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Classification of Aspergillus, Penicillium, Talaromyces And available online at www.studiesinmycology.org STUDIES IN MYCOLOGY 95: 5–169 (2020). Classification of Aspergillus, Penicillium, Talaromyces and related genera (Eurotiales): An overview of families, genera, subgenera, sections, series and species J. Houbraken1*, S. Kocsube2, C.M. Visagie3, N. Yilmaz3, X.-C. Wang1,4, M. Meijer1, B. Kraak1, V. Hubka5, K. Bensch1, R.A. Samson1, and J.C. Frisvad6* 1Westerdijk Fungal Biodiversity Institute, Utrecht, The Netherlands; 2Department of Microbiology, Faculty of Science and Informatics, University of Szeged, Szeged, Hungary; 3Department of Biochemistry, Genetics and Microbiology, Forestry and Agricultural Biotechnology Institute (FABI), University of Pretoria, P. Bag X20, Hatfield, Pretoria, 0028, South Africa; 4State Key Laboratory of Mycology, Institute of Microbiology, Chinese Academy of Sciences, No. 3, 1st Beichen West Road, Chaoyang District, Beijing, 100101, China; 5Department of Botany, Charles University in Prague, Prague, Czech Republic; 6Department of Biotechnology and Biomedicine Technical University of Denmark, Søltofts Plads, B. 221, Kongens Lyngby, DK 2800, Denmark *Correspondence: J. Houbraken, [email protected]; J.C. Frisvad, [email protected] Abstract: 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, Ela- phomycetaceae, 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 com- bination), 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 1187 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. Key words: Classification, Infrageneric classification, New taxa, Nomenclature, Phylogeny, Polythetic classes. Taxonomic novelties: New family; Penicillaginaceae Houbraken, Frisvad & Samson; New genera: Acidotalaromyces Houbraken, Frisvad & Samson, Ascospirella Houbraken, Frisvad & Samson, Evansstolkia Houbraken, Frisvad & Samson, Pseudohamigera Houbraken, Frisvad & Samson; New sections:inAspergillus: sect. Bispori S.W. Peterson, Varga, Frisvad, Samson ex Houbraken, 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, in Penicillium: sect. Alfrediorum 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; New series:inAspergillus: ser. Acidohumorum Houbraken & Frisvad, ser. Alliacei Houbraken & Frisvad, ser. Ambigui Houbraken & Frisvad, ser. Arxiorum Houbraken & Frisvad, ser. Aurantiobrunnei Houbraken & Frisvad, ser. Avenacei Houbraken & Frisvad, ser. Bertholletiarum Houbraken & Frisvad, ser. Biplani Houbraken & Frisvad, ser. Brevipedes Houbraken & Frisvad, ser. Brunneouniseriati 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. Circumdati Houbraken & Frisvad, ser. Conjuncti Houbraken & Frisvad, ser. Coremiiformes Houbraken & Frisvad, ser. Cremei Houbraken & Frisvad, ser. Deflecti Houbraken & Frisvad, ser. Egyptiaci Houbraken & Frisvad, ser. Fennelliarum Houbraken & Frisvad, ser. Flavi Houbraken & Frisvad, ser. Flavipedes Houbraken & Frisvad, ser. Fumigati Houbraken & Frisvad, ser. Funiculosi Houbraken & Frisvad, ser. Halophilici Houbraken & Frisvad, ser. Heteromorphi Houbraken & Frisvad, ser. Homomorphi Houbraken & Frisvad, ser. Implicati Houbraken & Frisvad, ser. Japonici Houbraken & Frisvad, ser. Kalimarum Houbraken & Frisvad, ser. Kitamyces Houbraken & Frisvad, ser. Leporum Houbraken & Frisvad, ser. Leucocarpi 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. Nomiarum Houbraken & Frisvad, ser. Noonimiarum Houbraken & Frisvad, ser. Ochraceorosei Houbraken & Frisvad, ser. Olivimuriarum Houbraken & Frisvad, ser. Penicillioides Houbraken & Frisvad, ser. Polypaecilum Houbraken & Frisvad, ser. Pulvini Houbraken & Frisvad, ser. Restricti Houbraken & Frisvad, ser. Rubri Houbraken & Frisvad, ser. Salinarum Houbraken & Frisvad, ser. Sclerotiorum Houbraken & Frisvad, ser. Sparsi Houbraken & Frisvad, ser. Spathulati Houbraken & Frisvad, ser. Spelaei Houbraken & Frisvad, ser. Speluncei Houbraken & Frisvad, ser. Stellati Houbraken & Frisvad, ser. Steyniorum Houbraken & Frisvad, ser. Tamarindosolorum Houbraken & Frisvad, ser. Teporium Houbraken & Frisvad, ser. Terrei Houbraken & Frisvad, ser. Thermomutati Houbraken & Frisvad, ser. Studies in Mycology Peer review under responsibility of Westerdijk Fungal Biodiversity Institute. © 2020 Westerdijk Fungal Biodiversity Institute. Production and hosting by ELSEVIER B.V. This is an open access article under the CC BY license (http://creativecommons.org/licenses/by/4.0/). 5 HOUBRAKEN ET AL. Unguium Houbraken & Frisvad, ser. Unilaterales Houbraken & Frisvad, ser. Usti Houbraken & Frisvad, ser. Versicolores Houbraken & Frisvad, ser. Viridinutantes Houbraken & Frisvad, ser. Vitricolarum Houbraken & Frisvad, ser. Wentiorum Houbraken & Frisvad, ser. Whitfieldiorum Houbraken & Frisvad, ser. Xerophili Houbraken & Frisvad, in Penicillium: ser. Adametziorum Houbraken & Frisvad, ser. Angustiporcata Houbraken & Frisvad, ser. Atramentosa Houbraken & Frisvad, ser. Brevicompacta Houbraken & Frisvad, ser. Buchwaldiorum Houbraken & Frisvad, ser. Cinnamopurpurea Houbraken & Frisvad, ser. Clavigera Houbraken & Frisvad, ser. Copticolarum Houbraken & Frisvad, ser. Corylophila Houbraken & Frisvad, ser. Costaricensia Houbraken & Frisvad, ser. Dalearum Houbraken & Frisvad, ser. Estinogena Houbraken & Frisvad, ser. Euglauca Houbraken & Frisvad, ser. Fortuita Houbraken & Frisvad, ser.
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