Article

Overview of Xylariales (Ascomycota) in Bulgaria

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Abstract

This article represents information about the known diversity of xylarialean fungi of Bulgaria. Fifty-seven species, including three varieties and one subspecies, are listed in five families of Xylariales in alphabetical order. Information on the species distribution by floristic regions and their host plants or substrata is included. Anthostoma decipiens is reported for the first time for Bulgaria, and Hypoxylon howeanum has been recorded on Diatrype stigma. The first molecular results are reported for Diatrypella quercina, Hypoxylon fuscum, H. fragiforme, and Jackrogersella cohaerens from Bulgaria, based on the examination of some recent Bulgarian collections.

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June 2024
Dimitar Stoykov · Pablo Alvarado
... Another six fungicolous taxa with minute ascomata (Cosmospora magnusiana s.l., Dialonectria episphaeria, Flamocladiella anomiae, Hypoxylon howeanum, Nectria decora and Tubeufia cerea), growing predominantly on old effete stromata of Diatrype sigma, Diatrypella quercina, Diatrypella sp., Quaternaria quaternata and Massaria anomia were reported recently by Stoykov et al. (2018), Lechat et al. (2019), Stoykov (2020Stoykov ( , 2021Stoykov ( , 2023, Stoykov and Alvarado (2023). However, only two of these identifications were supported by molecular data. ...
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Abstract A revised version of Chapter F of the International Code of Nomenclature for algae, fungi, and plants is presented, incorporating amendments approved by the Fungal Nomenclature Session of the 11th International Mycological Congress held in San Juan, Puerto Rico in July 2018. The process leading to the amendments is outlined. Key changes in the San Juan Chapter F are (1) removal of option to use a colon to indicate the sanctioned status of a name, (2) introduction of correctability for incorrectly cited identifiers of names and typifications, and (3) introduction of option to use name identifiers in place of author citations. Examples have been added to aid the interpretation of new Articles and Recommendations, and Examples have also been added to the existing Art. F.3.7 concerning the protection extended to new combinations based on sanctioned names or basionyms of sanctioned names (which has been re-worded), and to Art. F.3.9 concerning typification of names accepted in the sanctioning works.
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A multigene phylogeny was constructed including a significant number of representative species of the main lineages in the Xylariaceae and four DNA loci – the internal transcribed spacer region (ITS), the large subunit (LSU) of the nuclear rDNA; the second largest subunit of the RNA polymerase II (RPB2) and beta-tubulin (TUB2). Specimens were selected based on more than a decade of intensive morphological and chemotaxonomic work and cautious taxon sampling was performed to cover the major lineages of the Xylariaceae, however with emphasis on hypoxyloid species. The comprehensive phylogenetic analysis revealed a clear-cut segregation of the Xylariaceae into several major clades, which was well in accordance with previously established morphological and chemotaxonomic concepts. One of these clades contained Annulohypoxylon, Hypoxylon, Daldinia and other related genera that have stromatal pigments and a nodulisporium-like anamorph. They are accommodated in the family Hypoxylaceae, which is resurrected and emended. Representatives of genera with a nodulisporium-like anamorph and bipartite stromata, lacking stromatal pigments (i.e. Biscogniauxia, Camillea and Obolarina) appeared in a clade basal to the xylarioid taxa. As they clustered with Graphostroma platystomum, they are accommodated in the Graphostromataceae. The new genus Jackrogersella with J. multiformis as type species is segregated from Annulohypoxylon. The genus Pyrenopolyporus is resurrected for Hypoxylon polyporus and allied species. The genus Daldinia and its allies Entonaema, Rhopalostroma, Ruwenzoria and Thamnomyces appeared in two separate subclades, which may warrant further splitting of Daldinia in the future, and even Hypoxylon was divided in several clades. However, more species of these genera need to be studied before a conclusive taxonomic rearrangement can be envisaged. Epitypes were designated for several important species in which living cultures and molecular data are available, in order to stabilise the taxonomy of the Xylariales.
