires a new scientific name (B. J.
Coppins, in litt.).
Lit.: Purvis et al. (1992).
Melittiosporium
lichenicola = Diploschistes muscorum
Merismatium Zopf 1898
Ascomycota: Verrucariales: Verrucariaceae.
deminutum (Arnold) Cl. Roux & Nav.
Ros. 2002 1152
discrepans (J. Lahm) Triebel 1989 1154
nigritellum (Nyl.) Vouaux 1913 2110
lopadii = Merismatium discrepans
Lit.: Roux et al. (2002), andTriebel (1989).
Metasphaeria
cetrariicola = Lichenopeltella cetrariicola
stereocaulorum = Cercidospora stereocaulorum
tartarina = Sagediopsis campsteriana

212 THE LICHENOLOGIST
Vol. 35
Micarea Fr. 1825
Ascomycota: Lecanorales: Micareaceae.
inquinans (Tul.) Coppins 1992 1877
Lit.: Rambold & Triebel (1992).
Microcalicium Vain. 1927
Ascomycota: incertae sedis: Microcaliciaceae.
arenarium (Hampe ex A. Massal.) Tibell
1978 1951; B79: 43, 87: 81, 91: 59
subpedicellatum = Microcalicium disseminatum
Note: The distinctive pycnidial anamorphs are not named
independently, and are described in Hawksworth (1981).
Lit.: Hawksworth (1981), andTibell (1978).
Microdiplodia
ferruginea = Lichenodiplis lecanorae
lecanorae = Lichenodiplis lecanorae
Micropeltopsis
cetrariicola = Lichenopeltella cetrariicola
Microthelia
cookei = Muellerella pygmaea var. athallina
dispora = Polycoccum dzieduszyckii
dispora f. octospora = Polycoccum marmoratum
dissepta ? = Weddellomyces periphericus
dzieduszyckii = Polycoccum dzieduszyckii
exerrans = Endococcus exerrans
marmorata = Polycoccum marmoratum
Lit.: Hawksworth (1985), andTriebel (1989).
Milospium D. Hawksw. 1975
Conidial fungi.
graphideorum (Nyl.) D. Hawksw. 1975
892; B81: 35
Lit.: Hawksworth (1975b, 1979a, 1984).
Minutophoma D. Hawksw. 1981
Conidial fungi.
chrysophthalmae D. Hawksw. 1981 2111
Lit.: Hawksworth (1981).
Monodictys S. Hughes 1958
Conidial fungi.
anaptychiae (Lindau) D. Hawksw. 1975
2112
cellulosa S. Hughes 1958 2113
fuliginosa Etayo 1996 2114
lepraria = Monodictys cellulosa
Note: Monodictys anaptychiae and M. fuliginosa are probably
not congeneric with M. cellulosa nor the type
species of the genus, the non-lichenicolous M. putredinis
(Wallr.) S. Hughes 1958 which occurs on rotten
wood. See Laundon (1992) on the name M. cellulosa.
Lit.: Ellis (1976), Etayo & Diederich (1996b),
Hawksworth (1975b, 1979a, 1990), Laundon (1992),
Rao & de Hoog (1986), andWedin (1993).
Muellerella Hepp 1862
Ascomycota: Verrucariales: Verrucariaceae.
hospitans Stizenb. 1863 2115
lichenicola (Sommerf.) D. Hawksw.
1979 2116
polyspora Hepp ex Müll. Arg. 1862 2117
pygmaea (Körb.) D. Hawskw. 1979
2118
var. athallina (Müll. Arg.) Triebel 1989
2119
ventosicola (Mudd) D. Hawksw. 2003 7
2120
atricola ? = Muellerella lichenicola
haplotella = Muellerella polyspora
opegraphicola = Muellerella polyspora
pygmaea var. ventosicola = Muellerella ventosicola
Note: This generic name is often attributed to ‘Hepp ex
Müll. Arg.’, but Hepp provided the generic name to
Müller Argoviensis and this is a case of ‘in’ rather than
‘ex’ in the sense of Art. 46 Note 1 of the Code.
Muellerella ventosicola di#ers from M. pygmaea in the
ornamented ascospores (Matzer 1993b).
Lit.: Hawksworth (1975, 1979b, 1982a), Matzer
(1993b), andTriebel (1989).
Mycobacidia
arenicola = Arthrorhaphis grisea
flavovirescens = Arthrorhaphis citrinella
plumbina = Toninia plumbina
vermifera = Spirographa fusisporella
Mycobilimbia
endocarpicola ? = Toninia verrucarioides
killiasii = Bacidia killiasii
7 Muellerella ventosicola (Mudd) D. Hawksw.,
comb. nov. (basionym: Microthelia ventosicola Mudd,
Man. Br. Lich.: 307, 1861).

2003 Lichenicolous fungi of Great Britain and Ireland—Hawksworth 213
Mycocalicium
parietinum = Mycocalicium subtile (F)
Mycomelaspilea
leciographoides = Melaspilea leciographoides
Mycosphaerella
cookei = Muellerella lichenicola
Myxophora Döbbeler & Poelt 1978
Ascomycota: ? Dothideales: Pseudoperisporiaceae.
leptogiophila (Minks ex G. Winter) Nik.
Ho#m. & Hafellner 2000 2305
Lit.: Ho#mann & Hafellner (2000).
Nanostictis M.S. Christ. 1954
Ascomycota: Ostropales: Stictidaceae.
christiansenii Etayo 1996 2170; B 86:
50, 87: 82
Lit.: Alstrup (1985), Etayo & Diederich (1996b), and
Christiansen (1954)
Nectria
epicallopisma = records uncertain
hirta = Trichonectria hirta
indigens = Nectriopsis indigens
insidiosa = Hymenobia aporea
lecanodes = Nectriopsis lecanodes
parmeliae = Nectriopsis parmeliae
peltigerae = Pronectria robergei
rubifaciens = Nectriopsis rubifaciens
robergei = Pronectria robergei
santessonii = Pronectria santessonii
Nectriella
anisospora = Pronectria anisospora
robergei = Pronectria robergei
tenacis = Pronectria tenacis
tenuispora = Pronectria tenuispora
tincta auct. angl. = Pronectria santessonii
Nectriopsis Maire 1911
Ascomycota: Hypocreales: Bionectriaceae.
indigens (Arnold) Diederich & Schroers
1999 2123
lecanodes (Ces.) Diederich & Schroers
1999 2124; B89: 77
parmeliae (Berk. & M. A. Curtis) M. S.
