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Study on Cortinarius subgenus Telamonia section Hydrocybe in Europe, with especial emphasis on Mediterranean taxa

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Abstract

In this paper we have attempted to clarify the taxonomy and nomenclature of thirteen taxa of the genus Cortinarius subgenus Telamonia (sections Hydrocybe, Fraternii) well represented in the southwestern Mediterranean area of Europe (C. atrocoeruleus, C. bombycinus, C casimiri, C. contrarius, C. decipiens, C. fraternus, C. gallurae, C. hoffmannii, C. petroselineus, C. sertipes, C. subturibulosus, C. urdaibaiensis and C. vernus). To this end we have performed a combined study of morphological and molecular data (rDNA ITS sequences). The morphological analysis was carried out on 114 collections and the molecular analysis involved 31 of the 114 collections, including 11 type collections (types for C. casimiri and C. fraternus were not available). In addition, a study of spores under field emission scanning electron microscopy (FESEM) was conducted. The results of the combined analysis allowed us to asign the studied material to five species (C. casimiri s.l., C. decipiens s.l., C. gallurae, C. subturibulosus s.l. and C. vernus s.l.). Thus, all collections from more continental areas, which were originally identified as six different taxa (C. atrocoeruleus, C. contrarius, C. decipiens, C. fraternus, C. sertipes, C. flexipes fo. sertipes) corresponded to C. decipiens sensu lato, a widely distributed, genetically and morphologically variable species. Cortinarius casimiri is also found in such habitats, but it is confirmed as distinct taxon. Collections from Mediterranean sclerophyllous communities correspond to C. gallurae, C. vernus sensu lato and C. subturibulosus sensu lato. Due to close phylogenetic relationships we propose the new combinations C. casimiri var. hoffmannii (=C. decipiens var. hoffmannii non C. hoffmannii) and C. subturibulosus var. bombycinus (=C. bombycinus), and the new variety C. vernus var. nevadavernus (=C. vernus H. Lindstr. & Melot sensu auct.).

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... Cortinarius punctatisporus Garnica morphologically also resembles C. egonii but differs with a clearly inflated stipe. Moreover, the ITS sequence similarity of C. egonii and C. punctatisporus is < 97% [8,38] PILEUS CONTEXT 800-900 µm wide, formed by two layers, the uppermost with inflated hyphae elements with a diam. Of (11) 15-31 (45) µm (n = 37), spongy looking, hyaline, colorless, thin-walled, hyphal walls not encrusted. ...
... The lack of reference data shows that the South American Cortinarius diversity is still widely underexplored and it also suggests the presence of endemic Cortinarius lineages in this area. Up until now there is very little overlap between Cortinarius species described from Australia and New-Zealand [38] with the species reported from South American Nothofagaceae forests. Given the high diversity of Southern Hemisphere Cortinarius diversity and the lack of available reference sequences, it is not surprising that the phylogeny of Cortinarius species remains largely unresolved. ...
... The diversity of Cortinarius from the Northern Hemisphere and from New-Zealand and Australia have been extensively studied (e.g., [24,38,68,[70][71][72]). However, aside from the pioneering work from Moser and Horak [5,26] there are relatively few studies that have focused on South American Cortinarius diversity [25]. ...
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This paper is a contribution to the current knowledge of taxonomy, ecology and distribution of South American Cortinarius (Pers.) Gray. Cortinarius is among the most widely distributed and species-rich basidiomycete genera occurring with South American Nothofagaceae and species are found in many distinct habitats, including shrublands and forests. Due to their ectomycorrhizal role, Cortinarius species are critical for nutrient cycling in forests, especially at higher latitudes. Some species have also been reported as edible fungi with high nutritional quality. Our aim is to unravel the taxonomy of selected Cortinarius belonging to phlegmacioid and myxotelamonioid species based on morphological and molecular data. After widely sampling Cortinarius specimens in Patagonian Nothofagaceae forests and comparing them to reference collections (including holotypes), we propose five new species of Cortinarius in this work. Phylogenetic analyses of concatenated rDNA ITS-LSU and RPB1 sequences failed to place these new species into known Cortinarius sections or lineages. These findings highlight our knowledge gaps regarding the fungal diversity of South American Nothofagaceae forests. Due to the high diversity of endemic Patagonian taxa, it is clear that the South American Cortinarius diversity needs to be discovered and described in order to understand the evolutionary history of Cortinarius on a global scale.
... The most widely used gene region for studies at species rank is the internal transcribed spacer region (ITS), which has proven to be useful in Cortinarius in subgenus Telamonia by Kytö vuori et al. (2005), Lindström et al. (2008), Niskanen et al. (2009) and Suárez-Santiago et al. (2009), in subgenus Phlegmacium by Garnica et al. (2009), Frøslev et al. (2007 and Ortega et al. (2008) and in subgenus Myxacium by Peintner (2008). In most studies the majority of species differ with respect to the degree of intra-and interspecific variation, and most polymorphic species typically do not have closely related sister taxa; the amount of intraspecific variation reported usually was fewer than six base pairs (bp) while the interspecific variation was more than six bp . ...
... Nearly all the aforementioned studies found morphologically indistinguishable subgroups inside morphologically delimited species. Unlike most studies comparing taxa at species rank, Ortega et al. (2008) and Suárez-Santiago et al. (2009) also compared infraspecific varieties. In their study species differed by at least 10 diagnostic positions and varieties by 2-9 diagnostic positions. ...
... Base polymorphisms are marked with ambiguous IUB codes and length polymorphisms with N (information provided on request). Frøslev et al. (2007), Niskanen et al. (2009) and Suárez-Santiago et al. (2009) reported the intragenomic variation in Cortinarius. The sequences were compared with the material in public databases (GenBank: http://www. ...
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Cortinarius sect. Armillati (subgenus Telamonia) was studied extensively based on morphology and molecular data. A total of about 1000 specimens, mostly from Fennoscandia, were revised. The nomenclature of the species was confirmed by sequencing the type material. Phylogenetic relationships were inferred by analyses of ITS, and the results were compared with the morphological and ecological data. Based on macro- and micromorphological characters, as well as molecular data, section Armillati contains only the medium to large species with slightly hygrophanous pileus and ± reddish or in some species yellowish brown to rose brown universal veils. The other red-brown-veiled species, previously included in Armillati, seem to belong to at least seven different sections or clades: sect. Anthracini, sect. Boulderenses, sect. Brunneotincti p.p., sect. Cinnabarini, sect. Fulvescentes, /Fuscoperonatus, and /Praestigiosus. Our study recognized six Armillati species from northern Europe: C. armillatus, C. luteo-ornatus, C. paragaudis, and three species described as new, C. pinigaudis, C. roseoarmillatus, and C. suboenochelis. The former three also occur in North America. Two additional species, C. subarmillatus (Japan) and C. quercoarmillatus (Costa Rica), are known outside the area. Based on the phylogenetic analysis, the species associated with deciduous trees, C. armillatus, C. quercoarmillatus, and C. roseoarmillatus, all with dextrinoid, thick-walled spores, formed a separate group from the mainly conifer-associated species, C. luteo-ornatus, C. paragaudis, C. pinigaudis and C. suboenochelis, all with fairly thin to moderately thick-walled, indextrinoid to moderately dextrinoid spores. Descriptions of the northern European species are provided, the distribution is mapped and their taxonomy, ecology, distribution, and relationships are discussed. A total of 64 new sequences of 12 species are reported including 17 sequences from type material. Our study also suggests that ITS sequences are not always sufficiently variable for species-rank recognition (barcoding) in Cortinarius.
... Загальне поширення. Європа: Австрія, Велика Британія, Данія, Естонія, Ірландія, Іспанія, Італія, Люксембург, Нідерланди, Німеччина, Норвегія, Польща, Португалія, Росія, Словенія, Україна, Фінляндія, Франція, Швейцарія, Швеція; Азія: Росія (Сибір, Далекий Схід); Африка: Марокко; Північна Америка: Канада, США; Нова Зеландія (Nezdoiminogo, 1996;Suárez-Santiago et al., 2009;Niskanen et al., 2012; https://www.gbif.org/species/3348211). ...
... Наприкінці варто зауважити, що описані у статті таксони досить широко розповсюджені у багатьох європейських країнах, а якщо й трапляються спорадично, проте не належать до числа надто рідкісних (Breitenbach, Kränzlin, 2000;Suárez-Santiago et al., 2009;Niskanen et al., 2012). З високою ймовірністю в Україні вони теж трапляються частіше, аніж нам відомо на сьогодні. ...
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Data on some noteworthy finds of interesting representatives of the subgenus Telamonia of the genus Cortinarius in Ukraine are presented. Two of them, C. decipiens var. atrocoeruleus and C. flexipes var. flabellus, have been registered for the first time in Ukraine; the typical variety of the former species is still unknown in our country. Other two taxa are rarely reported in Ukraine; those are C. flexipes var. flexipes, known also from Left-Bank Polyssia, and C. diasemospermus, so far recorded only in the Ukrainian Carpathians. All taxa were found in Male Polissia National Nature Park (Khmelnytsky Region), C. diasemospermus was also collected in Shatsky National Nature Park. The data about their distribution in Ukraine and worldwide, detailed descriptions of macro- and microscopical features, as well as original photos of the fruit bodies and drawings of the spores are provided. It should be noted that all these taxa are quite common in appropriate habitats in Europe and usually are not considered rare. Probably in Ukraine they also occur more often than they are reported at present and new finds of these species and varieties can be expected.
... Gray is one of the largest genera within the Basidiomycetes with the number of species estimated at ca. 2000 (Kirk et al. 2008) and with a worldwide distribution. Although it is a well known genus, revisions (Garnica et al. 2002;Peintner et al. 2003;Frøslev et al. 2006a, b;Frøslev and Jeppesen 2008;Ortega et al. 2008Ortega et al. , 2009Niskanen et al. 2008Niskanen et al. , 2009Niskanen et al. , 2012aNiskanen et al. , b, 2013Vila et al. 2008;Suárez-Santiago et al. 2009;Bojantchev 2011a, b;Bojantchev and Davis 2011) have demonstrated that the diversity of the genus is greater than previously thought, which means that numerous new species are still to be expected. But the large number of taxa, together with the lack of reliable microscopic characters and the plasticity of the widely used macro-morphological characters make species delimitation within the genus difficult ). ...