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This work presents recently discovered country records of stromatic and non-stromatic pyrenomycetes. The size of the microscopic characters is given in the form of minimum and maximum values. Measurements under LM were usually taken in water. The microscopic features were examined in water and in solution of Cotton Blue in lactophenol. Photographs were taken with the help of Canon PS A460 under Boeco microscopes. The distribution of Diatrypella favacea, D. quercina (Xylariales), Gnomonia geranii-macrorrhizi, Gnomoniopsis comari s. lat., Mamiania fimbriata, Ophiognomonia rosae, Plagiostoma apiculatum (Diaporthales) and Pleospora herbarum (Pleosporales) is given according to the data published. Pleospora herbarum (Pleosporales) is found on dead twigs of Amorpha fruticosa and Buddleja davidii. All of the species except P. herbarum are reported from new localities. Seven pyrenomycetous fungi are reported from new localities (Forebalkan, Stara Planina Mts, Sofia region, Vitosha region, Rila and Rhodopi Mts), while D. quercina, P. apiculatum and P. herbarum are collected on new host-plants.
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Hypoxylon petriniae sp. nov. is described from a new combination of morphological characters of its teleomorph and anamorph. It was found in France and other countries of Western and Central Europe, mostly colonising Fraxinus excelsior. Its status is supported by a chemotaxonomic study by analytical HPLC profiling with diode array and mass spectrometric detection of secondary metabolites in several Hypoxylon spp. These analyses revealed the presence of binaphthalene tetrol (BNT) as major stromatal metabolite of H. petriniae. This widespread purple stromatal pigment of Xylariaceae was not found in H. rubiginosum, H. cercidicolum, and further, presumably related species. In contrast to H. rubiginosum and H. cercidicolum, H. petriniae was found devoid of mitorubrin. Orsellinic acid and rubiginosins, which are recently identified azaphilone pigments chemically related to mitorubrin were found as common stromatal metabolites of many of the aforementioned taxa, and the occurrence of particular azaphilones appears to be specific for some species. A key to European Hypoxylon spp. is provided.
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The genus Lopadostoma (Xylariaceae, Xylariales) is revised. Most species formerly assigned to Lopadostoma do not belong to the genus. Twelve species are herein recognised, of which two are only known from morphology. Ten species, of which six (L. americanum, L. fagi, L. insulare, L. lechatii, L. meridionale and L. quercicola) are newly described, are characterised by both morphology and DNA phylogeny using LSU, ITS and rpb2 sequences. Morphologically, ecologically and phylogenetically Lopadostoma is a well-defined genus comprising exclusively species with pustular pseudostroma development in bark of angiospermous trees. Phaeosperma ailanthi, Phaeosperma dryophilum and Sphaeria linosperma are combined in Lopadostoma. Lopadostoma gastrinum is neotypified and L. turgidum is lecto- and epitypified. Species with asci and ascospores similar to those of Lopadostoma but having perithecia immersed in wood, particularly those of Lopadostoma subg. Anthostomopsis have been determined to be unrelated to the genus. DNA data confirm that Anthostoma is unrelated to Lopadostoma. Its type and currently only confirmed species Anthostoma decipiens belongs to Diatrypaceae. DNA data also show that L. pouzarii and Barrmaelia macrospora are unrelated to Lopadostoma. A commentary is provided for names in Lopadostoma and those names in Anthostoma that may be putative species of Lopadostoma based on their protologues. Anthostoma insidiosum is an older name for Anthostomella (Diatrype) adusta.