Cole & D. Hawskw. 2001 2125
rubefaciens (Ellis & Everh.) M. S. Cole &
D. Hawksw. 2001 2126; B86: 50
Lit.: Cole & Hawksworth (2001), Hawksworth (1986),
Rossman et al. (1999), andSérusiaux et al. (1999).
Neocoleroa
lichenicola subsp. bouteillei = Wentiomyces lichenicola
subsp. bouteillei
Neolamya Theiss. & H. Syd. 1918
Ascomycota: incertae sedis: incertae sedis.
peltigerae (Mont.) Theiss. & H. Syd.
1918 2127
Lit.: Keissler (1930).
Nesolechia A. Massal. 1856
Ascomycota: Lecanorales: Parmeliaceae.
oxyspora (Tul.) A. Massal. 1856 2139
associata = Geltingia associata
cetrariicola ? = Phaeopyxis punctum
cladoniaria ? = Phaeopyxis punctum
insita = Steinia geophana (L)
intumescens = Rimularia insularis
lichenicola = Lecidea lichenicola (L)
neglecta = Rhymbocarpus neglectus
nitschkei = Skyttea nitschkei
puncta = Phaeopyxis punctum
vitellinaria = Carbonea vitellinaria
var. supersparsa = Carbonea supersparsa
Note: Alstrup & Hawksworth (1990) retained this
generic name for N. oxyspora and did not accept its
inclusion within Phacopsis because of di#erences in the
exciple, pigmentation, and ascospore shape. Molecular
studies support this separation (Persˇoh & Rambold
2002). The varieties recognized by Triebel et al. (1995)
are not accepted here following a study of additional
collections.
Lit.: Triebel et al. (1995).
Niesslia Auersw. 1869
Ascomycota: Hypocreales: Niessliaceae.
Anamorph: Monocillium.
cladoniicola D. Hawksw. & W. Gams
1975 2128
lobariae Etayo & Diederich 1996 2129;
B 80: 53
Note: The Monocillium anamorph found in N. cladoniicola
has not been seen in nature or separately named.
Lit.: Etayo & Diederich (1996b), Gams (1971), and
Hawksworth (1975a),

214 THE LICHENOLOGIST
Vol. 35
Nigromacula Etayo 2002
Conidial fungi.
uniseptata (D. Hawksw.) D. Hawksw.
2003 8 2262
Note: This parasite of Hypotrachyna spp. forms cupulate
conidiomata in irregular black patches on the thallus
surface. It di#ers from Vouauxiella in the structure of
the conidiomata, with the conidia all being formed at
one level at the surface, and also in the darker brown
conidia which are thickly 1-septate and distinctly perforate.
The type species of Nigromacula, N. hypotrachynae
Etayo 2002, described from Colombia, is a synonym of
N. uniseptata (isotype studied).
Lit.: Etayo (2002), and Hawksworth (1978).
Obryzum Wallr. 1825
Ascomycota: incertae sedis: Obryzaceae.
corniculatum Wallr. 1825 2130
Lit.: Eriksson (1981), and Ho#mann & Hafellner
(2000).
Odontotrema Nyl. 1858
Ascomycota: Ostropales: Odontotremataceae.
pertusariae Etayo, Diederich & Coppins
2002 2376
Note: Diederich et al. (2002) took up this generic name
for the lichenicolous fungi hitherto referred to Lethariicola
Grummann 1969 (Lumbsch & Hawksworth 1990)
but without examining material of the type species of
Odontotrema. Their decision should be tested by
molecular methods as Odontotrema otherwise contains
species which grow on plant leaves or wood.
Lit.: Diederich et al. (2002).
Omphalina
cupulatoides ? = Arrhenia peltigerina.
Opegrapha Ach. 1809
Ascomycota: Arthoniales: Roccellaceae.
brevis Coppins 1987 1841
glaucomaria (Nyl.) Källsten 1994 1976;
B 80: 53
lamyi (O. J. Rich. ex Nyl.) Triebel 1989
2131
8 Nigromacula uniseptata (D. Hawksw.) D.
Hawksw., comb. nov. (basionym: Vouauxiella uniseptata
D. Hawksw., Notes R. Bot. Gdn Edinb. 36: 195,
1978).
parasitica (A. Massal.) H. Olivier 1906
1842
pertusariicola Coppins & P. James 1979
1843
physciaria (Nyl.) D. Hawksw. & Coppins
1992 1953
pulvinata Rehm 1869 1954
rotunda Hafellner 1994 1067; B77: 40
rupestris Pers., 1794 2132
sphaerophoricola Isbrand & Alstrup
1992 2380
thelotrematis Coppins 1987 1844
zwackhii (A. Massal. ex Zwackh) Källsten
1993 2133
centrífuga = Opegrapha rupestris
lentiginosa = Melaspilea lentiginosa
maculans = Opegrapha glaucomaria
monspeliensis = Opegrapha parasitica
persoonii = Opegrapha rupestris
semicincta = Opegrapha parasitica
Lit.: Atienza (1992), Coppins (1987), Hafellner
(1994), Hawksworth (1980, 1994), Isbrand & Alstrup
(1992), andPurvis et al. (1992).
Ophiobolus
peltigerae = Neolamya peltigerae
Orbicula
tartaricola = Roselliniopsis tartaricola
Paranectria Sacc. 1878
Ascomycota: Hypocreales: Bionectriaceae.
a$nis (Grev.) Sacc. 1878 2134
oropensis (Ces.) D. Hawksw. & Piroz.
1977 2135; B86: 51, 89: 70
subsp. parvispora M. S. Cole & D.