... But the large number of taxa, together with the lack of reliable microscopic characters and the plasticity of the widely used macro-morphological characters make species delimitation within the genus difficult ). In recent years phylogenetic analyses based on molecular data have proved to be a useful and reliable method for the taxonomic delimitation of numerous species (Peintner et al. 2003;Niskanen et al. 2009Niskanen et al. , 2011Niskanen et al. , 2012aNiskanen et al. , b, 2013Ortega et al. 2008Ortega et al. , 2009Vila et al. 2008;Suárez-Santiago et al. 2009). ...
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We clarify the taxonomy and nomenclature of several taxa of the genus Cortinarius subgenus Phlegmacium. To this aim, we have used a combination of morphological and molecular data. The evolutionary relationships of the species were inferred by comparison of the nuITS by means of weighted maximum parsimony, maximum likelihood and two different types of Bayesian methods (with and without a priori alignments). Phylogenetic resolution and support of all or most of the species included in this study and their relationships were possible only when including the phylogenetic signal from ambiguously aligned regions in weighted maximum parsimony analyses (recoded INAASE characters) and when the analysis simultaneously optimized alignment and phylogeny (with BAli-phy). Three species are described as new, Cortinarius mediterraneensis, C. cistoglaucopus and C. palazonianus, and C. olivaecodionysae is proposed for C. dionysae f. olivaceus. Descriptions are provided for these taxonomic and nomenclatural novelties, along with discussions of morphological and phylogenetic affinities to closely related taxa. Scanning microphotographs of the basidiospores are provided for the discussed taxa, and color pictures of the basidiomes in their natural habitat are provided for C. cistoglaucopus, C. mediterraneensis and C. palazonianus.
... Gray i particularment del subgènere Telamonia (Fr.) Trog. Destaquem, dintre dels nombrosos estudis efectuats en aquest camp, els de Brandrud et al. (2018), Frøslev et al. (2005, Garnica et al. (2005), Harrower et al. (2011), Liimatainen et al. (2014, 2020, Peintner et al. (2004) i Suárez- Santiago et al. (2009). ...
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We describe Cortinarius aavae Liimat. & Niskanen, so far known from North America, which was recently discovered in a mixed forest of Pinus sylvestris and Quercus ilex in the Penyagolosa Massif, during the autumn of 2021. Ma cro- and microscopic descriptions, as well as iconography are provided for the studied species. In addition, nrITS sequence of the ribosomal DNA is compared with the closest ones found in the GenBank database.
... More recently, a number of studies employed molecular tools to unveil overlooked diversity in Mediterranean cortinarii and test the phylogenetic autonomy of previously published species described on the basis of morphology (Ortega et al. 2008, Suárez-Santiago et al. 2009, Clericuzio et al. 2017, Bellanger et al. 2018, Mahiques et al. 2018, Soop et al. 2019b). In the present paper, we investigate subg. ...
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Molecular phylogenies in the past decade have demonstrated that the described diversity of Cortinarius is still underestimated, especially outside continental and boreal ecoregions where the genus has been historically investigated. We tackled this issue by revisiting the so far unresolved subgenus Leprocybe, and focused on the largely unexplored Mediterranean hotspot of biodiversity. The sequencing and phylogenetic analysis of 161 vouchered collections from Austria, Cyprus, France, Germany, Italy and Spain, including 16 types, allowed for the delineation of 11 species in this lineage, three of them recognised as new to science and formally introduced as C. jimenezianus, C. selinolens and C. viridans spp. nov., respectively. Interestingly, the newly described species exhibit a strict Mediterranean distribution, and one of them is putatively endemic to the island of Cyprus, highlighting the remarkable potential of this neglected ecoregion to uncover further undescribed diversity of Cortinarius in the future. The present work also unveils 23 synonymies in this subgenus, as well as previously undetected crypticism within C. venetus. Next Generation Sequencing carried out on three old and contaminated holotypes, successfully decrypts their phylogenetic identity, including that of C. leproleptopus, finally settling the long-standing controversy over the taxonomic status of this species. A brief overview of each species in the subgenus is lastly provided and a key is proposed to facilitate the identification of presently known European taxa of Leprocybe in the field.
... Young specimens of C. pakistanicus may resemble several species in C. sect. Hydrocybe due to overall morphology, pigmentation and the presence of traces of veil in the surface of pilei (Suárez-Santiago et al. 2009), although our reconstructed phylogenetic tree showed that members of this section are phylogenetically unrelated to the new species. The type species of this section is C. decipiens Fr., which usually shows grayish lamellae and stipe cortex when young, traits that were not noticed on our specimens. ...
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The genus of basidiomycetous fungi Cortinarius occurs worldwide, from subtropical to boreal latitudes. Although molecular systematics has triggered the study of these fungi in the Americas and Europe in the last two decades, there is still limited research on its diversity in large portions of the planet, such as the high mountain ranges of Asia. Several collections of Cortinarius were made during mycological field trips conducted between 2014 and 2018 in pure oak forests in the Pakistan Himalayas. An integrative framework combining morphological and phylogenetic data was employed for their study. As a result, the two species C. pakistanicus and C. pseudotorvus are here described as new to science. Detailed macro-and micro-morphological descriptions, including SEM images of spores, and a molecular phylogenetic reconstruction based on nrITS sequence data are provided and used to discriminate the new species from morphologically and phylogenetically close taxa. Whereas our phylogenetic tree inference gave unequivocal support for the inclusion of C. pseudotorvus within C. sect. Telamonia, the assignment of C. pakistanicus to any known sections remained elusive. These species likely establish ectomycorrhizal associations with trees in the genus Quercus, making this type of forest in the Pakistan Himalayas a promising focus for future research on the diversity of Cortinarius. A peer-reviewed open-access journal MycoKeys Launched to accelerate biodiversity research RESEARCH ARTICLE Arooj Naseer et al. / MycoKeys 74: 91-108 (2020) 92
... Se dégage de ces résultats la nature fallacieuse du critère « présence/absence de teintes bleues » dans ces trois lignées, déjà relevée dans d'autres groupes de Telamonia (par ex. SUAREZ-SANTIAGO et al. 2009) et soupçonnée par certains mycologues bien avant les premiers travaux phylogénétiques (Melot, comm. pers.). ...
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Un travail récent de révision taxinomique des sections Bicolores et Saturnini du genre Cortinarius a conduit à redéfinir les contours de C. cinnamoviolaceus M.M. Moser. Cette espèce, jusqu’alors considérée comme un Bicolores typique, s’est avérée phylogénétiquement externe à cette section et morphologiquement intermédiaire entre Bicolores et Duracini. Les nombreux synonymes de C. cinnamoviolaceus identifiés par l’analyse ont permis de mieux cerner le concept de C. dolabratus Fr. et C. imbutus Fr. et de rattacher ces deux espèces aux sect. Bicolores et Saturnini, respectivement. Ce faisant, cette espèce « Canada dry » qui ressemble à un Bicolores, un Duracini ou à un Saturnini, mais qui n’appartient à aucune de ces trois sections, invite à réviser les critères définitoires de ces sections. Recent work on the taxonomic revision of Cortinarius sect. Bicolores and Saturnini has led to the redefinition of C. cinnamoviolaceus M.M. Moser. This species, previously considered a typical Bicolores, has been phylogenetically positioned outside this section and morphologically intermediate between Bicolores and Duracini. The numerous synonyms of C. cinnamoviolaceus identified by the analysis allowed better understanding of the concepts of C. dolabratus Fr. and C. imbutus Fr. and to assign these two species to sect. Bicolores and Saturnini, respectively. As such, this "Canada dry" species, which resembles a Bicolores, a Duracini or a Saturnini but does not belong to any of these three sections, invites us to revise the defining criteria of these three sections.
... The diversity of the genus Cortinarius (Pers.) Gray (1821: 627) in the Iberian Peninsula is slowly expanding due to the integration of phylogenetics within traditional morphological, anatomical and ecological studies (Ortega et al. 2008, Suárez-Santiago et al. 2009, Fernández-Brime et al. 2014. Particular attention has been given to Cortinarius species in protected areas such as the Cadí-Moixeró Natural Park. ...
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Cortinarius is one of the most species-rich, conspicuous and widespread genera of basidiomycete ectomycorrhizal fungi. While knowledge of the morphogenetic diversity of Cortinarius in central and northern Europe is rather extensive, less is known about the genus in other regions, such as the Iberian Peninsula. The present study assesses the taxonomical status of an unknown Cortinarius species from the Cadí-Moixeró Natural Park (north-eastern Iberian Peninsula), using morphological, phylogenetic and ecological data. As a result, C. uxorum is presented as new species in C. subg. Telamonia sect. Firmiores. It is phenotypically recognized by its medium to large-sized, stout basidiomata with persistent and copious whitish to silvery-white veil remnants on the pileus margin, and the ring-like stripe on the upper stipe. The restricted distribution of this species to a Natural Park highlights the need to consider fungi as a pivotal component of biodiversity in conservation policies.
... Por el contrario, valores bootstrap y PP más elevados se obtuvieron para nodos que principalmente conectan parejas de especies en los extremos del árbol. Este mismo escenario ha sido constatado en otros estudios filogenéticos, que trataban taxones de Telamonia (NISKANEN et al., 2008(NISKANEN et al., , 2011SUÁREZ-SANTIAGO et al., 2009;DIMA et al., 2014). Generalmente se acepta que valores de soporte estadístico bajo son resultado de muestreos de taxones pobres, señal filogenética baja del marcador molecular usado, baja calidad de los alineamientos de múltiples secuencias de ADN, etc. Para generar filogenias de Telamonia con alto soporte estadístico, tanto en nodos internos como externos, se deberían añadir al análisis más marcadores moleculares y más taxones. ...
Article
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The present work aims at proposing two novel Telamonia species that were discovered in the Cadí-Moixeró Natural Park, a protected area south of the Pyrenees. Cortinarius exsularis sp. nov. and Cortinarius garciae sp. nov. are hereby described based on conventional procedures as well as phylogenetic analyses using ITS molecular data. We also employ a method based on genetic distances to support our species delimitation approach.
... sections), bringing DNA sequence data as well as morphological and ecological data in accordance, as already done by several authors (e.g. Garnica et al. 2009Garnica et al. , 2011Suárez-Santiago et al. 2009;Niskanen et al. 2013a, b;Dima et al. 2014;Stensrud et al. 2014;Liimatainen et al. 2015). ...
