A review of recently introduced Aspergillus, Penicillium, Talaromyces and other Eurotiales species.

The order Eurotiales is diverse and includes species that impact our daily lives in many ways. In the past, its taxonomy was difficult due to morphological similarities, which made accurate identification of species difficult. This situation improved and stabilised with recent taxonomic and nomenclatural revisions that modernised Aspergillus, Penicillium and Talaromyces. This was mainly due to the availability of curated accepted species lists and the publication of comprehensive DNA sequence reference datasets. This has also led to a sharp increase in the number of new species described each year with the accepted species lists in turn also needing regular updates. The focus of this study was to review the 160 species described between the last list of accepted species published in 2020 until 31 December 2022. To review these species, single-gene phylogenies were constructed and GCPSR (Genealogical Concordance Phylogenetic Species Recognition) was applied. Multi-gene phylogenetic analyses were performed to further determine the relationships of the newly introduced species. As a result, we accepted 133 species (37 Aspergillus, two Paecilomyces, 59 Penicillium, two Rasamsonia, 32 Talaromyces and one Xerochrysium), synonymised 22, classified four as doubtful and created a new combination for Paraxerochrysium coryli, which is classified in Xerochrysium. This brings the number of accepted species to 453 for Aspergillus, 12 for Paecilomyces, 535 for Penicillium, 14 for Rasamsonia, 203 for Talaromyces and four for Xerochrysium. We accept the newly introduced section Tenues (in Talaromyces), and series Hainanici (in Aspergillus sect. Cavernicolarum) and Vascosobrinhoana (in Penicillium sect. Citrina). In addition, we validate the invalidly described species Aspergillus annui and A. saccharicola, and series Annuorum (in Aspergillus sect. Flavi), introduce a new combination for Dichlaena lentisci (type of the genus) and place it in a new section in Aspergillus subgenus Circumdati, provide an updated description for Rasamsonia oblata, and list excluded and recently synonymised species that were previously accepted. This study represents an important update of the accepted species lists in Eurotiales. Taxonomic novelties: New sections: Aspergillus section Dichlaena Visagie, Kocsubé & Houbraken. New series: Aspergillus series Annuorum J.J. Silva, B.T. Iamanaka, Frisvad. New species: Aspergillus annui J.J. Silva, M.H.P. Fungaro, Frisvad, M.H. Taniwaki & B.T. Iamanaka; Aspergillus saccharicola J.J. Silva, Frisvad, M.H.P. Fungaro, M.H. Taniwaki & B.T. Iamanaka. New combinations: Aspergillus lentisci (Durieu & Mont.) Visagie, Malloch, L. Kriegsteiner, Samson & Houbraken; Xerochrysium coryli (Crous & Decock) Visagie & Houbraken. Citation: Visagie CM, Yilmaz N, Kocsubé S, Frisvad JC, Hubka V, Samson RA, Houbraken J (2024). A review of recently introduced Aspergillus, Penicillium, Talaromyces and other Eurotiales species. Studies in Mycology 107: 1-66. doi: 10.3114/sim.2024.107.01.

Multi-locus phylogeny unmasks hidden species within the specialised spider-parasitic fungus, Gibellula (Hypocreales, Cordycipitaceae) in Thailand.

Over 80 species of hypocrealean fungi are reported as pathogens of spiders and harvestmen. Among these fungi, the genus Gibellula is highly regarded as a specialised spider-killer that has never been reported to infect other arthropods. While more than 20 species of Gibellula are known, few attempts to identify the infected spiders have been made despite the fact that the host specificity can help identify the fungal species. Here, we morphologically describe and illustrate eight new species of Gibellula and three new records from Thailand of known species along with the multi-gene phylogeny that clearly showed the segregation among the proposed species. Examination of the Gibellula-infected spider hosts identified Oxyopidae, Uloboridae and, for the first time, the ant-mimicking genus Myrmarachne. Taxonomic novelties: New species: Gibellula brevistipitata Kuephadungphan, Tasanathai & Luangsa-ard, G. longicaudata Tasanathai, Kuephadungphan & Luangsa-ard, G. longispora Kuephadungphan & Luangsa-ard, G. nigelii Kuephadungphan, Tasanathai & Luangsa-ard, G. parvula Kuephadungphan, Tasanathai & Luangsa-ard, G. pilosa Kuephadungphan, Tasanathai & Luangsa-ard, G. solita Kuephadungphan, Tasanathai & Luangsa-ard, G. trimorpha Tasanathai, Khonsanit, Kuephadungphan & Luangsa-ard. Citation: Kuephadungphan W, Petcharad B, Tasanathai K, Thanakitpipattana D, Kobmoo N, Khonsanit A, Samson RA, Luangsa-ard JJ (2022). Multi-locus phylogeny unmasks hidden species within the specialised spider-parasitic fungus, Gibellula (Hypocreales, Cordycipitaceae) in Thailand. Studies in Mycology 101: 245-286. doi: 10.3114/sim.2022.101.04.

