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

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, 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 In contrast, a series classification in Aspergillus and Penicillium is often outdated or lacking, but is still relevant, e.g., the allocation of a species to a series can be highly predictive in what functional characters the species might have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.

Introduction

The order Eurotiales harbours various economically important genera, such as Aspergillus, Penicillium, Rasamsonia and Talaromyces. Species classified in this order have diverse properties and include (one of) the most important food spoilage organisms (e.g., Aspergillus proliferans (eurotium morph), Paecilomyces variotii), mycotoxin producers (e.g., Aspergillus flavus; aflatoxins), human pathogens (Aspergillus fumigatus, A. flavus, Talaromyces marneffei) and indoor contaminants (e.g., Aspergillus versicolor, A. penicillioides, Penicillium chrysogenum) (Frisvad et al., 2019, Samson et al., 2019, van den Brule et al., 2019, and references therein). Besides their negative impact on human activities, these species are also used in food fermentations (e.g., A. oryzae: soy sauce, miso; P. roqueforti: blue-veined cheese (Roquefort); P. camemberti: Camembert cheese), in biotechnology to produce organic acids and enzymes (e.g., Aspergillus niger: citric acid; Rasamsonia emersonii; Thermomyces lanuginosus: enzyme production) and for the production of pharmaceuticals (e.g., Aspergillus terreus: lovastatin; Penicillium brevicompactum: mycophenolic acid; P. rubens: penicillin) (Houbraken et al., 2012a, Houbraken et al., 2014a, Frisvad et al., 2019).

The use of an infrageneric classification system has a long tradition in Aspergillus and Penicillium (Bainier, 1907, Biourge, 1923, Zaleski, 1927). Thom and his co-workers recognised distinct clusters of species in these genera and named those “groups” or “series” (Thom and Church, 1926, Thom and Raper, 1945, Raper and Thom, 1949, Raper and Fennell, 1965). However, their concept of “groups” does not have any nomenclatural status, and the “series” were wrongly introduced and therefore invalid (Art. 21.1 and 36.1). To avoid confusion and to promote taxonomic stability, a formal infrageneric classification system was needed. Pitt (1980) replaced the “group” structure in Penicillium by a subgeneric and sectional structure, and Gams et al. (1985) carried out the same changes to Aspergillus. The phenotype-based infrageneric classification system proposed in Aspergillus was primarily based on conidium colour, conidiophore morphology and growth rates on agar media. This classification system still has a large overlap with the current system that is based on molecular data (Raper and Fennell, 1965, Houbraken and Samson, 2011, Jurjević et al., 2015, Chen et al., 2016a). Pitt (1980) formally introduced an infrageneric classification system for Penicillium and its sexual morphs Talaromyces and Eupenicillium (10 sections, 21 series), and various other systems have been proposed afterwards. For example, Stolk & Samson (1983) introduced a sectional classification system in Eupenicillium with four sections, Stolk & Samson (1985) subdivided Penicillium into 10 sections and 18 series, and Frisvad & Samson (2004b) recognised five sections and 17 series in subgen. Penicillium. These phenotype-based sectional classifications are nowadays replaced by a system based on DNA sequence data (Houbraken and Samson, 2011, Houbraken et al., 2016). In contrast to Aspergillus, this DNA-based sectional classification system is often not congruent with the old, phenotype-based systems. Although subgeneric, sectional and/or series classifications have a long tradition in Aspergillus and Penicillium, they are not widely used in mycology; however, they are (being) implemented for some economically significant and speciose genera such as Trichoderma, Alternaria, Hebeloma and Talaromyces. The use of infrageneric ranks for phylogenetic clades is useful for managing large speciose genera, like Aspergillus, Penicillium and Talaromyces. Despite molecular reassessments over the last decade, the classification of Aspergillus, Penicillium and Talaromyces species into subgenera and sections has been rather stable. Aspergillus currently includes 25 sections, Penicillium 26 sections and Talaromyces seven sections. In contrast to the classical monographs on Aspergillus and Penicillium (Thom and Church, 1926, Thom and Raper, 1945, Raper and Fennell, 1965, Pitt, 1980), a series level classification based on DNA sequence data is lacking in these genera.

Subgenera, sections, subsections, series and subseries are useful categories between genus and species level and are official nomenclatural taxonomic ranks. Well-supported clades discovered by DNA sequence analyses often indicate natural groups of species that can be translated into a subgenus, section or series. Therefore, these ranks can be highly predictive in what functional characters the species might have. For example, Penicillium subgen. Penicillium sect. Roquefortorum ser. Roquefortorum is a clear clade based on DNA sequence data (Samson et al., 2004, Houbraken et al., 2010a, Houbraken and Samson, 2011). A character analysis of the species from this series shows that they have many characters in common: all grow well on 0.5 % acetic acid and on creatine sucrose agar, all have large globose conidia, rough-walled conidiophore stipes, all grow at elevated carbon dioxide levels, and all produce roquefortine C, noting that they are also polythetic in the production of other extrolites. Penicillium carneum, P. paneum, P. psychrosexuale and P. roqueforti (all members of sect. and ser. Roquefortorum) produce different combinations of mycophenolic acids, isofumigaclavines, patulin, botrydiploidin, marcfortines, penipalines, penipacids, penipanoids and eremofortines (Boysen et al., 1996, Frisvad and Samson, 2004b, Frisvad et al., 2004, Nielsen et al., 2006, O'Brien et al., 2006, Houbraken et al., 2010a, Li et al., 2011, Li et al., 2013, Li et al., 2014). Thus, the classification of a species, like the newly described species P. mediterraneum (Guevara-Suarez et al. 2020) to sect. and ser. Roquefortorum, is highly predictive in what functional characters they might have.

According to Gould (2000), Linnaeus’ binomial nomenclatural system for the species has survived to this day because the genus reflects the phylogeny and the species epithet reflects the functional phenotype and phylotype: “Linnaeus’s classification scheme can be visualised as a series of nested boxes in which the species is the irreducible category” (our italics). We concur, and a consequence of this is that subspecific levels such as subspecies, varieties, forma specialis etc. should not be used in taxonomy, at least not in a formal way (see Wilson & Brown 1953). However, some of the more interesting categories are those between the genus and the species: subgenera, sections, subsections, series and subseries (Kirk et al. 2008). Do these levels have a nomenclatural status and are they predictive for characters? Should they be formally used in taxonomy and cladonomy?

In this study, the families and genera of the Eurotiales were re-evaluated using a nine-gene sequence dataset. New names for lineages representing a new family and four new genera are proposed in the Taxonomy section of this article. The same dataset was used to study the currently defined subgeneric and sectional classification system in Aspergillus and Penicillium. The relationship within Aspergillus and Penicillium was studied using a 4-gene sequence dataset and a novel, sequence-based series classification is proposed. Finally, a list of accepted species in the Eurotiales (except Elaphomycetaceae) is given. This overview updates the Aspergillus (Samson et al. 2014), Penicillium (Visagie et al. 2014b) and Talaromyces (Yilmaz et al. 2014) lists and is expanded with data of other genera and species belonging to the Eurotiales.

Materials and methods

Strain selection for datasets

The phylogenies presented in this study are based on sequences obtained from the NCBI nucleotide database (GenBank), genome-sequenced strains (GenBank, DOE Joint Genome Institute (JGI)) and sequences newly generated in this study. A selection of strains was made to study the phylogenetic relationships within the Eurotiales. The selection aimed to include the current known diversity in the order. In most cases, the types of the species and genera were included. An overview of strains and species is given in Table S1 (Supplementary Information - online only). The phylogenetic relationship of the accepted Aspergillus and Penicillium species was determined with the aim to introduce a new series classification in those genera. We aimed to include all Aspergillus and Penicillium species from the list of accepted species (see below) that had tubulin (BenA), calmodulin (CaM) and/or RNA polymerase II second largest subunit (RPB2) sequences. Species belonging to the same subgenus were analysed together in one dataset, resulting in eight datasets (Aspergillus, Circumdati, Cremei, Fumigati, Nidulantes, Polypaecilum (in Aspergillus)); Aspergilloides and Penicillium (in Penicillium). Steenwyk et al. (2019), using a phylogenomic approach, showed that sect. Nigri does not belong to subgen. Circumdati and the species belonging to this section were therefore analysed in a separate dataset. Finally, in order to determine the taxonomic position of Aspergillus texensis and Penicillium cellarum, two separate datasets were constructed and analysed. Publicly available sequences on GenBank were supplemented with newly generated sequences of A. minisclerotigenes and P. aurantiogriseum strains (for the A. texensis and P. cellarum datasets, respectively) present in the CBS and DTO culture collection housed at the Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands.

DNA isolation, sequencing

Genomic DNA was extracted from cultures grown on malt extract agar (MEA) using the DNeasy® UltraClean® Microbial Kit (Qiagen, Germany) following the manufacturer's instructions. The following primers were used for PCR amplification: T10 (O'Donnell & Cigelnik 1997) or Bt2a (Glass & Donaldson 1995) & Bt2b (Glass & Donaldson 1995) for the partial beta-tubulin (BenA) gene region; Cmd5 (Hong et al. 2006) or CF1 (Peterson 2008) & Cmd6 (Hong et al. 2006) for the calmodulin (CaM) gene region; V9G (de Hoog & Gerrits van den Ende 1998) & LS266 (Masclaux et al. 1995) for the internal transcribed spacer regions (ITS), including 5.8S nrRNA gene region; LR0R & LR5 (Vilgalys & Sun 1994) for a part of the 28S nrDNA (large subunit rDNA, LSU), and NS1 & NS4 (White et al. 1990) for a part of the 18S nrDNA (small subunit rDNA, SSU). Parts of the Tsr1 (gene coding for a putative ribosome biogenesis protein), Cct8 (gene coding for the theta subunit of the TCP-1 chaperonin complex) and RPB1 (RNA polymerase II largest subunit) genes were amplified and sequenced using the methods described previously by Houbraken & Samson (2011). A part of the RPB2 gene was amplified and sequenced using the primers RPB2-F1 (GCITTYTTCYTIGGITAYATGG) & RPB2-7CR_1 (CATRGCYTGYTTRCCCATIGC). The PCR mixes containing dimethylsulfoxide (DMSO) were ran at an annealing temperature of 48 °C; the mixes containing bovine serum albumine (BSA) at 55 °C. Each of the amplicons was sequenced in both directions using the PCR primers and the BigDye Terminator v. 3.1 Cycle Sequencing Kit (Applied Biosystems, Foster City, CA, USA) following the manufacturer's instructions. Sequencing was performed with an Applied Biosystems™ 3730xl DNA Analyzer (ThermoFisher Scientific, CA, USA). Consensus sequences for each locus were assembled using SeqMan Pro v.15 (DNASTAR). Novel sequences generated in this study were deposited in the GenBank database under accession numbers MN431358–MN431418, MN969061–MN969442, MT024497–MT024529 and MT066177–MT066186.

Study on phylogenetic relationships above section level

The families and genera of the Eurotiales were re-evaluated using a nine-gene sequence dataset and the same dataset was used to study the currently defined subgeneric and sectional classification system in Aspergillus and Penicillium. The analysis included DNA sequences of nine loci (BenA, CaM, Cct8, ITS, LSU, RPB1, RPB2, SSU and Tsr1) from 263 species belonging to the order Eurotiales and 16 species from the order Onygenales as outgroup. The dataset was compiled using publicly and newly generated sequences listed in Table S1. Sequences of the RPB1, RPB2, Cct8, SSU and LSU loci were aligned using PRANK v. 140603 (Löytynoja 2014) with the -F option. As CaM, BenA, Tsr1 and the ITS loci are difficult to align, a guide tree based on a per-gene partitioned dataset of RPB1, RPB2, Cct8, SSU and LSU sequences was applied with the -F and -prunetree option. The guide tree was inferred by maximum likelihood (ML) using RAxML-NG v. 0.9.0 (Kozlov et al. 2019) under the GTR model with gamma-distributed rate heterogeneity. For the final inference, the best fitting model for each locus was determined by ModelTest-NG v. 0.1.4 (Darriba et al. 2019) based on the corrected Akaike Information Criterion (Sugiura, 1978, Hurvich and Tsai, 1989) with a maximum likelihood starting tree, set to choose between all models implemented in RAxML-NG with discrete gamma rate categories or FreeRate (Yang 1995) model. For model selection, CaM and BenA datasets were partitioned to exons and introns, while the ITS dataset was partitioned to ribosomal rDNA and ITS1-ITS2 regions. Alignments of CaM, BenA, Tsr1 and ITS datasets contained a high number of indels with important phylogenetic signal (Nagy et al. 2012), therefore gaps were recoded as absence/presence characters by 2matrix (Salinas & Little 2014) implementing the simple indel coding algorithm (Simmons & Ochoterena 2000). The four indel matrices were treated as a single partition and added to the concatenated dataset. As indel-based datasets do not contain constant sites, the ascertainment bias correction described by Lewis (2001) was used during the analysis. Branch supports of the best ML tree were estimated by 500 bootstrap replicates.

Phylogenetic analysis of series relationships within Aspergillus and Penicillium

Separate phylograms were made of each Aspergillus and Penicillium subgenus based on a combined dataset of BenA, CaM and RPB2 gene sequences. The combined datasets were made from sequences derived from representative Aspergillus and Penicillium species. An overview of species and their corresponding GenBank accession numbers can be found in the “list of accepted species” below. The separate sequence data sets were aligned using MAFFT v. 7.427 (Katoh & Standley 2013), and subsequently combined with BioEdit v. 7.0.5.3 (Hall 1999) into a three-locus dataset. Phylogenetic analyses were inferred from Maximum Likelihood (ML) and Bayesian inference (BI). Maximum Likelihood analysis was performed using RAxML-HPC2 on XSEDE v. 8.2.12 via the CIPRES Science Gateway (www.phylo.org) with the default GTRCAT model. Bayesian inference analysis was performed with MrBayes v. 3.2.6 (Ronquist et al. 2012) using a Markov Chain Monte Carlo (MCMC) algorithm. Hamigera avellanea CBS 295.48^T and Penicillium expansum CBS 325.48^T served as outgroup in the Aspergillus phylogenies; H. avellanea CBS 295.48^T and Aspergillus glaucus CBS 516.65^T were the outgroup species in the Penicillium phylogenies.

Extrolites

Secondary metabolite data from literature were used if they were based on reliable identifications. In some cases, isolates of newly described species were analyzed using high performance liquid chromatography with diode array detection (HPLC-DAD) and/or using UHPLC-DAD-MS-MS (Frisvad and Thrane, 1987, Nielsen et al., 2011, Kildgaard et al., 2014, Klitgaard et al., 2014).

Results and discussion

Study on phylogenetic relationships above section level

A phylogenetic analysis based on the combined nine-gene dataset was conducted to determine the relationship within the Eurotiales. A total of 279 species were included (incl. outgroup species) in the analysis and an overview of the best-fit models for each partition is given in Table 1. The full result of the phylogenetic analysis is shown in Supplementary Fig. S1, and two phylograms with collapsed nodes are shown here (Fig. 1, Fig. 2). Fig. 1 shows the relationship between families and genera within the Eurotiales. Five main lineages are present within the order, and those lineages are treated as families: Aspergillaceae, Elaphomycetaceae, Thermoascaceae, Trichocomaceae, and an unnamed lineage containing species originally described in Penicillium (P. kabunicum, P. moldavicum). Penicillium nodisitatum also belongs to this lineage (see results below) and this species was recently combined in Penicillago. This lineage is therefore named after this genus (Penicillaginaceae, see Taxonomy section).

Table 1.

The best-fit models for each partition proposed by ModelTest-NG based on the corrected Akaike Information Criterion.

Partition	Model
BenA exon	TVM+R4+F
BenA intron	TIM3+R4+F
CCT8	HKY+R4+F
CaM exon	TPM1uf+R4+F
CaM intron	HKY+R4+F
rDNA of ITS region	TrN+G4+F
ITS1-ITS2	GTR+G4+F
LSU	TIM3+G4+F
RPB1	TVM+R4+F
RPB2	TPM1uf+R4+F
SSU	TIM2+G4+F
TSR1	TrN+R4+F
Indel	BIN+ASC_LEWIS

Fig. 1.

Combined phylogeny using nine loci (RPB1, RPB2, Cct8, Tsr1, CaM, BenA, SSU, LSU, ITS). Clades in the phylogram are collapsed showing the relationship between genera and families in the Eurotiales. The phylogram is based on 263 species belonging to the order Eurotiales and 16 species from the order Onygenales (used an outgroup). The species used in the analysis can be found in Supplementary Fig. S1 and Supplementary Table S1.

Fig. 2.

Combined phylogeny using nine loci (RPB1, RPB2, Cct8, Tsr1, CaM, BenA, SSU, LSU, ITS). In the phylogram, only the subgenera and sections of Aspergillus and Penicillium are shown; the other genera are collapsed as one outgroup clade. The phylogram is based on 263 species belonging to the order Eurotiales and 16 species from the order Onygenales (used an outgroup). The species used in the analysis can be found in Supplementary Fig. S1 and Supplementary Table S1.

The accepted, known genera of the Eurotiales (e.g., Houbraken and Samson, 2011, Kocsubé et al., 2016, Guevara-Suarez et al., 2020) are indicated in Fig. 1. Hamigera striata and Talaromyces leycettanus form a unique lineage in the Aspergillaceae and Penicillium lignorum and Talaromyces luteus represent independent lineages in the Trichocomaceae. Hamigera striata is phylogenetically most closely related to Talaromyces leycettanus (88 % bootstrap (BS) support) and those two species are sister to a clade including taxa classified in Hamigera, Pseudopenicillium and Warcupiella (98 % BS). Penicillium lignorum is sister to a clade containing Sagenomella species (100 % BS) and Talaromyces luteus is sister to Thermomyces dupontii and Tm. lanuginosus (100 % BS). These four species are combined in new genera below and the relationship with other genera is discussed in the notes (see Taxonomy section).

Fig. 2 shows the relationship between subgenera and sections within Aspergillus and Penicillium. Six main lineages are present in Aspergillus, representing the subgenera in the genus (subgen. Aspergillus, Circumdati, Cremei, Fumigati, Nidulantes and Polypaecilum). Two clades (sections) are present in subgen. Aspergillus, ten in subgen. Circumdati, one in subgen. Cremei, four in subgen. Fumigati, nine in subgen. Nidulantes and one in subgen. Polypaecilum. The phylogenetic relationship within Aspergillus is well-resolved and the bootstrap values are generally higher than 95 % (Fig. 2). Exceptions are the nodes of sections Flavipedes (89 % BS), Ochraceorosei (94 % BS) and Restricti (61 % BS). In sect. Flavipedes, the relationship of A. neoniveus with the other taxa of the section is moderately supported, while the relationship of A. penicillioides (sect. Restricti) with other representatives of the section (A. conicus, A. restrictus, A. glabripes, A. halophilicus) is poorly supported (Supplementary Fig. S1).

Two main, well-supported lineages are present in Penicillium, representing subgenera Aspergilloides and Penicillium. Subgenus Aspergilloides was divided into two clades: one clade containing the majority of subgen. Aspergillioides taxa, the other including four species: Penicillium alfredii, P. cryptum, P. lagena and P. lassenii. New sections for these species are introduced in the Taxonomy section. Nineteen lineages (sections) are present in subgen. Aspergilloides and 13 lineages in subgen. Penicillium. The majority of branches gained good or full statistical support (>95 % BS) (Fig. 2). The main exception was a clade containing taxa classified in sections Fasciculata, Osmophila, Penicillium and Roquefortorum. The statistical support within this clade was generally moderate or poor.

Phylogenetic analysis of series relationships within Aspergillus and Penicillium

The phylogenetic relationships among members of Aspergillus and Penicillium were studied using a combined 3-gene dataset (BenA, CaM, RPB2). The number of included strains and the length of each partition is given in Table 2. The results of the phylogenetic analyses are discussed in the notes in the Taxonomy section.

Table 2.

Details on the combined datasets used in this study.

Figure number	Dataset	No. species	Length of dataset (bp)
			BenA	CaM	RPB2
Fig. 6	Penicillago	8	467	530	945
Fig. 11	Aspergillus subgen. Aspergillus	55	458	767	1007
Fig. 12	Aspergillus subgen. Circumdati	113	693	811	1017
Fig. 13	Aspergillus subgen. Cremei	19	424	812	1013
Fig. 14	Aspergillus subgen. Fumigati	76	594	717	1014
Fig. 15	Aspergillus subgen. Nidulantes	124	456	805	1014
Fig. 16	Aspergillus subgen. Nigri	32	562	654	1014
Fig. 17	Aspergillus subgen. Polypaecilum	17	538	642	977
Fig. 18	Penicillium subgen. Aspergilloides	331	711	922	801
Fig. 19	Penicillium subgen. Penicillium	145	531	622	978
Fig. 20	Aspergillus texensis	35	875	1194	n/a
Supplementary Fig. S2	Hamigera	12	488	591	1011
Supplementary Fig. S3	Talaromyces	170	633	894	852
Supplementary Fig. S4	Penicillium cellarum	45	429	492	767

Families in Eurotiales

The phylogenetic relationship of families and genera belonging to the Eurotiales is given in Fig. 1. Five lineages, representing families, were present in our phylogenetic analysis. Based on a 4-gene phylogeny and phenotypic characters, Houbraken & Samson (2011) segregated the Trichocomaceae in three families (Aspergillaceae, Thermoascaceae and Trichocomaceae). No representatives of the Elaphomycetaceae were included in that study. Based on a phylogenetic analysis of 320 orthologous clusters from selected species, Quandt et al. (2015) showed that Elaphomyces granulatus (Elaphomycetaceae) is a sister to Trichocomaceae. This relationship is confirmed in our analysis (Fig. 1, Supplementary Fig. S1). Elaphomyces species are ectomycorrhizal (like hypogeous truffles in the Pezizales) and produce subglobose, hypogeous 'truffle' fruiting bodies, which have an organised outer layer of tissue (peridium) that enclose the gleba or spore-bearing tissue (Trappe 1979). The position of Elaphomycetaceae in the Eurotiales is therefore remarkable, and this family represents one of the few independent lineages of the mycorrhizal symbiosis in Ascomycota (Tedersoo et al. 2010). The uniting character of the Elaphomycetaceae with the other families in the Eurotiales is the production of cleistothecia, although there are exceptions (e.g., Trichocoma). Furthermore, a lineage containing Penicillium kabunicum and P. moldavicum is sister to the other families in the Eurotiales and is named Penicillaginaceae below. This indicates that a penicillium-like conidiophore was the basal morphology in the Eurotiales and that this has been lost in Elaphomycetaceae. A comparative genome analysis, including the Penicillaginaceae, might shed insight into the evolution of the ectomycorrhizal association within the Eurotiales. Summarised, our analysis shows that the order Eurotiales contains five families: Aspergillaceae, Elaphomycetaceae, Penicillaginaceae, Thermoascaceae and Trichocomaceae.



Aspergillaceae Link, Abh. Königl. Akad. Wiss. Berlin: 165. 1826 [1824]. MycoBank MB80489.



Type: Aspergillus P. Micheli ex Haller



Description: See Houbraken & Samson (2011) (morphology, phylogeny); Fig. 1, this study (phylogeny).



Elaphomycetaceae Tul. ex Paol., in Saccardo, Syll. Fung. 8: 863. 1889. MycoBank MB80727.



Type: Elaphomyces T. Nees



Description: See Miller & Miller Jr (1984) (morphology), Castellano and Stephens, 2017, Paz et al., 2017 (morphology, phylogeny); Fig. 1, this study (phylogeny).



Penicillaginaceae Houbraken, Frisvad & Samson, fam. nov. MycoBank MB832568.



Etymology: This family is named after the sole genus in this family, Penicillago.



Type: Penicillago Guevara-Suarez, Gené & Dania García



Diagnosis: This family is phylogenetically distinct and sister to the families Aspergillaceae, Elaphomycetaceae, Thermoascaceae and Trichocomaceae in the order Eurotiales (Fig. 1). Conidiophores are penicillium-like and the phialides have a long, narrow neck.



Notes: The newly introduced family Penicillaginaceae includes one genus, Penicillago. This genus was thought to belong to the Aspergillaceae, and was named Penicillago (referring to the close phylogenetic relationship with Penicillium) (Guevara-Suarez et al. 2020). However, this genus forms a unique lineage in the Eurotiales and is rather distantly related to Penicillium in the Aspergillaceae (Fig. 1).



Thermoascaceae Apinis, Trans. Brit. Mycol. Soc. 50: 581. 1967. MycoBank MB81467.



Type: Thermoascus Miehe



Description: See (Apinis 1967) (morphology); Houbraken & Samson (2011) (morphology, phylogeny); Fig. 1, this study (phylogeny).



Trichocomaceae E. Fisch., in Engler & Prantl, Nat. Pflanzenfam. 1(1): 310. 1897. MycoBank MB81485.



Type: Trichocoma Jungh.



Description: See Houbraken & Samson (2011) (morphology, phylogeny); Fig. 1, this study (phylogeny).

Genera

Based on our phylogenetic analysis (Fig. 1), the Aspergillaceae includes 15 genera (Aspergillago, Aspergillus, Dichlaena, Evansstolkia, Hamigera, Leiothecium, Monascus, Penicilliopsis, Penicillium, Phialomyces, Pseudohamigera, Pseudopenicillium, Sclerocleista, Warcupiella, Xerochrysium and Xeromyces), the Thermoascaceae two (Paecilomyces, Thermoascus), the Trichocomaceae eight (Acidotalaromyces, Ascospirella, Dendrosphaera, Rasamsonia, Sagenomella, Talaromyces, Thermomyces, Trichocoma and the Penicillaginaceae one (Penicillago). No material of Dendrosphaera eberhardtii (type of genus Dendrosphaera) and Dichlaena lentisci (type of genus Dichlaena) were available for examination. We follow Pitt et al. (2000) and tentatively accept these genera. A taxonomic review of the Elaphomycetaceae is not part of this study. According to Paz et al. (2017), this family includes two genera, Elaphomyces and Pseudotulostoma; however, the taxonomic status of the latter genus is questionable and molecular data indicate that it is congeneric with Elaphomyces (Castellano et al. 2016). An overview of genera belonging to the Eurotiales is given below.



Acidotalaromyces Houbraken, Frisvad & Samson, gen. nov. MycoBank MB832551.



Etymology: This species requires a low pH for its growth and it was previously classified (as Penicillium lignorum) in Penicillium subgen. Biverticillium, which contains species that are nowadays mostly classified in Talaromyces.



Type: Penicillium lignorum Stolk



Diagnosis: Phylogenetically distinct (Fig. 1). Conidiophores talaromyces-like (Fig. 3). No or very slow growth on regular agar media (e.g., CYA, MEA) and moderate growth on acidified media (pH 3.5).



Fig. 3.

Drawing of Acidotalaromyces lignorum CBS 709.68 showing conidiophores and conidia; from Stolk (1969). Scale bar = 10 μm.

Notes: Acidotalaromyces is a monotypic genus and forms a unique lineage in the Trichocomaceae, phylogenetically related to Sagenomella. It requires acidified agar media (pH 3.5) for growth, as no or very limited growth occurs on regular media of slightly acidic or neutral pH. Acidotalaromyces is known from rotting wood in Europe and potentially produce biotechnologically interesting enzymes.



Ascospirella Houbraken, Frisvad & Samson, gen. nov. MycoBank MB832552.



Etymology: Named after the typical transverse to spiral ridges on the ascospores of the type species.



Type: Penicillium luteum Zukal



Diagnosis: Phylogenetically distinct; conidiophores typically biverticillate, but monoverticillate and irregular forms usually also present; ascomata yellow to orange; ascospores bearing 3–5 conspicuous transverse or spiral ridges or striations (Fig. 4).



Fig. 4.

Morphological characters of Ascospirella lutea. A. Colonies from left to right, after 7 d at 25 °C (top row) CYA, YES, MEA, OA; (bottom row) CYA reverse, YES reverse, DG18, CREA. B. Ascomata on OA after 8 wk at 25 °C. C. Ascoma. D. Asci and ascospores. E. Conidiophores. F. Conidia. G. Ascospores. H. SEM micrograph of ascus with ascospores. Scale bars: C–G = 10 μm; H = 1 μm.

Notes: Ascospirella is a monotypic genus in the Trichocomaceae and is phylogenetically most closely related to Thermomyces. Thermomyces contains thermophilic species (Tm. lanuginosus, Tm. dupontii), while the sole member in Ascospirella (i.e. Ascospirella lutea) is a mesophile. Ascospirella can be further distinguished from Thermomyces by the production of penicillium-like conidiophores and yellow to orange ascomata (Fig. 4). The production of ascospores with conspicuous transverse or spiral ridges or striations is a striking feature for Ascospirella. Similarly, ornamented ascospores are also produced in Trichocoma paradoxa and Talaromyces udagawae. Both species also belong to the Trichocomaceae, but are phylogenetically distinct. Ascospirella readily produces ascomata on agar media (e.g., OA, MEA), while ascoma formation by Trichocoma is only observed on the natural substrate. The ascospores of Ascospirella lutea resemble Talaromyces udagawae and these species were therefore thought to be closely related (Stolk & Samson 1972). These species differ in ascospores size and ornamentation, and ascomatal initials.



Aspergillago Samson et al., Stud. Mycol. 85: 211. 2016. MycoBank MB819186.



Type: Aspergillago clavatoflava (Raper & Fennell) Samson et al.



Notes: Aspergillago was introduced to accommodate Aspergillus clavatoflavus (Kocsubé et al. 2016). The phylogenetic relationship of Aspergillago with other genera was unresolved (Houbraken and Samson, 2011, Kocsubé et al., 2016); however, Fig. 1 shows that this is a sister genus of Penicilliopsis.



Aspergillus P. Micheli ex Haller, Hist. stirp. indig. Helv. inch.: 113. 1768. MycoBank MB7248.



Type: Aspergillus glaucus (L.) Link



Notes: The typical conidiophore structure in Aspergillus is the aspergillum, with a foot cell, non-septate stipe ending in a vesicle on which the metulae and/or phialides are borne. Houbraken & Samson (2011) demonstrated that the type species of Polypaecilum and Phialosimplex with a simpler structure are related to members of sections Cremei and Aspergillus, phylogenetically placing those genera within the classical concept of Aspergillus. Furthermore, Aspergillus paradoxus, A. malodoratus and A. crystallinus characterised by aspergillus-like structures were shown to belong to Penicillium sect. Paradoxa. With these new taxonomic insights based on phylogenetic relationships, the generic boundaries of Aspergillus are now well defined. Samson et al. (2014) recommended methods for the identification and characterization of Aspergillus creating the basis for a stable taxonomy of the genus.

 The genus contains sexual morphs with different structures. In the dual nomenclature era, these structures were recognised as separate sexual genera; however various studies have demonstrated that they are all within the monophyly of Aspergillus (Kocsubé et al., 2016, Steenwyk et al., 2019). The teleomorphic generic name (sexual morphs) are nowadays indicated as morphotypes: eurotium-type, neosartorya-type, emericella-type, petromyces-type, chaetosartorya-type, fennellia-type and neopetromyces-type (Houbraken & Samson 2017). The sexual morph found in sect. Nigri can be regarded as the saitoa-type and studies on the genus Dichlaena are underway to elucidate the relationship with Aspergillus.



Dendrosphaera Pat., Bull. Soc. Mycol. France 23: 69. 1907. MycoBank MB1455.



Type: Dendrosphaera eberhardtii Pat.



Notes: Dendrosphaera (Patouillard 1907) is typified with Dendrosphaera eberhardtii, the sole species in the genus. The genus is phenotypically related to Trichocoma and produces very small brushes of soft hyphae bearing asci and ascospores (Malloch 1985). Kobayasi & Yokoyama (1981) reported that the asexual morph is talaromyces-like (Penicillium subgen. Biverticillium), similar as in Trichocoma. The ascospores of Dendrosphaera germinate poorly or not at all on agar media and no cultures or sequences were available for this study (hence not included in our phylogenetic analysis). The exact taxonomic position of this genus needs to be elucidated, but until that time, we follow Pitt et al. (2000) and (tentatively) accept it in the Trichocomaceae.



Dichlaena Durieu & Mont., Expl. Sci. l'Algérie 1: 405. 1849. MycoBank MB1514.



Type: Dichlaena lentisci Durieu & Mont.



Notes: Dichlaena lentisci, the type of the genus, was described in 1849 and found as mature fruiting bodies on decayed leaves. Malloch & Cain (1972) isolated and studied a strain of this species (TRTC 45715), which produces uniseriate Aspergillus conidiophores and a close relationship with this genus is therefore likely. No material was available for the current study, but studies are underway to elucidate the relationship between Dichlaena and Aspergillus. At this moment, it is too premature to combine D. lentisci in Aspergillus and we therefore tentatively retain Dichlaena. Three other species are described in Dichlaena: D. bovina, D. indica and D. pterodontis. The asexual morph of D. pterodontis is not aspergillus-like, but a hyphomycete producing conidia in slimy heads (Ram 1971). The classification of this species in Dichlaena is doubtful and it is therefore not included in our list of species. Thielavia bovina was combined by Booth (1961) in Dichlaena as D. bovina. The original description of this species (Scalia 1900) is insufficient to support the classification in Dichlaena and the taxonomic status of this species is therefore doubtful. Dichlaena indica was reported to produce an aspergillus asexual morph (Index Fungorum, RecordID 127024) and we therefore tentatively accept this species in Dichlaena.



Evansstolkia Houbraken, Frisvad & Samson, gen. nov. MycoBank MB832553.



Etymology: Named after H.C. Evans and A.C. Stolk, the authors of the sole species in this genus.



Type: Penicillium leycettanum H.C. Evans & Stolk



Diagnosis: Phylogenetically distinct. Conidiophores paecilomyces-like; conidia brown; chlamydospores present, thick-walled; ascospores decorated with somewhat jagged, irregular, mostly longitudinal ridges of different length. Thermotolerant to thermophilic. Fig. 5.



Fig. 5.

Morphological characters of Evansstolkia leycettana (CBS 398.68^T). A. Colonies from left to right, after 7 d at 37 °C unless stated otherwise (top row) MEA 25 °C, MEA, YES, OA; (bottom row) CYA 25 °C, CYA, DG18, CREA. B–C. Conidiophores and conidia. D. SEM micrograph of ascospores. Scale bars: B–C = 10 μm; D = 1 μm.

Notes: Evansstolkia forms a single lineage and is a monotypic genus, containing the species Talaromyces leycettanus (basionym: Penicillium leycettanum). Talaromyces leycettanus was originally classified in Talaromyces because of the production of ascomata that are surrounded by a definite network of pale yellow hyphae and the production of asci in chains. In contrast to these observations, Houbraken & Samson (2011) noted that this species phylogenetically belongs to the Aspergillaceae, instead of the Trichocomaceae, and this observation is confirmed in this study. Talaromyces leycettanus is phylogenetically most closely related to Hamigera striata (88 % BS, Fig. 1). This species produces paecilomyces-like conidiophores, brown coloured conidia, thick-walled chlamydospores and ascospores that are decorated with somewhat jagged, irregular, mostly longitudinal ridges of different length. Furthermore, Tal. leycettanus is thermotolerant to thermophilic. This combination of characters is unique in the Aspergillaceae and we therefore decided to accommodate this species in a novel genus.



Hamigera Stolk & Samson, Persoonia 6: 342. 1971. MycoBank MB2215.



Type: Hamigera avellanea (Thom & Turesson) Stolk & Samson



Notes: The taxonomy of Hamigera and related genera has been subject of various studies. The genus Hamigera was erected for two Talaromyces species (Tal. avellaneus, Tal. striatus) that produce asci singly instead of in chains (Stolk & Samson 1971). Later, the genus Merimbla was introduced for the asexual morph of H. avellanea (Pitt 1979). von Arx (1986) transferred Warcupiella to Hamigera and treated Merimbla and Raperia as congeneric, giving priority to the latter. Peterson et al. (2010) revised the genus Hamigera using a multigene sequence-based approach and accepted seven species. They showed that Warcupiella (and the related asexual genus Raperia) and Hamigera striata (here combined to Pseudohamigera) do not belong to this genus. Merimbla and Hamigera resided in the same lineage, and after the introduction of a single name nomenclature, Merimbla ingelheimensis was transferred to Hamigera ingelheimensis (Igarashi et al. 2014). Talaromyces brevicompactus (asexual morph Merimbla brevicompacta, simultaneously published, identical holotype) is phenotypically similar to Hamigera avellanea (reported as Tal. avellaneus), but differs in their initials, ascospore ornamentation and conidiophore branching (Kong 1999). Samson et al. (2011c) indicated that Talaromyces brevicompactus represents a distinct species in Hamigera. The new combination, Hamigera brevicompacta, is made below. Hamigera avellanea var. alba is phylogenetically related to H. pallida (Supplementary Fig. S2), but the taxonomic status of both species needs further study. In total, nine species are currently accepted: Hamigera avellanea, H. brevicompacta, H. fusca, H. inflata, H. ingelheimensis, H. insecticola, H. pallida, H. paravellanea and H. terricola.



Leiothecium Samson & Mouch., Canad. J. Bot. 53: 1634. 1975. MycoBank MB2719.



Type: Leiothecium ellipsoideum Samson & Mouch.



Notes: Leiothecium is characterised by dark, glabrous ascomata with thin pseudoparenchymatous walls and by ellipsoidal, reticulate, hyaline ascospores (Samson & Mouchacca 1975a). Leiothecium is phylogenetically a sister of a cluster containing the genera Monascus, Xerochrysium and Xeromyces (Fig. 1). Monascus is also phenotypically related to Leiothecium, but differs in producing a thin, plectenchymatous ascomatal wall and smooth-walled ascospores (Samson and Mouchacca, 1975a, Barbosa et al., 2017). Furthermore, Monascus produces a basipetospora-type asexual morph, while an asexual morph is not observed in Leiothecium. Leiothecium also shows some similarity with Ascorhiza (considered a doubtful taxon below) and Hapsidospora (Hypocreales) (Samson & Mouchacca 1975a) because of the presence of cleistothecial ascomata and reticulate ascospores (Marin-Felix et al. 2014). The genus includes two species, Leiothecium ellipsoideum and L. cristatum.



Monascus Tiegh., Bull. Soc. Bot. France 31: 226. 1884. MycoBank MB3247.



Type: Monascus ruber Tiegh.



Notes: The genus Monascus was described by van Tieghem (1884) to accommodate the sexually reproducing species M. ruber and M. mucoroides. The genus is characterised by the production of colourless to light brown cleistothecia, in some species becoming dark brown in time, and smooth-walled ascospores. In the abandoned dual name nomenclature system, the genus Basipetospora was found to be the asexual morph of Monascus and was characterised by the production of aleurioconidia in a basipetal manner from undifferentiated conidiogenous cells that progressively shorten (retrogression, Cole & Samson 1979). The name Monascus was recommended over Basipetospora (Rossman et al. 2016). Monascus is phylogenetically sister to a clade containing the xerophilic genera Xerochrysium and Xeromyces (Fig. 1). Barbosa et al. (2017) conducted a phylogenetic analysis of this genus and resolved Monascus in nine species (Monascus argentinensis, M. flavipigmentosus, M. floridanus, M. lunisporas, M. mellicola, M. pallens, M. purpureus, M. recifensis and M. ruber) and two sections (sections Floridani and Rubri).



Paecilomyces Bainier, Bull. Soc. Mycol. France 23: 27. 1907. MycoBank MB9196.



Type: Paecilomyces variotii Bainier



Notes: Phylogenetic analysis of the 18S rDNA demonstrated that Paecilomyces sensuSamson (1974) is polyphyletic across two subclasses, Sordariomycetidae and Eurotiomycetidae. The type species of this genus, Paecilomyces variotii, and its thermophilic relatives belong in the Eurotiales (Luangsa-Ard et al. 2004). The ascomycete genus Byssochlamys is linked to Paecilomyces sensu stricto. The taxonomy of these genera was studied by Samson et al. (2009) and five Byssochlamys and four Paecilomyces species were accepted. One of their accepted species was Paec. saturatus. This species was based on the variety name Paecilomyces mandshuricus var. saturatus, while species names were also available (Penicillium viniferum, Paec. dactylethromorphus). Penicillium viniferum (1939) would have priority, but was invalidly described (without Latin diagnosis), and the correct name for Paec. saturatus is therefore Paec. dactylethromorphus. With the introduction of a single name nomenclature system, Paecilomyces got priority over Byssochlamys (Rossman et al. 2016). Paecilomyces is nowadays characterised by producing irregularly branched conidiophores bearing phialides with an inflated base and abruptly narrowing to a thin neck and producing olive-brown conidia in chains. They are thermotolerant and some species are able to produce a byssochlamys sexual morph and smooth-walled ellipsoidal ascospores. Rasamsonia is phenotypically most closely related and differs in having more regularly branched conidiophores with distinctly rough-walled structures. Penicillium and Talaromyces are phenotypically similar genera, but generally produce more regularly branched conidiophores, flask-shaped (Penicillium, Talaromyces) or lanceolate (Talaromyces) phialides and conidia in shades of green. Paecilomyces names are available for most Byssochlamys species, except Byssochlamys lagunculariae, and a new combination is proposed below (Paec. lagunculariae). After the taxonomic treatment of Samson et al. (2009), only one new species was described in Paecilomyces, Paec. tabacinus (Crous et al. 2016). The genus currently includes 10 species: Paecilomyces brunneolus, Paec. formosus, Paec. lagunculariae, Paec. dactylethromorphus, Paec. divaricatus, Paec. fulvus, Paec. niveus, Paec. tabacinus, Paec. variotii and Paec. zollerniae. Paecilomyces formosus is invalid, because it was based on the invalidly described species Monilia formosa (nom. inval., Art. 36.1). Paecilomyces maximus might be the correct name to use; however, we wait with this taxonomic change because this species (and therefore also Paec. formosus) might be a complex of at least three species.



Penicillago Guevara-Suarez et al., Fungal Syst. Evol. 5: 64. 2020. MycoBank MB822073.



Type: Penicillago nodositata (Valla) Guevara-Suarez et al.



Notes: The phylogenetic analysis (Fig. 6) shows that four species can be recognised in Penicillago. These species were all originally described in Penicillium (P. kabunicum, P. mirabile, P. moldavicum and P. nodositatum). Only Penicillium nodositatum was recently combined to Penicillago (Guevara-Suarez et al. 2020) as Penicillago nodositata. The others are transferred to Penicillago below. Morphologically, these species seem unrelated and the most striking unifying character is the production of ampulliform phialides that have a relatively long, narrowed neck. They also share the production of penicillium-like conidiophores, produce conidia in green shades, are unable to grow on CYA incubated at 37 °C and have moderate growth on CREA without acid production. A comparison of the macro- and micromorphology of these species is shown in Fig. 7, Fig. 8 and summarised in Table 3. Penicillago nodositata is macroscopically distinct. This species produces relatively smaller and more colourful colonies than the other three members of the genus. Penicillago mirabile (= Penicillium mirabile) grows better on CYAS than the other species in the genus. Penicillago kabunica (= Penicillium kabunicum) and P. moldavica (= Penicillium moldavicum) are phylogenetically closely related and can be differentiated on their conidium ornamentation (Ramírez 1982). The former produces smooth-walled conidia, and the conidia of the latter are echinulate.



Fig. 6.

Phylogenetic trees based on single and a combined data set of BenA, CaM and RPB2 sequences showing the relationship between Penicillago species. The BI posterior probability (pp) values and bootstrap percentages of the maximum likelihood (ML) analysis are presented at the nodes; fully supported branches are thickened. Values less than 70 % bootstrap support (ML) or less than 0.95 posterior probability (Bayesian analysis) are indicated with a hyphen or not shown. The bar indicates the number of substitutions per site.

Fig. 7.

Overview of macromorphology of species classified in Penicillago (Penicillaginaceae), 7-d-old cultures at 25 °C (unless mentioned otherwise). Columns, left to right: P. kabunica CBS 575.90, P. mirabilis CBS 624.72, P. moldavica CBS 574.90, P. nodositata CBS 333.90; rows, top to bottom: CYA 30 °C, CYA, YES, MEA, OA, CREA, DG18, CYAS.

Fig. 8.

Overview of micromorphology of species classified in Penicillago (Penicillaginaceae). A–C.P. kabunica CBS 575.90, conidiophores and conidia. D–F.P. mirabilis CBS 624.72, conidiophores and conidia. G–I.P. moldavica CBS 574.90, conidiophores and conidia. J–L.P. nodositata CBS 333.90, conidiophores and conidia. Scale bar = 10 μm.

Table 3.

Overview of colony diameters on various agar media and microscopic characters of Penicillago species.

Species	Colony diameter, 7 d, in mm								Conidiophore branching1	Shape conidia	Ornamentation conidia	Shape phialide
	CYA	CYAS	YES	MEA	CREA	DG18	CYA30°C	CYA37°C
P. kabunica	36–40	8–12	>60	>60	26–30	30–34	30–36	No growth	Mono- or biverticillate, divaricate	Globose to subglobose, with a pointed end	Smooth walled	Ampulliform, with a long, narrowed neck
P. mirabilis	38–42	18–22	51–59	47–53	23–27	34–38	31–35	No growth	Mono- or biverticillate, occasional monophialidic	Globose to subglobose	Echinulate	Ampulliform, with a conspicuously long, narrowed neck
P. nodositata	19–23	6–9	29–33	26–30	6–8	15–19	8–12	No growth	Predominantly biverticillate	Subglobose to ellipsoidal	Echinulate	Ampulliform, with a conspicuously long, narrowed neck
P. moldavica	32–36	16–19	46–52	60–70	15–19	17–21	23–27	No growth	Mono- or biverticillate, divaricate	Globose to subglobose	Echinulate	Ampulliform, with a long, narrowed neck

¹The examined P. kabunica and P. moldavica strains were degenerated and part of presented data are also based on Ramirez (1982).

Penicilliopsis Solms, Ann. Jard. Bot. Buitenzorg 6: 53. 1887. MycoBank MB3806.



Type: Penicilliopsis clavariiformis Solms



Notes: Penicilliopsis was described by Solms-Laubach (1887) on seeds of Diospyros macrophylla collected in the Botanical Garden of Bogor in Indonesia. This genus was originally introduced for sexually reproducing species and the genera Sarophorum, Stilbothamnium, Stilbodendron and Pseudocordyceps are associated asexual genera. The aforementioned genera were critically re-examined using morphological characters by Samson & Seifert (1985), but this group of fungi is to date not yet studied using molecular techniques. The type of Sarophorum, S. ledermannii, is considered to be conspecific with S. palmicola, and the latter species is regarded to be the asexual morph of Penicilliopsis clavariiformis (Samson & Seifert 1985). In a single name nomenclature system, Penicilliopsis (1887) will have priority over Sarophorum (1916). We therefore consider Sarophorum a synonym of Penicilliopsis, and Sarophorum ledermannii and S. palmicola synonyms of Penicilliopsis clavariiformis. Stilbodendron is typified with S. camerunense (= S. cervinum), and this species is referred to as the asexual morph of Penicilliopsis africana. Based on these data, Stilbodendron is also considered a synonym of Penicilliopsis. Furthermore, Stilbodendron camerunense, S. congoense and S. cervinum are conspecific (Samson & Seifert 1985) and treated there as synonyms of Penicilliopsis africana. Stilbothamnium (type S. togoensis (= Aspergillus togoensis)) was also thought to be related to Penicilliopsis, but Stilbothamnium is currently considered a synonym of Aspergillus (Samson and Seifert, 1985, Samson et al., 2014, Frisvad et al., 2019). Hsieh & Ju (2002) studied the taxonomic position of Pseudocordyceps and the sole species in this genus, Pseudocordyceps seminicola. Cultures obtained from ascospores of the undescribed Penicilliopsis yielded a culture of Pseudocordyceps seminicola. Based on this observation, the name Penicilliopsis pseudocordyceps was introduced for the sexual morph of Pseudocordyceps seminicola. Penicilliopsis, described in the year 1887, has priority over Pseudocoryceps (described in 1936).

 No (ex-)type material of Penicilliopsis africana and P. clavariiformis was included in our study, but CBS 257.33, a representative strain of Penicilliopsis clavariiformis (Samson & Seifert 1985), was used in our phylogenetic study (Fig. 1). Aspergillus zonatus clustered most closely with P. clavariiformis (Kocsubé et al. 2016), and this species was therefore combined in Penicilliopsis. Following Samson and Seifert, 1985, Hsieh and Ju, 2002 and Kocsubé et al. (2016), we include Penicilliopsis africana, P. clavariiformis, P. pseudocordyceps, and P. zonata in the species list below.



Penicillium Link, Mag. Ges. Naturf. Freunde Berlin 3: 16. 1809. MycoBank MB9257.



Type: Penicillium expansum Link



Notes: The generic concept of Penicillium is now well-defined following studies of Houbraken & Samson (2011) and Visagie et al. (2014b). The main conidiophore structure is the penicillus which can be mono- (simple), bi-, ter- or quarterverticillate. Houbraken & Samson (2011) synonymised the genera Torulomyces and Thysanophora with Penicillium. The sexual morphs which were formerly known as Eupenicillium, Chromocleista and Hemicarpenteles are regarded as synonyms on the basis of the single nomenclature. Visagie et al. (2014b) recommended methods for the identification and characterization of Penicillium creating the basis for a stable taxonomy of the genus.



Phialomyces P.C. Misra & P.H.B. Talbot, Canad. J. Bot. 42: 1287. 1964. MycoBank MB9341.



Type: Phialomyces macrosporus P.C. Misra & P.H.B. Talbot



Notes: The genus Phialomyces was described for species that produce large, dark, warted conidia on phialides whose apices are neither prolonged nor divergent (Misra & Talbot 1964). This genus phenotypically resembles Paecilomyces; however, species in this genus produce generally smaller, hyaline or slightly pigmented conidia. Furthermore, the phialides of Paecilomyces have a broad base and a long narrow neck. Based on a 4-gene phylogeny, Houbraken & Samson (2011) classified Phialomyces in the Aspergillaceae, with Sclerocleista being its most closely related sister. This relationship is confirmed in our multigene phylogenetic analysis (Fig. 1). Five species were described in this genus: Phialomyces fusiformis (Rodriguez & Decock 2003), Ph. macrosporus (Misra & Talbot 1964), Ph. microsporus (Zhang et al. 2010), Ph. striatus (Castañeda & Gams 1991) and Ph. taiwanensis (Matsushima 1985).

 Phialomyces striatus was, after Ph. macrosporus, the second species that was described in this genus. Sequences derived from the ex-type strain of this species (CBS 550.89^T) indicate a relationship with taxa in Talaromyces sect. Talaromyces. A new name for this species is given below (as Talaromyces striatoconidius). The taxonomic position of Ph. taiwanensis is uncertain. Mercado-Sierra et al. (1998) transferred this species to Thysanophora. Later, Thysanophora was synonymised with Penicillium and therefore Houbraken & Samson (2011) renamed this species Penicillium taiwanense. Until new information becomes available, we follow Houbraken & Samson (2011) and retain this species in Penicillium. The type culture of Ph. fusiformis (MUCL 43747^T) was not included in our study and no sequence data is present in public databases. Based on the description, this species is phenotypically similar to Ph. macrosporus, the generic type, and we therefore accept this species in the genus. In 2010, a small-spored Phialomyces species was described as Ph. microsporus (Zhang et al. 2010). Sequences obtained from the ex-type strain (DTO 413-G5^T) show that this species belongs to Penicillium sect. Canescentia, and it is most closely related to P. arizonense and P. yarmokense. An overview taxonomic study of this section is lacking, and we therefore wait with combining this species in Penicillium. Besides the two currently accepted species, our phylogenetic analysis (Supplementary Fig. S1) also shows that Penicillium arenicola and Merimbla humicoloides belong to Phialomyces. These species are combined in Phialomyces below and in total four species are treated in this genus: Ph. arenicola, Ph. fusiformis, Ph. humicoloides and Ph. macrosporus.



Pseudohamigera Houbraken, Frisvad & Samson, gen. nov. MycoBank MB832554.



Etymology: Named after Hamigera.



Type: Penicillium striatum Raper & Fennell



Diagnosis: Phylogenetically distinct. Conidiophores penicillium-like, monoverticillate or biverticillate; conidia pale grey; ascospores elliptical with walls bearing a series of wavy, longitudinal flanges or frills. Mesophilic.



Notes: Pseudohamigera is phylogenetically most closely related to Evansstolkia (Fig. 1), a newly erected genus to accommodate Talaromyces leycettanus (see above). The ascospores of Pseudohamigera somewhat resemble those of Evansstolkia (Fig. 5, Fig. 6). The main difference between those genera is their optimum and maximum growth temperature. Pseudohamigera is a mesophile with an optimum growth temperature between 25 and 30 °C and a maximum around 37 °C; Evansstolkia is thermotolerant to thermophilic (optimum around 40 °C; maximum around 55 °C). Hamigera and Pseudohamigera differ markedly from each other in the ornamentation of their ascospores and in their conidial state. The ascospores of Pseudohamigera have 8–12 longitudinal striations while those of Hamigera are pitted. Furthermore, the asexual morph of Pseudohamigera is penicillium-like, and Hamigera has a merimbla-type conidiophore. Stolk & Samson (1971) already mentioned that these differences would justify classifying H. striata (= P. striatum) and H. avellanea in separate genera, but this was deferred at that time. Warcupiella differs from Pseudohamigera by the production of conspicuously spiny ascospores, without a trace of a furrow or equatorial ridge. Furthermore, Warcupiella produces a raperia-type (aspergillus-like) asexual morph.



Pseudopenicillium Guevara-Suarez et al., Fungal Syst. Evol. 5: 66. 2020. MycoBank MB822076.



Type: Pseudopenicillium megasporum (Orpurt & Fennell) Guevara-Suarez, Cano & Gené



Notes: The genus Pseudopenicillium was recently introduced and two species previously classified in Penicillium, Penicillium megasporum and Penicillium giganteum, were combined in this genus. In the same article, Pseudopenicillium cervifimosum was newly described. Pseudopenicillium is characterised by its penicillium-like conidiophores and brown globose conidia with conspicuous disjunctors (Fig. 10). The genus is phylogenetically most closely related to Hamigera (Fig. 1, Guevara-Suarez et al. 2020). Pseudopenicillium differs morphologically from Hamigera by the production of large (6–9 μm), globose, dark or dull green conidia; the conidia of Hamigera are smaller (3–5 μm in length), (sub)ellipsoidal and in shades of brown (avellaneous). A sexual morph in Pseudopenicillium is unknown. Pseudopenicillium produces penicillium-like conidiophores, but can morphologically be differentiated from Penicillium and Penicillago by its short and often irregularly branched conidiophores producing large conidia in short chains. Furthermore, the colonies of Pseudopenicillium species on YES agar have a yeast-like appearance.

