The lichenicolous fungi of Great Britain and Ireland: an overview and annotated checklist

David L. Hawksworth

This list enumerates all lichenized and lichenicolous fungi known from the west coast of South Greenland between 60º10’N and 61º20’N. 744 lichen species and infraspecific taxa, 156 lichenicolous and six allied fungi are treated. New to Greenland are: Absconditella trivialis, Acarospora complanata, A. wahlenbergii, Agonimia gelatinosa, Anema nummularium, Arthonia nephromiaria, Arthrorhaphis aeruginosa, A. olivaceae, Bacidia beckhausii, Bellemerea diamarta, Biatora pullata, Botryolepraria lesdainii, Buellia arnoldii, B. erubescens, B. ocellata, B. thiopoliza, Caloplaca citrina, Cercidospora verrucosaria, Cliostomum pallens, Clypeococcum cetrariae, Dactylospora deminuta, Endococcus verrucisporus, E. verrucosus, Epibryon conductrix, Epicladonia simplex, Fuscidea mollis var. caesioalbescens, Hypogymnia vittata, Lecanora aitema, L. cadubriae, L.campestris, L. carpoides, L. subaurea, Lecidea turgidula, Lecidella anomaloides, Lepraria eburnea chemotype II, L. rigidula, Leucocarpia dictyospo...

A study of the lichenicolous fungi occurring on species of the lichenized genus Peltigera has resulted in six new species: Libertiella curvispora, L. didymospora, L. fennica Alstrup, Polycoccum superficiale, Roselliniella peltigericola, and Zwackhiomyces kiszkianus. A key to the five known species of Libertiella is included. This paper brings the number of fungi known on this host to 87, of which 61 are not known from any other host genus, providing additional evidence for the richness of Peltigera thalli as a host for novel fungi. The possible hypotheses to explain the richness of this host genus for lichenicolous fungi are enumerated; these are not mutually exclusive.

The lichenicolous fungi Abrothallus teloschistis, Ceratobasidium bulbillifaciens, Cladosporium licheniphilum, Cornutispora ciliata, Epicladonia sandstedei, Lichenohendersonia varians, Lichenothelia renobalesiana, Stigmidium clauzadei, Vouauxiella verrucosa, Zwackhiomyces berengerianus are reported for the first time for Ukraine. Descriptions, localities, ecology and distribution of the recorded species are provided.

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. 0024-2829/03/030191+42 $30.00/0 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 2003 The British Lichen Society. Published by Elsevier Ltd. All rights reserved. 192 THE LICHENOLOGIST (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 18571), 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. Vol. 35 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 T 1. Numbers of lichenicolous fungi recorded from Great Britain and Ireland Source Smith (1926) Watson (1948) Hawksworth et al. (1980) Hawksworth (1983a) Present work Accepted species* Increase on previous list (%) 55 148 183 218 403 – 169 24 16 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 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. Vol. 35 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 (Peršoh & 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 parasymbionts3. 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 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 (Rı́os 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, Rı́os & Grube (2000) found that the licheni- 195 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 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 Vol. 35 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 F. 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 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; Vol. 35 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 T 2. Surveys, indices and keys to lichenicolous fungi occurring on particular lichen hosts Host lichens Reference(s) Arthrorhaphis Baeomyces Brigantiaea Cladonia Dibaeis Haematomma Icmadophila Koerberiella Lepraria Lobaria Lobariaceae Omphalina Peltigera Hafellner & Obermayer (1995) Ihlen (1998) Hafellner (1995) Bachmann (1927–28) Ihlen (1998) Kalb et al. (1995) Ihlen (1998) Rambold et al. (1990) Kummerling et al. (1993) Etayo & Diederich (1996b) Wedin & Hafellner (1998) Santesson (1989) Hawksworth (1980b), Hawksworth & Mia˛dlikowska (1997) Kummerling & Alstrup (1992), and Martı́nez Moreno (1999) Martı́nez & Hafellner (1998). Kondratyuk & Galloway (1995), Kondratyuk et al. (1994) Navarro-Rosinés et al. (1994, 1995) Ihlen (1995) Hafellner & Berger (2000) Matzer (1996) Triebel (1989) Molitor & Diederich (1997) Keissler (1930), Oudemans (1918–24), Zopf (1896), Index of Fungi (1940 on) Peltigerales Pseudocyphellaria Squamarina Thamnolia Trapelia Foliicolous lichens Lecideoid lichens Pyrenocarpous aquatic lichens All lichens 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 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 Vol. 35 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; B 87: 76 2003 Lichenicolous fungi of Great Britain and Ireland—Hawksworth cetrariae Kotte 1909 2002 cladoniae R. Sant. & D. Hawksw. 1990 2003; B 82: 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; B 84: 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). 201 alpestris (Zopf) Theiss. & Syd. 1918 777; B 74: 54, 86: 42 interlatens (Arnold) Matzer & Hafellner 1990 2368; B 91: 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; B 86: 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). Acremonium Link 1809 Conidial fungi. Teleomorph: Nectriopsis and Pronectria. lichenicola W. Gams 1971 2374; B 76: 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), and Lowen (1995). Actinopeltis peltigericola = Lichenopeltella peltigericola Adelococcus Theiss. & Syd. 1918 Ascomycota: Verrucariales: Adelococcaceae. Arthonia Ach. 1806 Ascomycota: Arthoniales: Arthoniaceae. almquistii Vain. 1883 771; B 82: 42 amylospora Almq. 1880 1930 apotheciorum (A. Massal.) Almq. 1880 1501 cohabitans Coppins 1989 27; B 72: 47 diploiciae Calat. & Diederich 1995 2009; B 83: 47 epiphyscia Nyl. 1875 122; B 79: 38 fuscopurpurea (Tul.) R. Sant. 1960 775 gelidae R. Sant. 1986 1961 graphidicola Coppins 1989 735 intexta Almq. 1880 1933 invadens Coppins 1989 729; B 66: 27 molendoi (Heufl. ex Frauenf.) R. Sant. 1986 1934; B 88: 67 neglectula Nyl. 1874 2323; B 89: 70 202 THE LICHENOLOGIST peltigerae Th. Fr. 1866 1935; B 84: 46 phaeophysciae Grube & Matzer 1997 1982; B 85: 46 punctella Nyl. 1859 1929; B 86: 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; B 84: 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; B 75: 34 citrinella (Ach.) Poelt 1990 100 grisea Th. Fr. 1860 313; B 67: 38 muddii Obermayer 1994 1923 Vol. 35 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), and Santesson & 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), and Pitt 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. arachnoidea max. p. = Athelia 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 Lit.: Hawksworth (1979a). Bachmanniomyces D. Hawksw. 1981 Conidial fungi. uncialicola (Zopf) D. Hawksw. 1981 2016; B 86: 43 203 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 Lit.: Hawksworth (1981, 1990). Note: British material of B. badia generally has a welldeveloped autonomous thallus. 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 Lit.: Hafellner (1979a), and Hawksworth (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). Blarneya D. Hawksw., Coppins & P. James 1980 Conidial fungi. hibernica D. Hawksw., Coppins & P. James 1980 185; B 69: 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; B 72: 47, 70: 38, 79: 39, 87: 78 Carbonea (Hertel) Hertel 1983 Ascomycota: Lecanorales: Lecanoraceae. supersparsa (Nyl.) Hertel 1983 706; B 82: 44 vitellinaria (Nyl.) Hertel 1983 1880; B 82: 44 Lit.: Hertel (1967, 1983), and Triebel (1989). Catenomycopsis See Chaenothecopsis. Catillaria epiblastematica = Scutula epiblastematica episema = Toninia episema 204 THE LICHENOLOGIST 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; B 81: 33, 90: 79 Lit.: Hawksworth (1990), Triebel (1989), and Triebel & Rambold (1988). Cercidospora Körb. 1865 Ascomycota: incertae sedis: incertae sedis. cladoniicola Alstrup 1997 2024; B 82: 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), and Zhurbenko & Triebel (2003). Chaenothecopsis Vain. 1927 Ascomycota: Mycocaliciales: Mycocaliciaceae. Anamorphs: Asterophoma, Catenomycopsis, or