Nejvíce citovaný článek - PubMed ID 26779142
The Eurotiales is a relatively large order of Ascomycetes with members frequently having positive and negative impact on human activities. Species within this order gain attention from various research fields such as food, indoor and medical mycology and biotechnology. In this article we give an overview of families and genera present in the Eurotiales and introduce an updated subgeneric, sectional and series classification for Aspergillus and Penicillium. Finally, a comprehensive list of accepted species in the Eurotiales is given. The classification of the Eurotiales at family and genus level is traditionally based on phenotypic characters, and this classification has since been challenged using sequence-based approaches. Here, we re-evaluated the relationships between families and genera of the Eurotiales using a nine-gene sequence dataset. Based on this analysis, the new family Penicillaginaceae is introduced and four known families are accepted: Aspergillaceae, Elaphomycetaceae, Thermoascaceae and Trichocomaceae. The Eurotiales includes 28 genera: 15 genera are accommodated in the Aspergillaceae (Aspergillago, Aspergillus, Evansstolkia, Hamigera, Leiothecium, Monascus, Penicilliopsis, Penicillium, Phialomyces, Pseudohamigera, Pseudopenicillium, Sclerocleista, Warcupiella, Xerochrysium and Xeromyces), eight in the Trichocomaceae (Acidotalaromyces, Ascospirella, Dendrosphaera, Rasamsonia, Sagenomella, Talaromyces, Thermomyces, Trichocoma), two in the Thermoascaceae (Paecilomyces, Thermoascus) and one in the Penicillaginaceae (Penicillago). The classification of the Elaphomycetaceae was not part of this study, but according to literature two genera are present in this family (Elaphomyces and Pseudotulostoma). The use of an infrageneric classification system has a long tradition in Aspergillus and Penicillium. Most recent taxonomic studies focused on the sectional level, resulting in a well-established sectional classification in these genera. In contrast, a series classification in Aspergillus and Penicillium is often outdated or lacking, but is still relevant, e.g., the allocation of a species to a series can be highly predictive in what functional characters the species might have and might be useful when using a phenotype-based identification. The majority of the series in Aspergillus and Penicillium are invalidly described and here we introduce a new series classification. Using a phylogenetic approach, often supported by phenotypic, physiologic and/or extrolite data, Aspergillus is subdivided in six subgenera, 27 sections (five new) and 75 series (73 new, one new combination), and Penicillium in two subgenera, 32 sections (seven new) and 89 series (57 new, six new combinations). Correct identification of species belonging to the Eurotiales is difficult, but crucial, as the species name is the linking pin to information. Lists of accepted species are a helpful aid for researchers to obtain a correct identification using the current taxonomic schemes. In the most recent list from 2014, 339 Aspergillus, 354 Penicillium and 88 Talaromyces species were accepted. These numbers increased significantly, and the current list includes 446 Aspergillus (32 % increase), 483 Penicillium (36 % increase) and 171 Talaromyces (94 % increase) species, showing the large diversity and high interest in these genera. We expanded this list with all genera and species belonging to the Eurotiales (except those belonging to Elaphomycetaceae). The list includes 1 187 species, distributed over 27 genera, and contains MycoBank numbers, collection numbers of type and ex-type cultures, subgenus, section and series classification data, information on the mode of reproduction, and GenBank accession numbers of ITS, beta-tubulin (BenA), calmodulin (CaM) and RNA polymerase II second largest subunit (RPB2) gene sequences.