Fig. 10.

Morphological characters of Pseudopenicillium megasporum (CBS 256.55^NT). A. Colonies from left to right, after 7 d at 25 °C (top row) CYA, YES, MEA, OA; (bottom row) CYA reverse, YES reverse, DG18, CREA. B–C. Conidiophores and conidia. D. Conidia. Scale bars: B–D = 10 μm.

 Pitt (1980) introduced series Megaspora to accommodate Penicillium megasporum and P. asperosporum. These species could be differentiated based on conidium size: the conidia of P. megasporum are 6–7 μm, and those of P. asperosporum measure 4–5 μm. Pitt (1980) further noted that both species show little affinity with most other Penicillium species, with P. montanense being one of the exceptions. In a more recent taxonomic study by Houbraken et al. (2014b), they treated P. asperosporum as a synonym of P. montanense in section Aspergilloides. Based on sequence data of six loci, Peterson et al. (2010) depicted P. giganteum and P. megasporum as sibling species and accepted both species. CBS 144.69 (ex-type of P. giganteum) differs from CBS 256.55 (ex-type P. megasporum) in producing smaller colonies (CYA 8 mm; YES 15 mm; MEA 15 mm; DG18 10 mm vs CYA 18 mm; YES 21 mm; MEA 19 mm; DG18 17 mm). Furthermore, CBS 144.69 did not grow on CREA, while CBS 256.55 shows weak to moderate growth (7 mm) with weak acid production. We accept both species based on our observations and the phylogenetic data reported in Peterson et al. (2010).



Fig. 9.

Morphological characters of Pseudohamigera striata. A. Colonies from left to right, after 7 d at 25 °C (top row) CYA, YES, MEA, OA; (bottom row) CYA reverse, YES reverse, DG18, CREA. B. Ascomata on OA after 8 wk at 25 °C. C. Ascoma and asci. D. Asci and ascospores. E. Ascospores. F–G. Conidiophores. H. Conidia. I. SEM micrograph of an ascospore. Scale bars: C–H = 10 μm; I = 1 μm.

Rasamsonia Houbraken & Frisvad, Antonie van Leeuwenhoek 101: 411. 2011 [2012]. MycoBank MB519868.



Type: Rasamsonia emersonii (Stolk) Houbraken & Frisvad



Notes: Houbraken et al. (2012b) erected the genus Rasamsonia for the eurotialean Geosmithia species G. argillacea, G. cylindrospora, and G. eburnea (sexual morph, Talaromyces eburneus). The type of Geosmithia, Penicillium lavendulum, is a member of the Hypocreales and this name could therefore not be used for these eurotialean Geosmithia species. In the same publication, also the thermophiles Talaromyces emersonii and Tal. byssochlamydoides were transferred to Rasamsonia. Rasamsonia is characterised by producing olive-brown conidia, cylindrical phialides that usually gradually taper towards the apices and conidiophores with distinctly rough-walled stipes; ascomata, if present, have a scanty covering. Rasamsonia is phylogenetically a sister genus of Trichocoma (Houbraken and Samson, 2011, Houbraken et al., 2012b, Fig. 1). Rasamsonia phenotypically resembles Paecilomyces, and both genera contain thermotolerant species, produce olive brown conidia, and form ascomata with no or scarce ascomatal covering, but Rasamsonia differs from Paecilomyces in having more regularly branched conidiophores with distinctly rough-walled structures (Houbraken et al. 2012b). With the introduction of the genus, Houbraken et al. (2012b) combined five species in the genus (Rasamsonia argillacea, R. eburnea, R. cylindrospora, R. byssochlamydoides, R. emersonii) and described one new species (R. brevistipitata). Six new species were described afterwards, and all are accepted here: Rasamsonia composticola (Su & Cai 2013), Rasamsonia aegroticola, R. piperina (Houbraken et al. 2013), Rasamsonia pulvericola (Tanney & Seifert 2013), Rasamsonia columbiensis (Crous et al. 2016) and Rasamsonia frigidotolerans (Rodríguez-Andrade et al. 2020). This genus currently consists of 12 species. The genus originally only included thermotolerant or thermophilic species that have optimum growth temperatures above 30 °C and maximum growth temperatures above 45 °C (as defined by Cooney and Emerson, 1964, Crisan, 1973, Maheshwari et al., 2000). Interestingly, R. frigidotolerans and R. pulvericola are mesophiles, which means that the generic description should be expanded. The presence of a mesophilic species within this otherwise thermotolerant/thermophilic genus is an interesting subject for investigation of the evolution of thermophily.



Sagenomella W. Gams, Persoonia 10: 100. 1978. MycoBank MB9773.



Type: Sagenomella diversispora (J.F.H. Beyma) W. Gams



Notes: The genus Sagenomella was introduced by Gams (1978) to accommodate acremonium-like fungi and is characterised by the production of connected conidial chains and sympodially proliferating, often centrally swollen phialides. This genus is typified with Sagenomella diversispora, and Houbraken & Samson (2011) showed that this genus is phylogenetically unrelated to Penicillium s. str. Phylogenetically, this genus is most closely related to the new genus Acidotalaromyces (Fig. 1). In contrast to Sagenomella, the sole species in Acidotalaromyces produces penicillium-like conidiophores and has a preference for acidic substrates.

Gams (1978) described three species (Sagenomella alba, S. oligospora, S. verticillata) and combined six in Sagenomella (S. diversispora, S. griseoviridis, S. humicola, S. sagenomatis, S. sclerotialis, S. striatispora). Another six species and one variety were included in the genus afterwards: S. alba var. synnematosa, S. bohemica, S. chlamydospora, S. dimorphica, S. keratitidis, S. ocotl and S. ryukyuensis (Ueda and Udagawa, 1984, Fassatiová and Pecková, 1990, Gené et al., 2003, Seixas et al., 2005, Samson et al., 2011c). A taxonomic revision of the genus using molecular data is lacking.

 Sagenomella alba and S. oligospora do not belong to the Trichocomaceae and should be transferred to another genus. An ITS sequence (MH860843) of the type strain of S. alba (CBS 167.74^T) indicates a relationship with Crocicreas in Helotiales (Leotiomycetes). Comparison of LT633929, an ITS sequence of the type of S. oligospora (CBS 168.74^T) shows that this species probably represents a new genus or species in Sordariomycetes (near Phialemonium). Various species that were originally described in Sagenomella are combined in other genera. Sagenomella bohemica, S. ryukyuensis and S. sagenomatis belong to Talaromyces and the accepted names of those species are Talaromyces bohemicus, Tal. ryukyuensis and Tal. viride, respectively (Yilmaz et al. 2014). Sagenomella chlamydospora, S. keratitidis and S. sclerotialis are classified in Aspergillus as Aspergillus chlamydospora, A. keratitidis and A. sclerotialis, respectively (Samson et al. 2014). Sagenomella dimorphica was invalidly described (without type, Art. 40.1). Based on the data presented above, the following six species are accepted in Sagenomella: S. diversispora, S. griseoviridis, S. humicola, S. ocotl, S. striatispora and S. verticillata. The taxonomic position of S. alba var. synnematosa is unknown and it is unclear whether this variety should be raised to species level or should be considered a synonym of “Sagenomella alba”.



Sclerocleista Subram., Curr. Sci. 41: 757. 1972. MycoBank MB4928.



Type: Sclerocleista ornata (Raper, Fennell & Tresner) Subram.



Notes: The genus Sclerocleista was introduced to accommodate the sexually reproducing species Aspergillus ornatus (Subramanian 1972). Raper et al. (1953) noted that this species produces parenchymatous, purplish cleistothecia that differed markedly from all other Aspergilli. Based on these observations, they suggested that this species should be placed in a novel group of potentially ascosporic species. Another phenotypically closely related species is Aspergillus citrisporus (Raper et al. 1953). The name Sclerocleista thaxteri was introduced for this sexually reproducing Aspergillus species (Subramanian 1972). These two phenotypically similar species are also phylogenetically closely related and the only members of Sclerocleista to date. They form a distinct lineage in the Aspergillaceae, distantly related to Aspergillus, and sister to Phialomyces (Fig. 1).



Talaromyces C.R. Benj., Mycologia 47: 681. 1955. MycoBank MB5347.



Type: Talaromyces flavus (Klöcker) Stolk & Samson



Notes: Benjamin (1955) established the genus Talaromyces for sexual reproducing Penicillium species that produce soft-walled ascomata covered with interwoven hyphae. Several phylogenetic studies (e.g., LoBuglio et al., 1993, Berbee et al., 1995, Ogawa et al., 1997, Ogawa and Sugiyama, 2000, Wang et al., 2007a, Houbraken and Samson, 2011, Samson et al., 2011c) have shown the majority of species belonging to Penicillium subgen. Biverticillium reside in a clade with species producing a talaromyces sexual morph. This clade is distantly related to Penicillium and following the single nomenclature, these taxa were accommodated in Talaromyces (Samson et al. 2011c). Yilmaz et al. (2014) provided a monograph of Talaromyces and accepted 88 species placed in seven sections: Bacillispori, Helici, Islandici, Purpurei, Subinflati, Talaromyces and Trachyspermi. This monograph was the basis of the description of many new taxa and in recent years many new species were described all over the world. The number of species grew rapidly and have now reached more than 170 species. Interestingly, Rodríguez-Andrade et al. (2019) described new Talaromyces species with basipetospora-like conidiophores, which have not been previously described in Talaromyces. The phylogenetic relationships within Talaromyces is given in Supplementary Fig. S3 and an overview of the species within the genus is given below in the list.



Thermoascus Miehe, Selbsterhitz. Heus: 70. 1907. MycoBank MB5446.



Type: Thermoascus aurantiacus Miehe



Notes: The taxonomy of Thermoascus has changed over time and Dactylomyces and Coonemeria are currently considered synonyms (Apinis, 1967, Mouchacca, 1997). Thermoascus is characterised by the production of orange-yellow, brown or red-brown, soft cleistothecia formed in a more or less continuous crust-like layer with a pseudoparenchymatous wall. The species of the genus are thermophilic. The asexual morphs of Thermoascus differ significantly and can be absent, or paecilomyces-, or polypaecilum-like. Thermoascus is phylogenetically related to Paecilomyces (Fig. 1, Houbraken & Samson 2011). This link is illustrated by Byssochlamys verrucosa (= Paecilomyces verrucosus). This species was, based on phenotypic characters, described in Paecilomyces, but it phylogenetically belongs in Thermoascus. A new combination for this species as Th. verrucosus is proposed below. The addition of this species to Thermoascus further expands the phenotypic diversity of the genus.

 Six species (Th. aegyptiacus, Th. aurantiacus, Th. crustaceus, Th. isatschenkoi, Th. taitungiacus and Th. thermophilus) and two varieties (Th. crustaceus var. verrucosus and Th. aurantiacus var. levisporus) are described in Thermoascus. A taxonomic study using sequence data and dealing with all currently described species is lacking. Thermoascus isatschenkoi Malchevskaya (1939) is regarded as a doubtful species of which no satisfactory description exists and no material is available for examination. An LSU sequence from Thermoascus aurantiacus var. levisporus ATCC 46197 (obtained directly from the ATCC website, www.lgcstandards-atcc.org/) is identical to Thermoascus aurantiacus ATCC 204492 and NRRL 5861, and similar to CBS 398.64 (1 bp difference) and CBS 257.34 (2 bp difference). We therefore tentatively treat this species as a synonym of Thermoascus aurantiacus. Thermoascus crustaceus var. verrucosus is a distinct species (see below) and the name Th. yaguchii is introduced for this variety. Sequences show that Th. aegyptiacus is a distinct species, related to Th. crustaceus and Th. yaguchii; the relationship of Th. thermophilus is unresolved. The taxonomic status of Th. taitungiacus remains unknown. In total, eight Thermoascus species are included in the list (including the new combination Th. verrucosus and the new name Th. yaguchii).



Thermomyces Tsikl., Ann. Inst. Pasteur 13: 500. 1899. MycoBank MB10209.



Type: Thermomyces lanuginosus Tsikl.



Notes: Thermomyces is phylogenetically most closely related to Ascospirella, but is phenotypically distinct (see under Ascospirella) (Houbraken & Samson 2011, Fig. 1). Thermomyces is a thermophilic genus and species of the genus are commercially used for the production of various (thermostable) enzymes. The genera have different sexual morphs and five species are described since the introduction of Thermomyces: Th. dupontii, Th. ibadanensis, Th. lanuginosus (generic type), Th. stellatus and Th. verrucosus. Thermomyces stellatus and Th. verrucosus are classified in the Microascaceae and Chaetomiaceae, respectively, and Th. ibadanensis is a synonym of Th. lanuginosus (Houbraken et al., 2014a, Wang et al., 2019b).



Trichocoma Jungh., Praem. Fl. Crypt. Java: 9. 1838. MycoBank MB5551.



Type: Trichocoma paradoxa Jungh.



Notes: The monotypic genus Trichocoma is characterised by asci in hyphal masses or tufts that can measure up to 10–20 mm (Kominami et al., 1952, Malloch, 1985). The sexual morph is only observed on natural substrates and not seen on agar media; a talaromyces-like asexual morph can be present on agar media. Trichocoma is phylogenetically most closely related to Rasamsonia (Fig. 1, Houbraken and Samson, 2011, Houbraken et al., 2012b). The majority of Rasamsonia species are thermotolerant or thermophilic, while Trichocoma paradoxa is mesophilic. Furthermore, Rasamsonia produces scanty ascomatal coverings and distinctly ornamented conidiophore stipes.



Warcupiella Subram., Curr. Sci. 41: 757. 1972. MycoBank MB5762.



Type: Warcupiella spinulosa (Warcup) Subram.



Notes: Warcupiella is a monotypic genus (Subramanian 1972) that was introduced to accommodate the sexually reproducing Aspergillus spinosus (Raper & Fennell 1965). Raper & Fennell (1965) mentioned that the classification of A. spinulosus in Aspergillus was difficult because the ascosporic stage differed from all other ascosporic species known at that time in having large, spiny ascospores without any a trace of equatorial ridges or furrows. Besides the unique sexual morph, A. spinosus also produces an asexual morph that is not typical for Aspergillus. A typical Aspergillus conidiophore terminates in a vesicle on which several metulae or phialides develop synchronously. In contrast, the conidiophores of Warcupiella spinulosa (A. spinulosus) terminate in a subvesicle. This subvesicle is cut off by a septum into a shorter apical cell and a larger basal portion. The apical cell develops into a phialide, and several smaller phialides arising later from the part of the subvesicle below the septum (Subramanian & Rajendran 1979). This observation was sufficient evidence to introduce the new genus Raperia for the asexual morph of Warcupiella (type Raperia spinulosa) (Subramanian & Rajendran 1979). Our phylogenetic analysis confirms the unique position of this species outside Aspergillus. Based on the publication date, Warcupiella has priority over Raperia. This species is phylogenetically sister of a clade containing Pseudopenicillium and Hamigera (Fig. 1). One species is classified in this genus, Warcupiella spinulosa.



Xerochrysium Pitt, IMA Fungus 4: 236. 2013. MycoBank MB807003.



Type: Xerochrysium dermatitidis (A. Agostini) Pitt



Notes: Xerochrysium was erected for xerophilic species belonging to the Eurotiales that produce a chrysosporium-like asexual morph. Species are characterised by the production of aleurioconidia, but also by the formation of chlamydoconidia and arthroconidia (Pitt et al. 2013). Xeromyces, a genus containing the extreme xerophile X. bisporus, is phylogenetically related (Fig. 1) and the xerophilic nature of both indicates a close relationship. Xeromyces is distinguished from Xerochrysium because it is primarily a sexual genus, in which fresh isolates readily produce characteristic asci containing two D-shaped ascospores; unlike Xerochrysium, it does not produce chlamydo- or aleurioconidia, instead producing a rare fraseriella-type asexual morph (Pitt et al. 2013). Two species are described in this genus: Xerochrysium dermatitidis and X. xerophilum (Pitt et al. 2013).



Xeromyces L.R. Fraser, Proc. Linn. Soc. New South Wales 78: 245. 1954. MycoBank MB5830.



Type: Xeromyces bisporus L.R. Fraser



Notes: See also Xerochrysium. The genus Xeromyces has a single species, X. bisporus. This is an extreme xerophile and growth will not occur on media with a high water activity. Xeromyces bisporus produces colourless cleistothecia, with evanescent asci containing two ”D”-shaped ascospores.

Doubtful genera

Ascorhiza Lecht.-Trinka, Compt. Rend. Hebd. Séances Acad. Sci., 192: 499. 1931. MycoBank MB372.



Type: Ascorhiza leguminosarum Lecht.-Trinka



Notes: Ascorhiza is considered a doubtful genus. It was introduced by Lechtova-Trinka (1931) for a cleistothecial ascomycete parasitic in root tubers of Astragalus alopecuroides. No type or other material was available for this study, but from the description and illustrations in Lechtova-Trinka (1931), it is apparent that this fungus produces ascomata, subglobose asci and ellipsoidal, reticulate ascospores that measure 9–10 × 6 μm in size. The fungus was not cultivated, but was observed developing in root tubers, with the ascomata surrounded by the tissue of the tuber. Because of the incomplete description and absence of type material, the genus Ascorhiza is regarded as doubtful (Samson & Mouchacca 1975b).

Infrageneric classification (subgenera, sections and series) in Aspergillus and Penicillium

An updated infrageneric classification system is presented below for Aspergillus and Penicillium. Besides these two genera, an infrageneric classification is also present in Monascus and Talaromyces (Yilmaz et al., 2014, Barbosa et al., 2017). Monascus includes two sections (Floridani and Rubri; Fig. 1) and Talaromyces seven (Bacillispori, Helici, Islandici, Purpurei, Subinflati, Talaromyces and Trachyspermi; Supplementary Fig. S3). Monascus is relatively small with nine accepted species, and a further subdivision in series is not useful or needed. On the other hand, Talaromyces includes 170 species and a series classification of this species-rich genus could be useful. However, the phylogenetic relationships below section level are less resolved (see Supplementary Fig. S3) than for Aspergillus and Penicillium and we therefore decided to wait with the introduction of a series classification until more data becomes available, and/or more species are being described. The infrageneric classification proposed below is mainly based on phylogenetic data and supplemented with phenotype, physiology and extrolite data. As discussed above, these latter characters can be present in most, but not all members of the class (polythetic classes). Descriptions are given for all accepted series and the sections without a formal series classification. None of the series (and subseries) in Thom and Church, 1926, Thom, 1930, Thom and Raper, 1945, Raper and Thom, 1949 and Ramírez (1982) are validly published due to the form in which they are presented (Art. 21.1., 36.1). Only sections and series names are considered and compete; those from different ranks are not included in this study. Aspergillus is subdivided 75 series (73 new, one new combination and the autonym Aspergillus), and Penicillium in 89 series (57 new, six new combinations) (excl. the informally introduced series names, see Table 4, Table 5).



Table 4.

Overview of current infrageneric classification system in Aspergillus; the number of accepted species per section and series are mentioned between brackets².

Subgenus	Section	Series
Aspergillus	Aspergillus (32)	Aspergillus (11)
		Chevalierorum (7)
		Leucocarpi (1)
		Rubri (9)
		Tamarindosolorum (1)
		Teporium (1)
		Xerophili (2)
	Restricti (22, incl. A. tapirirae)	Halophilici (1)
		Penicillioides (9)
		Restricti (9)
		Vitricolarum (2)
Circumdati	Candidi (7)	Candidi1 (7)
	Circumdati (27)	Circumdati (11)
		Sclerotiorum (9)
		Steyniorum (7)
	Flavi (35)	Alliacei (5)
		Avenacei (1)
		Bertholletiarum (1)
		Coremiiformes (2)
		Flavi (16)
		Kitamyces (4)
		Leporum (3)
		Nomiarum (3)
	Flavipedes (15)	Flavipedes (9)
		Neonivei (1)
		Olivimuriarum (1)
		Spelaei (4)
	Janorum (4)	Janorum1 (4)
	Nigri (29)	Carbonarii (4)
		Heteromorphi (2)
		Homomorphi (1)
		Japonici (12)
		Nigri (10)
	Petersoniorum (4)	Petersoniorum1 (4)
	Robusti (1)	Robusti1 (1)
	Tannerorum (1)	Tannerorum1 (1)
	Terrei (17)	Ambigui (2)
		Nivei (6)
		Terrei (9)
Cremei	Cremei (17)	Arxiorum (1)
		Brunneouniseriati (2)
		Cremei (5)
		Inflati (3)
		Pulvini (1)
		Wentiorum (5)
Fumigati	Cervini (10)	Acidohumorum (1)
		Cervini (9)
	Clavati (8)	Clavati1 (8)
	Fumigati (59)	Brevipedes (4)
		Fennelliarum (5)
		Fumigati (10)
		Neoglabri (11)
		Spathulati (2)
		Thermomutati (3)
		Unilaterales (12)
		Viridinutantes (12)
	Vargarum (1)	Vargarum1 (1)
Nidulantes	Aenei (10)	Aenei1 (10)
	Bispori (1)	Bispori1 (1)
	Cavernicolarum (5)	Cavernicolarum (4)
		Egyptiaci (1)
	Nidulantes (74)	Aurantiobrunnei (2)
		Multicolores (6)
		Nidulantes (25)
		Speluncei (6)
		Stellati (13)
		Unguium (5)
		Versicolores (17)
	Ochraceorosei (3)	Funiculosi (1)
		Ochraceorosei (2)
	Raperorum (2)	Raperorum1 (2)
	Silvatici (1)	Silvatici1 (1)
	Sparsi (9)	Biplani (2)
		Conjuncti (4)
		Implicati (1)
		Sparsi (2)
	Usti (25)	Calidousti (12)
		Deflecti (5)
		Monodiorum (1)
		Usti (7)
Polypaecilum	Polypaecilum (16)	Canini (2)
		Kalimarum (2)
		Noonimiarum (5)
		Polypaecilum (2)
		Salinarum (4)
		Whitfieldiorum (1)

¹Informally introduced series (for details, see Notes).

²The taxonomic position of Aspergillus argenteus, A. beijingensis, A. collembolorum, A. crassihyphae, A. ellipsoideus, A. maritimus, A. qizutongii, A. raianus, A. subunguis, A. vinosobubalinus and A. wangduanlii is uncertain and are therefore not included in this Table.

Table 5.

Overview of current infrageneric classification system in Penicillium; the number of accepted species per section and series are mentioned between brackets.

Subgenus	Section	Series
Aspergilloides	Alfrediorum (1)	Alfrediorum1 (1)
	Aspergilloides (53)	Fortuita (1)
		Glabra (7)
		Hoeksiorum (2)
		Improvisa (1)
		Kiamaensia (1)
		Livida (3)
		Longicatenata (2)
		Pinetorum (9)
		Quercetorum (1)
		Saturniformia (1)
		Spinulosa (7)
		Sublectatica (3)
		Thiersiorum (1)
		Thomiorum (12)
		Verhageniorum (2)
	Charlesia (9)	Costaricensia (1)
		Fellutana (2)
		Indica (4)
		Phoenicea (2)
	Cinnamopurpurea (20)	Cinnamopurpurea (3)
		Idahoensia (12)
		Jiangxiensia (2)
		Nodula (3)
	Citrina (42)	Citrina (7)
		Copticolarum (3)
		Euglauca (4)
		Gallaica (1)
		Paxillorum (1)
		Roseopurpurea (2)
		Sheariorum (1)
		Sumatraensia (1)
		Westlingiorum (22)
	Crypta (1)	Crypta1 (1)
	Eremophila (1)	Eremophila1 (1)
	Exilicaulis (58)	Alutacea (2)
		Citreonigra (4)
		Corylophila (11)
		Erubescentia (18)
		Lapidosa (14)
		Restricta (9)
	Gracilenta (6)	Angustiporcata (1)
		Estinogena (1)
		Gracilenta (1)
		Macrosclerotiorum (3)
	Griseola (1)	Griseola1 (1)
	Inusitata (2)	Inusitata1 (2)
	Lanata-Divaricata (76)	Dalearum (13)
		Janthinella (24)
		Oxalica (3)
		Rolfsiorum (16)
		Simplicissima (20)
	Lasseniorum (1)	Lasseniorum1 (1)
	Ochrosalmonea (2)	Ochrosalmonea (2)
	Ramigena (6)	Georgiensia (1)
		Ramigena (5)
	Sclerotiorum (35)	Adametziorum (13)
		Herqueorum (5)
		Sclerotiorum (17)
	Stolkia (7)	Stolkia1 (7)
	Thysanophora (8)	Thysanophora1 (8)
	Torulomyces (15)	Torulomyces1 (15)
Penicillium	Brevicompacta (11)	Brevicompacta (5)
		Buchwaldiorum (2)
		Olsoniorum (3)
		Tularensia (1)
	Canescentia (16)	Atroveneta (5)
		Canescentia (11)
	Chrysogena (18)	Aethiopica (1)
		Chrysogena (12)
		Crustacea (3)
		Goetziorum (1)
		Persicina (1)
	Eladia (2)	Eladia1 (2)
	Fasciculata (32)	Camembertiorum (11)
		Corymbifera (9)
		Gladioli (1)
		Verrucosa (3)
		Viridicata (8)
	Formosana (1)	Formosana1 (1)
	Osmophila (2)	Osmophila (1)
		Samsoniorum (1)
	Paradoxa (9)	Atramentosa (6)
		Paradoxa (3)
	Penicillium (8)	Clavigera (2)
		Digitata (1)
		Italica (2)
		Penicillium (2)
		Sclerotigena (1)
	Ramosum (17)	Lanosa (6)
		Raistrickiorum (3)
		Scabrosa (1)
		Soppiorum (6)
		Virgata (1)
	Robsamsonia (14)	Claviformia (1)
		Glandicolarum (4)
		Robsamsonia (7)
		Urticicolae (2)
	Roquefortorum (5)	Roquefortorum (5)
	Turbata (4)	Turbata1 (4)

¹Informally introduced series (for details, see Notes).

Aspergillus subgen. Aspergillus [autonym], MycoBank MB701330.



Type: Aspergillus glaucus (L.) Link, Mag. Ges. Naturf. Freunde Berlin 3: 16. 1809.



Description: See Gams et al. (1985) (morphology), Houbraken and Samson, 2011, Kocsubé et al., 2016, this study (Fig. 2) (phylogeny).



Section Aspergillus [autonym], MycoBank MB548676.



Type: Aspergillus glaucus (L.) Link, Mag. Ges. Naturf. Freunde Berlin 3: 16. 1809.



Description: See Gams et al. (1985) (morphology), Kocsubé et al. (2016), this study (Fig. 2) (phylogeny), Chen et al., 2017, Sklenář et al., 2017 (morphology, phylogeny).



Series Aspergillus [autonym], MycoBank MB834209.



Type: Aspergillus glaucus (L.) Link, Mag. Ges. Naturf. Freunde Berlin 3: 16. 1809.



Diagnosis: Phylogeny: Series Aspergillus belongs to sect. Aspergillus, subgen. Aspergillus; this series forms a well-supported clade together with members of ser. Rubri (Fig. 11); the phylogenetic relationship of A. cibarius and A. endophyticus is unresolved and these species are tentatively classified in ser. Aspergillus. Morphology & physiology: Colonies restricted on MEA, spreading on low water activity media (e.g., M40Y), yellow or orange; conidiophores uniseriate; no growth on CY20S and M60Y at 37 °C, most species grow moderately on CY20S; rapid growth on M60Y at 25 °C, except A. neocarnoyi (restrictedly CY20S at 25 °C, 3–5 mm after 7 d). Sexual morph eurotium-type, homothallic, yellow; ascospores non-crested, with low crests (<0.5 μm) or irregular crests measuring 0.5–1 μm. Series description based on Chen et al., 2017, Visagie et al., 2017.



Fig. 11.

Combined phylogeny for BenA, CaM and RPB2 data sets showing the phylogenetic relation of species, series and sections within Aspergillus subgen. Aspergillus. The BI posterior probability (pp) values and bootstrap percentages of the maximum likelihood (ML) analysis are presented at the nodes; fully supported branches are thickened. Values less than 70 % bootstrap support (ML) or less than 0.95 posterior probability (Bayesian analysis) are indicated with a hyphen or not shown. The bar indicates the number of substitutions per site. The phylogram is rooted with Hamigera avellanea and Penicillium expansum.

Included species: Aspergillus aerius, A. aurantiacoflavus, A. brunneus, A. cibarius, A. endophyticus, A. glaucus, A. levisporus, A. megasporus, A. neocarnoyi, A. niveoglaucus, A. proliferans.



Extrolites: All species produce echinulins, neoechinulins, isoechinulins, auriglaucins and flavoglaucins in common with most other species in section Aspergillus. Mycophenolic acid and its precursor 5,7-dihydroxy-4-methylphthalide (Grove 1972) has been found in A. brunneus and A. niveoglaucus in series Aspergillus and in A. pseudoglaucus (identified as A. flavus by Grove (1972)) in ser. Rubri, but also in sect. Restricti (in six species classified in all four series of the section (Halophilici, Penicillioides, Restricti and Vitricolarum; Sklenář et al. (2017)).



Series Chevalierorum Houbraken & Frisvad, ser. nov. MycoBank MB832995.



Etymology: Named after the type species of this series, A. chevalieri.



Type: Aspergillus chevalieri (L. Mangin) Thom & Church, The Aspergilli: 111. 1926.



Diagnosis: Phylogeny: Series Chevalierorum belongs to sect. Aspergillus, subgen. Aspergillus; this series is phylogenetically sister to series Aspergillus and Rubri (Fig. 11). Morphology & physiology: Colonies restricted on MEA, spreading on low water activity media (e.g., M40Y), yellow or orange; conidiophores uniseriate; all species grow rapid on M60Y at 37 °C, and CY20S and M60Y at 25 °C; all except A. caperatus and A. costiformis grow on CY20S at 37 °C. Sexual morph eurotium-type, homothallic, yellow; ascospores with high crests (0.5 μm). Series description based on Chen et al. (2017).



Included species: Aspergillus caperatus, A. chevalieri, A. costiformis, A. cristatus, A. intermedius, A. montevidensis, A. porosus.



Extrolites: All species produce echinulins, neoechinulins, isoechinulins, auroglaucins and flavoglaucins in common with most other species in sect. Aspergillus.



Series Leucocarpi Houbraken & Frisvad, ser. nov. MycoBank MB832996.



Etymology: Named after the type species of this series, A. leucocarpus.



Type: Aspergillus leucocarpus Hadlok & Stolk, Antonie van Leeuwenhoek 35: 9. 1969.



Diagnosis: Phylogeny: Series Leucocarpi belongs to sect. Aspergillus, subgen. Aspergillus and is an early branch in sect. Aspergillus (together with ser. Teporium) (Fig. 11). Morphology & physiology: Colonies restricted; conidial colour en masse greyish green or dark green; conidiophores uniseriate; no growth on CY20S at 37 °C, moderate growth on M60Y at 37 °C. Sexual morph eurotium-type, homothallic, white (without the characteristic yellow colour seen in other series of sect. Aspergillus); ascospores in surface view globose to subglobose, spore bodies slightly verruculose, in side view lenticular, furrow pronounced, with scattered protuberances, crests 0.5 μm. Series description based on Chen et al. (2017).



Included species: Aspergillus leucocarpus.



Extrolites: Echinulins, epiheveadrides, neoechinulins (Chen et al. 2017). Aspergillus leucocarpus is the only species in sect. Aspergillus that does not produce auroglaucins and flavoglaucins.



Series Rubri Houbraken & Frisvad, ser. nov. MycoBank MB832997.



Etymology: Named after the type species of this series, A. ruber.



Type: Aspergillus ruber (Jos. König et al.) Thom & Church, Aspergillus: 112. 1926.



Diagnosis: Phylogeny: Series Rubri belongs to sect. Aspergillus, subgen. Aspergillus; this series forms a well-supported clade together with members of ser. Aspergillus (Fig. 11). Morphology & physiology: Colonies restricted on MEA, spreading on low water activity media (e.g., M40Y), yellow or orange; conidiophores uniseriate; no growth on CY20S at 37 °C, four species (A. appendiculatus, A. cumulatus, A. mallochii, A. sloanii) cannot grow on M60Y at 37 °C, rapid growth on M60Y at 25 °C, growth on CY20S at 25 °C (except A. appendiculatus). Sexual morph eurotium-type, homothallic; ascospores non-crested or with reduced crests, the only exception is A. cumulatus, which produces irregular, low (<0.5 μm) crests. Series description based on Chen et al. (2017).



Included species: Aspergillus appendiculatus, A. cumulatus, A. mallochii, A. microperforatus, A. pseudoglaucus, A. ruber, A. sloanii, A. tonophilus, A. zutongqii.



Extrolites: Like most other species in sect. Aspergillus, all species produce echinulins, isoechinulins, neoechinulins, auroglaucins and flavoglaucin (ascomatal metabolites) (Chen et al. 2017). Aspergillus pseudoglaucus in addition produces mycophenolic acid and asperentins (Séguin et al., 2014, Chen et al., 2017, Mouhamadou et al., 2017). Aspergillus ruber produces large amounts of rubrocristin and erythroglaucin, giving the species its red colour (Arai et al., 1989a, Wang et al., 2007b (misidentified as A. variecolor), Li et al. 2017) in addition to asperinines, asperflavin, emodin, physcion, but the latter three are also produced by many other species in sect. Aspergillus.



Series Tamarindosolorum Houbraken & Frisvad, ser. nov. MycoBank MB832998.



Etymology: Named after the type species of this series, A. tamarindosoli.



Type: Aspergillus tamarindosoli A.J. Chen et al., Stud. Mycol. 88: 123. 2017.



Diagnosis: Phylogeny: Series Tamarindosolorum belongs to sect. Aspergillus, subgen. Aspergillus; this series is sister to series Aspergillus, Chevalierorum and Rubri (Fig. 11). Morphology & physiology: Colonies restricted on MEA, spreading on low water activity media (e.g., M40Y); conidial colour en masse pale green to greyish green; conidiophores uniseriate; no growth on CY20S at 37 °C. Sexual morph eurotium-type, homothallic, yellow; ascospores hyaline, in surface view globose to subglobose, spore bodies verruculose, in side view lenticular, furrow present, crests irregular, 0.5–1.5 μm. Series description based on Chen et al. (2017).



Included species: Aspergillus tamarindosoli.



Extrolites: Asperflavin, auroglaucin, bisanthrons, echinulins, emodin, dihydroauroglaucin, epiheveadrides, flavoglaucin, isoechinulins, neoechinulins, physcion, tetrahydroauroglaucin (Chen et al. 2017).



Series Teporium Houbraken & Frisvad, ser. nov. MycoBank MB832999.



Etymology: Named after the type species of this series, A. teporis.



Type: Aspergillus teporis A.J. Chen et al., Stud. Mycol. 88: 123. 2017.



Diagnosis: Phylogeny: Series Teporium belongs to sect. Aspergillus, subgen. Aspergillus and is an early diverging lineage in sect. Aspergillus (together with ser. Leucocarpi) (Fig. 11). Morphology & physiology: Colonies on MEA restricted, spreading on media with low water activity; conidial colour en masse greyish green to dark green; conidiophores uniseriate; fast growth on CY20S and M60Y at 37 °C. Sexual morph eurotium-type, homothallic, cream yellow; ascospores in surface view globose to subglobose, spore bodies slightly verruculose, in side view lenticular, furrow pronounced, with scattered protuberances, crests 0.5 μm. Series description based on Chen et al. (2017).



Included species: Aspergillus teporis.



Extrolites: Echinulins, epiheveadrides, isoechinulins, neoechinulins.



Series Xerophili Houbraken & Frisvad, ser. nov. MycoBank MB833000.



Etymology: Named after the type species of this series, A. xerophilus.



Type: Aspergillus xerophilus Samson & Mouch., Antonie van Leeuwenhoek 41: 348. 1975.



Diagnosis: Phylogeny: Series Xerophili belongs to sect. Aspergillus, subgen. Aspergillus; this series contains early diverging species sister to series Aspergillus, Chevaleriorum, Rubri and Tamarindosolorum (Fig. 11). Morphology & physiology: No growth on CYA and MEA, colonies spreading on low water activity media; conidial colour undetermined, sporulation absent; conidiophores uniseriate; no growth on CY20S, rapid growth on M60Y; Aspergillus osmophilus grows rapidly on M60Y at 37 °C, while A. xerophilus does not grow under this condition. Sexual morph eurotium-type, homothallic, yellow; ascospores with low crests (<0.5 μm). Series description based on Chen et al. (2017).



Included species: Aspergillus osmophilus, A. xerophilus.



Extrolites: Asperflavin, auroglaucins, flavoglaucin, echinulins and neoechinulins are shared with most other series in sect. Aspergillus. Sulochrin is only produced by A. xerophilus in sect. Aspergillus but is also produced by other Aspergilli such as A. terreus and A. wentii (Curtis et al., 1970, Assante et al., 1980).



Notes on series in sect. Aspergillus: The production of yellow cleistothecia (except in ser. Leucocarpi), lenticular ascospores and uniseriate conidiophore heads are characters shared by members of section Aspergillus. The species in this section are osmo-, xero- or halotolerant, have a worldwide distribution and are common in indoor air, house dust, cereals, and food products with low water activity (Chen et al., 2017, Visagie et al., 2017). Hubka et al. (2013a) recognised three clades in sect. Aspergillus (A. chevalieri-clade, A. glaucus-clade, A. ruber-clade) and Chen et al. (2017) also included the “A. xerophilus-clade”. These four clades are treated as series in our study. Aspergillus cibarius and A. endophyticus belong to a large well-supported clade that includes series Aspergillus and Rubri. The exact relationship of these species remains unresolved because of weakly supported branching within this clade; however, we tentatively classify them in ser. Aspergillus. Aspergillus leucocarpus, A. tamarindosoli, A. teporis each form single-species clades and are here treated as separate series (Fig. 11). The features of the asexual morph and macromorphology of colonies were of less importance to distinguish the various series. Red hyphae were uniformly absent from species of the ser. Chevaleriorum but can be absent in members of other series as well (Hubka et al. 2013a).

Section Restricti

Type: Aspergillus restrictus G. Sm., J. Textile Inst. 22: 115. 1931.



Description: See Gams et al. (1985) (morphology), Kocsubé et al. (2016) (phylogeny), Sklenář et al. (2017) (morphology, phylogeny).



Series Halophilici Houbraken & Frisvad, ser. nov. MycoBank MB833001.



Etymology: Named after the type species of this series, A. halophilicus.



Type: Aspergillus halophilicus C.M. Chr. et al., Mycologia 51: 636. 1961.



Diagnosis: Phylogeny: Series Halophilici belongs to sect. Restricti, subgen. Aspergillus; the phylogenetic position of ser. Halophilici is unresolved in Fig. 11, but is closely related to series Restricti and Vitricolarum in Supplementary Fig. S1. Morphology & physiology: No growth on MEA, CYA and M40Y; conidiophores uniseriate, radiate conidial heads (sparsely produced); ser. Halophilici are not able to grow on agar media, including those with high sugar (M60Y) or salt (MEA + 10 % NaCl), growth is present on Czapek agar supplemented with 70 % sucrose. Sexual morph eurotium-type, homothallic, hyaline to pale yellow, globose to subglobose; ascospores hyaline, lenticular with two equatorial crests. Series description based on Sklenář et al. (2017).



Included species: Aspergillus halophilicus.



Extrolites: Asperphenamate, cristatin A, echinulin, mycophenolic acid (trace), preechinulin (Micheluz et al., 2016, Sklenář et al., 2017). The white ascomata of A. halophilicus do not contain auroglaucins, flavoglaucin or anthraquinones, explaining the absence of yellow colours in the ascomata.



Series Penicillioides Houbraken & Frisvad, ser. nov. MycoBank MB833002.



Etymology: Named after the type species of this series, A. penicillioides.



Type: Aspergillus penicillioides Speg., Revista Fac. Agron. Univ. Nac. La Plata 2: 246. 1896.



Diagnosis: Phylogeny: Series Penicillioides belongs to sect. Restricti, subgen. Aspergillus; the series is sister to the other series in this section, though with strong statistical support in ML (>95 % BS) and poor support in Bayesian analysis (<0.95 pp) (Fig. 11). Morphology & physiology: Colonies restricted on MEA and CYA or growth absent, moderate on M40Y; conidiophores uniseriate, with globose conidial heads and sometimes later becoming radiate, stipe surface (SEM) with hairs; no growth on CY20S at 37 °C, members of the series Penicillioides are the most xerophilic of the section. Sexual morph unknown. Series description based on Sklenář et al. (2017).



Included species: Aspergillus canadensis, A. clavatophorus, A. hordei, A. infrequens, A. magnivesiculatus, A. penicillioides, A. reticulatus, A. salinicola, A. tardicrescens.



Extrolites: Most species produce asperglaucide, while two species produce asperphenamate. One species produces mycophenolic acid and another species produces chrysogine. Three species produce echinulin and two species produce antarone A (Sklenář et al. 2017).



Series Restricti Houbraken & Frisvad, ser. nov. MycoBank MB833003.



Etymology: Named after the type species of this series, A. restrictus.



Type: Aspergillus restrictus G. Sm., J. Textile Inst. 22: 115. 1931.



Diagnosis: Phylogeny: Series Restricti belongs to sect. Restricti, subgen. Aspergillus and is a sister of series Vitricolarum (Fig. 11). Morphology & physiology: Colonies restricted on MEA and CYA or growth absent, moderate or spreading on M40Y; conidiophores uniseriate, with compact or loosely columnar heads, stipe surface (SEM) with hairs; no growth on CY20S at 37 °C (except A. pachycaulis). Sexual morph unknown. Series description based on Sklenář et al. (2017).



Included species: Aspergillus caesiellus, A. conicus, A. destruens, A. domesticus, A. gracilis, A. pachycaulis, A. pseudogracilis, A. restrictus, A. villosus.



Extrolites: All species produce asperphenamate, while two species produce asperglaucide. Four species produce clavatol, one orthosporins and one fulvic acid analog PI-4. Two species can produce mycophenolic acid (Sklenář et al. 2017).



Series Vitricolarum Houbraken & Frisvad, ser. nov. MycoBank MB833004.



Etymology: Named after the type species of this series, A. vitricola.



Type: Aspergillus vitricola [as “vitricolae”] Ohtsuki, Bot. Mag. (Tokyo) 75: 436. 1962.



Diagnosis: Phylogeny: Series Vitricolarum belongs to subgen. Aspergillus, sect. Restricti and is a sister of ser. Restricti (Fig. 11). Morphology & physiology: Colonies restricted on MEA and CYA or growth absent, moderate on M40Y; conidiophores uniseriate, with radiate heads, stipe surface (SEM) smooth; no growth on CY20S at 37 °C. Sexual morph unknown. Series description based on Sklenář et al. (2017).



Included species: Aspergillus glabripes, A. vitricola.



Extrolites: Aspergillus glabripes produces asperphenamate; A. vitricola produces asperglaucide, and one isolate produces an orthosporin (Sklenář et al. 2017).



Notes on series in sect. Restricti: Section Restricti members are xerophilic and grow optimally on low water activity substrates (containing high concentrations of sugar or salt). Sklenář et al. (2017) recognised five clades (A. conicus-, A. halophilicus-, A. penicillioides-, A. restrictus-, A. vitricola-clade) in their monographic study on sect. Restricti. In our study, we introduce four series for these five clades. The A. restrictus- and A. conicus-clades are combined in ser. Restricti because of their unresolved phylogenetic relationship (Fig. 11, Supplementary Fig. S1). The shape of conidial heads can be used to distinguish the series in sect. Restricti. Conidial heads of ser. Restricti form compact or loose columns, those of ser. Vitricolarum are radiate and ser. Penicillioides mainly have globose conidial heads, which might become radiate after prolonged incubation (Sklenář et al. 2017). Ascomata are only produced in ser. Halophilici. Furthermore, ser. Restricti species (e.g., A. caesiellus, A. pachycaulis, A. restrictus) are less xerophilic compared to ser. Penicillioides, which contains the most xerophilic species of the section (e.g., A. penicillioides).



Aspergillus subgen. Circumdati W. Gams et al., Adv. Pen. Asp. Syst.: 59. 1986 [1985]. MycoBank MB832507.



Type: Aspergillus alutaceus Berk. & M.A. Curtis, Grevillea 3: 108. 1875 (= Aspergillus ochraceus).



Description: See Gams et al. (1985) (morphology), Houbraken and Samson, 2011, Kocsubé et al., 2016, this study (Fig. 2) (phylogeny).



Section Candidi W. Gams et al., Adv. Pen. Asp. Syst.: 61. 1986 [1985]. MB832512.



Type: Aspergillus candidus Link, Mag. Ges. Naturf. Freunde Berlin 3: 16. 1809.



Description: Phylogeny: Series Candidi belongs to subgen. Circumdati, sect. Candidi and is sister to sect. Petersoniorum (Supplementary Fig. S1, Fig. 12). Morphology & physiology: Colonies restricted or moderate; conidial colour en masse white or yellow; conidiophores biseriate. Sexual morph unknown; sclerotia produced by some species, black or purple-black. Also see Peterson et al., 2008, Kocsubé et al., 2016 (phylogeny), Varga et al., 2007b, Hubka et al., 2018b (morphology, phylogeny).



Fig. 12.

Combined phylogeny for BenA, CaM and RPB2 data sets showing the phylogenetic relation of species, series and sections within Aspergillus subgen. Circumdati (excl. sect. Nigri, see Fig. 13). The BI posterior probability (pp) values and bootstrap percentages of the maximum likelihood (ML) analysis are presented at the nodes; fully supported branches are thickened. Values less than 70 % bootstrap support (ML) or less than 0.95 posterior probability (Bayesian analysis) are indicated with a hyphen or not shown. The bar indicates the number of substitutions per site. The phylogram is rooted with Hamigera avellanea.

Included species: Aspergillus campestris, A. candidus, A. dobrogensis, A. pragensis, A. subalbidus, A. taichungensis, A. tritici.



Extrolites: Most species in sect. Candidi produce the shikimic acid derived secondary metabolites chloroflavonins, terphenyllins, candidusins and xanthoascins (Hubka et al. 2018b), in addition to the terpene-derived taichunins (Kato et al. 2018) and the amino acid derived bicyclo [2.2.2]diazaoctane ring containing taichunamides (Kagiyama et al. 2016).



Notes on sect. Candidi: No subdivision of sect. Candidi is proposed, and ser. Candidi is only informally introduced here (see Table 4). Section Petersoniorum is phylogenetically distant but is the most closely related section (Fig. 2, Jurjević et al. 2015), and some species in this section also produce conidia in white or yellow shades. Section Candidi species produce predominantly globose vesicles commonly reaching or exceeding a diameter of 20 μm, while the vesicles in sect. Petersoniorum species do not exceed 20 μm in diam and are variable in shape (pyriform, subglobose, elongate near angular or penicillium-like). Furthermore, sect. Candidi species can produce black or purple-black sclerotia, while the sclerotia in sect. Petersoniorum are pale yellow to brown (Jurjević et al. 2015).



Section Circumdati W. Gams et al., Adv. Pen. Asp. Syst.: 59. 1986 [1985]. MycoBank MB832508.



Type: Aspergillus alutaceus Berk. & M.A. Curtis, Grevillea 3: 108. 1875 (= Aspergillus ochraceus).



Description: See Gams et al. (1985) (morphology), Visagie et al. (2014c) (morphology, phylogeny), Steenwyk et al. (2019) (genome).



Series Circumdati Houbraken & Frisvad, ser. nov. MycoBank MB832987.



Etymology: This series is based on the same type as sect. Circumdati, and therefore also named after this section.



Type: Aspergillus alutaceus Berk. & M.A. Curtis, Grevillea 3: 108. 1875 (= Aspergillus ochraceus).



Diagnosis: Phylogeny: Series Circumdati belongs to subgen. Circumdati, sect. Circumdati and is phylogenetically most closely related to ser. Sclerotiorum; the node is fully supported in the Bayesian analysis (1.00 pp, data not shown), but bootstrap support is lacking (<70 %, Fig. 12). Morphology & physiology: Colonies spreading; conidial colour en masse mostly light yellow to ochre; conidiophores biseriate; generally no growth at 37 °C or restricted (<20 mm, 7 d, CYA), some species grow more rapid (A. pallidofulvus, A. muricatus). Sexual morph generally not observed in culture, except in A. muricatus (homothallic, neopetromyces-type); sclerotia production common, white, cream or yellow. Series description based on Visagie et al. (2014c).



Included species: Aspergillus affinis, A. auricomus, A. cretensis, A. melleus, A. muricatus, A. ochraceus, A. ostianus, A. pallidofulvus, A. sesamicola, A. westerdijkiae, A. westlandensis.



Extrolites: Circumdatins and destruxins are only produced in ser. Circumdati.



Series Sclerotiorum Houbraken & Frisvad, ser. nov. MycoBank MB832581.



Etymology: Named after the type species of the series, Aspergillus sclerotiorum.



Type: Aspergillus sclerotiorum G. A. Huber, Phytopathology 23: 306. 1933.



Diagnosis: Phylogeny: Series Sclerotiorum belongs to subgen. Circumdati, sect. Circumdati and is phylogenetically sister to series Circumdati and Steyniorum, though statistical support is lacking (<70 % BS, Fig. 12); The Bayesian analysis posterior probability (pp) positions this series as sister to ser. Circumdati (1.00 pp, data not shown). Morphology & physiology: Colonies spreading; conidial colour en masse mostly light yellow to ochre; conidiophores biseriate; good growth at 37 °C > 20 mm (7 d, CYA), with exception of A. roseoglobulosus (Visagie et al. 2014c). Sexual morph unknown; sclerotia production common, white, cream or yellow.



Included species: Aspergillus bridgeri, A. fresenii, A. neobridgeri, A. persii, A. pseudosclerotiorum, A. roseoglobulosus, A. salwaensis, A. sclerotiorum, A. subramanianii.



Extrolites: This series includes a species, A. persii, producing aspernidines, cyclopenins, mevinolins (= lovastatins), and sclerotiumins (= aspersclerotiorones) (reported as A. sclerotiorum) (Phainuphong et al., 2016, Bao et al., 2017, Phainuphong et al., 2017b, Phainuphong et al., 2018b, Lebar et al., 2019). Cyclopenin, radarins, secalonic acid A, secopenitrem D and sulphinines is only found in species in ser. Sclerotiorum (Visagie et al. 2014c). No species in ser. Sclerotiorum produce mellein.



Series Steyniorum Houbraken & Frisvad, ser. nov. MycoBank MB832582.



Etymology: Named after the type species of the series, Aspergillus steynii.



Type: Aspergillus steynii Frisvad & Samson, Stud. Mycol. 50: 39. 2004.



Diagnosis: Phylogeny: Series Steyniorum belongs to subgen. Circumdati, sect. Circumdati and is sister to a clade containing ser. Circumdati, but this relationship is lacking statistical support (Fig. 12); Bayesian analysis places this series confidently (1.00 pp) as a sister to series Circumdati and Sclerotiorum (phylogram not shown). Morphology & physiology: Colonies spreading; conidial colour en masse mostly light yellow to ochre; conidiophores biseriate; generally, no growth at 37 °C or sometimes restricted growth (<20 mm, 7 d, CYA). Sexual morph unknown; sclerotia production common, white, cream or yellow. Series description based on Visagie et al. (2014c).



Included species: Aspergillus elegans, A. insulicola, A. occultus, A. ochraceopetaliformis, A. pseudoelegans, A. pulvericola, A. steynii.



Extrolites: Metabolites only produced by species in ser. Steyniorum include antibiotic Y, asteltoxins, cycloechinulin, insulicolides, N-methylepiamauuromine, ochrindols, quinolactacin, and verruculogen TR-2 (Visagie et al. 2014c). No species in ser. Steyniorum produce aspergamides, mellamides, neohydroxyaspergillic acids or petromurins (Visagie et al. 2014c).



Notes on series in sect. Circumdati: The extrolites produced in the three series of sect. Circumdati are similar and the mycotoxin ochratoxin A is produced in all series of the section. Extrolites include aspergamides (= stephacidins = sclerotiamides) (not yet found in species in ser. Steyniorum), aspochracins / sclerotiotides, aspyrones, circumdatins, mellamides, melleins, ochratoxins, orthosporins, penicillic acids, and xanthomegnins (Visagie et al. 2014c). The proposed series classification is based on the results of the multigene phylogeny (Fig. 12). Based on a phylogenetic analysis, Visagie et al. (2014c) recognised seven main clades in sect. Circumdati. One clade included A. robustus and this clade is raised to section level (sect. Robusti; see below) (Jurjević et al. 2015). The A. auricomus-, A. muricatus-, and A. ochraceus-clades fide Visagie et al. (2014c) are treated here as ser. Circumdati. Even though A. auricomus and A. westlandensis (together forming the A. auricomus-clade; Visagie et al. 2014c) form a sister clade to the other members of ser. Circumdati (Fig. 12), we did not find any additional evidence to treat these two species as a separate series. Series Circumdati includes species that produce aspochracins, mellamides, circumdatins and aspergamides; this series apparently lost the ability to produce aspochracins (Visagie et al. 2014c). Series Sclerotiorum represents the A. fresenii-clade fide Visagie et al. (2014c); ser. Steyniorum includes the A. ochraceopetaliformis-clade fide Visagie et al. (2014c), A. steynii and A. elegans. The latter two species were treated as a separate clade (Visagie et al. 2014c), and produce TR-2 and cycloechinulin, two extrolites not produced by any other species of sect. Circumdati.



Section Flavi W. Gams et al., Adv. Pen. Asp. Syst.: 60. 1986 [1985]. MycoBank MB832510.



Type: Aspergillus flavus Link, Mag. Ges. Naturf. Freunde Berlin 3: 16. 1809.



Description: See Gams et al. (1985) (morphology), Kocsubé et al. (2016), this study (Fig. 2) (phylogeny), Frisvad et al. (2019) (morphology, phylogeny).



Series Alliacei Houbraken & Frisvad, ser. nov. MycoBank MB832583.