Phialophora-like. epithallina Tibell 1975 1939 parasitaster (Bagl. & Carestı́a) 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; B 88: 68 vainioana (Nádv.) Tibell 1979 1833 viridireagens (Nádv.) A. F. W. Schmidt 1970 1942 Vol. 35 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), and Roberts (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 ciliata Kalb 1993 2035; B 90: 70 lichenicola D. Hawksw. & B. Sutton 1976 2036 triangularis Diederich & Etayo 1995 2324; B 89: 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; B 89: 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), and Wedin & Hafellner (1998). Corticium arachnoideum = Athelia arachnoidea centrifugum = Athelia arachnoidea Cryptomyces peltigerae = Corticifraga fuckelii Cyphelium Ach., 1815 Ascomycota: Lecanorales: Physciaceae marcianum de Lesd. 1908 1865; B 89: 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 205 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; B 79: 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. 20004 parasitica (Flörke ex Spreng.) Zopf 1896 1973 parellaria (Nyl.) Arnold 1877 2048; B 81: 33 purpurascens Triebel 1989 2049; B 81: 34 saxatilis (Schaer.) Hafellner 1979 2050; B 80: 49 lamyi = Opegrapha lamyi scapanaria5 (F) stygia = Dactylospora stygia (F) urceolata auct. angl. = Dactylospora frigida Lit.: Etayo (1991), Hafellner (1979a), Hawksworth (1990, 1994), and Triebel (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 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. 20036 Conidial fungi. pseudeverniae (Etayo & Diederich) D. Hawksw. 2003 2107; B 80: 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. Vol. 35 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), and Lumbsch (1989). Diplotomma Flot. 1850 Ascomycota: Lecanorales: Physciaceae. murorum (A. Massal.) Coppins 1980 317 pulverulentum (Anzi) D. Hawksw. 2002 1855; B 84: 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.: Etayo & Diederich (1996a). Lit.: Hawksworth (1978). Diederimyces See Phaeosphaerobolus. 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) 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). Echinodiscus Etayo & Diederich 2000 Ascomycota: ? Leotiales: incertae sedis. lesdainii (Vouaux) Etayo & Diederich 2000 2294; B 86: 46 Lit.: Etayo & Diederich (2000). Echinothecium glabrum = Sphaerellothecium araneosum 2003 Lichenicolous fungi of Great Britain and Ireland—Hawksworth Endocarpon crombiei = Thamnogalla crombiei Endococcus Nyl. 1855 Ascomycota: ? Dothideales: incertae sedis. apiciicola Nyl. 1855 2054 brachysporus (Zopf) M. Brand & Diederich 1999 2295; B 86: 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; B 77: 38, 83: 57 perpusillus Nyl. 1857 2059 propinquus (Körb.) D. Hawksw. 1979 2060; B 86: 47 rugulosus Nyl., 1855 2061 verrucosporus Alstrup 1994 2297; B 86: 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; B 77: 38 stenospora (Harm.) D. Hawksw. 1981 2064 Lit.: Hawksworth (1981, 1986). Epicymatica thallophila = Anisomeridium biforme (L) Lit: Hawksworth (1986). 207 Epilichen Clem. 1909 Ascomycota: Lecanorales: ? Rhizocarpaceae. scabrosus (Ach.) Clem. 1909 510; B 61: 27 Lit: Hafellner (1979a, b). Eurotium See Aspergillus. Everniicola D. Hawksw. 1982 Conidial fungi. flexispora D. Hawksw. 1982 2065; B 79: 40 Lit.: Alstrup & Hawksworth (1990), and Hawksworth (1982a). Fusarium Link 1809 Conidial fungi. peltigerae Westend. 1849 2288; B 85: 50 kuhnii auct. = ? Athelia arachnoidea Lit.: Hawksworth (1979a). Gelatinopsis Rambold & Triebel 1990 Ascomycota: Helotiales: Helotiaceae. ericetorum (Körb.) Rambold & Triebel 1990 2066; B 77: 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 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 Vol. 35 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), and Sikaroodi 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 Lit.: Braun (1988), and Hawksworth (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). 