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Tremella caloplacae (Zahlbr.) Diederich is a species complex including at least nine different species. Here, we formally describe the new species Tremella elegantis, T. nimisiana, T. parietinae, T. pusillae and T. sorediatae. Tremella elegantis induces galls in the hymenium of Rusavskia elegans and forms 2-celled basidia, where cells rarely elongate and sometimes give the appearance of two immature, independent basidia. Tremella nimisiana has small basidiomata (less than 1 mm diam.), narrowly ellipsoid to pyriform 2-celled, occasionally clavate to subcylindrical 3-celled basidia, and grows in the hymenium of Xanthocarpia species. Tremella parietinae is characterized by the exclusive growth in the hymenium of Xanthoria parietina, the broadly fusiform to ellipsoid probasidia, and the subspherical, pyriform or ellipsoid 2(–3)-celled basidia. Tremella pusillae has ellipsoidal probasidia, 2(–3)-celled pyriform or ellipsoidal basidia that sometimes are constricted at the septum, and grows only on Calogaya pusilla. Tremella sorediatae is characterized by inducing galls on the thallus of Rusavskia sorediata and by pyriform to ellipsoid basidia that sometimes are constricted at the septum. Three species are not formally described and are left unnamed as Tremella sp. 13 on Calogaya biatorina, Tremella sp. 14 on Calogaya decipiens and Tremella sp. 15 on Polycauliona sp. Tremella caloplacae in the strict sense is re-circumscribed as a species confined to Variospora species.
The lichenicolous ‘heterobasidiomycetes’ belong in the Tremellomycetes (Agaricomycotina) and in the Pucciniomycotina. In this paper, we provide an introduction and review of these lichenicolous taxa, focusing on recent studies and novelties of their classification, phylogeny and evolution. Lichen-inhabiting fungi in the Pucciniomycotina are represented by only a small number of species included in the genera Chionosphaera, Cyphobasidium and Lichenozyma. The phylogenetic position of the lichenicolous representatives of Chionosphaera has, however, never been investigated by molecular methods. Phylogenetic analyses using the nuclear SSU, ITS, and LSU ribosomal DNA markers reveal that the lichenicolous members of Chionosphaera form a monophyletic group in the Pucciniomycotina, distinct from Chionosphaera and outside the Chionosphaeraceae. The new genus Crittendenia is described to accommodate these lichen-inhabiting species. Crittendenia is characterized by minute synnemata-like basidiomata, the presence of clamp connections and aseptate tubular basidia from which 4–7 spores discharge passively, often in groups. Crittendenia, Cyphobasidium and Lichenozyma are the only lichenicolous lineages known so far in the Pucciniomycotina, whereas Chionosphaera does not include any lichenicolous taxa.
A new species of Synarthonia, S. leproidica, is described from Luxembourg. Phylogenetic analyses of mtSSU and RPB2 sequences were used to determine the generic affiliation of this sterile species. Synarthonia leproidica differs from all other species of the genus by the combination of a leproid thallus and the production of psoromic acid. It is the sister species to S. muriformis in our phylogenetic analyses. The discovery of the new species suggests that other strictly sorediate lichen species might have been overlooked in Europe, even in intensely explored countries such as Luxembourg. Phylogenetic analyses further confirm the placement of Reichlingia anombrophila in the genus Reichlingia and of Synarthonia astroidestera in the genus Synarthonia. Arthonia atlantica is transferred to the genus Reichlingia as R. dendritica.
A diversity of filamentous microfungi was discovered from thallus surfaces of epiphytic lichens preserved in Bitterfeld and Baltic amber. We report seven distinct morphologies of dematiaceous hyphomycetes, some of which closely resemble species of the extant genera Sporidesmium, Taeniolella s. lat. and Taeniolina. Both the placement of the fungi on their substrates and the exquisite preservation of delicate structures indicate that the fungi were fully developed before they were engulfed by fresh resin. The lichens probably grew on the trunks of resin producing trees and became embedded in resin flows together with their fungal associates. The findings demonstrate that a wide range of presumably specialised fungi have lived on living and decomposing lichen thalli at least since the Paleogene. The findings add an interesting new component to the as yet poorly known mycota of the ancient European amber forests.