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- Acidotalaromyces Houbraken, Frisvad & Samson, Acidotalaromyces lignorum (Stolk) Houbraken, Frisvad & Samson, Ascospirella Houbraken, Frisvad & Samson, Ascospirella lutea (Zukal) Houbraken, Frisvad & Samson, Aspergillus chaetosartoryae Hubka, Kocsubé & Houbraken, Classification, Evansstolkia Houbraken, Frisvad & Samson, Evansstolkia leycettana (H.C. Evans & Stolk) Houbraken, Frisvad & Samson, Hamigera brevicompacta (H.Z. Kong) Houbraken, Frisvad & Samson, Infrageneric classification, New combinations, series, New combinations, species, New genera, New names, New sections, New series, New taxa, Nomenclature, Paecilomyces lagunculariae (C. Ram) Houbraken, Frisvad & Samson, Penicillaginaceae Houbraken, Frisvad & Samson, Penicillago kabunica (Baghd.) Houbraken, Frisvad & Samson, Penicillago mirabilis (Beliakova & Milko) Houbraken, Frisvad & Samson, Penicillago moldavica (Milko & Beliakova) Houbraken, Frisvad & Samson, Phialomyces arenicola (Chalab.) Houbraken, Frisvad & Samson, Phialomyces humicoloides (Bills & Heredia) Houbraken, Frisvad & Samson, Phylogeny, Polythetic classes, Pseudohamigera Houbraken, Frisvad & Samson, Pseudohamigera striata (Raper & Fennell) Houbraken, Frisvad & Samson, Talaromyces resinae (Z.T. Qi & H.Z. Kong) Houbraken & X.C. Wang, Talaromyces striatoconidius Houbraken, Frisvad & Samson, Taxonomic novelties: New family, Thermoascus verrucosus (Samson & Tansey) Houbraken, Frisvad & Samson, Thermoascus yaguchii Houbraken, Frisvad & Samson, in Aspergillus: sect. Bispori S.W. Peterson, Varga, Frisvad, Samson ex Houbraken, in Aspergillus: ser. Acidohumorum Houbraken & Frisvad, in Aspergillus: ser. Inflati (Stolk & Samson) Houbraken & Frisvad, in Penicillium: sect. Alfrediorum Houbraken & Frisvad, in Penicillium: ser. Adametziorum Houbraken & Frisvad, in Penicillium: ser. Alutacea (Pitt) Houbraken & Frisvad, sect. Crypta Houbraken & Frisvad, sect. Eremophila Houbraken & Frisvad, sect. Formosana Houbraken & Frisvad, sect. Griseola Houbraken & Frisvad, sect. Inusitata Houbraken & Frisvad, sect. Lasseniorum Houbraken & Frisvad, sect. Polypaecilum Houbraken & Frisvad, sect. Raperorum S.W. Peterson, Varga, Frisvad, Samson ex Houbraken, sect. Silvatici S.W. Peterson, Varga, Frisvad, Samson ex Houbraken, sect. Vargarum Houbraken & Frisvad, ser. Alliacei Houbraken & Frisvad, ser. Ambigui Houbraken & Frisvad, ser. Angustiporcata Houbraken & Frisvad, ser. Arxiorum Houbraken & Frisvad, ser. Atramentosa Houbraken & Frisvad, ser. Aurantiobrunnei Houbraken & Frisvad, ser. Avenacei Houbraken & Frisvad, ser. Bertholletiarum Houbraken & Frisvad, ser. Biplani Houbraken & Frisvad, ser. Brevicompacta Houbraken & Frisvad, ser. Brevipedes Houbraken & Frisvad, ser. Brunneouniseriati Houbraken & Frisvad, ser. Buchwaldiorum Houbraken & Frisvad, ser. Calidousti Houbraken & Frisvad, ser. Canini Houbraken & Frisvad, ser. Carbonarii Houbraken & Frisvad, ser. Cavernicolarum Houbraken & Frisvad, ser. Cervini Houbraken & Frisvad, ser. Chevalierorum Houbraken & Frisvad, ser. Cinnamopurpurea Houbraken & Frisvad, ser. Circumdati Houbraken & Frisvad, ser. Clavigera Houbraken & Frisvad, ser. Conjuncti Houbraken & Frisvad, ser. Copticolarum Houbraken & Frisvad, ser. Coremiiformes Houbraken & Frisvad, ser. Corylophila Houbraken & Frisvad, ser. Costaricensia Houbraken & Frisvad, ser. Cremei Houbraken & Frisvad, ser. Crustacea (Pitt) Houbraken & Frisvad, ser. Dalearum Houbraken & Frisvad, ser. Deflecti Houbraken & Frisvad, ser. Egyptiaci Houbraken & Frisvad, ser. Erubescentia (Pitt) Houbraken & Frisvad, ser. Estinogena Houbraken & Frisvad, ser. Euglauca Houbraken & Frisvad, ser. Fennelliarum Houbraken & Frisvad, ser. Flavi Houbraken & Frisvad, ser. Flavipedes Houbraken & Frisvad, ser. Fortuita Houbraken & Frisvad, ser. Fumigati Houbraken & Frisvad, ser. Funiculosi