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For a monograph based on a polythetic concept, several thousands of herbarium specimens, and several hundreds of freshly collected and cultured specimens of Daldinia and allied Xylariaceae, originating from around the world, were studied for morphological traits, including by SEM, and chemically by HPLC profiles using UV-visible and mass spectrometric detection. Emphasis was given to tropical material, and importantly, ancient specimens, including as many types as possible, were tracked and studied to review earlier taxonomic concepts. An epitype of D. eschscholtzii was selected as representative of the morphochemotype that is most widely distributed in the tropics. Six new species of Daldinia from the tropics and the southern Hemisphere are described. Daldinia asphalatum is resurrected, and D. cudonia is regarded as its synonym. In addition, the following binomials are epi-, iso-, neo- and/or lectotypified: Daldinia asphalatum, D. caldariorum, D. clavata, D. cuprea, D. durissima, D. eschscholtzii, D. grandis, D. loculata, and D. vernicosa. Annellosporium and Versiomyces are regarded as synonyms of Daldinia. Many new synonymies in Daldinia are proposed, and some previously published names are rejected. In total, 47 taxa in Daldinia are recognised and a key is provided. Their biogeography, chorology, and ecology, as well as the importance of their secondary metabolites, are also discussed. The previous definition of the genus is emended. The species concept is based mainly on morphological and other phenotype-derived characters because, despite diligent search, no molecular data or cultures of several of the accepted species could be obtained. Daldinia is segregated into five major groups, based on phenotypic characteristics. Some unnamed but aberrant specimens were not found in good condition and are therefore not formally described as new species. However, they are illustrated in detail in a hope that this will facilitate the discovery of fresh material in future. A preliminary molecular phylogeny based on 5.8S/ITS nrDNA including numerous representatives of all hitherto described taxa for which cultures are extant, was found basically in agreement with the above mentioned segregation of the genus, based on morphological and chemotaxonomic evidence. In the rDNA based phylogenetic tree, Daldinia appears clearly distinct from members of the genera Annulohypoxylon and Hypoxylon; nevertheless, representatives of small genera of predominantly tropical origin (Entonaema, Phylacia, Ruwenzoria, Rhopalostroma, Thamnomyces) appear to have evolved from daldinioid ancestors and are nested inside the Daldinia clade. Interestingly, these findings correlate with chemotaxonomic characters to a great extent, especially regarding the distribution of marker metabolites in their mycelial cultures. Hence, the current study revealed for the first time that fungal secondary metabolite profiles can have taxonomic value beyond the species rank and even coincide with phylogenetic data. Taxonomic novelties: Daldinia andina sp. nov., D. australis sp. nov., D. hausknechtii sp. nov., D. rehmii sp. nov., D. starbaeckii sp. nov., D. theissenii sp. nov., D. cahuchosa comb. nov., D. nemorosa comb. nov.
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The genus Entonaema comprises Xylariaceae with hollow, gelatinous stromata that accumulate liquid. Some of its species, including the type species, appear related to Daldinia from a polyphasic approach, comprising morphological studies, comparisons of ribosomal DNA sequences, and high performance liquid chromatography (HPLC) profiles with diode array and mass spectrometric detection (HPLC-DAD-MS). This methodology was used to study Entonaema pallidum. Its major stromatal constituent was identified as xylaral, a secondary metabolite known from Xylaria polymorpha. This compound was detected in several Xylaria spp., including the tropical X. telfairii and morphologically similar taxa, whose stromata may also become hollow and filled with liquid. Cultures of E. pallidum resembled those of Xylaria, substantially differing from other Entonaema spp., in their morphology, 5.8S/ITS nrDNA sequences, and HPLC profiles. The type specimen of E. mesentericum was located in the spirit collection of the herbarium B and found to agree morphologically with the nomenclatorily younger E. pallidum. Traces of xylaral were even detected by HPLC-DAD-MS in the spirit in which the fungus had been preserved. Entonaema pallidum is thus regarded as a later synonym of E. mesentericum. Therefore, the latter name is transferred to Xylaria. A key to entonaemoid Xylariaceae is provided. Colour reactions (NH3, KOH) of the ectostroma were applied to a limited number of Xylaria spp., but metabolite profiles of cultures appear more promising as chemotaxonomic traits to segregate this genus. As xylaral was also found in Nemania and Stilbohypoxylon spp., while being apparently absent in Hypoxylon and allied genera, it may be a chemotaxonomic marker for Xylariaceae with Geniculosporium-like anamorphs.
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New records of Cosmospora magnusiana s.l. from Stara Planina Mts and Sofia region, associated with stromata of Diatrypella on dead oak and maple twigs, are reported. The two findings are presented with concise description and original colour illustrations. Information from the known published sources is summarized, and available data on the ecology and distribution is applied.