Hawksw. 2001 2377
superba D. Hawksw. 1982 2136
Lit.: Cole & Hawksworth (2001), Hawksworth
(1982a), andRossman et al. (1999).
Perigrapha Hafellner 1996
Ascomycota: Arthoniales: incertae sedis.
superveniens (Nyl.) Hafellner 1996 2021
Lit.: Hafellner (1996a).
Pezizella Fuckel 1870
Ascomycota: Helotiales: Helotiaceae.
epithallina (W. Phillips & Plowr.) Sacc.
1889 2137

2003 Lichenicolous fungi of Great Britain and Ireland—Hawksworth 215
Note: Calycina Nees ex Gray 1821 has been stated to be
an earlier generic name (Baral 1994) andPezizella is
not accepted in Eriksson et al. (2003); Baral is a
co-author of the latter work. Both generic names are
still in current use, but Pezizella contains many more
species and conservation of that name with a di#erent
type is to be preferred.
Lit.: Hawksworth (1980a).
Phacopsis Tul. 1852
Ascomycota: Lecanorales: Parmeliaceae.
huuskonenii Räsänen 1949 2138
oxyspora = Nesolechia oxyspora
var. fusca = Nesolechia oxyspora
Lit.: Hawksworth (1978), Persˇoh & Rambold (2002),
and Triebel et al. (1995).
Phaeopyxis Rambold & Triebel 1990
Ascomycota: Helotiales: incertae sedis.
punctum (A. Massal.) Rambold et al.
1990 1955
varia Coppins et al. 1990 1964
Lit.: Rambold & Triebel (1990).
Phaeospora Hepp ex Stein 1879
Ascomycota: Verrucariales: ? Verrucariaceae.
exoriens (Stirt.) A. L. Sm. 1926 2140
parasitica (Lönnr.) Zopf 1874 2141
rimosicola (Leight. ex Mudd) Hepp ex
Stein 1879 2142
caninae = Pyrenidium actinellum
epicallopisma = Weddellomyces epicallopisma
hetairizans = Pyrenidium hetairizans
parasitica var. dzieduszyckii = Polycoccum dzieduszyckii
parmeliarum = Cucurbidothis pithyophila var. cembrae
(F)
peripherica = Weddellomyces periphericus
supersparsa = Lasiosphaeriopsis supersparsa
vesicularia = incorrectly reported from Great Britain
and Ireland
Lit.: Hawksworth (1975a, 1983a).
Phaeosporobolus D. Hawksw. & Hafellner
1986
Conidial fungi.
Teleomorph: Diederimyces.
alpinus R. Sant. et al. 1990 2143;B77:41
usneae D. Hawksw. & Hafellner 1986
2144
Note: Diederimyces fuscideae Etayo 1995 was described
as a new genus and species for the teleomorph of P.
alpinus but the connection has not been proved culturally
and the fungus described by Etayo (1995) has not
yet been found in the British Isles.
Lit.: Alstrup & Hawksworth (1990), Hawksworth
(1990), andHawksworth & Hafellner (1986).
Pharcidia
aggregata = Stigmidium aggregatum
allogena = Arthopyrenia allogena
conoidea var. solorinaria = Stigmidium solorinarium
consociata = uncertain application
crombiei = Thamnogalla crombiei
dispersa = Zwackhiomyces dispersus
dubiella = Sphaerulina dubiella
ephebes = Stigmidium ephebes
epicymatica = Stigmidium congestum
gyrophorarum = Stigmidium gyrophorarum
lichenicola = Muellerella lichenicola
microspila = Stigmidium microspilum
punctilla = Stigmidium punctillum
superposita = Stigmidium superpositum
triphractoides ? = Phaeospora parasitica
Phoma Sacc. 1880
Conidial fungi.
cytospora (Vouaux) D. Hawksw. 1976
2146; B90: 84
everniae D. Hawskw. 1994 2147
lobariae Diederich & Etayo 1995 1984;
B 84: 51
physciicola Keissl. 1911 2150; B78: 62,
89: 70
abietinae = Lecanactis abietina (L)
lecanorae ? = Opegrapha sp. (L)
lichenis ? = Physconia distorta (L)
parmeliarum = Vouauxiomyces sp.
ramalinae = Vouauxiomyces ramalinae
truncata = Vouauxiomyces truncatus
uncialicola = Bachmanniomyces uncialicola
usneae = Pseudoseptoria usneae
Lit.: Etayo & Diderich (1995), Hawksworth (1976,
1981, 1994).
Phragmonaevia
fuckelii = Corticifraga fuckelii
peltigerae = Corticifraga peltigerae
Phragmothyrium
cetrariicola = Lichenopeltella cetrariicola

216 THE LICHENOLOGIST
Vol. 35
Phyllosticta
cytospora = Phoma cytospora
Physalospora
fimbriatae = Lichenosticta alcicornaria
psoromoides = Catapyrenium psoromoides (L)
Lit.: Ho#mann & Hafellner (2000).
Plectocarpon Fée 1825
Ascomycota: Arthoniales: Roccel
laceae.
lichenum (Sommerf.) D. Hawksw. 1984
2153; B88: 74
sampaianae Diederich & Etayo 1994
2154; B77: 41
scrobiculatae Diederich & Etayo 1994
2155; B84: 52
Lit.: Diederich & Etayo (1994), andHawksworth &
Galloway (1984).
Pleospilis
ascaridiella = Spirographa fusisporella
Pleospora Rabenh. ex Ces. & De Not.
1863
Ascomycota: Pleosporales: Pleosporaceae.
leptogiicola D. Hawksw. 1975 2156
addubitans = uncertain appliccation (F)
hookeri = Dacampia hookeri
peripherica = Weddellomyces periphericus
Note: The generic placement of P. leptogiicola needs
re-investigation.
Lit.: Hawksworth (1975a).