Article
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Arbutoid mycorrhizas of Comarostaphylis arbutoides (Arbutoidea, Ericaceae) from neotropical montane forests are rarely described. To date, only mycorrhizal associations with the fungal species Leccinum monticola, Leotia lubrica and Sebacina sp. are known from literature. The genus Cortinarius is one of the most species-rich ectomycorrhizal taxa with over 2000 assumed species. In this study, two sites in the Cordillera de Talamanca of Costa Rica were sampled, where Com. arbutoides is endemic and grows together with Quercus costaricensis. Using a combined method of rDNA sequence analysis and morphotyping, 33 sampled mycorrhizal systems of Cortinarius were assigned to the subgenera Dermocybe, Phlegmacium and Telamonia. Specific plant primers were used to identify the host plant. Here, we present the phylogenetic data of all found Cortinarii and describe four of the arbutoid mycorrhizal systems morphologically and anatomically.
... It is a mountain to subalpine species growing in deciduous woodlands primarily under Salix L. and Betula L., Populus L. and Quercus L. but also under conifers such as Pinus L. and Picea Link. C. albonigrellus can be distinguished from C. decipiens by its abundant white cortina which constitutes a differentiated ring on the stipe (Breitenbach and Krӓnzlin, 2000;Suarez-Santiago, 2009). ...
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Cortinarius albonigrellus J. Favre, C. assiduus Mahiques, A. Ortega & Bidaud and C. percavus J. Favre are reported from Bingöl province as new records for Turkish mycobiota. Short descriptions and the photographs of the taxa are provided and discussed briefly.Türkiye Cortinarius'ları için Bingöl yöresinden üç yeni kayıt Özet Cortinarius albonigrellus J. Favre, C. assiduus Mahiques, A. Ortega & Bidaud ve C. percavus J. Bingöl yöresinden Türkiye mikobiotası için yeni kayıt olarak rapor edilmiştir. Taksonların kısa deskripsiyonu ve fotografları verilmiş ve kısaca tartışılmıştır.
... Cortinarius is one of the largest genera of Agaricales, with more than 4800 epithets and more than 300 taxonomic subdivisions (subgenera, sections, stirps) as recorded in the Index Fungorum (http://www.indexfungorum.org). Phylogenetic analyses of the genus helped to delimit taxonomic entities within the genus Peintner et al. 2002Peintner et al. , 2003Frøslev et al. 2006;Garnica et al. 2009;Suarez-Santiago et al. 2009;Niskanen et al. 2011;Niskanen et al. 2013a, b), but the morphologically delimited subgenera Phlegmacium, Telamonia, Cortinarius, Myxacium, Sericeocybe, Dermocybe or Leprocybe as proposed by several authors could not be confirmed by molecular methods (Liu et al. 1997;Hoiland and Holst-Jensen 2000;Peintner et al. 2001Peintner et al. , 2002Peintner et al. , 2004Frøslev et al. 2005;Garnica et al. 2005). The subgenus Telamonia consists of Telamonia s.str., and several morphologically resembling lineages with unresolved phylogenetic relationships (Moser 1952;Melot 1990, Melot 1990;Garnica et al. 2003Garnica et al. , 2005Peintner et al. 2004). ...
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Cortinarius microglobisporus spec. nov., a taxon with small, roundish basidiospores, is here described from a submediterranean habitat in Italy (Apeninnes). A full description, colour pictures of basidiomata, and line drawings of microscopic features are provided. Phylogenetic relationships and species delimitations were investigated using rDNA ITS sequences, morphological, and ecological data. This taxon shares morphological characters with several groups of Cortinarii, e.g., with subgenus Telamonia and with section Anomali. Our phylogenetic analyses place C. microglobisporus in close relationship to the species complex of C. illuminus. The section "Illumini" was proposed for C. illuminus by Niskanen et al., in Niskanen (2008), and was not validated until now. Now, C. microglobisporus is the second species belonging to the section "Illumini", a distinct section, with unresolved basal relationship, but clearly not belonging to subgenus Telamonia s. str.
... 1960Moser (e.g. , 1969Moser (e.g. -1970Moser (e.g. , 1983 from Central Europe, Høiland (1984), Brandrud (e.g., 1996Brandrud (e.g., , 1998, Brandrud et al. (e.g., 1990Brandrud et al. (e.g., , 2012 and Niskanen et al. (e.g., 2009Niskanen et al. (e.g., , 2011aNiskanen et al. (e.g., , 2013a mainly from northern Europe, Frøslev et al. (e.g., 2006Frøslev et al. (e.g., , 2007 from northern and Central Europe, Orton (1955Orton ( , 1958 from Great Britain, and Ortega et al. (2008) and Suárez-Santiago et al. (2009) from mediterranean area. Selected papers of some of the major contributors to Cortinarius systematics in North America include Peck (1873; also see Gilbertson 1962), Kauffman (1918Kauffman ( , 1923Kauffman ( , 1932, Smith (1939Smith ( , 1942Smith ( , 1944, Ammirati (1972), Ammirati et al. (2013), Garnica et al. (2009), Liu et al. (1997), Moser & Ammirati (1996, 1999, Moser et al. (1995), Bojantchev (2011a, b) and Niskanen et al. (2013b, c). ...
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Cortinarius is a species-rich and morphologically challenging genus with a cosmopolitan distribution. Many names have not been used consistently and in some instances the same species has been described two or more times under separate names. This study focuses on subg. Phlegmacium as traditionally defined and includes species from boreal and temperate areas of the northern hemisphere. Our goals for this project were to: i) study type material to determine which species already have been described; ii) stabilize the use of Friesian and other older names by choosing a neo- or epitype; iii) describe new species that were discovered during the process of studying specimens; and iv) establish an accurate ITS barcoding database for Phlegmacium species. A total of 236 types representing 154 species were studied. Of these 114 species are described only once whereas 40 species had one ore more synonyms. Of the names studied only 61 were currently represented in GenBank. Neotypes are proposed for 21 species, and epitypes are designated for three species. In addition, 20 new species are described and six new combinations made. As a consequence ITS barcodes for 175 Cortinarius species are released.
... Molecular studies based on the ITS1 and ITS2 regions of the rDNA, in combination with morphological and ecological data, have been applied successfully for species delimitation in the Agaricales including the large genus Cortinarius (e.g., Eberhardt and Beker 2010;Garnica et al. 2011;Frøslev et al. 2007;Niskanen et al. 2011;2013a;Suárez-Santiago et al. 2009). Traditional subgeneric classification of Cortinarius based largely on morphological characters (e.g., Bidaud et al. 1994;Consiglio et al. 2007; Consiglio 2012) is not supported by recent molecular studies (e.g., Garnica et al. 2005;Peintner et al. 2004) and should be mostly discarded. ...
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Two new Cortinarius species are described from European Quercus forests and one new combination is made based on molecular and morphological data. Cortinarius uraceomajalis is a vernal species currently only known from Hungary, Bulgaria and Italy, but likely is common throughout central and south-eastern Europe. Cortinarius uraceonemoralis is a widely distributed species in Europe. The new combination Cortinarius nolaneiformis is based on Hydrocybe nolaneiformis Velen. and is a widespread vernal species in Europe. Cortinarius uraceomajalis and C. nolaneiformis are preliminarily placed in sect. Colymbadini, characterized by having a positive (yellow) UV reaction, while C. uraceonemoralis with a UV negative reaction is placed in sect. Uracei. A neotype is selected for C. colymbadinus and C. uraceus to stabilize the nomenclature. Taxonomic novelties: Cortinarius uraceomajalis Dima, Liimat., Niskanen & Bojantchev, Cortinarius uraceonemoralis Niskanen, Liimat., Dima, Kytöv., Bojantchev & H. Lindstr., Cortinarius nolaneiformis (Velen.) Dima, Niskanen & Liimat.
... In recent years, many species-level studies of Cortinarius, using morphological and molecular data, have been done in Europe (e.g., subgenus Telamonia: Kytövuori et al. 2005a;Lindström et al. 2008;Niskanen et al. 2009Niskanen et al. , 2011aSuárez-Santiago et al. 2009;subgenus Phlegmacium: Frøslev et al. 2007;Ortega et al. 2008;subgenus Myxacium: Peintner 2008;subgenus Cortinarius: Niskanen et al. 2012), but only few papers including North American material have been published (Moser and Peintner 2002;Matheny and Ammirati 2006;Garnica et al. 2009Garnica et al. , 2011Harrower et al. 2011). ...
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Five new Cortinarius species with medium to large basidiomata are described based on morphological and molecular data. Three of them, Cortinarius aavae, Cortinarius brunneocalcarius, and Cortinarius grosmorneënsis, belong to subgenus Telamonia and one, Cortinarius subfloccopus, to clade /Fulvescentes. The relationship of the fifth species, Cortinarius brunneotinctus, was not solved but it resembles species of clade /Anomali. Cortinarius brunneocalcarius and C. aavae do not have close relatives within the subgenus Telamonia; whereas, C. grosmorneënsis belongs to section Brunnei. Cortinarius grosmorneënsis is only known from eastern Canada, but the other four species have a wide distribution: C. brunneocalcarius and C. subfloccopus occur in North America and Europe, and C. brunneotinctus and C. aavae in western and eastern North America. The descriptions of the novel species are presented and comparison to similar species provided.
... In Funga Nordica, Knudsen and Vesterholt (2008) placed the section Colymbadini in subgenus Telamonia, C. cotoneus and relatives in the subgenus Cortinarius, section Veneti Korad & Maubl. Molecular studies in Cortinarius have been valuable for defining species as well as determining their phylogenetic relationships (Høiland and Holst-Jensen 2000; Garnica et al. 2003 Garnica et al. , 2005 Garnica et al. , 2009 Peintner et al. 2004; Frøslev et al. 2007; Ammirati et al. 2007; Niskanen et al. 2009; Suárez-Santiago et al. 2009). Garnica et al. (2003) developed a phylogenetic framework for species of Phlegmacium in Europe, and Ammirati et al. (2007) were able to place new species of Cortinarius from Costa Rica within a molecular phylogeny in relation to known species. ...
