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Previously described species of the acerosa complex treated here, in alphabetical order. A. Arrhenia acerosa s. str., O-189467, Norway, photo: Arne Aronsen. B-D. Arrhenia glauca. B. Epitype, WU-6564, Austria, photo: Anton Hausknecht. C. Lectotype, Germany, illustration: August Batsch. D. GB-0058855, Sweden, photo: Leif Strindvall. E-F. Arrhenia latispora. E. WU-22359, Austria, photo: Anton Hausknecht. F. Epitype, LIP-0401569, France, photo: Pierre-Arthur Moreau. G-I. Arrhenia subglobisemen. G. DAOM-981251, Canada, NL, photo: Andrus Voitk. H-I. Epitype, BBF-GC15100901, France, photo: Gilles Corriol. J. Arrhenia tillii, holotype, WU-18120, Austria, photo: Anton Hausknecht.

Previously described species of the acerosa complex treated here, in alphabetical order. A. Arrhenia acerosa s. str., O-189467, Norway, photo: Arne Aronsen. B-D. Arrhenia glauca. B. Epitype, WU-6564, Austria, photo: Anton Hausknecht. C. Lectotype, Germany, illustration: August Batsch. D. GB-0058855, Sweden, photo: Leif Strindvall. E-F. Arrhenia latispora. E. WU-22359, Austria, photo: Anton Hausknecht. F. Epitype, LIP-0401569, France, photo: Pierre-Arthur Moreau. G-I. Arrhenia subglobisemen. G. DAOM-981251, Canada, NL, photo: Andrus Voitk. H-I. Epitype, BBF-GC15100901, France, photo: Gilles Corriol. J. Arrhenia tillii, holotype, WU-18120, Austria, photo: Anton Hausknecht.

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A molecular genetic study of the Arrhenia acerosa complex using the ITS fungal barcoding marker revealed unexpected diversity along a cascading group supporting over 20 lineages. Among these, we identified five previously described species: A. acerosa s.str., A. glauca, A. latispora, A. subglobisemen, and Rhodocybe tillii (recombined as A. tillii)....

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... recognized species (in order of first description; more complete treatment under Taxonomy). When Batsch described Agaricus glaucus (Batsch 1786) he did not indicate a type but provided an illustration (Fig. 2C), which we designate below as lectotype. Collection WU-6564 ( Fig. 2B) is a perfect match for this lectotype, and fits the description in Batsch's protologue. It comes from a similar central European woodland habitat, about 400 km from the type location. Below we designate this collection as epitype for the species. It belongs in a ...
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... recognized species (in order of first description; more complete treatment under Taxonomy). When Batsch described Agaricus glaucus (Batsch 1786) he did not indicate a type but provided an illustration (Fig. 2C), which we designate below as lectotype. Collection WU-6564 ( Fig. 2B) is a perfect match for this lectotype, and fits the description in Batsch's protologue. It comes from a similar central European woodland habitat, about 400 km from the type location. Below we designate this collection as epitype for the species. It belongs in a large clade with collections from Scandinavia and central Europe, which ...
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... from a similar central European woodland habitat, about 400 km from the type location. Below we designate this collection as epitype for the species. It belongs in a large clade with collections from Scandinavia and central Europe, which has enjoyed consistently high support in all our phylogenetic analyses. Agaricus acerosus Fr., s. str.* ( Fig. 2A), is neotypified below by a collection (UPS-151993) from Femsjö, the type locality, fitting Fries' protologue morphologically, growing on woody debris in similar forest habitat. It was one of three collections from northern Europe, which formed a clade with consistently high support in all our phylogenetic analyses. Three collections ...
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... neotypified below by a collection (UPS-151993) from Femsjö, the type locality, fitting Fries' protologue morphologically, growing on woody debris in similar forest habitat. It was one of three collections from northern Europe, which formed a clade with consistently high support in all our phylogenetic analyses. Three collections of A. latispora* (Fig. 2D, F) from the European Alps agreed morphologically with the holotype, but because it was unavailable for sequencing, a collection from the toporegion (LIP-0401569) is designated as epitype. Although several other species in the complex also produce spores of similar width (e.g. A. subglobisemen, A. svalbardensis, A. tillii), in addition to ...