Taxonomy of Aspergillus series Versicolores: species reduction and lessons learned about intraspecific variability.

Aspergillus series Versicolores members occur in a wide range of environments and substrates such as indoor environments, food, clinical materials, soil, caves, marine or hypersaline ecosystems. The taxonomy of the series has undergone numerous re-arrangements including a drastic reduction in the number of species and subsequent recovery to 17 species in the last decade. The identification to species level is however problematic or impossible in some isolates even using DNA sequencing or MALDI-TOF mass spectrometry indicating a problem in the definition of species boundaries. To revise the species limits, we assembled a large dataset of 518 strains. From these, a total of 213 strains were selected for the final analysis according to their calmodulin (CaM) genotype, substrate and geography. This set was used for phylogenetic analysis based on five loci (benA, CaM, RPB2, Mcm7, Tsr1). Apart from the classical phylogenetic methods, we used multispecies coalescence (MSC) model-based methods, including one multilocus method (STACEY) and five single-locus methods (GMYC, bGMYC, PTP, bPTP, ABGD). Almost all species delimitation methods suggested a broad species concept with only four species consistently supported. We also demonstrated that the currently applied concept of species is not sustainable as there are incongruences between single-gene phylogenies resulting in different species identifications when using different gene regions. Morphological and physiological data showed overall lack of good, taxonomically informative characters, which could be used for identification of such a large number of existing species. The characters expressed either low variability across species or significant intraspecific variability exceeding interspecific variability. Based on the above-mentioned results, we reduce series Versicolores to four species, namely A. versicolor, A. creber, A. sydowii and A. subversicolor, and the remaining species are synonymized with either A. versicolor or A. creber. The revised descriptions of the four accepted species are provided. They can all be identified by any of the five genes used in this study. Despite the large reduction in species number, identification based on phenotypic characters remains challenging, because the variation in phenotypic characters is high and overlapping among species, especially between A. versicolor and A. creber. Similar to the 17 narrowly defined species, the four broadly defined species do not have a specific ecology and are distributed worldwide. We expect that the application of comparable methodology with extensive sampling could lead to a similar reduction in the number of cryptic species in other extensively studied Aspergillus species complexes and other fungal genera. Citation: Sklenár F, Glässnerová K, Jurjevic Z, Houbraken J, Samson RA, Visagie CM, Yilmaz N, Gené J, Cano J, Chen AJ, Nováková A, Yaguchi T, Kolarík M, Hubka V (2022). Taxonomy of Aspergillus series Versicolores: species reduction and lessons learned about intraspecific variability. Studies in Mycology 102 : 53-93. doi: 10.3114/sim.2022.102.02.

New and Interesting Fungi. 4.