Houbraken & Frisvad, ser. Gallaica Houbraken & Frisvad, ser. Georgiensia Houbraken & Frisvad, ser. Goetziorum Houbraken & Frisvad, ser. Gracilenta Houbraken & Frisvad, ser. Halophilici Houbraken & Frisvad, ser. Herqueorum Houbraken & Frisvad, ser. Heteromorphi Houbraken & Frisvad, ser. Hoeksiorum Houbraken & Frisvad, ser. Homomorphi Houbraken & Frisvad, ser. Idahoensia Houbraken & Frisvad, ser. Implicati Houbraken & Frisvad, ser. Improvisa Houbraken & Frisvad, ser. Indica Houbraken & Frisvad, ser. Japonici Houbraken & Frisvad, ser. Jiangxiensia Houbraken & Frisvad, ser. Kalimarum Houbraken & Frisvad, ser. Kiamaensia Houbraken & Frisvad, ser. Kitamyces Houbraken & Frisvad, ser. Lapidosa (Pitt) Houbraken & Frisvad, ser. Leporum Houbraken & Frisvad, ser. Leucocarpi Houbraken & Frisvad, ser. Livida Houbraken & Frisvad, ser. Longicatenata Houbraken & Frisvad, ser. Macrosclerotiorum Houbraken & Frisvad, ser. Monodiorum Houbraken & Frisvad, ser. Multicolores Houbraken & Frisvad, ser. Neoglabri Houbraken & Frisvad, ser. Neonivei Houbraken & Frisvad, ser. Nidulantes Houbraken & Frisvad, ser. Nigri Houbraken & Frisvad, ser. Nivei Houbraken & Frisvad, ser. Nodula Houbraken & Frisvad, ser. Nomiarum Houbraken & Frisvad, ser. Noonimiarum Houbraken & Frisvad, ser. Ochraceorosei Houbraken & Frisvad, ser. Olivimuriarum Houbraken & Frisvad, ser. Osmophila Houbraken & Frisvad, ser. Paradoxa Houbraken & Frisvad, ser. Paxillorum Houbraken & Frisvad, ser. Penicillioides Houbraken & Frisvad, ser. Phoenicea Houbraken & Frisvad, ser. Pinetorum (Pitt) Houbraken & Frisvad, ser. Polypaecilum Houbraken & Frisvad, ser. Pulvini Houbraken & Frisvad, ser. Quercetorum Houbraken & Frisvad, ser. Raistrickiorum Houbraken & Frisvad, ser. Ramigena Houbraken & Frisvad, ser. Restricti Houbraken & Frisvad, ser. Robsamsonia Houbraken & Frisvad, ser. Rolfsiorum Houbraken & Frisvad, ser. Roseopurpurea Houbraken & Frisvad, ser. Rubri Houbraken & Frisvad, ser. Salinarum Houbraken & Frisvad, ser. Samsoniorum Houbraken & Frisvad, ser. Saturniformia Houbraken & Frisvad, ser. Scabrosa Houbraken & Frisvad, ser. Sclerotigena Houbraken & Frisvad, ser. Sclerotiorum Houbraken & Frisvad, ser. Sheariorum Houbraken & Frisvad, ser. Simplicissima Houbraken & Frisvad, ser. Soppiorum Houbraken & Frisvad, ser. Sparsi Houbraken & Frisvad, ser. Spathulati Houbraken & Frisvad, ser. Spelaei Houbraken & Frisvad, ser. Speluncei Houbraken & Frisvad, ser. Spinulosa Houbraken & Frisvad, ser. Stellati Houbraken & Frisvad, ser. Steyniorum Houbraken & Frisvad, ser. Sublectatica Houbraken & Frisvad, ser. Sumatraensia Houbraken & Frisvad, ser. Tamarindosolorum Houbraken & Frisvad, ser. Teporium Houbraken & Frisvad, ser. Terrei Houbraken & Frisvad, ser. Thermomutati Houbraken & Frisvad, ser. Thiersiorum Houbraken & Frisvad, ser. Thomiorum Houbraken & Frisvad, ser. Unguium Houbraken & Frisvad, ser. Unilaterales Houbraken & Frisvad, ser. Usti Houbraken & Frisvad, ser. Verhageniorum Houbraken & Frisvad, ser. Versicolores Houbraken & Frisvad, ser. Virgata Houbraken & Frisvad, ser. Viridinutantes Houbraken & Frisvad, ser. Vitricolarum Houbraken & Frisvad, ser. Wentiorum Houbraken & Frisvad, ser. Westlingiorum Houbraken & Frisvad, ser. Whitfieldiorum Houbraken & Frisvad, ser. Xerophili Houbraken & Frisvad, series Tularensia (Pitt) Houbraken & Frisvad,
- Publikační typ
- časopisecké články MeSH