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... ITS sequence data, the species is set apart by its alpine habitat, larger size, Pelargonium odour and striking dark bluish-violet colour. Because the holotype for the pink-spored Rhodocybe tillii (WU-18120) fell into the acerosa complex it is transferred to Arrhenia as the sole representative of A. tillii* ( Krisai & Noordel.) Krisai & I. Saar (Fig. 2J). Arrhenia subglobisemen* (Fig. 2G, H, I), has the widest distribution of any species of the complex, with collections from Estonia, France, Norway, Newfoundland, and Labrador in arctic-alpine to woodland settings. These agreed morphologically with the type specimen, but because it was not available for sequencing, the species is ...
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... set apart by its alpine habitat, larger size, Pelargonium odour and striking dark bluish-violet colour. Because the holotype for the pink-spored Rhodocybe tillii (WU-18120) fell into the acerosa complex it is transferred to Arrhenia as the sole representative of A. tillii* ( Krisai & Noordel.) Krisai & I. Saar (Fig. 2J). Arrhenia subglobisemen* (Fig. 2G, H, I), has the widest distribution of any species of the complex, with collections from Estonia, France, Norway, Newfoundland, and Labrador in arctic-alpine to woodland settings. These agreed morphologically with the type specimen, but because it was not available for sequencing, the species is epitypfied below with a collection from the ...
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... exsiccatum. We have chosen no. 1761 as neotype. It contains material collected at Femsjö, the collecting area of Elias Fries (Petersen & Knudsen 2015) and thus the type locality for Ag. acerosus s. str., grows in similar habitat and on similar substrate as described in the protologue, and agrees well with the current description and our photo ( Fig. 2A) of the species. It differs widely from Fries' (1874a) illustration of A. acerosa, which shows yellow-brown, fanshaped specimens with fairly long, thin, lateral, tapering and strigose pseudostipes attached to moss, which Fries notes shows a dark variety (varietam fuscam) of Ag. acerosus, thus already noting the diversity in the complex. ...
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... references to illustrations by Lange (1936) and pl. 63B is indicated as representative of the material distributed in no. 1761, showing grey, pleurotoid, short stipitate specimens. In the phylogenetic tree (Fig. 1b) the neotype falls in a clade with a specimen from Estonia and one from Norway, Vestfold county in SE Norway. The latter specimen ( Fig. 2A) comes from similar habitat (pathside bare earth with moss) and has similar spores (6.8-8.7 × 3.5-4.4 µm, Q avg = 2.1, narrowly elliptical to slightly ...
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... Pleurotellus roseolus (Quél.) Kühner, based on Pleurotus roseolus Quél.) C o m m e n t . -Arrhenia tillii differs from AC-3 ( Fig. 4B 1-2 ) by a rudimentary or lacking stipe, lateral attachment, pink sporeprint and coniferous substrate. Additional finds of Arrhenia tillii in France and Switzerland share the same substrate (Pinus nigra) as the Austrian collections (Senn-Irlet 1986, Francini 2000. A pink specimen from Quebec (Ac-5, Fig 4G; Lamoureux 2015), identified as ...
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... finds of Arrhenia tillii in France and Switzerland share the same substrate (Pinus nigra) as the Austrian collections (Senn-Irlet 1986, Francini 2000. A pink specimen from Quebec (Ac-5, Fig 4G; Lamoureux 2015), identified as Clitopilus tillii, proved to be quite distant in the ITS tree (see discussion). The subglobose, obovoid to broadly elliptical spores make a striking discriminating character of A. tillii: seemingly smooth under the light microscope, but pustular-wavy in lateral and slightly angular-wavy in pole view with a large hilar apiculus under SEM, characters that previously led to its inclusion in the former genus Rhodocybe. ...
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... that the stipe of P. acerosus starts centrally, soon ceases to elongate, and because of inequilateral growth, ends up eccentric or lateral. Such transition from a central to lateral stipe within the acerosa complex was noted in A. svalbardensis, and is suggested by comparing the collection from northern Québec (Fig. 5C) and A. subglobisemen in Fig. 2G. The thick stipe and incomplete pileus of at least one Québec basidioma approaches the pleurotoid habitus and lateral stipe of the latter. Study of these omphalinoid species of the acerosa complex is an interesting and significant project, requiring, among other things, typification of A. griseopallida and study of its relationship to ...