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; B 68: 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 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; B 83: 49 supersparsa (Arnold ex Zopf) Triebel 1989 2079 209 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 centrı́fuga = Opegrapha rupestris inspersa auct. = Dactylospora parasitica lamyı́ = 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). Lit.: Eriksson & Santesson (1986), Hawksworth (1980a), and Triebel (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. 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 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; B 82: 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), and Hawksworth & Mia˛dlikowska (1997). Lichenochora Hafellner 1989 Ascomycota: Phyllachorales: Phyllachoraceae. aprica Hafellner & Nik. Ho#m. 2000 coarctatae (de Lesd.) Hafellner & F. Berger 2000 2328; B 88: 72 inconspicua Hafellner 1989 2086; B 73: 60 lecidellae Boqueras & Nav.-Ros. 1998 2284; B 84: 50 mediterraneae Calat., Nav.-Ros. & E. Calvo 2000 2319; B 88: 72, 89: 76 Vol. 35 obscuroides (Linds.) Triebel & Rambold 1992 2087 weillii (Werner) Hafellner & R. Sant. 1989 2089; B 74: 59 thallina = Lichenochora obscuroides Lit.: Calatyud et al. (2000), Hafellner (1989), NavarroRosinés et al. (1998), and Rambold & 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; B 74: 59 usneae (Anzi) D. Hawksw. 1977 2094 xanthoriae M.S. Christ. 1956 2095; B 74: 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; B 67: 39 lichenicola Dyko & D. Hawksw. 1979 2097; B 78: 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; B 88: 72 2003 Lichenicolous fungi of Great Britain and Ireland—Hawksworth ramalinae Etayo & Diederich 1997 2181; B 85: 52 Lit.: Aptroot et al. (1997), Earland-Bennett & Hawksworth (1999a), Hawkworth (1980b, 1982), Matzer (1996), and Spooner & 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; B 81: 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 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, chandiobasidium aurantiacum Diederich et al. (Basidiomycota: Ceratobasidiales) has not yet recorded in the British Isles but may have overlooked. Macrophomina pseudeverniae = Diederichia pseudeverniae Marchandiobasidium See Marchandiomyces. Mar2003 been been Lit.: Diederich (1990), Diederich et al. (2003), Etayo & Diederich (1996a), and Hawksworth (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; B 72: 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 Lit.: Diederich & Etayo (2000). Lophothelium acervatum = Polycoccum trypethelioides 211 lopadii = Merismatium discrepans Lit.: Roux et al. (2002), and Triebel (1989). Metasphaeria cetrariicola = Lichenopeltella cetrariicola stereocaulorum = Cercidospora stereocaulorum tartarina = Sagediopsis campsteriana 212 THE LICHENOLOGIST 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; B 79: 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), and Tibell (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), and Triebel (1989). Milospium D. Hawksw. 1975 Conidial fungi. graphideorum (Nyl.) D. Hawksw. 1975 892; B 81: 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 Vol. 35 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), and Wedin (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. 20037 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), and Triebel (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) Lit.: Cole & Hawksworth (2001), Hawksworth (1986), Rossman et al. (1999), and Sérusiaux et al. (1999). Mycomelaspilea leciographoides = Melaspilea leciographoides Neocoleroa lichenicola subsp. bouteillei = Wentiomyces lichenicola subsp. bouteillei 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; B 89: 77 parmeliae (Berk. & M. A. Curtis) M. S. Cole & D. Hawskw. 2001 2125 rubefaciens (Ellis & Everh.) M. S. Cole & D. Hawksw. 2001 2126; B 86: 50 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 (Peršoh & 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 Nigromacula Etayo 2002 Conidial fungi. uniseptata (D. Hawksw.) D. Hawksw. 20038 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). Vol. 35 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; B 77: 40 rupestris Pers., 1794 2132 sphaerophoricola Isbrand & Alstrup 1992 2380 thelotrematis Coppins 1987 1844 zwackhii (A. Massal. ex Zwackh) Källsten 1993 2133 centrı́fuga = 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), and Purvis 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; B 86: 51, 89: 70 subsp. parvispora M. S. Cole & D. Hawksw. 