Although Aspergillus fumigatus is the major agent of invasive aspergillosis, an increasing number of infections are caused by its cryptic species, especially A. lentulus and the A. viridinutans species complex (AVSC). Their identification is clinically relevant because of antifungal drug resistance and refractory infections. Species boundaries in the AVSC are unresolved since most species have uniform morphology and produce interspecific hybrids in vitro. Clinical and environmental strains from six continents (n = 110) were characterized by DNA sequencing of four to six loci. Biological compatibilities were tested within and between major phylogenetic clades, and ascospore morphology was characterised. Species delimitation methods based on the multispecies coalescent model (MSC) supported recognition of ten species including one new species. Four species are confirmed opportunistic pathogens; A. udagawae followed by A. felis and A. pseudoviridinutans are known from opportunistic human infections, while A. felis followed by A. udagawae and A. wyomingensis are agents of feline sino-orbital aspergillosis. Recently described human-pathogenic species A. parafelis and A. pseudofelis are synonymized with A. felis and an epitype is designated for A. udagawae. Intraspecific mating assay showed that only a few of the heterothallic species can readily generate sexual morphs in vitro. Interspecific mating assays revealed that five different species combinations were biologically compatible. Hybrid ascospores had atypical surface ornamentation and significantly different dimensions compared to parental species. This suggests that species limits in the AVSC are maintained by both pre- and post-zygotic barriers and these species display a great potential for rapid adaptation and modulation of virulence. This study highlights that a sufficient number of strains representing genetic diversity within a species is essential for meaningful species boundaries delimitation in cryptic species complexes. MSC-based delimitation methods are robust and suitable tools for evaluation of boundaries between these species.
Aspergillus section Restricti together with sister section Aspergillus (formerly Eurotium) comprises xerophilic species, that are able to grow on substrates with low water activity and in extreme environments. We adressed the monophyly of both sections within subgenus Aspergillus and applied a multidisciplinary approach for definition of species boundaries in sect. Restricti. The monophyly of sections Aspergillus and Restricti was tested on a set of 102 isolates comprising all currently accepted species and was strongly supported by Maximum likelihood (ML) and Bayesian inferrence (BI) analysis based on β-tubulin (benA), calmodulin (CaM) and RNA polymerase II second largest subunit (RPB2) loci. More than 300 strains belonging to sect. Restricti from various isolation sources and four continents were characterized by DNA sequencing, and 193 isolates were selected for phylogenetic analyses and phenotypic studies. Species delimitation methods based on multispecies coalescent model were employed on DNA sequences from four loci, i.e., ID region of rDNA (ITS + 28S), CaM, benA and RPB2, and supported recognition of 21 species, including 14 new. All these species were also strongly supported in ML and BI analyses. All recognised species can be reliably identified by all four examined genetic loci. Phenotype analysis was performed to support the delimitation of new species and includes colony characteristics on seven cultivation media incubated at several temperatures, growth on an osmotic gradient (six media with NaCl concentration from 0 to 25 %) and analysis of morphology including scanning electron microscopy. The micromorphology of conidial heads, vesicle dimensions, temperature profiles and growth parameters in osmotic gradient were useful criteria for species identification. The vast majority of species in sect. Restricti produce asperglaucide, asperphenamate or both in contrast to species in sect. Aspergillus. Mycophenolic acid was detected for the first time in at least six members of the section. The ascomata of A. halophilicus do not contain auroglaucin, epiheveadride or flavoglaucin which are common in sect. Aspergillus, but shares the echinulins with sect. Aspergillus.