An order, family and genus are validated, seven new genera, 35 new species, two new combinations, two epitypes, two lectotypes, and 17 interesting new host and / or geographical records are introduced in this study. Validated order, family and genus: Superstratomycetales and Superstratomycetaceae (based on Superstratomyces ). New genera: Haudseptoria (based on Haudseptoria typhae); Hogelandia (based on Hogelandia lambearum); Neoscirrhia (based on Neoscirrhia osmundae); Nothoanungitopsis (based on Nothoanungitopsis urophyllae); Nothomicrosphaeropsis (based on Nothomicrosphaeropsis welwitschiae); Populomyces (based on Populomyces zwinianus); Pseudoacrospermum (based on Pseudoacrospermum goniomae). New species: Apiospora sasae on dead culms of Sasa veitchii (Netherlands); Apiospora stipae on dead culms of Stipa gigantea (Spain); Bagadiella eucalyptorum on leaves of Eucalyptus sp. (Australia); Calonectria singaporensis from submerged leaf litter (Singapore); Castanediella neomalaysiana on leaves of Eucalyptus sp. (Malaysia); Colletotrichum pleopeltidis on leaves of Pleopeltis sp. (South Africa); Coniochaeta deborreae from soil (Netherlands); Diaporthe durionigena on branches of Durio zibethinus (Vietnam); Floricola juncicola on dead culm of Juncus sp. (France); Haudseptoria typhae on leaf sheath of Typha sp. (Germany); Hogelandia lambearum from soil (Netherlands); Lomentospora valparaisensis from soil (Chile); Neofusicoccum mystacidii on dead stems of Mystacidium capense (South Africa); Neomycosphaerella guibourtiae on leaves of Guibourtia sp. (Angola); Niesslia neoexosporioides on dead leaves of Carex paniculata (Germany); Nothoanungitopsis urophyllae on seed capsules of Eucalyptus urophylla (South Africa); Nothomicrosphaeropsis welwitschiae on dead leaves of Welwitschia mirabilis (Namibia); Paracremonium bendijkiorum from soil (Netherlands); Paraphoma ledniceana on dead wood of Buxus sempervirens (Czech Republic); Paraphoma salicis on leaves of Salix cf. alba (Ukraine); Parasarocladium wereldwijsianum from soil (Netherlands); Peziza ligni on masonry and plastering (France); Phyllosticta phoenicis on leaves of Phoenix reclinata (South Africa); Plectosphaerella slobbergiarum from soil (Netherlands); Populomyces zwinianus from soil (Netherlands); Pseudoacrospermum goniomae on leaves of Gonioma kamassi (South Africa); Pseudopyricularia festucae on leaves of Festuca californica (USA); Sarocladium sasijaorum from soil (Netherlands); Sporothrix hypoxyli in sporocarp of Hypoxylon petriniae on Fraxinus wood (Netherlands); Superstratomyces albomucosus on Pycnanthus angolensis (Netherlands); Superstratomyces atroviridis on Pinus sylvestris (Netherlands); Superstratomyces flavomucosus on leaf of Hakea multilinearis (Australia); Superstratomyces tardicrescens from human eye specimen (USA); Taeniolella platani on twig of Platanus hispanica (Germany), and Tympanis pini on twigs of Pinus sylvestris (Spain). Citation: Crous PW, Hernández-Restrepo M, Schumacher RK, Cowan DA, Maggs-Kölling G, Marais E, Wingfield MJ, Yilmaz N, Adan OCG, Akulov A, Álvarez Duarte E, Berraf-Tebbal A, Bulgakov TS, Carnegie AJ, de Beer ZW, Decock C, Dijksterhuis J, Duong TA, Eichmeier A, Hien LT, Houbraken JAMP, Khanh TN, Liem NV, Lombard L, Lutzoni FM, Miadlikowska JM, Nel WJ, Pascoe IG, Roets F, Roux J, Samson RA, Shen M, Spetik M, Thangavel R, Thanh HM, Thao LD, van Nieuwenhuijzen EJ, Zhang JQ, Zhang Y, Zhao LL, Groenewald JZ (2021). New and Interesting Fungi. 4. Fungal Systematics and Evolution 7: 255-343. doi: 10.3114/fuse.2021.07.13.

A re-evaluation of Penicillium section Canescentia, including the description of five new species.

A survey of Penicillium in the fynbos biome from South Africa resulted in the isolation of 61 species of which 29 were found to be new. In this study we focus on Penicillium section Canescentia, providing a phylogenetic re-evaluation based on the analysis of partial beta-tubulin (BenA), calmodulin (CaM) and RNA polymerase II second largest subunit (RPB2) sequence data. Based on phylogenies we show that five fynbos species are new and several previously assigned synonyms of P. canescens and P. janczewskii should be considered as distinct species. As such, we provide descriptions for the five new species and introduce the new name P. elizabethiae for the illegitimate P. echinatum. We also update the accepted species list and synonymies of section Canescentia species and provide a review of extrolites produced by these species. Citation: Visagie CM, Frisvad JC, Houbraken J, et al. 2021. A re-evaluation of Penicillium section Canescentia, including the description of five new species. Persoonia 46: 163-187. https://doi.org/10.3767/persoonia.2021.46.06.

Revisiting Metarhizium and the description of new species from Thailand.