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... Arrhenia-species are either obligatory bryophiles, e.g., A. retiruga and A. lobata, others are lichenophiles, e.g., A. peltigerina, still others are not strictly moss-bound at all, such as species of the acerosa-complex (Voitk et al. 2020(Voitk et al. , 2022. Recently, Voitk et al. (2022) have shown that the species diversity of sphagnophilous Arrhenia is much greater than originally thought and that some are obligate and others facultative sphagnophiles, i.e., A. bigelowii, A. gerardiana, A. telmatiaea, and A. philonotis, respectively. ...
... However, unlike the sphagnophilous and most other mossdwelling Arrhenia, A. bryophthora causes a necrosis on its host and must therefore be considered a true parasite (Voitk et al. 2022). To our knowledge, a similarly destructive lifestyle is only known from the lichenophilic A. peltigerina (Diederich et al. 2022), which is macroscopically similar, but microscopically distinct by the presence of clamp connections and phylogenetically related to species of the auriscalpium/spathulata complex (Voitk et al. 2020). ...
... In the phylogenetic tree based on ITS and LSU sequences (Fig. 1), all specimens of A. bryophthora group in the same moderately supported clade that is the most basal one of all sequences assigned to Arrhenia specimens. Considering the phylogenetic results presented in Fig. 1 and in the literature (Voitk et al. 2020(Voitk et al. , 2022Zhang et al. 2022) and the fact that the current understanding of the genus Arrhenia includes species with facultative and obligate bryophilous, lichenophilous, and saprotrophic lifestyles as well as cyphelloid, pleurotoid, and omphalinoid habitus; it is plausible that this taxonomic group should be revised and studied in more detail in the future. This includes careful morphological and phylogenetic studies based on multi-locus sequencing and comprehensive sampling. ...
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The very hot summers of recent years have led to an increase in the number of large forest fires in Europe. We investigated four large fire sites in Brandenburg and Saxony (Germany) up to 4 years after the fires with a focus on studying the post-fire fungal communities. In this context, we documented two species of Agaricomycetes associated with mosses, which are common but particularly emerge on burnt areas, i.e., Arrhenia bryophthora sp. nov. and Bryopistillaria clavarioides sp. nov. The former is an agaric with omphalinoid habit that causes the dieback of the common moss Ceratodon purpureus ; the latter is a clavarioid fungus associated with either Ceratodon purpureus or another common moss, Funaria hygrometrica . Both fungal species appear to be restricted to recently burnt areas and have otherwise not been observed on or in close vicinity of these mosses. Herein, we describe these fungi macro- and micromorphologically as well as on a molecular basis and discuss their taxonomic position and potential lifestyles.
... The genus Arrhenia currently comprises almost 100 species (Index Fungorum, Myco-Bank) and new species are continuously added (e.g., Voitk et al. 2020Voitk et al. , 2022. Lodge et al. (2014) formally established the monophyletic tribe Arrheniae Lücking, comprising the genera Arrhenia, Dictyonema C. Agardh ex Kunth, Cora Fr., and Acantholichen P.M. Jørg. ...
... Initial blast searches in GenBank revealed that the most similar sequences are found in the Arrhenia spathulata group, which corresponds to the second well-supported clade of Voitk et al. (2020). We therefore downloaded sequences of 11 representatives of the A. spathulata group from GenBank to investigate the nrITS variation in relation to the segregation of species and the relations of our samples within this clade ( (Kumar et al. 2018). ...
... . One sequence of A. obscurata (D.A. Reid) Redhead, Lutzoni, Moncalvo and Vilgalys was downloaded and used as the outgroup(Voitk et al. 2020). Calculations of genetic distances (observed p-distance) were done in Geneious and phylo-Table 1. ...
Article
The new species Arrhenia schistidicola Øvstedal & L.Lindblom is described from Western Norway, growing on the moss Schistidium crassipilum on vertical mortar walls. Arrhenia schistidicola is characterized by the lack of a stipe, molecular evidence, and its host moss. It is morphologically similar to A. retiruga, however, molecular analyses show that it is more closely related to A. spathulata.