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Cortinarius parkeri, a new vernal species in subgenus Cortinarius, section Veneti, is described from the Pacific Northwest of North America. Within its subgenus, C. parkeri is unique in that its membranous universal veil is attached to the pileus edge of young mushrooms and extends as a covering over the pileus surface. At maturity, the universal veil forms a sheathing membranous volva-like structure that flares out above the stipe base. Sequences of the nuclear ribosomal DNA internal transcribed spacer region (ITS1-5.8s-ITS2) determined for 13 collections of C. parkeri were all identical, except for a polymorphic two base pair indel. In a phylogeny including other related species from North America and Europe, C. parkeri is monophyletic. In contrast to intraspecific distances, the interspecific genetic distance between C. parkeri and other Veneti species sequences was large. Being dikaryotic, mushrooms of C. parkeri contain genomes from two parents. As expected in a dikaryon in an interbreeding population, the polymorphic indel was homozygous in some collections but heterozygous in three collections, with conflicting sequences resulting from mixed parental types. Taken together, morphological and molecular results strongly support C. parkeri as a new and distinctive species.
... This level of similarity worked well for species recognition in other basidiomycetes (Ryberg et al. 2008; Hughes et al. 2009; Wright et al. 2009 ). However, intraspecific polymorphism in Cortinarius ITS regions can be up to 6 bp, about 1%, while interspecific variation can be low, as little as 3 bp (<1%) (Frøslev et al. 2007; Garnica et al. 2009; Niskanen et al. 2009; Suarez-Santiago et al. 2009). Further, instead of a " barcode gap " , a clear dividing line between interspecific and intraspecific variation, Harrower (2010) found continuous variation within and among Cortinarius species. ...
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Throughout the world, the diversity of fungi remains poorly characterized and Cortinarius is a classical example of a difficult, species-rich, and under-researched mushroom genus. Here, we sequenced and analyzed ribosomal internal transcribed spacer (ITS) sequence barcodes from herbarium specimens to improve understanding of Cortinarius species diversity in British Columbia (B.C.), Canada. Starting with 962 specimen sequences, 617 from B.C. herbaria, we present a maximum likelihood tree showing 179 putative Cortinarius species in British Columbia. As a working definition, we considered a “species” to be a monophyletic clade that included a reliably identified reference sequence, with a maximum of 3% ITS sequence variation. If no reference sequence was available, “species” were groups sharing 97% or more sequence identity. By these criteria, 110 putative B.C. species matched European species and 12 B.C. species matched species exclusively found in the Americas. Of the 56 B.C. species that did not match an identified reference sequence, some may be new to science, while others likely represent described species without available sequences. By depositing sequences from B.C. specimens into GenBank and BOLD, and by providing our alignment to TreeBASE, we have supplied the resources necessary to improve accuracy in identifications of Cortinarius in future systematic and ecological studies.
... Molecular methods have provided new tools for understanding the diversity and relationships of Cortinarius species, consequently an increasing number of studies on Cortinarius have been published in recent years (e.g. Frøslev et al. 2007;Ortega et al. 2008; Garnica et al. 2009;Suárez-Santiago et al. 2009;Niskanen et al. 2011aNiskanen et al. , 2012a. The studies by Kytö vuori et al. (2005a) and Niskanen et al. (2008Niskanen et al. ( , 2009Niskanen et al. ( , 2011bNiskanen et al. ( , 2012b in particular have examined the brown Telamonia species. ...
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Cortinarius bovinus and morphologically similar conifer-associated species were studied using material mainly from northern Europe. To stabilize the nomenclature, relevant types were examined. Phylogenetic relationships and species limits were investigated with rDNA ITS and nuclear rpb2 sequences as well as morphological data. We recognize seven species: C. bovinus (neotypified) and six new species, C. anisochrous, C. bovinaster, C.bovinatus, C. fuscobovinus, C. fuscobovinaster and C. oulankaënsis. Their taxonomy, ecology, distribution and relationships are discussed, and a key to species is provided. Based on our phylogeny and morphological data the species were placed in section Bovini.
... The most extensive taxonomic studies have concentrated on European species (e.g. Frøslev et al. 2007;Ortega et al. 2008;Niskanen et al. 2009Niskanen et al. , 2011Suárez-Santiago et al. 2009). Fewer studies have been completed for North American species and these have focused primarily on subgenus Phlegmacium (Moser and Peintner 2002;Garnica et al. 2009Garnica et al. , 2011. ...
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The North American species of Cortinarius section Sanguinei were studied using morphological characters and ITS and RPB2 sequence data. Several type collections were also examined. Four species were identified: C. harrisonii sp. nov, C. neosanguineus sp. nov., C. sanguineus, and C. sierraensis comb. nov. Of these, C. sanguineus also occurs in Europe together with C. puniceus, a fifth member of the section. Typical features of these species include ± red, fairly small basidiomata, stipe basal mycelium often with yellow to reddish yellow tints, amygdaloid to ellipsoid spores, and aniline-red lamellar trama and pileipellis hyphae when mounted in KOH. Two other species with red lamellae C. marylandensis comb. nov. and C. smithii stat. nov. & nom. nov. are also discussed.
... Recently, the use of molecular data has facilitated taxonomic work. Several studies on Telamonia species combining morphological and molecular data have been published by, e.g., Ammirati et al. (2007), Kytövuori et al. 2005, Lindström et al. (2008, Matheny and Ammirati (2006), Niskanen et al. (2006, 2008b), and Suárez-Santiago et al. (2009. The molecular markers used have been the rDNA ITS1 and ITS2 regions, which have also proved to be suitable species-level markers in other genera of Agaricales (e.g. ...
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A new species, Cortinarius badiolaevis Niskanen, Liimat., Mahiques, Ballarà, Kytöv. in subgenus Telamonia, is described based on morphological and molecular data. Typical for the species is a dark brown cap, pale flesh, indistinct universal veil, small amygdaloid to ellipsoid spores [7.0–8.5 × 4.5–5.0(−5.5) µm], and dark exsiccata. It occurs in mesic to dryish coniferous forests, and grows in calcareous soil presumably with Pinus and Picea. So far, it is only known from Spain and Sweden, and considered rare. Based on the phylogenetic analyses of the ITS regions, it does not belong to any of the known sections, nor does it have any very closely related sister species. KeywordsTaxonomy–ITS–DNA
... The introduction of DNA sequence characteristics in fungal taxonomy has been a significant improvement. The most widely used regions for studies at species rank are rDNA ITS1 and ITS2, which have proven useful for species delimitation in Cortinarius by for example Frøslev et al. (2007), Garnica et al. (2009), Kytö vuori et al. (2005), Lindströ m et al. (2008, Niskanen et al. (2009, 2011), Ortega et al. (2008 and Suárez-Santiago et al. (2009). These regions also have been proposed as species-identifier sequences (barcodes) in Cortinarius (Frøslev et al. 2007, Ortega et al. 2008. ...
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The red species of Cortinarius subgenus Dermocybe in Europe were studied based on morphological and molecular data. Three completely red species were recognized: C. sanguineus (syn. C. sanguineus var. aurantiovaginatus), C. puniceus (syn. C. cruentus, C. rubrosanguineus) and C. vitiosus comb. nov. Cortinarius sanguineus has dusky red to red pileus, reddish yellow mycelium and lacking or with only slightly encrusted hyphae in pileipellis. It occurs in mesic to damp forests with Picea, often on rich soil in the boreal and montane areas of Europe, presumably also in eastern Canada. Cortinarius puniceus differs from C. sanguineus by its stronger purplish red, narrower spores and spot-like encrusted hyphae in pileipellis. It grows with deciduous trees in the temperate zone of Europe. Cortinarius vitiosus is known only from Fennoscandia and occurs in dry to mesic coniferous forests. It has fairly thin, often zonate, dark red to dark reddish brown pileus, pale red mycelium, small spores and encrusted lamellar trama and pileipellis hyphae. In addition to these three species C. fervidus and C. phoeniceus occasionally have red basidiomes. The relationships of the species were inferred by analysis of ITS sequences. Our study suggests that the section Sanguinei, as earlier defined, is polyphyletic. Here the section is limited to include C. sanguineus, C. puniceus and North American D. sierraensis. The relationships with other red species were not determined. Section Dermocybe, including C. cinnamomeus, C. croceus and C. uliginosus, formed a monophyletic group, and the section Malicoriae had some support. A total of 34 new sequences are published including nine from type specimens.
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Alonso, J., Rigueiro, A., (2020). Catálogo da macromi-cobiota das montañas do Courel (Galicia, NO España). Monografías do Ibader - Serie Cadernos da Estación Científica do Courel. Ibader. Universidade de Santiago de Compostela. Lugo. Resumo Os traballos de catalogación da biodiversidade son imprescindibles como base de coñecemento para garantir a sustentabilidade do medio natural, así como para a definición e aplicación de proxectos de xestión e desenvolvemento dos territorios. Este traballo é unha aproximación ao coñecemento e catalogación da macromicobiota correspondente ás divisións Basidiomycota e Ascomycota das montañas do Courel, inventariando e describindo 410 taxons para o territorio, que xunto con 126 distintos taxons previamente citados por outros autores, e tamén recollidos neste traballo, fan un total de 536 taxons referenciados, un importante número reflexo da gran biodiversidade fúnxica das montañas do Courel, o que se corresponde coa diversidade xeolóxica, climática e florística deste territorio excepcional. Dos 410 taxons descritos, 322 son primeiras citas publicadas para o territorio das montañas do Courel, 109 para a provincia de Lugo, 48 para Galicia, 5 para a península ibérica, e unha é primeira cita europea: Simocybe serrulata. O estudo realizouse durante algo máis de 3 anos (outono de 2016 ata finais 2019) de periódico traballo de prospección micolóxica no territorio, complementado nos laboratorios da Estación Científica do Courel e do Departamento de Produción Vexetal e Proxectos de Enxeñaría da Escola Politécnica Superior de Enxeñaría (USC-Campus Terra de Lugo). Palabras clave micobiota, Basidiomycota, Ascomycota, biodiversidade fúnxica, Lugo. Abstract Cataloguing works about biodiversity are essential as a knowledge base in order to guarantee the natural environment sustainability, as well as for the definition and application of projects of management and development of the territories. This work is an approach to the knowledge and cataloguing of the macromycobiota of the divisions Basidiomycota and Ascomycota of the Courel montains, describing 410 taxa for the territory, which together with 126 taxa previously record by other authors, and also collected in this work, make a total of 536 referenced taxa, an important number reflecting the great fungal biodiversity of the Courel mountains, which corresponds to the geological, climatic and floristic diversity of this exceptional territory. Of the 410 descripted taxa, 323 are first time published records in the territory of Courel mountains, 109 in the province of Lugo, 48 in Galicia, 5 in the Iberian Peninsula, and one in Europe: Simocybe serrulata. The study was made during a bit more than 3 years (autumn 2016, end of 2019) via periodic work of mycologic prospection in the territory, complemented in the labs of the Courel Scientific Station and the Department of Crop Production of the Higher Polytechnic Engineering School (USC-Campus Terra, Lugo). Keywords mycobiota, Basidiomycota, Ascomycota, fungal biodiversity, Lugo.
