Lichenicolous Genus Research Articles

Clypeococcum wedinii (Dothideomycetes), a new lichenicolous fungus on Bunodophoron, with an updated key to species of Clypeococcum

AbstractClypeococcum wedinii, a pyrenocarpous ascomycete growing on the lichen genus Bunodophoron in Chile and New Zealand, is described as new to science. It differs from other Clypeococcum species in the combination of the following characteristics: a conspicuous gall formation, scattered, loose to dense stromatic growths sometimes looking like a clypeus, an ascomatal wall composed of both textura intricata and angularis in surface view, a non-amyloid hymenium, an absence of ostiolar filaments, 8-spored asci, and 1-septate ascospores arranged biseriately in the ascus. An updated key to the species of the lichenicolous genus Clypeococcum is provided.

Taxonomic and Phylogenetic Reassessment of Pyrgidium (Mycocaliciales) and Investigation of Ascospore Morphology.

Mycocaliciales comprise non-lichenized either saprotrophic or lichenicolous fungi which occur in temperate and tropical regions. The mazaediate, saprotrophic and monospecific genus, Pyrgidium, is currently assigned to this order, yet the phylogenetic placement of the genus has remained uncertain due to the absence of molecular data. In order to investigate the systematic position of Pyrgidium, two specimens collected in Brazil and Thailand, respectively, were used to generate mtSSU, SSU, LSU and ITS sequences. However, given that most other representatives of this order only have LSU and ITS sequences available, the phylogenetic reconstruction was limited to these two markers. The phylogenetic analyses confirmed placement of the genus within Mycocaliciales, the genus possessing a sister group relationship with the lichenicolous genus Sphinctrina. Detailed morphological descriptions and illustrations are provided, including those for type specimens of the various synonyms subsumed under the hitherto only accepted species, Pyrgidium montellicum (Beltr.) Tibell. The ascospore morphology was investigated using compound and scanning electronic microscopy (SEM). Principal component analysis (PCA) was performed for the ascospore size using PC-ORD 7. The molecular data and re-examination of the type specimens support the monospecific nature of this genus.

Morphological and phylogenetic reassessment of Sclerococcum simplex from China

Sclerococcum is a species rich lichenicolous genus in Ascomycota. However, many species of Sclerococcum have been described based only on morphology. This study provides sequences for the large-subunit ribosomal RNA (LSU), internal transcribed spacer (ITS) and mitochondrial small-subunit ribosomal RNA (mtSSU) for the first time for Sclerococcum simplex, and the first geographical record of the genus from China. Phylogenetic analyses confirmed that S. simplex clustered within Sclerococcum sensu stricto while the genus was recovered as paraphyletic in Dactylosporaceae. Our asexual specimen of Sclerococcum simplex was collected on a corticolous Pertusaria thallus. The taxonomic affinity of Sclerococcum simplex is investigated based on phylogenetic and morphological evidence.

Ajaysinghia dendriscostictae a new genus and species of lichenicolous fungus growing on Dendriscosticta praetextata (Peltigerales: Peltigeraceae) in India

The new commensalistic lichenicolous genus Ajaysinghia , is described, illustrated, and compared with other morphologically similar fungi. The type species, A. dendriscostictae , grows on the thallus of the corticolous lichen Dendriscosticta praetextata in temperate and subalpine regions of Central Himalaya (Uttarakhand, India). It is distinguished from other lichenicolous ascomycetes described so far in having pseudoperithecia growing in clumps making it appear like galls, (0–)1-septate, brown ascospores [(12–)14.2–17.3(–18) × (3–)3.7–4.8(–5) μm, l/b = (2.8–)3.3–4.2(–4.6) μm], with rounded to pointed ends, slightly constricted at the septum and a cyanobacterial host ( Dendriscosticta praetextata ).

<p><strong><em>Kukwaea pubescens gen</em></strong><strong>. et<em> sp. nova </em>(Helotiales, incertae sedis), a new lichenicolous fungus on <em>Cetraria islandica</em>, and a key to the lichenicolous fungi occurring on <em>Cetraria </em>s. str.</strong></p>

A new lichenicolous genus Kukwaea is introduced for a species discovered on Cetraria islandica from coniferous forests in European and Asian parts of Russia. The new fungus is characterized by its cupulate, brown ascomata with grey to blackish disc surrounded by brownish grey hairs, exciple of textura angularis type, with crystals in the lower part, with granulose excipular hairs obtuse at the tips, simple to forked paraphyses, Calycina-type asci, and hyaline, aseptate ascospores. The DNA sequence data confirmed its placement in Helotiales, but the exact affiliation remains open. A worldwide key for lichenicolous fungi occurring on Cetraria s. str. is provided.