Etymology: Named after the type species of the series, Aspergillus alliaceus.



Type: Aspergillus alliaceus Thom & Church, Aspergilli: 163. 1926.



Diagnosis: Phylogeny: Series Alliacei belongs to subgen. Circumdati, sect. Flavi and is sister to a large clade containing series Bertholletiarum, Coremiiformes, Flavi, Kitamyces and Nomiarum (Fig. 12). Morphology & physiology: Colonies spreading; conidial colour en masse yellow; conidiophores biseriate; growth at 37 °C, no or poor growth at 42 °C; reverse on AFPA (Aspergillus Flavus and Parasiticus Agar) cream. Sexual morph generally not observed in culture, present in A. alliaceus, petromyces-type, homothallic; sclerotia often present, black. Series description based on Frisvad et al. (2019).



Included species: Aspergillus alliaceus, A. lanosus, A. magaliesburgensis, A. neoalliaceus, A. vandermerwei.



Extrolites: Certain secondary metabolites, such as altersolanols, asperlicins, burnettienes / phaeospelides, burnettramic acid, griseofulvins, mevinolins, nalgiovensins / allianthrones, and ochratoxins are only found in ser. Alliacei species and not in any other species of sect. Flavi (Goetz et al., 1985, Liesch et al., 1985, Liesch et al., 1988, Mandelare et al., 2018, Frisvad et al., 2019, Li et al., 2019a, Morishita et al., 2019).



Series Avenacei Houbraken & Frisvad, ser. nov. MycoBank MB832584.



Etymology: Named after the type species of the series, Aspergillus avenaceus.



Type: Aspergillus avenaceus G. Sm., Trans. Brit. Mycol. Soc. 26: 24. 1943.



Diagnosis: Phylogeny: Series Avenacei belongs to subgen. Circumdati, sect. Flavi, encompassing the earlier diverging species to all other series of sect. Flavi (Fig. 12). Morphology & physiology: Colonies spreading; conidial colour en masse beige; conidiophores biseriate; growth at 37 °C, no growth at 42 °C; reverse on AFPA cream. Sexual morph unknown; sclerotia often present, large, black. Series description based on Frisvad et al. (2019).



Included species: Aspergillus avenaceus.



Extrolites: Aspergillus avenaceus is the only species in sect. Flavi that does not produce kojic acid, but isolates in the species can produce avenaciolides, and in common with other species in sect. Flavi altersolanols, aspirochlorins, pseurotin A and 3-nitropropionic acid (Frisvad et al. 2019).



Series Bertholletiarum Houbraken & Frisvad, ser. nov. MycoBank MB832988.



Etymology: Named after the type species of the series, Aspergillus bertholletiae.



Type: Aspergillus bertholletiae [as “bertholletius”] Taniwaki et al., PLoS ONE 7: e42480, 6. 2012.



Diagnosis: Phylogeny: Series Bertholletiarum belongs to subgen. Circumdati, sect. Flavi and is sister to a clade containing series Flavi, Kitamyces and Nomiarum (Fig. 12). Morphology & physiology: Colonies spreading; conidial colour en masse brown; conidiophores biseriate; good growth at 37 °C, no growth at 42 °C; reverse on AFPA cream. Sexual morph unknown; sclerotia not observed in culture. Species associated with coconut trees. Series description based on Frisvad et al. (2019).



Included species: Aspergillus bertholletiae.



Extrolites: In common with species from other series in sect. Flavi, A. bertholletiae produces cyclopiazonic acid, kojic acid, 3-O-methylsterigmatocystin, parasiticolides, tenuazonic acid and ustilaginoidin C (Frisvad et al. 2019).



Series Coremiiformes Houbraken & Frisvad, ser. nov. MycoBank MB832585.



Etymology: Named after the type species of the series, Aspergillus coremiiformis.



Type: Aspergillus coremiiformis Bartoli & Maggi, Trans. Brit. Mycol. Soc. 71: 386. 1979.



Diagnosis: Phylogeny: Series Coremiiformes belongs to subgen. Circumdati, sect. Flavi and is sister to a clade containing series Bertholletiarum, Flavi, Kitamyces and Nomiarum (Fig. 12). Morphology & physiology: Colonies spreading; conidial colour en masse orange-brown; conidiophores biseriate; No growth at 37 °C, synnemata present; reverse on AFPA cream. Sexual morph unknown; sclerotia not observed in culture. Series description based on Frisvad et al. (2019).



Included species: Aspergillus coremiiformis, A. togoensis.



Extrolites: In common with species in other series in sect. Flavi, A. togoensis produces aflatoxin B₁, a bisiderin, paspaline, paspalinine, paxillin and sterigmatocystin, while the other species in the series (A. coremiiformis) is quite different and only produces unknown indole alkaloids (Frisvad et al. 2019).



Series Flavi Houbraken & Frisvad, ser. nov. MycoBank MB832989.



Etymology: Named after the type species of the series, Aspergillus flavus.



Type: Aspergillus flavus Link, Mag. Ges. Naturf. Freunde Berlin 3: 16. 1809.



Diagnosis: Phylogeny: Series Flavi belongs to subgen. Circumdati, sect. Flavi and is sister to ser. Kitamyces (Fig. 12). Morphology & physiology: Colonies spreading; conidial colour en masse mostly yellow-green, occasionally brown (A. oryzae), or brownish green (A. parasiticus); conidiophores uni- and/or biseriate; good growth at 37 °C, generally growth at 42 °C, except A. mottae and A. subflavus; reverse on AFPA orange, except in A. oryzae. Sexual morph generally not observed in culture, except in A. flavus and A. parasiticus, petromyces-type; heterothallic; sclerotia often present, black. Most species are primarily associated with nuts and oil-seeds, but will also grow on foods from domesticated plants such as cereals (e.g. maize) and dry fruits. Aspergillus oryzae is the domesticated form of A. flavus, and A. sojae of A. parasiticus; both are used in food fermentations. Series description based on Frisvad et al. (2019).



Included species: Aspergillus aflatoxiformans, A. arachidicola, A. austwickii, A. cerealis, A. flavus, A. krugeri, A. minisclerotigenes, A. mottae, A. novoparasiticus, A. oryzae, A. parasiticus, A. pipericola, A. sergii, A. sojae, A. subflavus, A. transmontanensis.



Extrolites: Most species produces aflatoxins (B and G types) (and precursors such as versicolorins and sterigmatocystins), aflatrems, aflavarins, aflavazols, aflaviniones, asparasones, asperfurans, aspergillic acids, aspergillomarasmins, aspirochlorins, chrysogines, citreoisocoumarins, cyclopiamides, cyclopiazonic acids (and the related speradins), ditryptophenalines, kojic acids, kojistatins, leporins, miyakamides (= oryzamides), 3-nitropropionic acid, parasitenone, parasiticolides, parasiticols, parasperones, penicillins, sporogens, and ustilaginoidins (Frisvad et al. 2019).



Series Kitamyces Houbraken & Frisvad, ser. nov. MycoBank MB832586.



Etymology: Named after G. Kita, who described A. tamarii, the type species of this series.



Type: Aspergillus tamarii Kita, Centralbl. Bakteriol. 2. Abth. 37: 433. 1913.



Diagnosis: Phylogeny: Series Kitamyces belongs to subgen. Circumdati, sect. Flavi and is sister to ser. Flavi (Fig. 12). Morphology & physiology: Colonies spreading; conidial colour en masse in shades of brown; conidiophores biseriate; good growth at 37 °C, no growth at 42 °C; reverse on AFPA dark brown. Sexual morph not observed in culture; sclerotia occasionally present, black. Series description based on Frisvad et al. (2019).



Included species: Aspergillus caelatus, A. pseudocaelatus, A. pseudotamarii, A. tamarii.



Series Leporum Houbraken & Frisvad, ser. nov. MycoBank MB832587.



Etymology: Named after the type species of the series, Aspergillus leporis.



Type: Aspergillus leporis States & M. Chr., Mycologia 58: 738. 1966.



Diagnosis: Phylogeny: Series Leporum belongs to subgen. Circumdati, sect. Flavi and is sister to a large clade containing series Alliacei, Bertholletiarum, Coremiiformes, Flavi, Kitamyces and Nomiarum. Morphology & physiology: Colonies spreading; conidial colour en masse yellow-green with a shade of beige, beige or olive; conidiophores biseriate; growth at 37 °C, no growth at 42 °C; reverse on AFPA cream. Sexual morph not observed in culture; sclerotia often present, large, black. Aspergillus leporis is dung-associated, while A. aspearensis and A. hancockii have been reported to be soil-borne. It is not known whether the two latter species are actually dung-associated. Series description based on Frisvad et al. (2019).



Included species: Aspergillus aspearensis, A. hancockii, A. leporis.



Extrolites: Aflavarins, aflavinines, antibiotic Y, clavatols, dehydroterrestric acid, eupenifeldin, fumitremorgins, hancockiamides, 7-hydroxytrichothecolone, kojic acid, leporines, leporizines, mevinolins, onychocins, paspalines, pseurotins, speradins (Frisvad et al. 2019). The aflavarins, aflavinines, paspalinines, pseurotins, speradins and kojic acid have been found in other series in sect. Flavi, but antibiotic Y, dehydroterrestric acid, eupenifeldin, fumitremorgins, hancockiamides, 7- hydroxytrichothecolone, leporizines, mevinolins and onychocins have only been found in ser. Leporum.



Series Nomiarum Houbraken & Frisvad, ser. nov. MycoBank MB832588.



Etymology: Named after the type species of the series, Aspergillus nomiae.



Type: Aspergillus nomiae Kurtzman et al., Antonie van Leeuwenhoek 53: 151. 1987.



Diagnosis: Phylogeny: Series Nomiarum belongs to subgen. Circumdati, sect. Flavi and is sister to a large clade containing series Alliacei, Bertholletiarum, Coremiiformes, Flavi, Kitamyces and Nomiarum (Fig. 12). Morphology & physiology: Colonies spreading; conidial colour en masse (dark) yellow-green; conidiophores biseriate; good growth at 37 °C, no growth at 42 °C; reverse on AFPA cream orange. Sexual morph generally not observed in culture, present in A. nomiae, petromyces-type, heterothallic; sclerotia often present, bullet-shaped, black. Species primarily associated to bees. Series description based on Frisvad et al. (2019).



Included species: Aspergillus luteovirescens, A. nomiae, A. pseudonomiae.



Extrolites: Aflatoxins (B and G type) (and precursors), altersolanols, anominine, aspernomine, aspergillic acids, chrysogines, kojic acid, miyakamides, paspaline, paspalinine, pseurotins, sporogens, and tenuazonic acid (Frisvad et al., 2018, Frisvad et al., 2019).



Notes on series in sect. Flavi: Using a multigene phylogenetic analysis, Frisvad et al. (2019) recognised eight clades in sect. Flavi. These clades are treated here as separate series. The majority of species belonging to the phylogenetically related series Flavi, Nomiarum and Kitamyces produce aflatoxin B and G. Non-aflatoxin producers are A. oryzae, A. sojae and A. subflavus in ser. Flavi, A. caelatus and A. tamarii in ser. Kitamyces, and A. pseudotamarii only produces aflatoxin B. Series Bertholletiarum can produce the aflatoxin precursor O-methylsterigmatocystin, but not aflatoxins. Series Coremiiformes is phylogenetically a sister to series Flavi, Kitamyces, Bertholletiarum (Fig. 12) and includes a species (A. togoensis) that produces aflatoxin B (Frisvad et al. 2019). Species in ser. Alliacei are able to produce ochratoxins, unlike species in other series of sect. Flavi. Other extrolites only found in sect. Circumdati ser. Alliacei are altersolanols, asperlicins, burnettiene, burnettramic acid, griseofulvin. Series Avenacei and Leporum are early branching clades in sect. Flavi (Fig. 7, Frisvad et al. 2019), but do not have the ability to produce aflatoxins or ochratoxins. The sole species in ser. Avenacei (A. avenaceus) does not produce kojic acid, an extrolite produced by the majority of species in sect. Flavi (Frisvad et al. 2019).



Section Flavipedes W. Gams et al., Adv. Pen. Asp. Syst.: 59. 1986 [1985]. MycoBank MB832506.



Type: Aspergillus flavipes (Bainier & Sartory) Thom & Church, Aspergilli: 155. 1926.



Description: See Hubka et al. (2015) (morphology, phylogeny), Kocsubé et al. (2016) (phylogeny).



Series Flavipedes Houbraken & Frisvad, ser. nov. MycoBank MB832990.



Etymology: Named after the type species of the series, Aspergillus flavipes.



Type: Aspergillus flavipes (Bainier & Sartory) Thom & Church, Aspergilli: 155. 1926.



Diagnosis: Phylogeny: Series Flavipedes belongs to subgen. Circumdati, sect. Flavipedes and is phylogenetically sister to series Olivimuriarum and Spelaei (Fig. 12). Morphology & physiology: Colonies growing moderately fast; conidial colour en masse white, pale brown, or yellow-brown; conidiophores biseriate; good growth at 37 °C, some species grow on CYA at 40 °C (e.g., A. ardalensis, A. neoflavipes, A. templicola (reported under A. mangaliensis), all grow on M40Y at 40 °C. Sexual morph generally not observed in culture, except in A. neoflavipes, fennellia-type; sclerotia not observed in culture. Series description based on Hubka et al. (2015).



Included species: Aspergillus ardalensis, A. capensis, A. flavipes, A. iizukae, A. micronesiensis, A. neoflavipes, A. suttoniae, A. templicola, A. urmiensis.



Extrolites: Aspochalasins, cytochalasins, flavipins, flaviphenalenones and geodins.



Series Neonivei Houbraken & Frisvad, ser. nov. MycoBank MB832589.



Etymology: Named after the type species of the series, Aspergillus neoniveus.



Type: Aspergillus neoniveus Samson et al., Stud. Mycol. 69: 53. 2011.



Diagnosis: Phylogeny: Series Neonivei belongs to subgen. Circumdati, sect. Flavipedes; the phylogenetic relationship of this series is unresolved (more information, see Notes section Flavipedes). Morphology & physiology: Colonies growing moderately fast; conidial colour en masse white; conidiophores biseriate. Sexual morph fennellia-type, yellow, orange-yellow. Series description based on Samson et al. (2011a) and Hubka et al. (2015).



Included species: Aspergillus neoniveus.



Extrolites: Aspochalamins, citreoviridin and paspalinine are produced by A. neoniveus, the sole species in the series (Samson et al. 2011a).



Series Olivimuriarum Houbraken & Frisvad, ser. nov. MycoBank MB835555.



Etymology: Named after the type species of the series, Aspergillus olivimuriae.



Type: Aspergillus olivimuriae S.W. Peterson & S. Crognale, Int. J. Syst. Evol. Microbiol. 69: 2901. 2019.



Diagnosis: Phylogeny: Series Olivimuriarum belongs to subgen. Circumdati, sect. Flavipedes and is phylogenetically sister to ser. Spelaei, though statistical support is lacking (Fig. 12); Bayesian analysis confidently shows that this series is sister to series Flavipedes and Spelaei (1.00 pp, data not shown). Morphology & physiology: Colonies growing moderately fast; conidial colour en masse avellaneous; conidiophores biseriate; moderate growth at 37 °C, no growth on CYA at 40 °C. Sexual morph unknown; sclerotia not observed in culture. Series description based onet al. Crognale et al. (2019).



Included species: Aspergillus olivimuriae.



Extrolites: Extrolite production by the sole species in the series has not been performed.



Series Spelaei Houbraken & Frisvad, ser. nov. MycoBank MB832590.



Etymology: Named after the type species of the series, Aspergillus spelaeus.



Type: Aspergillus spelaeus A. Nováková et al., Mycologia 107: 194. 2015.



Diagnosis: Phylogeny: Series Spelaei belongs to subgen. Circumdati, sect. Flavipedes and is phylogenetically sister to series Olivimuriarum and Flavipedes (Fig. 12). Morphology & physiology: Colonies growing moderately fast; conidial colour en masse white, pale brown, or yellow-brown; conidiophores biseriate; no or moderate growth at 37 °C, generally no growth on CYA and M40Y at 40 °C. Sexual morph unknown; sclerotia not observed in culture. Series description based on Hubka et al. (2015).



Included species: Aspergillus luppiae, A. movilensis, A. polyporicola, A. spelaeus.



Extrolites: Aspochalasins and curvularins can be produced by ser. Spelaei taxa.



Notes on series in sect. Flavipedes: Hubka et al. (2015) recognised two main clades in sect. Flavipedes and named them the A. flavipedes- and A. spelaeus-clade. These results are confirmed here and the series names Flavipedes and Spelaei are introduced for those phylogenetically related clades (Fig. 12). These two series in sect. Flavipedes can be differentiated based on growth rates on CYA incubated at 37 and 40 °C. Furthermore, extrolites from species in ser. Flavipedes include aspochalasins, cytochalasins, flavipins, geodins and flaviphenalenones and ser. Spelaei species can produce aspochalasins and curvularins. The phylogenetic position of ser. Neonivei is uncertain. Our phylogenetic analysis based on BenA, CaM and RPB2 sequences (Fig. 12) shows that this series is basal in sections Terrei and Flavipedes; however, this relationship lacks bootstrap support, but is fully supported in the Bayesian analysis. Our nine-gene phylogeny (Supplementary Fig. S1) positions ser. Neonivei in sect. Flavipedes (89 % BS). In the past, A. neoniveus (syn. Fennellia nivea) was classified in sections Janorum, Terrei and Flavipedes (Peterson, 2008, Peterson et al., 2008, Hubka et al., 2015, Jurjević et al., 2015), but never with high statistical support. We decided to include ser. Neonivei in sect. Flavipedes based on the result of our 9-gene phylogeny and future research using genome sequence data will probably resolve the exact position of this series.



Section Janorum [as “Jani”] Hubka et al., Mycologia 107: 197. 2015. MycoBank MB832532.



Type: Aspergillus janus Raper & Thom, Mycologia 36: 556. 1944.



Description: Phylogeny: Section Janorum belongs to subgen. Circumdati. Morphology & physiology: Colonies growing restricted; conidial colour en masse in shades of green and white; conidiophores variable, 1) tall, white sporulating, biseriate conidiophores with pyriform to clavate vesicles, producing smooth, uncoloured conidia; 2) lower, green sporulating, biseriate conidiophores with commonly pyriform vesicles producing green, echinulate conidia; 3) micro- to semimacronematous conidiophores producing globose, elliptical or clavate conidia, that are commonly truncate; no (A. brevijanus) or restricted (A. janus) growth at 37 °C, no growth at 40 °C. Sexual morph unknown. Also see Hubka et al. (2015) (morphology, phylogeny), Kocsubé et al. (2016) (phylogeny).



Included species: Aspergillus brevijanus, A. janus, A. trisporus, A. yunnanensis.



Extrolites: Asperphenamate, brevicompanins and janoxepin are found in sect. Janorum.



Notes: Because no subdivision of sect. Janorum is proposed, ser. Janorum is only informally introduced here. Section Janorum is phylogenetically related to sections Terrei and Flavipedes (Fig. 2). Species belonging to this section produce three types of conidiophores and conidia, and colonies have green and white sectors making them distinctive (Hubka et al. 2015).



Section Nigri W. Gams et al., Adv. Pen. Asp. Syst.: 60. 1986 [1985]. MycoBank MB832511.



Type: Aspergillus niger Tiegh., Ann. Sci. Nat., Bot., ser. 5, 8: 240. 1867; nom. cons. (Kozakiewicz et al. 1992).



Description: See Gams et al. (1985) (morphology), Samson et al., 2007b, Varga et al., 2011 (morphology, phylogeny), Kocsubé et al. (2016) (phylogeny), Vesth et al., 2018, Steenwyk et al., 2019 (genome analysis).



Notes: The phylogenomic analysis of Steenwyk et al. (2019) shows that section Nigri does not belong to subgen. Circumdati and that it is more closely related to subgen. Nidulantes. The species in subgen. Nidulantes are phenotypically distinct from sect. Nigri species, indicating that this section represents a separate subgenus. Based on phenotypic and extrolite data, and our phylogenetic analysis (Fig. 1, Fig. 2), we decided to maintain sect. Nigri in subgen. Circumdati until more (genome) data supporting the analysis of Steenwyk et al. (2019) becomes available.



Series Carbonarii Houbraken & Frisvad, ser. nov. MycoBank MB832591.



Etymology: Named after the type species of the series, Aspergillus carbonarius.



Type: Aspergillus carbonarius (Bainier) Thom, J. Agric. Res. 7: 12. 1916.



Diagnosis: Phylogeny: Series Carbonarii belongs to subgen. Circumdati, sect. Nigri and is phylogenetically sister to ser. Nigri (Fig. 13). Morphology & physiology: Colonies spreading; conidial colour en masse black or blackish brown; conidiophores biseriate; generally weak or no growth at 37 °C (except A. ibericus); conidia rough-walled, large, 6–9 μm. Sexual morph generally not produced in culture, except in A. sclerotiicarbonarius, saitoa-type, heterothallic, orange to red-brown. Series description based on Samson et al. (2007b) and Varga et al. (2011).



Fig. 13.

Combined phylogeny for BenA, CaM and RPB2 data sets showing the phylogenetic relation of species and series within Aspergillus sect. Nigri. The BI posterior probability (pp) values and bootstrap percentages of the maximum likelihood (ML) analysis are presented at the nodes; fully supported branches are thickened. Values less than 70 % bootstrap support (ML) or less than 0.95 posterior probability (Bayesian analysis) are indicated with a hyphen or not shown. The bar indicates the number of substitutions per site. The phylogram is rooted with Hamigera avellanea and Penicillium expansum.

Included species: Aspergillus carbonarius, A. ibericus, A. sclerotiicarbonarius, A. sclerotioniger.



Extrolites: Citric acid, oxalic acid, gluconic acid, atromentins, ochratoxins, funalenone, naphtho-γ-pyrones and pyranonigrins are shared by two or more species in ser. Carbonarii.



Series Heteromorphi Houbraken & Frisvad, ser. nov. MycoBank MB832592.



Etymology: Named after the type species of the series, Aspergillus heteromorphus.



Type: Aspergillus heteromorphus Bat. & H. Maia, Anais Soc. Biol. Pernambuco 15: 200. 1957.



Diagnosis: Phylogeny: Series Heteromorphi belongs to subgen. Circumdati, sect. Nigri and is phylogenetically sister to a clade containing series Carbonarii and Nigri (Fig. 13). Morphology & physiology: Colonies spreading; conidial colour en masse black or blackish brown; conidiophores biseriate; no growth at 37 °C; conidia 3–5 μm. Sexual morph unknown; sclerotia can be induced. Series description based on Samson et al. (2007b) and Varga et al. (2011).



Included species: Aspergillus ellipticus, A. heteromorphus.



Extrolites: Atromentins, austdiol, candidusins, terphenyllins, and xanthoascin.



Series Homomorphi Houbraken & Frisvad, ser. nov. MycoBank MB832593.



Etymology: Named after the type species of the series, Aspergillus homomorphus.



Type: Aspergillus homomorphus Steiman et al. ex Samson & Frisvad, Stud. Mycol. 50: 58. 2004.



Diagnosis: Phylogeny: Series Homomorphi belongs to subgen. Circumdati, sect. Nigri and is phylogenetically sister to ser. Japonici (Fig. 13). Morphology & physiology: Colonies spreading; conidial colour en masse black or blackish brown; conidiophores biseriate; growth at 37 °C, no growth at 40 °C; conidia 5–7 μm, with spiny ornamentation. Sexual morph unknown; sclerotia not observed in culture. Series description based on Steiman et al., 1995, Samson et al., 2007b and Varga et al. (2011).



Included species: Aspergillus homomorphus.



Extrolites: Asperflavin, atromentins, decaturins, dehydrocarolic acid, homomorphosins, styrylpyrones.



Series Japonici Houbraken & Frisvad, ser. nov. MycoBank MB834295.



Etymology: Named after the type species of the series, Aspergillus japonicus.



Type: Aspergillus japonicus Saito, Bot. Mag. (Tokyo) 20: 61. 1906.



Diagnosis: Phylogeny: Series Japonici belongs to subgen. Circumdati, sect. Nigri and is phylogenetically sister to ser. Homomorphi (Fig. 13). Morphology & physiology: Colonies spreading; conidial colour en masse black or blackish brown; conidiophores uniseriate; no growth at 37 °C; conidia 3–6 μm. Sexual morph generally not produced in culture, except in a species described as “Saitoa japonica” (Rajendran & Muthappa 1980); sclerotia often present. Series description based on Samson et al., 2007b, Varga et al., 2011 and Hubka & Kolarik (2012).



Included species: Aspergillus aculeatinus, A. aculeatus, A. assiutensis (accepted species, unpubl. data, XC Wang), A. brunneoviolaceus, A. floridensis, A. indologenus, A. japonicus, A. labruscus, A. saccharolyticus, A. serratalhadensis, A. trinidadensis, A. uvarum.



Extrolites: Aculene A, aflavinins, asperflavin, aspergillimide, calbistrin C, emodin, neopyranopnigrin, neoxaline, okaramin X, pre-aurantiamin, secalonic acid D.



Series Nigri Houbraken & Frisvad, ser. nov. MycoBank MB832991.



Etymology: Named after the type species of the series, Aspergillus niger.



Type: Aspergillus niger Tiegh., Ann. Sci. Nat., Bot., ser. 5, 8: 240. 1867; nom. cons. (Kozakiewicz et al. 1992).



Diagnosis: Phylogeny: Series Nigri belongs to subgen. Circumdati, sect. Nigri and is phylogenetically sister to ser. Carbonarii (Fig. 13). Morphology & physiology: Colonies spreading; conidial colour en masse black or blackish brown; conidiophores biseriate; good growth at 37 and 40 °C; conidia (2.5–)3–5 μm. Sexual morph generally not produced in culture, except in A. tubingensis, saitoa-type, heterothallic (Horn et al. 2013); sclerotia produced in most species. Series description based on Samson et al. (2007b) and Varga et al. (2011).



Included species: Aspergillus brasiliensis, A. costaricensis, A. eucalypticola, A. luchuensis, A. neoniger, A. niger, A. piperis, A. tubingensis, A. vadensis, A. welwitschiae.



Extrolites: Citric acid, oxalic acid, gluconic acid, tensyuic acids, atromentins, ochratoxins, funalenone, fumonisins, kotanins, yanuthones, naphtho-γ-pyrones, tensidols, malformins, nigragillins, pyranonigrins, asperazines, aflavinins (only in sclerotia) are shared by two or more species in ser. Nigri.



Notes on series in sect. Nigri: Frisvad et al. (2007) suggested a series (and subseries) classification for section Nigri using morphological, chemical and physiological features. Their suggested provisional series classification fits with our suggested phylogenetic-based classification. Fig. 13 shows the phylogenetic relationship among the species of section Nigri based on partial BenA, CaM and RPB2 gene sequencing. These relationships are similar to those presented in Vesth et al. (2018) using genome sequence data. In their manuscript, they distinguished the A. niger- and the A. tubingensis-clade. In contrast, Varga et al. (2011) treated these two clades as one, named the A. niger-clade. We follow Varga et al. (2011) in our concept of ser. Nigri, because species of this series share the ability to grow well at 37 and 40 °C and form biseriate conidiophores and similar-sized conidia measuring (2.5–)3–5 μm. Furthermore, the extrolites shared by two or more species in series Nigri are citric acid, oxalic acid, gluconic acid, tensyuic acids, atromentins, ochratoxins, funalenone, fumonisins, kotanins, yanuthones, naphtho-γ-pyrones, tensidols, malformins, nigragillins, pyranonigrins, asperazines, aflavinins (only in sclerotia) (Nielsen et al., 2009, Varga et al., 2011, Frisvad et al., 2018, Vesth et al., 2018). Series Carbonarii is characterised by a generally weak or no growth at 37 °C (except A. ibericus) and production of large conidia measuring 6–9 μm. Extrolites shared by two or more species in ser. Carbonarii include citric acid, oxalic acid, gluconic acid, atromentins, ochratoxins, funalenone, naphtho-γ-pyrones and pyranonigrins (Nielsen et al., 2009, Varga et al., 2011). Series Japonici is a sister series of ser. Homomorphi. Series Homomorphi include species that have biseriate conidiophores and produce the extrolites homomorphosins and decaturins, while ser. Japonici species have uniseriate conidiophores and produce aculenes, asperparalines, calbistrins, neoxalines, okaramins, pre-aurantiamine, and/or secalonic acids. The phylogenetic relationship of A. labruscus and A. saccharolyticus with the other species of ser. Japonici is unclear (Fig. 13). In contrast to our results, Fungaro et al. (2017) classified A. labruscus, A. homomorphus and A. saccharolyticus in the A. homomorphus-clade and a similar result was shown in Crous et al. (2018b, Fungal Planet 720). Based on genome sequence analysis, A. saccharolyticus is sister to the other ser. Japonici species, confirming the result of our 3-gene phylogeny. Unfortunately, A. labruscus and A. serratalhadensis were not included in the genome study of Vesth et al. (2018) and genome sequencing of these species might reveal the correct classification of these species in the future.



Section Petersoniorum [as “Petersonii”] Jurjević & Hubka, Pl. Syst. Evol. 301: 2449. 2015. MycoBank MB832533.



Type: Aspergillus petersonii Jurjević & Hubka, Pl. Syst. Evol. 301: 2454. 2015.



Description: Phylogeny: Section Petersoniorum belongs to subgen. Circumdati and the phylogenetic relationship with other Aspergillus sections needs to be resolved (see Notes below). Morphology & physiology: Colonies restricted; conidial colour en masse in shades of green; conidiophores biseriate; no growth 40 °C. Sexual morph unknown; sclerotia produced in most species (except A. asclerogenus), globose to ellipsoidal, pale yellow to brown. Also see Jurjević et al. (2015) (morphology, phylogeny).



Included species: Aspergillus arenarioides, A. asclerogenus, A. petersonii, A. peyronelii.



Notes on sect. Petersoniorum: Because no subdivision of sect. Petersoniorum is proposed, ser. Petersoniorum is only informally introduced here. In Fig. 12, this section is an early diverging clade in subgen. Circumdati; however, a more thorough analysis places this section most close to sect. Candidi (Fig. 2) confirming the results of Jurjević et al. (2015). Sections Petersoniorum and Candidi can be differentiated by their differences in conidial and sclerotial colour (for more details, see sect. Candidi).



Section Robusti Jurjević & Hubka, Pl. Syst. Evol. 301: 2460. 2015. MycoBank MB814443.



Type: Aspergillus robustus M. Chr. & Raper, Mycologia 70: 200. 1978.



Description: Phylogeny: Section Robusti belongs to subgen. Circumdati and is phylogenetically sister to sections Tannerorum and Circumdati. Morphology & physiology: Colonies restricted; conidial colour en masse in shades of yellow; conidiophores biseriate; no growth 37 °C. Sexual morph unknown; sclerotia produced, black (Christensen & Raper 1978). See also: Jurjević et al. (2015) (morphology, phylogeny).



Included species: Aspergillus robustus.



Notes: Because no subdivision of sect. Robusti is proposed, ser. Robusti is only informally introduced here. Section Robusti is a single species section. This section is phylogenetically sister to sections Tannerorum and Circumdati (Fig. 2, Fig. 12). It differs from both sections by the production of black coloured sclerotia and phototropic conidiophores (Visagie et al. 2014c).



Section Tannerorum [as “Tanneri”] Jurjević & Hubka, Pl. Syst. Evol. 301: 2460. 2015. MycoBank MB832534.



Type: Aspergillus tanneri Kwon-Chung et al., J. Clin. Microbiol. 50: 3312. 2012.



Description: Phylogeny: Section Tannerorum belongs to subgen. Circumdati and is sister to sect. Circumdati. Morphology & physiology: Colonies restricted; sporulation sparse; conidiophores biseriate; good growth 37 °C. Sexual morph unknown; sclerotia not observed in culture. Also see Jurjević et al. (2015) (morphology, phylogeny).



Included species: Aspergillus tanneri.



Extrolites: No extrolites are reported for A. tanneri, the sole species in this section.



Notes: No subdivision of sect. Tannerorum is proposed and ser. Tannerorum is only informally introduced here (Table 4). Section Tannerorum is a single species section. This section is phylogenetically sister to sect. Circumdati (Fig. 2, Fig. 12). It differs from this series by its small pyriform vesicles, lack of sclerotia, very poor sporulation, uncoloured reverse of colonies without production of soluble pigments and better growth at 37 °C than at 25 °C (Jurjević et al. 2015).



Section Terrei W. Gams et al., Adv. Pen. Asp. Syst.: 59. 1986 [1985]. MycoBank MB832505.



Type: Aspergillus terreus Thom, Amer. J. Bot. 5: 85. 1918.



Description: See Gams et al. (1985) (morphology), Samson et al. (2011a) (morphology, phylogeny), Kocsubé et al. (2016) (phylogeny).



Series Ambigui Houbraken & Frisvad, ser. nov. MycoBank MB832594.



Etymology: Named after the type species of the series, Aspergillus ambiguus.



Type: Aspergillus ambiguus Sappa, Allionia 2: 254. 1955.



Diagnosis: Phylogeny: Series Ambigui belongs to subgen. Circumdati, sect. Terrei and is phylogenetically sister to series Nivei and Terrei. Morphology & physiology: Colonies restricted; conidial colour en masse white, cream, dull yellow or grey-green; conidiophores biseriate. Sexual morph unknown. Series description based on Raper & Fennell (1965).



Included species: Aspergillus ambiguus, A. microcysticus.



Extrolites: Butryolactone, terrequinone A.



Series Nivei Houbraken & Frisvad, ser. nov. MycoBank MB832595.



Etymology: Named after the type species of the series, Aspergillus niveus.



Type: Aspergillus niveus Blochwitz, Ann. Mycol. 27: 205. 1929.



Diagnosis: Phylogeny: Series Nivei belongs to subgen. Circumdati, sect. Terrei and is phylogenetically most closely related to ser. Terrei. Morphology & physiology: Colonies growing moderately fast; conidial colour en masse white, vinaceous fawn, blue-green; conidiophores biseriate; moderate growth at 37 °C. Sexual morph unknown. Series description based on Raper & Fennell (1965) and Samson et al. (2011a).



Included species: Aspergillus allahabadii, A. bicephalus, A. carneus, A. iranicus, A. neoindicus, A. niveus.



Extrolites: Aszonalenins, citrinins are shared by ser. Nivei species.



Series Terrei Houbraken & Frisvad, ser. nov. MycoBank MB832992.



Etymology: Named after the type species of the series, Aspergillus terreus.



Type: Aspergillus terreus Thom, Amer. J. Bot. 5: 85. 1918.



Diagnosis: Phylogeny: Series Terrei belongs to subgen. Circumdati, sect. Terrei and is phylogenetically most closely related to ser. Nivei. Morphology & physiology: Colonies spreading; conidial colour en masse (light) olive-brown; conidiophores biseriate; good growth at 37 °C. Sexual morph generally not produced in culture, except in A. terreus, heterothallic, fennellia-type. Series description based on Samson et al. (2011a).



Included species: Aspergillus alabamensis, A. aureoterreus, A. citrinoterreus, A. floccosus, A. heldtiae, A. hortae, A. neoafricanus, A. pseudoterreus, A. terreus.



Extrolites: Series Terrei extrolites include acetylaranotins, ardeemins, aspergillamides, aspergillicins, aspulvinones, asterriquinones, aszonalenins, butyrolactones, citreoviridins, citrinins, cytochalasins, geodins, gregatins, mevinolins, terrecyclic acids, terreic acid, terreins, terremides, terrequinones, terretonins and territrems (Samson et al. 2011a).



Notes on series in sect. Terrei: Three series are introduced in sect. Terrei: series Ambigui, Nivei and Terrei. Series Terrei and Nivei are sister series and the species in those series differ in their conidial colour. Species in ser. Terrei generally produce conidia in brown shades, while ser. Nivei are in shades of yellow, vinaceous fawn or white. The two species of ser. Ambigui grow slower than the taxa of series Terrei and Nivei.



Aspergillus subgen. Cremei Samson et al., Stud. Mycol. 85: 210. 2016. MycoBank MB819182.



Type: Aspergillus cremeus Kwon-Chung & Fennell, Gen. Aspergillus: 418. 1965.



Description: See Kocsubé et al. (2016) (morphology, phylogeny).



Section Cremei W. Gams et al., Adv. Pen. Asp. Syst.: 61. 1986 [1985]. MycoBank MB832513.



Type: Aspergillus cremeoflavus Samson & W. Gams, Adv. Pen. Asp. Syst.: 37. 1986 [1985]. MycoBank MB114701 (= Aspergillus cremeus).



Description: See Gams et al. (1985) (morphology, and partial sect. Wentii, see Notes below), Kocsubé et al. (2016), Fig. 2, this study (phylogeny).



Series Arxiorum Houbraken & Frisvad, ser. nov. MycoBank MB833044.



Etymology: Named after the type species of this series, A. arxii.



Type: Aspergillus arxii (Fort & Guarro) Houbraken et al., Stud. Mycol. 78: 154. 2014.



Diagnosis: Phylogeny: Series Arxiorum belongs to sect. Cremei, subgen. Cremei and is phylogenetically sister to a clade containing series Brunneouniseriati, Cremei and Wentiorum, though without statistical support (Fig. 14). Morphology & physiology: Colonies restricted; conidiophores and conidia not observed in culture. Sexual morph chaetosartorya-type, homothallic; ascospores hyaline, ellipsoidal, with two equatorial ridges, convex surfaces finely rugose. Series description based on Fort & Guarro (1984).



Fig. 14.

Combined phylogeny for BenA, CaM and RPB2 data sets showing the phylogenetic relation of species and series within Aspergillus subgen. Cremei. The BI posterior probability (pp) values and bootstrap percentages of the maximum likelihood (ML) analysis are presented at the nodes; fully supported branches are thickened. Values less than 70 % bootstrap support (ML) or less than 0.95 posterior probability (Bayesian analysis) are indicated with a hyphen or not shown. The bar indicates the number of substitutions per site. The phylogram is rooted with Hamigera avellanea and Penicillium expansum.

Included species: Aspergillus arxii.



Extrolites: No known extrolites have been found in Aspergillus arxii.



Series Brunneouniseriati Houbraken & Frisvad, ser. nov. MycoBank MB833045.



Etymology: Named after the type species of this series, A. brunneouniseriatus.



Type: Aspergillus brunneouniseriatus Suj. Singh & B.K. Bakshi, Trans. Brit. Mycol. Soc. 44: 160. 1961.



Diagnosis: Phylogeny: Series Brunneouniseriati belongs to sect. Cremei, subgen. Cremei and is phylogenetically sister to a clade containing series Cremei and Wentiorum, though with moderate (BS = 70 %) or poor (pp < 0.95) statistical support (Fig. 14). Morphology & physiology: Colonies growing rapidly; conidia en masse grey, (dark) olive-brown or grey-green; conidiophores uniseriate, stipes hyaline, smooth. Sexual morph unknown. Series description based on Raper & Fennell (1965) and Crous et al. (2015).



Included species: Aspergillus brunneouniseriatus, A. citocrescens.



Extrolites: No known extrolites have been found in series Brunneouniseriati.



Series Cremei Houbraken & Frisvad, ser. nov. MycoBank MB833046.



Etymology: Named after the type species of this series, A. cremeus.



Type: Aspergillus cremeus Kwon-Chung & Fennell, Gen. Aspergillus: 418. 1965.



Diagnosis: Phylogeny: Series Cremei belongs to sect. Cremei, subgen. Cremei and is phylogenetically sister to ser. Wentiorum (Fig. 14). Morphology & physiology: Colonies varying from restricted to spreading, more rapid growth on agar media with reduced water activity; conidia en masse in shades of green; conidiophores biseriate (A. cremeus, A. gorakhpurensis), uni- and biseriate (A. stromatoides) or strictly uniseriate (A. itaconicus), stipes hyaline, smooth, long. Sexual morph unknown (A. gorakhpurensis, A. itaconicus, A. stromatoides) or observed in culture (A. chaetosartoryae, A. cremeus), ascomata consisting of several layers of thick-walled hyphae, the outer layer becoming dematiaceous, homothallic; ascospores with prominent equatorial ridges, convex surface with spines. Series description based on Raper and Fennell, 1965, Kamal and Bhargava, 1969 and Wiley & Simmons (1973).



Included species: Aspergillus chaetosartoryae, A. cremeus, A. gorakhpurensis, A. itaconicus, A. stromatoides.



Extrolites: Only A. itaconicus has been examined for secondary metabolites according to the literature. This species can produce chrysogine (reported here), itaconic acid (Kinoshita, 1932, Steiger et al., 2013), itaconitin (Nakajima et al. 1964) and sorbicillins (reported here).



Series Inflati (Stolk & Samson) Houbraken & Frisvad, comb. nov. MycoBank MB833047.

Basionym: Penicillium ser. Inflata Stolk & Samson, Adv. Pen. Asp. Syst.: 174. 1986 [1985].



Type: Penicillium inflatum Stolk & Malla, Persoonia 6: 197. 1971. (syn. Aspergillus inflatus).



Diagnosis: Phylogeny: Series Inflati belongs to sect. Cremei, subgen. Cremei and is phylogenetically sister to all other series in sect. Cremei. Morphology & physiology: Colonies restricted (A. inflatus) or spreading (A. koreanus), conidia en masse grey-green, greyish olive or grey-brown; conidiophores bi- or triseriate, or penicillium-like and biverticillate divaricate branched, stipe hyaline or pale reddish, smooth or finely roughened. Sexual morph unknown. Series description based on (Stolk and Malla, 1971, Bissett and Widden, 1984, Hyde et al., 2016).



Extrolites: Aspergillus inflatus and a putative new species tentatively named A. oregonensis (CBS 576.95A&B) in ser. Inflati can produce sterigmatocystin (Rank et al. 2011). Otherwise sterigmatocystin (and aflatoxins) has only been found in the subgenera Circumdati (sect. Flavi) and Nidulantes in the genus Aspergillus (Chen et al., 2016a, Hubka et al., 2016a, Frisvad et al., 2019).



Included species: Aspergillus inflatus, A. koreanus, A. tardus.



Series Pulvini Houbraken & Frisvad, ser. nov. MycoBank MB833048.



Etymology: Named after the type species of this series, A. pulvinus.



Type: Aspergillus pulvinus Kwon-Chung & Fennell, Gen. Aspergillus: 455. 1965.



Diagnosis: Phylogeny: Series Pulvini belongs to sect. Cremei, subgen. Cremei and is phylogenetically sister to a clade containing series Arxiorum, Brunneouniseriati, Cremei and Wentiorum, though without statistical support (Fig. 14). Morphology & physiology: Colonies spreading, conidia en masse blue-green; conidiophores biseriate, stipes brownish pigmented, roughened, long, thick-walled. Sexual morph unknown. Series description based on Raper & Fennell (1965).



Included species: Aspergillus pulvinus.



Extrolites: No known extrolites have been found in Aspergillus pulvinus.



Series Wentiorum Houbraken & Frisvad, ser. nov. MycoBank MB833049.



Etymology: Named after the type species of this series, A. wentii.



Type: Aspergillus wentii Wehmer, Centralbl. Bakteriol., 2. Abth., 2: 149. 1896.



Diagnosis: Phylogeny: Series Wentiorum belongs to sect. Cremei, subgen. Cremei and is phylogenetically sister to ser. Cremei. Morphology & physiology: Colonies growing restrictedly on MEA and CYA, growing moderately or rapidly on agar media with reduced water activity; conidia en masse in shades of yellow-brown, olive-brown or greyish yellow; conidiophores biseriate, sometimes uniseriate (A. flaschentraegeri), stipes hyaline, smooth or at most slightly roughened, often long and thick-walled; no growth at 37 °C. Sexual morph unknown (A. europaeus, A. dimorphicus, A. flaschentraegeri, A. wentii) or present (A. chrysellus), chaetosartorya-type, homothallic; ascospores hyaline, lenticular, with two equatorial ridges, convex surfaces with spines. Series description based on Raper and Fennell, 1965, Tuthill and Christensen, 1986 and Hubka et al. (2016b).



Included species: Aspergillus chrysellus, A. dimorphicus, A. europaeus, A. flaschentraegeri, A. wentii.



Extrolites: All species in ser. Wentiorum produce asperflavin, emodin, physcion, emodin bianthrone, physcion bianthrone (and other bianthrons), sulochrin and other sulochrins, and wentilacton A and B, except A. flaschentraegeri which only produces asperflavin, physcion and physcion bisanthron. These and many related extrolites have been reported from A. wentii and A. europaeus (Wells et al., 1975, Assante et al., 1979, Assante et al., 1980, Dorner et al., 1980, Xu et al., 2015, Hubka et al., 2016b, Du et al., 2018, Li et al., 2018b, Form et al., 2019). The metabolites have not been detected yet in any species in the other series in subgen. Cremei.



Notes on sect. Cremei and included series: Raper & Fennell (1965) introduced the A. wentii and the A. cremeus group and later Gams et al. (1985) formally introduced these as sections Wentii and Cremei. All species classified by Raper & Fennell (1965) in sect. Cremei (A. cremeus, A. chrysellus, A. flaschentraegeri, A. itaconicus, A. stromatoides) are still here accepted in this section. Phylogenetic data demonstrated that sect. Wentii was superfluous (Peterson, 1995, Peterson, 2008). Aspergillus wentii is included in sect. Cremei and the other members of the A. wentii group (fide Raper & Fennell 1965) belong to sect. Flavi: A. terricola (= A. tamarii), A. terricola var. americana (= A. parasiticus), A. terricola var. indicus (= A. tamarii), A. thomii (= A. flavus) (Frisvad et al. 2019). Mainly based on molecular data, sect. Cremei expanded to 17 species, which are classified in six series (Arxiorum, Brunneouniseriati, Cremei, Inflati, Pulvini, Wentiorum). Series Inflati includes three species and two of those produce penicillium-like conidiophores. Aspergillus inflatus was originally described in Penicillium. Another species of this series, A. tardus, was assigned to the A. versicolor group (Bissett & Widden 1984), but the original description mentioned the non-synchronously production of metulae, a character frequently observed in Penicillium and not common in Aspergillus. Series Pulvini includes a species that produces blue-green coloured conidia, a unique feature in sect. Cremei. Series Cremei and Wentiorum are phylogenetically and phenotypically related. The conidia in ser. Cremei are more often in shades of green, while those of ser. Wentiorum are more often in shades of yellow-brown. Both series contain species that reproduce sexually. The sole taxon classified in series Arxiorum also reproduces sexually, but an asexual morph is not described. Series Brunneouniseriati includes two species that both have uniseriate conidial heads. Uniseriate heads are also produced by two species in other series: A. flaschentraegeri (ser. Wentiorum) and A. itaconicus (series Cremei). Regarding extrolites the six series in subgenus and section Cremei are remarkably different.



Aspergillus subgen. Fumigati W. Gams, M. Chr., Onions, Pitt & Samson, Adv. Pen. Asp. Syst.: 56. 1986 [1985]. MycoBank MB832495.



Type: Aspergillus fumigatus Fresen., Beitr. Mykol. 3: 81. 1863.



Description: See Gams et al. (1985) (morphology); Samson et al. (2007a) (morphology, phylogeny); Peterson et al., 2008, Kocsubé et al., 2016, Fig. 1, this study (phylogeny).



Section Clavati W. Gams et al., Adv. Pen. Asp. Syst.: 57. 1986 [1985]. MycoBank MB832500.



Type: Aspergillus clavatus Desm., Ann. Sci. Nat., Bot., ser. 2, 2: 71. 1834.



Description: Phylogeny: Series Clavati belongs to subgen. Fumigati. Morphology & physiology: Colonies growing restrictedly or moderately rapid; conidial colour en masse blue-green; conidiophores uniseriate, often with clavate vesicles; restricted growth at 37 °C, except A. acanthosporus (no growth). Sexual morph not observed in culture, or present, neosartorya-type, homothallic (A. acanthosporus) or heterothallic (A. clavatus). Section description based on Varga et al. (2007a) and Ojeda-López et al. (2018).



Included species: Aspergillus acanthosporus, A. clavatonanicus, A. clavatus, A. giganteus, A. longivesica, A. posadasensis, A. rhizopodus, A. seifertii∗ [∗ not included in Fig. 15; for more details, see Visagie & Houbraken (2020)].



Fig. 15.

Combined phylogeny for BenA, CaM and RPB2 data sets showing the phylogenetic relation of species, series and sections within Aspergillus subgen. Fumigati. The BI posterior probability (pp) values and bootstrap percentages of the maximum likelihood (ML) analysis are presented at the nodes; fully supported branches are thickened. Values less than 70 % bootstrap support (ML) or less than 0.95 posterior probability (Bayesian analysis) are indicated with a hyphen or not shown. The bar indicates the number of substitutions per site. The phylogram is rooted with Hamigera avellanea.

Extrolites: At least six species (no data for A. posadasensis and A. seifertii) in the section produce ribotoxins (Varga & Samson 2008) and tryptoquivalines / tryptoquivalones (Varga et al. 2007a). Kotaninins are produced by four of six species (A. acanthosporus, A. clavatonanicus, A. clavatus, A. rhizopodus), patulin by three (A. clavatus, A. giganteus, A. longivesica) and antafumicins by four (A. clavatonanicus, A. clavatus, A. giganteus, A. longivesica). Pyripyropens are produced by two species (A. longivesica and A. giganteus). Ribotoxins are shared with several species in sect. Fumigati, including A. fischeri, A. fumigatus, A. neoglaber and A viridinutans (Martínez-Ruiz et al., 1999, Olombrada et al., 2014), while the isolate of A. restrictus producing restrictocin and regulin was an A. fumigatus (NRRL 3050 = ATCC 34475). Except for a claim that A. oryzae can produce ribotoxins (Machida et al. 2005), producers of these small insecticidal and antifungal proteins (Olombrada et al., 2014, Citores et al., 2018) are concentrated in subgen. Fumigati, sect. Clavati and ser. Fumigati.



Notes: No subdivision of sect. Clavati is proposed, and ser. Clavati is only informally introduced here (Table 4).



Section Cervini W. Gams et al., Adv. Pen. Asp. Syst.: 56. 1986 [1985]. MycoBank MB832497.



Type: Aspergillus cervinus Massee, Bull. Misc. Inform. Kew 1914: 158. 1914.



Description: See Gams et al. (1985) (morphology), Kocsubé et al. (2016) (phylogeny), Chen et al. (2016c) (morphology, phylogeny).



Series Acidohumorum Houbraken & Frisvad, ser. nov. MycoBank MB832596.



Etymology: Named after the type species of the series, Aspergillus acidohumus.



Type: Aspergillus acidohumus A.J. Chen et al., Stud. Mycol. 85: 71. 2016.



Diagnosis: Phylogeny: Series Acidohumorum belongs to sect. Cervini, subgen. Fumigati. Morphology & physiology: Colonies restricted; conidial colour en masse dark fawn; conidiophores uniseriate; no growth at 37 °C. Sexual morph unknown. Series description based on Chen et al. (2016c).



Included species: Aspergillus acidohumus.



Extrolites: No extrolites have been found in the species in ser. Acidohumorum.



Series Cervini Houbraken & Frisvad, ser. nov. MycoBank MB832993.



Etymology: Named after the type species of the series, Aspergillus cervinus.



Type: Aspergillus cervinus Massee, Bull. Misc. Inform. Kew 1914: 158. 1914.



Diagnosis: Phylogeny: Series Cervini belongs to sect. Cervini, subgen. Fumigati. Morphology & physiology: Colonies growing moderately fast; conidial colour en masse fawn; conidiophores uniseriate; generally no or very poor growth at 37 °C. Sexual morph unknown. Series description based on Chen et al. (2016c).



Included species: Aspergillus cervinus, A. christenseniae, A. kanagawaensis, A. novoguineensis, A. nutans, A. parvulus, A. subnutans, A. transcarpaticus, A. wisconsinensis.



Extrolites: Six of nine species produce terremutin, four of nine produce aspervenone, and three of nine produce 4-hydroxymellein (Chen et al. 2016c).



Notes: Aspergillus acidohumus is the sole species in ser. Acidohumorum. This series clearly belongs to section Cervini, but is phylogenetically distant from ser. Cervini. Series Acidohumorum is phenotypically distinct from other species by its very slow growth rate.



Section Fumigati W. Gams et al., Adv. Pen. Asp. Syst.: 56. 1986 [1985]. MycoBank MB832496.



Type: Aspergillus fumigatus Fresen., Beitr. Mykol. 3: 81. 1863.



Description: See Gams et al. (1985) (morphology); Peterson et al., 2008, Kocsubé et al., 2016, Fig. 1, Fig. 2, this study (phylogeny); Samson et al. (2007a) (morphology, phylogeny).



Series Brevipedes Houbraken & Frisvad, ser. nov. MycoBank MB832597.



Etymology: Named after the type species of the series, Aspergillus brevipes.



Type: Aspergillus brevipes G. Sm., Trans. Brit. Mycol. Soc. 35: 241. 1952.



Diagnosis: Phylogeny: Series Brevipedes belongs to sect. Fumigati, subgen. Fumigati; the series is well-supported; however, the phylogenetic relationship of the series with other series of the section remains unresolved (Fig. 15). Morphology & physiology: Colonies growing moderately fast or spreading; conidial colour en masse blue-green; conidiophores uniseriate; good growth at 37 °C. Sexual morph not observed in culture, or present, neosartorya-type, homothallic, white, yellowish white or pale yellow. Series description based on Samson et al. (2007a) and Hubka et al. (2017).



Included species: Aspergillus brevipes, A. duricaulis, A. quadricinctus, A. tsurutae.



Extrolites: Asperdurin, asperpentyn, cyclopaldic acids, duricaulic acid, fumagillin, meleagrin, pseurotins, roquefortine C, viriditoxin. Asperdurin, asperpentyn, cyclopaldic acids, duricaulic acid and meleagrin has only been found in ser. Brevipedes in sect. Fumigati.



Series Fennelliarum Houbraken & Frisvad, ser. nov. MycoBank MB832598.



Etymology: Named after the type species of the series, Aspergillus fennelliae.



Type: Aspergillus fennelliae Kwon-Chung & S.J. Kim, Mycologia 66: 629. 1974.