Polyblastia
armericola = Launderlindsaya borreri
deminutum = Merismatium deminutum
nigritella = Merismatium nigritellum
Polycarpella
cookei = Muellerella lichenicola
Polycoccum Saut. ex Körb. 1865
Ascomycota: incertae sedis: Dacampiaceae.
arnoldii (Hepp) D. Hawksw. 1979 2157;
B 90: 85
crassum Vězda 1970 2158
dzieduszyckii (Boberski) D. Hawksw.
1980 2159
kerneri J. Steiner 1893 2160
marmoratum (Kremp.) D. Hawksw.
1980 2161
microcarpon Diederich & Etayo 1998
2162
microsticticum (Leight.) Arnold 1891
1483; B78: 62,79:45
opulentum (Th. Fr. & Almq.) Arnold
1874 2163
peltigerae (Fuckel) Vězda 1969 2164
pulvinatum (Eitner) R. Sant. 1993 2165
slaptoniense D. Hawksw. 1994 2166
sporastatiae (Anzi) Arnold 1874 2167
squamarioides (Mudd) Arnold 1874
2168
trypethelioides (Th. Fr.) R. Sant. 1960
2169
dannenbergii auct. angl. ? = Roselliniella atlantica
p. p. and Peridiothelia fuliguncta p. p. (F)
epicrassum = Clypeococcum epicrassum
galligenum = Polycoccum pulvinatum
gelidaria = Roselliniopsis gelidaria
vermicularium = Not correctly reported
Note: The name P. opulentum has been applied to
several di#erent species, and the identity of the material
from the British Isles requires a reassessment (Atienza
et al. 2003, Navarro-Rosinés & Roux 1990).
Lit.: Atienza et al. (2003), Diederich & Etayo (1998),
Hawksworth (1978, 1994), Hawksworth & Diederich
(1988), and Vězda (1970).
Pronectria Clem. 1911
Ascomycota: Hypocreales: Bionectriaceae.
Anamorph: Acremonium or ? Illosporium.
anisospora (Lowen) Lowen 1990 2171
Anamorph: Acremonium pedatum.
echinulata Lowen 1999 2290; B87: 84
fissuriprodiens Etayo 1996 2172
pertusariicola Lowen 1999 2298; B 86:
53, 87: 84, 89: 70
robergei (Mont. & Desm.) Lowen 1990
2173; B77: 40
santessonii (Lowen & D. Hawksw.)
Lowen 1990 2174
tenacis (Vouaux) Lowen 1990 2175
tenuispora (D. Hawksw.) Lowen 1990
2176

2003 Lichenicolous fungi of Great Britain and Ireland—Hawksworth 217
xanthoriae Lowen & Diederich 1990
2177
streimannii = Xenonectriella streimannii
Note: Illosporium carneum has often been considered the
anamorph of P. robergei but the connection has not been
proved by ascospore cultures. Acremonium spp. usually
arise from ascospore cultures and not all those found
have been given separate binomials.
Lit.: Hawksworth (1990, 1994), Lowen (1989),
Lowen & Diederich (1990), Lowen & Hawksworth
(1986), andRossman et al. (1999).
Psammina Sacc. & M. Rousseau ex E.
Bommer & M. Rousseau 1891
Conidial fungi.
inflata Earl.-Benn. & D. Hawksw. 1999
2278
simplex Earl.-Benn. & D. Hawksw. 1999
2279
stipitata D. Hawksw. 1979 2178
Lit.: Earland-Bennett & Hawksworth (1999b), and
Hawksworth (1981).
Pseudoseptoria Speg. 1910
Conidial fungi.
usneae (Vouaux) D. Hawksw. 1981 2151
Note: The British record requires confirmation.
Lit.: Hawksworth (1981).
Pycnopsammina Etayo & Diederich
1995
Conidial fungi.
lobariae Etayo & Diederich 1995 1852;
B 84: 52
Lit.: Earland-Bennett & Hawksworth (1999b), and
Etayo & Diederich (1995).
Pyrenidium Nyl. 1865
Ascomycota: incertae sedis: Dacampiaceae.
actinellum Nyl. 1865 2179
hetairizans (Leight.) D. Hawksw. 1986
2180; B76: 55
Note: Pyrenidium actinellum may be treated too broadly
at present, and eventually require division.
Lit.: Hawksworth (1980a, 1983b, 1986).
Pyreniococcus
exoriens = Phaeospora exoriens
Pyrenopeziza
lettaui = Unguiculariopsis lettaui
thallophila = Unguiculariopsis thallophila
Pyrenulella
endococcoidea = Phaeospora rimosicola
Raciborskiomyces Siemaszko 1925
Ascomycota: ? Dothideales: Pseudoperisporiaceae.
peltigericola (D. Hawksw.) M. E. Barr
1997 2271
Note: Transferred from Wentiomyces because of the
simple and acute peridial setae.
Lit.: Barr (1997), and Hawksworth (1980a).
Ramboldia Kantvilas & Elix 1994
Ascomycota: Lecanorales: Lecanoraceae.
insidiosa (Th. Fr.) Hafellner 1995 1741
Note: A lichenicolous lichen which takes over the
photobiont of its host.
Lit.: Hawksworth (1982a), andPoelt (1974).
Ramularia
peltigericola = Hawksworthiana peltigericola
Refractohilum D. Hawksw. 1977
Conidial fungi.
galligenum D. Hawksw. 1977 2182
pluriseptatum Etayo & Cl. Roux 1997
2183
Lit.: Hawksworth (1977a, 1979a), and Roux et al. (1997).
Rhagadostoma Körb. 1865
Ascomycota: Sordariales: Nitschkiaceae.
lichenicola (De Not.) Keissl. 1930 2184;
B 56: 34, 61: 27
rugosum Nav.- Ros. & Hladun 1994 2185
Lit.: Hawksworth (1980b), Keissler (1930), Navarro-
Rosinés et al. (1999), Navarro-Rosinés & Hladun
(1994), andVězda (1970).
Rhizocarpon Ramond ex DC. 1805
Ascomycota: Lecanorales: Rhizocarpaceae.