The 2018 classification and checklist of lichenicolous fungi, with 2000 non-lichenized, obligately lichenicolous taxa

Lichenicolous fungi represent a highly specialized and successful group of organisms that live exclusively on lichens, most commonly as host-specific parasites, but also as broad-spectrum pathogens, saprotrophs or commensals. We present here the most recent update to the classification of lichenicolous fungi in the Ascomycota and Basidiomycota to genus level, arranged phylogenetically according to published classifications. For each genus, all known lichenicolous taxa (obligately lichenicolous taxa, lichenicolous lichens, and facultatively lichenicolous taxa) are listed, along with information about types, synonyms, pertinent literature and whether or not molecular data are available for any of the listed species. The number of accepted lichenicolous fungi is now 2319, with 2000 obligately lichenicolous species, subspecies or varieties, 257 lichenicolous lichens and 62 facultatively lichenicolous taxa. These species are found in 10 different classes of Fungi (Ascomycota and Basidiomycota), 55 orders, 115 families and 397 genera. The 2319 total taxa is an increase from the 1559 total species reported in the last published catalogue in 2003, and a larger number than the approximately 1800 reported in the most recent online checklist (www.lichenicolous.net) posted in January 2018. Of the total number of taxa, 2219 (96%) are ascomycetes and 100 (4%) are basidiomycetes. Of the 397 genera containing lichenicolous species, c. 50% (198) are entirely lichenicolous. In addition, six families (Abrothallaceae, Adelococcaceae, Cyphobasidiaceae, Obryzaceae, Polycoccaceae, Sarcopyreniaceae) and two orders (Abrothallales, Cyphobasidiales) are entirely lichenicolous. Sequence information is available for lichenicolous species in 128 (32%) of the 397 genera containing lichenicolous species, and in 56 (28%) of the 198 entirely lichenicolous genera. Many species are known from only one host lichen, but it is likely that broader host ecologies will be discovered as new sequence information is obtained from ongoing microbiome studies. Phaeopyxis Rambold & Triebel is considered as a new synonym of Bachmanniomyces D.Hawksw., resulting in five new combinations B. australis (Rambold & Triebel) Diederich & Pino-Bodas (≡ P. australis), B. carniolicus (Arnold) Diederich & Pino-Bodas (≡ Biatora carniolica), B. muscigenae (Alstrup & E.S.Hansen) Diederich & Pino-Bodas (≡ P. muscigenae), B. punctum (A.Massal.) Diederich & Pino-Bodas (≡ Nesolechia punctum) and B. varius (Coppins, Rambold & Triebel) Diederich & Pino-Bodas (≡ P. varia). As a consequence of a phylogenetic analysis including new sequences, Dactylospora Korb. is regarded as a new synonym of Sclerococcum Fr. : Fr., resulting in one new name (S. acarosporicola Ertz & Diederich) and 46 new combinations. Sclerococcaceae Reblova, Unter. & W.Gams is considered as a new synonym of Dactylosporaceae Bellem. & Hafellner. The new Sclerococcum ophthalmizae Coppins is described. Sclerophyton occidentale Herre is lectotypified on the lichenicolous fungus present in the type specimen and becomes a younger synonym of Sclerococcum parasiticum. A replacement name is Arthonia polydactylonis Diederich & Ertz (≡ A. ceracea). Further new combinations are Abrothallus lobariae (Diederich & Etayo) Diederich & Ertz (≡ Phoma lobariae), A. psoromatis (Zhurb. & U. Braun) Diederich & Zhurb. (≡ P. psoromatis), Asteroglobulus pyramidalis (Etayo) Diederich (≡ Cornutispora pyramidalis), Didymocyrtis grumantiana (Zhurb. & Diederich) Zhurb. & Diederich (≡ Phoma grumantiana), Epithamnolia atrolazulina (Etayo) Diederich (≡ Hainesia atrolazulina), Gyalolechia epiplacynthium (Etayo) Diederich (≡ Fulgensia epiplacynthium), Nesolechia doerfeltii (Alstrup & P.Scholz) Diederich (≡ Phacopsis doerfeltii), N. falcispora (Triebel & Rambold) Diederich (≡ P. falcispora), N. oxyspora var. fusca (Triebel & Rambold) Diederich (≡ P. oxyspora var. fusca), Preussia peltigerae (Brackel) Diederich (≡ Sporormiella peltigerae), Scutula curvispora (D.Hawksw. & Miądl.) Diederich (≡ Libertiella curvispora), S. didymospora (D.Hawksw. & Miądl.) Diederich (≡ L. didymospora), Stigmidium haesitans (Nyl.) Diederich (≡ Verrucaria haesitans), and S. parvum (Henssen) Diederich (≡ Pharcidia parvum).