Diagnosis: Phylogeny: Series Fennelliarum belongs to sect. Fumigati, subgen. Fumigati; Fig. 15 shows a close phylogenetic relationship with ser. Unilaterales, though statistical support for this relationship is lacking. Morphology & physiology: Colonies spreading; conidial colour en masse blue-green; conidiophores uniseriate; good growth at 37 °C. Sexual morph neosarotya-type, homo- or heterothallic, white, yellowish white or pale yellow. Series description based on Samson et al. (2007a) and Hubka et al. (2017).



Included species: Aspergillus denticulatus, A. fennelliae, A. huiyaniae, A. similanensis, A. sublevisporus.



Extrolites: Antafumicins, asperfuran, aszonalenins, aszonapyrones, chevalones, fumigaclavines, fumigatins, gliotoxin, isocoumarins, pyripyropenes, reticulol, similanamide, similanpyrones, viridicatumtoxin, viriditoxin. Asperfuran, the isocoumarins, reticulol and viridicatumtoxin are only found in this series in sect. Fumigati.



Series Fumigati Houbraken & Frisvad, ser. nov. MycoBank MB832994.



Etymology: Named after the type species of the series, Aspergillus fumigatus.



Type: Aspergillus fumigatus Fresen., Beitr. Mykol. 3: 81. 1863.



Diagnosis: Phylogeny: Series Fumigati belongs to sect. Fumigati, subgen. Fumigati, and is phylogenetically most closely related to ser. Viridinutantes (Fig. 15). Morphology & physiology: Colonies spreading; conidial colour en masse blue-green; conidiophores uniseriate; growth at 37 and 50 °C. Sexual morph not observed in culture or present, neosartorya-type, homo- or heterothallic, white or yellowish white. Series description based on Samson et al. (2007a) and Hubka et al. (2017).



Included species: Aspergillus fischeri, A. fumigatiaffinis, A. fumigatus, A. fumisynnematus, A. laciniosus, A. lentulus, A. novofumigatus, A. oerlinghausensis, A. spinosus, A. takakii.



Extrolites: Extrolite families found in species of ser. Fumigati include ardeemins, asnovolins, asperfumigatin, asperfumin (asperfumoid), aszonalenins, aszonapyrones, aurantines, avenaciolides, cephalimycins, chevalones, chloroanthraquinones, chrysogine, cottoquinazolines, cycloechinuline, cyclopiazonic acids, cyclotryprostatins, fiscalins, fischerins, fumagillins, fumicyclines, fumigachlorin, fumigaclavines, fumigatins, fumimycins, fumiquinazolines, fumitremorgins, gliotoxins, helvolic acids, lentulins, neosartoricins, novoamauromins, novobenzomalvins, novofumigatamide, palitantins, pseurotins, pyripyropenes, sartorenol, sartorypyrones, sesterfischeric acids, setosusin, sphingofungins, takakiamide, terreins, (territrems), trypacidins, tryprostatins, tryptoquivalines, wortmannins, xanthocillins. Even though ser. Fumigati species produce the largest diversity of different secondary metabolites, certain secondary metabolites are only found in other series of sect. Fumigati, for example anishidiol, antafumicins, asperdurin, asperfurans, asperpentyns, avenaciolides, cyclopaldic acids, cytochalasins, gancidin, glabramycins, kotanins, meleagrin, monochaetin, mycophenolic acid, phomaligins, reticulol, sartoryglabrins, sartorymensin, tatenoic acid, viridicatumtoxins, viriditins, viriditoxins, wasabidienones, wortmannins.



Series Neoglabri Houbraken & Frisvad, ser. nov. MycoBank MB832599.



Etymology: Named after the type species of the series, Aspergillus neoglaber.



Type: Aspergillus neoglaber Kozak., Mycol. Pap. 161: 56. 1989.



Diagnosis: Phylogeny: Series Neoglabri belongs to sect. Fumigati, subgen. Fumigati; Fig. 15 shows a phylogenetic relationship with series Fennelliarum and Unilaterales, though statistical support for this relationship is lacking. Morphology & physiology: Colonies spreading; conidial colour en masse blue-green; conidiophores uniseriate; good growth at 37 °C. Sexual morph not observed in culture or present, neosartorya-type, homothallic, white, yellowish white or pale yellow. Series description based on Samson et al. (2007a) and Hubka et al. (2017).



Included species: Aspergillus auratus, A. australensis, A. brevistipitatus, A. conversis, A. elsenburgensis∗, A. galapagensis, A. neoglaber, A. papuensis, A. shendaweii, A. solicola, A. stramenius (∗ not included in Fig. 15).



Extrolites: Extrolite families found in ser. Neoglabri include antafumicins, asperpentyns, aszonalenins, avenaciolides, chrysogines, clavatols, fellutanines, fumigatin, glabramycins, gregatins, helvolic acid, kotanins, sartoryglabramides, sartoryglabrin, takakiamide, tryptoquivalines, wortmannins. The asperpentyns, chrysogines, fellutanins, glabramycins, gregatins, sartoryglabramides, sartoryglabrin, takakiamide have only been found in ser. Neoglabri.



Series Spathulati Houbraken & Frisvad, ser. nov. MycoBank MB832600.



Etymology: Named after the type species of the series, Aspergillus spathulatus.



Type: Aspergillus spathulatus Takada & Udagawa, Mycotaxon 24: 396. 1985.



Diagnosis: Phylogeny: Series Spathulati belongs to sect. Fumigati, subgen. Fumigati; this section is phylogenetically unique and Fig. 15 shows a relationship with series Fennelliarum, Neoglabri and Unilaterales, though statistical support for this relationship is lacking. Morphology & physiology: Colonies spreading; conidial colour en masse blue-green; conidiophores uniseriate; good growth at 37 °C. Sexual morph neosartorya-type, homothallic, pale yellow; ascospores with large equatorial crests, convex surface smooth. Series description based on Samson et al. (2007a) and Hubka et al. (2017).



Included species: Aspergillus spathulatus, A. takadae∗ (∗ not included in Fig. 15).



Extrolites: Aszonalenins, xanthocillins.



Series Thermomutati Houbraken & Frisvad, ser. nov. MycoBank MB832601.



Etymology: Named after the type species of the series, Aspergillus thermomutatus.



Type: Aspergillus thermomutatus (Paden) S.W. Peterson, Mycol. Res. 96: 549. 1992.



Diagnosis: Phylogeny: Series Thermomutati belongs to sect. Fumigati, subgen. Fumigati; the phylogenetic relationship with other series of the section remains unresolved (Fig. 15). Morphology & physiology: Colonies spreading; conidial colour en masse blue-green; conidiophores uniseriate; good growth at 37 °C. Sexual morph neosartorya-type, homothallic, white, yellowish white. Series description based on Samson et al. (2007a) and Hubka et al. (2017).



Included species: Aspergillus delicatus, A. tatenoi, A. thermomutatus.



Extrolites: Extrolite families found in ser. Thermomutati include aszonalenins, aszonapyrones, brasilianamides, eurochevalierine, fischerindoline, gliotoxins, helvolic acids, pseudofischerine, pyripyropenes, tatenoic acid, tryptoquivalines (R & S). The brasilianamides, eurochevelierine, fischerindole, pseudofischerine and tatenoic acid has only been found in species in ser. Thermomutati in sect. Fumigati.



Series Unilaterales Houbraken & Frisvad, ser. nov. MycoBank MB832602.



Etymology: Named after the type species of the series, Aspergillus unilateralis.



Type: Aspergillus unilateralis Thrower, Austral. J. Bot. 2: 355. 1954.



Diagnosis: Phylogeny: Series Unillateralis belongs to sect. Fumigati, subgen. Fumigati; Fig. 15 shows a close phylogenetic relationship with ser. Fennelliarum, though statistical support for this relationship is lacking. Morphology & physiology: Colonies spreading; conidial colour en masse blue-green; conidiophores uniseriate, sometimes with nodding heads; maximum growth temperature around 42 to 45 °C, sometimes lower (A. tasmanicus) or higher (A. assulatus, A. marvanovae, A. nishimurae, A. turcosus, and A. waksmanii, 47 °C). Sexual morph not observed in culture, or present, neosartorya-type, homo- or heterothallic, white, yellowish white or pale yellow. Series description based on Samson et al. (2007a) and Hubka et al. (2017).



Included species: Aspergillus assulatus, A caatingaensis, A. hiratsukae, A. marvanovae, A. multiplicatus, A. nishimurae, A. pernambucoensis, A. tasmanicus, A. tsunodae, A. turcosus, A. unilateralis, A. waksmannii.



Extrolites: Anishidiol, aszonalenins, aszonapyrones, avenaciolides, gliotoxins, helvolic acid, kotanins, monochaetin, mycophenolic acid and sartorypyrones. Anishidiol has only been found in series Unilaterales, but the kotanins have also been found in ser. Neoglabri.



Series Viridinutantes Houbraken & Frisvad, ser. nov. MycoBank MB832603.



Etymology: Named after the type species of the series, Aspergillus viridinutans.



Type: Aspergillus viridinutans Ducker & Thrower, Austal. J. Bot. 2: 355. 1954.



Diagnosis: Phylogeny: Series Viridinutantes belongs to sect. Fumigati, subgen. Fumigati and is phylogenetically most closely related to ser. Fumigati (Fig. 15). Morphology & physiology: Colonies spreading, some species growing moderately fast (A. viridinutans, A. frankstonensis); conidial colour en masse blue-green; conidiophores uniseriate, generally with nodding heads; good growth at 37 °C, maximum growth temperature of 42 or 45 °C. Sexual morph not observed in culture, or present, neosartorya-type, homo- or heterothallic, white, yellowish white. Series description based on Talbot et al. (2017) and Hubka et al. (2018a).



Included species: Aspergillus acrensis, A. arcoverdensis, A. aureolus, A. bezerrae, A. curviformis, A. felis, A. frankstonensis, A. pseudoviridinutans, A. siamensis, A. udagawae, A. viridinutans, A. wyomingensis. Aspergillus curviformis is tentatively included in this series based on its original description; no sequence data or material was available for this study.



Extrolites: Extrolite families found in ser. Viridinutantes include ardeemins, aszonapyrones, chevalones, clavatols, cytochalasins, fiscalins, fumigaclavines, fumagillins, fumigatins, fumiquinazolins, gancidin, helvolic acid, neosartoryadines, neosartoryones, phomaligins, pseurotins, pyripyropenes, trypacidins, sartorymensin, tryptoquivalines, viriditin, viriditoxin, wasabidienones. The cytochalasins, gancidin, neosartoryadines, neosartoryones, phomaligins, sartorymensin, and viriditin have only been found in ser. Viridinutantes in sect. Fumigati.



Notes on series in sect. Fumigati: Hubka et al. (2017) studied the phylogenetic relationship of sect. Fumigati species and recognised eight clades (A. brevipes-, A. fennelliae-, A. fumigatus-, A. neoglaber-, A. spinosus-, A. tatenoi-, A. unilateralis- and A. viridinutans-clade). More recently, Hubka et al. (2018a) recognised three additional clades: the A. thermomutatus-, A. spathulatus- and A. auratus-clade. Samson et al. (2007a) were the last who studied the taxonomy of the whole section. Based on this information, and data in more recent publications, it is difficult to find good characters to delimit series in this section. The series classification is therefore mainly based on (published) phylogenetic data (Hubka et al., 2017, Hubka et al., 2018a, this study). Extrolites in sect. Fumigati (with its eight families): ardeemins, asnovolins, aszonalenins, avenaciolides, cephalimycins, chaetominines, chevalones, chrysogines, clavatols, cycloechinulins, cyclopiazonic acids, cytochalasins, expansolides, fiscalins, fischerins, fumagillins,fumicyclins, fumigaclavines, fumigatins, fumigatonins, fumiquinazolines, fumitremorgins, gangicins, glabramycins, gliotoxins, helvolic acids, lentulins, neosartorins, novoamaauromins, novobenzomalvins, pseurotins, pyripyropenes, sartorypyrones, sphingofungins terreins trypacidins, tryprostatins, tryptoquivalines, viridicatumtoxins, viriditoxins, wortmannins and several more (see above) (Hong et al., 2005, Larsen et al., 2007, Samson et al., 2007a, Hong et al., 2008, Hubka et al., 2013b, Hubka et al., 2017, Hubka et al., 2018a, Frisvad and Larsen, 2015, Tamiya et al., 2015, Bessa et al., 2016, May Zin et al., 2016, Rajachan et al., 2016, Yu et al., 2016, Bang et al., 2019, Xu et al., 2019a, Yu et al., 2019). Among these, the aszonalenin biosynthetic family (BF) is shared by six species, the pyrones (aszonapyrone, chavalones, sartorypyrones) are shared by five species, the helvolic acid biosynthetic family is shared by five species, the pyripyropene BF is shared by four species, the gliotoxin BF is shared by four species, the fumigatin BF is shared by 4 species and the tryptoquivaline BF is shared by four species. Since the secondary metabolite biosynthetic family members are distributed as polythetic characters, not every member of the species series may produce these extrolites.



Section Vargarum Houbraken & Frisvad, sect. nov. MycoBank MB832604.



Etymology: In honour of Janos Varga, a prominent Aspergillus researcher and advocate of a broad monophyletic Aspergillus including polypaecilum-type morphs.



Type: Aspergillus cejpii (Milko) Samson et al., Stud. Mycol. 78: 155. 2014.



Diagnosis: Phylogeny: Section Vargarum belongs to subgen. Fumigati and is phylogenetically most closely related to sect. Clavati (Fig. 2, Fig. 15) Morphology & physiology: Colonies growing moderately fast; conidiophores polypaecilum-like; growth at 37 °C. Sexual morph neosartorya-type, homothallic, yellowish white.



Extrolites: Isolates in ser. Vargarum can produce gliotoxin, rubratoxins, tryptoquivalones and xanthocillins (Varga et al. 2007a). These extrolites are also produced by other series in subgen. Fumigati, except rubratoxins, which have been found only in this series in Aspergillus, but is also produced by Talaromyces purpurogenus (Yilmaz et al. 2012) outside Aspergillus.



Included species: Aspergillus cejpii.



Notes: The polypaecilum-like asexual morph present in ser. Vargarum is unique in subgen. Fumigati. This morphology type is also found in species belonging to Aspergillus subgen. Polypaecilum.



Aspergillus subgen. Nidulantes W. Gams et al., Adv. Pen. Asp. Syst: 57. 1986 [1985]. MycoBank MB832501.



Type: Aspergillus nidulellus Samson & W. Gams, Adv. Pen. Asp. Syst.: 44. 1986 [1985] (= Aspergillus nidulans).



Description: See Gams et al. (1985) (morphology); Kocsubé et al. (2016), Fig. 1, Fig. 2, this study (phylogeny); Chen et al. (2016a) (morphology, phylogeny).



Section Aenei Varga & Samson, IMA Fungus 1: 203. 2010. MycoBank MB517672.



Type: Aspergillus aeneus Sappa, Allionia 2: 84. 1954.



Description: Phylogeny: Section Aenei belongs to subgen. Nidulantes and is phylogenetically sister to sect. Nidulantes (Fig. 2, Fig. 16) Morphology & physiology: Colonies growing moderately or fast, conidia en masse in shades of green or olive-brown; conidiophores biseriate, stipes brown pigmented, Hülle cells abundant (except in A. heyangensis), often in crusts, globose, subglobose or pyriform; no growth at or above 40 °C. Sexual morph not observed in culture, or present, emericella-type, homothallic; ascospore convex smooth or delicately roughened, with two equatorial crests. Section description based on Varga et al. (2010a).



Fig. 16.

Combined phylogeny for BenA, CaM and RPB2 data sets showing the phylogenetic relation of species, series and sections within Aspergillus subgen. Nidulantes. The BI posterior probability (pp) values and bootstrap percentages of the maximum likelihood (ML) analysis are presented at the nodes; fully supported branches are thickened. Values less than 70 % bootstrap support (ML) or less than 0.95 posterior probability (Bayesian analysis) are indicated with a hyphen or not shown. The bar indicates the number of substitutions per site. The phylogram is rooted with Hamigera avellanea and Penicillium expansum.

Included species: Aspergillus aeneus, A. bicolor, A. coloradensis∗, A. crustosus, A. discophorus, A. eburneocremeus, A. foeniculicola, A. heyangensis, A. karnatakaensis, A. spectabilis [∗ not included in Fig. 16; details on classification, see Sklenář et al. (2020)].



Extrolites: Sterigmatocystin (5/9), decaturins (3/9), asperugins (2/9), asteltoxin (2/9), karnatakafurans (2/9), Mer-NF8054X (2/9), emeheteron (1/9), fumitremorgins (1/9), physcion (1/9), PR-toxin (1/9), quinolactacin (1/9), stellatin (1/9), terrein (1/9) (Manniche et al., 2004, Varga et al., 2010a).



Notes: Hubka et al. (2016a) included the current sect. Aenei in sect. Nidulantes as the Aspergillus aeneus-clade. Our nine-gene phylogenetic analysis locates representatives of sect. Aenei with statistical confidence outside sect. Nidulantes, confirming the results of previous reports (Varga et al., 2010a, Chen et al., 2016a). Phenotypically, the sexual species in sect. Aenei (A. bicolor, A. discophorus, A. foeniculicola, A. spectabilis) produce similar ascospores as taxa in the A. nidulans-clade, but none of them are able to grow at 40 °C (Chen et al. 2016a). There are no data indicating a subdivision of sect. Aenei into more than one series, and ser. Aenei is therefore only informally introduced here (see Table 4).



Section Bispori S.W. Peterson et al. ex Houbraken, sect. nov. MycoBank MB833244.

Synonym: Aspergillus sect. Bispori Peterson et al., Aspergillus in the genomic era: 42. 2008; sect. inval.



Etymology: Named after the type species of this section, A. bisporus.



Type: Aspergillus bisporus Kwon-Chung & Fennell, Mycologia 63: 479. 1971.



Diagnosis: Phylogeny: Section Bispori belongs to subgen. Nidulantes and is phylogenetically sister to a large clade containing sections Nidulantes, Aenei, Usti, Cavernicolarum, Raperorum and Silvatici. Morphology & physiology: Colonies restricted or growing moderately fast, conidia en masse olive to dark brown; conidiophores uniseriate, stipes slightly brown pigmented, smooth, occasionally showing granular pigment deposits; colonies growing more rapid at 37 °C than at 25 °C. Sexual morph unknown. Section description based on Kwon-Chung & Fennell (1971) (under A. bisporus).



Included species: Aspergillus bisporus.



Extrolites: There are no known extrolites from A. bisporus.



Notes: Aspergillus bisporus could not be satisfactorily fitted in any of the groups delineated by Raper & Fennell (1965). Based on morphological similarities, Samson (1979) classified the species in sect. Cervini. Molecular studies revealed that A. bisporus belongs to subgen. Nidulantes (Peterson, 2000, Peterson, 2008, Peterson et al., 2008, Chen et al., 2016a). Section Bispori was informally introduced (Peterson et al. 2008), and this section is validated here. This section includes one species, A. bisporus, and introducing ser. Bispori would therefore be superfluous; ser. Bispori is only informally introduced here.



Section Cavernicolarum [as “Cavernicolus”’] A.J. Chen et al., Stud. Mycol. 84: 112. 2016. MycoBank MB832535.



Type: Aspergillus cavernicola Lörinczi, Contrtii bot. Univ. Babes-Bolyai, Cluj, Grad. bot.: 341. 1969.



Description: See Chen et al. (2016a) (morphology, phylogeny).



Series Cavernicolarum Houbraken & Frisvad, ser. nov. MycoBank MB833245.



Etymology: Named after the type species of this series, A. cavernicola.



Type: Aspergillus cavernicola Lörinczi, Contrtii bot. Univ. Babes-Bolyai, Cluj, Grad. bot.: 341. 1969.



Diagnosis: Phylogeny: Series Cavernicolarum belongs to sect. Cavernicolarum, subgen. Nidulantes and is phylogenetically sister to a clade containing ser. Egyptiaci. Morphology & physiology: Colonies growing restrictedly or moderately rapid, conidia en masse in shades of green or brown; conidiophores biseriate, short, occasionally long (in A. californicus), stipes uncoloured or in brown shades, smooth, Hülle cells often present, usually globose or subglobose; no growth at 37 °C. Sexual morph unknown. Series description based on Raper & Fennell (1965) and Chen et al. (2016a).



Included species: Aspergillus californicus, A. cavernicola, A. kassunensis, A. subsessilis.



Extrolites: Red azaphilones of the same constitution as monascorubramin and N-glutarylrubropunctamin and similar extrolites found in Talaromyces albobiverticillius, Tal. atroroseus and Tal. purpurogenus (Yilmaz et al., 2012, Frisvad et al., 2013b, Venkatachalam et al., 2018) have been detected in A. cavernicola, including monasnicotinic acid (Antipova et al. 2018a). Aspergillus kassunensis produces asperugins and Mer-NF8054X and A. californicus arugosins (Samson et al. 2011b). No extrolites are common for the four species.



Series Egyptiaci Houbraken & Frisvad, ser. nov. MycoBank MB833246.



Etymology: Named after the type species of this series, A. egyptiacus.



Type: Aspergillus egyptiacus Moub. & Mustafa, Egypt. J. Bot. 15: 153. 1972.



Diagnosis: Phylogeny: Series Egyptiaci belongs to sect. Cavernicolarum, subgen. Nidulantes and is phylogenetically sister to a clade containing ser. Cavernicolarum. Morphology & physiology: Colonies restricted, conidia en masse in shades of green; conidiophores, mostly not arranged in typical Aspergillus heads when grown at 25 °C, biseriate, short, smooth; often also poorly developed conidiophores present: phialides solitary or occurring in little groups along the conidiophores resembling penicillium-like structures; Hülle cells produced, varying from globose to almost cylindrical; good growth on malt extract agar with 20 % or 40 % sucrose at 35 °C, minimum growth temperature 15 °C, optimum 35 °C and maximum 45 °C. Sexual morph unknown. Series description based on Samson & Mouchacca (1974).



Included species: Aspergillus egyptiacus.



Extrolites: Fumitremorgin A & B, verruculogen and biosynthetically related indole-terpene-alkaloids are produced by A. egyptiacus (Samson et al. 2011b). These compounds are produced by species in both Aspergillus (different sections) and Penicillium (also different sections), but also by one species in sect. Nidulantes, ser. Stellati, namely A. caespitosus (Schroeder et al., 1975, Steyn et al., 1981).



Notes: Section Cavernicolarum contains five species previously assigned to sect. Usti and share the production of short stipes. Series Egyptiaci is phylogenetically distinct and the sole species in this section that grows well at 37 °C; ser. Cavernicolarum species do not grow at this temperature.



Section Nidulantes W. Gams et al., Adv. Pen. Asp. Syst.: 57. 1986 [1985]. MycoBank MB832502.



Type: Aspergillus nidulellus Samson & W. Gams, Adv. Pen. Asp. Syst.: 44. 1986 [1985]. MycoBank MB114711 (= Aspergillus nidulans).



Description: See Gams et al., 1985, Chen et al., 2016a (morphology, phylogeny), Kocsubé et al. (2016) (phylogeny).



Series Aurantiobrunnei Houbraken & Frisvad, ser. nov. MycoBank MB833247.



Etymology: Named after the type species of this series, A. aurantiobrunneus.



Type: Aspergillus aurantiobrunneus (G.A. Atkins et al.) Raper & Fennell, Gen. Aspergillus: 511. 1965.



Diagnosis: Phylogeny: Series Aurantiobrunnei belongs to sect. Nidulantes, subgen. Nidulantes and is phylogenetically sister of ser. Speluncei. Morphology & physiology: Colonies restricted, sporulation absent or sparse; conidiophores biseriate, smooth, stipes hyaline to pale brown; Hülle cells present, hyaline to pale brown, globose, subglobose or ovoid; no growth at 37 °C. Sexual morph emericella-type, homothallic; ascospores globose to subglobose, with two equatorial crests, smooth convex. Series description based on Chen et al. (2016a).



Included species: Aspergillus aurantiobrunneus, A. purpureus.



Extrolites: Epurpurins (including emerin), variecolactones (including emericolins, stellatic acid, variecoacetal, variecolin, variecolol), and sterigmatocystin (including versicolorins) are shared by the two species in the ser. Aurantiobrunnei. Eremophiline has been found in A. aurantiobrunneus and calbistrins and shamixanthones has been found in A. purpureus (Chen et al. 2016a).



Series Multicolores Houbraken & Frisvad, ser. nov. MycoBank MB833248.



Etymology: Named after the type species of this series, A. multicolor.



Type: Aspergillus multicolor Sappa, Allionia 2: 87. 1954.



Diagnosis: Phylogeny: Series Multicolores belongs to sect. Nidulantes, subgen. Nidulantes and is phylogenetically sister of ser. Nidulantes. Morphology & physiology: Colonies growing moderately fast or spreading, conidia en masse in shades of green; conidiophores biseriate, smooth, stipes hyaline to yellowish brown; Hülle cells absent (A. mulundensis, A. purpureocrustaceus) or present (A. incahuasiensis, A. multicolor, A. pluriseminatus, A. tumidus), pale yellowish brown, orange, brown to pink, globose, subglobose or ovoid. Sexual morph not observed in culture, or present (A. pluriseminatus), emericella-type, homothallic; ascospores lenticular, convex surface tuberculate under SEM, with two conspicuously pleated, stellate and striate equatorial crests. Series description based on Stchigel and Guarro, 1997, Chen et al., 2016a, Crous et al., 2018b, Piontelli et al., 2019.



Included species: Aspergillus incahuasiensis∗, A. multicolor, A. mulundensis, A. pluriseminatus, A. purpureocrustaceus∗, A. tumidus [∗ not included in Fig. 16; details on classification, see Sklenář et al. (2020)].



Extrolites: Aspergillus multicolor, A. mulundensis and A. pluriseminatus all produce asticolorins or closely related dibenzofurans. Aspergillus mulundensis produces mulundocandins and emericellamide, and azaphilones in common with A. pluriseminatus (Bills et al., 2016, Chen et al., 2016a), while A. multicolor is the only species in the series that has been reported to produce sterigmatocystins (Hamasaki et al. 1980).



Series Nidulantes Houbraken & Frisvad, ser. nov. MycoBank MB833249.



Etymology: Named after the type species of this series, A. nidulellus.



Type: Aspergillus nidulellus Samson & W. Gams, Adv. Pen. Asp. Syst.: 44. 1986 [1985]. MycoBank MB114711 (= Aspergillus nidulans).



Diagnosis: Phylogeny: Series Nidulantes belongs to sect. Nidulantes, subgen. Nidulantes and is phylogenetically sister of ser. Multicolores. Morphology & physiology: Colonies spreading, conidia en masse green; conidiophores biseriate, stipes hyaline to yellowish brown pigmented, smooth, occasionally with surface protuberances; Hülle cells present, globose; good growth at 37 °C and 40 °C, growth at 45 °C, except for A. botswanensis, A. fruticulosus, A. latilabiatus and A. recurvatus. Sexual morph generally present (only not observed in A. recurvatus), emericella-type, homothallic; ascospores irregularly wrinkled, finely pitted, rugulose or echinulate, with two equatorial crests (except for four crests in A. quadrilineatus). Series description based on Chen et al. (2016a).



Included species: Aspergillus amethystinus∗, A. aurantiopurpureus, A. botswanensis, A. corrugatus, A. desertorum, A. dipodomyus, A. falconensis, A. foveolatus, A. fruticulosus, A. jaipurensis, A. latilabiatus, A. navahoensis, A. nidulans, A. omanensis, A. pachycristatus, A. quadrilineatus, A. recurvatus, A. rugulosus, A. savannensis, A. spinulosporus, A. stercorarius, A. striatus, A. sublatus, A. sulphureoviridis, A. violaceus [∗ not included in Fig. 16; details on classification, see Sklen]? et al. (2020)].



Extrolites (number of species producing compound / total species in series): Asperthecin (14/22), asperugins (10/22), austinols (2/22), austalides (5/22), calbistrins (2/22), cordycepin (1/22), cyclopaldic acid (1/22), desertorins (6/22), echinocandins (3/22), emericellin (9/22), emerin/epurpurins (2/22), emestrin (6/22), emindols (7/22), falconensins (6/22), falconensons (5/22), gregartins (2/22), 2-ω-hydroxyemodin (4/22), isocoumarins (1/22), paxillin (6/22), quadrilineatin (and the related nidulol & silvaticol) (2/22), shamixanthones (10/22), sterigmatocystin (14/22, A. latilabiatus producing only versicolorins), terrequinone A (4/22), violaceols (16/22), viridicatumtoxin (1/22) (Chen et al. 2016a).



Series Speluncei Houbraken & Frisvad, ser. nov. MycoBank MB833250.



Etymology: Named after the type species of this series, A. spelunceus.



Type: Aspergillus spelunceus Raper & Fennell [as “speluneus”], Gen. Aspergillus: 457. 1965.



Diagnosis: Phylogeny: Series Speluncei belongs to sect. Nidulantes, subgen. Nidulantes and is phylogenetically sister of ser. Aurantiobrunnei. Morphology & physiology: Colonies restricted or moderately fast, sporulation sparse or moderate, en masse in shades of green; conidiophores biseriate, stipes hyaline or yellowish brown, smooth; Hülle cells absent or present (A. askiburgiensis, A. asperescens, A. spelunceus), hyaline, globose, subglobose or ovoid; no growth at 37 °C (except A. asperescens). Sexual morph unknown. Series description based on Chen et al. (2016a).



Included species: Aspergillus askiburgiensis, A. asperescens, A. aureolatus, A. spelunceus, A. varians, A. viridicatenatus.



Extrolites (number of species producing compound / total species in series): Azaphilones (1/6), austinols (1/6), calbistrins (1/6), desertorins (1/6), emerin / epurpurins (2/6), 2-ω-hydroxyemodin (1/6), a phthalide (1/6), shamixanthones (1/6), sterigmatocystin (5/6, A. varians only produces the versicolorin precursors), violaceols (1/6), viridicatins and cyclopenols (1/6) (Chen et al. 2016a).



Series Stellati Houbraken & Frisvad, ser. nov. MycoBank MB833251.



Etymology: Named after the type species of this series, A. stellatus.



Type: Aspergillus stellatus Curzi, Atti Reale Accad. Naz. Lincei, Rendiconti Cl. Sci. Fis. 19: 428. 1934.



Diagnosis: Phylogeny: Series Stellati belongs to sect. Nidulantes, subgen. Nidulantes and is phylogenetically sister of a clade including series Multicolores, Nidulantes and Unguium. Morphology & physiology: Colonies spreading, conidia en masse green, conidiophores biseriate, stipes hyaline to yellowish brown, smooth; Hülle cells present, globose; no growth at 40 °C. Sexual morph present (except in A. caespitosus), emericella-type, homothallic; ascospores globose, stellate or appendaged. Series description based on Chen et al. (2016a).



Included species: Aspergillus angustatus, A. astellatus, A. caespitosus, A. dromiae, A. filifer, A. miraensis, A. olivicola, A. qinqixianii, A. stella-maris, A. stellatus, A. stelliformis∗, A. undulatus, A. venezuelensis [∗ not included in Fig. 16; details on classification, see Sklenář et al. (2020)].



Extrolites (number of species producing compound / total species in series): Aflatoxin B₁ (4/12), asperlicine (1/12), asperthecin (6/12), asperugines (3/12), astellolide (1/12), asteltoxin (4/12), austinol (1/12), curvularins (1/12), desertorins (6/12), emericellins (8/12), emerin / epurpurin (1/12), fischerin (1/12), gregatins (1/12), 2-ω-hydroxyemodin (8/12), Mer-NF-8054X (1/12), secalonic acid D 781/12), shamixanthones (10/12), sterigmatocystin (5/12), terrein (3/12), varitriol (4/12), verruculogens and fumitremorgins (1/12), violaceols (1/12) (Chen et al. 2016a).



Series Unguium Houbraken & Frisvad, ser. nov. MycoBank MB833253.



Etymology: Named after the type species of this series, A. unguis.



Type: Aspergillus unguis (Émile-Weill & L. Gaudin) Thom & Raper, Mycologia 31: 667. 1939.



Diagnosis: Phylogeny: Series Unguium belongs to sect. Nidulantes, subgen. Nidulantes and is phylogenetically sister of series Multicolores and Nidulantes. Morphology & physiology: Colonies growing restrictedly or moderately rapid; conidia en masse in shades of green (yellow-green, olive-green); conidiophores biseriate; stipes hyaline to yellowish brown, smooth; Hülle cells absent (A. israelensis, A. unguis) or present (A. croceus), globose, subglobose or pyriform; growth at 37 °C absent (A. croceus, A. israelensis) or restricted (A. unguis), some A. unguis strains grow restrictedly at 40 °C). Sexual morph unknown, A. unguis NRRL 2393 was reported to tardily produce ascospores. Series description based on Fennell and Raper, 1955, Chen et al., 2016a, Hubka et al., 2016a.



Included species: Aspergillus croceiaffinis∗, A. croceus, A. israelensis, A. longistipitatus∗, A. unguis [∗ not included in Fig. 16; details on classification, see Sklenář et al. (2020)].



Extrolites: Aspergillus unguis produces asperunguisones (and 3-ethyl-5,7-dihydroxy-3,6-dimethyl phthalide), penicillin G, unguinols (= nidulins), aspergillusidones, aspergicides, agonodepsides, emeguisins, folipastatins, haiderin, nasrin, rubidin, shirin, yasimin (= unguinol = tridechloronidulin), unguisins, unguispyrones, ustilaginoidin C and violaceols / orcinols (Chen et al., 2016a, Phainuphong et al., 2017a, Morshed et al., 2018, Phainuphong et al., 2018a), while A. croceus produces desertorins / kotanins and sterigmatocystin and A. israelensis produce emindols (Chen et al. 2016a), showing there is no extrolites in common between the three species in series Unguium.



Series Versicolores Houbraken & Frisvad, ser. nov. MycoBank MB833254.



Etymology: Named after the type species of this series, A. versicolor.



Type: Aspergillus versicolor (Vuill.) Tirab., Ann. Bot. (Roma) 7: 9. 1908.



Diagnosis: Phylogeny: Series Versicolores belongs to sect. Nidulantes, subgen. Nidulantes; the phylogenetic relationship with other series in the section is unresolved. Morphology & physiology: Colonies restricted or moderately fast, conidia en masse in shades or green or brown; conidiophores biseriate, stipes hyaline or pale brown, smooth or with tubercles; Hülle cells absent or present, hyaline, globose, subglobose, ellipsoidal or pyriform; no or poor growth at 37 °C. Sexual morph unknown. Series description based on Jurjević et al. (2012) and Chen et al. (2016a).



Included species: Aspergillus amoenus, A. austroafricanus, A. creber, A. cvjetkovicii, A. fructus, A. griseoaurantiacus, A. hongkongensis, A. jensenii, A. pepii, A. protuberus, A. puulaauensis, A. subversicolor, A. sydowii, A. tabacinus, A. tennesseensis, A. venenatus, A. versicolor.



Extrolites: Most species produce sterigmatocystin (14/16) and precursors such as versicolorins, averufin, averufanin, and norsolorinic acid ((Jurjević et al., 2013, Chen et al., 2016a, Jakšić Despot et al., 2017). Other extrolites from series Versicolores include aniduquinolones (2/14), brevianamides F, J, K, Q, R, T, U (1/14), calbistrins (3/14), deoxybrevianamides (2/14), isocoumarins (2/14), cyclopenols / viridicatols (2/14), emericellin / arugosins (1/14), shamixanthones (4/14), insulicolides (2/14), mangrovamides (1/14), psychrophilin E-H (1/14), sydowic acids (5/14), sydowinins (1/14), versicolamides (and notoamides, stephacidins) (7/14), versiols (2/14), violaceols (13/14), WIN64745 (1/14) (Chen et al. 2016a).



Notes on sect. Nidulantes and included series: Section Nidulantes species share the production of more or less brown-pigmented, smooth conidiophores (occasionally with surface protuberances) with globose, subglobose or subclavate vesicles. The conidia are generally globose and echinulate and are en masse green coloured. Chen et al. (2016a) recognised seven clades in sect. Nidulantes and these were named the A. aurantiobrunneus-, A. multicolor-, A. nidulans-, A. spelunceus-, A. stellatus-, A. unguis- and A. versicolor-clade. These seven clades are here treated as series. These series were primarily based on multigene phylogenetic analysis and certain shared characters, such as growth rates at 25, 37 and 40 °C and ascospore ornamentation (if produced).



Section Ochraceorosei Frisvad & Samson, Syst. Appl. Microbiol. 28: 451. 2005. MycoBank MB500165.



Type: Aspergillus ochraceoroseus Bartoli & Maggi, Trans. Brit. Mycol. Soc. 71: 393. 1978.



Description: See Frisvad et al. (2005) (morphology), Kocsubé et al. (2016) (phylogeny).



Series Funiculosi Houbraken & Frisvad, ser. nov. MycoBank MB833255.



Etymology: Named after the type species of this series, A. funiculosus.



Type: Aspergillus funiculosus G. Sm., Trans. Brit. Mycol. Soc. 39: 111. 1956.



Diagnosis: Phylogeny: Series Funiculosi belongs to sect. Ochraceorosei, subgen. Nidulantes and is phylogenetically sister ser. Ochraceorosei (Fig. 16). Morphology & physiology: Colonies growing moderately; conidia en masse yellow-green, olive-brown to deep brownish purple; conidiophores uniseriate, stipes hyaline or faintly coloured just below the vesicle, smooth; Hülle cells absent. Sexual morph unknown. Series description based on Smith (1956) and Raper & Fennell (1965).



Included species: Aspergillus funiculosus.



Extrolites: Aspergillus funiculosus produces kojic acid (Siddhardha et al. 2010) and funicin = ethericin B (König et al., 1978, Hamasaki et al., 1980, König et al., 1980, Nakamura et al., 1984) and is chemically different from A. ochraceoroseus and A. rambellii in series Ochraceorosei. Chemically these diphenylether antibiotic extrolites are shared with Aspergillus sydowii and other species in series Versicolores (Li et al. 2015a) and kojic acid is shared with some species in section Nidulantes (Frisvad and Samson, 2004a, Frisvad et al., 2005, Chen et al., 2016a) and Flavi (Frisvad et al. 2019).



Series Ochraceorosei Houbraken & Frisvad, ser. nov. MycoBank MB833256.



Etymology: Named after the type species of this series, A. ochraceoroseus.



Type: Aspergillus ochraceoroseus Bartoli & Maggi, Trans. Brit. Mycol. Soc. 71: 393. 1979 [1978].



Diagnosis: Phylogeny: Series Ochraceorosei belongs to sect. Ochraceorosei, subgen. Nidulantes and is phylogenetically sister ser. Funiculosi. Morphology & physiology: Colonies growing moderately or fast on MEA, conidia en masse in shades of yellow; conidiophores biseriate, stipes hyaline, smooth, long; Hülle cells absent; no growth at 37 °C. Sexual morph unknown. Series description based on Bartoli & Maggi (1978) and Frisvad et al. (2005).



Included species: Aspergillus ochraceoroseus, A. rambellii.



Extrolites: Aflatoxin B₁, B₂, sterigmatocystin, 3-O-methylsterigmatocystin (Frisvad et al. 2005). Many other species produce aflatoxin and the related sterigmatocystins, but species in ser. Ochraceorosei do not produce kojic acid like species in sect. Flavi (Frisvad et al. 2019), but produce unique secondary metabolites, not as yet structure elucidated, that only occur in this section. No extrolites in common with the other species in sect. Nidulantes (Chen et al. 2016a).



Notes on series in sect. Ochraceorosei: The taxonomic position of A. funiculosus was discussed several times in the past (Smith, 1956, Raper and Fennell, 1965, Peterson, 2008, Peterson et al., 2008, Hubka et al., 2016a). Smith (1956) could not satisfactorily classify this species and mentioned a possible relationship with A. versicolor, A. flavus and A. glaucus, and Raper & Fennell (1965) accepted this species in sect. Sparsi (as A. sparsus group). Using sequence data, A. funiculosus took a basal position in sect. Sparsi in the phylogenetic analysis of Peterson et al. (2008), while this species was more similar to A. ochraceoroseus in other analyses (Houbraken and Samson, 2011, Chen et al., 2016a). Based on our nine-gene phylogeny, A. funiculosus is sister to Aspergillus ochraceoroseus, A. rambellii of ser. Ochraceorosei (Fig. 2). The sole species in ser. Funiculosi is uniseriate, in contrast to the biseriate species in ser. Ochraceorosei. Aspergillus funiculosus does not produce aflatoxins or sterigmatocystins, but kojic acid and funicin (= ethericin B) (König et al., 1978, Hamasaki et al., 1980, König et al., 1980, Nakamura et al., 1984, Siddhardha et al., 2010).



Section Raperorum S.W. Peterson, Varga, Frisvad, Samson ex Houbraken, sect. nov. MycoBank MB833258.

Synonym: Aspergillus sect. Raperi Peterson et al., Aspergillus in the genomic era: 42. 2008; sect. inval.



Etymology: Named after the type species of this section, A. raperi.



Type: Aspergillus raperi Stolk & J.A. Meyer, Trans. Brit. Mycol. Soc. 40: 190. 1957.



Diagnosis: Phylogeny: Section Raperi belongs to subgen. Nidulantes and is phylogenetically sister to a clade containing sections Nidulantes, Aenei, Usti and Cavernicolarum (Fig. 16). Morphology & physiology: Colonies growing restrictedly, moderately or fast, conidia en masse in shades of yellow or green; conidiophores uniseriate (A. raperi) or biseriate (A. ivorensis), stipes hyaline, smooth (A. raperi) or rough (A. ivorensis), long; Hülle cells present, globose to subglobose, pyriform or elongate. Sexual morph unknown. Section description based on Bartoli & Maggi (1978) and Stolk & Meyer (1957).



Included species: Aspergillus ivoriensis, A. raperi.



Extrolites: No known extrolites have been reported.



Notes: Section Raperorum is introduced to accommodate A. ivoriensis and A. raperi. These species differ in growth rates and conidiophore structure. However, we tentatively keep both species in one series. Based on phenotypic characters, Aspergillus raperi was first placed in the A. versicolor group (= ser. Versicolores) (Stolk & Meyer 1957) and later in the A. ornatus group (Raper & Fennell 1965). This species forms, together with A. ivoriensis, a unique lineage in subgen. Nidulantes, and sect. Raperi was informally introduced to accommodate these species (Peterson et al. 2008). This section is formally introduced here as sect. Raperorum. Section Raperorum is not subdivided in series and therefore ser. Raperorum is only informally introduced here.



Section Silvatici S.W. Peterson, Varga, Frisvad, Samson ex Houbraken, sect. nov. MycoBank MB833259.

Synonym: Aspergillus sect. Silvati Peterson et al., Aspergillus in the genomic era: 44. 2008; sect. inval.



Etymology: Named after the type species of this section, A. silvaticus.



Type: Aspergillus silvaticus Fennell & Raper, Mycologia 47: 83. 1955.



Diagnosis: Phylogeny: Section Silvatici belongs to subgen. Nidulantes and is phylogenetically sister to a large clade containing sections Nidulantes, Aenei, Usti, Cavernicolarum and Raperorum (Fig. 16). Morphology & physiology: Colonies growing moderate to fast, conidia en masse in shades of green; conidiophores biseriate, stipes brownish pigmented, smooth, seldom exceeding 300 μm; Hülle cells present, globose to subglobose. Sexual morph unknown. Section description based on Raper & Fennell (1965).



Included species: Aspergillus silvaticus.



Extrolites: Naphthalic anhydride, phthalides (silvaticol, O-methylsilvaticol, ethyl 3-methylorsellinate, 6-hydroxy-4-methoxy-5-methylphtalimidine, 3,6-dimethyl-4-hydroxy-2-methoxybenzaldehyde, nidulol, quadrilineatin), shamixanthones (arugosin A, B & E, silvaticamide), silvathione and dithiosilvatin (Homma et al., 1980, Yamazaki et al., 1981a, Fujita et al., 1985, Maebayashi and Yamazaki, 1985, Kawahara et al., 1986, Kawahara et al., 1987, Kawahara et al., 1988). Naphthalic anhydride and the silvatins are unique to this section.



Notes: Section Silvatici (as Silvati) was informally introduced to accommodate A. silvaticus (Peterson et al. 2008) and this section is formally introduced here. Only one species is accommodated in sect. Silvatici. Introducing ser. Silvatici would therefore be superfluous, and the series name Silvatici is therefore only informally introduced.



Section Sparsi W. Gams, M. Chr., Onions, Pitt & Samson, Adv. Pen. Asp. Syst.: 61. 1986 [1985]. MycoBank MB832514.



Type: Aspergillus sparsus Raper & Thom, Mycologia 36: 572. 1944.



Description: See Gams et al. (1985) (morphology), Kocsubé et al. (2016) (phylogeny).



Series Biplani Houbraken & Frisvad, ser. nov. MycoBank MB833260.



Etymology: Named after the type species of this series, A. biplanus.



Type: Aspergillus biplanus Raper & Fennell, Gen. Aspergillus: 434. 1965.



Diagnosis: Phylogeny: Series Biplani belongs to sect. Sparsi, subgen. Nidulantes and is phylogenetically sister to ser. Sparsi (Fig. 16). Morphology & physiology: Colonies growing restrictedly or moderately rapid, conidia en masse in shades of green (dark green, blueish green); conidiophores biseriate, stipes brown, smooth, long; fragmentary conidial structures of varying size and without definite pattern borne near the agar surface; Hülle cells absent. Sexual morph unknown. Series description based on Raper & Fennell (1965).



Included species: Aspergillus biplanus, A. diversus.



Extrolites: Auroglaucin is shared with sections Aspergillus and Restricti, also with A. conjuctus in ser. Conjuncti (Varga et al. 2010b).



Series Conjuncti Houbraken & Frisvad, ser. nov. MycoBank MB833261.



Etymology: Named after the type species of this series, A. conjunctus.



Type: Aspergillus conjunctus Kwon-Chung & Fennell, Gen. Aspergillus: 552. 1965.



Diagnosis: Phylogeny: Series Conjuncti belongs to sect. Sparsi, subgen. Nidulantes and is phylogenetically sister to ser. Implicati (Fig. 16). Morphology & physiology: Colonies grow restrictedly or moderately rapid; conidia en masse in shades of brown (brown, red-brown, olive brown); conidiophores biseriate, stipes hyaline or (pale) brown pigmented, smooth; reduced conidial structures produced near colony surface absent; Hülle cells absent (A. amazonicus, A. anthodesmis) or present (A. conjunctus, A. panamensis), elongate and/or variously curved or twisted. Sexual morph unknown. Series description based on Raper and Fennell, 1965, Bartoli and Maggi, 1978.



Included species: Aspergillus amazonicus, A. anthodesmis, A. conjunctus, A. panamensis.



Extrolites (number of species producing compound / total number of species in series): An aszonalenin (1/4), auroglaucin (1/4), gregatins (2/4), and siderin (3/4) have been found in species in ser. Conjuncti.



Series Implicati Houbraken & Frisvad, ser. nov. MycoBank MB833262.



Etymology: Named after the type species of this series, A. implicatus.



Type: Aspergillus implicatus Persiani & Maggi, Mycol. Res. 98: 871. 1994.



Diagnosis: Phylogeny: Series Implicati belongs to sect. Sparsi, subgen. Nidulantes and is phylogenetically sister to ser. Conjuncti (Fig. 16). Morphology & physiology: Colonies growing moderately, conidia en masse white or pale yellow; conidiophores biseriate, stipes hyaline, smooth, long, surrounded by parallel sterile hyphae, slightly echinulate, originating from the base growing up to the vesicle and later then branching to build the hyphal tangle; reduced, penicillate conidiophores present; Hülle cells absent. Sexual morph unknown. Series description based on Maggi & Persiani (1994).



Included species: Aspergillus implicatus.



Extrolites: Only a versicolorin has been detected in A. implicatus (Varga et al. 2010b), indicating that this species may produce either sterigmatocystin and aflatoxins or austocystins, as versicolorins are precursors for both biosynthetic families; however, this has not been confirmed. Since species in sect. Sparsi are not known for producing aflatoxins or austocystins, and since A. implicatus was found in the same habitat as A. ochraceoroseus, that actually produce aflatoxins.



Series Sparsi Houbraken & Frisvad, ser. nov. MycoBank MB833263.



Etymology: Named after the type species of this series, A. sparsus.



Type: Aspergillus sparsus Raper & Thom, Mycologia 36: 572. 1944.



Diagnosis: Phylogeny: Series Sparsi belongs to sect. Sparsi, subgenus Nidulantes and is phylogenetically sister to ser. Biplani (Fig. 16). Morphology & physiology: Colonies spreading, conidia en masse reddish brown; conidiophores biseriate, stipes dark brown pigmented, smooth (A. haitiensis) or rough-walled (A. sparsus), long; small, malformed, proliferating conidial structures produced near colony surface; Hülle cells absent. Sexual morph unknown. Series description based on Raper & Fennell (1965) and Varga et al. (2010b).



Included species: Aspergillus haitiensis, A. sparsus.



Extrolites: Some extrolites of unknown structure are common for the two species in ser. Sparsi. The only known extrolites detected in this series are gregatins and siderin (found in A. haitiensis NRRL 4569), also found in species in ser. Conjuncti (Varga et al. 2010b).



Notes on sect. Sparsi and included series: Fennell & Raper (1955) created the Aspergillus sparsus group to accommodate four species that possess certain characters, such as pigmented conidiophore stipes and production of smaller conidium forming structures (besides typical aspergillus-like conidiophores). Gams et al. (1985) formally introduced sect. Sparsi for this group of species. Peterson et al. (2008) included seven species in this section (A. anthodesmis, A. biplanus, A. conjunctus, A. diversus, A. funiculosus, A. panamensis, A. sparsus). Later, the taxonomy of this section was studied using a polyphasic approach and the section was expanded with A. amazonicus, A. haitiensis, A. implicatus (Varga et al. 2010b). All species except A. funiculosus (sect. Ochraceorosei, ser. Funiculosi) are accepted in this section in the current study (Fig. 16). The species in ser. Implicati produce white or pale-yellow conidia; a unique feature in sect. Sparsi. Hülle cells are present in some species of ser. Conjuncti and those are predominantly elongate. These resemble the Hülle cells produced by sect. Usti members, rather than the (sub)globose or pyriform Hülle cells in sections Aenei, Cavernicolarum, Nidulantes, Raperorum and Silvatici. Species in series Biplani, Implicati and Sparsi produce reduced, sometimes penicillate, conidiophores.



Section Usti W. Gams et al., Adv. Pen. Asp. Syst.: 58. 1986 [1985]. MycoBank MB832504.



Type: Aspergillus ustus (Bainier) Thom & Church, Aspergilli: 152. 1926.



Description: See Gams et al. (1985) (morphology), Houbraken et al., 2007, Samson et al., 2011b (morphology, phylogeny), Kocsubé et al. (2016) (phylogeny).



Series Calidousti Houbraken & Frisvad, ser. nov. MycoBank MB833264.



Etymology: Named after the type species of this series, A. calidoustus.



Type: Aspergillus calidoustus Varga, Houbraken & Samson, Eukaryot. Cell 7: 636. 2008.



Diagnosis: Phylogeny: Series Calidousti belongs to sect. Usti, subgen. Nidulantes and is phylogenetically sister of ser. Usti (Fig. 16). Morphology & physiology: Colonies growing moderately, conidia en masse greyish yellow, brownish grey, greyish brown or greyish green; conidiophores biseriate, stipes hyaline, (yellow-)brown, smooth; Hülle cells absent (A. germanicus, A. keveioides, A. pseudodeflectus) or present (A. asper, A. calidoustus, A. carlsbadensis, A. contaminans, A. fuscicans, A. keveii, A. thesauricus), generally irregularly elongate, ovoid, curved to coiled, sometimes globose (A. carlsbadensis). Sexual morph unknown. Series description based on Raper and Fennell, 1965, Samson et al., 2011b, Nováková et al., 2012, Visagie et al., 2014a, Jurjević and Peterson, 2016.



Included species: Aspergillus asper, A. calidoustus, A. carlsbadensis, A. contaminans, A. fuscicans, A. germanicus, A. insuetus∗, A. keveii, A. keveioides, A. pseudodeflectus, A. sigurros, A. thesauricus (∗ not included in Fig. 16).



Extrolites: Of the five species examined for extrolites, four produce drimans, two produce asperugins, two produce ophiobolins (G & H), one produces brevianamide A, one produces an arugosin, one produces nidulol and one produces TMC120-B (Samson et al., 2011b, Kozlovsky et al., 2017).



Series Deflecti Houbraken & Frisvad, ser. nov. MycoBank MB833266.



Etymology: Named after the type species of this series, A. deflectus.



Type: Aspergillus deflectus Fennell & Raper, Mycologia 47: 83. 1955.



Diagnosis: Phylogeny: Series Deflecti belongs to sect. Usti, subgen. Nidulantes and is phylogenetically sister to a clade containing series Caldidousti, Monodiorum and Usti (Fig. 16). Morphology & physiology: Colonies grow restrictedly, conidia en masse orange-yellow to grey-green, mycelium on CYA pinkish or orange-shaded; conidiophores biseriate, stipes hyaline or in shades of (reddish) brown or yellowish orange, smooth; Hülle cells absent (A. collinsii, A. lucknowensis) or present (A. deflectus, A. elongatus, A. turkensis), predominantly (irregularly) elongate, straight or curved. Sexual morph unknown. Series description based on Raper and Fennell, 1965, Samson et al., 2011b, Jurjević and Peterson, 2016.



Included species: Aspergillus collinsii, A. deflectus, A. elongatus, A. lucknowensis, A. turkensis.



Extrolites: Of the four species examined for extrolites in this series, two produce emerin, two produce deflectins, two produce shamixanthones, one produces fumitremorgin C, one produces notoamide E and one produces desacetylferriacetylfusigen B (Anke, 1977, Anke et al., 1981, Samson et al., 2011b, Kozlovsky et al., 2017).



Series Monodiorum Houbraken & Frisvad, ser. nov. MycoBank MB833267.



Etymology: Named after the type species of this series, A. monodii.



Type: Aspergillus monodii (Locq.-Lin.) Varga et al., Stud. Mycol. 69: 91. 2011.