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Mycologue emblématique de la région, Georges Chevassut (1923-2003) a incontestablement marqué l'histoire de la SHHNH de la fin du siècle dernier, au moins de notre section Mycologie. Il s'est imposé en trois décennies comme le spécialiste incontournable des cortinaires méditerranéens, dont il a décrit de nombreuses espèces nouvelles pour la science. Nous proposons ici un inventaire inédit ainsi qu'une analyse qualitative détaillée de cette production, à la lumière des travaux récents de taxonomie moléculaire qui redéfinissent les contours et le contenu du genre Cortinarius Abstract: Georges Chevassut (1923-2003), an emblematic mycologist of our Region, undoubtedly marked the history of the SHHNH at the end of the last century, at least of our Mycology section. Across three decades, he became the leading specialist of Mediterranean cortinarias, of which he described many species new for science. We propose here an unpublished inventory as well as a detailed qualitative analysis of this production, in light of recent molecular taxonomy works which define new boundaries and content of the genus Cortinarius
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Cortinarius is probably the largest genus among macromycetes, with an estimated number of species being more than 3,000. The taxonomy is problematical with even experts having difficulty in identifying to species level or group. Most of the literature also provides a level of detail that beginners find excessive and confusing. «Guida introduttiva al genere Cortinarius in Europa» is aimed at those interested in these fungi, whether as a keen mycologist or from casual interest. It will enable them to identify the species described, using dichotomous keys, detailed descriptions and accompanying illustrations. The excellent artwork is by the talented Italian mushroom illustrator Riccardo Mazza. The treatment follows traditional lines but incorporates the results of recent phylogenetic studies, cross referenced to an extensive bibliography.
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Extended areas on the island of Cyprus including the vast majority of forested land, are occupied by Pinus brutia, an ecologically important tree forming ectomycorrhizal associations with a wide array of fungi. In this third installment of macromycete inventories from selected ecosystems in Cyprus, the basidiomycete diversity within P. brutia communities is reported following a 12-y-inventory. Two hundred and thirty-one taxa are compiled in an annotated checklist, 95 of which constitute new records for the country. Most notable of these are Clitopilus daamsii, Hygrophorocybe nivea, Lepiota lepida, Leucoagaricus georginae, Rhodocybe matesina, Russula werneri, Tephrocybe striaepilea, Tephroderma fuscopallens, and Tricholoma chrysophyllum, which are rarely reported in published literature. Notes on the altitude, phenology and estimated abundance are provided, accompanied by selected imagery and a review of previously published records.
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So far approximately 144,000 species of fungi have been named but sequences of the majority of them do not exist in the public databases. Therefore, the quality and coverage of public barcode databases is a bottleneck that hinders the study of fungi. Cortinarius is the largest genus of Agaricales with thousands of species worldwide. The most diverse subgenus in Cortinarius is Telamonia and its species have been considered one of the most taxonomically challenging in the Agaricales. Its high diversity combined with convergent, similar appearing taxa have earned it a reputation of being an impossible group to study. In this study a total of 746 specimens, including 482 type specimens representing 184 species were sequenced. Also, a significant number of old types were successfully sequenced, 105 type specimens were over 50 years old and 18 type specimens over 100 years old. Altogether, 20 epi-or neotypes are proposed for recently commonly used older names. Our study doubles the number of reliable DNA-barcodes of species of C. subgenus Telamonia in the public sequence databases. This is also the first extensive phylogenetic study of the subgenus. A majority of the sections and species are shown in a phylogenetic context for the first time. Our study shows that nomenclatural problems, even in difficult groups like C. subgenus Telamonia, can be solved and consequently identification of species based on ITS barcodes becomes an easy task even for non-experts of the genus.
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ALONSO DIAZ, J. (2016): Algunas especies de macromicetos presentes en un jaral de Cistus ladanifer L. en el municipio de A Rúa de Valdeorras (Ourense, NO península ibérica) III. Micolucus 3: 7-20. ABSTRACT: In this article previous studies on macromycetes present in a cistus maquis of Cistus ladanifer L. located in the municipality of A Rúa de Valdeorras (Ourense, Galicia, Spain, NW Iberian Peninsula) are extended with the records and descriptions of 9 taxa, one of them specifically associated with Cistus spp.: Hebeloma cistophilum Maire, and the others are 8 non exclusive taxa of this habitat: Amanita curtipes f. curtipes E.-J. Gilbert, Amanita pantherina (DC.) Krombh., Astraeus hygrometricus (Pers.) Morgan, Boletus aereus Bull., Cortinarius gallurae D. Antonini, M. Antonini & Consiglio, Cerioporus meridionalis (A. David) Zmitr. & Kovalenko, Scleroderma polyrhizum (J.F. Gmel.) Pers. and Thelephora terrestris Ehrh., being Cortinarius gallurae and Cerioporus meridionalis the first records for Galicia.
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New morphological and molecular data are provided for several representatives of subg. Phlegmacium sect. Calochroi. We propose a detailed differential diagnosis for each considered species and a key to them, based on material from the Iberian Peninsula. Various algorithms of phylogenetic analysis and species delimitation were used to analyse the molecular dataset. Finally, we suggest the use of an integrative taxonomy approach to the general study of Cortinarius.
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This dataset contains the specimen records from the catalogue of the nomenclature types of fungi and lichens in the Herbarium of the University of Granada, Spain. These herbarium specimens are included in the GDA and GDAC collections, acronyms from Index Herbariorum (Thiers 2014). At this time, the type collection of fungi and lichens contains 88 type specimens of 49 nominal taxa, most from Agaricales and the genus Cortinarius, described from the western Mediterranean, mainly Spain, by the following authors: V.Antonin, J.Ballarà, A.Bidaud, G.F.Bills, M.Bon, C.Cano, M.Casares, G.Chevassut, M.Contu, F.Esteve-Raventós, R.Galán, L.Guzmán-Dávalos, R.Henry, E.Horak, R.Mahiques, G.Malençon, P.Moënne-Loccoz, G.Moreno, A.Ortega, F.Palazón, V.N.Suárez.-Santiago, A.Vêzda, J.Vila, and M.Villareal. For each specimen, the locality indication, species name, observation date, collector, type status, related information, associated sequences, other catalogue numbers related to each type, and image URL are recorded. The dataset is associated with an image collection named “Colección de imágenes de los tipos nomenclaturales de hongos, líquenes, musgos y algas incluidos en el Herbario de la Universidad de Granada (GDA y GDAC)” (Vizoso and Quesada 2013) which is housed and accessible at the Global Biodiversity Information Facility in Spain (GBIF.ES) Hosting and Publishing Service “Biodiversity Image Portal of Spanish collections” and is also available at the Herbarium of University of Granada institutional web (Vizoso 2014a, Vizoso 2014b). That image collection contains 113 images, of which 56 correspond to the nomenclature types of 49 taxa (47 fungi, 2 lichens), the rest of the images in this collection correspond to documents and specimens or microscopy photographs which are included in the herbarium specimens of fungi. These complement and document the process of the typification.
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Cortinarius is an ectomycorrhizal Agaricales genus with high diversity of which rDNA se- quences of 86 species together with four outgroup taxa were investigated phylogenetically by aid of Maximum Likelihood and Bayesian analyses. The Cortinarius data set represents 81 taxa from the Northern Hemisphere showing the main variation spectrum among the species. In addition, five species from the Southern Hemisphere are included. The phy- logenetic tree of Cortinarius gives statistical support to twelve monophyletic groups in the upper level. They are discussed in context of morphology, chemistry (secondary com- pounds), and ecology. The phylogenetic tree lacks, however, satisfactory support for its backbone. Several species could not be included in any group, especially those forming the basal framework of the tree. Of special interest is a “superclade” comprising eight of our monophyletic clades and two singletons. Here we find the majority of species with soluble pigments of octaketide origin, all species with compounds of nonaketide origin, the major- ity of species with hygrophaneous pileus, few species with viscid pileus, and no species with bulbous stipe base. Moreover, all species except one have duplex pileus cuticle. The morphological traits are not indicative for any clade, although some are more frequent in some clades than others. During the evolution they have been gained and lost several times. The chemical characteristics are – to a certain degree – more indicative for the clades. The evolution and ecological role of these compounds are discussed. Concerning the North European species, there are ecological differences between the clades, especially between clades specializing to rich or calcareous forests and clades specializing to poor forests or arctic-alpine environments.
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This paper presents a study about three taxa of the genus Cortinarius (Myxacium), little known in the mediterranean area of peninsular Spain: Cortinarius barbatus (Batsch: Fr.) Fr., Cortinarius croceocaeruleus (Pers.: Fr.) Fr. and Cortinarius croceocaeruleus var. meridionalis Bidaud, Ortega & Mahiques var. nov.
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Species of Cortinarius sect. Calochroi show a high grade of morphological variability in the Mediterranean region with regard to vicariant populations in the Eurosiberian region. This has contributed to confusion on taxonomy and nomenclature, especially when the variability affects the characters traditionally used for taxonomic delimitation. In this paper, we have attempted to clarify the taxonomy and nomenclature of several Calochroi taxa in the Mediterranean area by using macromorphological and spore analyses, as well as phylogenetic analysis of ITS sequences. We have demonstrated the usefulness of ITS sequences for species rank identification in Cortinarius (barcoding). We have been able to assign the studied material to eight species using ITS sequences (C. haasii, C. parasuaveolens, C. platypus, C. rickenianus, C. sancti-felicis, C. selandicus, C. splendidior and C. subgracilis), and to detect misidentified collections that were used by their authors to describe a new species (C. violaceipes). Moreover, molecular data have been useful to test the taxonomic value of morphological and spore characters. The morphological analysis has enabled us to detect morphological variability previously unreported in several taxa. A new variety, C. haasii var. quercus-ilicicola, is proposed on the basis of morphological and molecular data. Also, the taxonomic status of C. rickenianus is discussed. Finally, our results increase both the distribution area as well as the ecological preferences of five studied species, and several important biogeographical disjunctions are reported (C. selandicus and C. subgracilis).