Disentangling Phialophora section Catenulatae: disposition of taxa with pigmented conidiophores and recognition of a new subclass, Sclerococcomycetidae (Eurotiomycetes)

A new genus Rhopalophora is described for Phialophora clavispora, a lignicolous species formerly placed in Phialophora section Catenulatae that possesses pigmented conidiophores, phialides with a single conidiogenous locus that occasionally appear as schizophialides, and clavate, aseptate conidia arranged in chains or sometimes in heads. Sexual morphs are not known for this taxon. Phylogenetic analysis of DNA sequences from five loci (nucSSU, ITS, nucLSU, mitSSU, rpb1 and rpb2) of this and related fungi supports the introduction of a new family, Sclerococcaceae, for which we establish the order Sclerococcales. This order belongs to the new subclass Sclerococcomycetidae, a strongly supported clade within the Eurotiomycetes that is basal to a lineage containing the Chaetothyriomycetidae, Coryneliomycetidae and Eurotiomycetidae. Rhopalophora clavispora fits in this new family and is closely related to an isolate of Fusichalara minuta. The Sclerococcales also encompass marine, lignicolous species of Dactylospora, two species of the lichenicolous genus Sclerococcum, and a lineage comprised of strains from the digestive tracts of Neotropical wood-inhabiting beetles. We confirm that Dactylospora is polyphyletic; the phylogenetic placement of D. parasitica, the generic type, remains unknown.

Hidden diversity of marine borderline lichens and a new order of fungi: Collemopsidiales (Dothideomyceta)

The fungal genus Collemopsidium comprises species that develop so-called borderline lichen symbioses with algae or cyanobacteria. Together with morphologically similar pyrenocarpous fungi it has been assigned to the family Xanthopyreniaceae. The adscription of this family to higher taxonomic ranks remain uncertain. Using sequence data of five nuclear genomic regions (nuLSU, nuSSU, tef1-α, rpb1 and rpb2) and one mitochondrial locus (mtSSU) we found that the studied representatives of this family are placed in the Dothideomyceta, yet relationships with the classes Dothideomycetes and Arthoniomycetes remain uncertain. We describe the new order Collemopsidiales to accommodate the genus Collemopsidium (paraphyletic as currently understood) and the lichenicolous genus Zwackhiomyces. Using five fungal fossils as calibrations points, we infer an age of c. 230 Mya for the crown of Collemopsidiales. Based on two molecular markers, we also provide insight into the global diversity of marine species of the genus Collemopsidium. According to the species delimitation algorithm GMYC, c. 26 putative species exist, far more than the six species recognized hitherto. We have confirmed this result by comparing the two alternative species models by means of Bayes factors, using path sampling and stepping-stone sampling algorithms to estimate the marginal likelihood of each model. Finally, our observations suggest rock-boring ability evolved in parallel in the different lineages within this group of fungi.

Phylogenetic placement of the lichenicolous, anamorphic genus Lichenodiplis and its connection to Muellerella-like teleomorphs

Lichenicolous fungi are a specialized group of taxa which inhabit lichens and develop diverse degrees of specificity and parasitic behaviour towards their hosts. They are recognized only by their phenotypic symptoms and sexual or asexual spore-producing structures on the lichen thalli. Only recently, molecular data and culture dependent approaches have helped in uncovering the species diversity and in verifying the phylogenetic position and anamorph–teleomorph relationships of some taxa. Here, we studied the phylogenetic placement of representative taxa of two lichenicolous genera, the coelomycete Lichenodiplis and the ascomycete Muellerella. We obtained molecular data for three nuclear and mitochondrial loci (28S, 18S, and 16S), both from fresh collected specimens and culture isolates. Our multilocus phylogeny places Lichenodiplis and Muellerella samples in one monophyletic, fully supported clade, sister to Epibryon (Epibryaceae) in Chaetothyriales (Eurotiomycetes). Morphological analyses of axenically cultured fungi show the formation of conidiomata and conidiospores in both Lichenodiplis and Muellerella isolates. We suggest that the species Lichenodiplis lecanorae and Muellerella atricola represent, respectively, the anamorphic and teleomorphic stages of the same fungus and discuss their relationships with the other fungal families in Chaetothyriomycetidae.