218 THE LICHENOLOGIST
Vol. 35
advenulum (Leight.) Hafellner & Poelt
1976 1956
ochrolechiae (Poelt & Nimis) Hafellner
1992 1892
Lit.: Poelt (1985), andPoelt & Hafellner (1982).
Rhizoctonia
See Athelia.
Rhymbocarpus Zopf 1896
Ascomycota: Helotiales: incertae sedis.
cruciatus (Sherwood, D. Hawksw. &
Coppins) Etayo & Diederich 2000
2205; B84: 52
neglectus (Vain.) Diederich & Etayo
2000 2106; B81: 36
pubescens (Etayo & Diederich)
Diederich & Etayo 2000 2302
Lit.: Diederich & Etayo (2000).
Rimularia Nyl. 1868
Ascomycota: Agyriales: Agyriaceae.
insularis (Nyl.) Hertel & Rambold 1985
736; 82 50
Note: The species first behaves as a lichenicolous fungus,
capturing algae from its host (Ríos et al. 2002).
Lit.: Fryday (1999), Hertel (1970), and Hertel &
Rambold (1990).
Roselliniella Vain. 1921
Ascomycota: Sordariales: incertae sedis.
atlantica Matzer & Hafellner 1990 2186
cladoniae (Anzi) Matzer & Hafellner
1990 2187
microthelia (Wallr.) Nik. Ho#m. &
Hafellner 2000 2306; B91: 62
nephromatis (P. Crouan) Matzer &
Hafellner 1990 2189
Lit.: Ho#mann & Hafellner (2000), and Matzer &
Hafellner (1990).
Roselliniopsis Matzer & Hafellner 1990
Ascomycota: Sordariales: incertae sedis.
gelidaria (Mudd) Matzer 1993 2188
tartaricola (Linds.) Matzer 1993 2190
Lit.: Matzer (1993a), andMatzer & Hafellner (1990).
Rosellinula R. Sant. 1986
Ascomycota: ? Dothideales: incertae
sedis.
haplospora (Th. Fr. & Almq. ex Th. Fr.)
R. Sant. 1986 2333; B89: 79, 91: 62
Lit.: Matzer & Hafellner (1990).
Sagediopsis (Sacc.) Vain. 1921
Ascomycota: Verrucariales: Adelococcaceae.
aquatica (Stein) Triebel 1989 2191; B74:
64, 77: 64
barbara (Th. Fr.) R. Sant. & Triebel
1989 2192; B82: 51, 89: 79
campsteriana (Linds.) D. Hawksw. &
R. Sant. 1990 2193; B88: 76
lomnitzensis (Stein) Orange 2002 2013
tartarina = Sagediopsis campesteriana
Note: The subgenus Hawksworthiella Hafellner 1993
may merit generic status.
Lit.: Alstrup & Hawksworth (1990), Hafellner (1993),
Orange (2002), andTriebel (1989, 1993).
Sarcopyrenia Nyl. 1858
Ascomycota: ? Sordariales: incertae
sedis.
beckhausiana (J. Lahm) M. B. Aguirre
et al. 1990 1985; B84: 52
cylindrospora (P. Crouan) M. B. Aguirre
1990 1847; B73: 63
gibba Nyl. 1857 1307; B52: 37, 65: 27,
66: 29, 74: 64, 79: 47, 82: 51, 91: 62
Lit.: Aguirre-Hudson (1991), Navarro-Rosinés &
Hladun (1990), and Tretiach & Navarro-Rosinés
(1996).
Sclerococcum Fr. 1819
Conidial fungi.
montagnei Hafellner 1996 2194; B 81:
38, 89: 80
normandinae Diederich & Etayo 1995
2291; B87: 85
simplex D. Hawksw. 1979 2195
sphaerale (Ach.) Fr. 1825 1848
Lit.: Etayo & Calatayud (1998), Etayo & Diederich
(1995), Hafellner (1996b), and Hawksworth (1975a,
1979a).

2003 Lichenicolous fungi of Great Britain and Ireland—Hawksworth 219
Scutula Tul. 1852
Ascomycota: Lecanorales: Micareaceae.
Anamorphs: Karsteniomyces and Libertiella.
aggregata Bagl. & Carestía 1889 2196
cristata (Leight.) Sacc. & D. Sacc. 1906
2197
epiblastematica (Wallr.) Rehm 1890 2198
Anamorph: Libertiella a#. malymedyensis.
epicladonia (Nyl.) Zopf 1906 2199
krempelhuberi Körb. 1865 2200; B85:
57
miliaris (Wallr.) Trevis. 1853 2201
Anamorph: Karsteniomyces peltigerae.
solorinaria (Nyl.) P. Karst. 1885 2202;
B 78: 64
stereocaulorum (Anzi) Körb. 1865 2203
episema = Toninia episema
peltigerea ? = Scutula epiblastematica
Note: The genus is badly in need of a critical
revision. The Karsteniomyces peltigerae (P. Karst.) D.
Hawksw. 1981 anamorph has not been specifically
reported from Great Britain and Ireland but is to be
expected.
Lit.: Hawksworth (1986), andTriebel et al. (1997).
Sirococcus
lichenicola = Bachmanniomyces uncialicola
Sirothecium
lichenicola = Vouauxiella lichenicola
Skyttea Sherwood, D. Hawksw. & Coppins
1981
Ascomycota: ? Helotiales: incertae sedis.
buelliae Sherwood, D. Hawksw. &
Coppins 1981 2204
caesii Diederich & Etayo 2000 2207
elachistophora (Nyl.) Sherwood & D.
Hawksw. 1981 2206; B72: 51
gregaria Sherwood, D. Hawksw. &
Coppins 1981 2208; B85: 57
lecanorae Diederich & Etayo 2000 2301
nitschkei (Körb.) Sherwood, D. Hawksw.
& Coppins 1981 1342
pyrenulae Diederich, Etayo & Coppins
2000 2300; B89: 80, 91: 62
spinosa D. Hawksw. & Coppins 1982
2209
viridis D. Hawksw. & Coppins 1982 2211
cruciata = Rhymbocarpus cruciatus
lettaui = Unguiculariopsis lettaui
refractiva = Unguiculariopsis refractiva
tephromelarum = Skyttea elachistophora
thallophila = Unguiculariopsis thallophila
Lit.: Diederich & Etayo (2000), Hawksworth (1982a),
and Sherwood et al. (1981).
