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Line drawing of Hebeloma grandisporum collection KRAM F-47189 (HJB13114) (holotype): A. Basidia 6 1000. B. Cheilocystidia 6 1000, bar 5 10 mm. C. Spores 6 1600, scale bar 5 mm.
Source publication
Between 2002 and 2012 regular visits to the Carpathians were made and a number of Hebeloma spp. were collected from the alpine area. In total 44 collections were made that represent 11 species, two of which, Hebeloma grandisporum and H. oreophilum, are described here as new. Of the 11 species, four (H. alpinum, H. marginatulum and the two species d...
Citations
... Newly generated sequences were accessioned to GenBank and received accession numbers MT157306, OP160001, OP162978-OP163006, OP163129-OP163165, and OP168761. Details are listed in SUPPLEMENTARY TABLE 3. Previously published sequences (Argüelles-Moyao et al. 2017;Bandini et al. 2019Bandini et al. , 2021aBandini et al. , 2021bBandini et al. , 2022aBandini et al. , 2022bBeker et al. 2010Beker et al. , 2013Beker et al. , 2016Beker et al. , 2018Bidartondo and Read 2008;Bizio and Castellan 2017;Bowman and Arnold 2021;Brock et al. 2009;Brown et al. 2022;Cervini et al. 2020;Chen et al. 2018;Cho et al. 2016;Christ et al. 2011;Clausing and Polle 2020;Cripps et al. 2019;Crous et al. 2018;Csizmár et al. 2021;Eberhardt et al. 2009Eberhardt et al. , 2013Eberhardt et al. , 2016aEberhardt et al. , 2016bEberhardt et al. , 2018Eberhardt et al. , 2020aEberhardt et al. , 2020bEberhardt et al. , 2021Eberhardt et al. , 2022aEberhardt et al. , 2022bEberhardt et al. , 2022cFan and Bau 2018;Frings et al. 2020;Garrido-Benavent et al. 2020;Grilli et al. 2016;Guzman-Davalos et al. 2003;Hallen et al. 2003;Harrower et al. 2011;Hashimoto et al. 2012;Holec et al. 2014Holec et al. , 2016Hughes et al. 2009;Hyde et al. 2016;Jabeen and Khalid 2020;Kasuya and Hosaka 2017;Katanić et al. 2016;Kennedy et al. 2011;Kranabetter et al. 2015;Krisai-Greilhuber et al. 2018;Kropp et al. 2013;Krüger et al. 2012;Landry et al. 2021;Larsson et al. 2009Larsson et al. , 2014Latha et al. 2016;Malysheva and Kiyashko 2011;Malysheva et al. 2016;Marchetti et al. 2014;Matheny 2005;Matheny and Bougher 2017;Matheny et al. 2002Matheny et al. , 2006Matheny et al. , 2007Matheny et al. , 2015Matheny et al. , 2020Niskanen et al. 2011Niskanen et al. , 2012Olchowik et al. 2021;Osmundson et al. 2013;Peintner et al. 2004;Rodríguez-Gutíerrez et al. 2020;Ryberg et al. 2008Ryberg et al. , 2010Schoch et al. 2012Schoch et al. , 2014Seger et al. 2017;Sesli 2021;Soop et al. 2019;Stensrud et al. 2014;Suz et al. 2014;Tedersoo et al. 2003Tedersoo et al. , 2006Tedersoo et al. , 2020Thorn et al. 1996;Tian and Matheny 2021;van der Walt et al. 2020;Vašutová et al. 2018;Vauras and Larsson 2020;Vesterholt et al. 2014;Vu et al. 2019;Walther et al. 2005;Yang et al. 2005;Zhang et al. 2017) Vašutová et al. 2018Vauras and Larsson 2020;Vesterholt et al. 2014;Vu et al. 2019;Walther et al. 2005;Yang et al. 2005;Zhang et al. 2017) used in this study are summarized in SUPPLEMENTARY TABLE 4. To determine the taxonomic relationships of sequences from collections that were not Hebeloma, BLAST searches were carried out against GenBank (Johnson et al. 2008), UNITE (Kõljalg et al. 2005), and BOLD (Ratnasingham and Hebert 2007) databases. BLAST searches against our own data were done in Geneious R10 (Biomatters, Auckland, New Zealand) with default settings. ...