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- Aspergillus canadensis Visagie, Yilmaz, F. Sklenar & Seifert, Aspergillus clavatophorus F. Sklenar, S.W. Peterson & Hubka, Aspergillus destruens Zalar, F. Sklenar, S.W. Peterson & Hubka, Aspergillus domesticus F. Sklenar, Houbraken, Zalar & Hubka, Aspergillus glabripes F. Sklenar, Ž. Jurjević & Hubka, Aspergillus hordei F. Sklenar, S.W. Peterson & Hubka, Aspergillus infrequens F. Sklenar, S.W. Peterson & Hubka, Aspergillus magnivesiculatus F. Sklenar, Zalar, Ž. Jurjević & Hubka, Aspergillus pachycaulis F. Sklenar, S.W. Peterson, Ž. Jurjević & Hubka, Aspergillus penicillioides, Aspergillus pseudogracilis F. Sklenar, Ž. Jurjević & Hubka, Aspergillus restrictus, Aspergillus reticulatus F. Sklenar, Ž. Jurjević, S.W. Peterson & Hubka, Aspergillus salinicola Zalar, F. Sklenar, Visagie & Hubka, Aspergillus tardicrescens F. Sklenar, Houbraken, Zalar, & Hubka, Aspergillus villosus F. Sklenar, S.W. Peterson & Hubka, Eurotium, food spoilage, indoor fungi, linear discriminant analysis, multigene phylogeny, multispecies coalescent model, sick building syndrome, xerophilic fungi,
- Publikační typ
- časopisecké články MeSH
Aspergillus section Aspergillus (formerly the genus Eurotium) includes xerophilic species with uniseriate conidiophores, globose to subglobose vesicles, green conidia and yellow, thin walled eurotium-like ascomata with hyaline, lenticular ascospores. In the present study, a polyphasic approach using morphological characters, extrolites, physiological characters and phylogeny was applied to investigate the taxonomy of this section. Over 500 strains from various culture collections and new isolates obtained from indoor environments and a wide range of substrates all over the world were identified using calmodulin gene sequencing. Of these, 163 isolates were subjected to molecular phylogenetic analyses using sequences of ITS rDNA, partial β-tubulin (BenA), calmodulin (CaM) and RNA polymerase II second largest subunit (RPB2) genes. Colony characteristics were documented on eight cultivation media, growth parameters at three incubation temperatures were recorded and micromorphology was examined using light microscopy as well as scanning electron microscopy to illustrate and characterize each species. Many specific extrolites were extracted and identified from cultures, including echinulins, epiheveadrides, auroglaucins and anthraquinone bisanthrons, and to be consistent in strains of nearly all species. Other extrolites are species-specific, and thus valuable for identification. Several extrolites show antioxidant effects, which may be nutritionally beneficial in food and beverages. Important mycotoxins in the strict sense, such as sterigmatocystin, aflatoxins, ochratoxins, citrinin were not detected despite previous reports on their production in this section. Adopting a polyphasic approach, 31 species are recognized, including nine new species. ITS is highly conserved in this section and does not distinguish species. All species can be differentiated using CaM or RPB2 sequences. For BenA, Aspergillus brunneus and A. niveoglaucus share identical sequences. Ascospores and conidia morphology, growth rates at different temperatures are most useful characters for phenotypic species identification.