Skyttella D. Hawksw. & R. Sant. 1988
Ascomycota: ? Helotiales: incertae sedis.
mulleri (Willey) D. Hawksw. & R. Sant.
1988 2212; B72: 51, 78: 64
Lit.: Hawksworth & Santesson (1988).
Sphaerellothecium Zopf 1897
Ascomycota: ? Dothideales: Mycosphaerellaceae.
araneosum (Rehm) Zopf 1897 2213
minutum Hafellner 1993 2214; B73: 63
propinquellum (Nyl.) Cl. Roux & Triebel
1994 2215; B77: 41
Lit.: Hafellner (1993), Hawksworth (1994), Roux &
Triebel (1994), andTriebel (1989).
Sphaeria
cerinaria ? = Lichenodiplis lecanorae
Sphaerulina Sacc. 1878
Ascomycota: ? Dothideales: Mycosphaerellaceae.
dolichotera (Nyl.) Vouaux 1913 2216
dubiella (Nyl.) Keissl. ex Walt. Watson
1948 2217
chlorococca = Lauderlindsaya acroglypta (L)
corniculata = Obryzum corniculatum
endococcoidea = Phaeospora rimosicola
stereocaulorum = Cercidospora stereocaulorum
Note: The position and status of both these species
requires re-investigation.
Sphinctrina Fr. 1825
Ascomycota: Mycocaliciales: Sphinctrinaceae.
anglica Nyl. 1860 1957
leucopoda Nyl. 1860 1958

220 THE LICHENOLOGIST
Vol. 35
tubiformis A. Massal. 1853 1959; B73:
63
turbinata (Pers.) De Not. 1846 1261
gelasinata = Sphinctrina turbinata
kylemorensis = Sphinctrina leucopoda
microcephala = Sphinctrina tubiformis
microcephala auct. angl. = Sphinctrina anglica
pedata = Sphinctrina leucopoda
tubaeformis = Sphinctrina tubiformis
Lit.: Löfgren & Tibell (1979).
Spiloma
Note: S. auratum Sm. 1809 may be the earliest name for
a conidial fungus on Dirina that currently lacks an
appropriate generic name (B. J. Coppins, in litt.).
Spilomela
ascaridiella = Spirographa fusisporella
vermifera = Spirographa fusisporella
Spilomium
graphideorum = Milospium graphideorum
Spirographa Zahlbr. 1903
Ascomycota: ? Helotiales: incertae sedis.
fusisporella (Nyl.) Zahlbr. 1903 2218;
B 86: 55
vinosa Holien & Triebel 1996 2219
Lit.: Hawksworth (1980b), and Holien & Triebel
(1996).
Sporotrichum
lichenicola = ? Lecanocillium lecanii (F)
Stegia
vermicularis = Thamnogalla crombiei
Stigmidium Trevis., 1860
Ascomycota: Dothideales: Mycosphaerellaceae.
aggregatum (Mudd) D. Hawksw. 1975
2220
arthoniae (Arnold) Hafellner 1994 2221;
B 80: 57, 88: 76
congestum (Körb.) Triebel 1991 2222;
B 77: 41
degelii R. Sant. 1993 2223
ephebes (Henssen) D. Hawksw. 1975
2225
epiramalina (Vouaux) Hafellner 1994
2224; 80: 57
fuscatae (Arnold) R. Sant. 1993 2226
gyrophorarum (Arnold) D. Hawksw.
1975 2057
hageniae (Rehm) Hafellner 1988 2227
lecidellae Triebel et al. 1995 2308; B87:
85
leucophlebiae Cl. Roux & Triebel 1994
2228
marinum (Deakin) Swinscow 1965 2229
microspilum (Körb.) D. Hawksw. 1975
1963
mitchellii Cl. Roux & Bricaud 1994 2230
mycobilimbiae Cl. Roux et al. 1994 2231
peltideae (Vain.) R. Sant. 1960 2232
pumilum (Lettau) Matzer & Hafellner
1990 2234; B;80: 57, 89: 80
punctillum (Arnold) D. Hawksw. 1975
2233
rivulorum (Kernst.) Cl. Roux & Nav.
Ros. 1994 1986; B84: 53
solorinarium (Vain.) D. Hawksw. 1983
2235
superpositum (Nyl.) D. Hawksw. 1975
2236
tabacinae (Arnold) Triebel 1989 2237;
80: 57
xanthoparmeliarum Hafellner 1994 2238
allogenum = Arthopyrenia allogena
dispersum = Zwackhiomyces dispersus
schaereri auct. = Stigmidium congestum
Note: Many of the ‘species’ in this genus may be too
narrowly circumscribed and a critical re-evaluation is
required.
Lit.: Hawksworth (1975a, 1983a, 1994), Roux &
Triebel (1994), Santesson (1993b), and Triebel
(1989).
Synaptospora
tartaricola = Roselliniopsis tartaricola
Syzygospora G. W. Martin 1937
Basidiomycota: Tremellales: Syzgosporaceae.
bachmannii Diederich & M. S. Christ.
1996 2239
physciacearum Diederich 1996 2240;
B 82: 51, 91: 63
Lit.: Diederich (1996).

2003 Lichenicolous fungi of Great Britain and Ireland—Hawksworth 221
Taeniolella S. Hughes 1958
Conidial fungi.
beschiana Diederich 1992 2309; B87: 85
delicata M. S. Christ. & D. Hawksw.
1979 2241
phaeophysciae D. Hawksw. 1979 2242;
B 80: 57
punctata M. S. Christ. & D. Hawksw.
1979 2243
rolfii Diederich & Zhurb. 1997 2327
scripta = Taeniolina scripta
Lit.: Diederich (1992), Diederich & Zhurbenko (1991),
and Hawksworth (1979a, 1990).