Over the last two decades the molecular phylogeny and classification of Metarhizium has been widely studied. Despite these efforts to understand this enigmatic genus, the basal lineages in Metarhizium are still poorly resolved. In this study, a phylogenetic framework is reconstructed for the Clavicipitaceae focusing on Metarhizium through increased taxon-sampling using five genomic loci (SSU, LSU, tef, rpb1, rpb2) and the barcode marker ITS rDNA. Multi-gene phylogenetic analyses and morphological characterisation of green-spored entomopathogenic Metarhizium isolates from Thailand and soil isolates of M. carneum and M. marquandii reveal their ecological, genetic and species diversity. Nineteen new species are recognised in the Metarhizium clade with narrow host ranges: two new species are found in the M. anisopliae complex - M. clavatum on Coleoptera larvae and M. sulphureum on Lepidoptera larvae; four new species are found in the M. flavoviride complex - M. biotecense and M. fusoideum on brown plant hoppers (Hemiptera), M. culicidarum on mosquitoes, M. nornnoi on Lepidoptera larvae; three new species M. megapomponiae, M. cicadae, M. niveum occur on cicadas; five new species M. candelabrum, M. cercopidarum, M. ellipsoideum, M. huainamdangense M. ovoidosporum occur on planthoppers, leafhoppers and froghoppers (Hemiptera); one new species M. eburneum on Lepidoptera pupae; and four new species M. phuwiangense, M. purpureum, M. purpureonigrum, M. flavum on Coleoptera . Of these 19 new species, seven produce a sexual morph (M. clavatum, M. eburneum, M. flavum, M. phuwiangense, M. purpureonigrum, M. purpureum, and M. sulphureum) and asexual morphs are found in the remaining new species and also in M. sulphureum, M. purpureonigrum and M. purpureum. Metarhizium blattodeae, M. koreanum and M. viridulum are new records for Thailand. An alternative neotype for Metarhizium anisopliae is proposed based on multi-gene and 5'tef analyses showing that CBS 130.71 from Ukraine is more suitable, being from a much closer geographical location to Metchnikoff's Metarhizium anisopliae. This isolate is distinct from the neotype of Metarhizium anisopliae var. anisopliae proposed by M. Tulloch from Ethiopia (ARSEF 7487). Six new genera are established for monophyletic clades subtending the core Metarhizium clade, including Keithomyces, Marquandomyces, Papiliomyces, Purpureomyces, Sungia, and Yosiokobayasia. Metarhizium carneum, M. aciculare, and M. neogunnii are combined in Keithomyces and one new combination for M. marquandii in Marquandomyces is proposed. Purpureomyces is introduced for species producing purple stromata including a new combination for M. khaoyaiense and two new species P. maesotensis and P. pyriformis. Papiliomyces contains two new combinations for M. liangshanense and Metacordyceps shibinensis. The genus Sungia is proposed for the Korean species M. yongmunense on Lepidoptera pupa and Yosiokobayasia for the Japanese species M. kusanagiense also on Lepidoptera pupa. A synoptic and dichotomous key to the accepted taxa is provided together with tables listing distinguishing morphological characters between species, host preferences, and geography.

Classification of Aspergillus, Penicillium, Talaromyces and related genera (Eurotiales): An overview of families, genera, subgenera, sections, series and species.

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.

Phylogenetic re-evaluation of Thielavia with the introduction of a new family Podosporaceae.

The genus Thielavia is morphologically defined by having non-ostiolate ascomata with a thin peridium composed of textura epidermoidea, and smooth, single-celled, pigmented ascospores with one germ pore. Thielavia is typified with Th. basicola that grows in close association with a hyphomycete which was traditionally identified as Thielaviopsis basicola. Besides Th. basicola exhibiting the mycoparasitic nature, the majority of the described Thielavia species are from soil, and some have economic and ecological importance. Unfortunately, no living type material of Th. basicola exists, hindering a proper understanding of the classification of Thielavia. Therefore, Thielavia basicola was neotypified by material of a mycoparasite presenting the same ecology and morphology as described in the original description. We subsequently performed a multi-gene phylogenetic analyses (rpb2, tub2, ITS and LSU) to resolve the phylogenetic relationships of the species currently recognised in Thielavia. Our results demonstrate that Thielavia is highly polyphyletic, being related to three family-level lineages in two orders. The redefined genus Thielavia is restricted to its type species, Th. basicola, which belongs to the Ceratostomataceae (Melanosporales) and its host is demonstrated to be Berkeleyomyces rouxiae, one of the two species in the "Thielaviopsis basicola" species complex. The new family Podosporaceae is sister to the Chaetomiaceae in the Sordariales and accommodates the re-defined genera Podospora, Trangularia and Cladorrhinum, with the last genus including two former Thielavia species (Th. hyalocarpa and Th. intermedia). This family also includes the genetic model species Podospora anserina, which was combined in Triangularia (as Triangularia anserina). The remaining Thielavia species fall in ten unrelated clades in the Chaetomiaceae, leading to the proposal of nine new genera (Carteria, Chrysanthotrichum, Condenascus, Hyalosphaerella, Microthielavia, Parathielavia, Pseudothielavia, Stolonocarpus and Thermothielavioides). The genus Canariomyces is transferred from Microascaceae (Microascales) to Chaetomiaceae based on its type species Can. notabilis. Canariomyces is closely related to the human-pathogenic genus Madurella, and includes three thielavia-like species and one novel species. Three monotypic genera with a chaetomium-like morph (Brachychaeta, Chrysocorona and Floropilus) are introduced to better resolve the Chaetomiaceae and the thielavia-like species in the family. Chrysocorona lucknowensis and Brachychaeta variospora are closely related to Acrophialophora and three newly introduced genera containing thielavia-like species; Floropilus chiversii is closely related to the industrially important and thermophilic species Thermothielavioides terrestris (syn. Th. terrestris). This study shows that the thielavia-like morph is a homoplastic form that originates from several separate evolutionary events. Furthermore, our results provide new insights into the taxonomy of Sordariales and the polyphyletic Lasiosphaeriaceae.