... last accessed March 2022) lists 83 taxa, 78 species, and 5 varieties, with worldwide distribution; most species have been described from temperate regions. The genus is believed to have a mutualistic relationship with bryophytes, although several members of the A. acerosa complex were noted to have a saprobic relationship with dead plant matter (Voitk et al., 2020). Arrhenia was placed in Hygrophoraceae by Lodge et al. (2013), but an encompassing phylogenetic analysis of the genus has not been done. ...
... The pileus of most species of Arrhenia is greyish, blackish, bluish or brown-grey (Redhead et al., 2002), and can be clearly distinguished from A. nivea. Although there are some near-white species of Arrhenia, e.g., A. eburnea , and some whitish members of the Arrhenia acerosa complex (Voitk et al., 2020) their well-developed gills, and often at least a rudimentary stipe, prevent them from being mistaken for A. nivea. Of white cyphelloid species from (Uzun et al., 2018). ...
... DNA fragmentation in ancient herbarium samples is a well-documented phenomenon [1,78], including in lichen-forming fungi [16,[18][19][20]79,80]. Considering this, and other postmortem damage known to take place in collected specimens [16,80], the success rate for the samples studied here is remarkably high, which may, in part, be explained by the generally higher sequencing success for Basidiomycota among fungal collections [24,25]. ...
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Lichen collected worldwide for centuries have resulted in millions of specimens deposited in herbaria that offer the potential to assess species boundaries, phenotypic diversification, ecology, and distribution. The application of molecular approaches to historical collections has been limited due to DNA fragmentation, but high-throughput sequencing offers an opportunity to overcome this barrier. Here, we combined a large dataset of ITS sequences from recently collected material and historical collections, obtained through Sanger, 454, or Illumina Sequencing, to test the performance of ITS barcoding in two genera of lichenized Basidiomycota: Cora and Corella. We generated new sequence data for 62 fresh specimens (from 2016) and 274 historical collections (collected between 1888 and 1998), for a dataset of 1325 sequences. We compared various quantitative approaches to delimit species (GMYC, bPTP, ASAP, ABGD) and tested the resolution and accuracy of the ITS fungal barcoding marker by comparison with a six-marker dataset. Finally, we quantitatively compared phylogenetic and phenotypic species delimitation for 87 selected Cora species that have been formally described. Our HTS approach successfully generated ITS sequences for 76% of the historical collections, and our results show that an integrative approach is the gold-standard for understanding diversity in this group.
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Novel species of fungi described in this study include those from various countries as follows: Australia, Aschersonia mackerrasiae on whitefly, Cladosporium corticola on bark of Melaleuca quinquenervia, Penicillium nudgee from soil under Melaleuca quinquenervia, Pseudocercospora blackwoodiae on leaf spot of Persoonia falcata, and Pseudocercospora dalyelliae on leaf spot of Senna alata. Bolivia, Aspicilia lutzoniana on fully submersed siliceous schist in high-mountain streams, and Niesslia parviseta on the lower part and apothecial discs of Erioderma barbellatum onatwig. Brazil, Cyathus bonsai on decaying wood, Geastrum albofibrosum from moist soil with leaf litter, Laetiporus pratigiensis on a trunk of a living unknown hardwood tree species, and Scytalidium synnematicum on dead twigs of unidentified plant. Bulgaria, Amanita abscondita on sandy soil in a plantation of Quercus suber. Canada, Penicillium acericola on dead bark of Acer saccharum, and Penicillium corticola on dead bark of Acer saccharum. China, Colletotrichum qingyuanense on fruit lesion of Capsicum annuum. Denmark, Helminthosphaeria leptospora on corticioid Neohypochnicium cremicolor. Ecuador (Galapagos), Phaeosphaeria scalesiae on Scalesia sp. Finland, Inocybe jacobssonii on calcareouss oils in dry forests and park habitats. France, Cortinarius rufomyrrheus on sandy soil under Pinus pinaster, and Periconia neominutissima on leaves of Poaceae. India, Coprinopsis fragilis on decaying