William Alphonso Murrill was an American mycologist of the early 20th century. He described 1453 new species of Agaricales, Boletales, and Polyporales. Within these were 44 taxa that he described as Hebeloma or that he recombined into Hebeloma. Additionally, there are five species, of which we are aware, that Murrill described within other genera that should be referred to the genus Hebeloma. A further three species described from northern America by J. P. F. C. Montagne, and transferred to Hebeloma by Saccardo, were commented on by Murrill and not accepted within the genus. These 52 taxa are analyzed here, both morphologically and molecularly, as far as possible. For 18 of his types, internal transcribed spacer (ITS) sequences were generated. For two species (H. harperi and H. subfastibile), which were mixed collections, lectotypes are designated. Twenty-three of the taxa analyzed are Hebeloma, as the genus is recognized today, and six of these (H. australe, H. harperi, H. paludicola, H. subaustrale, H. subfastibile, and H. viscidissimum) are regarded as current, i.e., they are names that should be accepted and used. Hebeloma paludicola is an earlier name for H. hygrophilum, described from Europe. Gymnopilus viscidissimus is synonymous with H. amarellum but has priority and is here recombined into Hebeloma. The remaining 17 Hebeloma taxa are synonymized with other species that have priority. The remaining 29 species belong to a range of genera; molecularly supported were Agrocybe, Cortinarius, Inocybe, Inosperma, Phlegmacium, Pholiota, Pseudosperma, and Pyrrhulomyces. Recombinations and synonymizations are made as appropriate and necessary. The names H. alachuanum and H. vatricosum, respectively Inocybe vatricosa, are considered doubtful and should be avoided.
... Within our project we have embraced in various combinations two variable domains of the mitochondrial small subunit of the ribosomal RNA genes (mitSSU), referred to as V6 and V9, as well as part of the gene for the second largest subunit of RNA polymerase II (RPB2), a partial sequence of the translation elongation factor 1 α (TEF1a) and the protein coding gene MCM7 (Minichromosome Maintenance Complex Component 7). Technical details can be found in Eberhardt (2012), Eberhardt et al. (2016Eberhardt et al. ( , 2021a, Vesterholt et al. (2014) and Cripps et al. (2019) and references therein. ...
The genus Hebeloma is renowned as difficult when it comes to species determination. Historically, many dichotomous keys have been published and used with varying success rate. Over the last 20 years the authors have built a database of Hebeloma collections containing not only metadata but also parametrized morphological descriptions, where for about a third of the cases micromorphological characters have been analysed and are included, as well as DNA sequences for almost every collection. The database now has about 9000 collections including nearly every type collection worldwide and represents over 120 different taxa. Almost every collection has been analysed and identified to species using a combination of the available molecular and morphological data in addition to locality and habitat information. Based on these data an Artificial Intelligence (AI) machine-learning species identifier has been developed that takes as input locality data and a small number of the morphological parameters. Using a random test set of more than 600 collections from the database, not utilized within the set of collections used to train the identifier, the species identifier was able to identify 77% correctly with its highest probabilistic match, 96% within its three most likely determinations and over 99% of collections within its five most likely determinations.
... Newly generated sequences were accessioned to Gen-Bank (ON167764-ON167898, ON168958-ON168966, ON202494-ON202614 and ON237944-ON237985), Suppl. material 1: Table S1 summarizes all sequences used in the analyses, including those previously published in the context of a number of publications (Eberhardt et al. 2009(Eberhardt et al. , 2015(Eberhardt et al. , 2016a(Eberhardt et al. , 2016b(Eberhardt et al. , 2021a(Eberhardt et al. , 2022a(Eberhardt et al. , 2022bEberhardt and Beker 2010;Beker et al. 2010Beker et al. , 2013Beker et al. , 2016Schoch et al. 2012;Cripps et al. 2019). ...