Two new genera and four new species of bulbil-forming basidiomycetes are described. Phylogenetic analyses of nuLSU and ITS sequences place them in Cantharellales. A facultative lichenicolous species with yellow to orange-yellow bulbils from South America groups with the type of Burgella and is consequently described as B. lutea. The new species and genus Burgellopsis nivea is introduced for material from Scotland with white bulbils overgrowing saxicolous lichens. An obligate lichenicolous species with particularly large, applanate bulbils developing over Peltigerales in South America could not be placed accurately using ITS sequences and is described as the new species and genus Bulbilla applanata. A European species with brown, facultatively lichenicolous bulbils grouped with Ceratobasidium and Thanatephorus species and is described as the new Ceratobasidium bulbillifaciens.
Based on morphological, anatomical, chemical, ecological and molecular evidence, Blarneya is synonymized here with Tylophoron. The molecular phylogeny derived from sequences obtained from sporodochia of Blarneya places this genus, described to accommodate an anamorphic lichen with white cushion-shaped sporodochia, within Tylophoron. This conclusion is further supported by the discovery of Tylophoron-type ascomata emerging directly from thalli with Blarneya-type sporodochia and producing identical hyaline conidia. In one specimen pycnidia were also observed. This represents a surprising variety of morphologically different conidiomata. A different anamorphic type was previously reported from Tylophoron, and this is confirmed here by molecular analysis for T. moderatum: besides thalli with ascomata this species has anamorphic thalli with an irregularly delimited brown sporodochial felt and brown conidia. Ascomata are not known from these entirely anamorphic thalli, whereas they do occur infrequently in Tylophoron species with Blarneya-type sporodochia. A key to all currently accepted species of Tylophoron is provided. In addition to the corticolous Tylophoron hibernicum, confined to humid forests, two saxicolous species with Blarneya-type sporodochia are described here as new: T. galapagoense, known only from Galapagos, differs from T. hibernicum by a thicker, more compact, beige rather than white, more strongly C+ red thallus, growing below sheltered rock overhangs in dry forests; T. stalactiticum has a C− thallus with stipitate, white, C+ red sporodochia; the species is known only from a single locality in Tenerife, on a large slope with volcanic boulders.
The new lichenicolous Llimoniella phaeophysciae, known from Italy, Spain and the USA (Arizona) on Phaeophyscia, resembles Geltingia associata in the subcylindrical asci with uniseriate, shortly ellipsoid ascospores, but is distinguished by several important characters. Gelatinopsis acarosporicola, G. heppiae, Geltingia groenlandiae and Psorotichia terricola (syn. nov. Gelatinopsis leptogii) are considered to be congeneric with L. phaeophysciae, although some of them have elongate clavate and not subcylindrical asci and they are consequently combined in Llimoniella. These five species are distinguished from Llimoniella s.str. by a different excipular and epihymenial pigmentation not reacting with KOH and are treated as the informal L. phaeophysciae group. The new L. placopsidis is described from Placopsis in New Zealand. More complex pigmentation patterns are documented for L. pyrenulae and L. ramalinae. A revised key to the species of Llimoniella is presented. Gelatinopsis ericetorum and G. roccellae are combined in Rhymbocarpus. A redescription of Geltingia associata is given.
The lichenicolous Phoma species on Cladonia are revised. A statistical analysis of the conidial size suggests that three species are involved: P. cladoniicola sp. nov. a common and widespread species in the Northern Hemisphere occurring on a wide range of Cladonia species; P. foliaceiphila sp. nov. known from C. foliacea and C. fimbriata in Europe; and P. grumantiana sp. nov. known from C. symphycarpia and C. mateocyatha in Svalbard and the USA.
Opegrapha species with muriform ascospores are revised and two species, differing only in the shape of their excipulum, are accepted: O. arabica, a widespread coastal species, and O. varians, known from mainland Yemen and Socotra. The genus Dictyographa, previously distinguished from Opegrapha by its muriform ascospores, is reduced into synonymy with Opegrapha. Dictyographa angolensis and D. sandwicensis are new synonyms of O. arabica; the type of D. cinerea was found to be a mixture of different species of the Hysteriaceae; and D. contortuplicata and D. psyllocarpa are excluded from the Arthoniales. Lectotypes are designated for D. angolensis, D. contortuplicata and D. sandwicensis.