Reassessment of the Lichenicolous Hyphomycete GenusTalpapellis

Heuchert, B., Zhurbenko, M. P. & Braun, U. 2014. Reassessment of the lichenicolous hyphomycete genus Talpapellis. — Herzogia 27: 83–92.Based on a re-examination of type material of Talpapellis peltigerae, type species of the monotypic genus Talpapellis, conidiogenesis and structure of conidiophores and conidia have been reassessed, which led to a confirmation of Talpapellis as a separate, recognized lichenicolous genus with superficially Cladosporium-, Fusicladium- and Pseudocladosporium-like morphology. Owing to similar structures of conidiogenous loci and conidial hila as well as the characteristic structure of the walls of conidiophores and conidia, the lichenicolous genus Verrucocladosporium is considered to be morphologically similar to and confusable with Talpapellis, but differs with regard to proliferation of conidiophores and formation of conidiogenous cells. Therefore, Talpapellis and Verrucocladosporium are maintained as separate genera. The descriptions of Talpapellis and T. peltigerae are amended, and the new variety T. peltigerae var. rossica is introduced, illustrated and discussed, supplemented by a key to the recognized taxa of Talpapellis and Verrucocladosporium.

The obligately lichenicolous genus Lichenoconium represents a novel lineage in the Dothideomycetes

Lichenicolous fungi are obligately lichen-associated organisms that have evolved many times throughout the Ascomycota and Basidiomycota. Approximately 20 % of lichenicolous ascomycetes are recognized only from asexual (anamorphic) characteristics, so the phylogenetic position of many groups has never been resolved. Here we present the first molecular phylogeny of Lichenoconium, a genus of strictly asexual, obligately lichenicolous species with broad geographic distributions and diverse host ecologies. We obtained nuclear and mitochondrial rDNA sequences from fungal cultures isolated from four species in the genus, including a new species, Lichenoconium aeruginosum sp. nov., collected in France, Luxembourg and Netherlands. Our multilocus phylogeny supports the monophyly of fungi in the genus Lichenoconium, and places the genus in the Dothideomycetes, an ascomycete class made up mainly of saprobes and plant-associated endophytes and pathogens. There are only a few recognized groups of lichen-formers in the Dothideomycetes, but Lichenoconium is not supported as being closely related to any of these, nor to any other recognized order within the Dothideomycetes. Given that Lichenoconium is but one of over 100 genera of anamorphic lichenicolous fungi, most of which have never been studied phylogenetically, we suggest that asexual lichenicolous fungi may represent novel and evolutionarily significant phylogenetic groups in the Kingdom Fungi.

Phacopsis — A lichenicolous genus of the family Parmeliaceae

SSU nrDNA studies of two representatives of the lichenicolous genus Phacopsis revealed that they belong to the family Parmeliaceae (Lecanorales) and therefore represent lichenicolous lichens with an endokapylic thallus. Because they are the only lichenicolous taxa within this family, it is suggested that their transition from a foliose and/or fruticose precursor lichen might have been due to a unique (or rare) evolutionary one step event(s). Phylogenetic analyses of the ITS nrDNA of the type species of Phacopsis (P. vulpina) and Nesolechia (N. oxyspora = P. oxyspora) could neither confirm nor reject a monophyletic origin of these two Parmeliacean genera, as previously implied by their synonymization. However, it is considered premature to draw nomenclatorial consequences.

THE DISTRIBUTION OF SELECTED DIAGNOSTIC CHARACTERS IN THE LECANORALES

Traditional classification concepts in lichenology are often, but not always, supported by molecular results. Molecular data should be compared and correlated with micro-morphological and ultrastructural information before systematic rearrangements are undertaken. Visualization of the distribution of morphological and other characters in specified groups is considered as a desirable resultper se, but it is also important to discover whether correlating characters are dependent on each other or not; and if not, whether their distribution in a group might support existing classification concepts. A data set for lecanoralean and other lichenized and lichenicolous genera, comprising 90—mostly multi-state—characters was used to store morphological, chemical and ecological data, and to test character correlations. Several examples of such analyses are presented. The following pairs of characters show some degree of dependence: ascospore septation and number per ascus, ascospore wall type and pigmentation, ascospore and epihymenium pigmentation. Several authors postulated that ascus types are good phylogenetic markers. Ascus types have been widely used for classification concepts of the Lecanorales. Two-dimensional correlation queries of ascus types with the following morphologcal characters were made: substratum preference, thallus growth form and ascospore septation. These correlations supply characteristic profiles for the various ascus types, which have to be compared with forthcoming phylogenetic hypotheses based on molecular data.