The species of Hebeloma have been little studied in Mexico, but have received attention as edibles and in trials to enhance production of edible fungi and tree growth through inoculation of seedlings with ectomycorrhizal fungi. Here we describe three new species of Hebeloma that are currently known only from Mexico. These species belong to separate sections of the genus: H. ambustiterranum is a member of H. sect. Hebeloma, H. cohaerens belongs to H. sect. Theobromina, while H. magnicystidiatum belongs to H. sect. Denudata. All three species were collected from subtropical pine-oak woodland; all records of H. cohaerens came from altitudes above 2500 m. Hebeloma ambustiterranum is commonly sold in the local markets of Tlaxcala as a prized edible mushroom. An additional nine species are reported from Mexico, of which eight are new records for the country: H. aanenii , H. eburneum , H. excedens , H. ingratum , H. neurophyllum , H. sordidulum , H. subaustrale and H. velutipes . First modern descriptions of H. neurophyllum and H. subaustrale , originally described from the USA, are given here.
... The spore size would indicate that this is what we used to call Hebeloma clavulipes in, e.g. Eberhardt et al. (2015Eberhardt et al. ( , 2021a, Beker et al. (2016Beker et al. ( , 2018 and Cripps et al. (2019), but it should be noted that H. palustre Peck (1899), described below, should also be referred to the same species and that the name Hebeloma palustre has priority. ...
... In order to begin unravelling the then taxonomically rather chaotic genus Hebeloma, we (in Eberhardt et al. 2013Eberhardt et al. , 2015Eberhardt et al. , 2016Beker et al. 2016, Grilli et al. 2016 decided to first tackle the taxa described from Europe before applying ourselves to the numerous species and varieties described from North America. The only exception we made was for H. incarnatulum Grilli et al. 2016), because the name was already in use in Europe (e.g. ...
Charles Horton Peck described some 2700 species of North American fungi in the 19th and early 20th centuries. Among these were 31 species that he described as Hebeloma or that later authors recombined into Hebeloma. These 31 taxa have been analyzed morphologically and molecularly, as far as possible. For six of these species, lectotypes are designated. For twelve species, ITS sequences (some partial) were generated. Thirteen of the species analyzed are Hebeloma, as the genus is delimited today. Of these 13, nine are regarded as 'current', i.e. are names that should be accepted and used. Of the remaining four, three are synonymized with earlier Peck species and one with the generic type H. mesophaeum. Numerous Hebeloma species described from America are synonymized with some of Peck's species, such as H. albidulum, H. album, H. colvinii, H. excedens, H. palustre, H. sordidulum, and H. velatum; Peck's H. album, H. palustre, and H. velatum are earlier names for H. fragilipes, H. clavulipes, and H. dunense, respectively. All three names were in current use and described from Europe. The 18 species that are not Hebeloma belong to a range of genera: Agrocybe, Hemistropharia, Inocybe, Inosperma, Naucoria, and Pholiota; three species that were not previously recombined into their respective genera are here recombined and one species, Hebeloma commune is synonymized with Pholiota lenta. Two taxa, that are not Hebeloma, remain unresolved. Sixty later Hebeloma taxa described from North America are revised and synonymized with Peck species and seven with H. mesophaeum, 36 of these supported by ITS (some partial) sequence data. Updates on two species, H. petrakii and H. remyi, from Europe, are also given, and a lectotype and epitype selected for the latter.
... Phenotypic characters are combined with phylogenetic analysis produced by using DNA sequences of nuclear ribosomal internal transcribed spacers (nrITS) and nuclear large subunits of ribosomal RNA gene (nrLSU) regions. nrITS region was selected because of availability of universal primers, high PCR success rate, a high percent of correct identification, a considerable number of sequences available in NCBI, and its superior resolution at infrageneric classification level for Hebeloma genus (Aanen et al. 2000, Aanen et al. 2001, Vesterholt et al. 2014, Eberhardt et al. 2015a, Eberhardt et al. 2015b, Tekpinar & Kalmer 2019. The nrLSU region, located immediately downstream of the nrITS, has also been widely used for phylogenetic assignments (Tekpinar & Kalmer 2019, Lücking et al. 2020. ...