Redefining Humicola sensu stricto and related genera in the Chaetomiaceae.

The traditional concept of the genus Humicola includes species that produce pigmented, thick-walled and single-celled spores laterally or terminally on hyphae or minimally differentiated conidiophores. More than 50 species have been described in the genus. Species commonly occur in soil, indoor environments, and compost habitats. The taxonomy of Humicola and morphologically similar genera is poorly understood in modern terms. Based on a four-locus phylogeny, the morphological concept of Humicola proved to be polyphyletic. The type of Humicola, H. fuscoatra, belongs to the Chaetomiaceae. In the Chaetomiaceae, species producing humicola-like thick-walled spores are distributed among four lineages: Humicola sensu stricto, Mycothermus, Staphylotrichum, and Trichocladium. In our revised concept of Humicola, asexual and sexually reproducing species both occur. The re-defined Humicola contains 24 species (seven new and thirteen new combinations), which are described and illustrated in this study. The species in this genus produce conidia that are lateral, intercalary or terminal on/in hyphae, and conidiophores are not formed or are minimally developed (micronematous). The ascospores of sexual Humicola species are limoniform to quadrangular in face view and bilaterally flattened with one apical germ pore. Seven species are accepted in Staphylotrichum (four new species, one new combination). Thick-walled conidia of Staphylotrichum species usually arise either from hyphae (micronematous) or from apically branched, seta-like conidiophores (macronematous). The sexual morph represented by Staphylotrichum longicolleum (= Chaetomium longicolleum) produces ascomata with long necks composed of a fused basal part of the terminal hairs, and ascospores that are broad limoniform to nearly globose, bilaterally flattened, with an apical germ pore. The Trichocladium lineage has a high morphological diversity in both asexual and sexual structures. Phylogenetic analysis revealed four subclades in this lineage. However, these subclades are genetically closely related, and no distinctive phenotypic characters are linked to any of them. Fourteen species are accepted in Trichocladium, including one new species, twelve new combinations. The type species of Gilmaniella, G. humicola, belongs to the polyphyletic family Lasiosphaeriaceae (Sordariales), but G. macrospora phylogenetically belongs to Trichocladium. The thermophilic genus Mycothermus and the type species My. thermophilum are validated, and one new Mycothermus species is described. Phylogenetic analyses show that Remersonia, another thermophilic genus, is sister to Mycothermus and two species are known, including one new species. Thermomyces verrucosus produces humicola-like conidia and is transferred to Botryotrichum based on phylogenetic affinities. This study is a first attempt to establish an inclusive modern classification of Humicola and humicola-like genera of the Chaetomiaceae. More research is needed to determine the phylogenetic relationships of "humicola"-like species outside the Chaetomiaceae.

Taxonomy of Aspergillus section Flavi and their production of aflatoxins, ochratoxins and other mycotoxins.