2001 2377 superba D. Hawksw. 1982 2136 Lit.: Cole & Hawksworth (2001), (1982a), and Rossman et al. (1999). Hawksworth 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 Note: Calycina Nees ex Gray 1821 has been stated to be an earlier generic name (Baral 1994) and Pezizella 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), Peršoh & 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 215 alpinus R. Sant. et al. 1990 2143; B 77: 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), and Hawksworth & 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 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 Phoma Sacc. 1880 Conidial fungi. cytospora (Vouaux) D. Hawksw. 1976 2146; B 90: 84 everniae D. Hawskw. 1994 2147 lobariae Diederich & Etayo 1995 1984; B 84: 51 physciicola Keissl. 1911 2150; B 78: 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). Lit.: Hawksworth (1975a, 1983a). Phaeosporobolus D. Hawksw. & Hafellner 1986 Conidial fungi. Teleomorph: Diederimyces. Phragmonaevia fuckelii = Corticifraga fuckelii peltigerae = Corticifraga peltigerae Phragmothyrium cetrariicola = Lichenopeltella cetrariicola 216 THE LICHENOLOGIST 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; B 88: 74 sampaianae Diederich & Etayo 1994 2154; B 77: 41 scrobiculatae Diederich & Etayo 1994 2155; B 84: 52 Lit.: Diederich & Etayo (1994), and Hawksworth & 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 Vol. 35 kerneri J. Steiner 1893 2160 marmoratum (Kremp.) D. Hawksw. 1980 2161 microcarpon Diederich & Etayo 1998 2162 microsticticum (Leight.) Arnold 1891 1483; B 78: 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; B 87: 84 fissuriprodiens Etayo 1996 2172 pertusariicola Lowen 1999 2298; B 86: 53, 87: 84, 89: 70 robergei (Mont. & Desm.) Lowen 1990 2173; B 77: 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 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), and Rossman 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; B 76: 55 Note: Pyrenidium actinellum may be treated too broadly at present, and eventually require division. 217 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), and Poelt (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), NavarroRosinés et al. (1999), Navarro-Rosinés & Hladun (1994), and Vězda (1970). Rhizocarpon Ramond ex DC. 1805 Ascomycota: Lecanorales: Rhizocarpaceae. 218 THE LICHENOLOGIST advenulum (Leight.) Hafellner & Poelt 1976 1956 ochrolechiae (Poelt & Nimis) Hafellner 1992 1892 Vol. 35 Rosellinula R. Sant. 1986 Ascomycota: ? Dothideales: incertae sedis. haplospora (Th. Fr. & Almq. ex Th. Fr.) R. Sant. 1986 2333; B 89: 79, 91: 62 Lit.: Poelt (1985), and Poelt & Hafellner (1982). Rhizoctonia See Athelia. Rhymbocarpus Zopf 1896 Ascomycota: Helotiales: incertae sedis. cruciatus (Sherwood, D. Hawksw. & Coppins) Etayo & Diederich 2000 2205; B 84: 52 neglectus (Vain.) Diederich & Etayo 2000 2106; B 81: 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 (Rı́os 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; B 91: 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), and Matzer & Hafellner (1990). Lit.: Matzer & Hafellner (1990). Sagediopsis (Sacc.) Vain. 1921 Ascomycota: Verrucariales: Adelococcaceae. aquatica (Stein) Triebel 1989 2191; B 74: 64, 77: 64 barbara (Th. Fr.) R. Sant. & Triebel 1989 2192; B 82: 51, 89: 79 campsteriana (Linds.) D. Hawksw. & R. Sant. 1990 2193; B 88: 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), and Triebel (1989, 1993). Sarcopyrenia Nyl. 1858 Ascomycota: ? Sordariales: incertae sedis. beckhausiana (J. Lahm) M. B. Aguirre et al. 1990 1985; B 84: 52 cylindrospora (P. Crouan) M. B. Aguirre 1990 1847; B 73: 63 gibba Nyl. 1857 1307; B 52: 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; B 87: 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 Scutula Tul. 1852 Ascomycota: Lecanorales: Micareaceae. Anamorphs: Karsteniomyces and Libertiella. aggregata Bagl. & Carestı́a 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; B 85: 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. 