In the current study, a new fungus species Hebeloma bingolense sp. nov. is described and illustrated based on materials collected in Bingöl, Turkey. Drawings of microscopic characters, scanning microphotographs of the basidiospores, and color pictures of the basidiomes in their natural habitats are provided. Molecular analysis was also carried out to characterize the new species reliably by using DNA sequences of nuclear ribosomal internal transcribed spacers (nrITS) and nuclear large subunit of ribosomal RNA gene (nrLSU). The phylogenetic position and evolutionary relationships of H. bingolense with other species of Hebeloma are inferred by means of Maximum Likelihood (ML) and Bayesian Inferences (BI) of the concatenated dataset. The novel taxon is located within a well-supported clade formed by members of Hebeloma section Hebeloma and it is closely related to H. dunense and H. alpinicola.
... In addition to these characters, the ribosomal internal transcribed spacers (ITS) comprising ITS1, 5.8S rDNA, and ITS2 sub-regions and the large subunit nuclear ribosomal RNA gene (nrLSU) sequences were also used to be sure of species delimitation and to determine the phylogenetic perspective of the local species with respective to the other species within the genus. Recent phylogenetic studies (Aanen, Kuyper, Boekhout & Hoekstra, 2000;Aanen, Kuyper & Hoekstra, 2001;Vesterholt, Eberhardt & Beker, 2014;Eberhardt, Beker & Vesterholt 2015a;Eberhardt, Ronikier, Schütz & Beker, 2015b) demonstrate that ITS region is very useful marker for studying infrageneric classification of Hebeloma genus and some mycologists (Shimono, Kato & Takamatsu, 2004;Vizzini, Ercole & Contu, 2012;Demirel, 2016) favoured LSU as a phylogenetic marker since it is easily amplified, sequenced, and aligned. Therefore, sequences of these two regions were selected to study the phylogeny. ...
Aim of Study: The specimen identified as Hebeloma subtortum based on both morphological and
molecular characterizations.
Area of Study: Samples were collected from Bingöl province and the study was conducted at the
Department of Molecular Biology and Genetics in Van Yüzüncü Yıl University.
Material and Methods: Characters of pileus, stipe, lamellae and basidia, cystidia, spores were used as
macroscopic and microscopic features, respectively. DNA sequences of two loci including the internal
transcribed spacer (ITS) region and the large subunit (LSU) of nuclear ribosomal RNA gene were used
to show the evolutionary relationship and taxonomic position of the species within Hebeloma genus. The
DNA sequences of the above-mentioned regions of H. subtortum were compared to those of the same and
different species of the genus downloaded from NCBI.
Main results: In phylogenetic analyses, H. subtortum distinctly clustered with its representatives
retrieved from NCBI with high bootstrap value. The ITS tree was more informative compared to LSU.
Hebeloma subtortum closely grouped with H. mesophaeum in the ITS tree.
Research highlights: Hebeloma subtortum has been described and illustrated as a new record from
Turkey.
... Numerous taxa of Hebeloma have been reported in association with Persicaria, Betula, Salix, and Dryas from arctic-alpine habitats including those in the Alps (Favre 1955;Bon 1986;Bruchet 1974;Debaud et al. 1981;Kühner and Lamoure 1986;Senn-Irlet 1990;Senn-Irlet 1993;Jamoni 2008;Graf 1994;Brunner et al. 2017), Iceland (Eyjolfsdottir 2009), Scandinavia (Vesterholt 2005(Vesterholt , 2008Knudsen and Vesterholt 2008), Svalbard (Hutinen 1987;Ohenoja 1971;Gulden and Torkelsen 1996;Beker et al. 2018), Pyrenees (Corriol 2008), and the Carpathians (Eberhardt et al. 2015b). In North America, there are reports from Greenland (Lange 1957;Borgen 2006;Borgen et al. 2006), Canada (Ohenoja andOhenoja 1993, 2010), Alaska (Miller 1998), and the Rocky Mountains (Miller and Evenson 2001;Cripps and Horak 2008;Beker et al. 2010). ...