Aflatoxins and ochratoxins are among the most important mycotoxins of all and producers of both types of mycotoxins are present in Aspergillus section Flavi, albeit never in the same species. Some of the most efficient producers of aflatoxins and ochratoxins have not been described yet. Using a polyphasic approach combining phenotype, physiology, sequence and extrolite data, we describe here eight new species in section Flavi. Phylogenetically, section Flavi is split in eight clades and the section currently contains 33 species. Two species only produce aflatoxin B1 and B2 (A. pseudotamarii and A. togoensis), and 14 species are able to produce aflatoxin B1, B2, G1 and G2: three newly described species A. aflatoxiformans, A. austwickii and A. cerealis in addition to A. arachidicola, A. minisclerotigenes, A. mottae, A. luteovirescens (formerly A. bombycis), A. nomius, A. novoparasiticus, A. parasiticus, A. pseudocaelatus, A. pseudonomius, A. sergii and A. transmontanensis. It is generally accepted that A. flavus is unable to produce type G aflatoxins, but here we report on Korean strains that also produce aflatoxin G1 and G2. One strain of A. bertholletius can produce the immediate aflatoxin precursor 3-O-methylsterigmatocystin, and one strain of Aspergillus sojae and two strains of Aspergillus alliaceus produced versicolorins. Strains of the domesticated forms of A. flavus and A. parasiticus, A. oryzae and A. sojae, respectively, lost their ability to produce aflatoxins, and from the remaining phylogenetically closely related species (belonging to the A. flavus-, A. tamarii-, A. bertholletius- and A. nomius-clades), only A. caelatus, A. subflavus and A. tamarii are unable to produce aflatoxins. With exception of A. togoensis in the A. coremiiformis-clade, all species in the phylogenetically more distant clades (A. alliaceus-, A. coremiiformis-, A. leporis- and A. avenaceus-clade) are unable to produce aflatoxins. Three out of the four species in the A. alliaceus-clade can produce the mycotoxin ochratoxin A: A. alliaceus s. str. and two new species described here as A. neoalliaceus and A. vandermerwei. Eight species produced the mycotoxin tenuazonic acid: A. bertholletius, A. caelatus, A. luteovirescens, A. nomius, A. pseudocaelatus, A. pseudonomius, A. pseudotamarii and A. tamarii while the related mycotoxin cyclopiazonic acid was produced by 13 species: A. aflatoxiformans, A. austwickii, A. bertholletius, A. cerealis, A. flavus, A. minisclerotigenes, A. mottae, A. oryzae, A. pipericola, A. pseudocaelatus, A. pseudotamarii, A. sergii and A. tamarii. Furthermore, A. hancockii produced speradine A, a compound related to cyclopiazonic acid. Selected A. aflatoxiformans, A. austwickii, A. cerealis, A. flavus, A. minisclerotigenes, A. pipericola and A. sergii strains produced small sclerotia containing the mycotoxin aflatrem. Kojic acid has been found in all species in section Flavi, except A. avenaceus and A. coremiiformis. Only six species in the section did not produce any known mycotoxins: A. aspearensis, A. coremiiformis, A. lanosus, A. leporis, A. sojae and A. subflavus. An overview of other small molecule extrolites produced in Aspergillus section Flavi is given.

Cladosporium species in indoor environments.

As part of a worldwide survey of the indoor mycobiota about 520 new Cladosporium isolates from indoor environments mainly collected in China, Europe, New Zealand, North America and South Africa were investigated by using a polyphasic approach to determine their species identity. All Cladosporium species occurring in indoor environments are fully described and illustrated. Fourty-six Cladosporium species are treated of which 16 species are introduced as new. A key for the most common Cladosporium species isolated from indoor environments is provided. Cladosporium halotolerans proved to be the most frequently isolated Cladosporium species indoors.

Cephalotrichum and related synnematous fungi with notes on species from the built environment.

A recent taxonomic revision of Microascaceae with an emphasis on synnematous fungi enabled re-identification of previously isolated indoor strains of Cephalotrichum. All available Cephalotrichum strains from the culture collection of the Westerdijk Institute were studied, 20 originating from the built environment. Phylogenetic relationships were inferred from DNA sequence data from the internal transcribed spacer 1 and 2 and intervening 5.8S nrDNA (ITS), and parts of ß-tubulin (tub2) and translation elongation factor 1-α (tef1) genes. Additionally, herbarium material of 14 Cephalotrichum species described from soil in China was studied, and the taxonomy of C. album, not considered in recent revisions, was reevaluated. Sixteen phylogenetic species in Cephalotrichum are distinguished, five described as new species: C. domesticum, C. lignatile, C. telluricum, C. tenuissimum and C. transvaalense. Five Cephalotrichum species occur in the built environment: C. domesticum, C. gorgonifer (formerly known as Trichurus spiralis), C. microsporum, C. purpureofuscum, and C. verrucisporum. Based on the number of isolates, C. gorgonifer (nine strains) is the most common indoor species. The study of the Chinese herbarium material resulted in the acceptance of three additional Cephalotrichum species: C. casteneum, C. ellipsoideum, and C. spirale. Four species are considered nomena dubia (C. cylindrosporum, C. macrosporum, C. ovoideum, and C. robustum), five are placed in synonymy with other Cephalotrichum species (C. acutisporum, C. inflatum, C. longicollum, C. oblongum, C. terricola) and one species, C. verrucipes, is probably a synonym of Penicillium clavigerum. Cephalotrichum columnare, former Doratomyces columnaris, is transferred to Kernia. Cephalotrichum album, formerly known as Doratomyces putredinis, is transferred to Acaulium and redescribed.

Phylogeny of xerophilic aspergilli (subgenus Aspergillus) and taxonomic revision of section Restricti.