The Lichenicolous Genus Lettauia in North America

The genus Lettauia is reported in North America from Tennessee, with the species L. santessonii Ihlen & Tensberg sp. nov. and L. cladoniicola D. Hawksw. & R. Sant. The new species Lettauia santessonii, lichenicolous on Loxospora pustulata, is distinct from L. cladoniicola by the shorter and broader ascospores and larger apothecia. During a field trip to eastern North America in 1992, the junior author collected a lichenicolous fungus on Loxosporapustulata (Brodo & W. L. Culb.) R. C. Harris, which proved to represent an undescribed species of Lettauia D. Hawksw. & R. Sant., a recently described genus based on the single species L. cladoniicola (Hawksworth & Santesson 1990). The aim of the present paper is to describe this new species, and to report on the occurrence of L. cladoniicola in North America. Lettauia was recently reported new to North America by Esslinger and Egan (1995). MATERIALS AND METHODS Fungal hyphae were stained blue with lactoglycerol-aniline-blue (LGAB). Amyloid reactions were carried out, after pretreatment with 10% KOH, by a modified Lugol's solution in which H20 was replaced by lactic acid. The scatter diagram is based on the length and width of 40 spores from four apothecia of each of the type specimens of Lettauia cladoniicola and L. santessonii, and plotted in 0.5 Mm increments (i.e., rounded to the nearest 0.5 Mm). Lettauia D. Hawksw. & R. Sant. Bibliotheca Lichenologica 38: 136 (1990). Type species: L. cladoniicola D. Hawksw. & R. Sant. The asci and ascospores ofLettauia suggest affinites with the Fuscidaceae Hafellner (Hawksworth & Santesson 1990), which, with a question mark, has been assigned to the Teloschistales (Eriksson & Hawksworth 1993). LETTAUIA SANTESSONII Ihlen & Tonsberg sp. nov. (FIG. 1-2) Ascomycetes lichenicola. Lettauiae cladoniicolae D. Hawksw. & R. Sant. similis, sed ascosporis brevioribus (14-20 pm versus 18-25 Mm) et latioribus (3-5 ~m versus 2.5-3.0 Mm) et apotheciis majoribus (0.2-0.4 mm diam. versus 0.15-0.20 mm). Ascomata apothecia, 0.2-0.4 mm in diameter, slightly immersed in host thallus, scattered or in groups; disc more or less flat, pale brownish; exciple d stinct, uneven, paler than disc, consisting of radiating hyphae. Hypothecium hyaline. Hymenium hyaline, 37-54 tm high, K/I+ blue; epithecium hyaline to pale orange. Hamathecium formed by paraphyses; paraphyses slightly branched or unbranch d, not anastomosed, up to 2 pm in diameter, with slightly swollen apices; apices up to 2.5 pm. A ci elongate-clavate to broadly clavate, 42-52 x 10-12 pm, upper part of innermost layer and periascal gel K/I+ blue, 6-8 spored. Ascospores + distichously arranged, narrowly ellipsoid to ? cylindric-fusiform, with rounded ends, hyaline, (1-)2-3 septate, (12.5-)14.5-18.7(-20) x (2-)2.9-4.5(-5) pm (length: mean = 16.6, SD = 2.1; width: mean = 3.7, SD = 0.8). TYPE: U.S.A. TENNESSEE. Monroe Co., Cherokee Nat. Forest, E of Tellico, S of Road FR 210, Bald River Falls, on Loxospora pustulata growing on Betula lenta, 16 September 1992, Tensberg 18085 (holotype, BG; isotypes, DUKE, UPS). Lettauia santessonii was found lichenicolous on the thallus of Loxospora pustulata growing corticolous on Betula lenta. The new species clearly belongs in Lettauia, as circumscribed by Hawksworth and Santesson (1990), based on apothecioid ascomata, persistent exciple, simple or branched paraphyses, elongate-clavate ascus with an amyloid reaction in the periascal gel and upper part of the innermost layer, and 1-3 septate, cylindric-fusiform, hyaline ascospores (see Hawksworth & Santesson 1990 for further details). Lettauia santessonii differs from L. cladoniicola by the shorter and broader ascospores (Fig. 2), in L. cladoniicola they are (-17)18.8-24.2(-30) x (1.5-) 2.1-3.3(-4) pm (length: mean = 21.5, SD = 2.7; width: mean = 2.7, SD = 0.6). The new species is further distinct from L. cladoniicola by the broader apothecia (0.15-0.20 mm in diameter in L. cladoniicola) and its host choice (the hosts of L. cladoniicola are given below). The color of the infected parts of the host thallus 0007-2745/96/32-33$0.35/0 This content downloaded from 157.55.39.59 on Sat, 15 Oct 2016 04:35:14 UTC All use subject to http://about.jstor.org/terms 1996] IHLEN & TONSBERG: LETTAUIA IN NORTH AMERICA 33