... Aanen and Kuyper 1999, 2004, Aanen and Kuyper 2004 also gave some evidence with regard to the limits of biological species. As described earlier (Eberhardt et al. 2015a, 2015b, Grilli et al. 2016, species definitions based on several lines of evidence may share ITS or other loci' haplotypes, presumably as a result of incomplete linage sorting, hybridization or other population processes. The molecular distance between some species is so small that we assume that not all groups we recognize as species had sufficient time to reach monophyly in all loci. ...
... Hebeloma vaccinum is known to occur in low elevation dunes and woodlands with Salix; it is widespread in Northern Europe. Other arctic-alpine collections are from the European Alps, the Carpathians in Slovakia, and Greenland, always with Salix species Eberhardt et al. 2015b). It could be recognized in the Rocky Mountains by its association with dwarf Salix, small size, lack of a veil, and distinct spores and cystidia; compare with H. aurantioumbrinum. ...
Numerous taxa of Hebeloma have been reported in association with Salix, Dryas, and Betula in arctic-alpine habitats. However, species are notoriously difficult to delineate because morphological features overlap, and previously there was little reliable molecular data available. Recent progress in ITS-sequencing within the genus, coupled with an extensive database of parametrically described collections, now allows comparisons between species and their distributions. Here we report 16 species of Hebeloma from the Rocky Mountain alpine zone from some of the lowest latitudes (latitude 36°–45°N) and highest elevations (3000–4000 m) for arctic-alpine fungi in the northern hemisphere. Twelve of these species have been reported from arctic-alpine habitats in Europe and Greenland and are now molecularly confirmed from the Middle and Southern Rockies, greatly expanding their distribution. These are: Hebeloma alpinum, H. aurantioumbrinum, H. dunense, H. hiemale, H. marginatulum, H. mesophaeum, H. nigellum, H. oreophilum, H. subconcolor,
H. spetsbergense, H. vaccinum, and H. velutipes. Hebeloma hygrophilum is known from subalpine habitats in Europe, but was never recorded in arctic-alpine ecology. Three species recorded from the Rockies, but as yet not reported from Europe, are H. alpinicola, H. avellaneum, and H. excedens. The last two have never previously been reported from an arctic-alpine habitat. For all three of these species, the holotypes have been studied morphologically and molecularly, and have been incorporated into the analysis.
... Each collection has been given a H.J. Beker (HJB) Database Record number; these are also cited, where relevant, as we intend to publish this database at some future point and we believe these numbers will therefore be useful for future researchers. Species identification was primarily by morphology and anatomy, aided by ITS and, for some collections treated in earlier publications Eberhardt, Beker, & Vesterholt, 2015;Eberhardt, Ronikier, Schütz, & Beker, 2015;Grilli et al., 2016), by sequence data of additional loci. The data from other arctic or alpine areas used for comparison, includes 510 collections that have been described by Beker et al. (2016), and a further 160 records that have recently been added to our database, many from Chukotka in the Russian Federation. ...
... In Beker et al. (2016) this issue is discussed at length. Many Hebeloma species have been exclusively reported from alpine and arctic habitats (e.g., Beker et al., 2016;Bruchet, 1970;Eberhardt, Ronikier et al., 2015;Vesterholt, 2005). It is also interesting that a high number of otherwise typically lowland species can be found in arctic and alpine environments; of the 25 species recorded in alpine and arctic areas , 11 have also been recorded in lowland habitats. ...
Between 1960 and 2009, regular visits to Svalbard were made and a number of Hebeloma spp. were collected by several mycologists. We have studied 249 such collections, many from the Herbarium in Oslo (O) and from the private herbarium of the first author. The collections represent 17 different species. Of the 17 species, five (H. louiseae, H. pallidolabiatum, H. perexiguum, H. pubescens and H. spetsbergense) are only known from Svalbard. In this paper we discuss the habitat and apparent associates of the 17 species recorded, as well as their frequency of occurrence within our sample, and compare this with our data from other regions of the northern hemisphere. A further analysis compares the records we have from the collection of basidiomes with data published from root analysis of mycorrhizal associates on Svalbard. Root associate data strongly suggest the occurrence of an unknown Hebeloma species, not close to any of the infrageneric groups known to occur in arctic habitats.