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Studies of the Cortinarius saniosus (Fr.: Fr.) Fr. complex and a new closely related species, C. aureovelatus (Basidiomycota, Agaricales) The variation of the widely spread Cortinarius saniosus is described with respect to morphological, ecological and molecular characteristics. C. chrysomallus D. Lamoure and C. aureomarginatus P.D. Orton are regarded as synonymous with C. saniosus based on morphological and molecular data. A new closely related species C. aureovelatus Bendiksen, K. Bendiksen & H. Lindstr., growing in coniferous and mountain birch forests, is described. A taxonomical and nomenclatural treatment of the complex, based upon present knowledge, is proposed.
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Internal transcribed spacer (ITS) nuclear ribosomal DNA (rDNA) sequences of 27 species of Cortinarius representing the subgenera Cortinarius, Dermocybe, Leprocybe, Myxacium, Phlegmacium, Sericeocybe, and Telamonia were studied and compared with ITS rDNA of Rozites caperatus and three outgroup taxa: Gymnopilus sapineus, Hypholoma capnoides, and Tricholoma equestre. With two different gap treatments, the sequence data were used to reconstruct the phylogeny of Cortinarius using maximum parsimony (PAUP). Alternative outlines of the subgenera compiled from literature were used to create phylogenetic constraints which were subjected to parsimony analysis using the ITS data. Unconstrained and constrained parsimony trees were compared using a maximum likelihood method (DNAML/PHYLIP). In the ITS based phylogenies Cortinarius s. l. was consistently divided into two lineages, one centered around the members of the subgenus Telamonia, the other comprising the majority of ingroup taxa including Rozites. Because additional data are needed before nomenclatorial changes are considered, we present an outline of what may be regarded as two separate genera, Cortinarius s. s. and "Telamonia". The status of Rozites will depend on similar future studies.
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The genus Cortinarius Fr. (Cortinariaceae, Agaricales) is divided into four or more subgenera. Dermocybe (Fr.) Sacc. has been recognized as either a subgenus of Cortinarius or a separate genus, distinguished in part by the presence of various anthraquinonic pigments. Nucleotide sequences of ribosomal DNA 5.8S and internal transcribed spacers were used to investigate the phylogenetic relationships among species of Dermocybe and selected taxa from subgenera of Cortinarius. Sequence data from 47 herbarium specimens representing 31 taxa (28 species plus 3 varieties) of Dermocybe and Cortinarius were analyzed using parsimony, maximum likelihood, and neighbor joining. In general, molecular data support the morphological groupings of the taxa, although they more closely correspond to biochemical (anthraquinone and other) analyses. Phylogenetic trees showed that, while the sections Dermocybe and Malicoriae are monophyletic, and the concolorous or almost concolorous red species (section Sanguineae, such as D. sanguinea and relatives) together formed a coherent clade, the subgenus Dermocybe sensu lato itself is polyphyletic. Cortinarius californicus clusters with taxa in Cortinarius, subgenus Telamonia, section Armillati. Dermocybe olivaceopicta is more closely related to other subgenera of Cortinarius than to Dermocybe. Within the genus Cortinarius, certain of the subgenera may actually represent coherent genera. Of the subgenera examined, Telamonia, Phlegmacium, and possibly Sericeocybe appear to represent well defined taxonomic groupings. However, current assignments of taxa within Leprocybe and Myxacium were inconsistent with the molecular data. Reorganization of some taxa and taxonomic groups is suggested.
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Ten new species of Cortinarius are described from Costa Rican neotropical montane forests based on morphological and molecular phylogenetic data. Morphological features of the basidiomes and nuclear rDNA sequence analyses place these species in the following clades: Anomali (Cortinarius sericeolazulinus), Cotoneus (Cortinarius aureopigmentatus), Limonii (Cortinarius aurantiobrunneus), Lustrati (Cortinarius comarostaphylii, Cortinarius comparioides, Cortinarius ovreboi), Myxacium (Cortinarius costaricensis), and Telamonia (Cortinarius carranzae, Cortinarius quercoarmillatus, Cortinarius savegrensis). Species delimitations, diagnostic characters, sequence pair wise comparisons, and affinities to closely related species are included for each species. These Cortinarii occur with several species of Quercus with the exception of C. comarostaphylii, which is associated with Comarostaphylis (Ericaceae) in the páramo and in mixed forests with Quercus and Comarostaphylis.
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Phylogenetic relationships in the genus Cortinarius with emphasis on European species were studied by integrating a molecular phylogenetic analysis of nuclear rDNA sequences with morphological, chemical, and ecological data. Internal transcribed spacers (ITS1 and ITS2, including 5.8S) and the D1–D2 regions of nuclear rDNA of 262 Cortinarius species including most of the recognised subgenera from Europe, South America, Australia, Tasmania, New Zealand, and Asia were sequenced and analysed by maximum likelihood and neighbour joining. Our morphological and molecular analyses showed considerable disagreement with earlier morphology-based hypotheses concerning the infrageneric delimitations in Cortinarius. We propose a new base for a more natural classification system in Cortinarius involving the taxonomic rearrangement of the species into the following major lineages that we informally designate as “clades” at this time: (1) phlegmacioid species are included in Alluti, Amarescentes, Arguti, Calochroi, Caerulescentes, Percomes, Phlegmacioides, Phlegmacium, Purpurascentes, Scauri, and Vulpini; (2) the Heterocliti clade includes species of the subgenera Leprocybe and Cystogenes; (3) most of the Myxacium species are included in the Delibuti and Myxacium clades; (4) part of Rozites are included in the Rozites clade; (5) the north-hemispheric species of subgenus Dermocybe are included in the Dermocybe clade and part of the south-hemispheric species in the Splendidi clade; (6) part of Sericeocybe are integrated in the Anomali and Telamonia clades; (7) members of subgenus Cortinarius are assigned to the Cortinarius clade; (8) telamonioid taxa are included in the Obtusi, Renidentes, and Telamonia clades. Several Cortinarius species with peculiar patterns of morphology and basidiome coloration had isolated positions in the sequence analysis, and the evolutionary relationships among the major lineages remained unresolved. Our results suggest the convergent evolution of certain morphological traits across phylogenetically divergent lineages in Cortinarius. However, combinations of features such as pileipellis anatomy, spore shape, veil coloration, basidiome pigments, and stipe shape appear useful to circumscribe clades. Our molecular phylogenetic analysis allows insights in global geographical patterns of species distribution within the major lineages in the genus Cortinarius. The clade structure proposed here could be used as backbone for further more detailed systematic studies.
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The paper presents the results of a study of four species of the genus Cortinarius associated to Alnus glutinosa in montane communities. The study is complemented with drawings of micro-anatomical characters, SEM micrographs of basidiospores and colour photographs of basidiomata in their natural habitat.
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Phylogenetic analyses of non-protein-coding nucleotide sequences such as ribosomal RNA genes, internal transcribed spacers, and introns are often impeded by regions of the alignments that are ambiguously aligned. These regions are characterized by the presence of gaps and their uncertain positions, no matter which optimization criteria are used. This problem is particularly acute in large-scale phylogenetic studies and when aligning highly diverged sequences. Accommodating these regions, where positional homology is likely to be violated, in phylogenetic analyses has been dealt with very differently by molecular systematists and evolutionists, ranging from the total exclusion of these regions to the inclusion of every position regardless of ambiguity in the alignment. We present a new method that allows the inclusion of ambiguously aligned regions without violating homology. In this three-step procedure, first homologous regions of the alignment containing ambiguously aligned sequences are delimited. Second, each ambiguously aligned region is unequivocally coded as a new character, replacing its respective ambiguous region. Third, each of the coded characters is subjected to a specific step matrix to account for the differential number of changes (summing substitutions and indels) needed to transform one sequence to another. The optimal number of steps included in the step matrix is the one derived from the pairwise alignment with the greatest similarity and the least number of steps. In addition to potentially enhancing phylogenetic resolution and support, by integrating previously nonaccessible characters without violating positional homology, this new approach can improve branch length estimations when using parsimony.
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The aim of the present study was to investigate the phylogeny and evolution of sequestrate fungi (with gastroid or partially exposed basidiomes) in relation to their gilled relatives from the Cortinariaceae (Basidiomycetes). Phylogenetic analyses of 151 ITS sequences from 77 gilled species and 37 sequestrate taxa were performed using maximum parsimony and maximum likelihood methods. Results show that sequestrate basidiome forms occur in all three major ectomycorrhizal lineages of Cortinariaceae: the clades Cortinarius, Hebeloma/Hymenogaster/Naucoria, and Descolea. However, these forms do not appear within the saprobic outgroup Gymnopilus, indicating multiple origins of sequestrate forms from ectomycorrhizal ancestors. Additionally, within the Cortinarius clade sequestrate forms have multiple origins: emergent Cortinarius spp., Thaxterogaster, Quadrispora, Protoglossum, and two Hymenogaster spp. (H. remyi, H. sublilacinus) share common ancestors with Cortinarius spp., but these sequestrate genera are not closely related to each other (with exception of Thaxterogaster and Quadrispora). Hymenogaster sensu stricto, Setchelliogaster, and Descomyces were placed in the two other major clades. Thus, sequestrate taxa evolved independently many times within brown-spored Agaricales. Furthermore, emergent, secotioid, and gastroid forms have evolved independently from each other, and so are not necessarily intermediate forms. After their establishment, these apparently morphologically stable taxa show a tendency to radiate.