Diagnosis: Phylogeny: Series Monodiorum belongs to sect. Usti, subgen. Nidulantes and is phylogenetically sister to a clade containing series Caldidousti and Usti (Fig. 16). Morphology & physiology: Colonies restricted; conidiophores not observed on various media after cultivation at different temperatures; Hülle cells present, surrounding the ascomata, globose to ellipsoidal. Sexual morph fennellia-type, homothallic; ascospores hyaline, with two equatorial ridges, convex smooth. Series description based on Samson et al. (2011b).



Included species: Aspergillus monodii.



Extrolites: Terrein has been detected in A. monodii (Samson et al. 2011b).



Series Usti Houbraken & Frisvad, ser. nov. MycoBank MB833268.



Etymology: Named after the type species of this series, A. ustus.



Type: Aspergillus ustus (Bainier) Thom & Church, Aspergilli: 152. 1926.



Diagnosis: Phylogeny: Series Usti belongs to sect. Usti, subgen. Nidulantes and is phylogenetically sister to a clade containing ser. Caldidousti (Fig. 16). Morphology & physiology: Colonies growing moderately or fast, conidia en masse in shades of brown; conidiophores biseriate, stipes (pale) brown or reddish brown pigmented, smooth and occasionally having warts; Hülle cells generally present (absent in A. pseudoustus), predominantly elongate, occasionally irregularly globose or ovoid (A. granulosus). Sexual morph unknown, except in A. heterothallicus, emericella-type, heterothallic; ascospores orange-brown, with two equatorial ridges, convex smooth. Series description based on Raper and Fennell, 1965, Samson et al., 2011b, Nováková et al., 2012, Visagie et al., 2014a.



Included species: Aspergillus baeticus, A. granulosus, A. heterothallicus, A. porphyreostipitatus, A. pseudoustus, A. puniceus, A. ustus.



Extrolites: Of the six species examined chemically in this series four produce asperugins, three produce ustic acid, three produce nidulol, two produce austocystins, two produce drimans, one produces austamide, one produces austdiol, one produces austocystins (and versicolorins), one produces deoxybrevianamides, one produces emethallicin, one produces emeheterone, one produces emesterone, one produces Mer-NF8054X, one produces ophiobolin G & H, one produces stellatin, and one produces sterigmatocystin (Samson et al. 2011b, and references therein).



Notes on sect. Usti and included series: No asexual morph is observed in ser. Monodiorum and only the sexual morph is produced (A. monodii is homothallic). Sexual reproduction is also present in one member in ser. Usti (A. heterothallicus, heterothallic); all taxa in series Calidousti and Deflecti reproduce strictly asexually. Species classified in series Deflecti and Monodiorum grow more restricted than those in series Calidousti and Usti. Series Calidousti and Usti cannot be differentiated using phenotypic characters and are primary separated based on phylogenetic data. Members of these series grow moderately fast, produce conidia in shades of brown and have biseriate conidiophores. Some species produce Hülle cells or are able to grow at 37 °C, though these characters cannot be linked to a specific series. Future studies might give more insight whether certain phenotypic characters are shared by the taxa of these series.



Aspergillus subgen. Polypaecilum Samson et al., Stud. Mycol. 85: 211. 2016. MycoBank MB819184.



Type: Polypaecilum insolitum G. Sm., Trans. Brit. Mycol. Soc. 44: 437. 1961 (= Aspergillus insolitus).



Description: See Kocsubé et al., 2016, Tanney et al., 2017(morphology, phylogeny),



Section Polypaecilum Houbraken & Frisvad, sect. nov. MycoBank MB833038.



Etymology: Named after the genus Polypaecilum.



Type: Polypaecilum insolitum G. Sm., Trans. Brit. Mycol. Soc. 44: 437. 1961 (= Aspergillus insolitus).



Diagnosis: fideKocsubé et al. (2016): Conidia formed on reduced phialides (as in Phialosimplex salinarum; Greiner et al. 2014, appearing as phialide collula only), small phialides with long collula often with a thickened centre part (like in Phialosimplex caninus; Sigler et al. (2010)) or on polyphialides (as in Polypaecilum insolitum; Smith (1961)), with the common theme of a thin, long collulum producing chains of conidia that are large compared to the diameter of the collulum. Aspergillus conidiophores not produced. The species are halophilic or osmophilic (Wheeler et al., 1988, Wheeler and Hocking, 1993, Greiner et al., 2014, Piñar et al., 2015, Piñar et al., 2016). The sect. Polypaecilum includes species of the previously known genera Polypaecilum and Phialosimplex.



Series Canini Houbraken & Frisvad, ser. nov. MycoBank MB833039.



Etymology: Named after the type of the series, Phialosimplex caninus.



Type: Phialosimplex caninus Sigler et al., Med. Mycol. 48: 338. 2010 (= Aspergillus caninus).



Diagnosis: Phylogeny: Series Canini belongs to sect. Polypaecilum, subgen. Polypaecilum and is phylogenetically sister to ser. Salinarum (Fig. 17). Morphology & physiology: Colonies on CYA restricted, growth on yeast malt extract agar (YMA) moderate, growth on YMA with 20 % NaCl absent, conidia en masse yellowish white to orange grey; conidiogenous cells phialides (monophialidic; sometimes proliferating form a second opening, polyphialidic), simple, borne laterally on the vegetative hyphae or occasionally on short, unbranched conidiophores, 3–16 μm long; conidia in heads or chains; chlamydospores absent (A. caninus) or present (A. chlamydosporus), on short unbranched or branched stalks; growth on MEA at 37 °C. Sexual morph unknown, sclerotia not observed in culture. Xerotolerant and having potential to cause opportunistic disseminated mycoses in dogs. Series description based on Sigler et al., 2010, Martinelli et al., 2017, Tanney et al., 2017.



Fig. 17.

Combined phylogeny for BenA, CaM and RPB2 data sets showing the phylogenetic relation of species and series within Aspergillus subgen. Polypaecilum. The BI posterior probability (pp) values and bootstrap percentages of the maximum likelihood (ML) analysis are presented at the nodes; fully supported branches are thickened. Values less than 70 % bootstrap support (ML) or less than 0.95 posterior probability (Bayesian analysis) are indicated with a hyphen or not shown. The bar indicates the number of substitutions per site. The phylogram is rooted with Hamigera avellanea and Penicillium expansum.

Included species: Aspergillus caninus, A. chlamydosporus.



Extrolites: No extrolites are reported for the species in this series.



Series Kalimarum Houbraken & Frisvad, ser. nov. MycoBank MB833040.



Etymology: Named after the type of the series, A. kalimae.



Type: Aspergillus kalimae Tanney et al., Stud. Mycol. 88: 249. 2017.



Diagnosis: Phylogeny: Series Kalimarum belongs to sect. Polypaecilum, subgen. Polypaecilum and is phylogenetically sister to series Canini and Salinarum (Fig. 17). Morphology & physiology: No growth or at most germination on CYA or MEA, growth on YMA absent or very restricted, optimal growth on YMA with 15 % NaCl, growth present on YMA with 25 % NaCl; conidia en masse white, typical Aspergillus conidiophores lacking; conidiogenous cells monophialidic, polyphialides sometimes present, simple, solitary, borne laterally or terminally on hyphae, 3–30 μm long; conidia solitary, in chains or (small) heads; chlamydospores absent (A. kalimae) or present (A. atacamensis); no growth at 37 °C, halotolerant. Sexual morph unknown; sclerotia absent. Series description based on Martinelli et al., 2017, Tanney et al., 2017.



Included species: Aspergillus atacamensis, A. kalimae.



Extrolites: No extrolites are reported for the species in this series.



Series Noonimiarum Houbraken & Frisvad, ser. nov. MycoBank MB833041.



Etymology: Named after the type of the series, A. noonimiae.



Type: Aspergillus noonimiae Tanney et al., Stud. Mycol. 88: 252. 2017.



Diagnosis: Phylogeny: Series Noonimiarum belongs to sect. Polypaecilum, subgen. Polypaecilum and is phylogenetically sister to ser. Polypaecilum (Fig. 17). Morphology & physiology: Colonies restricted on CYA and MEA, growth absent or only germination on MEA with 20 % NaCl; growth at 37 °C; conidia, when produced, white en masse; typical Aspergillus conidiophores lacking; conidiogenous cells monophialidic, polyphialides sometimes present, simple, solitary, borne laterally or terminally on hyphae, 2.5–25 μm long; conidia solitary or in chains, with persistent membranous sheath. Sexual morph unknown; sclerotia absent or present in culture (A. sclerotialis, A. keratitidis). Series description based on Sigler et al., 2010, Tanney et al., 2017.



Included species: Aspergillus keratitidis, A. noonimiae, A. sclerotialis, A. thailandensis, A. waynelawii.



Extrolites: No extrolites are reported for the species in this series.



Series Polypaecilum Houbraken & Frisvad, ser. nov. MycoBank MB833037.



Etymology: Named after the genus Polypaecilum.



Type: Polypaecilum insolitum G. Sm., Trans. Brit. Mycol. Soc. 44: 437. 1961 (= Aspergillus insolitus).



Diagnosis: Phylogeny: Series Polypaecilum belongs to sect. Polypaecilum, subgen. Polypaecilum and is phylogenetically sister to ser. Noonimiarum (Fig. 17). Morphology & physiology: Colonies restricted on CYA and MEA, growth absent or only germination on MEA with 20 % NaCl; conidia en masse white; typical Aspergillus conidiophores lacking, reproductive structures polyphialides, born solitary on short conidiophores, polyphialides large and complex, 15–60 μm long; chlamydospores absent (A. pisce) or present (A. insolitus). Sexual morph unknown; sclerotia absent (A. insolitus) or present (A. pisce). Isolated from hypersaline habitats. Series description based on Smith, 1961, Pitt and Hocking, 2009, Tanney et al., 2017.



Included species: Aspergillus insolitus, A. pisce.



Extrolites: No extrolites are reported for the species in this series.



Series Salinarum Houbraken & Frisvad, ser. nov. MycoBank MB833042.



Etymology: Named after the type of the series, A. salinarum.



Type: Aspergillus salinarum (Greiner et al.) Zalar & Greiner, Extremophiles 21: 762. 2017.



Diagnosis: Phylogeny: Series Salinarum belongs to sect. Polypaecilum, subgen. Polypaecilum and is phylogenetically sister to ser. Canini (Fig. 17). Morphology & physiology: No growth or only germination on CYA, growth on YMA absent or very restricted, optimal growth on YMA with 15 % NaCl, growth present on YMA with 25 % NaCl; conidia en masse white, typical Aspergillus conidiophores lacking, conidiogenous cells monophialidic, polyphialides absent, simple, solitary, borne laterally or terminally on hyphae, 3–30 μm long; conidia solitary, in chains or (small) heads; chlamydospores absent (A. baarnensis, A. loretoensis) or present (A. salinarum, A. salisburgensis); halotolerant, or (obligate) halophilic. Sexual morph unknown; sclerotia absent. Series description based on Martinelli et al., 2017, Tanney et al., 2017, González-Martínez et al., 2019.



Included species: Aspergillus baarnensis, A. loretoensis∗, A. salinarum, A. salisburgensis [∗not included in Fig. 17; details on classification, see González-Martínez et al. (2019)].



Extrolites: No extrolites are reported for the species in this series.



Series Whitfieldiorum Houbraken & Frisvad, ser. nov. MycoBank MB833043.



Etymology: Named after the type of the series, A. whitfieldii.



Type: Aspergillus whitfieldii Tanney et al., Stud. Mycol. 88: 258. 2017.



Diagnosis: Phylogeny: Series Whitfieldiorum belongs to sect. Polypaecilum, subgen. Polypaecilum and is phylogenetically related to series Canini, Kalimarum and Salinarum; the exact phylogenetic relationship with those series is unresolved (Fig. 17). Morphology & physiology: Colonies restricted, no growth on MEA with 20 % NaCl, no growth at 37 °C; sporulation sparse; typical Aspergillus conidiophores lacking, conidiogenous cells mono- to polyphialidic, solitary, borne laterally or terminally on vegetative hyphae, sometimes occurring in hyphal networks resembling branched conidiophores, 4–36 μm; conidia solitary or in chains, with persistent membranous sheath; chlamydospores absent. Sexual morph unknown; sclerotia not observed in culture. Series description based on Tanney et al. (2017).



Included species: Aspergillus whitfieldii.



Extrolites: No extrolites are reported for the species in this series.



Notes on sect. Polypaecilum and included series: We did not find significant differences within subgen. Polypaecilum that would warrant it to split the subgenus in sections. Section Polypaecilum is therefore the sole section in subgen. Polypaecilum, hence, both have the same description. However, there are few differences within sect. Polypaecilum and we choose to introduce six series (series Canini, Kalimarum, Noonimiarum, Polypaecilum, Salinarum and Whitfieldiorum). Series Polypaecilum forms a strongly supported clade and can be distinguished from the other series by their large and more complexly branched polyphialides. Sclerotia are produced by species in series Noonimiarum (A. sclerotialis, A. keratitidis) and Polypaecilum (A. pisce), and were not observed in the other series. The conidial chains of some species of subgen. Polypaecilum adhered within a persistent membranous sheath visible by SEM and light microscopy. This sheath is present in series Noonimiarum (A. keratitidis, A. noonimiae, A. sclerotialis, A. waynelawii) and Kalimarum (A. kalimae). The production of chlamydospores is distributed over subgen. Polypaecilum and not restricted to any specific series. The two species accommodated in ser. Canini were previously classified in the genus Phialosimplex. These species share the ability to grow at 37 °C and have the potential to cause opportunistic disseminated mycoses in dogs. Furthermore, ser. Canini species are unable to grow on MY10-12, and all other species in subgen. Polypaecilum can (Tanney et al. 2017). These species are xerotolerant, while the members of the phylogenetically related series Salinarum and Kalimarum are considered halotolerant (Sigler et al., 2010, Martinelli et al., 2017, Tanney et al., 2017). The phylogenetic position of A. whitfieldii (the sole species in ser. Whitfieldiorum) is unresolved and more research is needed to determine its exact phylogenetic relationship with the other series of subgen. Polypaecilum (Fig. 17, Tanney et al. 2017).

 Like in many other extremophilic fungi, secondary metabolite production is not common in species of subgen. (and sect.) Polypaecilum. For example, Xeromyces bisporus does not produce a single family of secondary metabolites (Leong et al. 2015). On the other hand, species in sections Aspergillus and Restricti produce a large number of secondary metabolites, as do other species in most of the sections of Aspergillus.



Penicillium subgen. Aspergilloides Dierckx, Ann. Soc. Sci. Bruxelles. 25: 85. 1901. MycoBank MB833420.



Type: Penicillium aurantiobrunneum Dierckx, Ann. Soc. Sci. Bruxelles 25: 86. 1901 (= Penicillium glabrum).



Description: See Houbraken & Samson (2011) (morphology, phylogeny); Fig. 1, this study (phylogeny).



Section Alfrediorum Houbraken & Frisvad, sect. nov. MycoBank MB834239.



Etymology: Named after the type species of the series, Penicillium alfredii.



Type: Penicillium alfredii Visagie et al., Stud. Mycol. 78: 116. 2014.



Diagnosis: Phylogeny: Section Alfrediorum belongs to subgen. Aspergilloides and is phylogenetically most closely related to sect. Lasseniorum (Fig. 2, Fig. 18). Morphology & physiology: Colonies restricted; conidial colour en masse greyish green; conidiophores monoverticillate, smooth; no growth at 37 °C. Sexual morph unknown; sclerotia not produced in culture. Section description based on Visagie et al. (2014a).



Fig. 18.

Combined phylogeny for BenA, CaM and RPB2 data sets showing the phylogenetic relation of species, series and sections within Penicillium subgen. Aspergilloides. The BI posterior probability (pp) values and bootstrap percentages of the maximum likelihood (ML) analysis are presented at the nodes; fully supported branches are thickened. Values less than 70 % bootstrap support (ML) or less than 0.95 posterior probability (Bayesian analysis) are indicated with a hyphen or not shown. The bar indicates the number of substitutions per site. The phylogram is rooted with Aspergillus glaucus and Hamigera avellanea.

Included species: Penicillium alfredii.



Extrolites: No known extrolites detected.



Notes: Penicillium alfredii is the sole species in sect. Alfredii. The phylogenetic position of this species remained unresolved and it could previously not be classified properly in any of the known sections (Visagie et al. 2014a). Penicillium alfredii colonies resemble those of species in sect. Torulomyces; however, sect. Torulomyces species generally produce conidiophores that have solitary phialides and these are not observed in P. alfredii (Visagie et al. 2014a). A subdivision of the section cannot be made and therefore ser. Alfrediorum is only informally introduced here.



Section Aspergilloides Pitt, The Genus Penicillium: 169. 1980 [1979]. MycoBank MB832951.

Synonym: Eupenicillium sect. Pinetorum (Pitt) Stolk & Samson, Stud. Mycol. 23: 88. 1983.



Type: Penicillium aurantiobrunneum Dierckx, Ann. Soc. Sci. Bruxelles 25: 86. 1901 (= Penicillium glabrum).



Description: See Houbraken and Samson, 2011, Houbraken et al., 2014b (phylogeny, morphology).



Series Glabra Pitt, The Genus Penicillium: 169. 1980 [1979]. MycoBank MB832952.



Type: Penicillium glabrum (Wehmer) Westling, Ark. Bot. 11: 131. 1911.



Diagnosis: Phylogeny: Series Glabra belongs to subgen. Aspergilloides, sect. Aspergilloides and forms a unique lineage sister to series Kiamaensia, Spinulosa and Thomiorum (Fig. 18). Morphology & physiology: Colonies spreading on CYA, MEA and YES, texture velvety; conidial colour en masse dark green; conidiophores monoverticillate with vesiculate apex, smooth; conidia ornamented, finely to distinctly rough-walled, globose to subglobose; on CREA often weak growth (except P. armarii) and moderate acid production. Sexual morph unknown; sclerotia not observed in culture. Series description based on Houbraken et al. (2014b).



Included species: Penicillium armarii, P. bussumense, P. frequentans, P. glabrum, P. pulvis, P. purpurescens, P. rudallense.



Extrolites: The very common species P. glabrum (as Citromyces glaber) has been reported to produce citric acid (Wehmer, 1893, Raper and Thom, 1949) and members of the geodin biosynthetic family, including asterric acid, bis-dechlororogeodin, questin, questinol, and sulochrin (Mahmoodian & Stickings 1964). We did not detect any members of the sulochrin biosynthetic family in P. frequentans, but a series of extrolites with unique UV spectra. Penicillium rudallense has been reported to produce a large number of austalides (Wang et al. 2019a), but we detected nigragillin and pyranonigrins in this species. Asterric acid, sulochrin and pyranonigrin A was detected in P. bussumense; P. pulvis produced asterric acid, some red anthraquionones, sulochrin, pyranonigrins and spinulosin.

 Frequentin production was reported from a fungus identified as an atypical strain of P. frequentans CBS 345.51 (Curtis et al. 1951), but this strain is correctly identified as P. subspinulosum (Houbraken et al. 2014b). Species in this series have not been systematically studied chemotaxonomically. Austalides have also been reported in P. thomii in ser. Thomiorum and geodins were reported from P. lividum from ser. Livida (Sobolevskaya et al. 2014), but the identity of those isolates has not been confirmed.



Series Fortuita Houbraken & Frisvad, ser. nov. MycoBank MB834240.



Etymology: Named after the type species of the series, Penicillium fortuitum.



Type: Penicillium fortuitum Visagie & Seifert, Persoonia 41: 387. 2018.



Diagnosis: Phylogeny: Series Fortuita belongs to subgen. Aspergilloides, sect. Aspergilloides and forms a single species lineage sister to ser. Sublectatica, though statistical support for this relationship is lacking. Morphology & physiology: Colonies restricted; conidial colour en masse greyish green; conidiophores predominantly monoverticillate, occasionally divaricate, smooth; no growth at 37 °C. Sexual morph unknown; sclerotia not observed in culture. Series description based on Crous et al. (2018a).



Included species: Penicillium fortuitum.



Extrolites: No chemotaxonomic data on Penicillium fortuitum is known.



Series Hoeksiorum Houbraken & Frisvad, ser. nov. MycoBank MB834241.



Etymology: Named after the type species of the series, Penicillium hoeksii.



Type: Penicillium hoeksii Houbraken, Stud. Mycol. 78: 423. 2014.



Diagnosis: Phylogeny: Series Hoeksiorum belongs to subgen. Aspergilloides, sect. Aspergilloides and is sister to ser. Livida (Fig. 18). Morphology & physiology: Colonies growing moderately; conidial colour en masse grey or blue green; conidiophores monoverticillate, in older parts divaricate, smooth, generally shorter than 250 μm; conidia ellipsoidal, finely rough-walled; no growth at 30 and 37 °C. Sexual morph unknown; sclerotia not observed in culture. Series description based on Houbraken et al. (2014b).



Included species: Penicillium hoeksii, P. zhuangii.



Extrolites: Penicillium hoeksii produces fulvic acids and haenamindole, but no chemotaxonomic data is available for P. zhuangii.



Series Improvisa Houbraken & Frisvad, ser. nov. MycoBank MB834242.



Etymology: Named after the type species of the series, Penicillium improvisum.



Type: Penicillium improvisum Visagie et al., Persoonia 36: 256. 2016.



Diagnosis: Phylogeny: Series Improvisa belongs to subgen. Aspergilloides sect. Aspergilloides and the phylogenetic relationship of this single species is unresolved (Fig. 18). Morphology & physiology: Colonies growing moderately; conidial colour en masse greyish turquoise to greyish green; conidiophores predominantly monoverticillate, sometimes divaricate, smooth; conidia globose to subglobose, smooth; growth at 37 °C absent. Sexual morph unknown; sclerotia not observed in culture. Series description based on Crous et al. (2016).



Included species: Penicillium improvisum.



Extrolites: No chemotaxonomical data on P. improvisum is available.



Series Kiamaensia Houbraken & Frisvad, ser. nov. MycoBank MB834243.



Etymology: Named after the type species of the series, Penicillium kiamaense.



Type: Penicillium kiamaense Houbraken & Pitt, Stud. Mycol. 78: 426. 2014.



Diagnosis: Phylogeny: Series Kiamaensia belongs to subgen. Aspergilloides, sect. Aspergilloides and is, with weak statistical support, sister to series Spinulosa and Thomiorum (Fig. 18). Morphology & physiology: Colonies moderate to spreading; conidial colour en masse dark green; conidiophores predominantly monoverticillate, occasionally with a short branch, finely rough-walled; conidia globose or subglobose, ornamented with striations; growth at 30 °C, absent at 37 °C. Sexual morph unknown; sclerotia not observed in culture. Series description based on Houbraken et al. (2014b).



Included species: Penicillium kiamaense.



Extrolites: Penicillium kiamense produces some anthraquinones and some extrolites that are not yet structure elucidated.



Series Livida Houbraken & Frisvad, ser. nov. MycoBank MB834244.



Etymology: Named after the type species of the series, Penicillium lividum.



Type: Penicillium lividum Westling, Ark. Bot. 11: 134. 1911.



Diagnosis: Phylogeny: Series Livida belongs to subgen. Aspergilloides, sect. Aspergilloides and is sister to ser. Hoeksiorum (Fig. 18). Morphology & physiology: Colonies growing moderately; conidial colour en masse (dark) blue-green; conidiophores monoverticillate, rough-walled, longer than 250 μm; conidia broadly ellipsoidal or ellipsoidal, distinctly roughened, often striate; growth at 30 °C, absent at 37 °C. Sexual morph unknown; sclerotia not produced in culture. Series description based on Pitt (1980) and Houbraken et al. (2014b).



Included species: Penicillium kananaskense, P. lividum, P. odoratum.



Extrolites: Penicillium odoratum has been reported to produce citrinin (Nakajima & Nozawa 2004), and this has been confirmed here. Penicillium kananaskense produces citreoisocoumarins and a large number of unique extrolites that are not yet structure elucidated. Penicillium lividum has been reported to produce austalides, daldinin D, peniciraistin C, questin and sulochrin (Sobolevskaya et al., 2014, Zhuravleva et al., 2014a, Zhuravleva et al., 2014b, Sobolevskaya et al., 2016b). We could not confirm this, but found some unique not yet structure elucidated extrolites for P. lividum. The three species in ser. Livida have no extrolites in common.



Series Longicatenata Houbraken & Frisvad, ser. nov. MycoBank MB834245.



Etymology: Named after the type species of the series, Penicillium longicatenatum.



Type: Penicillium longicatenatum Visagie et al., Stud. Mycol. 78: 429. 2014.



Diagnosis: Phylogeny: Series Longicatenata belongs to subgen. Aspergilloides, sect. Aspergilloides; the two species in the series are related in the phylogeny of Houbraken et al. (2014b) with moderate statistical support (81 % BS, 0.96 pp); however, their relationship is unresolved in this study (Fig. 18). Morphology & physiology: Colonies growing moderate or spreading; conidial colour en masse in shades of green (greyish green, dull green or dark green); colony texture velvety or floccose; conidiophores monoverticillate, smooth; conidia (sub)globose, finely or distinctly rough-walled; generally no growth at 37 °C or sometimes restricted growth (<4 mm, 7 d, CYA). Sexual morph unknown, sclerotia absent (P. vagum) or present (P. longicatenatum). Series description based on Houbraken et al. (2014b).



Included species: Penicillium longicatenatum, P. vagum.



Extrolites: Penicillium longicatenatum produced kotanins, palitantin, spinulosin and a versicolorin, while the ex-type culture of P. vagum produces citrinin. Other strains supposed to be P. vagum were chemotaxonomically very different: DTO 038-E7 and DTO 056-I6 produced spinulosin and daldinins, while DTO 120-B1 and DTO 120-B4 produced asperfuran, dehydrocarolic acid, frequentin and palitantin. This indicates that P. vagum may be split into three species.



Series Pinetorum (Pitt) Houbraken & Frisvad, comb. nov. MycoBank MB834246.

Basionym: Eupenicillium ser. Pinetorum Pitt, The Genus Penicillium: 105. 1980 [1979]. MycoBank MB832947.



Type: Eupenicillium pinetorum Stolk, Antonie van Leeuwenhoek 34: 37. 1968 (= Penicillium fuscum).



Diagnosis: Phylogeny: Series Pinetorum belongs to subgen. Aspergilloides, sect. Aspergilloides and is unresolved, but forms a sister lineage to series Glabra, Kiamaensia, Spinulosa and Thomiorum (Fig. 18). Morphology & physiology: Colonies growing restrictedly or moderately rapid; conidial colour en masse variable (in shades of green); conidiophores monoverticillate, generally smooth, sometimes rough-walled (P. clavistipitatum), short; conidia globose to subglobose, distinctly rough-walled, thick; no growth at 37 °C. Sexual morph generally not produced in culture, except for P. fuscum, eupenicillium-type, homothallic, greyish yellow; ascospores ellipsoidal, with two longitudinal flanges, convex smooth-walled; sclerotia generally absent, except in P. flavisclerotiatum (yellow) and P. tsitsikammaense (white). Series description based on Pitt (1980) and Houbraken et al. (2014b).



Included species: Penicillium ardesiacum, P. athertonense, P. brunneoconidiatum, P. clavistipitatum, P. flavisclerotiatum, P. fuscum, P. montanense, P. tsitsikammaense, P. turcosoconidiatum.



Extrolites: The species in ser. Pinetorum are chemotaxonomically quite different. Penicillium athertonense produces viridicatins, P. flavisclerotiatum DTO 184-D8 produces burnettienes, while the isolates DTO 180-I1 and DTO 181-I9 produce asperfuran. Penicillium fuscum produces asperfuran, while isolates allocated to P. lapatayae produce kotanins, lapatins and spinulosins indicating P. lapatayae is a separate species. However, we follow Houbraken et al. (2014b) and consider this species as a synonym of P. fuscum based on molecular data. Penicillium montanense produces unique extrolites, while P. turcosoconidiatum (DTO 181-A3) produces citreoviridin and haenamindole.



Series Quercetorum Houbraken & Frisvad, ser. nov. MycoBank MB834247.



Etymology: Named after the type species of the series, Penicillium quercetorum.



Type: Penicillium quercetorum Baghd., Novosti Sist. Nizsh. Rast. 5: 110. 1968.



Diagnosis: Phylogeny: Series Quercetorum belongs to subgen. Aspergilloides, sect. Aspergilloides and is sister to series Hoeksiorum and Livida (Fig. 18). Morphology & physiology: Colonies growing moderately; conidial colour en masse dull green; conidiophores monoverticillate, smooth-walled; conidia globose, smooth; growth at 37 °C absent. Sexual morph not observed in culture; sclerotia orange-brown. Series description based on Houbraken et al. (2014b).



Included species: Penicillium quercetorum.



Extrolites: No chemotaxonomic data available for P. quercetorum.



Series Saturniformia Houbraken & Frisvad, ser. nov. MycoBank MB834248.



Etymology: Named after the type species of the series, Penicillium saturniforme.



Type: Penicillium saturniforme (L. Wang & W.Y. Zhuang) Houbraken & Samson, Stud. Mycol. 70: 48. 2011.



Diagnosis: Phylogeny: Series Saturniformia belongs to subgen. Aspergilloides, sect. Aspergilloides and is sister to ser. Verhageniorum (Fig. 18). Morphology & physiology: Colonies growing moderate; conidial colour en masse green to greyish olive; conidiophores biverticillate; no growth on CYA incubated at 30 °C; conidia (broadly) ellipsoidal, finely rough-walled. Sexual morph eupenicillium-type, homothallic, pinkish brown; ascospores ellipsoidal, with two very closely appressed equatorial ridges, convex smooth with sparsely scattered fine warts or irregular ribs along the outer areas. Series description based on Wang & Zhuang (2009) and Houbraken et al. (2014b).



Included species: Penicillium saturniforme.



Extrolites: No chemotaxonomic data available for P. saturniforme.



Series Spinulosa Houbraken & Frisvad, ser. nov. MycoBank MB834249.



Etymology: Named after the type species of the series, Penicillium spinulosum.



Type: Penicillium spinulosum Thom, U.S.D.A. Bur. Animal Industr. Bull. 118: 76. 1910.



Diagnosis: Phylogeny: Series Spinulosa belongs to subgen. Aspergilloides, sect. Aspergilloides and is sister to ser. Thomiorum (Fig. 18). Colonies spreading on CYA, MEA and YES, texture (slightly) floccose; conidial colour en masse mostly pure or dull green; conidiophores monoverticillate with vesiculate apex, smooth; conidia ornamented, finely to distinctly rough-walled, globose to subglobose; on CREA poor or good growth, acid production often absent or poor. Sexual morph unknown; sclerotia not observed in culture. Series description based on Houbraken et al. (2014b).



Included species: Penicillium grancanariae, P. palmense, P. roseomaculatum, P. spinulosum, P. sterculiniicola, P. subspinulosum, P. trzebinskii.



Extrolites: Penicillium spinulosum has been reported to produce spinulosin (Birkinshaw and Raistrick, 1931, Anslow and Raistrick, 1938, Pettersson, 1965). The original producing strain, IMI 091950, did not match the description of P. spinulosum (results reported here). Penicillium subspinulosum produces frequentin and palitantin (Houbraken et al. 2014b), in common with P. trzebinskii; P. spinulosum produces asperfuran, while P. sterculiniicola produces asperfuran and 12,13-deoxybrevianamide E.



Series Sublectatica Houbraken & Frisvad, ser. nov. MycoBank MB834250.



Etymology: Named after the type species of the series, Penicillium sublectaticum.



Type: Penicillium sublectaticum Houbraken et al., Stud. Mycol. 78: 436. 2014.



Diagnosis: Phylogeny: Series Sublectatica belongs to subgen. Aspergilloides, sect. Aspergilloides and is sister to ser. Fortuita, though statistical support for this relationship is lacking (Fig. 18). Morphology & physiology: Colonies growing moderately fast or spreading; conidial colour en masse in various shades of green (greyish green, dark green, dull green); conidiophores monoverticillate; growth on CYA incubated at 30 °C, no growth at 37 °C. Sexual morph unknown; sclerotia not observed in culture. Series description based on Houbraken et al. (2014b).



Included species: Penicillium infra-aurantiacum, P. malmesburiense, P. sublectaticum.



Extrolites: Penicillium infra-aurantiacum produces citrinin, while P. malmesburiense produces unique extrolites that have not been structure elucidated yet.



Series Thiersiorum Houbraken & Frisvad, ser. nov. MycoBank MB834251.



Etymology: Named after the type species of the series, Penicillium thiersii.



Type: Penicillium thiersii S.W. Peterson et al., Mycologia 96: 1283. 2004.



Diagnosis: Phylogeny: Series Thiersiorum belongs to subgen. Aspergilloides, sect. Aspergilloides and is sister to all other series in the section (Fig. 18). Morphology & physiology: Colonies spreading; conidial colour en masse dark bluish grey; conidiophores monoverticillate, smooth or slightly roughened; conidia ellipsoidal, smooth; growth absent at 37 °C. Sexual morph eupenicillium-type, homothallic, pale brown; ascospores ellipsoidal, with equatorial ridge, smooth or finely roughened convex. Series description based on Peterson et al. (2004) and Houbraken et al. (2014b).



Included species: Penicillium thiersii.



Extrolites: Penicillium thiersii produces thiersinines and their precursors such as 1’-O-acetylpaxilline, dehydroxypaxilline, paxilline, paspaline, PC-M5’ and PC-M6 (Li et al. 2002), decaturin B, C and D, 15-deoxyoxalicine A, oxalicine A and B (Zhang et al., 2003, Li et al., 2005) and emindole SB and thiersindole A-C (Li et al., 2002, Li et al., 2003).



Series Thomiorum Houbraken & Frisvad, ser. nov. MycoBank MB834252.



Etymology: Named after the type species of the series, Penicillium thomii.



Type: Penicillium thomii Maire, Bull. Soc. Hist. Nat. Afrique N. 8: 189. 1917.



Diagnosis: Phylogeny: Series Thomiorum belongs to subgen. Aspergilloides, sect. Aspergilloides and is sister to ser. Spinulosa (Fig. 18). Morphology & physiology: Colonies spreading on CYA, MEA and YES; conidial colour en masse dull green; conidiophores monoverticillate with a vesiculate apex, rough-walled, conidia ellipsoidal or fusiform; growth on CYA incubated at 30 °C, (5–)15–35(–45), no growth at 37 °C. Sexual morph unknown; sclerotia commonly produced, in shades of pink (orange-pink, brownish pink). Series description based on Houbraken et al. (2014b).



Included species: Penicillium aurantioviolaceum, P. austroafricanum, P. cartierense, P. contaminatum, P. crocicola, P. fusisporum, P. grevilleicola, P. jejuense, P. roseoviride, P. thomii, P. valentinum, P. yezoense.



Extrolites: Penicillium thomii has been reported to produce N-acetylphenylalaninol and the related 2(S)-acetamido-3-phenylpropylacetate, austalides, furan-2-carboxylic acid derivatives, guaidiol A and 4,10,11-trihydroxyguaiane, pallidopenillines, penistinraistin C and the related daldinin D, sargassopenillines, thomimarides, thomimarines, VM55599 and zesteropenillines (Sobolevskaya et al., 2014, Zhuravleva et al., 2014a, Zhuravleva et al., 2014b, Sobolevskaya et al., 2016a, Sobolevskaya et al., 2016b, Sobolevskaya et al., 2018, Afiyatullov et al., 2017a, Afiyatullov et al., 2017b, Afiyatullov et al., 2018). Penicillium austroafricanum produces fumagillin and P. auirantioviolaceum produces spinulosins. Five species in series Thomiorum produce haenamindole: P. cartierense, P. contaminatum, P. crocicola, P. roseoviride and P. yezoense; P. contaminatum also produces palitantin.



Series Verhageniorum Houbraken & Frisvad, ser. nov. MycoBank MB834253.



Etymology: Named after the type species of the series, Penicillium verhagenii.



Type: Penicillium verhagenii Houbraken, Stud. Mycol. 78: 443. 2014.



Diagnosis: Series Verhageniorum belongs to subgen. Aspergilloides, sect. Aspergilloides and is sister to ser. Saturniformia (Fig. 18). Morphology & physiology: Colonies growing moderately; conidial colour en masse variable, in green shades with a blue element; conidiophores biverticillate, sometimes becoming divaricate by sympodial branching of the stipe at the apex, smooth or finely roughened; conidial shape variable, rough-walled; no growth on CYA incubated at 30 °C Sexual morph unknown; sclerotia not observed in culture. Series description based on Houbraken et al. (2014b).



Included species: Penicillium ranomafanaense, P. verhagenii.



Extrolites: Penicillium ranomafanaense produces andrastin A, asterric acid, fulvic acids, gregatins and geodin; P. verhagenii produces a quinone of unknown structure.



Notes on sect. Aspergilloides and included series: Section Aspergilloides was introduced by Pitt (1980) to accommodate Penicillium species that predominantly produce monoverticillate conidiophores in which at least a portion of the stipes terminate in vesicular swellings. The phenotype-based infrageneric classification systems proposed in Penicillium are generally loosely corresponding with those based on phylogenetic inference using sequence data. Houbraken & Samson (2011) proposed a sectional classification system based on the phylogenetic analysis of a combined four-gene dataset and re-circumscribed section Aspergilloides. The majority of species belonging to this re-circumscribed section grow moderately or fast on agar media and are predominantly monoverticillate. The section was subsequently revised, and the 51 accepted species were distributed over 12 clades (Houbraken et al. 2014b). These clades are here treated as series. The relationship of P. kiamaense was unresolved and was therefore not accommodated in a clade; we introduced ser. Kiamaensia for this species. After 2014, three new species were described in sect. Aspergilloides (P. fortuitum, P. improvisum, and P. jejuense). The former two species form unique lineages in the section and are therefore accommodated in unique, separate series (Fortuita, Improvisa).

 The series classification is primarily based on the phylogenetic relationships of the species within the section, and this is often supported by morphology and physiology data. Growth rate, the ability to grow at 30 °C, conidiophore branching pattern and conidial shape and ornamentation were useful characters to differentiate the series of sect. Aspergilloides. The phylogenetic support was low or absent for series Fortuita, Improvisa and Kiamaensia. The phylogenetic distance of the former two series was sufficient to accommodate them in separate series. In addition, ser. Fortuita grows restrictedly, a feature shared with the distantly related ser. Pinetorum. Series Kiamaensia is a sister series of series Spinulosa and Thomiorum, though statistical support for this is weak. Series Spinulosa and Thomiorum are phylogenetically and phenotypically distinct. Series Kiamaensia is introduced in order to maintain monophyletic series. The relationship between the two species in ser. Longicatenata is moderately supported in the phylogram of Houbraken et al. (2014b). These species are phenotypically unrelated and this suggests that they might belong to two separate series. The discovery of more species related to this clade might show that they are actually more than one series; however, we prefer at this moment a conservative approach and maintain both species in one series.



Section Charlesia Houbraken & Samson, Stud. Mycol. 70: 33. 2011. MycoBank MB563125.



Type: Penicillium charlesii G. Sm., Trans. Brit. Mycol. Soc. 18: 90. 1933.



Description: See Peterson et al. (2005) and Houbraken & Samson (2011) (morphology, phylogeny); a modern taxonomic study on this section is lacking.



Series Costaricensia Houbraken & Frisvad, ser. nov. MycoBank MB834254.



Etymology: Named after the type species of the series, Penicillium costaricense.



Type: Penicillium costaricense Visagie et al., Persoonia 36: 263. 2016.



Diagnosis: Series Costaricensia belongs to subgen. Aspergilloides, sect. Charlesia and is sister to series Fellutana, Indica and Phoenicea. Morphology & physiology: Colonies restricted; conidial colour en masse turquoise to dull green; conidiophores monoverticillate, smooth; conidia subglobose, smooth-walled; growth at 37 °C absent. Sexual morph unknown; sclerotia not observed in culture. Series description based on Visagie et al. (2016b).



Included species: Penicillium costaricense.



Extrolites: Andrastin A & C (Visagie et al. 2016b).



Series Fellutana Pitt, The Genus Penicillium: 263. 1980 [1979]. MycoBank MB832961.



Type: Penicillium fellutanum Biourge, Cellule 33: 262. 1923.



Diagnosis: Series Fellutana belongs to subgen. Aspergilloides, sect. Charlesia and is sister to ser. Indica (Fig. 18). Morphology & physiology: Colonies growing restricted; conidial colour en masse dark green; conidiophores monoverticillate or furcate, smooth; conidia globose or ellipsoidal, finely or distinctly rough-walled; growth at 37 °C absent. Sexual morph unknown; sclerotia not observed in culture. Series description based on Pitt (1980) and Peterson et al. (2005).



Included species: Penicillium charlesii, P. fellutanum.



Extrolites: Penicillium charlesii in ser. Fellutana produces carolic acids (Clutterbuck et al., 1934, Clutterbuck et al., 1935a, Clutterbuck et al., 1935b, Clutterbuck et al., 1935c), an uracil nucleoside (Maynard & Gander 1966) and exopolysaccharides (Haworth et al. 1935). Penicillium fellutanum has been reported to produce different secondary metabolites such as fellutamides (Shigemori et al. 1991), fellutanine A-E and isofellutanine B & C (Kozlovsky et al., 1997a, Kozlovsky et al., 1997b, Kozlovsky et al., 2000), cyclosporine (Anjum et al. 2012) and peniphenylanes (Zhang et al. 2016).



Series Indica Houbraken & Frisvad, ser. nov. MycoBank MB834255.



Etymology: Named after the type species of the series, Penicillium indicum.



Type: Penicillium indicum D.K. Sandhu & R.S. Sandhu, Canad. J. Bot. 41: 1273. 1963.



Diagnosis: Series Indica belongs to subgen. Aspergilloides, sect. Charlesia and is sister to ser. Fellutana (Fig. 18). Morphology & physiology: Colonies growing moderately fast or spreading; conidial colour en masse dull green or grey green; conidiophores predominantly monoverticillate, conspicuously vesiculate, smooth; conidia subglobose to ellipsoidal, smooth-walled; growth at 37 °C present (P. chermesinum, P. indicum) or absent (P. lunae). Sexual morph unknown; sclerotia produced in P. indicum, white to cream. Series description based on Pitt (1980) and Crous et al. (2019).



Included species: Penicillium chermesinum, P. cuddlyae∗, P. indicum, P. lunae∗ [∗not included in Fig. 18; more info on their phylogenetic relationship, see Crous et al. (2019)].



Extrolites: Penicillium chermesinum is reported to produce chermesins (Liu et al. 2016b), penicilliumolides (Darsih et al. 2015), PR-toxins (Darsih et al. 2015), chermesinones and terphenyllins (Huang et al. 2011) and costaclavin (Agurell 1964), and also to secrete the ribotoxins proteins (Hwu et al. 2001).



Series Phoenicea Houbraken & Frisvad, ser. nov. MycoBank MB834256.



Etymology: Named after the type species of the series, Penicillium phoeniceum.



Type: Penicillium phoeniceum J.F.H. Beyma, Zentralbl. Bakteriol. Parasitenk., Abt. 2 88: 136. 1933.



Diagnosis: Series Phoenicea belongs to subgen. Aspergilloides, sect. Charlesia and is sister to series Fellutana and Indica (Fig. 18). Morphology & physiology: Colonies growing restrictedly or moderately rapid; conidial colour en masse dull green or dull greyish blue; conidiophores monoverticillate, vesiculate, smooth; conidia globose, smooth-walled; growth at 37 °C present (P. phoeniceum) or absent (P. coffeae). Sexual morph unknown; sclerotia not observed in culture. Series description based on Pitt (1980) and Peterson et al. (2005).



Included species: Penicillium coffeae, P. phoeniceum.



Extrolites: Penicillium phoenicum has been reported to produce phoenicin (Friedheim, 1938, Posternak, 1938, Curtin et al., 1940, Posternak et al., 1943, Steiner et al., 1974).



Notes on series of sect. Charlesia: Peterson et al. (2005) studied the phylogenetic relationship of P. coffeae within the genus Penicillium. They showed that this species is related to P. charlesii, P. chermesinum, P. coffeae, P. fellutanum, P. indicum and P. phoeniceum; all species currently classified in sect. Charlesia. The phenotypic similarity between P. charlesii, P. fellutanum (ser. Fellutana), P. chermesinum, P. indicum (ser. Indica), and P. coffeae and P. phoeniceum (ser. Phoenicea) was also indicated and these groups of species (here treated as series) could be distinguished using colony growth rates and conidiophore complexity. Penicillium lunae and P. costaricense were described after Peterson et al. (2005); the former species belongs to ser. Indica and the latter represents a single species series.



Section Cinnamopurpurea Houbraken & Samson, Stud. Mycol. 70: 34. 2011. MycoBank MB563128.



Type: Penicillium cinnamopurpureum Udagawa, J. Agric. Food Sci., Tokyo 5: 1. 1959.



Description: See Houbraken and Samson, 2011, Peterson et al., 2015 (morphology, phylogeny).



Series Cinnamopurpurea Houbraken & Frisvad, ser. nov. MycoBank MB834257.



Etymology: Named after the type species of the series, Penicillium cinnamopurpureum.



Type: Penicillium cinnamopurpureum Udagawa, J. Agric. Food Sci., Tokyo 5: 1. 1959.



Diagnosis: Phylogeny: Series Cinnamopurpurea belongs to subgen. Aspergilloides, sect. Cinnamopurpurea and is a sister series of Idahoensia and Nodula (Fig. 18). Morphology & physiology: Colonies restricted; conidial colour en masse blue-green, grey-green or pale green; conidiophores monoverticillate; stipes smooth, short, often less than 50 μm in length; conidia globose to subglobose, sometimes (broadly) ellipsoidal, smooth; growth at 37 °C absent (P. gravinicasei, P. parvulum) or present (P. cinnamopurpureum). Sexual morph not observed in culture (P. gravinicasei, P. parvulum) or present (P. cinnamopurpureum), eupenicillium-type, homothallic, pinkish cinnamon to brown; ascospores ellipsoidal, with two close equatorial ridges, valves (finely) rough-walled, warted viewed by SEM; sclerotia not observed in culture. Series description based on Pitt, 1980, Stolk and Samson, 1983, Peterson and Horn, 2009 and Anelli et al. (2018).



Included species: Penicillium cinnamopurpureum, P. gravinicasei, P. parvulum.



Extrolites: Penicillium cinnamopurpureum and P. parvulum produce some red anthraquinones of unknown structure.



Series Idahoensia Houbraken & Frisvad, ser. nov. MycoBank MB834258.



Etymology: Named after the type species of the series, Penicillium idahoense.



Type: Penicillium idahoense Paden, Mycopathol. Mycol. Appl. 43: 259. 1971.



Diagnosis: Phylogeny: Series Idahoensia belongs to subgen. Aspergilloides, sect. Cinnamopurpurea and is sister to ser. Nodula (Fig. 18). Morphology & physiology: Colonies restricted; conidial colour en masse in shades of green, grey-green, blue-green or pale green; conidiophores monoverticillate, occasionally with an additional branch; stipes short, smooth, often vesiculate; conidia varying from globose to ellipsoidal, mostly smooth or finely roughened, sometimes conspicuously spinulose (P. malacaense); growth at 37 °C generally absent, sometimes present (P. idahoense, (Visagie et al. 2014a). Sexual morph generally not observed in culture, only present in P. idahoense, eupenicillium-type, homothallic, (dark) brown; ascospores ellipsoidal, with two close equatorial ridges, valves (finely) smooth-walled under light microscope, warted viewed by SEM; sclerotia absent or present (P. fluviserpens, P. lemhiflumine), brown. Series description based on Paden, 1971, Ramírez, 1982, Stolk and Samson, 1983, Visagie et al., 2014a and Peterson et al. (2015).



Included species: Penicillium colei, P. cvjetkovicii, P. ellipsoideosporum, P. fluviserpens, P. idahoense, P. infrapurpureum, P. lemhiflumine, P. malacaense, P. minnesotense∗, P. monsgalena, P. monsserratidens, P. salmoniflumine [∗ not included in Fig. 18, for details on their phylogenetic relationship, see Crous et al. (2019)].



Extrolites: Red anthraquinones possibly related to roseopurpurin (Peterson et al. 2015); P. colei and P. monsserratidens produce citreoviridin (Peterson et al. 2015).



Series Jiangxiensia Houbraken & Frisvad, ser. nov. MycoBank MB834259.



Etymology: Named after the type species of the series, Penicillium jiangxiense.



Type: Penicillium jiangxiense H.Z. Kong & Z.Q. Liang, Mycosystema 22: 4. 2003.



Diagnosis: Phylogeny: Series Jiangxiensia belongs to subgen. Aspergilloides, sect. Cinnamopurpurea and is sister to the other series of sect. Cinnamopurpurea. Morphology & physiology: Colonies growing slowly; sporulation poor, conidial colour en masse grey-green, blueish grey; conidiophores predominantly monoverticillate, occasionally with an additional branch, stipes smooth; conidia globose to subglobose or ellipsoidal, smooth; growth at 37 °C reported in P. jiangxiense. Sexual morph unknown; sclerotia absent (P. jiangxiense) or present (P. pusillum), brownish. Series description based on Smith, 1939, Pitt, 1980 and Kong & Liang (2003).



Included species: Penicillium jiangxiense, P. pusillum.



Extrolites: No chemotaxonomic data available for these species.



Series Nodula Houbraken & Frisvad, ser. nov. MycoBank MB834260.



Etymology: Named after the type species of the series, Penicillium nodulum.



Type: Penicillium nodulum H.Z. Kong & Z.T. Qi, Mycosystema 1: 108. 1988.



Diagnosis: Phylogeny: Series Nodula belongs to subgen. Aspergilloides, sect. Cinnamopurpurea and is phylogenetically sister to ser. Idahoensia (Fig. 18). Morphology & physiology: Colonies growing restricted; conidial colour en masse dull green to olive green (P. nodulum, P. shennongjianum) or uncoloured (P. incoloratum); conidiophores predominantly monoverticillate, occasionally branched; stipes smooth, short, less than 50 μm in length; conidia globose to subglobose (P. incoloratum, P. shennongjianum) or ellipsoidal (P. nodulum), smooth-walled; growth at 37 °C absent. Sexual morph unknown; sclerotia not produced in culture. These species are to date only reported from China. Series description based on Kong & Qi (1988) and Huang & Qi (1994).



Included species: Penicillium incoloratum, P. nodulum, P. shennongjianum.



Extrolites: Penicillium nodulum produces griseofulvin, but there are no chemotaxonomic data available for the other species in ser. Nodula.



Notes on series of sect. Cinnamopurpurea: Peterson et al. (2015) studied the species within sect. Cinnamopurpurea. They noted that this group of species are morphologically quite similar, all producing subglobose to ellipsoidal smooth to finely roughened spores, monoverticillate to divaricate biverticillate smooth-walled conidiophores and quite slow-growing colonies, often with a brown reverse on some media (Peterson et al. 2015). It is difficult to find good characters to delimit series in this section and the current series classification is therefore based on phylogenetic data.



Section Citrina Houbraken & Samson, Stud. Mycol. 70: 40. 2011. MycoBank MB563132.



Type: Penicillium citrinum Thom, U.S.D.A. Bur. Animal Industr. Bull. 118: 61. 1910.



Description: See Houbraken & Samson (2011) and Houbraken et al. (2011a) (morphology, phylogeny).



Series Citrina Raper & Thom ex Pitt, The Genus Penicillium: 290. 1980 [1979]. MycoBank MB832965.

Synonym: Penicillium ser. Implicata Raper & Thom ex Pitt, The Genus Penicillium: 191. 1980 [1979].



Type: Penicillium citrinum Thom, U.S.D.A. Bur. Animal Industr. Bull. 118: 61. 1910.



Diagnosis: Phylogeny: Series Citrina belongs to subgen. Aspergilloides, sect. Citrina, and the phylogenetic relationship with other series is unknown (Fig. 18). Morphology & physiology: Colonies growing moderately or fast; conidial colour en masse variable, (blueish) grey-green, dull green or pure green; conidiophores biverticillate, smooth; conidia globose, subglobose or broadly ellipsoidal, smooth or finely roughened; growth at 37 °C variable (absent: P. gorlenkoanum, P. steckii, P. tropicoides, P. tropicum; variable: P. hetheringtonii, P. sizovae; present: P. citrinum). Sexual morph unknown, or present (P. tropicoides, P. tropicum), eupenicillium-type, orange-tan, becoming (brownish) grey; ascospores ellipsoidal, with two narrow closely appressed equatorial ridges, convex smooth or finely roughened; sclerotia absent. Series description based on Houbraken et al., 2010b, Houbraken et al., 2011a.



Included species: Penicillium citrinum, P. gorlenkoanum, P. hetheringtonii, P. sizovae, P. steckii, P. tropicoides, P. tropicum.



Extrolites: Quinolactacin is produced by 4/7 species in series Citrina (P. citrinum, P. heteringtonii, P. sizovae, P. steckii), citrinin is produced by 3/7 species (P citrinum, P. gorlenkoanum, P. heteringtonii), citriquinones by 3/7 species (P. gorlenkoanum, P. citrinum, P. steckii), citrinalin by 2/7 species (P. hetheringtonii, P. tropicoides), isochromantoxins by 2/7 species (P. steckii, P. tropicoides), tanzawaic acid by 2/7 species (P. sizovae, P. steckii), and chanoclasvin by 1/7 species (P. gorlenkoanum) (Houbraken et al., 2011a, El-Neketi et al., 2013, Lai et al., 2013, Ranji et al., 2013).



Series Copticolarum Houbraken & Frisvad, ser. nov. MycoBank MB834261.



Etymology: Named after the type species of the series, Penicillium copticola.



Type: Penicillium copticola Houbraken et al., Stud. Mycol. 70: 88. 2011.



Diagnosis: Phylogeny: Series Copticolarum belongs to subgen. Aspergilloides, sect. Citrina, and is phylogenetically related to ser. Sumatraensia, though without statistical support (Fig. 18). Morphology & physiology: Colonies growing moderately; conidial colour en masse dull (grey-)green or pure green; conidiophores symmetrically biverticillate, smooth, finely or distinctly rough-walled; conidia broadly ellipsoidal, smooth; growth present at 30 °C (up to 23 mm), absent at 37 °C. Sexual morph unknown; sclerotia not observed in culture. Series description based on Houbraken et al. (2011a).



Included species: Penicillium copticola, P. dokdoense, P. terrigenum.



Extrolites: Penicillium copticola produces sporogen AO1 and related terpenes in addition to penicillimides / penicillithiophenols and chimeric products of those families of secondary metabolites (Bu et al., 2015, Daengrot et al., 2015).