219 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; B 72: 51, 78: 64 Lit.: Hawksworth & Santesson (1988). Sphaerellothecium Zopf 1897 Ascomycota: ? Dothideales: Mycosphaerellaceae. araneosum (Rehm) Zopf 1897 2213 minutum Hafellner 1993 2214; B 73: 63 propinquellum (Nyl.) Cl. Roux & Triebel 1994 2215; B 77: 41 Lit.: Hawksworth (1986), and Triebel et al. (1997). Lit.: Hafellner (1993), Hawksworth (1994), Roux & Triebel (1994), and Triebel (1989). Sirococcus lichenicola = Bachmanniomyces uncialicola Sphaeria cerinaria ? = Lichenodiplis lecanorae Sirothecium lichenicola = Vouauxiella lichenicola Sphaerulina Sacc. 1878 Ascomycota: ? Dothideales: Mycosphaerellaceae. dolichotera (Nyl.) Vouaux 1913 2216 dubiella (Nyl.) Keissl. ex Walt. Watson 1948 2217 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; B 72: 51 gregaria Sherwood, D. Hawksw. & Coppins 1981 2208; B 85: 57 lecanorae Diederich & Etayo 2000 2301 nitschkei (Körb.) Sherwood, D. Hawksw. & Coppins 1981 1342 pyrenulae Diederich, Etayo & Coppins 2000 2300; B 89: 80, 91: 62 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 tubiformis A. Massal. 1853 1959; B 73: 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 Vol. 35 fuscatae (Arnold) R. Sant. 1993 2226 gyrophorarum (Arnold) D. Hawksw. 1975 2057 hageniae (Rehm) Hafellner 1988 2227 lecidellae Triebel et al. 1995 2308; B 87: 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; B 84: 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 Taeniolella S. Hughes 1958 Conidial fungi. beschiana Diederich 1992 2309; B 87: 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; B 90: 86 Lit.: Hawksworth (1975a), and Ho#mann & Hafellner (2000). Thamnogalla D. Hawksw. 1980 Ascomycota: Ostropales: Odontotremataceae. crombiei (Mudd) D. Hawksw. 1980 2245 Lit.: Hawksworth (1980b), and Ho#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), and Salisbury (1966). Tichothecium calcaricola = Muellerella lichenicola cerinarium ? = Lichenodiplis lecanorae 221 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; B 80: 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; B 75: 40, 91: 63 hypogymniae Diederich & M.S. Christ. 1996 2247 lichenicola Diederich 1986 2248 lobariacearum Diederich & M.S. Christ. 1996 2249; B 82: 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 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), and Rossman 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; B 90: 86 lettaui (Grummann) Coppins 1990 2258; B 82: 52, 89: 81 manriquei Etayo 1996 1997; B 87: 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; B 72: 52, 73: 65, 86: 56 advenula = Phaeospora parasitica aspiciliae = Verrucaria aspiciliicola chlorococca = Lauderlindsaya acroglypta (L) elachistophora = Skyttea elachistophora fumosaria ? = Endococcus rugulosus Vol. 35 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), and Morgan-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 are other cases of genera which include cephalothecoid and non-cephalothecoid species (e.g. Cercophora; Lundqvist 1972). peroccidentalis J. C. Hawksw. 1995 2273 David 223 & D. Lit.: David & Hawksworth (1995). 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; B 76: 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), and Roux et al. (1994). Xanthoriicola D. Hawksw. 1973 Conidial fungi. physciae (Kalchbr.) D. Hawksw. 1973 2272 Lit.: Hawksworth (1979a), Punithalingam (1973). and Hawksworth & 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), and Rossman et al. (1999). Xylographa andrewii = Geltingia associata Zevadia J. C. David & D. Hawksw. 1995 Conidial fungi. Zwackhiomyces Grube & Hafellner 1990 Ascomycota: incertae sedis: Xanthopyreniaceae. berengerianus (Arnold) Grube & Triebel 1990 2274 coepulonus (Norman) Grube & R. Sant. 1990 2320; B 88: 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; B 82: 47 Lit.: Grube & Hafellner (1990), Ho#mann & Hafellner (2000), and Orange (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 Rainbow for technical and secretarial support, respectively. The Institute’s photographers, the late David W. 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