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 ß-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.

Polyphasic taxonomy of Aspergillus section Aspergillus (formerly Eurotium), and its occurrence in indoor environments and food.

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 ß-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.

Phylogenetic analysis of Monascus and new species from honey, pollen and nests of stingless bees.

The genus Monascus was described by van Tieghem (1884) to accommodate M. ruber and M. mucoroides, two species with non-ostiolate ascomata. Species delimitation in the genus is still mainly based on phenotypic characters, and taxonomic studies that include sequence data are limited. The genus is of economic importance. Species are used in fermented Asian foods as food colourants (e.g. 'red rice' (ang-kak, angka)) and found as spoilage organisms, and recently Monascus was found to be essential in the lifecycle of stingless bees. In this study, a polyphasic approach was applied combining morphological characters, ITS, LSU, ß-tubulin, calmodulin and RNA polymerase II second largest subunit sequences and extrolite data, to delimit species and to study phylogenetic relationships in Monascus. Furthermore, 30 Monascus isolates from honey, pollen and nests of stingless bees in Brazil were included. Based on this polyphasic approach, the genus Monascus is resolved in nine species, including three new species associated with stingless bees (M. flavipigmentosus sp. nov., M. mellicola sp. nov., M. recifensis sp. nov., M. argentinensis, M. floridanus, M. lunisporas, M. pallens, M. purpureus, M. ruber), and split in two new sections (section Floridani sect. nov., section Rubri sect. nov.). Phylogenetic analysis showed that the xerophile Monascus eremophilus does not belong in Monascus and monophyly in Monascus is restored with the transfer of M. eremophilus to Penicillium (P. eremophilum comb. nov.). A list of accepted and excluded Monascus and Basipetospora species is given, together with information on (ex-)types cultures and barcode sequence data.

Scopulariopsis and scopulariopsis-like species from indoor environments.

Scopulariopsis-like species are often reported from the indoor environment, as well as from clinical samples. The lack of type isolates and thorough phylogenetic studies in the Microascaceae hampered the correct identification of these isolates. Based on recent phylogenetic studies, which resulted in multiple name changes, the aim is to molecularly identify the Scopulariopsis and scopulariopsis-like species which occur in the indoor environment and give an overview of the current species in these genera and their habitats. Strains from the CBS culture collection were supplemented with almost 80 indoor strains of which the internal transcribed spacer 1 and 2 and intervening 5.8S nrDNA (ITS), beta-tubulin (tub2) and translation elongation factor 1-alpha (tef1) gene regions were sequenced for phylogenetic inference. The multi-gene phylogenies recognise 33 Microascus species and 12 Scopulariopsis species and showed that the recently established genus Fuscoannellis, typified by Scopulariopsis carbonaria, should be synonymized with the genus Yunnania. Seven new Microascus species, four new Scopulariopsis species, and one new Yunnania species, are described, and a new name in Microascus and two new name combinations (one in Microascus, and one in Yunnania) are proposed. In the indoor environment 14 Microascus species and three Scopulariopsis species were found. Scopulariopsis brevicaulis (22 indoor isolates) and Microascus melanosporus (19 indoor isolates) are the most common indoor species, in number of isolates, followed by M. paisii (8 indoor isolates) and S. candida (7 indoor isolates). A genus phylogeny based on the ITS, tef1 and the large subunit 28S nrDNA (LSU) of the type or representative isolates of all here recognised species is provided depicting all species habitats. No correlation between phylogenetic relationship and habitat preference could be observed. Ten species which are found indoor are also found in relation with human-derived samples. A table showing recent name changes and a key to common species of Scopulariopsis and scopulariopsis-like genera found indoors is included.

Taxonomic re-evaluation of species in Talaromyces section Islandici, using a polyphasic approach.

The taxonomy of Talaromyces rugulosus, T. wortmannii and closely related species, classified in Talaromyces sect. Islandici, is reviewed in this paper. The species of Talaromyces sect. Islandici have restricted growth on MEA and CYA, generally have yellow mycelia and produce rugulosin and/or skyrin. They are important in biotechnology (e.g. T. rugulosus, T. wortmannii) and in medicine (e.g. T. piceus, T. radicus). The taxonomy of sect. Islandici was resolved using a combination of morphological, extrolite and phylogenetic data, using the Genealogical Concordance Phylogenetic Species Recognition (GCPSR) concept, with special focus on the T. rugulosus and T. wortmannii species complexes. In this paper, we synonymise T. variabilis, Penicillium concavorugulosum and T. sublevisporus with T. wortmannii, and introduce four new species as T. acaricola, T. crassus, T. infraolivaceus and T. subaurantiacus. Finally, we provide a synoptic table for the identification of the 19 species classified in the section.