Series Euglauca Houbraken & Frisvad, ser. nov. MycoBank MB834262.



Etymology: Named after the type species of the series, Penicillium euglaucum.



Type: Penicillium euglaucum J.F.H. Beyma, Antonie van Leeuwenhoek 6: 269. 1940.



Diagnosis: Phylogeny: Series Euglauca belongs to subgen. Aspergilloides, sect. Citrina; however, the phylogenetic relationship within this section is unresolved (Fig. 18). Morphology & physiology: Colonies growing restrict or moderately fast; conidial colour en masse grey-green; conidiophores predominant monoverticillate or biverticillate, smooth; conidia globose to subglobose, smooth or finely roughened; growth at 37 °C absent or present. Sexual morph eupenicillium-type, brown or brownish grey; ascospores ellipsoidal, with two appressed distinct ridges, convex slightly roughened with warts and small ridges (P. anatolicum) or reticulate (P. argentinense, P. euglaucum). Series description based on Houbraken et al. (2011a).



Included species: Penicillium anatolicum, P. argentinense, P. euglaucum, P. vascosobrinhous (recently described species, not included in Fig. 18).



Extrolites: Curvularin and dehydrocurvularin are produced by P. anatolicum and P. argentinense. Furthermore, P. anatolicum produces anthraquinones, bisanthrons and sorbicillins, and P. euglaucum produces terrain (Houbraken et al. 2011a).



Series Gallaica Houbraken & Frisvad, ser. nov. MycoBank MB834263.



Etymology: Named after the type species of the series, Penicillium gallaicum.



Type: Penicillium gallaicum Ramírez et al., Mycopathol. 72: 30. 1980.



Diagnosis: Phylogeny: Series Gallaica belongs to subgen. Aspergilloides, sect. Citrina and is phylogenetically basal to other series of sect. Citrina (Fig. 18). Morphology & physiology: Colonies growing moderately fast; conidial colour en masse dull or pale grey-green; conidiophores monoverticillate, occasionally with additional branch, smooth, short; conidia globose or subglobose, smooth; growth at 37 °C absent or very restricted (5 mm). Sexual morph unknown; sclerotia produced by P. gallaicum, orange-brown. Series description based on Houbraken et al. (2011a).



Included species: Penicillium gallaicum.



Extrolites: Citreoviridin, and several uncharacterised compounds unique for this series in sect. Citrina (“KOKSO”, “3-S”, “VYL”) (Houbraken et al. 2011a).



Series Paxillorum Houbraken & Frisvad, ser. nov. MycoBank MB834264.



Etymology: Named after the type species of the series, Penicillium paxilli.



Type: Penicillium paxilli Bainier, Bull. Soc. Mycol. France 23: 95. 1907.



Diagnosis: Phylogeny: Series Paxillorum belongs to subgen. Aspergilloides, sect. Citrina, and is phylogenetically related to ser. Sheariorum, though without statistical support (Fig. 18). Morphology & physiology: Colonies growing moderately to fast; conidial colour en masse dull (blue-)green; conidiophores predominantly symmetrically biverticillate, rough-walled; conidia subglobose, smooth; good growth at 30 °C, no growth at 37 °C. Sexual morph unknown; sclerotia not observed in culture. Series description based on Houbraken et al. (2011a).



Included species: Penicillium paxilli.



Extrolites: Penicillium paxilli produces paxilline (Cole et al. 1974, Springer et al., 1975, Fan et al., 2018a) and pyrenocins (Fan et al., 2018a, Fan et al., 2018b, under the name P. camemberti).



Series Roseopurpurea Houbraken & Frisvad, ser. nov. MycoBank MB834265.



Etymology: Named after the type species of the series, Penicillium roseopurpureum.



Type: Penicillium roseopurpureum Dierckx, Ann. Soc. Sci. Bruxelles 25: 86. 1901.



Diagnosis: Phylogeny: Series Roseopurpurea belongs to subgen. Aspergilloides, sect. Citrina; the series is well-supported, but phylogenetic relationship with other series within sect. Citrina is undetermined (Fig. 18). Morphology & physiology: Colonies restricted; conidial colour en masse pale grey-green; conidiophores monoverticillate or furcate, smooth, generally short; conidia globose to subglobose, smooth or finely roughened; growth at 30 °C absent or restricted (13 mm), at 37 °C absent. Sexual morph unknown; sclerotia not observed in culture. Series description based on Houbraken et al. (2011a).



Included species: Penicillium roseopurpureum, P. sanguifluum.



Extrolites: The species in ser. Roseopurpurea produce the anthraquionone carviolin (= roseopurpurin) and related anthraquinones (Hind, 1940a, Hind, 1940b, Posternak, 1940). In addition, P. sanguifluum produces aculeatusquinones, citreofuran, citridones, curvularins, neobrugarones, penilactone, roseopurpurins A-I (not related to roseopurpurin), sulfimarin, and trichodimerol (Aly et al., 2011, Shang et al., 2016).



Series Sheariorum Houbraken & Frisvad, ser. nov. MycoBank MB834266.



Etymology: Named after the type species of the series, Penicillium shearii.



Type: Penicillium shearii Stolk & D.B. Scott, Persoonia 4: 396. 1967.



Diagnosis: Phylogeny: Series Sheariorum belongs to subgen. Aspergilloides, sect. Citrina, and is phylogenetically related to ser. Paxillorum, though without statistical support (Fig. 18). Morphology & physiology: Colonies growing moderately fast; conidial colour en masse grey-green; conidiophores predominantly biverticillate, smooth; conidia subglobose to broadly ellipsoidal, smooth; growth at 37 °C. Sexual morph eupenicillium-type, dark grey; ascospores ellipsoidal with two equatorial ridges, convex roughened, warted. Series description based on Houbraken et al. (2011a).



Included species: Penicillium shearii.



Extrolites: Paxillin, paspalinine, shearinins (Belofsky et al., 1995, Houbraken et al., 2011a, Ariantari et al., 2019). The production of paxillin indicates a relationship to P. paxilli (ser. Paxillorum) and P. thiersii (ser. Thiersiorum).



Series Sumatraensia Houbraken & Frisvad, ser. nov. MycoBank MB834267.



Etymology: Named after the type species of the series, Penicillium sumatraense.



Type: Penicillium sumatraense Szilvinyi, Archiv. Hydrobiol.14 Suppl. 6: 535. 1936.



Diagnosis: Phylogeny: Series Sumatraensia belongs to subgen. Aspergilloides, sect. Citrina and is phylogenetically related to ser. Copticolarum, though without statistical support (Fig. 18). Morphology & physiology: Colonies growing moderately or fast; conidial colour en masse blue-green, dull green or dark green; conidiophores predominantly biverticillate, smooth; conidia subglobose to broadly ellipsoidal, finely roughened; growth at 37 °C absent. Sexual morph unknown; sclerotia not observed in culture. Series description based on Houbraken et al. (2011a).



Included species: Penicillium sumatraense.



Extrolites: Penicillium sumatraense produces curvularins such as curvularin, dehydrocurvularin, sumalactone A-D, sumalarins and citridones E-G (Vesonder et al., 1976, Malmstrom et al., 2000, Meng et al., 2013, de Castro et al., 2016, Ha et al., 2017, Wu et al., 2017, Xu et al., 2019b). This species has been reported to produce a blue mould rot of Vitus vinifera and Sparassis crispa (Mahdian and Zafari, 2016, Liu et al., 2018b).



Series Westlingiorum Houbraken & Frisvad, ser. nov. MycoBank MB834268.



Etymology: Named after the type species of the series, Penicillium westlingii.



Type: Penicillium westlingii K.W. Zaleski, Bull. Int. Acad. Polon. Sci., Sér. B., Sci. Nat. 1927: 473. 1927.



Diagnosis: Phylogeny: Series Westlingiorum belongs to subgen. Aspergilloides, sect. Citrina; the phylogenetic position of this series in sect. Citrina remains unknown (Fig. 18). Morphology & physiology: Colonies growing variable, restrictly (e.g., P. wellingtonense, P. nothofagi), moderately or rapidly (P. decaturense, P. quebecense); conidial colour en masse blue-green to greyish green; conidiophores predominantly biverticillate, generally smooth, except in certain strains of P. manginii and P. atrofulvum; conidia globose, subglobose or (broadly) ellipsoidal; growth at 37 °C absent. Sexual morph unknown; sclerotia absent or present, mostly in shades of orange-brown (P. aurantiacobrunneum, P. cairnsense, P. manginii, P. miczynskii, P. pasqualense, P. quebecense), sometimes black (P. atrofulvum). Series description based on Houbraken et al. (2011a).



Included species: Penicillium atrofulvum, P. aurantiacobrunneum, P. cairnsense, P. christenseniae, P. chrzaszczii, P. cosmopolitanum, P. decaturense, P. godlewskii, P. manginii, P. miczynskii, P. neomiczynskii, P. nothofagi, P. pancosmium, P. pasqualense, P. quebecense, P. raphiae, P. sucrivorum, P. ubiquetum, P. vancouverense, P. waksmanii, P. wellingtonense, P. westlingii.



Extrolites: Citrinin is produced by 13/21 species, terrein is produced by 9/21 species, citreoviridin is produced by 8/21 species, decaturins are produced by 5/21 species, okaramins are produced by 3/21 species, phoenicin is produced by 3/21 species, quinolactacin is produced by species, quinolactacin is produced by 2/21 species, benzomalvins are produced by 2/21 species, territrems are produced by 2/21 species, perinadins are produced by 2/21 species, daldinins are produced by 2/21 species, citrinalins are produced by 1/21 species, curvularins are produced by 1/21 species, cyclopiazonic acid is produced by 1/21 species, meleagrin is produced by 1/21 species, and pyrenocin is produced by 1/21 species (Houbraken et al. 2011a).



Notes on series of sect. Citrina: The phylogenetic relationships within sect. Citrina were studied in detail with partial β-tubulin and calmodulin sequences (Houbraken et al. 2011a). Nine lineages were recognised and are treated here as series. The deeper nodes in sect. Citrina are mostly without any statistical support and therefore the phylogenetic relationship between the various series remains uncertain (Fig. 18). Six sect. Citrina species form cleistothecia: all taxa of series Euglauca (three species) and Sheariorum (one species), and two species of ser. Citrina (P. tropicum and P. tropicoides). Sclerotium production is present in ser. Gallaica and seven species of ser. Westlingiorum. The growth rate at 30 and 37 °C is also informative at series level. Series Westlingiorum species generally have maximum growth temperatures at or below 30 °C (with exception of P. pasqualense, P. quebecense and P. decaturense; growth at 30 °C or 33 °C). In contrast, ser. Citrina species have higher optimum and maximum growth temperatures. With exception of P. tropicoides, all species were able to grow at 33 °C. Series Sheariorum species grow well at 37 °C. The majority of sect. Citrina taxa produce symmetrically branched biverticillate conidiophores. Exceptions are all species classified in series Roseopurpurea (P. roseopurpureum, P. sanguifluum) and Gallaica (P. gallaicum) that predominantly produce monoverticillate conidiophores. Conidiophore stipes are generally smooth, with exception of ser. Sheariorum species and certain isolates of P. manginii and P. atrofulvum of ser. Westlingiorum (Houbraken et al. 2011a).



Section Crypta Houbraken & Frisvad, sect. nov. MycoBank MB834269.



Etymology: Named after the type species of the series, Penicillium cryptum.



Type: Penicillium cryptum Goch., Mycotaxon 26: 349. 1986.



Diagnosis: Phylogeny: Section Crypta belongs to subgen. Aspergilloides and is phylogenetically related to sect. Torulomyces. Morphology & physiology: Colonies growing very restrictedly; conidial colour en masse pale yellow-green; conidiophores biverticillate, occasionally terverticillate; stipes short (8–50(–90) μm), smooth; conidia globose to subglobose, smooth to finely roughened. Sexual morph eupenicillium-type, pale beige; ascospores ellipsoidal, with two well separated ridges, convex smooth-walled. Series description based on Gochenaur & Cochrane (1986).



Included species: Penicillium cryptum.



Extrolites: No chemotaxonomic data available.



Notes: Houbraken & Samson (2011) accommodated P. cryptum in sect. Torulomyces. More recently, Visagie et al. (2016a) showed that this species is actually distantly related to the other sect. Torulomyces taxa and suggested that it might not belong to this section. This observation was confirmed by phenotypic characters: P. cryptum predominately produces biverticillate conidiophores, while sect. Torulomyces members produce solitary phialides. Furthermore, P. cryptum produces smooth-walled conidia, in contrast to the ornamented conidia in sect. Torulomyces. Penicillium cryptum is the sole species in sect. Crypta and a subdivision of the section can therefore not be made. Series Crypta is only informally introduced here.



Section Eremophila Houbraken & Frisvad, sect. nov. MycoBank MB834270.



Etymology: Named after the type species of the series, Penicillium eremophilum.



Type: Penicillium eremophilum (A.D. Hocking & Pitt) Houbraken et al., Stud. Mycol. 86: 47. 2017.



Diagnosis: Phylogeny: Section Eremophila belongs to subgen. Aspergilloides and is most closely related to sect. Charlesia. Morphology & physiology: Xerophilic, no growth on high water activity media; asexual morph not produced; growth at 37 °C absent. Sexual morph monascus-like, brownish orange; ascospores subglobose to ellipsoidal, formed in pairs, smooth-walled. Series description based on Hocking and Pitt, 1988, Leong et al., 2011 and Barbosa et al. (2017).



Included species: Penicillium eremophilum.



Extrolites: This species has not been examined for extrolites.



Notes: Penicillium eremophilum is the sole species in sect. Eremophila and is unlike any other Penicillium species, an obligate xerophile. The asexual morph of this species is not known. In addition, the formation of two-spored asci is also not shared with other Penicillium species and this feature, together with its xerophily, is shared with the phylogenetically distant species Xeromyces bisporus (Barbosa et al. 2017). Earlier studies (Park et al., 2004, Pettersson et al., 2011, Houbraken et al., 2014a, Barbosa et al., 2017) repeatedly position P. eremophilum (as Monascus eremophilus) in Penicillium, supporting the results of our phylogenetic analysis. Penicillium eremophilum was confidently positioned as a sister lineage of a clade containing P. charlesii and P. fellutanum (both sect. Charlesia) (Houbraken et al. 2014a), and this result is confirmed in our study (Fig. 2, Fig. 18).



Section Exilicaulis Pitt, The Genus Penicillium: 205. 1980 [1979]. MycoBank MB832954.

Synonym: Eupenicillium sect. Lapidosa (Pitt) Stolk & Samson, Stud. Mycol. 23: 55. 1983.



Type: Penicillium restrictum J.C. Gilman & E.V. Abbott, Iowa State Coll. J. Sci. 1: 297. 1927.



Description: See Houbraken and Samson, 2011, Visagie et al., 2016c (morphology, phylogeny).



Series Alutacea (Pitt) Houbraken & Frisvad comb. nov. MycoBank MB834271.

Basionym: Eupenicillium ser. Alutacea Pitt, The Genus Penicillium: 54. 1980 [1979].



Type: Eupenicillium alutaceum D.B. Scott, Mycopathol. Mycol. Appl. 36: 17. 1968.



Diagnosis: Phylogeny: Series Alutacea belongs to subgen. Aspergilloides, sect. Exilicaulis and is phylogenetically related to series Citreonigra, Corylophila, Lapidosa and Restricta; the most closely related series could not be determined (Fig. 18). Morphology & physiology: Colonies restricted or growing moderately fast; conidial colour en masse grey-green or dull green; conidiophores monoverticillate, short and generally 20–60 μm; conidia (broadly) ellipsoidal, smooth-walled; growth at 37 °C generally present. Sexual morph not produced in culture (P. decumbens) or present (P. alutaceum), eupenicillium-type, pale brown; ascospores ellipsoidal, yellow, with two, sometimes four, longitudinal flanges, convex smooth to finely roughened. Series description based on Pitt (1980).



Included species: Penicillium alutaceum, Penicillium decumbens.



Extrolites: Penicillium alutaceum produces andrastin A and fulvic acid. Penicillium decumbens produces the volatiles thujopsene, nerolidol (both terpenes), 1-octen-3-ol, 3-octanone and phenylethylalcohol (Halim et al., 1975, Polizzi et al., 2011), and decumbenones and calbistrins (Fujii et al. 2002, results reported here). Penicillium striatisporum also produces calbistrins (Brill et al., 1993, Jackson et al., 1993) (both as P. restrictum, Stewart et al. 2005) and citromycins and citromycetins (Capon et al. 2007). Other extrolites reported include cyclocitrinols (Lin et al. 2014), cyclopenicillone (Lin et al. 2011) trichopyrone, sorbicillin, penicillone A and 3,11-dihydroxy-6,8-dimethyldodecanoic acid (Lin et al. 2018), and diisooctylphthalate (Amer et al. 2019). We also detected andrastin A and C in P. decumbens. Decumbin, reported from P. decumbens was shown to be brefeldin A (and produced by Penicillium brefeldianum) (Singleton et al., 1958, Betina, 1992). Peniproline A and chrysotriazoles and related compounds could not be detected in P. decumbens by us (see Wang et al. 2017b, the ITS sequence indicates the producer is Penicillium limosum), but such compounds have been found in Penicillium paneum (Li et al. 2011). In ser. Alutacea only andrastin A seems to be in common between the two species. A full genome sequenced isolate, with 28 predicted gene clusters for secondary metabolites was first identified as P. decumbens, but was later shown to be P. oxalicum (Liu et al., 2013, Houbraken et al., 2014a). This was confirmed by the fact that the isolate of P. oxalicum produced roquefortine C and meleagrin as other isolates of P. oxalicum (Steyn & Vleggaar 1983).



Series Citreonigra Pitt, The Genus Penicillium: 218. 1980 [1979]. MycoBank MB832956.



Type: Penicillium citreonigrum Dierckx, Ann. Soc. Sci. Bruxelles 25: 86. 1901.



Diagnosis: Phylogeny: Series Citreonigra belongs to subgen. Aspergilloides, sect. Exilicaulis and is phylogenetically related to series Alutacea, Corylophila, Lapidosa and Restricta; the most closely related series could not be determined (Fig. 18). Morphology & physiology: Colonies growing restrictedly to moderately rapid; conidial colour en masse grey-green, dull green or dark green; conidiophores monoverticillate, occasionally biverticillate; stipe smooth-walled; conidia smooth, globose; growth at 37 °C absent (P. cinerascens, P. citreonigrum, P. fundyense) or restricted (P. citreosulfuratum). Sexual morph unknown. Series description based on Visagie et al. (2016c).



Included species: Penicillium cinerascens, P. citreonigrum, P. citreosulfuratum, P. fundyense.



Extrolites: Three species in the series Citreonigra produce citreoviridin, P. cinerascens, P. citreonigrum (Sakabe et al. 1964, as P. citreoviride) and P. citreosulphuratum. Penicillium citreonigrum also produces citreoindol (closely related to haemindole) (Matsunaga et al., 1991, Song et al., 2016) and dipicolinic acid (Kalle & Khandekar 1983). Penicillium cinerascens was also reported to produce gliotoxin and dehydrocarolic acid (Bracken & Raistrick 1947). An atlantinone, sclerotiorin (several sclerotiorins) and pencolide were reported from P. citreonigrum; however, this fungus is correctly identified as P. hirayamae (Wang et al. 2010). These metabolites are typical for members of Penicillium series Sclerotiorum. A strain identified as P. citreonigrum (XN 10) was reported to produce sclerotiamine, three chromones, several eremophilans such as PR-amide, citreopenin and 3-epi-isopetasol, and mycophenolic acid (Yuan et al., 2014, Yuan et al., 2015, Yuan et al., 2017). We have not been able to detect those extrolites in P. citreonigrum. The isolate SP-6 of P. citreonigrum was reported to produce (-)-dichlorodiaportal in addition to an unusual diketopiperazine and N-(3-acetamidopropyl)-3-hydroxy-4-methoxybenzamide and a related compound (Huang et al. 2018). We have not been able to confirm production of those extrolites in P. citreonigrum either.



Series Corylophila Houbraken & Frisvad, ser. nov. MycoBank MB834272.



Type: Penicillium corylophilum Dierckx, Ann. Soc. Sci. Bruxelles 25: 86. 1901.



Etymology: Named after the type species of the series, Penicillium corylophilum.



Diagnosis: Phylogeny: Series Corylophila belongs to subgen. Aspergilloides, sect. Exilicaulis and is phylogenetically related to ser. Lapidosa (Fig. 18). Morphology & physiology: Colonies generally spreading, sometimes growing moderately rapid or restrictedly (P. cravenianum, P. pagulum); conidial colour en masse in shades of green (greyish, dull or dark green); conidiophores predominantly biverticillate, sometimes mono- or terverticillate; stipe smooth or rough-walled; conidia globose to broadly ellipsoidal, smooth, finely or distinctly roughened; growth at 37 °C absent or present and often restricted (P. momoii, P. pagulum, P. repensicola, P. subturcoseum). Sexual morph unknown. Series description based on Visagie et al. (2016c).



Included species: Penicillium atrolazulinum, P. consobrinum, P. corylophilum, P. cravenianum, P. fagi, P. momoii, P. pagulum, P. punicae, P. repensicola, P. rubefaciens, P. subturcoseum.



Extrolites: Penicillium corylophilum produces andrastins and the related citreohybridinol, citreoisocoumarin and the related (+)-orthosporin, phomenone, isochromans and α-pyrones (Lai et al. 1991 (reported under the synonym P. citreovirens), Malmstrom et al., 2000, McMullin et al., 2014a, McMullin et al., 2014b, Yadav et al., 2014). Production of decarestrictins and epoxyagroclavine-I has also been reported from P. corylophilum (Grabley et al. 1992), but the identity of the strain has not been confirmed. Furan-2-carboxylic acid has been reported from P. corylophilum (Turner & Aldridge 1983). Penicillium fagi produces andrastin A, citrinalin, mycophenolic acid, a paraherquamide and pulvilloric acid. Penicillium rubefaciens produces a curvulic acid related extrolite. The other species have not yet been examined for extrolites, but there seem to be few extrolites in common between the species in ser. Corylophila.



Series Erubescentia (Pitt) Houbraken & Frisvad, comb. nov. MycoBank MB834273.

Basionym: Eupenicillium ser. Erubescentia Pitt, The Genus Penicillium: 70. 1980 [1979].



Type: Eupenicillium erubescens D.B. Scott, Mycopathol. Mycol. Appl. 36: 14. 1968. MycoBank MB330727.



Diagnosis: Phylogeny: Series Erubescentia belongs to subgen. Aspergilloides sect. Exilicaulis and is phylogenetically sister to all other series of this section (Fig. 18). Morphology & physiology: Colonies generally restricted, sometimes moderately fast; conidial colour en masse variable, olive grey, blue grey, grey-green or dull green; conidiophores monoverticillate; stipes short, smooth; conidial shape variable, globose, subglobose, ovoid, or (broadly) ellipsoidal, smooth or rough-walled; growth at 37 °C generally present, sometimes absent (e.g., P. hermansii, P. nepalense). Sexual morph unknown (12 species) or present (six species) (see list of accepted species), eupenicillium-type, brown, orange-brown or pinkish brown; ascospores (broadly) ellipsoidal, with one or two pairs of longitudinal flanges, convex smooth, roughened or spinose. Series description based on Pitt, 1980, Peterson et al., 1999, Visagie et al., 2016c and Houbraken et al. (2019).



Included species: Penicillium canis, P. catenatum, P. dimorphosporum, P. dravuni, P. erubescens, P. guttulosum, P. hermansii, P. labradorum∗, P. laeve, P. menonorum, P. nepalense, P. ovatum, P. parvofructum, P. parvum, P. pimiteouiense, P. rubidurum, P. striatisporum, P. vinaceum [∗ not included in Fig. 18; see Rothacker et al. (2020)].



Extrolites: Penicillium striatisporum produces calbistrins, striatisporins, hexylitaconic acids and striatisporolide A (Brill et al., 1993, Jackson et al., 1993, Stewart et al., 2005). Penicillium erubescens has been reported to produce anhydrofulvic acid and the related myxotrichin B and citromycins, GKK1032B, penialidin D, pyranochromones, secalonic acid A and SPF-3059-30 and related chromones including erubescensoic acid (Kumla et al., 2018, Kumla et al., 2019). Penicillium parvum has been reported to produce mycophenolic acid and many derivatives and precursors (including euparvic acid and euparvilactone), citromycetin, euparvione, 7-hydroxy-2,5-dimethyl-4H-chromen-4-one and azadirachtin (Habib et al., 2008, Kusari et al., 2012, León et al., 2013) and P. dravuni produces dictyosphaeric acid A & B and carviolin = roseopurpurin (Bugni et al., 2004, Burns et al., 2010). Penicillium vinaceum has been reported to produce vinaxanthones (Aoki et al., 1991, Řezanka et al., 2008) and (-)-(1R,4R)-1,4-(2,3)indolmethane-1-methyl-2,4-dihydro-1H-pyrazino-[2,1-b]-quinazoline.3,6-dione (Zheng et al. 2012), and penicillivinacine, citreoisocoumarin, indol-3-carbaldehyde, α-cyclopiazonic acid, terretrione A, brevianamide F and its diastereomer cyclo-D-Trp-L-Pro (Asiri et al. 2015). The latter seven metabolites are apparently produced by Penicillium rubens (or closely related species) according to the reported ITS sequence. Other metabolites reported from P. vinaceum was based on the strain DQ25 (Wei et al. 2009), which, based on the reported ITS sequence, is also a P. rubens. Penicillium parvum and P. erubescens share secondary metabolites from the citromycetin biosynthetic family, but else species in ser. Erubescentia shares coloured polyketide metabolites, that are somewhat different.



Series Lapidosa (Pitt) Houbraken & Frisvad, comb. nov. MycoBank MB834517.

Basionym: Eupenicillium ser. Lapidosa Pitt, The Genus Penicillium: 129. 1980 [1979].



Type: Eupenicillium lapidosum D.B. Scott & Stolk, Antonie van Leeuwenhoek 33: 298. 1967. MycoBank MB330733.



Diagnosis: Phylogeny: Series Lapidosa belongs to subgen. Aspergilloides, sect. Exilicaulis and is phylogenetically related to ser. Corylophila (Fig. 18). Morphology & physiology: Colonies growing moderate, sometimes fast; conidial colour en masse variable, in shades of green, dark green, greyish green or dull green; conidiophores biverticillate, sometimes monoverticillate; conidia variable shaped (globose to ellipsoidal), generally smooth-walled; growth at 37 °C variable. Sexual morph generally not observed in culture, present in P. lapidosum and P. terrenum, eupenicillium-type, orange-brown or light brownish yellow; ascospores hyaline to pale yellow, ellipsoidal, with two prominent longitudinal flanges, convex smooth or spinose. Series description based on Pitt (1980) and Visagie et al. (2016c).



Included species: Penicillium aotearoae, P. atrosanguineum, P. burgense, P. diabolicalicense, P. hemitrachum, P. lapidosum, P. maclennaniae, P. melinii, P. namyslowskii, P. raciborskii, P. smithii, P. terrenum, P. velutinum, P. xanthomelinii.



Extrolites: Penicillium smithii produces a metabolite related to phoenicin, in addition to citreoviridin, citreoisocoumarins and paxillin, and P. atrosanguineum and P. maclennaniae share quinone secondary metabolites (phoenicin and spinulosin, respectively) (Christensen et al. 1999). Furthermore, P. atrosanguineum produces some tryptoquivalines. Penicillium lapidosum was reported to produce patulin (Myrchink 1967) and lapidosin (Turner 1978). We detected pulvilloric acid in P. raciborskii and P. melinii. In addition, P. melinii produces andrastin A and daldinin D. Penicillium namyslowskii produces a polyene not yet structure elucidated and haenamindole. Most species in ser. Lapidosa have not yet been studied in any detail chemically.



Series Restricta Raper & Thom ex Pitt, The Genus Penicillium: 205. 1980 [1979]. MycoBank MB832955.



Type: Penicillium restrictum J.C. Gilman & E.V. Abbott, Iowa State Coll. J. Sci. 1: 297. 1927.



Diagnosis: Phylogeny: Series Restricta belongs to subgen. Aspergilloides, sect. Exilicaulis and is phylogenetically related to series Alutacea, Corylophila, Lapidosa and Citreonigra; the most closely related series could not be determined (Fig. 18). Morphology & physiology: Colonies growing restrictedly or moderately rapid; conidial colour en masse grey-green, blue-green or dull green; conidiophores generally monoverticillate; stipe smooth or finely roughened, generally short; conidia globose, subglobose, or (broadly) ellipsoidal, smooth or roughened; growth at 37 °C generally present. Sexual morph generally not observed in culture (15 species), sometimes present (P. katangense, P. meridianum, P. philippinense), eupenicillium-type, greyish orange or light brownish orange; ascospores ellipsoidal, yellow, with two small appressed longitudinal flanges, sometimes with a second smaller pair, convex smooth-walled or finely roughened. Series description based on Pitt, 1980, Stolk and Samson, 1983 and Visagie et al. (2016c).



Included species: Penicillium arabicum, P. chalabudae, P. cinereoatrum, P. heteromorphum, P. katangense, P. kurssanovii, P. meridianum, P. philippinense, P. restrictum.



Extrolites: Penicillium restrictum has been reported to produce pestalotin, hydroxypestalotin, LLP880γ and 5,6-dihydro-4-methoxy-6-(1-oxopentyl)-2H-pyran-2-one (Geiger et al. 2013), emodin, emodic acid and other anthraquinones (Figueroa et al. 2014), andrastin A and phomenone (Antipova et al. 2018b), gliotoxin and dehydrocarolic acid (Sankhala 1968), 2,3-dihydro-3,6-dihydroxy-2-methyl-4-pyrone, curvularin and dehydrocurvularin (as P. gilmanii, Raistrick and Rice, 1971, Rice and Chen, 1984), and restricticins (Hensens et al., 1991, Schwartz et al., 1991). Production of patulin and penicillic acid by P. restrictum (Martín et al. 2004) could not be confirmed.



Notes: Visagie et al. (2016c) studied the phylogenetic relationship between sect. Exilicaulis species and recognised six main lineages: the P. citreonigrum-, P. corylophilum-, P. decumbens-, P. melinii-, P. parvum- and P. restrictum-clades. These clades are treated here as series: Citreonigra, Corylophila, Alutacea, Lapidosa, Erubescentia and Restricta, respectively. These series are primary based on phylogenetic data. Visagie et al. (2016c) indicated that the branching pattern is informative. The phylogenetically related series Corylophila and Lapidosa contain species with biverticillate conidiophores, and monoverticillate species were present in the four other series (Alutacea, Citreonigra, Erubescentia and Restricta). Species belonging to sect. Exilicaulis have different growth rates and species belonging to series Alutacea, Citreonigra, Erubescentia and Restricta grow restrictedly or moderately fast, while species of series Lapidosa and Corylophila tend to be more spreading. Growth at 37 °C is generally present in species of series Alutacea, Erubescentia and Restricta and absent in Citreonigra and Corylophila, though exceptions are present in most series; species of ser. Lapidosa show variable growth at 37 °C. At this moment, it is not possible to distinguish the series on phenotypic characters and this could be subject of future studies.



Section Inusitata Houbraken & Frisvad, sect. nov. MycoBank MB834274.



Type: Penicillium inusitatum D.B. Scott, Mycopathol. Mycol. Appl. 36: 20. 1968.



Etymology: Named after the type species of the section, P. inusitatum.



Diagnosis: Phylogeny: Section Inusitata belongs to subgen. Aspergilloides and is phylogenetically sister to sect. Exilicaulis (Fig. 18). Morphology & physiology: Colonies restricted; conidial colour en masse dull green or glaucous grey; conidiophores monoverticillate (P. fractum) or biverticillate and divaricate (P. inusitatum); conidia ellipsoidal or cylindrical, smooth; growth at 37 °C present. Sexual morph eupenicillium-type, yellow, yellowish brown or orange-brown; ascospores globose, spinose (without flanges or furrows). Section description based on Pitt (1980) and Houbraken & Samson (2011).



Included species: Penicillium fractum, P. inusitatum.



Extrolites: No extrolites have been detected in the two species in ser. Inusitata yet.



Notes: Taxa belonging to this section were previously classified in Penicillium sect. Fracta. Section Fracta was (incorrectly) typified with Penicillium ornatum, a member of sect. Ramigena. Section Fracta is therefore considered a synonym of sect. Ramigena and the new section Inusitata is here proposed. A subdivision of sect. Inusitata is not made here and Penicillium ser. Inusitata is only informally introduced here. Eupenicillium ser. Fracta (MycoBank MB832945) (Pitt 1980) was formally introduced and is the basionym of this tentative series.



Section Gracilenta Houbraken & Samson, Stud. Mycol. 70: 40. 2011. MycoBank MB563131.



Type: Penicillium gracilentum Udagawa & Y. Horie, Trans. Mycol. Soc. Japan 14: 373. 1973.



Description: See Houbraken & Samson (2011), this study (phylogeny); a modern overview of this section is lacking.



Series Angustiporcata Houbraken & Frisvad, ser. nov. MycoBank MB834275.



Type: Penicillium angustiporcatum Takada & Udagawa, Trans. Mycol. Soc. Japan 24: 143. 1983.



Etymology: Named after the type species of the series, Penicillium angustiporcatum.



Diagnosis: Phylogeny: Series Angustiporcata belongs to subgen. Aspergilloides, sect. Gracilenta and is phylogenetically sister to series Estinogena, Gracilenta and Macrosclerotiorum (Fig. 18). Morphology & physiology: Colonies restricted to moderately fast; conidial colour en masse undetermined; conidiophores predominantly biverticillate, occasionally monoverticillate, smooth; conidia broadly ellipsoidal, smooth; growth at 37 °C absent. Sexual morph eupenicillium-type, flesh-coloured or pale yellowish brown, small (50–150 μm); ascospores broadly ellipsoidal, with two prominent equatorial ridges, convex rugose, with several low ribs. Series description based on Takada & Udagawa (1983).



Included species: Penicillium angustiporcatum.



Extrolites: No extrolites have been found in P. angustiporcatum.



Series Estinogena Houbraken & Frisvad, ser. nov. MycoBank MB834276.



Type: Penicillium estinogenum A. Komatsu & S. Abe ex G. Sm., Trans. Brit. Mycol. Soc. 46: 335. 1963.



Etymology: Named after the type species of the series, Penicillium estinogenum.



Diagnosis: Phylogeny: Series Estinogena belongs to subgen. Aspergilloides, sect. Gracilenta and is phylogenetically sister to ser. Macrosclerotiorum (Fig. 18). Morphology & physiology: Colonies spreading; reverse dark olive; conidial colour en masse blue-green; conidiophores symmetrically biverticillate, rough-walled; conidia globose or ellipsoidal; smooth or finely roughened. Sexual morph unknown; sclerotia not observed in culture. Series description based on Ramírez (1982).



Included species: Penicillium estinogenum.



Extrolites: Penicillium estinogenum produces asterric acid, erdin, estin, geodin geodin hydrate in addition to dehydocurvularin and patulin. Production of verruculogen (Day et al. 1980) could not be confirmed, and the original strain producing this secondary metabolite was found to be a new species (not yet described).



Series Gracilenta Houbraken & Frisvad, ser. nov. MycoBank MB834277.



Type: Penicillium gracilentum Udagawa & Y. Horie, Trans. Mycol. Soc. Japan 14: 373. 1973.



Etymology: Named after the type species of the series, Penicillium gracilentum.



Diagnosis: Phylogeny: Series Gracilenta belongs to subgen. Aspergilloides, sect. Gracilenta and is phylogenetically sister to series Estinogena and Macrosclerotiorum (Fig. 18). Morphology & physiology: Colonies growing moderately fast; conidial colour en masse dull blue-green to greenish grey; conidiophores predominantly monoverticillate, sometimes divaricate; conidia (broadly) ellipsoidal, smooth or finely roughened; growth at 37 °C absent. Sexual morph eupenicillium-type, greyish yellow-brown; ascospores ellipsoidal, with two equatorial sinuous ridges, convex scattered with small spines. Series description based on Udagawa & Horie (1973) and Pitt (1980).



Included species: Penicillium gracilentum.



Extrolites: No known extrolites were found in P. gracilentum.



Series Macrosclerotiorum Houbraken & Frisvad, ser. nov. MycoBank MB834278.



Type: Penicillium macrosclerotiorum L. Wang et al., Mycol. Res. 111: 1244. 2007.



Etymology: Named after the type species of the series, Penicillium macrosclerotiorum.



Diagnosis: Phylogeny: Series Macrosclerotiorum belongs to subgen. Aspergilloides, sect. Gracilenta and is phylogenetically sister to ser. Estinogena (Fig. 18). Morphology & physiology: Colonies growing moderately to fast; conidial colour en masse variable, grey-green to olive-grey; conidiophores monoverticillate, smooth; conidia globose, smooth or finely roughened; growth at 37 °C absent (P. macrosclerotiorum) or present (P. apimei, P. aquaticum). Sexual morph unknown; sclerotia absent (P. apimei) or present (P. aquaticum, P. macrosclerotiorum), white when young, becoming ivory in age. Series description based on Wang et al., 2007a, Barbosa et al., 2018 and Wanasinghe et al. (2018).



Included species: Penicillium apimei, P. aquaticum, P. macrosclerotiorum.



Extrolites: Penicillium macrosclerotium produces asterric acid, erdin, estin, geodin geodin hydrate like P. estinogenum in the sister ser. Estinogena.



Notes: Comparison of the phenotypic characters at section level did not reveal many significant similarities, except that all produce an olive-brown to brown reverse on agar media (Houbraken and Samson, 2011, Barbosa et al., 2018). The inability to grow at 37 °C was also mentioned as a shared character of the species belonging to this section. However, P. apimei and P. aquaticum were recently added to sect. Gracilenta and both are able to grow at this temperature (Barbosa et al., 2018, Wanasinghe et al., 2018). Section Gracilenta has not yet been subject of a taxonomic study; however, the phylogenetic data shows there are four main lineages within the section. These lineages are here treated as series. Species of ser. Macrosclerotiorum share the production of monoverticillate conidiophores and globose conidia, and this combination of characters is not observed in the other sect. Gracilenta series. Series Angustiporcata and Gracilenta are monotypic series. Species in these series are the only ones in the section that reproduce sexually. They differ in their conidiophore branching complexity (monoverticillate vs biverticillate) and convex ornamentation of the ascospores (bars vs spines). Penicillium estinogenum is the sole member of ser. Estinogena. This strictly asexual reproducing species grows rapidly and produces rough-walled, biverticillate conidiophores. These features are not observed in the other taxa of sect. Gracilenta.



Section Griseola Houbraken & Frisvad, sect. nov. MycoBank MB834279.



Type: Penicillium griseolum G. Sm., Trans. Brit. Mycol. Soc. 40: 485. 1957.



Etymology: Named after the type species of the section, Penicillium griseolum.



Diagnosis: Phylogeny: Section Griseola belongs to subgen. Aspergilloides and is phylogenetically sister of sections Aspergilloides, Charlesia, Eremophila and Sclerotiorum (Fig. 2, Fig. 18). Morphology & physiology: Colonies growing moderately quickly, thin and transparent on Czapek agar; texture funiculose; conidial colour en masse grey; conidiophores monoverticillate, smooth, short; conidia globose, conspicuously echinulate, with conspicuous connectives. Sexual morph unknown; sclerotia not observed in culture. Section description based on Smith (1957).



Included species: Penicillium griseolum.



Extrolites: No known extrolites have been reported from P. griseolum.



Notes: Penicillium griseolum was considered to be closely related to P. terlikowskii (= P. glabrum), from which it differs in its very poor growth on Czapek agar, its lack of green colour on malt extract agar, its different reverse colour, and in the curious tendency to form oidium-like chains of very slow ripening conidia (Smith 1957). Later, Pitt (1980) assigned this species to synonymy with P. restrictum based on phenotypic similarity, while Ramírez (1982) and Pitt et al. (2000) considered the species distinct. Using sequence data, Peterson & Horn (2009) showed that P. griseolum is a distinct species, though the phylogenetic position in the genus remained unresolved. In the phylogenetic analysis of the genus, Houbraken & Samson (2011) placed this species sister to sections Aspergilloides, Charlesia and Sclerotiorum; however, statistical support was generally poor. In this study, this relationship is confidently supported (Fig. 2). In order to maintain monophyly, sect. Griseola is introduced for this species. A subdivision of the section cannot be made and therefore ser. Griseola is only informally introduced here (Table 5).



Section Lanata-Divaricata Thom, The Penicillia: 157, 328. 1930. MycoBank MB834002.

Synonyms: Penicillium sect. Funiculosa Thom, The Penicillia: 157, 358. 1930.

Penicillium sect. Divaricata [as “Divaricatum”] Raper & Thom ex Pitt, The Genus Penicillium: 238. 1980 [1979].

Penicillium sect. Furcata [as “Furcatum”] Pitt, The Genus Penicillium: 272. 1980 [1979].

Eupenicillium sect. Javanica (Pitt) Stolk & Samson, Stud. Mycol. 23: 55. 1983.



Type: Penicillium janthinellum Biourge, Cellule 33: 258. 1923.



Description: See Houbraken and Samson, 2011, Visagie et al., 2015 (phylogeny, morphology).



Series Dalearum Houbraken & Frisvad, ser. nov. MycoBank MB834280.



Type: Penicillium daleae K.W. Zaleski, Bull. Int. Acad. Polon. Sci., Sér. B., Sci. Nat. 1927: 495. 1927.



Etymology: Named after the type species of the series, Penicillium daleae.



Diagnosis: Phylogeny: Series Dalearum belongs to subgen. Aspergilloides, sect. Lanata-Divaricata and is phylogenetically sister to series Rolfsiorum and Simplicissima (represented by P. subrubescens and P. simplicissimum in Supplementary Fig. S1, resp.). Morphology & physiology: Colonies growing moderately fast or spreading; conidial colour en masse dull green, blueish green, grey-green; conidiophores monoverticillate, biverticillate and divaricate, occasionally terverticillate, generally smooth, sometimes finely rough-walled; conidia variable shaped (globose, subglobose, (broadly) ellipsoidal), smooth or rough-walled, spinose; growth at 37 °C variable. Sexual morph generally not observed in culture, sometimes present, eupenicillium-type, homothallic, yellow-brown, ochre or yellowish cream; ascospores ellipsoidal, with two (shallow) longitudinal flanges (P. abidjanum, P. zonatum) or traces of an inconspicuous equatorial ridge (P. vanderhammenii), valves (finely) spinose).



Included species: Penicillium abidjanum, P. amphipolaria, P. austrosinense, P. daleae, P. griseopurpureum, P. guaibinense, P. jianfenglingense, P. penarojense, P. rubriannulatum, P. singorense, P. vanderhammenii, P. viridissimum, P. zonatum.



Extrolites: Penicillium abidjanum produces N-methylgliovirin. Penicillium daleae produces 3-(dimethylaminobenzyl)-N-(1,1-ddimethyl-2-propenyl)indol (Lam et al. 1994), JBIR-54 (Mukai et al. 2009), penicilliols (Kimura et al. 2009), isopenicins (Tang et al. 2019b), andrastin A, antarone A, curvulic acid, gregatins, penicillifuranone A, viomellein, vioxanthin, and xanthomegnin. Penicillium griseopurpureum produces andrastin A, curvulic acid and xanthoepocin. Penicillium penarojense produces paxillin, paspaline, and shearinins/janthitrems (Houbraken et al. 2011b), P. vanderhammenii produces paxillin, paspaline, penicilic acid and shearinins/janthitrems (Houbraken et al. 2011b), P. zonatum produces brefeldin A, curvulic acids, shearinins/janthitrems, viomellein, vioxanthin and xanthomegnin. The species in the series thus share several extrolites. Some of these are also produced by other species in sect. Lanata-Divaricata.



Series Janthinella Thom ex Pitt, The Genus Penicillium: 239. 1980 [1979]. MycoBank MB832959.

Synonym: Eupenicillium ser. Javanica Pitt, The Genus Penicillium: 113. 1980 [1979].



Type: Penicillium janthinellum Biourge, Cellule 33: 258. 1923.



Diagnosis: Phylogeny: Series Janthinella belongs to subgen. Aspergilloides, sect. Lanata-Divaricata and is phylogenetically sister to series Dalearum, Rolfsiorum and Simplicissima (represented by P. abidjanum, P. subrubescens and P. simplicissimum in Supplementary Fig. S1, resp.). Morphology & physiology: Colonies spreading; conidial colour en masse in shades of green, often grey-green; conidiophores biverticillate or divaricate, sometimes monoverticillate; stipes smooth-walled, sometimes roughened (P. javanicum); conidia variable in shape and ornamentation: globose, subglobose or (broadly) ellipsoidal and smooth, finely rough or rough-walled; growth at 37 °C generally present, sometimes absent (P. meloforme, P. yunnanense). Sexual morph not observed in culture (11 species) or present (13 species) (for details, see below in list of accepted species), eupenicillium-type, homothallic, creamish, yellow, yellow-brown or brown; ascospores ellipsoidal, with two distinct or inconspicuous longitudinal flanges, sometimes lacking equatorial ridges, convex ornamented or spinose. Series description based on Pitt, 1980, Stolk and Samson, 1983, Houbraken et al., 2011b and Visagie et al. (2015).



Included species: Penicillium brefeldianum, P. caperatum, P. cluniae, P. coeruleum, P. cremeogriseum, P. curticaule, P. ehrlichii, P. elleniae, P. glaucoroseum, P. janthinellum, P. javanicum, P. koreense, P. levitum, P. limosum, P. lineolatum, P. ludwigii, P. malacosphaerulum, P. meloforme, P. ortum, P. raperi, P. reticulisporum, P. setosum, P. uruguayense, P. yunnanense.



Extrolites: Penicillium brefeldianum produces bredinin (Mizuno et al. 1974), brefeldin A (Härri et al., 1963, Jouda et al., 2016), citromycetin and fulvic acid (Jouda et al., 2014, Jouda et al., 2016), eupenifeldin (Mayerl et al. 1993), hydroxyphenylglyoxaladoxime (Jouda et al. 2016), palitantin (Demetriadou et al. 1985) and penialidins (Jouda et al., 2014, Jouda et al., 2016, Cheng et al., 2018). Penicillium caperatum produces viridicatumtoxin and aplora indoloterpenes, and P. cluniae produces brefeldin A, janthitrems/sharinins, paraherquamides and cyclic dipeptides (diketopiperazines) (López-Gresa et al. 2006). Penicillium cremeogriseum produces brefeldin A, janthitrems/shearinins and fulvic acid; P. elleniae was reported to produce paxillin and sorbicillins (Houbraken et al. 2011b); P. glaucoroseum produces brefeldin A, fulvic acid and viomellein, vioxanthin and xanthomegnin; P. janthinellum and P. levitum produces viomellein, vioxanthin and xanthomegnin. Brefeldin A, fulvic acid, paspaline and palitantin are produced by P. ludwigii and P. reticulisporum produces xanthoepocin. These data show that the species in ser. Janthinella have many extrolites in common.



Series Oxalica Raper & Thom ex Pitt, The Genus Penicillium: 273. 1980 [1979]. MycoBank MB832963.



Type: Penicillium oxalicum Currie & Thom, J. Biol. Chem. 22: 289. 1915.



Diagnosis: Phylogeny: Series Oxalica belongs to subgen. Aspergilloides, sect. Lanata-Divaricata and is phylogenetically sister to all other series of sect. Lanata-Divaricata (Fig. 2, Fig. 18). Morphology & physiology: Colonies spreading; conidial colour en masse yellowish green, grey-green or dull green; texture strictly velutinous and crustose; conidiophores biverticillate, occasionally mono- or terverticillate, smooth; conidia ellipsoidal, smooth or finely roughened; growth at 37 °C present. Sexual morph unknown; sclerotia not observed in culture.



Included species: Penicillium diatomitis, P. oxalicum, P. soosanum.



Extrolites: Penicillium oxalicum produces oxaline, meleagrin, glandicolin A & B, and roquefortine C (Nagel et al., 1974, Nagel et al., 1976, Steyn and Vleggaar, 1983, Chen et al., 2015), secalonic acids, paecilin C and penicillixanthone (Steyn, 1970, Li et al., 2010, Kim et al., 2012, Bao et al., 2013, Wang et al., 2013b, Chen et al., 2015, Liu et al., 2015, Chen et al., 2019), oxalic acid (Currie & Thom 1915), benzenedicarboxylic acid and benzimidazole (Ahmad et al. 2019), (Z)-3-(3,4-dihydroxyphenyl)-2-foramidoacrylate and decaturins and oxalicins (Ubillas et al., 1989, Wang et al., 2013a, Li et al., 2015b, Zhang et al., 2015a), 2,2’,4,4’-tetrahydroxy-8’-methyl-6-methoxy-acyl-ethyl-diphenylmethanone (Liu et al., 2015, Liu et al., 2016a), hydroxyscytalone (Ji et al. 2014), coniochaetones and penicillones (Bao et al. 2014), gymnemagenin (Parthasarathy & Sathiyabama 2014), 2-(4-hydroxybenzoyl) quinazolin-4(3H)-one, 2-(4-hydroxybenzyl) quinazolin-4(3H)-one, methyl 4-hydroxyphenylacetate (= penipanoid C), rubinaphthin, citreorosein, emodin, isorhodoptilometrin and endocrocin (Li et al., 2010, Bao et al., 2013, Shen et al., 2013, Chen et al., 2015), oxalicumones (Sun et al., 2012, Sun et al., 2013, Zhang et al., 2013), 6,8-dihydroxy-3-methyl-9-oxo-9H-xanthene-1-carboxylic acid, 5-hydroxy-2-methoxybenzoic acid and 2-phenylacetic acid (Li et al. 2010),and vermiculidiol (Kim et al. 2012). Penicillium soosanum and P. diatomitis appear to be the producers of penioxalamine, as the ITS sequence (KJ101590) reported by Hu et al. (2014) is different from that of P. oxalicum sensu stricto (Kubátová et al. 2019).



Series Rolfsiorum Houbraken & Frisvad, ser. nov. MycoBank MB834281.



Type: Penicillium rolfsii Thom, Penicillia: 489. 1930.



Etymology: Named after the type species of the series, Penicillium rolfsii.



Diagnosis: Phylogeny: Series Rolfsiorum belongs to subgen. Aspergilloides, sect. Lanata-Divaricata and is phylogenetically sister to ser. Simplicissima (represented by P. simplicissimum in Supplementary Fig. S1). Morphology & physiology: Colonies spreading; conidial colour en masse grey-green, dull green; conidiophores terminally biverticillate, sometimes terverticillate or divaricate, rough-walled; conidia generally globose to subglobose, occasionally ellipsoidal or fusiform, smooth, finely roughened or rough; growth at 37 °C variable. Sexual morph unknown; sclerotia generally not observed in culture, present in P. rolfsii (white or pink) and P. soliforme (light yellow).



Included species: Penicillium annulatum, P. bissettii, P. camponotum, P. excelsum, P. flaviroseum, P. fructuariae-cellae, P. hainanense, P. ochrochloron, P. piscarium, P. pulvillorum, P. rolfsii, P. soliforme, P. subrubescens, P. svalbardense, P. terrarumae, P. vasconiae.



Extrolites: Penicillium bissettii produces aurantioclavine, neooxaline, meleagrin, roquefortine C and other precursors, oxalicine B and penicillic acid (Visagie et al. 2016b), P. camponotum produces andrastin A-D, citrinalin, mangrovamides, marcfortine A and B and patulin (Visagie et al. 2016b, and results reported here), P. excelsum produces andrastin A, curvulic acid, penicillic acid and xanthoepocin (Taniwaki et al. 2015), P. ochrochloron produces andrastin A, okaramine A-R, penicillic acid, penitrems including penitrem A, viridicatumtoxin and xanthoepocin (Hayashi et al., 1988, Hayashi et al., 1989, Hayashi et al., 1991a, Hayashi et al., 1995, Murao et al., 1988, Hayashi and Sakaguchi, 1998, Shiono et al., 1999, Shiono et al., 2000, Tuthill et al., 2001, Nielsen and Smedsgaard, 2003), P. piscarium produces haenamindole, janthitrems/shearinins, pulvilloric acid and xanthoepocin (Tuthill et al. 2001 and results reported here) and results reported here), P. pulvillorum produces andrastin A, curvulic acids, penicilic acid, pulvilloric acid, and xanthoepocin (Brian et al., 1957, Barber et al., 1986, Tuthill et al., 2001), P. rolfsii produces decaturins/oxalicins, paraherquamides, patulin, penicillic acid, viridicatumtoxin and xanthoepocin (data reported here), P. subrubescens and P. svalbardense produce xanthoepocin (Sonjak et al., 2007, Mansouri et al., 2013), and P. vasconiae produces curvulic acid, janthitrems/shearinins and pyripyropenes (results reported here). Species in ser. Rolfsiorum share many secondary metabolites, including andrastins, penicillic acid, curvulic acid, pulvilloric acid, viridicatumtoxin, janthitrems/shearinins, decaturins/oxalicins. However, not all species in the series produce these metabolites and several metabolites are found in other series in sect. Lanata-Divaricata as well.



Series Simplicissima Houbraken & Frisvad, ser. nov. MycoBank MB834282.



Type: Penicillium simplicissimum (Oudem.) Thom, Penicillia: 335. 1930.



Etymology: Named after the type species of the series, Penicillium simplicissimum.



Diagnosis: Phylogeny: Series Simplicissima belongs to subgen. Aspergilloides, sect. Lanata-Divaricata and is phylogenetically sister to ser. Rolfsiorum (represented by P. subrubescens in Supplementary Fig. S1). Morphology & physiology: Colonies spreading; conidial colour en masse grey-green or dull green; conidiophores biverticillate or divaricate, sometimes terverticillate; stipes smooth or rough-walled; conidia variable in size and shape, globose, subglobose, (broadly) ellipsoidal, ornamentation smooth, finely rough or rough-walled, sometimes in spiral pattern; growth at 37 °C variable. Sexual morph unknown; sclerotia generally not observed in culture, sometimes present, e.g., in P. mariae-cruscis, P. tanzanicum (yellowish), P. araracuaraense (yellow-brown), P. griseoflavum (greyish orange).