Taeniolina M. B. Ellis 1976
Conidial fungi.
scripta (P. Karst.) P. M. Kirk 1981 2244
Note: This species also occurs directly on bark and
wood.
Lit.: Ellis (1976).
Telogalla Nik. Ho#m. & Hafellner 2000
Ascomycota: Verrucariales: Verrucariaceae.
olivieri (Vouaux) Nik. Ho#m. &
Hafellner 2000 2068; B90: 86
Lit.: Hawksworth (1975a), andHo#mann & Hafellner
(2000).
Thamnogalla D. Hawksw. 1980
Ascomycota: Ostropales: Odontotremataceae.
crombiei (Mudd) D. Hawksw. 1980 2245
Lit.: Hawksworth (1980b), andHo#mann & Hafellner
(2000).
Thelidium
lacustre = Zwackhiomyces lacustris
Thelocarpon Nyl. 1853
Ascomycota: incertae sedis: Thelocarpaceae.
epibolum (Leight.) G. Salisb. 1953 1398
epibolum var. epithallinum = Thelocarpon epibolum
epithallinum = Thelocarpon epibolum
Lit.: Ahti (1973), andSalisbury (1966).
Tichothecium
calcaricola = Muellerella lichenicola
cerinarium ? = Lichenodiplis lecanorae
erraticum = Muellerella lichenicola
subsp. microphorum ? = Endococus sp.
gemmiferum auct. = Endococcus propinquus
lichenicola = Muellerella lichenicola
perpusillum = Endococcus perpusillus
pygmaeum = Muellerella pygmaea
var. ecatonosporum = Muellerella pygmaea
var. erraticum = Muellerella lichenicola
var. ventosicola = Muellerella ventosicola
rimosicola = Phaeospora rimosicola
rugulosum = Endococcus rugulosus
squamarioides = Polycoccum squamarioides
stigma auct. = Endococcus perpusillus
vermicularium = Endococcus vermicularius (not correctly
reported)
Toninia A. Massal. 1852
Ascomycota: Lecanorales: Ramalinaceae.
episema (Nyl.) Timdal 1991 1904; B80:
57, 85: 57
plumbina (Anzi) Hafellner & Timdal
1991 1907
verrucarioides (Nyl.) Timdal 1991 1418;
B 65: 37
kolax = Toninia verrucarioides
Lit.: Timdal (1991).
Torula
cyanescens = Kalchbrenneriella cyanescens
Tremella Pers. 1794
Basidiomycota: Tremellales: Tremellaceae.
cetrariicola Diederich & Coppins 1996
2246
coppinsii Diederich & G. Marson 1988
1917; B75: 40, 91: 63
hypogymniae Diederich & M.S. Christ.
1996 2247
lichenicola Diederich 1986 2248
lobariacearum Diederich & M.S. Christ.
1996 2249; B82: 51
normandinae Diederich 1996 2250
pertusariae Diederich 1996 2251
phaeographidis Diederich et al. 1996
2252
phaeophysciae Diederich & M.S. Christ.
1996 2253
protoparmeliae Diederich & Coppins
1996 2254
Lit.: Diederich (1996).

222 THE LICHENOLOGIST
Vol. 35
Trichoconis Clem. 1909
Conidial fungi
lichenicola D. Hawksw. 1980 2255
Lit.: Hawksworth (1980a).
Trichonectria Kirschst. 1907
Ascomycota: Hypocreales: Bionectriaceae.
hirta (A. Bloxam) Petch 1937 2256
aculeata = Trichonectria hirta
rubifaciens = Nectriopsis rubifaciens
Lit.: Hawksworth (1978), andRossman et al. (1999).
Trichothecium Link 1809
Conidial fungi.
roseum (Pers.) Link 1809 2257
Note: Probably fortuitously lichenicolous.
Trichothyrina
cetrariicola = Lichenopeltella cetrariicola
Unguiculariopsis Rehm 1909
Ascomycota: Helotiales: Helotiaceae.
groenlandiae (Alstrup & D. Hawksw.)
Etayo & Diederich 2000 2105
lesdainii (Vouaux) Etayo & Diederich
2000 2303; B90: 86
lettaui (Grummann) Coppins 1990 2258;
B 82: 52, 89: 81
manriquei Etayo 1996 1997; B87: 86
refractiva (Coppins) Coppins 1990 2259
thallophila (P. Karst.) W. Y. Zhuang
1988 2260
Lit.: Diederich & Etayo (2000).
Verrucaria Schrad. 1794
Ascomycota: Verrucariales: Verrucariaceae.
aspiciliicola R. Sant. 1984 1478
conturmatula Nyl. 1879 2346
latericola Erichsen 1943 1620; B72: 52,
73: 65, 86: 56
advenula = Phaeospora parasitica
aspiciliae = Verrucaria aspiciliicola
chlorococca = Lauderlindsaya acroglypta (L)
elachistophora = Skyttea elachistophora
fumosaria ? = Endococcus rugulosus
insularis auct. angl. = Verrucaria latericola
larbalestieri = Endococcus rugulosus
Note: Orange (2002) showed that the type of V. conturmatula
is a lichenicolous Verrucaria and not a Polycoccum.
Lit.: Orange (2002), Purvis et al. (1992), and
Zehetleitner (1978).
Vouauxiella Petr. & Syd. 1927
Conidial fungi.
lichenicola (Linds.) Petr. & Syd. 1927 2261
verrucosa (Vouaux) Petr. & Syd. 1927 2263
uniseptata = Nigromacula uniseptata
Lit.: Hawksworth (1981), andMorgan-Jones (1971).
Vouauxiomyces Dyko & D. Hawksw.
1979
Conidial fungi.
Teleomorph: Abrothallus.
ramalinae (Nordin) D. Hawksw. 1981
2264
Teleomorph: A. suecicus.
santessonii D. Hawksw. 1981 2265
Teleomorph: A. cf. parmeliarum.
truncatus (de Lesd.) Dyko & D. Hawksw.
1979 2266
Anamorph: A. microspermus
Note: Not all Abrothallus anamorphs have been given
independent binomials in Vouauxiomyces.