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Phylogenetic relationships of Rozites, Cuphocybe, and Rapacea were assessed using molecular phylogenetic approaches. These three genera are placed in Cortinariaceae and have been regarded as closely related to Cortinarius. Rozites includes more than 20 species, which are characterized by having both a membranaceous partial veil in the form of a persistent annulus and a membranaceous universal veil. Cuphocye (4 species) lacks an annulus or cortina, but has pigmented veil fibrils or scales. The monotypic genus Rapacea accommodates a distinct taxon with pale, nearly smooth and thick-walled basidiospores. We analyzed 56 sequences of the internal transcribed spacer region (ITS1, ITS2, and the intervening 5.8S rRNA gene) for nine species of Rozites, three species of Cuphocybe, 28 species of Cortinarius, Rapacea mariae and Protoglossum luteum. Two species of Hebeloma were used as outgroup. Large subunit (LSU) rDNA sequences from selected taxa were also analyzed. The results clearly demonstrate that Rozites species are nested within the clade/Cortinarius, and that Rozites is polyphyletic, suggesting that membranaceous veils have evolved several times in the genus Cortinarius. Also Rapacea and Cuphocybe are nested within Cortinarius, making the latter genus paraphyletic. Based on phylogenetic studies, Rozites, Cuphocybe and Rapacea are artificial genera and do not reflect natural relationships.
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Phylogenetic relationships of 54 European Phlegmacium species, including members of most of the sections of classical systematics, were studied, integrating macro-, micromorphological and chemical characters of the basidiomes, as well as molecular phylogenetic analysis of nuclear rDNA sequences. Microscopical structures of the basidiomes were studied by light microscopy. Basidiospore morphology was examined by scanning electron microscopy. Internal-transcribed spacers (ITS 1 and 2, including the 5.8S) and the D1/D2 (LSU) regions of nuclear rDNA were sequenced and analyzed with a Bayesian Markov chain Monte Carlo approach. Many subgroups detected by the molecular analysis are related to groups known from classical systematical concepts. Among others, these subgroups were significantly supported: i) a group containing most of the members of section Fulvi ss. Brandrud and the species Cortinarius arcuatorum, C. dibaphus and C. multiformis; ii) a group comprising taxa of section Calochroi ss. Brandrud and the species C. fulvocitrinus and C. osmophorus; iii) a group containing species of section Glaucopodes ss. Brandrud and C. caerulescens; iv) a group including members of section Phlegmacioides ss. Brandrud; v) a group that includes the species C. cephalixus, C. nanceiensis and C. mussivus. Stipe shape, color of flesh, pigment contents, KOH reaction on pileipellis and gelatinous layer, degree of development of a gelatinous layer on the pileipellis, and pileipellis structure were useful characters in delimiting subgroups in Phlegmacium, while basidiospore morphology was significant at species level. With the exception of C. glaucopus, C. infractus and C. scaurus, ITS and D1/D2 sequences obtained from collections of the same species from different geographical origins showed very little variation. Our molecular and morphological analyses suggest revisions of the traditional concepts of the subgenus Phlegmacium in Europe.
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Research on the molecular systematics of Cortinarius, a species-rich mushroom genus with nearly global distribution, is just beginning. The present study explores infrageneric relationships using rDNA ITS and LSU sequence data. One large dataset of 132 rDNA ITS sequences and one combined da-taset with 54 rDNA ITS and LSU sequences were generated. Hebeloma was used as outgroup. Bayesian analyses and maximum-likelihood (ML) analyses were carried out. Bayesian phylogenetic inference performed equally well or better than ML, especially in large datasets. The phylogenetic analysis of the combined dataset with species representing all currently recognized subgenera recovered seven well-supported clades (Bayesian posterior probabilities BPP > 90%). These major clades are: /Myxacium s.l., /subg. Cortinarius, the /phlegmacioid clade (including the subclades /Phlegmacium and /Delibuti), the /calochroid clade (/Calochroi, /Ochroleuci and /Allutus), the /telamonioid clade (/Telamonia, /Orellani, /Anomali), /Dermocybe s.l. and /Myxotelamonia. Our results show that Cortinarius consists of many lineages, but the relationships among these clades could not be elucidated. On one hand, the low divergence in rDNA sequences can be held responsible for this; on the other hand, taxon sampling is problematic in Cortinarius phylogeny. Because of the incredibly high diversity (~2000 Cortinarius species), our sampling included <5% of the known species. By choosing type species of subgenera and sections, our sampling is strongly biased toward Northern Hemisphere taxa. More extensive taxon sampling, especially of species from the Southern Hemisphere, is essential to resolve the phylogeny of this important genus of ectomycorrhizal fungi.
Article
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We sampled and analyzed approximately 2900bp across the three loci from 54 taxa belonging to a taxonomically difficult group of Cortinarius subgenus Phlegmacium. The combined analyses of ITS and variable regions of RPB1 and RPB2 greatly increase the resolution and nodal support for phylogenies of these closely related species belonging to clades that until now have proven very difficult to resolve with the ribosomal markers, nLSU and ITS. We present the first study of the utility of variable regions of the genes encoding the two largest subunits of RNA polymerase II (RPB1 and RPB2) for inferring the phylogeny of mushroom-forming fungi in combination with and compared to the widely used ribosomal marker ITS. The studied region of RPB1 contains an intron of the size and variability of ITS along with many variable positions in coding regions. Though almost entirely coding, the studied region of RPB2 is more variable than ITS. Both RNA polymerase II genes were alignable across all taxa. Our results indicate that several sections of Cortinarius need redefinition, and that several taxa treated at subspecific and varietal level should be treated at specific level. We suggest a new section for the two species, C. caesiocortinatus and C. prasinocyaneus, which constitute a well-supported separate lineage. We speculate that sequence information from RNA polymerase II genes have the potential for resolving phylogenetic problems at several levels of the diverse and taxonomically very challenging genus Cortinarius.
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We have conducted a taxonomic study of Cortinarius belleri, C. cistohelvelloides, C. impolitus and C. scobinaceus from material collected in Spain, France, Italy, Portugal and Morocco. Based on anatomy, morphology, ecology and distribution (geographical data) we recognize two species, C. impolitus and C. scobinaceus, and the new combination C. scobinaceus var. cistohelvelloides is made. Cortinarius belleri, C. cistohelvelloides, C. impolitus and C. scobinaceus have been typed, and a key for identification of these taxa is included.
Article
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Cortinarius is the most species rich genus of mushroom forming fungi with an estimated 2000 spp. worldwide. However, species delimitation within the genus is often controversial. This is particularly true in the section Calochroi (incl. section Fulvi), where the number of accepted taxa in Europe ranges between c.60 and c.170 according to different taxonomic schools. Here, we evaluated species delimitation within this taxonomically difficult group of species and estimated their phylogenetic relationships. Species were delimited by phylogenetic inference and by comparison of ITS sequence data in combination with morphological characters. A total of 421 ITS sequences were analyzed, including data from 53 type specimens. The phylogenetic relationships of the identified species were estimated by analyzing ITS data in combination with sequence data from the two largest subunits of RNA polymerase II (RPB1 and RPB2). Seventy-nine species were identified, which are believed to constitute the bulk of the diversity of this group in Europe. The delimitation of species based on ITS sequences is more consistent with a conservative morphological species concept for most groups. ITS sequence data from 30 of the 53 types were identical to other taxa, and most of these can be readily treated as synonyms. This emphasizes the importance of critical analysis of collections before describing new taxa. The phylogenetic separation of species was, in general, unambiguous and there is considerable potential for using ITS sequence data as a barcode for the group. A high level of homoplasy and phenotypic plasticity was observed for morphological and ecological characters. Whereas most species and several minor lineages can be recognized by morphological and ecological character states, these same states are poor indicators at higher levels.
Article
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We announce the release of the fourth version of MEGA software, which expands on the existing facilities for editing DNA sequence data from autosequencers, mining Web-databases, performing automatic and manual sequence alignment, analyzing sequence alignments to estimate evolutionary distances, inferring phylogenetic trees, and testing evolutionary hypotheses. Version 4 includes a unique facility to generate captions, written in figure legend format, in order to provide natural language descriptions of the models and methods used in the analyses. This facility aims to promote a better understanding of the underlying assumptions used in analyses, and of the results generated. Another new feature is the Maximum Composite Likelihood (MCL) method for estimating evolutionary distances between all pairs of sequences simultaneously, with and without incorporating rate variation among sites and substitution pattern heterogeneities among lineages. This MCL method also can be used to estimate transition/transversion bias and nucleotide substitution pattern without knowledge of the phylogenetic tree. This new version is a native 32-bit Windows application with multi-threading and multi-user supports, and it is also available to run in a Linux desktop environment (via the Wine compatibility layer) and on Intel-based Macintosh computers under the Parallels program. The current version of MEGA is available free of charge at (http://www.megasoftware.net).
Article
The recently-developed statistical method known as the "bootstrap" can be used to place confidence intervals on phylogenies. It involves resampling points from one's own data, with replacement, to create a series of bootstrap samples of the same size as the original data. Each of these is analyzed, and the variation among the resulting estimates taken to indicate the size of the error involved in making estimates from the original data. In the case of phylogenies, it is argued that the proper method of resampling is to keep all of the original species while sampling characters with replacement, under the assumption that the characters have been independently drawn by the systematist and have evolved independently. Majority-rule consensus trees can be used to construct a phylogeny showing all of the inferred monophyletic groups that occurred in a majority of the bootstrap samples. If a group shows up 95% of the time or more, the evidence for it is taken to be statistically significant. Existing computer programs can be used to analyze different bootstrap samples by using weights on the characters, the weight of a character being how many times it was drawn in bootstrap sampling. When all characters are perfectly compatible, as envisioned by Hennig, bootstrap sampling becomes unnecessary; the bootstrap method would show significant evidence for a group if it is defined by three or more characters.
Article
We have conducted a taxonomic study of Cortinarius belleri, C. cistohelvelloides, C. impolitus and C. scobinaceus from material collected in Spain, France, Italy, Portugal and Morocco. Based on anatomy, morphology, ecology and distribution (geographical data) we recognize two species, C. impolitus and C. scobinaceus, and the new combination C. scobinaceus var. cistohelvelloides is made. Cortinarius belleri, C. cistohelvelloides, C. impolitus and C. scobinaceus have been typed, and a key for identification of these taxa is included.