Phylogenetic reassessment of the Chaetomium globosum species complex.

Chaetomium globosum, the type species of the genus, is ubiquitous, occurring on a wide variety of substrates, in air and in marine environments. This species is recognised as a cellulolytic and/or endophytic fungus. It is also known as a source of secondary metabolites with various biological activities, having great potential in the agricultural, medicinal and industrial fields. On the negative side, C. globosum has been reported as an air contaminant causing adverse health effects and as causal agent of human fungal infections. However, the taxonomic status of C. globosum is still poorly understood. The contemporary species concept for this fungus includes a broadly defined morphological diversity as well as a large number of synonymies with limited phylogenetic evidence. The aim of this study is, therefore, to resolve the phylogenetic limits of C. globosum s.str. and related species. Screening of isolates in the collections of the CBS-KNAW Fungal Biodiversity Centre (The Netherlands) and the China General Microbiological Culture Collection Centre (China) resulted in recognising 80 representative isolates of the C. globosum species complex. Thirty-six species are identified based on phylogenetic inference of six loci, supported by typical morphological characters, mainly ascospore shape. Of these, 12 species are newly described here. Additionally, C. cruentum, C. mollipilium, C. rectum, C. subterraneum and two varieties of C. globosum are synonymised under C. globosum s.str., and six species are resurrected, i.e. C. angustispirale, C. coarctatum, C. cochliodes, C. olivaceum, C. spiculipilium and C. subglobosum. Chaetomium ascotrichoides is segregated from C. madrasense and the genus name Chaetomidium is rejected. Five species, including C. globosum s.str., are typified here to stabilise their taxonomic status. A further evaluation of the six loci used in this study as potential barcodes indicated that the 28S large subunit (LSU) nrDNA and the internal transcribed spacer regions and intervening 5.8S nrRNA (ITS) gene regions were unreliable to resolve species, whereas ß-tubulin (tub2) and RNA polymerase II second largest subunit (rpb2) showed the greatest promise as DNA barcodes for differentiating Chaetomium species. This study provides a starting point to establish a more robust classification system for Chaetomium and for the Chaetomiaceae.

A taxonomic review of Penicillium species producing conidiophores with solitary phialides, classified in section Torulomyces.

The genus Torulomyces was characterised by species that typically have conidiophores consisting of solitary phialides that produce long chains of conidia connected by disjunctors. Based on the phylogenetic position of P. lagena (generic ex-neotype), the genus and its seven species were transferred to Penicillium and classified in sect. Torulomyces along with P. cryptum and P. lassenii. The aim of this study was to review the species currently classified in sect. Torulomyces using morphology and phylogenies of the ITS, BenA, CaM and RPB2 regions. Based on our results, we accept 16 species in sect. Torulomyces, including 12 new species described as P. aeris, P. austricola, P. cantabricum, P. catalonicum, P. oregonense, P. marthae-christenseniae, P. riverlandense, P. tubakianum, P. variratense, P. williamettense, P. wisconsinense and P. wollemiicola. In addition, we reclassify P. laeve and P. ovatum in sect. Exilicaulis and correct the typification of P. lagena. We provide descriptions and notes on the identification of the species.

Generic hyper-diversity in Stachybotriaceae.

The family Stachybotriaceae was recently introduced to include the genera Myrothecium, Peethambara and Stachybotrys. Members of this family include important plant and human pathogens, as well as several species used in industrial and commercial applications as biodegraders and biocontrol agents. However, the generic boundaries in Stachybotriaceae are still poorly defined, as type material and sequence data are not readily available for taxonomic studies. To address this issue, we performed multi-locus phylogenetic analyses using partial gene sequences of the 28S large subunit (LSU), the internal transcribed spacer regions and intervening 5.8S nrRNA (ITS), the RNA polymerase II second largest subunit (rpb2), calmodulin (cmdA), translation elongation factor 1-alpha (tef1) and ß-tubulin (tub2) for all available type and authentic strains. Supported by morphological characters these data resolved 33 genera in the Stachybotriaceae. These included the nine already established genera Albosynnema, Alfaria, Didymostilbe, Myrothecium, Parasarcopodium, Peethambara, Septomyrothecium, Stachybotrys and Xepicula. At the same time the generic names Melanopsamma, Memnoniella and Virgatospora were resurrected. Phylogenetic inference further showed that both the genera Myrothecium and Stachybotrys are polyphyletic resulting in the introduction of 13 new genera with myrothecium-like morphology and eight new genera with stachybotrys-like morphology.