Included species: Penicillium alagoense, P. araracuaraense, P. brasilianum, P. cataractarum, P. echinulonalgiovense, P. globosum, P. griseoflavum, P. guangxiense, P. infrabuccalum, P. laevigatum, P. mariae-crucis, P. onobense, P. panissanguineum, P. paraherquei, P. pedernalense, P. simplicissimum, P. skrjabinii, P. spinuliferum, P. tanzanicum, P. wotroi.



Extrolites: Penicillium araracuarense produces pulvilloric acid, P. brasilianum produces aspterric acid, austins, brasilianoids and berkeleytrione (Hayashi et al., 1994, Matsuda et al., 2016, Zhang et al., 2018, Zhang et al., 2019), brasiliamides (Fujita et al., 2002, Fujita and Hayashi, 2004, Fill Taicia et al., 2009), fischerin, fumitremorgsins, verruculogen, spirotryprostatins and cyclotryprostatins (Frisvad, 1989, Zhang et al., 2019), JBIR 113, 114 & 115 (Koyama et al. 2012), neosartorin, penicillic acid (Frisvad et al., 1989, Kang and Kim, 2004, Schürmann et al., 2010), viridicatumtoxin and spirohexalines (Frisvad et al., 1989, Hayashi et al., 1994, Inokoshi et al., 2013, Inokoshi et al., 2016), and xanthoepocin (Tuthill et al., 2001, Bazioli et al., 2017), P. cataractum produces andrastin A-D, pulvilloric acid and trichodermamide A and C (Visagie et al. 2016b), P. echinulonalgiovense produces andrastin A and pulvilloric acid, P. mariae-crucis produces xanthomegnin and viomellein, P. onobense produces brefeldin A, janthitrems/shearinins, and 2-(4-hydroxyphenyl)-2-oxo acetaldehyde oxime, P. paraherquei produces fumitremorgins (Yoshizawa et al. 1976), paraherquonin (Okuyama et al. 1983), and paraherquamides (Yamazaki et al. 1981b), P. panissanguineum produces andrastins and pulvilloric acid (Visagie et al. 2016b), P. simplicissimum produces andrastin A, austalides (referred to as chromanols), paraherquamides, and xanthoepocin (Tuthill et al. 2001), P. tanzanicum produces bisdechlorogeodin, fiscalin C and pulvilloric acid (Visagie et al. 2016b) and P. wotroi produces pulvilloric acid (Houbraken et al. 2011b). In ser. Simplicissima many extrolites are in common, but they are also common in series Rolfsiorum, Dalearum and Janthinella. Series Oxalica species generally have least extrolites in common with the other series in sect. Lanata-Divaricata.



Notes on series of sect. Lanata-Divaricata: The deeper nodes within sect. Lanata-Divaricata are often poorly supported in Fig. 18, and the relationships between the series are therefore based on representative species in Supplementary Fig. S1. Using a multigene phylogenetic approach, Visagie et al. (2015) studied the relationship within sect. Lanata-Divaricata. Their new species resolved in three consistent, well-supported clades, named the P. janthinellum-, P. javanicum- and P. rolfsii-clade. In this study, sect. Lanata-Divaricata is subdidived in five series: Janthinella, Dalearum, Oxalica, Rolfsiorum and Simplicissima. The proposed P. janthinellum- and P. javanicum-clades in Visagie et al. (2015) are combined and equivalent to ser. Janthinella, and ser. Rolfsiorum includes the taxa of the P. rolfsii-clade. The majority of species outside the three clades of Visagie et al. (2015) belong to series Dalearum, Oxalica and Simplicissima, and exceptions are P. ehrlichii, P. meloforme, P. coeruleum, P. levitum (all ser. Janthinella), and P. vasconiae (ser. Rolfsiorum). The series classification in sect. Lanata-Divaricata is primarily based on the presented phylogeny and the series are difficult to distinguish phenotypically. Series Oxalica is distinct: species in this series produce spreading colonies, which are strictly velutinous and crustose and the conidia are ellipsoidal. None of the species in series Oxalica (three species), Rolfsiorum (16 species) and Simplicissima (21 species) are known to produce a sexual morph. In contrast, 13 out of the 24 ser. Janthinella species and three out of 13 ser. Dalearum species produce a sexual morph. The majority of species of series Oxalica and Janthinella are able to grow at 37 °C, while growth is more variable in the species of the section. Furthermore, ser. Rolfsiorum species produce rough-walled conidiophores and mostly globose conidia; however, these characters are shared with species in other series.



Section Lasseniorum Houbraken & Frisvad, sect. nov. MycoBank MB834283.



Type: Penicillium lassenii Paden, Mycopathol. Mycol. Appl. 43: 266. 1971.



Etymology: Named after the type species of the section, Penicillium lassenii.



Diagnosis: Phylogeny: Section Lasseniorum belongs to subgen. Aspergilloides and is phylogenetically most closely related to sect. Alfrediorum. Morphology & physiology: Colonies growing very restricted; conidial colour en masse pale grey-green; conidiophores monoverticillate or terminal biverticillate, smooth; conidia subglobose to ellipsoidal, smooth; growth at 37 °C absent. Sexual morph eupenicillium-type, homothallic, yellow-brown or tan; ascospores ellipsoidal, with two closely spaced ridges, convex smooth. Section description based on Paden (1971) and Pitt (1980).



Included species: Penicillium lassenii.



Extrolites: Penicillium lassenii produce extrolites which have not yet been structure elucidated.



Notes: Based on a multigene phylogenetic analysis, Houbraken & Samson (2011) accommodated P. lassenii, together with P. cryptum, in sect. Torulomyces. Visagie et al. (2016a) questioned the placement of P. lassenii (and P. cryptum) in sect. Torulomyces and this observation is confirmed here. Penicillium lassenii produces monoverticillate or terminal biverticillate conidiophores and smooth-walled conidia, while sect. Torulomyces species produce conidiophores with solitary phialides and typically roughened conidia (Paden, 1971, Visagie et al., 2016a). A series subdivision of the section is not proposed and therefore ser. Lasseniorum is only informally introduced here (Table 5).



Section Ochrosalmonea Houbraken & Samson, Stud. Mycol. 70: 33. 2011. MycoBank MB563127.



Type: Penicillium ochrosalmoneum Udagawa, J. Agric. Sci. Tokyo Nogyo Daig. 5: 10. 1959.



Description: See Houbraken & Samson (2011) (morphology, phylogeny), and here (under ser. Ochrosalmonea).



Series Ochrosalmonea Stolk & Samson, Adv. Pen. Asp. Syst.: 177. 1986 [1985]. MycoBank MB832728.



Type: Penicillium ochrosalmoneum Udagawa, J. Agric. Sci. Tokyo Nogyo Daig. 5: 10. 1959.



Diagnosis: Phylogeny: Series Ochrosalmonea belongs to subgen. Aspergilloides, sect. Ochrosalmonea and the series (and section) are phylogenetically related to Thysanophora. Morphology & physiology: Colonies growing restrictedly (P. isariiforme) or rapidly (P. ochrosalmoneum); conidial colour en masse greyish green or dull green; conidiophores predominantly biverticillate, occasionally with additional branches, smooth; conidia subglobose to apiculate (P. ochrosalmoneum) or ellipsoidal (P. isariiforme), smooth; growth at 37 °C absent (P. isariiforme) or present (P. ochrosalmoneum). Sexual morph produced in P. ochrosalmoneum, eupenicillium-type, homothallic, bright yellow or orange; ascospores ellipsoidal, with two longitudinal ridges, convex smooth to roughened or spinulose; sclerotia absent. Description based on Pitt (1980) and Houbraken & Samson (2011).



Included species: Penicillium isariiforme, P. ochrosalmoneum.



Extrolites: Both species in the series produce citreoviridin (Wicklow & Cole 1984).



Notes on series and section Ochrosalmonea: Penicillium ochrosalmoneum and P. isariifrome are phenotypically unrelated, but both are accommodated in sect. Ochrasalmonea based on the result of our phylogenetic analysis. For more details, see Houbraken & Samson (2011).



Section Ramigena Thom, The Penicillia: 156, 225. 1930. MycoBank MB834004.

Synonym: Penicillium sect. Fracta Houbraken & Samson, Stud. Mycol. 70: 35. 2011.



Type: Penicillium cyaneum (Bainier & Sartory) Biourge, Cellule 33: 102. 1923.



Description: See Houbraken & Samson (2011) (morphology, phylogeny); a taxonomic study on the section is lacking.



Series Georgiensia Houbraken & Frisvad, ser. nov. MycoBank MB834284.



Type: Penicillium georgiense S.W. Peterson & B.W. Horn, Mycologia 101: 79. 2009.



Etymology: Named after the type species of the series, Penicillium georgiense.



Diagnosis: Phylogeny: Series Georgiensia belongs to subgen. Aspergilloides, sect. Ramigena and is phylogenetically sister to ser. Ramigena. Morphology & physiology: Colonies restricted; conidial colour en masse pale to greyish green; conidiophores divaricate; stipes smooth; conidia ellipsoidal or subglobose, smooth-walled; growth at 37 °C present. Sexual morph unknown. Series description based on Peterson & Horn (2009).



Included species: Penicillium georgiense.



Extrolites: This species has not been examined for extrolites.



Series Ramigena Houbraken & Frisvad, ser. nov. MycoBank MB834285.



Type: Penicillium cyaneum (Bainier & Sartory) Biourge, Cellule 33: 102. 1923.



Etymology: Named after the type species of the series, Penicillium cyaneum.



Diagnosis: Phylogeny: Series Ramigena belongs to subgen. Aspergilloides, sect. Ramigena and is phylogenetically sister to ser. Georgiensia. Morphology & physiology: Colonies restricted; conidial colour en masse in shades of green (blue-green, dull green or grey-green); conidiophores strictly or predominantly monoverticillate, stipes smooth-walled; conidia generally ellipsoidal or pyriform, sometimes globose to subglobose (P. ornatum), smooth-walled; growth at 37 °C present or absent. Sexual morph generally not observed in culture, present in P. ornatum, eupenicillium-type, homothallic, buff to brown; ascospores broadly ellipsoidal, with two and sometimes four longitudinal flanges, convex smooth-walled. Series description based on Pitt (1980) and Houbraken & Samson (2011).



Included species: Penicillium capsulatum, P. cyaneum, P. hispanicum, P. ornatum, P. ramusculum.



Extrolites: Penicillium capsulatum, P. ornatum and P. ramusculum produce many unknown secondary metabolites, while P. cyaneum and P. hispanicum share production of deoxybrevianamide E. Penicillium hispanicum also produces asperflavin and bisanthrons.



Notes: Penicillium sect. Fracta is typified with Penicillium ornatum, a member sect. Ramigena. Section Fracta is therefore considered synonym of sect. Ramigena. Section Ramigena was introduced by Thom (1930) and the section was accepted in the classification proposed by Houbraken & Samson (2011). A taxonomic study of this section is lacking and the taxonomic status of P. cyaneum, P. dierckxii and P. sublateritium is unknown. Following Pitt, 1980, Houbraken and Samson, 2011 accepted these three species, even though RPB2 sequence data showed that these species are very closely related. This close relationship is supported by BenA and CaM sequences (>99.5 % homology) and we therefore treat P. dierckxii and P. sublateritium as synonyms of P. cyaneum. Penicillium georgiense is the sole member of ser. Georgiensia. Using sequence data, Peterson & Horn (2009) suggested a relationship of P. georgiense with P. thiersii (sect. Aspergilloides) and Houbraken & Samson (2011) classified this species in sect. Aspergilloides. While studying the taxonomy of sect. Aspergilloides, Houbraken et al. (2014b) showed that it is not part of this section and we here confidently classify P. georgiense in sect. Ramigena. Penicillium georgiense (ser. Georgiensia) differs from the species in the related ser. Ramigena by the production of divaricate branched conidiophores, while ser. Ramigena members are strictly or predominantly monoverticillate.



Section Sclerotiorum [as “Sclerotiora”] Houbraken & Samson, Stud. Mycol. 70: 32. 2011. MycoBank MB585167.



Type: Penicillium sclerotiorum J.F.H. Beyma, Zentralbl. Bakteriol. Parasitenk., Abt. 2 96: 418. 1937.



Description: See Houbraken and Samson, 2011, Visagie et al., 2013 (morphology, phylogeny).



Series Adametziorum Houbraken & Frisvad, ser. nov. MycoBank MB834286.



Type: Penicillium adametzii K.W. Zaleski, Bull. Int. Acad. Polon. Sci., Sér. B., Sci. Nat. 1927: 507. 1927.



Etymology: Named after the type species of the series, P. adametzii.



Diagnosis: Phylogeny: Series Adametziorum belongs to subgen. Aspergilloides, sect. Sclerotiorum and is phylogenetically sister to the other series of sect. Sclerotiorum. Morphology & physiology: Colonies restricted or moderately; conidial colour en masse dull green, grey-green; conidiophores monoverticillate, smooth-walled; conidia globose, subglobose or ellipsoidal, smooth or finely roughened; growth at 37 °C variable, absent or present. Sexual morph unknown; sclerotia absent or present, white (P. alexiae). Series description based on Visagie et al. (2013).



Included species: Penicillium adametzii, P. adametzioides, P. alexiae, P. amaliae, P. angulare, P. arianeae, P. bilaiae, P. brocae, P. jugoslavicum, P. lilacinoechinulatum, P. mellis, P. reconvexovelosoi∗, P. restingae [∗ not included in Fig. 18; for more information on the phylogenetic relationship, see Crous et al. (2019)].



Extrolites: Series Adametziorum species general produce dithiodiketopirazines such as brocazines, gliotoxins, epicoccins, phomazines, penicibrocazines, and spirobrocazines. Penicillium adametzioides produces glyanthrypine and lapatins, kotanins and spinulosins (Larsen et al., 2005, Liu et al., 2014b), Penicillium bilaiae produces citromycins, citromycetins, bilains, pistillarin and cyclic dipeptides (diketopiperazines) (Capon et al. 2007), hyalodendrins, 2-hydroxy-3,6-dimethylbenzaldehyde, dibutylphthalate, and 4-hydroxy-3,6-dimethyl-2H-pyran-2-one (Savard et al. 1994), penipratynolene and picolinic acids (Nakahara et al. 2004) and citric acid & oxalic acid (Cunningham & Kuiack 1992). Penicillium brocae produces brocazines, spirobrocazines, penibrocazines, phenopyrrozines, brocaeloids, brocaenols, pyranonigrin A, and brocapyrones (Bugni et al., 2003, Meng et al., 2014, Meng et al., 2015a, Meng et al., 2015b, Meng et al., 2016, Meng et al., 2017, Zhang et al., 2015b). Penicillium lilacinoechinulatum produces gliotoxins, phenopyrrozin, spinulosin and dehydrocarolic acid.



Series Herqueorum Houbraken & Frisvad, ser. nov. MycoBank MB834287.



Type: Penicillium herquei Bainier & Sartory, Bull. Soc. Mycol. France 28: 121. 1912.



Etymology: Named after the type species of the series, Penicillium herquei.



Diagnosis: Phylogeny: Series Herqueorum belongs to subgen. Aspergilloides, sect. Sclerotiorum and phylogenetically sister to ser. Sclerotiorum. Morphology & physiology: Colonies growing moderately fast; conidial colour en masse dull green, grey-green; conidiophores generally biverticillate, mono- and terverticillate also occasionally produced, smooth or rough-walled; conidia ellipsoidal, smooth or rough-walled; growth at 37 °C absent. Sexual morph generally not observed in culture, present in P. malachiteum, eupenicillium-type, homothallic, malachite-green; ascospores ellipsoidal, with two-equatorial ridges; sclerotia absent or present (P. herquei), cream. Series description based on Yaguchi et al., 1993, Visagie et al., 2013 and Wang et al. (2017a).



Included species: Penicillium choerospondiatis, P. herquei, P. malachiteum, P. sanshaense, P. verrucisporum.



Extrolites: Species in ser. Herqueorum produce atrovenetin, herqueinone, herqueichrysin, peniciherqueinone, peniciphenalines, sclerodin, scleroderolide, sclerodione, sculenonone A & B, and xanthoherquein (Stodola et al., 1951, Galarraga et al., 1955, Harman et al., 1955, Barton et al., 1959, Narasimhachari et al., 1963, Narasimhachari and Vining, 1963, Narasimhachari and Ramaswami, 1966, Narasimhachari and Vining, 1972, Simpson, 1976, Robinson et al., 1992, Trotter, 1992, Tansakul et al., 2014, Nishikori et al., 2016, Li et al., 2018a), herqulines (Furusaki et al., 1980, Enomoto et al., 1996, Chiba et al., 2017, Yang et al., 2018), neocyclocitrinol (Marinho et al. 2009), peniciherquamides (Nishikori et al. 2016), hualyzin (Ding et al. 2008), penicilquei A-D (Zhou et al. 2019), penicillic acid, and these metabolites have been found in both P. herquei and P. malachiteum. There are no secondary metabolites in common with series Adametziorum or Sclerotiorum.



Series Sclerotiorum Houbraken & Frisvad, ser. nov. MycoBank MB834288.



Type: Penicillium sclerotiorum J.F.H. Beyma, Zentralbl. Bakteriol. Parasitenk., Abt. 2 96: 418. 1937.



Etymology: Named after the type species of the series, Penicillium sclerotiorum.



Diagnosis: Phylogeny: Series Sclerotiorum belongs to subgen. Aspergilloides, sect. Sclerotiorum and is phylogenetically sister to ser. Herqueorum. Morphology & physiology: Colonies growing moderately fast; conidial colour en masse dull green or grey-green; conidiophores monoverticillate, smooth to rough; conidia globose, subglobose or ellipsoidal, smooth to finely rough; growth at 37 °C variable, absent or present. Sexual morph produced in P. hirayamae, eupenicillium-type, homothallic, orange or brown; ascospores ellipsoidal, with two small closely appressed longitudinal flanges, convex rough-walled; sclerotia produced in P. acidum, P. austrosinicum, P. johnkrugii, P. sclerotiorum, P. vanoranjei, orange, white/grey, cream to yellow. Series description based on Rivera and Seifert, 2011, Visagie et al., 2013 and Wang et al. (2017a)



Included species: Penicillium acidum, P. austrosinicum, P. cainii, P. circulare, P. daejeonium, P. exsudans, P. fernandesiae, P. guanacastense, P. hirayamae, P. jacksonii, P. johnkrugii, P. mallochii, P. maximae, P. meliponae, P. sclerotiorum, P. vanoranjei, P. viticola.



Extrolites: All species examined in ser. Sclerotiorum produce sclerotiorins, rotiorin, penazaphilones, penicilazaphilones (Curtin and Reilly, 1940, Maccurin and Reilly, 1940, Birkinshaw, 1952, Jackman et al., 1958, Arai et al., 1995, Matsuzaki et al., 1995a, Matsuzaki et al., 1995b, Pairet et al., 1995, Nam et al., 2000, Lucas et al., 2007, Arunpanichlert et al., 2010, Hemtasin et al., 2016, Zhou et al., 2016, Wang et al., 2018, Jia et al., 2019, Tang et al., 2019a, Wu et al., 2019), penicilisorin, (+)-4,6-dimetylocta-2,4-dienoic acid and 5,6-dihydro-3,5,6-trimethylpyran-2-one (Arunpanichlert et al. 2010), pencolide (Birkinshaw et al., 1963, Lucas et al., 2007), carotenes (Mase et al. 1957), multicolic acid, multicolosic acid and multicolanic acid (Gudgeon et al., 1974, Gedge and Pattenden, 1979, Gudgeon et al., 1979, Holker et al., 1987) and atlantinone A. In general, all species with a reddish reverse in ser. Sclerotiorum produce azaphilones such as sclerotiorin.



Notes on series of sect. Sclerotiorum: Visagie et al. (2013) divided sect. Sclerotiorum in three main clades. These clades are also observed in our phylogenetic study (Fig. 18), and we treat those clades here as series. Specific phenotypic characters that define the three series of the section could not be identified, but there are some polythetic features for each series (Visagie et al. 2013). Generally, species in ser. Herqueorum produce biverticillate conidiophores, in contrast with species of series Sclerotiorum and Adametziorum, which are monoverticillate. Taxa in ser. Sclerotiorum have colonies in orange colours and lack the strongly coloured, soluble pigments such as those generally observed in ser. Adametziorum species.



Section Stolkia Houbraken & Samson, Stud. Mycol. 70: 38. 2011. MycoBank MB563130.



Type: Penicillium stolkiae D.B. Scott, Mycopathol. Mycol. Appl. 36: 8. 1968.



Diagnosis: Phylogeny: Section Stolkia belongs to subgen. Aspergilloides and is phylogenetically related sister to sect. Lanata-Divaricata. Morphology & physiology: Colonies restricted or growing moderately fast; conidial colour en masse variable (dark green, greyish green, grey, blue green); conidiophores monoverticillate (P. alogum, P. boreae, P. canariense, P. donkii, P. pullum, P. stolkiae, P. subarcticum) or biverticillate (P. boreae, P. canariense), smooth, with some brown pigmentation; conidia globose, smooth or (finely) roughened; growth at 37 °C absent. Sexual morph not observed in culture, or present (P. stolkiae), eupenicillium-type, homothallic, orange-brown; ascospores ellipsoidal, with two well-defined longitudinal ridges, convex conspicuously roughened; sclerotia not produced. Section description based on Pitt, 1980, Peterson and Sigler, 2002, Houbraken and Samson, 2011 and Visagie et al. (2016b).



Included species: Penicillium alogum, P. boreae, P. canariense, P. donkii, P. pullum, P. stolkiae, P. subarcticum.



Extrolites: The are no chemotaxonomic data available for species in sect. Stolkia.



Notes: See Houbraken & Samson (2011). A series subdivision of the section is not proposed and therefore ser. Stolkia is only informally introduced here (see Table 5).



Section Thysanophora Houbraken & Samson, Stud. Mycol. 70: 33. 2011. MycoBank MB563126.



Type: Sclerotium glaucoalbidum Desm., Ann. Sci. Nat. Bot. ser. 3, 16: 329. 1851. MycoBank MB212120 (= Penicillium glaucoalbidum).



Diagnosis: Phylogeny: Section Thysanophora belongs to subgenus Aspergilloides and is phylogenetically sister to sect. Ochrosalmonea. Morphology & physiology: Colonies growing restrictedly; conidial colour en masse dark brown; conidiophores mono-, bi- or terverticillate, often with secondary, subapical formation of a penicillius (resulting in a chain of sympodial penicilli), dark brown pigmented, smooth or distinctly ornamented; conidia subglobose, ellipsoidal, fusiform or ovoidal, hyaline or in shades or brown, smooth or rough-walled. Sexual morph unknown; sclerotia pale brown, dark brown. Species are often associated with leaves of various trees (pine needles). Section description based on Ellis, 1971, Mercado-Sierra et al., 1998 and Houbraken & Samson (2011).



Included species: Penicillium asymmetricum∗, P. coniferophilum∗, P. glaucoalbidum∗, P. hennebertii, P. longisporum∗, P. melanostipe∗, P. taiwanense∗, P. taxi (∗ not included in Fig. 18, no sequence data available for these species).



Extrolites: There are no data for secondary metabolites in section Thysanophora.



Notes: A modern taxonomic study of this section is lacking and needed, and we therefore follow Houbraken & Samson (2011). A series subdivision of the section is not proposed and therefore ser. Thysanophora is only informally introduced here (Table 5).



Section Torulomyces (Delitsch) Stolk & Samson, Adv. Pen. Asp. Syst.: 169. 1986 [1985]. MycoBank MB832720.



Type: Penicillium lagena (Delitsch) Stolk & Samson, Stud. Mycol. 23: 100. 1983.



Diagnosis: Phylogeny: Section Torulomyces belongs to subgen. Aspergilloides and is phylogenetically sister to sect. Crypta. Morphology & physiology: Colonies restricted; conidial colour en masse greyish green, greenish white; conidiophores as solitary phialides, smooth; conidia globose or subglobose, occasionally broadly ellipsoidal (P. catalonicum, P. variratense), rough-walled; growth at 37 °C generally absent, present in P. aeris, P. cantabricum and P. riverlandense. Sexual morph generally not observed in culture, present in P. wollemiicola, eupenicillium-type, homothallic, colourless; ascospores ellipsoidal with two widely separated ridges, convex smooth-walled; sclerotia generally absent. Section description based on Visagie et al. (2016a).



Included species: Penicillium aeris, P. austricola, P. cantabricum, P. catalonicum, P. lagena, P. marthae-christenseniae, P. oregonense, P. parviverrucosum∗, P. porphyreum, P. riverlandense, P. tubakianum, P. variratense, P. williamettense, P. wisconsinense, P. wollemiicola (∗ not included in Fig. 18, no sequence data available for this species).



Extrolites: There are no chemotaxonomic data available for section Torulomyces.



Notes: Visagie et al. (2016a) studied the taxonomy of sect. Torulomyces and accepted 18 species. Penicillium cryptum (sect. Crypta), P. lassenii (sect. Lasseniorum), P. laeve and P. ovatum (sect. Exilicaulis) are transferred here to other sections, and P. parviverrucosum is tentatively accepted. By excluding P. cryptum, P. lassenii, P. laeve and P. ovatum, sect. Torulomyces becomes more homogenous and all species produce conidiophores with solitary phialides and rough-walled conidia. A series subdivision of the section is not proposed and therefore ser. Torulomyces is only informally introduced here (Table 5).



Penicillium subgen. Penicillium [autonym]. MycoBank MB92187.

Synonym: Penicillium subgen. Eupenicillium Dierckx, Ann. Soc. Sci. Bruxelles 25: 86. 1901.



Type: Penicillium expansum Link, Mag. Ges. Naturf. Freunde Berlin 3: 16. 1809.



Description: See Frisvad & Samson (2004b) (extrolites, morphology), Houbraken & Samson (2011) (morphology, phylogeny), Houbraken et al. (2016).



Section Brevicompacta Thom, The Penicillia: 157, 289. 1930. MycoBank MB834006.

Synonym: Penicillium sect. Coronata Pitt, The Genus Penicillium: 392. 1980 [1979].



Type: Penicillium brevicompactum Dierckx, Ann. Soc. Sci. Bruxelles 25: 88. 1901.



Description: See Frisvad & Samson (2004b) (as sect. “Coronata”; extrolites, morphology), Houbraken & Samson (2011) (morphology, phylogeny).



Series Brevicompacta Houbraken & Frisvad, ser. nov. MycoBank MB834482.



Etymology: Named after the type species of the series, Penicillium brevicompactum.



Type: Penicillium brevicompactum Dierckx, Ann. Soc. Sci. Bruxelles 25: 88. 1901.



Diagnosis: Phylogeny: Series Brevicompacta belongs to subgen. Penicillium, sect. Brevicompacta and is phylogenetically most closely related to ser. Olsoniorum. Morphology & physiology: Colonies restricted, colony texture velvety; conidiophores terverticillate, sometimes biverticillate or multiramulate, smooth-walled, wide; conidia subglobose or ellipsoidal, smooth or finely roughened. Sexual morph unknown; sclerotia present in P. neocrassum, brown. Series description based on Frisvad & Samson (2004b) and Serra & Peterson (2007).



Included species: Penicillium bialowiezense, P. brevicompactum, P. fennelliae, P. kongii, P. neocrassum.



Extrolites: The Raistrick phenols and mycophenolic acid are only found in ser. Brevicompacta within sect. Brevicompacta. Botryodiploidin is found in one species in ser. Brevicompacta (P. kongii). Xanthoepocin is produced by three out of five species in ser. Brevicompacta and by all species in ser. Olsoniorum. The breviones have been found in two species in ser. Brevicompacta, one species in ser. Spathulata and one species in ser. Olsoniorum. Quinolactacin is produced by one species in ser. Brevicompacta (P. bialowiezense) and one species in ser. Spathulata.



Series Buchwaldiorum Houbraken & Frisvad, ser. nov. MycoBank MB833972



Etymology: Named after the type species of the series, Penicillium buchwaldii.



Type: Penicillium buchwaldii Frisvad & Samson, FEMS Microbiol. Lett. 339: 86. 2013.



Diagnosis: Phylogeny: Series Buchwaldiorum belongs to subgen. Penicillium, sect. Brevicompacta and is phylogenetically most closely related to ser. Tularensia, though statistical support for this relationship is lacking (Fig. 19). Morphology & physiology: Colonies growing restrictedly (P. spathulatum) or moderately rapid (P. buchwaldii), colony texture velvety; conidiophores terverticillate, sometimes bi- or quarterverticillate, smooth to finely roughened; conidia globose or subglobose, rough-walled. Sexual morph unknown; sclerotia not observed in culture. Series description based on Frisvad et al. (2013a).



Fig. 19.

Combined phylogeny for BenA, CaM and RPB2 data sets showing the phylogenetic relation of species, series and sections within Penicillium subgen. Penicillium. The BI posterior probability (pp) values and bootstrap percentages of the maximum likelihood (ML) analysis are presented at the nodes; fully supported branches are thickened. Values less than 70 % bootstrap support (ML) or less than 0.95 posterior probability (Bayesian analysis) are indicated with a hyphen or not shown. The bar indicates the number of substitutions per site. The phylogram is rooted with Aspergillus glaucus and Hamigera avellanea.

Included species: Penicillium buchwaldii, P. spathulatum.



Extrolites: The extrolites only found in this series (in sect. Brevicompacta) include asperentins, communesins, paraherquamide, benzomalvins, cyclopenol and 2-chloro-6-[2’(S)-hydroxypropyl]-1,3,8-trihydroxy-anthraquinone (Frisvad et al. 2013a, Del Valle et al. 2016).



Series Olsoniorum [as “Olsonii” Pitt, The Genus Penicillium (London): 392. 1980 [1979]. MycoBank MB833052.



Type: Penicillium olsonii Bainier & Sartory, Ann. Mycol. 10: 398. 1912.



Diagnosis: Phylogeny: Series Olsoniorum belongs to subgen. Penicillium, sect. Brevicompacta and is phylogenetically related to ser. Brevicompacta. Morphology & physiology: Colonies growing restrictedly or moderately rapid, colony texture velvety; conidiophores multiramulate, smooth-walled, long, wide; conidia ellipsoidal, smooth-walled or finely roughened. Sexual morph unknown; sclerotia observed in some strains of P. olsonii and P. salami, pale to light yellow or greyish brown. Series description based on Frisvad and Samson, 2004b, Serra and Peterson, 2007 and Perrone et al. (2015).



Included species: Penicillium astrolabium, P. olsonii, P. salamii.



Extrolites: Extrolites in ser. Olsoniorum include some that are only found in this series in sect. Brevicompacta (chrysogine, 2-(4-hydroxyphenyl)-2-oxo acetaldehyde oxime, meleagrin, siderin and verruculone (Perrone et al. 2015), showing a close relationship between those two series.



Series Tularensia (Pitt) Houbraken & Frisvad, comb. nov. MycoBank MB833973.

Basionym: Eupenicillium ser. Tularensia Pitt, The Genus Penicillium: 98. 1980 [1979]. MycoBank MB832946.



Type: Eupenicillium tularense Paden, Mycopathol. Mycol. Appl. 43: 262. 1971 (= Penicillium tularense).



Diagnosis: Phylogeny: Series Tularensia belongs to subgen. Penicillium, sect. Brevicompacta and is phylogenetically related to ser. Buchwaldiorum, though statistical support is lacking (Fig. 19). Morphology & physiology: Colonies restricted, colony texture velvety; conidiophores predominant terverticillate, occasionally biverticillate, smooth-walled; conidia globose to broadly ellipsoidal, smooth. Sexual morph eupenicillium-type, homothallic, light brown to pale tan; ascospores ellipsoidal, with two longitudinal flanges and rugose walls. Series description based on Paden, 1971, Pitt, 1980, Stolk and Samson, 1983 and Frisvad et al. (2013a).



Included species: Penicillium tularense.



Extrolites: Apart from asperphenamate, shared with nearly all other species in sect. Brevicompacta, P. tularense produces a series of indoloterpenes unique for this species, including paxillins, and the related paspalines and janthitrems / shearinins, in addition to emindoles (Andersen and Frisvad, 2004, Frisvad et al., 2013a).



Notes: Section Brevicompacta species produce terverticillate or multiramulate branched conidiophores that superficially resemble Aspergillus in the stereomicroscope. The position of Penicillium fennelliae in sect. (and ser.) Brevicompacta is remarkable, because this species produces penicillic acid, an extrolite not shared by any other species in the section. Furthermore, it does not produce asperphenamate, a compound shared by all other sect. Brevicompacta species (Frisvad et al. 2013a). The section is split in four series: Brevicompacta, Buchwaldiorum, Olsoniorum and Tularensia. Series Brevicompacta and Olsoniorum can be differentiated by their conidiophore branching pattern: the former has terverticillate conidiophores; the latter multiramulate ones. Species in ser. Tularensia produce a sexual morph and this is not observed in the series Brevicompacta, Buchwaldiorum and Olsoniorum. Sclerotia have been found in some strains of P. olsonii and P. salami (of ser. Olsoniorum).



Section Canescentia Houbraken & Samson, Stud. Mycol. 70: 46. 2011. MycoBank MB563135.



Type: Penicillium canescens Sopp, Skr. Vidensk.-Selsk. Christiana Math.-Nat. Kl. 11: 181. 1912.



Description: See Houbraken & Samson (2011), this study (phylogeny); a modern overview including phenotypic data is not yet published.



Series Atroveneta Stolk & Samson, Adv. Pen. Asp. Syst.: 175. 1986 [1985]. MycoBank MB832726.



Type: Penicillium atrovenetum G. Sm., Trans. Brit. Mycol. Soc. 39: 112. 1956.



Diagnosis: Phylogeny: Series Atroveneta belongs to subgen. Penicillium, sect. Canescentia and is phylogenetically related to ser. Canescentia. Morphology & physiology: Colonies growing moderately fast to fast, colony texture velutinous, exudate often formed on CYA; conidia en masse from dark green to turquoise; conidiophores biverticillate and appressed with an occasional additional branch, smooth- or rough-walled; conidia globose to ellipsoidal, smooth-walled or roughened; conidia 2.5–3.5 μm; no growth at 37 °C. Sexual morph unknown; sclerotia produced by some species.



Included species: Penicillium antarcticum, P. atrovenetum, P. coralligerum, P. novae-zeelandiae, P. nucicola.



Extrolites: Species in ser. Atroveneta produce different extrolites than those in ser. Canescentia: Asperentins, atrovenetins, benzomalvins, citreoviridin, cycloaspeptides, naphthalic anhydride, 3-nitropropionic acid, patulin, while chrysogines, communesins, penitrems and xanthoepocin are shared with some species in ser. Canescentia (Turner and Aldridge, 1971, Frisvad and Filtenborg, 1990, Shiono et al., 2008, Vansteelandt et al., 2012, Geiger et al., 2013, Takahashi et al., 2017).



Series Canescentia Raper & Thom ex Pitt, The Genus Penicillium: 251. 1980 [1979]. MycoBank MB832960.



Type: Penicillium canescens Sopp, Skr. Vidensk.-Selsk. Christiana Math.-Nat. Kl. 11: 181. 1912.



Diagnosis: Phylogeny: Series Canescentia belongs to subgen. Penicillium, sect. Canescentia and is phylogenetically related to ser. Atroveneta. Morphology & physiology: Colonies growing moderately fast, colony texture floccose; conidiophores formed mostly on aerial mycelium, biverticillate with an occasional additional branch, divaricate, smooth- or rough-walled; conidia globose to ellipsoidal, smooth-walled or roughened; no growth at 37 °C. Sexual morph unknown. All species are soil- and litter-borne.



Included species: Penicillium arizonense, P. canescens, P. corvianum, P. dunedinense, P. griseoazureum, P. janczewskii, P. jensenii, P. murcianum, P. nigricans, P. radiatolobatum, P. yarmokense.



Extrolites: Acetylaranotins, austalides, curvulinic acids, fumagillins, griseofulvin, pseurotins, pyripyropens, trichodermamides and tryptoquivalines (Turner and Aldridge, 1971, Turner and Aldridge, 1983, Grijseels et al., 2016) are produced by some species in ser. Canescentia, but not by any species in ser. Atroveneta.



Notes: Two phylogenetically related series are accepted in sect. Canescentia: Canescentia and Atroveneta. These series can be differentiated by different extrolite profiles and by their colony texture, which is typically floccose in species in ser. Canescentia and velutinous in ser. Atroveneta.



Section Chrysogena Frisvad & Samson, Stud. Mycol. 49: 17. 2004. MycoBank MB700796.



Type: Penicillium chrysogenum Thom, U.S.D.A. Bur. Animal Industr. Bull. 118: 58. 1910.



Description: See Frisvad et al. (2004) (extrolites, morphology), Houbraken & Samson (2011) (phylogeny), Houbraken et al. (2012a) (morphology, phylogeny), Houbraken et al. (2016) (extrolites, morphology, phylogeny).



Series Aethiopica Frisvad & Samson, Stud. Mycol. 49: 19. 2004. MycoBank MB700800.



Type: Penicillium aethiopicum Frisvad, Mycologia 81: 848. 1989 (current name: Penicillium lanosocoeruleum).



Diagnosis: Phylogeny: Series Aethiopica belongs to subgen. Penicillium, sect. Chrysogena and is phylogenetically most closely related to ser. Chrysogena. Morphology & physiology: Colonies growing moderately fast, colony texture fasciculate, copious production of exudate droplets on CYA; conidiophores terverticillate, rough-walled; conidia ellipsoidal, smooth-walled; growth at 37 °C, slower growth on CYA with 5 % NaCl (CYAS) than on CYA. Sexual morph unknown; sclerotia not observed in culture. Series description based on Frisvad & Samson (2004b) and Houbraken et al. (2012a).



Included species: Penicillium lanosocoeruleum.



Extrolites: The only species in ser. Aethiopica, P. lanosocoeruleum (= P. aethiopicum), produces the extrolites griseofulvin, tryptoquialanines and viridicatumtoxin, and these extrolites are not produced by other species of sect. Chrysogena (Frisvad & Samson 2004b).



Series Chrysogena Raper & Thom ex Stolk & Samson, Adv. Pen. Asp. Syst.: 180. 1986 [1985]. MycoBank MB832730.

Synonym: Penicillium ser. Mononematosa Frisvad, Int. Mod. Meth. Pen. Asp. Clas. 269. 2000.



Type: Penicillium chrysogenum Thom, U.S.D.A. Bur. Animal Industr. Bull. 118: 58. 1910.



Diagnosis: Phylogeny: Series Chrysogena belongs to subgen. Penicillium, sect. Chrysogena and is phylogenetically most closely related to ser. Aethiopica. Morphology & physiology: Colonies growing moderately fast, colony texture velutinous, exudate droplets often present on CYA; conidiophores ter- to quarterverticillate, divergent, smooth; conidia globose to ellipsoidal, smooth; no or poor growth at 37 °C; growth on CYAS similar or more rapid than on CYA. Sexual morph generally not observed in culture, present under specific conditions in P. rubens (reported as P. chrysogenum by Böhm et al. (2013)), eupenicillium-type, heterothallic. Series description based on Frisvad & Samson (2004b) and Houbraken et al. (2012a).



Included species: Penicillium allii-sativi, P. chrysogenum, P. confertum, P. desertorum, P. dipodomyis, P. flavigenum, P. halotolerans, P. mononematosum, P. nalgiovense, P. rubens, P. tardochrysogenum, P. vanluykii.



Extrolites: Most species in ser. Chrysogena produce andrastins, chrysogine, meleagrin and roquefortine C, penicillin, sorbicillins (species with yellow exudate), while few species produce bioxanthracenes, citreoisocoumarins, chrysogenamide, cyclopaldic acids, dipodazin, haenamindole, nalgiovensins, penitrems, PR-toxins, secalonic acids, verrucosidins, verruculogen (and fumitremorgins), viriditoxins, xanthocillins, yanuthones Frisvad and Samson, 2004b, Frisvad et al., 2004 and Houbraken et al. (2012a).



Series Goetziorum Houbraken & Frisvad, ser. nov. MycoBank MB833974.



Etymology: Named after the type species of the series, Penicillium goetzii.



Type: Penicillium goetzii J. Rogers et al., Persoonia 29: 92. 2012.



Diagnosis: Phylogeny: Series Goetziorum belongs to subgen. Penicillium, sect. Chrysogena and is phylogenetically sister to a clade containing series Aethiopica, Chrysogena and Persicina. Morphology & physiology: Colonies growing moderately fast, colony texture velvety to slightly floccose; conidiophores ter- to quarterverticillate, smooth-walled; conidia broadly ellipsoidal, smooth; no growth at 37 °C; growth on CYAS similar to CYA. Sexual morph eupenicillium-type, homothallic, creamish brown; ascospores ellipsoidal, with two distinct equatorial ridges, valves reticulate. Series description based on Houbraken et al. (2012a).



Included species: Penicillium goetzii.



Extrolites: Aflavinines, andrastin A, citreoisocoumarins (Wang et al., 1995, Houbraken et al., 2012a).



Series Crustacea (Pitt) Houbraken & Frisvad, comb. nov. MycoBank MB833975.

Basionym: Eupenicillium ser. Crustacea Pitt, The Genus Penicillium: 139. 1980 [1979]. MycoBank MB832950.



Type: Eupenicillium crustaceum F. Ludw., Lehrb. Nied. Krypt.: 263. 1892 (current name: Penicillium kewense).



Diagnosis: Phylogeny: Series Crustacea belongs to subgen. Penicillium, sect. Chrysogena and is phylogenetically sister to a clade containing series Aethiopica, Chrysogena, Goetziorum and Persicina. Morphology & physiology: Colonies growing restrictedly or moderately fast, colony texture velvety or floccose; colony morphology dominated by sclerotia or ascomata, conidiophores predominantly terverticillate, sometimes bi- or quarterverticillate, appressed, smooth-walled; conidia globose to subglobose, smooth-walled. Sexual morph eupenicillium-type, homothallic, creamish, avellaneous or ochraceous; ascospores broadly ellipsoidal, with two distinct equatorial flanges, valves smooth or roughened. Series description based on Stolk & Samson (1983).



Included species: Penicillium egyptiacum, P. kewense, P. sinaicum.



Extrolites: Macrophorins, penicillic acid, secalonic acid D, xanthocillin X (Wang et al., 1995, Fujimoto et al., 2001).



Series Persicina Frisvad & Samson, Stud. Mycol. 49: 19. 2004. MycoBank MB700803.



Type: Penicillium persicinum L. Wang et al., Antonie van Leeuwenhoek 86: 177. 2004.



Diagnosis: Phylogeny: Series Persicina belongs to subgen. Penicillium, sect. Chrysogena and is phylogenetically sister to a clade containing series Aethiopica and Chrysogena. Morphology & physiology: Colonies growing moderately fast, colony texture velvety; conidiophores terverticillate, sometimes quarterverticillate; conidia ellipsoidal or cylindrical, smooth; growth at 37 °C, growth CYAS > CYA. Sexual morph unknown; sclerotia not observed in culture. Series description based on Frisvad & Samson (2004b).



Included species: Penicillium persicinum.



Extrolites: Penicillium persicinum, included in ser. Persicina, produces chrysogines, griseofulvins and roquefortines (Wang et al. 2004).



Notes: The majority of sect. Chrysogena species are characterised by the formation of smooth-walled, divergent, ter- to quarterverticillate conidiophores with relatively small phialides. The species in this section are tolerant to salt and the majority produce colonies with a velvety texture. This section is divided in five series: Aethiopica, Chrysogena, Crustacea, Goetziorum and Persicina. All species in series Goetziorum and Crustacea produce a sexual morph in a homothallic manner and this is not observed in the other series. Penicillin production is restricted to ser. Chrysogena. Series Aethiopica and Persicina are monotypic series and both share the ability to produce griseofulvin, an extrolite not produced by other sect. Chrysogena species. Series Aethiopica is characterised by the production of fasciculate colonies and rough-walled conidiophores; ser. Persicina produces appressed, smooth-walled conidiophores and ellipsoidal to cylindrical conidia, a combination of features not observed in series Chrysogena and Aethiopica (Frisvad & Samson 2004b).



Section Eladia (G. Smith) Stolk & Samson, Adv. Pen. Asp. Syst.: 171. 1986 [1985]. MycoBank MB832721.



Type: Penicillium sacculum E. Dale, Ann. Mycol. 24: 137. 1926.



Diagnosis: Phylogeny: Section Eladia belongs to subgen. Penicillium and is phylogenetically most closely related to sect. Canescentia. Morphology & physiology: Colonies restricted or growing moderately fast, colony texture velvety; conidiophores monoverticillate, occasionally biverticillate; conidia globose to subglobose, rough-walled, often with prominent connectives; phialides with a swollen base and very short neck, also born irregularly on the stipe. Sexual morph absent (P. sacculum) or present (P. senticosum), eupenicillium-type, homothallic, pale brown; ascospores ellipsoidal with two longitudinal flanges and echinulate walls. Section description based on Pitt, 1980, Stolk and Samson, 1983, Stolk and Samson, 1985 (morphology), Houbraken & Samson (2011) (morphology, phylogeny).



Included species: Penicillium sacculum, P. senticosum.



Extrolites: Penicillium senticosum produces curvularins and sorbicillins.



Notes: This section is phylogenetically most closely related to sect. Canescentia. The production of ornamented conidia (often with prominent connectives), phialides with very short necks and the irregularly position of these phialides on the conidiophore stipes confirms the distinct phylogenetic position of this series in Penicillium (Pitt, 1980, Stolk and Samson, 1983, Stolk and Samson, 1985). Section Eladia is not subdivided in series and therefore ser. Eladia is informally introduced in this article.



Section Fasciculata Thom, The Penicillia: 158, 374. 1930. MycoBank MB834008.

Synonyms: Penicillium sect. Lanata-typica Thom, The Penicillia: 157, 305. 1930.

Penicillium sect. Viridicata Frisvad & Samson, Stud. Mycol. 49: 27. 2004.



Type: Penicillium hirsutum Dierckx, Ann. Soc. Sci. Bruxelles 25: 89. 1901.



Description: See Frisvad & Samson (2004b) (as sect. Viridicata; morphology, extrolites), Houbraken & Samson (2011) (phylogeny), Houbraken et al. (2016) (morphology, phylogeny).



Series Camembertiorum [as “Camembertii”] Raper & Thom ex Pitt, The Genus Penicillium: 358. 1980 [1979]. MycoBank MB833061.

Synonym: Penicillium series Solita Frisvad, Int. Mod. Meth. Pen. Asp. Clas.: 279. 2000.



Type: Penicillium camemberti Thom, U.S.D.A. Bur. Animal Industr. Bull. 82: 33. 1906.



Diagnosis: Phylogeny: Series Camembertiorum belongs to subgen. Penicillium, sect. Fasciculata and forms a clade with series Corymbifera, Verrucosa and Viridicata. Morphology & physiology: Colonies growing moderately fast, colony texture floccose, granular or crustose, but not distinctly fasciculate; conidiophores terverticillate, rough-walled; conidia ellipsoidal or globose to subglobose, smooth or rough-walled (P. cavernicola, P. discolor, P. echinulatum, P. solitum); good growth and (delayed) base production on creatine agar (CREA). Sexual morph unknown; sclerotia not observed in culture. Species typically occur on proteinaceous and lipid-containing foods. Series description based on Frisvad & Samson (2004b).



Included species: Penicillium biforme, P. camemberti, P. caseifulvum, P. cavernicola, P. commune, P. crustosum, P. discolor, P. echinulatum, P. palitans, P. solitum, P. speluncae.



Extrolites: Most of the species are able to produce viridicatins, cyclopenins and palitantin, while some species produce cyclopiazonic acids, cyclopaldic acid, territrems and arisugacins, compactins, penitrems, andrastins and atlantinones, asteltoxin, aurantiamine, dipodazine, glyanthrypine, roquefortine, clavatols, penitrems and terrestric acids (the latter four only by P. crustosum), burnettienes, geosmin, fumigaclavines, chaetoglobosins, daldinin D, and rugulovasines (Frisvad & Samson 2004b).



Series Corymbifera Frisvad, Int. Mod. Meth. Pen. Asp. Clas.: 275. 2000. MycoBank MB700801.



Type: Penicillium hirsutum var. albocoremium Frisvad, Mycologia 81: 856. 1990 (current name: Penicillium albocoremium).



Diagnosis: Phylogeny: Series Corymbifera belongs to subgen. Penicillium, sect. Fasciculata and forms a clade with series Camembertiorum, Verrucosa and Viridicata; the position of P. venetum in this series is uncertain. Morphology & physiology: Colonies growing moderately fast, colony texture fasciculate or coremiform; most species produce coloured exudate droplets on CYA and MEA and grows moderately well on CREA, and acid is produced on CREA; conidiophores terverticillate, conspicuously roughened (except P. hordei, smooth to finely roughened), appressed; conidia globose or subglobose, smooth-walled (roughened in P. hordei). Sexual morph unknown; sclerotia not produced in culture. Species are mainly associated with flower bulbs and occasionally other plant roots. Series description based on Frisvad & Samson (2004b).



Included species: Penicillium albocoremium, P. allii, P. hirsutum, P. hordei, P. mali-pumilae, P. psychrotrophicum, P. radicicola, P. tulipae, P. venetum.



Extrolites: Roquefortine C and D are present in most species and meleagrin in some species. Cyclopenins and viridicatins are also present in several species in ser. Corymbifera. Atrovenetins and terrestic acids are also common in the series. Barceloneic acids, chrysogine, citrinin, compactins, daldinin D, ergometrine, fulvic acid, penicillic acid, penitrems are produced by some species in ser. Corymbifera (Overy and Frisvad, 2003, Frisvad and Samson, 2004b, Frisvad et al., 2004, Overy et al., 2005a, Overy et al., 2005b, Overy et al., 2005c, Hallas-Moller et al., 2018).



Series Gladioli Raper & Thom ex Stolk & Samson, Adv. Pen. Asp. Syst.: 183. 1986 [1985]. MycoBank MB832732.



Type: Penicillium gladioli L. McCulloch & Thom, Science 67: 217. 1928.



Diagnosis: Phylogeny: Series Gladioli belongs to subgen. Penicillium, sect. Fasciculata and is phylogenetically basal to the other series of the section. Morphology & physiology: Colonies growing restrictedly to moderately rapid, colony texture floccose to slightly fasciculate; conidiophores terverticillate, occasionally more complexly branched, rough-walled; conidia subglobose, smooth-walled; growth on CREA moderate to good. Sexual morph not observed in culture; sclerotia produced, avellaneous, pale brown. Series Gladioli is associated with gladiolus bulb rot (Frisvad & Samson 2004b).



Included species: Penicillium gladioli.



Extrolites: The only species in the series produce atrovenetins, gladiolic acids, glyanthrypine and patulin (Frisvad & Samson 2004b).



Series Verrucosa Frisvad, Int. Mod. Meth. Pen. Asp. Clas.: 269. 2000. MycoBank MB700806.



Type: Penicillium verrucosum Dierckx, Ann. Soc. Sci. Bruxelles 25: 88. 1901.



Diagnosis: Phylogeny: Series Verrucosa belongs to subgen. Penicillium, sect. Fasciculata and forms a clade with series Camembertiorum, Corymbifera and Viridicata. Morphology & physiology: Colonies restricted, colony texture velvety, floccose, or weakly fasciculate; conidiophores terverticillate; conidia globose to subglobose, rough-walled; weak growth on CREA, and no acid production; the three species in ser. Verrucosa grow well on nitrite-sucrose agar. Sexual morph unknown; sclerotia not observed in culture. Species are associated with stored cereal grains (P. verrucosum) and dried or salted meat products; they can also grow on cheese (Frisvad & Samson 2004b).



Included species: Penicillium nordicum, P. thymicola, P. verrucosum.



Extrolites: All species in Verrucosa produce verruculones and ochratoxins (Frisvad and Samson, 2004b, Nguyen et al., 2016). Some species produce alantrypinones, anacines, burnettienes, citrinin, daldinin D, fumiquinazolines, lumpidin, pyranonigrins, sclerotigenin, verrucins and viridic acid (Larsen et al., 1998, Ariza et al., 2001, Rahbæk et al., 2003, Frisvad and Samson, 2004b, Zhelifonova et al., 2012, Ma et al., 2017a, Tang et al., 2018).



Series Viridicata Raper & Thom ex Pitt, The Genus Penicillium: 334. 1980 [1979]. MycoBank MB832967.



Type: Penicillium viridicatum Westling, Ark. Bot. 11: 88. 1911.



Diagnosis: Phylogeny: Series Viridicata belongs to subgen. Penicillium, sect. Fasciculata and forms a clade with series Camembertiorum, Corymbifera and Verrucosa. Morphology & physiology: Colonies growing moderately fast, colony texture granular or velvety; conidiophores terverticillate; rough-walled, growth on CREA weak to moderate, no base production; conidia ellipsoidal or globose to subglobose, smooth or very finely roughened (except echinulate in P. neoechinulatum). Sexual morph unknown; sclerotia not observed in culture. Many species in ser. Viridicata are associated with stored cereal grains. Series Viridicata is a typical example of a polythetic series regarding extrolites as no metabolite is shared among all species (Frisvad and Samson, 2004b, Hallas-Moller et al., 2018). Most species produce a dark brown halo on Raulin-Thom agar (Frisvad & Samson 2004b).



Included species: Penicillium aurantiogriseum, P. cyclopium, P. freii, P. melanoconidium, P. neoechinulatum, P. polonicum, P. tricolor, P. viridicatum.



Extrolites: Several species in ser. Viridicata produce 4-hydroxy-3,6-dimethyl-2H-pyran-2-one, penicillic acids, cyclopenins / viridicatins, verrucofortines / fructigenines / puberulines, verrucosidins, xanthomegnins, while rather few species produce anacine, aspermytin A / peaurantiogriseols, aspterric acid, asteltoxin, auranthine, aurantiomides, aurantiamine / viridamine, brevianamide A, 4-(9-hydroxy-10-butynoxy)benzoic acid / penipratynolenes, chrysogine, moniliformin, penicyrones, penitrem A, pseurotins, puberulonic acid, terrestric acids, viridic acid (Frisvad and Samson, 2004b, Ma et al., 2015, Bu et al., 2016, Hallas-Moller et al., 2018, Hamed et al., 2019).