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- A. aurantiacoflavus Hubka, A.J. Chen, Jurjević & Samson, A. caperatus A.J. Chen, Frisvad & Samson, A. endophyticus Hubka, A.J. Chen, & Samson, A. levisporus Hubka, A.J. Chen, Jurjević & Samson, A. porosus A.J. Chen, Frisvad & Samson, A. tamarindosoli A.J. Chen, Frisvad & Samson, A. teporis A.J. Chen, Frisvad & Samson, A. zutongqii A.J. Chen, Frisvad & Samson, Ascomycota, Aspergillaceae, Aspergillus aerius A.J. Chen, Frisvad & Samson, Aspergillus proliferans, Eurotiales, Eurotium amstelodami, Extrolites, Multi-gene phylogeny,
- Publikační typ
- časopisecké články MeSH
Aspergillus is one of the economically most important fungal genera. Recently, the ICN adopted the single name nomenclature which has forced mycologists to choose one name for fungi (e.g. Aspergillus, Fusarium, Penicillium, etc.). Previously two proposals for the single name nomenclature in Aspergillus were presented: one attributes the name "Aspergillus" to clades comprising seven different teleomorphic names, by supporting the monophyly of this genus; the other proposes that Aspergillus is a non-monophyletic genus, by preserving the Aspergillus name only to species belonging to subgenus Circumdati and maintaining the sexual names in the other clades. The aim of our study was to test the monophyly of Aspergilli by two independent phylogenetic analyses using a multilocus phylogenetic approach. One test was run on the publicly available coding regions of six genes (RPB1, RPB2, Tsr1, Cct8, BenA, CaM), using 96 species of Penicillium, Aspergillus and related taxa. Bayesian (MrBayes) and Ultrafast Maximum Likelihood (IQ-Tree) and Rapid Maximum Likelihood (RaxML) analyses gave the same conclusion highly supporting the monophyly of Aspergillus. The other analyses were also performed by using publicly available data of the coding sequences of nine loci (18S rRNA, 5,8S rRNA, 28S rRNA (D1-D2), RPB1, RPB2, CaM, BenA, Tsr1, Cct8) of 204 different species. Both Bayesian (MrBayes) and Maximum Likelihood (RAxML) trees obtained by this second round of independent analyses strongly supported the monophyly of the genus Aspergillus. The stability test also confirmed the robustness of the results obtained. In conclusion, statistical analyses have rejected the hypothesis that the Aspergilli are non-monophyletic, and provided robust arguments that the genus is monophyletic and clearly separated from the monophyletic genus Penicillium. There is no phylogenetic evidence to split Aspergillus into several genera and the name Aspergillus can be used for all the species belonging to Aspergillus i.e. the clade comprising the subgenera Aspergillus, Circumdati, Fumigati, Nidulantes, section Cremei and certain species which were formerly part of the genera Phialosimplex and Polypaecilum. Section Cremei and the clade containing Polypaecilum and Phialosimplex are proposed as new subgenera of Aspergillus. The phylogenetic analysis also clearly shows that Aspergillus clavatoflavus and A. zonatus do not belong to the genus Aspergillus. Aspergillus clavatoflavus is therefore transferred to a new genus Aspergillago as Aspergillago clavatoflavus and A. zonatus was transferred to Penicilliopsis as P. zonata. The subgenera of Aspergillus share similar extrolite profiles indicating that the genus is one large genus from a chemotaxonomical point of view. Morphological and ecophysiological characteristics of the species also strongly indicate that Aspergillus is a polythetic class in phenotypic characters.
- Klíčová slova
- Aspergillago Samson, Houbraken & Frisvad, gen. nov., Aspergillago clavatoflava (Raper & Fennell) Samson, Houbraken & Frisvad, comb. nov., Aspergillus, Aspergillus subgenus Cremei, subgen. nov., Aspergillus subgenus Polypaecilum, subgen. nov., Monophyly, Multigene phylogeny, Nomenclature, Penicilliopsis zonatus (Kwon-Chung & Fennell) Samson, Houbraken & Frisvad, comb. nov., Teleomorphs,
- Publikační typ
- časopisecké články MeSH