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A large database of invasive forest pathogens ( IFP s) was developed to investigate the patterns and determinants of invasion in Europe. Detailed taxonomic and biological information on the invasive species was combined with country‐specific data on land use, climate, and the time since invasion to identify the determinants of invasiveness, and to differentiate the class of environments which share territorial and climate features associated with a susceptibility to invasion. IFP s increased exponentially in the last four decades. Until 1919, IFP s already present moved across Europe. Then, new IFP s were introduced mainly from North America, and recently from Asia. Hybrid pathogens also appeared. Countries with a wider range of environments, higher human impact or international trade hosted more IFP s. Rainfall influenced the diffusion rates. Environmental conditions of the new and original ranges and systematic and ecological attributes affected invasiveness. Further spread of established IFP s is expected in countries that have experienced commercial isolation in the recent past. Densely populated countries with high environmental diversity may be the weakest links in attempts to prevent new arrivals. Tight coordination of actions against new arrivals is needed. Eradication seems impossible, and prevention seems the only reliable measure, although this will be difficult in the face of global mobility.
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The decline in Ulmus pollen frequencies that occurred ca. 5000 14C years ago before present (BP) is a key biostratigraphic marker horizon in northwest European pollen diagrams, although its causes are still a subject of debate. To investigate this event further, fungal spore analyses were carried out across the Ulmus decline at Moel y Gerddi, north Wales, United Kingdom. The Ulmus decline was in three phases, with a primary decline with low cereal and Rumex pollen records as the only agricultural indicators. This was followed by a more significant decline, with general forest opening and the grassland/pasture indicator Plantago lanceolata. A third, less significant, decline was again accompanied by cereal-type pollen. Tree pollen frequencies subsequently recovered, with cereal-type pollen remaining well represented. The fungal data recorded woodland taxa and a background level of the obligate dung fungus Sporormiella, a proxy for local herbivore abundance. Sporormiella frequencies increased greatly after the main Ulmus decline, around which there were high percentages of the ascospores of the wood rot fungus Kretzschmaria deusta. The neoecology of Kretzschmaria deusta, and the behavior of its spore curve suggests the colonisation of local populations of already severely wounded trees. At this site Ulmus, and perhaps Tilia were infected at a time of markedly increased inferred herbivore concentrations. Neolithic farming techniques could have provided mortally wounded trees while enhancing livestock grazing, although the role of disease must also be considered. Elevated Kretzschmaria deusta values may be of ecological significance in the interpretation of the causes and nature of the Ulmus decline and similar forest disturbance events, and the indicator role of this fungus in forest paleoecology requires further study.
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Entonaema, Pulveria, Phylacia, Rhopalostroma, Sarcoxylon and Thamnomyces are relatively small and poorly studied genera of Xylariaceae. Their affinities to the mainstream of the family largely remain to be evaluated. Secondary metabolite profiles of type materials, recently collected specimens, and cultures, were generated to address this problem from a chemotaxonomic point of view. Micro-scale extraction and subsequent analytical HPLC with uv/visual (diode array) and ms detection in the positive and negative Electrospray mode were carried out, employing spectral libraries and standardised gradients that had been optimised to detect characteristic pure compounds in species of allied genera. Surprisingly, the characteristic metabolites had frequently remained stable even in specimens collected up to 190 years ago. Hence, this methodology not only proved valuable to establish the conspecificity of type materials with recent records, but also revealed some interesting correlations: (1) Stromatal pigments of Entonaema cinnabarina, E. globosum and E. liquescens are mitorubrins and other characteristic compounds also prevailing in particular species of Hypoxylon; (2) Rhopalostroma, Phylacia, Pulveria and Thamnomyces contain binapththalenes and other compounds typical of Daldinia and Hypoxylon; (3) Sarcoxylon, as well as E. dengii, E. moluccanum and E. pallida, contained none of these pigments, but characteristic yet unknown lipophilic metabolites were detected in their stromatal extracts; (4) Cultures of E. cinnabarina and Rhopalostroma indicum, obtained for the first time, produced essentially the same secondary metabolites that are also typical of Daldinia, but absent in Biscogniauxia, Hypoxylon and other xylariaceous genera. Hence, chemotaxonomic characters reflected the adaptive radiation of this family. Correlations between the evolution of morphological/anatomical characters are discussed.
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