Article
Phylogenetic analyses of non-protein-coding nucleotide sequences such as ribosomal RNA genes, internal transcribed spacers, and introns are often impeded by regions of the alignments that are ambiguously aligned. These regions are characterized by the presence of gaps and their uncertain positions, no matter which optimization criteria are used. This problem is particularly acute in large-scale phylogenetic studies and when aligning highly diverged sequences. Accommodating these regions, where positional homology is likely to be violated, in phylogenetic analyses has been dealt with very differently by molecular systematists and evolutionists, ranging from the total exclusion of these regions to the inclusion of every position regardless of ambiguity in the alignment. We present a new method that allows the inclusion of ambiguously aligned regions without violating homology.In this three-step procedure, first homologous regions of the alignment containing ambiguously aligned sequences are delimited. Second, each ambiguously aligned region is unequivocally coded as a new character, replacing its respective ambiguous region. Third, each of the coded characters is subjected to a specific step matrix to account for the differential number of changes (summing substitutions and indels) needed to transform one sequence to another.The optimal number of steps included in the step matrix is the one derived from the pairwise alignment with the greatest similarity and the least number of steps. In addition to potentially enhancing phylogenetic resolution and support, by integrating previously nonaccessible characters without violating positional homology,this new approach can improve branch length estimations when using parsimony.
Article
A nomenclatural study of the names associated with type collections deposited in the mycological collection of IB was undertaken. Many of these names were invalidly published (Art. 34.2.) or had never been officially recombined. Our Mycological Type Collection Database can now be searched on the world-wide-web, and thus we decided to validate or recombine the following 18 species epithets in Cortinarius. Cortinarius alneus, C. atrocoeruleus, C. atropurpureus, C. betuletorum, C. caesiostramineus var. amarissimus, C. chrysophthalma, C. corynecystis, C. duramarus, C. fagetorum, C. flavotomentosus, C. heterochromus, C. multicolor, C. nothovenetus, C. obscurooliveus, C. obscurooliveus var. brunneus, C. olivipes, C. subtriumphans var. cephalixoides, C. viridipes. We give an overview of the valid names of nomenclatural types regarded as synonyms and of nomenclatural types, which have already been recombined. Three names proposed in an earlier paper turned out to be later Cortinarius homonyms. Therefore, they are replaced by the new names C. singerianus C. glabriceps and C. deminutus.
Article
Phylogenetic relationships of two sections from subgenus Myxacium in the brown-spored agaric genus Cortinarius were studied using nuclear DNA sequence data from the internal transcribed spacers (ITS1 and ITS2) and the 5.8S rRNA gene of 23 species of Cortinarius, including seven subgenera. Of these, ten species represent the focal study group from sections Defibulati and Myxacium, three species represent three additional sections within subgenus Myxacium, and ten species represent six remaining subgenera within Cortinarius. Phylogenies inferred using parsimony and maximum likelihood strongly supported the two sections as monophyletic, but did not support the subgenus Myxacium as a distinct lineage. Neither method of analysis resolved relationships below the sectional level. Morphological traits were examined for the two monophyletic groups traditionally classified as sections Defibulati and Myxacium. Absence of clamp connections is a synapomorphy of section Defibulati. Cladistic analyses confirm that members sampled from subgenus Myxacium in other sections are more closely allied with other subgenera, suggesting the current taxonomy needs revision to reflect phylogenetic relationships.
Book
— We studied sequence variation in 16S rDNA in 204 individuals from 37 populations of the land snail Candidula unifasciata (Poiret 1801) across the core species range in France, Switzerland, and Germany. Phylogeographic, nested clade, and coalescence analyses were used to elucidate the species evolutionary history. The study revealed the presence of two major evolutionary lineages that evolved in separate refuges in southeast France as result of previous fragmentation during the Pleistocene. Applying a recent extension of the nested clade analysis (Templeton 2001), we inferred that range expansions along river valleys in independent corridors to the north led eventually to a secondary contact zone of the major clades around the Geneva Basin. There is evidence supporting the idea that the formation of the secondary contact zone and the colonization of Germany might be postglacial events. The phylogeographic history inferred for C. unifasciata differs from general biogeographic patterns of postglacial colonization previously identified for other taxa, and it might represent a common model for species with restricted dispersal.
Article
The recently-developed statistical method known as the "bootstrap" can be used to place confidence intervals on phylogenies. It involves resampling points from one's own data, with replacement, to create a series of bootstrap samples of the same size as the original data. Each of these is analyzed, and the variation among the resulting estimates taken to indicate the size of the error involved in making estimates from the original data, In the case of phylogenies, it is argued that the proper method of resampling is to keep all of the original species while sampling characters with replacement, under the assumption that the characters have been independently drawn by the systematist and have evolved independently. Majority-rule consensus trees can be used to construct a phylogeny showing all of the inferred monophyletic groups that occurred in a majority of the bootstrap samples. If a group shows up 95% of the time or more, the evidence for it is taken to be statistically significant. Existing computer programs can be used to analyze different bootstrap samples by using weights on the characters, the weight of a character being how many times it was drawn in bootstrap sampling. When all characters are perfectly compatible, as envisioned by Hennig, bootstrap sampling becomes unnecessary; the bootstrap method would show significant evidence for a group if it is defined by three or more characters.
Article
The taxonomy of Cortinarius section Phlegmacium in Europe is treated, including 18 taxa, with emphasis on an analysis of character variation and differentiation as a tool for species delimitation and the selection of diagnostic characters. Based on the character differentiation between closely related taxa, spore size, veil structure, veil pigment and pileipellis structure appeared to be the most taxonomic significant characters in the group, whereas characters related to carpophore size and shape were of little value. A numerical (multivariate) analysis was performed including four taxa in subsect. Arguti. The PCO and cluster analysis distinguished more or less completely the four taxa, and a character analysis revealed discontinuous variation in at least one character between the taxa. The spore morphology characters were the most distinguishing. The taxonomic implications of the character variation are discussed. The species of sect. Phlegmacium have very specialized habitat requirements, with strict mycorrhizal associations. Of the 17 species here accepted, 16 appear to be associated with a single host genus in Europe, and most are associated solely with a single host species. Many species are calciphilous, xerophilous and thermophilous, and some have a strictly southern distribution in Europe. The majority of taxa are widely distributed, frequently following the natural distribution of their host trees.
Article
The section Brunnei was extensively studied based on material from North Europe. To stabilise the nomenclature we studied the relevant types of taxa included in this section. Phylogenetic relationships and species limits were investigated using rDNA ITS sequences and the results were compared with the morphological data. We recognised 11 species: Cortinarius brunneus, C. clarobrunneus comb. nov., C. coleoptera, C. ectypus, C. gentilis, C. glandicolor (neotypified), C. pseudorubricosus, and four species described as new C. caesiobrunneus, C. albogaudis, C. carabus, and C. cicindela. They are described here and their taxonomy, ecology, distribution, and relationships are discussed. In addition, a key to species of the section Brunnei is provided. A total of 77 new sequences of 11 species are published including nine type sequences. Also the taxonomic assignments of sequences in the public databases belonging to the section Brunnei are revised.
Article
Panax, the ginseng genus, is one of the most medicinally important genera in the Orient and demonstrates a classical eastern Asian and eastern North American disjunct distributional pattern. Sequences of the internal transcribed spacers (ITS) and the 5.8S coding region of the nuclear ribosomal DNA repeat were obtained for the 12 species of Panax to reconstruct phylogenetic relationships. Of the 2 eastern North American species, P. quinquefolius and P. trifolius, P. quinquefolius was suggested to be more closely related to the eastern Asian species in the ITS tree, while P. trifolius was phylogenetically isolated. Monophyly of the three medicinally most important species, P. ginseng, P. notoginseng, and P. quinquefolius, suggested by previous workers, was not supported by the ITS data. A close phylogenetic relationship between Panax and Aralia was supported. Several biogeographical implications were inferred: (1) two divergence events have produced the eastern Asian and eastern North American disjunct distribution in Panax, (2) no intercontinental species pairs are found in Panax; (3) a discrepancy between the sequence divergence pattern and the phylogenetic pattern was observed in Panax, suggesting the need for caution in using sequence divergence data alone in inferring biogeographical patterns; (4) the Himalayas and central and western China are the current centers of diversity of the ginseng genus; and (5) the low ITS sequence divergence and a close relationship among species in that region suggest that rapid evolutionary radiation may have created such a diversity of Panax in the Himalayas and in central and western China.
Article
Three new Cortinarius species, Cortinarius conopileus, C. keralensis, and C. phlegmophorus spp. nov., are described from Kerala State in southern India. This is the first record of ectomycorrhizal Cortinarius spp. in the tropical part of India. In addition to distinct morphological characters, the comparative analysis of rDNA ITS sequences of the collections from India and morphologically similar species support the recognition of these taxa as new species. Phylogenetic analyses demonstrate that the three Indian Cortinarius spp. belong to both larger subclades of the genus Cortinarius, clade/cortinarius and clade/telamonia. As supported by morphological and molecular data, C. phlegmophorus belongs to Cortinarius subgen. Myxacium sect. Defibulati. Based on classical morphological characters, both C. keralensis and C. conopileus are representatives of subgen. Telamonia. However, C. conopileus belongs to clade/obtusi, which is a well-supported subclade of clade/cortinarius. Thus, in contrast to classical taxonomy, the clade/obtusi represents an independent evolutionary origin of telamonioid taxa. This result is also reflected by the distinct morphological characters of taxa of clade/obtusi, namely the lamellar trama with ellipsoid inflated hyphae and the presence of cystidia. In contrast, C. keralensis is a typical member of clade/telamonia. Within/telamonia, only relationships of closely related taxa are resolved due to the low genetic divergence found in ITS sequences. Based on morphological and molecular criteria, C. keralensis is a distinct taxon of sect. Saturnini.
Article
We describe seven new European species of Cortinarius. All species are based on analyses of morphological and DNA sequence data. They all belong to a well-supported clade comprising most species traditionally treated in Cortinarius subgenus Phlegmacium sections Fulvi and Calochroi (i.e. the/Calochroi clade). All taxa are either fulvoid (containing anthraquinoid pigments) or calochroid (without these pigments). Morphological and ecological data are presented for all species and compared with similar species. A dichotomous key is presented for C. calochrous and similar species, including all six newly described calochroid species. The calochroid species C. albertii, C. chailluzii, C. cisticola, C. sancti-felicis, C. selandicus and C. vesterholtii spp. nov., and the fulvoid species C. langeorum sp. nov. are described.
Some interesting species of the genus Cortinarius Fr. in Andalusia (Spain), Part 2
  • A Ortega
Ortega A, Mahiques R, 1995b. Some interesting species of the genus Cortinarius Fr. in Andalusia (Spain), Part 2. Documents Mycologiques XXV (98–100): 295–304.