The new lichenicolous fungus, Gelatinopsis leptogii, is described from Belgium and Germany, where it grows on Leptogium byssinum. It differs from G. ericetorum, which grows on Dibaeis, in the shorter and broader ascospores and different ascomatal pigments, and from G. roccellae, which grows on Roccella, in the shorter ascospores and larger ascomata.
The new genus Lambinonia Sérus. & Diederich is introduced for the lichenicolous hyphomycete Melanconium strigulae Elenkin & Woron. This species forms dark sporodochia on the surface or at the margin of the thalli, or extruding out of the perithecia and pycnidia of the foliicolous Strigula buxi, S. nemathora and S. nitidula. It is so far known from the western Caucasus (Georgia and Russia), southern Italy, Tenerife (Canary Islands), St-Lucia (West Indies), and Burundi and Rwanda in Africa. The new combination Lambinonia strigulae (Elenkin & Woron.) Sérus. & Diederich is introduced and a neotype collection is designated.
The new lichenicolous hyphomycete, Monodictys epilepraria, is described from thalli of several Lepraria species. It is known from the Czech Republic, Great Britain, Lithuania, Poland, Spain and Sweden, and appears to be widespread, but probably much overlooked. The new species is characterized by relatively small, brown, muriform, smooth-walled conidia developing on decolourized patches of the host.
The anamorphic basidiomycete genus Marchandiomyces presently includes two common lichenicolous (lichen-inhabiting) species, M. corallinus and M. aurantiacus (teleomorph Marchandiobasidium aurantiacum). We describe here a new species, M. lignicola sp. nov., that is similar to M. corallinus in the colour of its sclerotia, but differs in having a wood-inhabiting (lignicolous) habit. The phylogenetic position of this lignicolous fungus was compared with the lichenicolous species of Marchandiomyces and related species currently placed in the basidiomycetous families Corticiaceae and Ceratobasidiaceae using parsimony, likelihood, and Bayesian analyses of complete sequences of the nuclear small subunit and internal transcribed spacers ribosomal DNA, and a portion of the nuclear large subunit ribosomal DNA. These DNA sequences were obtained from isolated cultures of freshly collected specimens. Significant Bayesian posterior probabilities, as well as maximum likelihood and parsimony analyses, indicate that the new lignicolous species is closely related to M. corallinus, the type species of Marchandiomyces. In most analyses these two species are monophyletic with the lichenicolous M. aurantiacus, although this relationship is not strongly supported. Since M. lignicola is more closely related to M. corallinus than to M. aurantiacus, either a transition to the lignicolous habit occurred recently within an ancestral lichenicolous group or, more likely, transition to the lichenicolous habit arose recently and in parallel from an ancestral lignicolous habit.
The name Marchandiobasidium aurantiacum gen. sp. nov. is introduced for the teleomorph of Marchandiomyces aurantiacus. Dolipore septa and septal pore caps of the closely related Marchandiomyces corallinus are typical of the Ceratobasidiales, and the basidiomatal characters of Marchandiobasidium aurantiacum are reminiscent of those of the monotypic genus Waitea. Morphological, ultrastructural and molecular data suggest that Marchandiobasidium should not be included in Waitea, but should be treated as a distinct genus.
The genus Lethariicola is shown to be a synonym of Odontotrema. Fifteen lichenicolous species are recognized: O. bryoriae sp. nov. (on Bryoria), O. cuculare comb. nov. (on Parmeliopsis hyperopta), O. figulinum comb. nov. (on Lecanora polytropa) O. intermedium sp. nov. (on Thamnolia), O. lecanorae sp. nov. (on Lecanora), O. melaneliae sp. nov. (on Melanelia), O. ochrolechiae sp. nov. (on Ochrolechia), O. pertusariae sp. nov. (on Pertusaria hymenea), O. rhizocarpicola sp. nov. (on Rhizocarpon) O. santessonii sp. nov. (on Thamnolia), O. sipei comb. nov. (on Letharia), O. thamnoliae sp. nov. (on Thamnolia), and three undescribed species on Lecanora, Mycoblastus and Usnea. The type of O. cuculare has two distinct lichenicolous species of Odontotrema, and the name is lectotypified on the species growing on Parmeliopsis. A key is given to all lichenicolous species of Odontotrema and Paralethariicola.