... The genus Hebeloma, traditionally a difficult genus in which species morphological boundaries are often fuzzy and the relationship between species difficult to ascertain, has much benefited from the adoption of molecular techniques of phylogenetic reconstruction as a support of morphology. Over the last few years, the new approach has brought about the publication of a good number of studies, among which especially important those by the late J. Vesterholt's research team and co-workers:BEkER & al (2010),EBERhARDT & BEkER (2010),EBERhARDT & al (2009),EBERhARDT & al (2013),EBERhARDT & al (2015a),EBERhARDT & al (2015b),EBERhARDT & al (2015c),EBERhARDT & al (2016), GRILLI & al (2016) and VESTERhOLT & al. (2014, the main result of which has been the publication of a monograph devoted to the European species of the genus (BEkER & al, 2016). It is to be hoped that now that this much needed monograph is available, a greater number of mycologists will devote themselves to the study of this fascinating genus, even if morphological species identification remains difficult because of the requirement of specific observational expertise and many accurate measurements of spores and cheilocystidia. ...
Hebeloma lindae in Sicily, firSt report for the italian territory Edmondo Grilli via Tiburtina Valeria 55/A, 65026 POPOLI (PE) Emanuele Brugaletta Via Carlo Pisacane, 6 – 97100 RAGUSA (RG) Grilli E., Brugaletta E., 2016: Hebeloma lindae in Sicilia, prima segnalazione per l'Italia. abstract: Hebeloma lindae, a species of subsect. Scabrispora recently established on material from Spain and Greece, is reported for the first time from Italy. The species is fully described and illustrated, macro and microscopically, on the basis of two collections from Sicilian Mediterranean maquis. riassunto: Hebeloma lindae, taxon della sottosez. Scabrispora pubblicato recentemente sulla base di materiale raccolto in Spagna e Grecia, è segnalato per la prima volta in Italia. La specie è esaurientemente descritta ed illustrata, nei suoi caratteri macro-e microscopici, sulla base di due raccolte provenienti da ambiente mediterraneo siculo.
... The genus Hebeloma, traditionally a difficult genus in which species morphological boundaries are often fuzzy and the relationship between species difficult to ascertain, has much benefited from the adoption of molecular techniques of phylogenetic reconstruction as a support of morphology. Over the last few years, the new approach has brought about the publication of a good number of studies, among which especially important those by the late J. Vesterholt's research team and co-workers: , , EBERhARDT & al (2009), EBERhARDT & al (2013, EBERhARDT & al (2015a), EBERhARDT & al (2015b), EBERhARDT & al (2015c), , GRILLI & al (2016) and VESTERhOLT & al. (2014), the main result of which has been the publication of a monograph devoted to the European species of the genus . ...
Hebeloma lindae in Sicily, firSt report for the italian territory Edmondo Grilli via Tiburtina Valeria 55/A, 65026 POPOLI (PE) Emanuele Brugaletta Via Carlo Pisacane, 6 – 97100 RAGUSA (RG) Grilli E., Brugaletta E., 2016: Hebeloma lindae in Sicilia, prima segnalazione per l'Italia. abstract: Hebeloma lindae, a species of subsect. Scabrispora recently established on material from Spain and Greece, is reported for the first time from Italy. The species is fully described and illustrated, macro and microscopically, on the basis of two collections from Sicilian Mediterranean maquis. riassunto: Hebeloma lindae, taxon della sottosez. Scabrispora pubblicato recentemente sulla base di materiale raccolto in Spagna e Grecia, è segnalato per la prima volta in Italia. La specie è esaurientemente descritta ed illustrata, nei suoi caratteri macro-e microscopici, sulla base di due raccolte provenienti da ambiente mediterraneo siculo.