bark of logs, Filoboletus keralensis on unidentified woody substrate, Penicillium sankaranii from soil, Physisporinus tamilnaduensis on the trunk of Azadirachta indica, and Poronia nagaraholensis on elephant dung. Iran, Neosetophoma fic on infected leaves of Ficus elastica. Israel, Cnidariophoma eilatica (incl. Cnidariophoma gen. nov.) from Stylophora pistillata. Italy, Lyophyllum obscurum on acidic soil. Namibia, Aureobasidium faidherbiae on dead leaf of Faidherbia albida, and Aureobasidium welwitschiae on dead leaves of Welwitschia mirabilis. Netherlands, Gaeumannomycella caricigena on dead culms of Carex elongata, Houtenomyces caricicola (incl. Houtenomyces gen. nov.) on culms of Carex disticha, Neodacampia ulmea (incl. Neodacampia gen. nov.) on branch of Ulmus laevis, Niesslia phragmiticola on dead standing culms of Phragmites australis, Pseudopyricularia caricicola on culms of Carex disticha, and Rhodoveronaea nieuwwulvenica on dead bamboo sticks. Norway, Arrhenia similis half-buried and moss-covered pieces of rotting wood in grass-grownpath. Pakistan, Mallocybe ahmadii on soil. Poland, Beskidomyces laricis (incl. Beskidomyces gen. nov.) from resin of Larix decidua ssp. polonica, Lapidomyces epipinicola from sooty mould community on Pinus nigra, and Leptographium granulatum from a gallery of Dendroctonus micans on Picea abies. Portugal, Geoglossum azoricum on mossy areas of laurel forest areas planted with Cryptomeria japonica, and Lunasporangiospora lusitanica from a biofilm covering a bio deteriorated limestone wall. Qatar, Alternaria halotolerans from hypersaline sea water, and Alternaria qatarensis from water sample collected from hypersaline lagoon. South Africa, Alfaria thamnochorti on culm of Thamnochortus fraternus, Knufia aloeicola on Aloe gariepensis, Muriseptatomyces restionacearum (incl.Muriseptatomyces gen. nov.) on culms of Restionaceae, Neocladosporium arctotis on nest of cases of bagworm moths(Lepidoptera, Psychidae) on Arctotis auriculata, Neodevriesia scadoxi on leaves of Scadoxus puniceus, Paraloratospora schoenoplecti on stems of Schoenoplectus lacustris, Tulasnella epidendrea from the roots of Epidendrum × obrienianum, and Xenoidriella cinnamomi (incl. Xenoidriella gen. nov.) on leaf of Cinnamomum camphora. South Korea, Lemonniera fraxinea on decaying leaves of Fraxinus sp. frompond. 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Ukraine, Xenosphaeropsis corni on recently dead stem of Cornus alba. USA, Nothotrichosporon aquaticum (incl. Nothotrichosporon gen. nov.) from water, and Periconia philadelphiana from swab of coil surface. Morphological and culture characteristics for these new taxa are supported by DNA barcodes.
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Molecular studies of sphagnicolous arrhenias in Newfoundland and Labrador created four clades in Arrhenia , three obligate sphagnophiles (two scaly-capped and one smooth-capped) and one facultative (smooth-capped). Nomenclatural review recovered 16 names applied to omphalinoid sphagnicolous taxa in the past. Critical review of these left five as suitable to this group. One scaly obligate sphagnophilic clade contained the type for Arr. gerardiana and the other the type for Clitocybe gerardiana var. fusca ; the latter we introduce as the novel species Arr. bigelowii . It differed from the first by longer spores and a darkening reaction in 10% of collections. The smooth-capped third obligate sphagnophilic clade contained the types of Ag. telmatiaeus and Omphalina fusconigra ; we recombined it as Arr. telmatiaea . This is the darkest species of the group, with a more northern distribution. The facultative sphagnophile was identified as Arr. philonotis , a lighter smooth-capped species also with a more northern distribution. In addition, we collected an unidentified smooth-capped facultatively sphagnophilic species of Omphalina of the O. pyxidata complex. All five species are distributed in both Europe and North America. We describe each species of Arrhenia with a sequenced type, providing new type material where needed, and add an informal description of the Omphalina . Overall, this study adds new sequences from 82 specimens of sphagnicolous arrhenias to the two existing in GenBank when we began, 11 new sequences of the unidentified species of Omphalina , and several other arrhenias.
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