Lit.: Hawksworth (1981), and Hawksworth & Dyko
(1979).
Weddellomyces D. Hawksw. 1986
Ascomycota: incertae sedis: Dacampiaceae.
epicallopisma (Wedd.) D. Hawksw.
1986 2267; B 61: 27
macrosporus D. Hawksw. et al. 1990
2268
periphericus (Taylor) Alstrup & D.
Hawksw. 1990 2269
epicallopismum = Weddellomyces epicallopisma
peripherica = Weddellomyces periphericus
Note: Navarro-Rosinés & Roux (1995) only treated
species with cephalothecoid ascomata here, but there

2003 Lichenicolous fungi of Great Britain and Ireland—Hawksworth 223
are other cases of genera which include cephalothecoid
and non-cephalothecoid species (e.g. Cercophora;
Lundqvist 1972).
Lit.: Caltayud & Navarro-Rosinés (1998), Hawksworth
(1986, 1990), Navarro-Rosinés et al. (2001), and
Navarro-Rosinés & Roux (1995, 1997).
Wentiomyces Koord. 1907
Ascomycota: ? Dothideales: Pseudoperisporiaceae.
lichenicola subsp. bouteillei Cl. Roux
et al. 1994 2270; B76: 58, 77: 42
peltigericola = Raciborskiomyces peltigericola
Note: Barr (1997) took up the later name Neocoleroa
Petr. 1935 for this genus as the type had been reported
as described from sparse and immature material. She
has been followed by Eriksson et al. (2003) but this is
not an acceptable reason to reject a long-established
name as an epitype can be designated to retain its use if
necessary.
Lit.: Hawksworth (1980a), Matzer (1996), andRoux
et al. (1994).
Xanthoriicola D. Hawksw. 1973
Conidial fungi.
physciae (Kalchbr.) D. Hawksw. 1973
2272
Lit.: Hawksworth (1979a), and Hawksworth &
Punithalingam (1973).
Xenonectriella Weese 1919
Ascomycota: Hypocreales: Nectriaceae.
streimannii (S. Y. Kondr., Coppins &
D. J. Galloway) Rossman 1999 2367;
B 91: 56
Note: Perhaps not congeneric with the type species, X.
lutescens (Arnold) Weese 1919, which has muriform
ascospores.
Lit.: Kondratyuk (1996), andRossman et al. (1999).
Xylographa
andrewii = Geltingia associata
Zevadia J. C. David & D. Hawksw. 1995
Conidial fungi.
peroccidentalis J. C. David & D.
Hawksw. 1995 2273
Lit.: David & Hawksworth (1995).
Zwackhiomyces Grube & Hafellner 1990
Ascomycota: incertae sedis: Xanthopyreniaceae.
berengerianus (Arnold) Grube & Triebel
1990 2274
coepulonus (Norman) Grube & R. Sant.
1990 2320; B88: 77
dispersus (J. Lahm ex Körb.) Triebel &
Grube 1990 2275
immersae (Arnold) Grube & Triebel
1990 2276
lacustris (Arnold) Orange 2002 2378
lecanorae (Stein) Nik. Ho#m. 2000 2152
sphinctrinoides (Zwackh) Grube &
Hafellner 1990 2277; B82: 47
Lit.: Grube & Hafellner (1990), Ho#mann & Hafellner
(2000), andOrange (2002).
I am indebted to the help and support of numerous
colleagues around the world in my explorations of
lichenicolous fungi, who have sent specimens,
literature, commented on drafts of my papers, and
often collaborated as co-authors; in particular I wish
to acknowledge in this regard Brian J. Coppins,
Paul Diederich, Ove E. Eriksson, Josef Hafellner,
Eliyathamby Punithalingam, Rolf Santesson, and Brian
C. Sutton, and also the late M. Skytte Christiansen,
Martin Ellis, and Josef Poelt. For specimens from Great
Britain and Ireland, I am most grateful to many
lichenologists and other mycologists, especially Don S.
Chapman, Tom W. Chester, Brian J. Coppins, Howard
F. Fox, Peter M. Earland-Bennett, Vince J. Giavarini,
Oliver L. Gilbert, Christopher J. B. Hitch, George
Salisbury, Mark R. D. Seaward, Pauline B. Topham,
and the late Humphrey J. M. Bowen, Malcolm C.
Clark, Brian W. Fox, and T. ‘Dougal’ V. Swinscow.
I have also greatly benefitted from having various
graduate students and post-doctoral researchers working
alongside me at various times, especially Begoña
Aguirre-Hudson, Vagn Alstrup, Violeta Atienza,
Mariette S. Cole, Rosalind S. Lowen, and Martha A.
Sherwood-Pike. I was further priviledged to have access
to the collections and libraries of the Royal Botanic
Gardens Kew and The Natural History Museum
London through the courtesy of and often with assistance
from the pertinent section heads, including
Reginald W. G. Dennis, Derek A. Reid, David M.
Pegler, and Brian M. Spooner at Kew, and Peter W.
James and Jack R. Laundon in London.
Much of the data that constitutes the background for
this new checklist was compiled while I served at the
International Mycological Institute, where I was continuously
supported by Christine Thatcher and Marilyn

224 THE LICHENOLOGIST
Vol. 35
Rainbow for technical and secretarial support, respectively.
The Institute’s photographers, the late David W.
Fry and his successor Georgina Godwin, assisted with
the colour photographs presented here. During the
preparation of this checklist, I also benefited from
access to the machine-readable checklist maintained on
the British Lichen Society’s website by Jeremy M. Gray.
Paul M. Kirk generously also assisted in the determination
and checking of author citations of scientific
names, and Brian J. Coppins also noted various points
in proof.
The final text was completed while I was in receipt of
an award under the Programa Ramón y Cajal of the
Ministerio de Ciencias y Tecnologia of Spain (held at
the Facultad de Farmacia, Universidad Complutense
de Madrid) and with the understanding and constant
support of my wife, Patricia Taylor-Hawksworth.
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