The new lichenicolous fungus Gelatinopsis roccellae is described from NW Spain, where it grows on Roccella fuciformis and R. phycopsis. It differs from G. ericewrum, which grows on Dibaeis, by the shorter ascospores, smaller ascomata, and the absence of an olivaceous excipular and epihymenial pigment.
The biological status of Chiodecton epiphyllum (Arthoniales: Roccellaceae), described as a foliicolous lichen from Papua New Guinea, is reassessed, based on additional material from Brazil, Costa Rica, and Papua New Guinea. The species is a lichenicolous fungus growing on the foliicolous lichen Coenogonium flavicans (Gyalectales: Gyalectaceae). Furthermore, its ascomatal anatomy and lichenicolous growth habit suggest placement in the related genus Plectocarpon, instead of Chiodecton. We therefore propose the new combination Plectocarpon epiphyllum [Bas.: Chiodecton epiphyllum]. The species is further lectotypified on its ascomata since parts of the original description which denote vegetative characteristics refer to the sterile thallus of the host lichen.
The genus Skyttea is characterized by urceolate ascomata, with a narrow pore when young, a greenish or brownish, rarely reddish exciple of conglutinate hyphae with subglobose to ± cylindrical lumina, bordered near the margin by hyaline to greenish or brownish, usually smooth, straight hairs, the absence of periphyses, rarely branched and apically hardly thickened paraphyses, asci and a hymenium that are entirely I — and KI —, a thin lateral and a very thick apical ascus wall with a small ocular chamber, and normally aseptate ascospores. It is considered to belong to the Leotiales and not to the Ostropales. It comprises at least 17 species, including S. caesii sp. nov., S. carboneae sp. nov., S. lecanorae sp. nov., S. mayrhoferi sp. nov., S. pyrenulae sp. nov., S. radiatilis comb, nov., S. tavaresae. sp. nov. and S. thelotrematis sp. nov. Rhymbocarpus is shown to be a distinct leotialean genus, distinguished from Llimoniella by a different excipular structure, different pigments and the presence of excipular hairs, which, in some species, are embedded in the excipular gel and thus not visible macroscopically. In addition to the type species, R. geographici, of which a neotype is designated, it includes R. boomii sp. nov.,R. cruciatus comb, nov., R. fuscoatrae comb, nov.,R. makarovae sp. nov.,R. neglectus comb, nov., R. pertusariae sp. nov.,R. pubescens comb. nov. and R. stereocaulorum comb. nov. Llimoniella s. str. comprises at least 6 species, including the type, L. scabridula, L. adnata, L. pertusariae sp. nov., L. pyrenulae sp. nov., L. ramalinae comb. nov. and L. vinosa comb. nov. Several species formerly assigned to Skyttea or Llimoniella are now included in the leotialean genus Unguiculariopsis, and the following new combinations are proposed: Unguiculariopsis acrocordiae comb, nov., U. groenlandiae comb. nov. and U. lesdainii comb. nov. A lectotype is chosen for Lichen stictoceros, which is shown to be a synonym of Evemia prunastri. Keys are given for the species of Skyttea, Llimoniella and Rhymbocarpus, and for the lichenicolous genera of Leotiales and Ostropales.
Twenty species of lichenicolous ascomycetes are recorded for the western Pyrenees. Capronia hypotrachynae Etayo & Diederich (on Hypotrachyna species, also known from the Canary Islands and from Papua New Guinea), Llimoniella pubescens Etayo & Diederich (on Lepraria species, also from Scotland and Papua New Guinea), Polycoccum microcarpum Diederich & Etayo (on Cladonia species, also from Scotland), Skyttea megalosporae Etayo & Diederich (on Megalospora tuberculosa), Sphaerellothecium cinerascens Etayo & Diederich (on Cladonia parasitica) and S. parmeliae Diederich & Etayo (on Parmelia s. str., also from Finland) are described as new.