Notes: Species in sect. Fasciculata are able to grow well at 15 °C and low water activities, and they produce rough-walled conidiophores. This section is phylogenetically most closely related to sect. Penicillium (Fig. 19, Steenwyk et al. 2019). Section Fasciculata is divided in five series: Camembertiorum, Corymbifera, Verrucosa, Viridicata and Gladioli. The former four series are phylogenetically very closely related, and ser. Gladioli takes a sister position to those (Fig. 19). We tentatively place P. venetum in ser. Corymbifera. This species shares features with taxa in ser. Corymbifera but is phylogenetically closer to series Viridicata and Verrucosa. Penicillium cavernicola, P. discolor, P. echinulatum and P. solitum share the production of rough-walled conidia and these species were therefore previously placed in ser. Solita (Frisvad & Samson 2004b). Phylogenetic analysis shows that P. cavernicola is closely related to P. camemberti and related species. The other three species are phylogenetically related and these form a clade close to ser. Camembertiorum. The species previously classified in ser. Solita grow well on CREA, can use creatine as the sole N-source, and typically occur on proteinaceous and lipid-containing foods and these features are shared with ser. Camembertiorum. We therefore decided to expand ser. Camembertiorum with P. cavernicola, P. discolor, P. echinulatum and P. solitum, and treat ser. Solita (Frisvad & Samson 2004b) as a synonym of ser. Camembertiorum.



Section Formosana Houbraken & Frisvad, sect. nov. MycoBank MB833976.



Etymology: Named after the type species of the section, Penicillium formosanum.



Type: Penicillium formosanum H.M. Hsieh et al., Trans. Mycol. Soc. Rep. China 2: 159. 1987.



Diagnosis: Phylogeny: Section Formosana belongs to subgen. Penicillium and is phylogenetically basal to sections Chrysogena, Fasciculata, Osmophila, Penicillium, Samsoniorum and Roquefortorum. Morphology & physiology: Colonies growing moderately fast, colony texture fasciculate, producing yellow synnemata; conidiophores terverticillate, smooth-walled; conidia subglobose to broadly ellipsoidal, smooth-walled; no growth at 37 °C, weak growth and no acid production on CREA. Sexual morph unknown; sclerotia not observed in culture. Series description on Hsieh et al. (1987) and Frisvad & Samson (2004b).



Included species: Penicillium formosanum.



Extrolites: Asteltoxin and patulin (Frisvad & Samson 2004b).



Notes: Penicillium formosanum is the only species in sect. and ser. Formosana. This species forms a unique lineage in subgen. Penicillium and therefore treated as a separate section and series (Fig. 2, Fig. 19). Penicillium formosanum produces yellow synnemata on MEA and oatmeal agar (Frisvad & Samson 2004b), a feature not observed in species of related sections. A series classification is lacking in sect. and ser. Formosana is therefore only informally introduced in this article.



Section Osmophila Houbraken & Frisvad, Persoonia 36: 309. 2016. MycoBank MB815869.



Type: Penicillium osmophilum Stolk & Veenb.-Rijks, Antonie van Leeuwenhoek 40: 1. 1974.



Description: See Houbraken et al. (2016) (morphology, phylogeny).



Series Osmophila Houbraken & Frisvad, ser. nov. MycoBank MB833977.



Etymology: Named after the type species of the series, Penicillium osmophilum.



Type: Penicillium osmophilum Stolk & Veenb.-Rijks, Antonie van Leeuwenhoek 40: 1. 1974.



Diagnosis: Phylogeny: Series Osmophila belongs to subgen. Penicillium, sect. Osmophila and is phylogenetically most closely related to ser. Samsoniorum. Morphology & physiology: Colonies restricted, colony texture floccose; conidiophores bi- and terverticillate, smooth-walled; conidia broadly ellipsoidal or subglobose, smooth-walled; osmophilic. Sexual morph eupenicillium-type, homothallic, avellaneous; ascospores ellipsoidal, with two equatorial flanges, valves rough-walled. Series description based on Pitt (1980) and Houbraken et al. (2016).



Included species: Penicillium osmophilum.



Extrolites: Andrastin A and meleagrin.



Series Samsoniorum Houbraken & Frisvad, ser. nov. MycoBank MB833978.



Etymology: Named after R.A. Samson, a Dutch mycologist studying Penicillium taxonomy.



Type: Penicillium samsonianum L. Wang et al., Persoonia 36: 313. 2016.



Diagnosis: Phylogeny: Series Samsoniorum belongs to subgen. Penicillium, sect. Osmophila and is most closely related to ser. Osmophila. Morphology & physiology: Colonies restricted, colony texture lanose; conidiophores terverticillate, occasionally bi- or quarterverticillate, smooth-walled; conidia globose, smooth-walled; poor growth and no acid production on CREA, psychrotrophic. Sexual morph unknown; sclerotia not produced in culture. Series description based on Houbraken et al. (2016).



Included species: Penicillium samsonianum.



Extrolites: Berkbenzofuranthioester, dimethylphthalides (3,5-dimethyl-6-hydroxyphthalide and 3,5-dimethyl-6-methoxyphthalide), haenamindole, mycophenolic acid, patulin, penitrems, penitremones, phomfuranone, phomopsolides, phompyrone, roquefortine C, sclerotigenin (Stierle et al., 1997, Stierle et al., 2014 (strain misidentified as P. clavigerum); (Houbraken et al. 2016).



Notes: Section Osmophila was introduced to accommodate P. osmophilum and P. samsonianum. These two species are phylogenetically related in our three-gene phylogenetic analysis (Fig. 19), confirming the results of Houbraken et al. (2016). In contrast, these species did not form a well-supported clade in our nine-gene phylogenetic analysis (Fig. 2). The two species included in sect. Osmophila are phenotypically different and share a few characters (Houbraken et al. 2016). We therefore decided to treat these as separate series (Osmophila and Samsoniorum).



Section Paradoxa Houbraken & Samson, Stud. Mycol. 70: 43. 2011. MycoBank MB563134.



Type: Aspergillus paradoxus Fennell & Raper, Mycologia 47: 69. 1955.



Description: See Houbraken & Samson (2011) (morphology, phylogeny), Kocsubé et al. (2016) (phylogeny).



Series Atramentosa Houbraken & Frisvad, ser. nov. MycoBank MB833979.



Etymology: Named after the type species of the series, Penicillium atramentosum.



Type: Penicillium atramentosum Thom, U.S.D.A. Bur. Animal Industr. Bull. 118: 65. 1910.



Diagnosis: Phylogeny: Series Atramentosa belongs to subgen. Penicillium sect. Paradoxa and is phylogenetically most closely related to ser. Paradoxa. Morphology & physiology: Colonies growing moderately fast, brown reverse colour on CYA and YES, colony texture velvety; conidiophores predominantly terverticillate, sometimes bi- or quarterverticillate, thin, smooth-walled, good growth on CREA, but no acid production; conidia globose to subglobose or (broadly) ellipsoidal, smooth-walled. Sexual morph unknown. Series description based on Frisvad and Samson, 2004b, Visagie et al., 2014b and Guevara-Suarez et al. (2020).



Included species: Penicillium atramentosum, P. balearicum, P. fimosum, P. ibericum, P. magnielliptisporum, P. mexicanum.



Extrolites: Andrastin A, haenamindole, meleagrin, oxaline, roquefortine C and D, rugulovasine A and B.



Series Paradoxa Houbraken & Frisvad, ser. nov. MycoBank MB833980.



Etymology: Named after the type species of the series, Aspergillus paradoxus.



Type: Aspergillus paradoxus Fennell & Raper, Mycologia 47: 69. 1955 (current name: Penicillium paradoxum).



Diagnosis: Phylogeny: Series Paradoxa belongs to subgen. Penicillium, sect. Paradoxa is phylogenetically most closely related to ser. Atramentosa. Morphology & physiology: Colonies growing moderately fast or fast; conidiophores aspergillus-like, smooth-walled or roughened; biseriate, occasionally “triseriate”; conidia globose, subglobose or (broadly) ellipsoidal, smooth-walled or roughened. Sexual morph not observed in culture (P. crystallinum, P. malodoratum) or present (P. paradoxum), eupenicillium-type, homothallic, buff to vinaceous buff; ascospores ellipsoidal, with two equatorial flanges, valves smooth-walled; sclerotia present (P. malodoratus), buff to brown coloured. Series Paradoxa species produce an unpleasant smell and the species are mostly found on dung. Series description based on Raper and Fennell, 1965, Sarbhoy and Elphick, 1968 and Samson et al. (2014).



Included species: Penicillium crystallinum, P. malodoratum, P. paradoxum.



Extrolites: Andrastin A, chrysogine, chrysophanic acid, pachybasin, meleagrin, oxaline, sorbicillins (Frisvad and Samson, 2004b, Visagie et al., 2014b). Andrastin A, chrysogine, meleagrin and sorbicillins are shared with several species in ser. Chrysogena.



Notes: Section Paradoxa is phylogenetically related to sect. Turbata (Fig. 2, Fig. 19). Both sections include species that are associated with dung (Tuthill and Frisvad, 2002, Samson et al., 2014, Guevara-Suarez et al., 2020). Two series are recognised in sect. Paradoxa: Atramentosa and Paradoxa. These series are phylogenetically and phenotypically distinct. Series Paradoxa includes species that have an aspergillus-type asexual morph, while ser. Atramentosa species produce typical penicillium-type conidiophores. Penicillium atramentosum was previously classified in ser. Camembertiorum, due to its ability to grow on creatine as sole N-source. This species differs from other series in this section by the production of smooth-walled conidiophores and its alkali-tolerant nature (Frisvad & Samson 2004b).



Section Penicillium [autonym]. MycoBank MB549140.

Synonyms: Penicillium sect. Bulliardium Dierckx, Ann. Soc. Sci. Bruxelles 25: 85. 1901.

Penicillium sect. Cylindrosporum Pitt, The Genus Penicillium: 381. 1980 [1979].

Penicillium sect. Digitata Frisvad & Samson, Stud. Mycol. 49: 26. 2004.



Type: Penicillium expansum Link, Mag. Ges. Naturf. Freunde Berlin 3: 16. 1809.



Description: See Houbraken & Samson (2011) (phylogeny), Houbraken et al. (2016) (morphology, phylogeny).



Series Clavigera Houbraken & Frisvad, ser. nov. MycoBank MB833981.



Etymology: Named after the type species of the series, Penicillium clavigerum.



Type: Penicillium clavigerum Demelius, Verh. Zool.-Bot. Ges. Wien 72: 74. 1923.



Diagnosis: Phylogeny: Series Clavigera belongs to subgen. Penicillium, sect. Penicillium and is phylogenetically most closely related to ser. Sclerotigena. Morphology & physiology: Colonies restricted, colony texture coremiform, indeterminate synnemata (synnemata covered over nearly the entire length with conidiophores); conidiophores terverticillate, appressed, smooth-walled; conidia ellipsoidal, smooth-walled; poor growth on CYA supplemented with 5 % NaCl (CYAS). Sexual morph unknown; sclerotia not observed in culture. Series description based on Frisvad & Samson (2004b) and Crous et al. (2014).



Included species: Penicillium clavigerum, P. coccotrypicola.



Extrolites: Ascladiol, asperfuran, bioxanthracenes, cyclopiazonic acids, glandicolin A, isofumigaclavine A, norlichexanthone, patulin, penitrems, Raistrick phenols, roquefortin C, sclerotigenin, TAN-1612, viomellein, vioxanthin, xanthomegnin are all produced by P. clavigerum (Frisvad & Samson 2004b).



Series Digitata Raper & Thom ex Stolk & Samson, Adv. Pen. Asp. Syst.: 183. 1986 [1985]. MycoBank MB832731.



Type: Penicillium digitatum (Pers.) Sacc., Fungi. Italica Autogr. Delin: tab. 894. 1881.



Diagnosis: Phylogeny: Series Digitata belongs to subgen. Penicillium, sect. Penicillium and is phylogenetically related to series Italica and Penicillium; but the relationship between these three series remains unresolved. Morphology & physiology: Colonies spreading, colony texture velvety, conidial colour en masse olive; conidiophores terverticillate, appressed, smooth-walled; conidia ellipsoidal or cylindrical, large, measuring 6–9(–14) μm in length, smooth-walled; no growth on Czapek agar, poor growth and no acid production on CREA. Sexual morph unknown; sclerotia not observed in culture. Causing rot in citrus (Frisvad & Samson 2004b).



Included species: Penicillium digitatum.



Extrolites: Penicillium digitatum produces tryptoquialanines and tryptoquialanones (Ariza et al., 2002, Frisvad and Samson, 2004b).



Series Italica Fassatiová, Acta Univ. Carol., Biol. 12: 324. 1977. MycoBank MB832970.

Synonym: Penicillium series Italica Raper & Thom ex Pitt, The Genus Penicillium: 381. 1980 [1979].



Type: Penicillium italicum Wehmer, Hedwigia 33: 211. 1894.



Diagnosis: Phylogeny: Series Italica belongs to subgen. Penicillium, sect. Penicillium and is phylogenetically related to series Digitata and Penicillium; but the relationship between these three series remains unresolved. Morphology & physiology: Colonies restricted (P. ulaiense) or fast (P. italicum), colony texture velvety or fasciculate; conidiophores terverticillate, appressed, smooth-walled; conidia ellipsoidal or cylindrical, smooth-walled. Sexual morph not observed in culture; sclerotia can be present, colourless to light brown. Causing rot in citrus fruits. The extrolite deoxybrevianamide E is produced by both species in series Italica (Frisvad & Samson 2004b).



Included species: Penicillium italicum, P. ulaiense.



Extrolites: Italicic acids, PI-3, PI-4, dehydrofulvic acid, deoxybrevianamides, formylxanthocillin X, verrucolones including 5,6-dihydro-4-methoxy-2H-pyran-2-one, 4-methoxy-6-n-propenyl-2-pyrone, 5-hydroxymethylfuroic acid. Of these, P. ulaiense has only been shown to produce the deoxybrevanamides (deoxybrevianamide E, 12,13-dehydrodeoxybrevianamide E, 10,20-dehydro[12,13-dehydropropyl]-2-(1’,1’-dimethylallyltryptophanyl)diketo-piperazine) (Arai et al., 1989b, Frisvad and Samson, 2004b).



Series Penicillium [autonym]. MycoBank MB549141.

Synonym: Penicillium ser. Expansa Raper & Thom ex Fassatiová, Acta Univ. Carol., Biol. 12: 324. 1977.



Type: Penicillium expansum Link, Mag. Ges. Naturf. Freunde Berlin 3: 16. 1809.



Diagnosis: Phylogeny: Series Penicillium belongs to subgen. Penicillium, sect. Penicillium and is phylogenetically related to series Digitata and Italica; but the relationship between these three series remains unresolved. Morphology & physiology: Colonies growing restrictedly or moderately rapid, colony texture velvety to fasciculate; conidiophores terverticillate, divergent, smooth; conidia globose, subglobose or ellipsoidal, smooth; good growth and acid production (and base production) on CREA. Sexual morph unknown; sclerotia not observed in culture.



Included species: Penicillium expansum, P. marinum.



Extrolites: Andrastins, aurantioclavine∗, chaetoglobosins, citrinin, communesins∗, expansolides, 3,5-dimethyl-6-hydroxyphthalide and 3,5-dimethyl-6-methoxyphthalide, patulin∗, penochalasins, penostatins, pyrrolocins∗, and roquefortine C∗ are produced by ser. Penicillium species (Andersen et al., 2004, Frisvad and Samson, 2004b). The extrolites marked with an asterisk are produced by both species in ser. Penicillium.



Series Sclerotigena Houbraken & Frisvad, ser. nov. MycoBank MB833982.



Etymology: Named after the type species of the series, Penicillium sclerotigenum.



Type: Penicillium sclerotigenum W. Yamam., Sci. Rep. Hyogo Univ. Agric. 1: 69. 1955.



Diagnosis: Phylogeny: Series Sclerotigena belongs to subgen. Penicillium, sect. Penicillium and is phylogenetically most closely related to ser. Clavigera. Morphology & physiology: Colonies spreading, colony texture velvety; conidiophores bi- and terverticillate, appressed, smooth; conidia ellipsoidal, smooth. Sexual morph not observed in culture; sclerotia orange-brown. Causing rot in yam tubers (Frisvad & Samson 2004b).



Included species: Penicillium sclerotigenum.



Extrolites: The extrolites griseofulvins, gregatins, patulin and sclerotigenin are produced by the sole species in ser. Sclerotigena.



Notes: The species belonging to sect. Penicillium have a strongly fasciculate or synnematous colony texture, produce smooth-walled conidiophore stipes and conidia. Section Penicillium is poorly supported (ML < 70 % BS; BI < 0.95 pp) (Fig. 19), and a similar result was obtained by Houbraken et al. (2016). Five series are accepted in the phenotypically diverse sect. Penicillium: Clavigera, Digitata, Italica, Penicillium and Sclerotigena. Series Digitata, Italica and Penicillium are phylogenetically related and form a well-supported clade (Fig. 19); series Clavigerum and Sclerotigena form a related sister clade, but statistical support is lacking. Series Clavigerum and Sclerotigena are related but differ phenotypically. Series Sclerotigena species grow rapidly and form sclerotia, while ser. Clavigera species grow restrictedly and do not produce sclerotia. Series Italica and Digitata include species causing rot of citrus fruits, but these series are phenotypically distinct: ser. Italica species produce blue-green conidia, while olive-green conidia are produced in ser. Digitata.



Section Ramosum Stolk & Samson, Adv. Pen. Asp. Syst.: 179. 1986 [1985]. MycoBank MB832722.



Type: Penicillium lanosum Westling, Ark. Bot. 11: 97. 1911.



Description: See Stolk & Samson (1985) (morphology), Houbraken & Samson (2011) (phylogeny); a modern taxonomic overview of the section using phenotypic data is lacking.



Series Lanosa Stolk & Samson, Adv. Pen. Asp. Syst.: 180. 1986 [1985]. MycoBank MB834032.



Type: Penicillium lanosum Westling, Ark. Bot. 11: 97. 1911.



Diagnosis: Phylogeny: Series Lanosa belongs to subgen. Penicillium, sect. Ramosum and is phylogenetically most closely related to series Raistrickiorum and Scabrosa. Morphology & physiology: Colonies restricted, colony texture floccose; conidiophores biverticillate, twice biverticillate, or terverticillate, stipes smooth-walled or finely roughened; conidia globose to subglobose, smooth-, or rough-walled. Sexual morph unknown; sclerotia not observed in culture. All species are psychrotolerant.



Included species: Penicillium beceitense, P. jamesonlandense, P. kojigenum, P. lanosum, P. ribium, P. swiecickii.



Extrolites: Asperfuran, chrysogine, compactins, cycloaspeptides, griseofulvins,2-(4-hydroxyphenyl)-2-oxoacetaldehyde oxime, kojic acid, penicillic acid, pseurotins, psychrophilins, pyripyropenes, sclerotigenin, tryptoquivalines and viridicatumtoxin are produced by ser. Lanosa species (Frisvad et al. 2006).



Series Raistrickiorum Houbraken & Frisvad, ser. nov. MycoBank MB833983.



Etymology: Named after the type species of the series, Penicillium raistrickii.



Type: Penicillium raistrickii G. Sm., Trans. Brit. Mycol. Soc. 18: 90. 1933.



Diagnosis: Phylogeny: Series Raistrickiorum belongs to subgen. Penicillium, sect. Ramosum and is phylogenetically most closely related to ser. Scabrosa. Morphology & physiology: Colonies spreading, colony texture velutinous; conidiophores bi- or terverticillate, stipes rough-walled, inflated at the apex; conidia globose, smooth-walled; no growth at 37 °C, poor growth on CREA. Sexual morph not observed in culture; sclerotia produced in P. raistrickii and P. simile.



Included species: Penicillium raistrickii, P. sajarovii, P. simile.



Extrolites: Atrovenetins, chrysogine, griseofulvins, penicillic acid, peniciketals, peniciraistrins, pestafolide and desmethylcandidusins are produced by ser. Raistrickiorum species (Belofsky et al., 1998, Ma et al., 2012, Ma et al., 2017b, Liu et al., 2014a, Liu et al., 2018a, Li et al., 2019b).



Series Scabrosa Houbraken & Frisvad, ser. nov. MycoBank MB833984.



Etymology: Named after the type species of the series, Penicillium scabrosum.



Type: Penicillium scabrosum Frisvad, Samson & Stolk, Persoonia 14: 177. 1990.



Diagnosis: Phylogeny: Series Scabrosa belongs to subgen. Penicillium, sect. Ramosum and is phylogenetically most closely related to ser. Raistrickiorum. Morphology & physiology: Colonies growing restrictedly to moderately rapid, colony texture velvety; conidiophores biverticillate, sometimes with additional branch, rough-walled; conidia globose to subglobose, rough-walled. Sexual morph unknown; sclerotia not observed in culture.



Included species: Penicillium scabrosum.



Extrolites: The only species in ser. Scabrosa produces aurantiamine, asperpentyns, cyclopenins & viridicatins, gliovictins, penigequinolones, peniprequinolones, fumagillin, penicillic acid, pseurotins (Frisvad and Filtenborg, 1990, Larsen et al., 1998).



Series Soppiorum Houbraken & Frisvad, ser. nov. MycoBank MB833985.



Etymology: Named after the type species of the series, Penicillium soppii.



Type: Penicillium soppii K.W. Zaleski, Bull. Int. Acad. Polon. Sci., Cl. Sci. Math., Sér. B., Sci. Nat. 1927: 476. 1927.



Diagnosis: Phylogeny: Series Soppiorum belongs to subgen. Penicillium, sect. Ramosum and is phylogenetically sister to a clade with series Lanosa, Raistrickiorum and Scabrosa, though statistical support for this relationship is lacking; we place P. lusitanum in ser. Soppiorum, though statistical support for this position is poor (<70 % BS, 0.95 pp). Morphology & physiology: Colonies growing slow to moderately fast, colony texture velutinous, conidiophores on aerial hyphae or from substrate; conidiophores 100–500 μm in length; smooth-walled or finely roughened, conidia globose, subglobose or broadly ellipsoidal, smooth-walled to finely roughened; no growth at 30 °C and 37 °C, poor growth and no acid production on CREA. Sexual morph not observed in culture; sclerotia produced by P. soppii.



Included species: Penicillium americanum, P. chroogomphum, P. lenticrescens, P. lusitanum, P. soppii, P. tunisiense.



Extrolites: Asperentins, asperphenamate, benzomalvins, cycloaspeptides, fumagillin, griseofulvin, pseurotins and terrein are extrolites found in P. soppii (Frisvad et al. 2006).



Series Virgata Houbraken & Frisvad, ser. nov. MycoBank MB833986.



Etymology: Named after the type species of the series, Penicillium virgatum.



Type: Penicillium virgatum Nirenberg & Kwasna, Mycol. Res. 109: 977. 2005.



Diagnosis: Phylogeny: Series Virgata belongs to subgen. Penicillium, sect. Ramosum and is phylogenetically basal to the other series of sect. Ramosum. Morphology & physiology: Colonies restricted, colony texture velvety to floccose; conidiophores biverticillate, with additional branches, smooth; conidia globose to subglobose, rough-walled; poor growth and no acid production on CREA. Sexual morph unknown; sclerotia not observed in culture. Series description based on Kwaśna & Nirenberg (2005).



Included species: Penicillium virgatum.



Extrolites: Andrastin A.



Notes: Five well-supported lineages, representing series Lanosa, Raistrickiorum, Scabrosa, Soppiorum and Virgata are recognised in sect. Ramosum. This series classification is mainly based on the phylogenetic analysis presented in Fig. 19. Series Scabrosa is phylogenetically most close to ser. Raistrickiorum and these two series from a clade together with ser. Lanosa. Series Soppiorum is sister to these series, but the statistical support for this absent (ML = <75 % BS) or low (BI < 0.95 pp). Series Virgata takes a basal position in sect. Ramosum. A taxonomic study of sect. Ramosum is not yet performed and might give more data supporting the proposed series classification.



Section Robsamsonia Houbraken & Frisvad, Persoonia 36: 309. 2016. MycoBank MB815870.



Type: Penicillium robsamsonii Frisvad & Houbraken, Persoonia 36: 313. 2016.



Description: See Houbraken et al. (2016) (morphology, phylogeny).



Series Claviformia Raper & Thom ex Stolk et al., Mod. Conc. Pen. Asp. Clas.: 132. 1990. MycoBank MB834533.



Type: Penicillium vulpinum (Cooke & Massee) Seifert & Samson, Adv. Pen. Asp. Syst.: 144. 1986 [1985].



Diagnosis: Phylogeny: Series Claviformia is here tentatively classified in subgen. Penicilium, sect. Robsamsonia; statistical support for this classification is lacking (Fig. 19, see also Notes on series in section below). Morphology & physiology: Colonies restricted, sometimes growing moderately fast, having capitulate synnemata in concentric zones; conidiophores terverticillate, sinoid, smooth-walled; conidia ellipsoidal, smooth. Sexual morph unknown; sclerotia not observed in culture.



Included species: Penicillium vulpinum.



Extrolites: Andrastins, (-)-3-butyl-7-hydroxyphthalides, cyclopenins, cyclopiamine, meleagrin, 2-methoxymethyl-3-pentylphenol, 2-methyl-hydroquinone, oxalicin, oxalin, pachybasin, patulin, pintulin, roquefortine C & D viridicatins (Frisvad & Samson 2004b).



Series Glandicolarum Houbraken & Frisvad, ser. nov. MycoBank MB834600.



Etymology: Named after the type species of the series, Coremium glandicola.



Type: Coremium glandicola Oudem. Ned. Kruidk. Arch. 2: 918. 1903 (current name: Penicillium glandicola).



Diagnosis: Phylogeny: Series Glandicolarum belongs to subgen. Penicillium, sect. Robsamsonia and is phylogenetically basal to the other series of sect. Robsamsonia (except ser. Claviformia). Morphology & physiology: Colonies restricted; colony texture strongly fasciculate; conidiophores terverticillate, tuberculate stipes; conidia ellipsoidal, smooth-walled; good growth and acid production on CREA. Sexual morph unknown; sclerotia not observed in culture. Penicillium glandicola has been found on acorns and in dung.



Included species: Penicillium geumsanense, P. glandicola, P. granulatum, P. synnematicola.



Extrolites: Andrastin A, asperfuran, meleagrins, roquefortine C & D, patulidin, penitrems are produced by P. glandicola (Frisvad et al. 2004).



Series Robsamsonia Houbraken & Frisvad, ser. nov. MycoBank MB833987.



Etymology: Named after the type species of the series, Penicillium robsamsonii.



Type: Penicillium robsamsonii Frisvad & Houbraken, Persoonia 36: 313. 2016.



Diagnosis: Phylogeny: Series Robsamsonia belongs to subgen. Penicillium, sect. Robsamsonia and is phylogenetically related to ser. Urticicola, though statistical support is poor (Fig. 19). Morphology & physiology: Colonies restricted, sometimes growing moderately fast, colony texture fasciculate or synnematous; conidiophores terverticillate, smooth- or rough-walled; conidia ellipsoidal, smooth-walled; good growth on CREA and no or some acid production. Sexual morph unknown.



Included species: Penicillium brevistipitatum, P. compactum, P. concentricum, P. coprobium, P. coprophilum, P. fimorum, P. robsamsonii.



Extrolites: Alternariol, andrastins, chaetoglobosins, citreoisocoumarins, clavatols, cyclopiamins, griseofulvins, meleagrin, palitantin, patulin, patulodin, pyripyropenes, quinolactacins, roquefortine C, xanthoepocin. Pyripyropens are shared by ser. Robsamsonia species (Houbraken et al. 2016).



Series Urticicola Fassatiová, Acta Univ. Carol., Biol. 12: 324. 1977. MycoBank MB834534.



Type: Penicillium urticae Bainier, Bull. Soc. Mycol. France 23: 15. 1907. (current name: Penicillium griseofulvum).



Diagnosis: Phylogeny: Series Urticicola belongs to subgen. Penicillium, sect. Robsamsonia and is phylogenetically related to ser. Robsamsonia, though statistical support is poor (Fig. 19). Morphology & physiology: Colonies growing restrictedly or moderately rapid, colony texture weakly fasciculate; conidiophores bi-, ter-, or quarterverticillate, divergent, smooth-walled; phialides very short, less than 6 μm in length; conidia broadly ellipsoidal, smooth-walled. Poor growth and no acid production on CREA. Sexual morph unknown.



Included species: Penicillium dipodomyicola, P. griseofulvum.



Extrolites: Cyclopiamins, cyclopiazonic acids∗, fulvic acids, griseofulvins∗, mycelianamide, patulin∗, patulodin, roquefortine C. The extrolites with an asterisk are produced by both species in ser. Urticicola.



Notes: Species in sect. Robsamsonia grow moderately fast on CYA incubated at 25 °C and no or slow growth is present on CYA incubated at 30 °C. Section Robsamsonia includes four series: Claviformia, Glandicolarum, Robsamsonia and Urticicola. Series Robsamsonia and Urticicola are phylogenetically related, but the relationship of the series Claviformia (Penicillium vulpinum) and Glandicolarum (P. glandicola) is unresolved. In Houbraken et al. (2016), series Claviformia and Glandicolarum taxa took a basal position in sect. Robsamsonia, but lacked support in the ML and Bayesian analysis. A similar result was obtained in this study (Fig. 19). Penicillium vulpinum was included in the genome scale phylogenetic study of Steenwyk et al. (2019) and based on this analysis, this species is sister to P. coprophilum (ser. Robsamsonia) and P. griseofulvum (ser. Urticicola). Series Claviformia, Glandicolarum and Robsamsonia include species that occur on dung, and ser. Urticicola occurs mainly on dry cereals and seeds. Furthermore, series Claviformia and Glandicolarum include species that produce synnematous structures and these structures are also produced by certain members of ser. Robsamsonia (e.g., P. coprophilum). Based on similarity in habitat (dung) and morphology and the phylogenetic relationship using genome data, we decided to accommodate series Claviformia and Glandicolarum in sect. Robsamsonia. Stolk et al. (1990) listed Penicillium ser. Granulata Raper & Thom ex Fassatiová in their classification of the terverticillate Penicillia (type P. granulatum). We could not find Fassatiová’s original description of this series and regard this series as doubtful. Series Glandicolarum is introduced for P. glandicola and related species.



Section Roquefortorum [as “Roqueforti”] Frisvad & Samson, Stud. Mycol. 49: 16. 2004. MycoBank MB701527.



Description: See series Roquefortorum below.



Series Roquefortorum [as “Roqueforti”] Raper & Thom ex Frisvad, Int. Mod. Meth. Pen. Asp. Clas.: 277. 2000. MycoBank MB701528.



Type: Penicillium roqueforti Thom, U.S.D.A. Bur. Animal Industr. Bull. 82: 35. 1906.



Diagnosis: Phylogeny: Section Roquefortorum belongs to subgen. Penicillium; the phylogenetic relationship with other sections is unresolved (Fig. 2, Fig. 19). Morphology & physiology: Colonies spreading; texture velutinous; conidiophores terverticillate, with tuberculate stipes; conidia globose to subglobose, smooth-walled; growth at low pH (e.g., media containing 0.5 % acetic acid), high alcohol concentrations and elevated CO₂ levels. Good growth but no acid production on CREA. Sexual morph not observed in culture (P. carneum, P. paneum) or present, eupenicillium-type, homothallic (P. psychrosexuale) or heterothallic (P. roqueforti, Ropars et al. (2014)), pale orange-brown. Section description based on Frisvad & Samson (2004b) (extrolites, phylogeny), Houbraken et al. (2010a) (morphology, extrolites), Houbraken & Samson (2011) (phylogeny).



Included species: Penicillium carneum, P. mediterraneum, P. paneum, P. psychrosexuale, P. roqueforti.



Extrolites: Andrastins, botryodiplidin, chryzothiazoles, citreoisocoumarins, isofumigaclavines, marcfortines, mycophenolic acid, patulin, penipacids, penipanoids, PR-toxins, roquefortine C & D∗ (Frisvad and Samson, 2004b, Nielsen et al., 2005, Li et al., 2011, Li et al., 2013, Li et al., 2014, An et al., 2013 (P. paneum misidentified as P. chrysogenum), Xu et al. 2014; P. paneum misidentified as P. oxalicum).



Notes: Even though ser. Roquefortorum is the sole series in the section, we accept this series because it was formally introduced (Frisvad et al. 2000). The phylogenetic relationship of this section with other sections is unresolved (Fig. 2, Fig. 19). The extrolites marcfortines, botryodiploidin and isofumigaclavine are produced by members of this series, and are not detected in the related sections Chrysogena, Fasciculata, Penicillium and Robsamsonia (Houbraken et al. 2016).



Section Turbata Houbraken & Samson, Stud. Mycol. 70: 43. 2011. MycoBank MB563133.



Type: Penicillium turbatum Westling, Ark. Bot. 11: 128. 1911.



Diagnosis: Phylogeny: Section Turbata belongs to subgen. Penicillium and is phylogenetically related to sect. Paradoxa. Morphology & physiology: Colonies growing moderately fast, texture velutinous; conidiophores monoverticillate, (symmetrically biverticillate), or terverticillate; conidia globose, subglobose or ellipsoidal, smooth or rough-walled in older cultures. Sexual morph not observed in culture (P. madriti) or present (P. bovifimosum, P. turbatum (syn. Eup. baarnense)), eupenicillium-type, homothallic, in brown shades. Series description based on Houbraken & Samson (2011) (phylogeny).



Included species: Penicillium bovifimosum, P. caprifimosum, P. madriti, P. turbatum.



Extrolites: Species belonging to this section can produce penicillic acid and a fumagillin-like compound (Tuthill & Frisvad 2002).



Notes: No series classification is proposed for sect. Turbata, and ser. Turbata is therefore only informally introduced in this article.

New combinations and names for species in Eurotiales

Four new genera are introduced above (Acidotalaromyces, Ascospirella, Evansstolkia and Pseudohamigera). New combinations are made below for the species belonging to these new genera. Furthermore, new combinations (or names) are introduced for the following incorrectly classified species: Byssochlamys verrucosa, Chaetosartorya stromatoides, Merimbla humicoloides, Penicillium arenicola, P. kabunicum, P. mirabile, P. moldavicum, P. resinae, Phialomyces striatus, Talaromyces brevicompactus and Thermoascus crustaceus var. verrucosus. No Paecilomyces name was available for Byssochlamys lagunculariae and this is introduced below.



Acidotalaromyces lignorum (Stolk) Houbraken, Frisvad & Samson, comb. nov. MycoBank MB832555.

Basionym: Penicillium lignorum Stolk, Antonie van Leeuwenhoek 35: 264. 1969.



Notes: Stolk (1969) classified this species in sect. Biverticillata-Symmetrica (subgen. Biverticillium). Many of the species previously belonging to subgen. Biverticillium were transferred to Talaromyces (Samson et al. 2011c). Phylogenetically, this species clearly does not belong to Talaromyces sensu stricto and is the sole species in the newly introduced genus Acidotalaromyces.



Ascospirella lutea (Zukal) Houbraken, Frisvad & Samson, comb. nov. MycoBank MB832556.

Basionym: Penicillium luteum Zukal, Sitzungsber. Kaiserl. Akad. Wiss. Wien, Math.-Naturwiss. Cl., Abt. 1, 98: 561. 1890.

Synonyms: Gymnoascus luteus (Zukal) Sacc., Syll. Fung. 11: 437. 1895.

Talaromyces luteus (Zukal) C.R. Benj., Mycologia 47: 681. 1955.

Talaromyces luteus (Zukal) Stolk & Samson, Stud. Mycol. 2: 23. 1972.



Notes: Pitt (1980) treated Tal. udagawae as a synonym of Ascospirella luteum (= Tal. luteus). These species produce similarly ornamented ascospores, but are phylogenetically distinct (Yilmaz et al. 2014). Ascospirella lutea produces luteusins A-E (Fujimoto et al., 1990, Yoshida et al., 1996a, Yoshida et al., 1996b), which are different from the azaphilones produced by Penicillium, Aspergillus and Talaromyces (Gao et al. 2013).



Aspergillus chaetosartoryae Hubka, Kocsubé & Houbraken, nom. nov. MycoBank MB832557.

Replaced synonym: Chaetosartorya stromatoides B.J. Wiley & E.G. Simmons, Mycologia 65: 935. 1973, non Aspergillus stromatoides Raper & Fennell.



Etymology: Referring to Chaetosartorya, the teleomorph genus in which this species was originally described.



Notes: Chaetosartorya stromatoides was described as the sexual morph of Aspergillus stromatoides. This species was typified with QM 8944 (= CBS 265.73 = ATCC 24480 = IMI 171880 = NRRL 5501). Molecular analysis showed that Aspergillus stromatoides (IMI 123750^T = CBS 500.65 = NRRL 4519) and Chaetosartorya stromatoides are related, but distinct species (Peterson, 1995, Peterson, 2008). The name Aspergillus stromatoides is already occupied and therefore the name Aspergillus chaetosartoryae is introduced.



Evansstolkia leycettana (H.C. Evans & Stolk) Houbraken, Frisvad & Samson, comb. nov. MycoBank MB832558.

Basionym: Talaromyces leycettanus H.C. Evans & Stolk, Trans. Brit. Mycol. Soc. 56: 45. 1971.

Synonyms: Penicillium leycettanum H.C. Evans & Stolk, Trans. Brit. Mycol. Soc. 56: 45. 1971.

Paecilomyces leycettanus (H.C. Evans & Stolk) Stolk et al., Stud. Mycol. 2: 51. 1972.



Note: See above, under generic description of Evansstolkia.



Hamigera brevicompacta (H.Z. Kong) Houbraken, Frisvad & Samson, comb. nov. MycoBank MB832579.

Basionym: Talaromyces brevicompactus H.Z. Kong, Mycosystema 18: 9. 1999.



Notes: The taxonomy of Hamigera has been subject of various studies (Stolk and Samson, 1971, Pitt, 1980, Peterson et al., 2010). The genus has been molecularly revised (Peterson et al. 2010); however, Tal. brevicompactus was not included. The original description mentioned that this species is phenotypically similar to Hamigera avellanea (reported as Tal. avellaneus), but differs in ascomatal initials, ascospore ornamentation and conidiophore branching (Kong 1999). Samson et al. (2011c) indicated that Tal. brevicompactus represents a distinct species in Hamigera and our phylogenetic analysis (Supplementary Fig. S2) supports this observation. The new combination H. brevicompacta is proposed here.



Paecilomyces lagunculariae (C. Ram) Houbraken, Frisvad & Samson, comb. nov. MycoBank MB832559.

Basionym: Byssochlamys nivea var. lagunculariae C. Ram, Nova Hedwigia 16: 311. 1968.

Synonym: Byssochlamys lagunculariae (C. Ram) Samson et al., Persoonia 22: 18. 2009.



Notes: Ram (1968) described Byssochlamys nivea var. lagunculariae, without the description of the Paecilomyces asexual morph. In a later study on the taxonomy of Byssochlamys and its Paecilomyces asexual morphs, Samson et al. (2009) elevated this variety to species level, but also did not formally describe the Paecilomyces morph. The type species of Paecilomyces, Paec. variotii, and the type species of Byssochlamys, B. nivea, were shown to be congeneric through molecular sequence analyses (Luangsa-Ard et al., 2004, Samson et al., 2009). Anticipating on the change to one scientific name for fungi (McNeill et al. 2012), Rossman et al. (2016) recommended the use of Paecilomyces over Byssochlamys, and we therefore formally introduce Paec. lagunculariae here.



Penicillago kabunica (Baghd.) Houbraken, Frisvad & Samson, comb. nov. MycoBank MB832560.

Basionym: Penicillium kabunicum Baghd., Novosti Sist. Nizsh. Rast. 5: 98. 1968.



Notes: See under Penicillago. Penicillium kabunicum (and P. moldavicum) were treated by Ramírez (1982) in the “P. brasilianum-series” and Pitt (1980) treated this species as a synonym of P. janthinellum. These observations indicate a relationship with Penicillium sect. Lanata-Divaricata; however, phylogenetic analysis does not support this placement. Penicillago kabunica produces cycloaspeptide A and D.



Penicillago mirabilis (Beliakova & Milko) Houbraken, Frisvad & Samson, comb. nov. MycoBank MB832561.

Basionym: Penicillium mirabile Beliakova & Milko, Mikol. Fitopatol. 6: 145. 1972.



Notes: See also above, under Penicillago (under genera). The identity of this species is controversial and needs further study. Samson et al. (2011c) and Yilmaz et al. (2014) showed in their phylogenetic analyses that P. mirabile belongs to Talaromyces sect. Trachyspermi. However, Samson et al. (2011c) had doubts about these results and they therefore did not combine this species in Talaromyces. Furthermore, they noted that the type strain is in poor condition. Two sequences obtained from IMI 167383 (ex-type of P. mirabile) (KC992257, BenA; KC962096, ITS) are present in GenBank and comparison of those sequences indicate that this strain is a Penicillium corylophilum. In the original description of P. mirabile, it is mentioned that the species produces echinulate conidia with conspicuous connectives. This feature does not fit with P. corylophilum and the sequences derived from IMI 167383 are incorrect. Pitt (1980) placed P. mirabile in Penicillium subgen. Biverticillium (nowadays Talaromyces pro parte). However, he also reported production of smooth-walled conidia and the strain he examined therefore also does not fit with the original description. Re-examination of CBS 624.72, the ex-type of P. mirabile, shows that this strain fits well with the description of Ramírez (1982). Similar to the observation of Ramírez (1982), deep brown to black coloured sclerotia were not observed in CBS 624.72. Phylogenetic analysis using BenA, CaM, ITS and RPB2 sequences shows that this strain is related to P. kabunicum, P. moldavicum and P. nodositatum in the genus Penicillago (Fig. 6).



Penicillago moldavica (Milko & Beliakova) Houbraken, Frisvad & Samson, comb. nov. MycoBank MB832562.

Basionym: Penicillium moldavicum Milko & Beliakova, Novosti Sist. Nizsh. Rast. 4: 255. 1967.



Notes: See above, under Penicillago, and in Guevara-Suarez et al. (2020). Penicillago moldavica produces alternatriol.



Penicillago nodositata (Valla) Guevara-Suarez et al., Fungal Syst. Evol. 5: 64. 2020. MycoBank MB822074.

Basionym: Penicillium nodositatum Valla, Pl. & Soil 114: 146. 1989.



Notes: Based on sequence data, P. nodositatum was tentatively placed in synonymy with P. bilaiae by Houbraken & Samson (2011). This conclusion was drawn based on incorrect accession numbers in the CBS culture collection (Visagie et al. 2013). Visagie et al. (2013) subsequently re-examined CBS 333.90, the ex-type of P. nodositatum, and reported that this strain is related to P. kabunicum, outside Penicillium sensu stricto. Penicillago nodositata is unique in that it forms myconodules in roots of Alnus trees (Valla et al. 1989). We detected an altersolanol, cycloaspeptide A and griseofulvin in this species.



Phialomyces arenicola (Chalab.) Houbraken, Frisvad & Samson, comb. nov. MycoBank MB832563.

Basionym: Penicillium arenicola Chalab., Not. Syst. Crypt. Inst. bot. Acad. Sci. USSR: 162. 1950.

Synonym: Penicillium canadense G. Sm., Trans. Brit. Mycol. Soc. 39: 113. 1956.



Notes: Merimbla humicoloides, P. arenicola and P. canadense are phylogenetically related to Phialomyces macrosporus, the type species of Phialomyces (Fig. 1) (Misra & Talbot 1964). Based on a multigene phylogenetic analysis, M. humicoloides proved not to belong to the Hamigera-clade. To maintain a monophyletic genus, Peterson et al. (2010) decided to combine this species in the genus Penicillium, as P. humicoloides. On the other hand, they also noted a close relationship of this species with P. arenicola and P. canadensis, two species outside Penicillium sensu stricto (Houbraken & Samson 2011). The species belonging to the Phialomyces clade produce penicillium-like conidiophores and gold-brown conidia, a feature not observed in Penicillium sensu stricto. Based on phylogenetic data and morphology, we combine Merimbla (= Penicillium) humicoloides, and P. arenicola in Phialomyces. Pitt (1980) treated P. canadense as a synonym of P. arenicola. A phylogenetic analysis of six loci resolved P. canadense as a close relative of P. arenicola (Peterson et al. 2010) but since no phenotypic differences exist between these species we regard Penicillium canadense as a synonym of Phialomyces arenicola. Phialomyces arenicola has been reported to produce canadensolides (McCorkindale et al. 1968), chlorogentisylalcohol (McCorkindale et al. 1972) and asperphenamate (McCorkindale et al. 1978).



Phialomyces humicoloides (Bills & Heredia) Houbraken, Frisvad & Samson, comb. nov. MycoBank MB832564.

Basionym: Merimbla humicoloides Bills & Heredia, Mycol. Res.105: 1276. 2001.

Synonym: Penicillium humicoloides (Bills & Heredia) S.W. Peterson et al., Mycologia 102: 858. 2010.



Note: See Ph. arenicola.



Pseudohamigera striata (Raper & Fennell) Houbraken, Frisvad & Samson, comb. nov. MycoBank MB832565.

Basionym: Penicillium striatum Raper & Fennell, Mycologia 40: 521. 1948.

Synonyms: Talaromyces striatus (Raper & Fennell) C.R. Benj., Mycologia 47: 682. 1955.

Hamigera striata (Raper & Fennell) Stolk & Samson, Persoonia 6: 347. 1971.

Byssochlamys striata (Raper & Fennell) Arx, Mycotaxon 26: 120. 1986.

Penicillium lineatum Pitt, The Genus Penicillium: 485. 1980.

Paecilomyces lineatus (Pitt) Arx, Mycotaxon 26: 120. 1986.



Notes: The taxonomic position of Pseudohamigera striata was discussed several times in history. The sexual morph of Pseudohamigera striata was previously classified in Talaromyces, Hamigera and Byssochlamys and the asexual morph in Penicillium and Paecilomyces (Raper and Fennell, 1948, Stolk and Samson, 1971, Pitt, 1980, von Arx, 1986), showing the difficulty in classifying this species. Phylogenetic data (Supplementary Fig. S1) show that this species is distinct from Byssochlamys, Hamigera, Paecilomyces, Penicillium and Talaromyces.



Talaromyces resinae (Z.T. Qi & H.Z. Kong) Houbraken & X.C. Wang, comb. nov. MycoBank MB833989.

Basionym: Penicillium resinae Z.T. Qi & H.Z. Kong, Acta Mycol. Sin. 1: 103. 1982.



Notes: Based on an incorrect RPB2 sequence, Houbraken et al. (2014b) considered P. resinae a synonym of Penicillium purpurescens (sect. Aspergilloides). Penicillium purpurescens predominantly produced simple, monoverticillate conidiophores, while P. resinae was originally described to predominantly produce symmetrical biverticillate conidiophores. Qi & Kong (1982) classified P. resinae in the Penicillium funiculosum series, near Penicillium piceum). Both P. funiculosum and P. piceum are currently classified in Talaromyces and it is likely that this species also belongs to this genus. The ex-type strain of P. resinae (CBS 324.83) was re-examined and new sequence data was generated. Comparison of these sequences show a relationship with Tal. brasiliensis and Tal. subericola in sect. Trachyspermi (Supplementary Fig. S3).



Talaromyces striatoconidius Houbraken, Frisvad & Samson, nom. nov. MycoBank MB832566.

Replaced synonym: Phialomyces striatus R.F. Castañeda & W. Gams, Mycotaxon 42: 239. 1991, non Talaromyces striatus (Raper & Fennell) C.R. Benj. 1955.



Etymology: The epithet refers to the typical striate conidia produced by the species.



Notes: This species was originally described as Phialomyces striatus. A connection with Penicillium was made and it was suggested that the species takes an intermediate position between Phialomyces and Penicillium (Castañeda & Gams 1991). Molecular data shows that the species belongs to Talaromyces sect. Talaromyces and it is phylogenetically most closely related to Tal. galapagensis, Tal. indigoticus and Tal. rubicundus (Supplementary Fig. S3). Talaromyces striatoconidius can be easily differentiated from these related species by its production of striate conidia, a rare feature for most other Talaromyces species. The name Talaromyces striatus (= Pseudohamigera striata) is already occupied and therefore the name T. striatoconidius is proposed.



Thermoascus verrucosus (Samson & Tansey) Houbraken, Frisvad & Samson, comb. nov. MycoBank MB832567.

Basionym: Byssochlamys verrucosa Samson & Tansey, Trans. Brit. Mycol. Soc. 65: 512. 1975.

Synonym: Paecilomyces verrucosus Samson & Tansey, Trans. Brit. Mycol. Soc. 65: 512. 1975.



Notes: Samson & Tansey (1975) described B. verrucosa in Byssochlamys based on phenotypic characters. The conidial state of this species resembles Paecilomyces fulvus and both produce asci from croziers. However, based on an ITS phylogeny, Samson et al. (2009) showed that this species is related to Thermoascus. This observation is confirmed in our phylogenetic analysis (Supplementary Fig. S1) and we therefore combine this species in Thermoascus. Samson & Tansey (1975) already noted that B. verrucosa produced simple coiled initials, similar to those of Tal. leycettanus and Tal. purpureus, indicating a position outside Byssochlamys. Thermoascus crustaceus produces, like Th. verrucosus, a paecilomyces-type asexual morph and asci from croziers, and this also indicates a relationship with the genus Thermoascus.



Thermoascus yaguchii Houbraken, Frisvad & Samson, stat. et nom. nov. MycoBank MB833988.

Replaced synonym: Thermoascus crustaceus var. verrucosus Yaguchi, Someya & Udagawa, Mycoscience 36: 151. 1995, non Thermoascus verrucosus (Samson & Tansey) Houbraken, Frisvad & Samson (this study).



Etymology: In honour of Takashi Yaguchi, a prominent taxonomist studying various genera of Eurotiales and one of the authors that described Thermoascus crustaceus var. verrucosus.

Synonym: Coonemeria verrucosa (Yaguchi et al.) Mouch., Cryptog. Mycol. 18: 32. 1997.



Notes: Therrnoascus crustaceus var. verrucosus was introduced as a variety because of its verrucose ascospore ornamentation, in contrast to the echinulate ornamentation in Th. crustaceus var. crustaceus. Sequence data show that this variety represents a distinct species in Thermoascus. Comparison of partial BenA and CaM sequences of CBS 181.67 (a representative strain of Th. crustaceus) and PF-1160^T (= CBS 146343 = IFM 66000), the type of Th. crustaceus var. verrucosus, shows a homology of 95.6 and 96.9 %, respectively. The name Thermoascus verrucosus is already occupied and therefore the name Thermoascus yaguchii is proposed.

Invalid and not accepted species described after 2013

Since the “2014 Aspergillus/Penicillium/Talaromyces lists”, around 300 new names have been described in Penicillium, Aspergillus and Talaromyces. Re-evaluation of these names showed that some of those recently described species were not new to science, incorrectly described or described in the wrong genus. An overview of the not accepted species described after 2013 is given below.



Aspergillus capsici (J.F.H. Beyma) Houbraken et al., Stud. Mycol. 78: 154. 2014.



Notes: The genus Polypaecilum was synonymised with Aspergillus and therefore Polypaecilum capsici was incorrectly combined in Aspergillus. The correct name of Aspergillus capsici (basionym Scopulariopsis capsici J.F.H. Beyma) is Leuconeurospora capsici (J.F.H. Beyma) Malloch et al.



Aspergillus chinensis Samson et al., Stud. Mycol. 78: 155. 2014.



Notes: The name Aspergillus appendiculatus was already occupied and therefore the new name Aspergillus chinensis was introduced for Emericella appendiculata (Samson et al. 2014). Phylogenetic and morphologic examination showed that Aspergillus chinensis is identical with Aspergillus filifer. We follow Matsuzawa et al., 2012, Hubka et al., 2016a and Chen et al. (2016a) in correcting the name A. chinensis to Aspergillus filifer Zalar et al.



Aspergillus cicutus P. Singh & P.J. Cotty, Int. J. Food Microbiol. 289: 148. 2018, nom. inval.



Notes: This species is invalidly described: Arts 38.1 (without a description or diagnosis of the taxon), 40.1 (without designation of type specimen), F.5.1 (no identifier cited). None of the markers commonly used in Aspergilllus taxonomy (BenA, CaM or RPB2) were available for the representative isolates of A. cicutus (NRRL 66829, NRRL 66830, NRRL 66831) and only niaD (nitrate reductase) and aflR (aflatoxin pathway transcription factor) sequences were studied (Singh & Cotty 2019). This species could represent a novel species or it is a synonym of a sect. Flavi member (e.g., Aspergillus austwickii).



Aspergillus delacroxii Samson et al., Stud. Mycol. 78: 155. 2014.



Notes: From Hubka et al. (2016a): The name Aspergillus echinulatus was already occupied and therefore the name Aspergillus delacroxii was introduced for Aspergillus nidulans var. echinulatus Fennell & Raper. The epithet ‘‘delacroxii’’ was derived from the name of E.G. Delacroix and thus the correct Latin form is “delacroixii”. The spelling ‘‘delacroxii’’ could be treated as a correctable orthographical error [Art. 60.1; McNeill et al. (2012)], but the name A. delacroixii was validly proposed twice before, making the name A. delacroxii Samson et al. illegitimate. Even if the epithet ‘‘delacroxii’’ was grammatically correct, its similarity to the epithet “delacroixii” would likely cause confusion [(Art. 53.3; McNeill et al. (2012)]. We follow Hubka et al. (2016a) and the correct name for A. nidulans var. echinulatus is Aspergillus spinulosporus Hubka et al.



Aspergillus ferenczii (Varga & Samson) Samson et al., Stud. Mycol. 78: 155. 2014.



Notes: Samson et al. (2007a) described Neosartorya ferenczii as a new species in their monograph on Aspergillus sect. Fumigati and due to the introduction of the single name nomenclature, the combination in Aspergillus was subsequently made in 2014 (Samson et al. 2014). In their monograph, Samson et al. (2007a) were unable to obtain the ex-type cultures of Neosartorya indohii, N. nishimurae, N. sublevispora, N. takakii, and N. tsurutae, and these species were therefore treated as doubtful. However, comparison of the BenA, CaM and RPB2 sequences of the ex-type strains on A. ferenczii (CBS 121594) and A. sublevisporus (CBS 128796 = IFM 53598) showed that these are similar (BenA, EF669833 vs AB488759: 100 %; CaM, EF669903 vs AB488767: 99.6 %; ITS, EF669977 vs MN431376: 100 %; RPB2, EF669764 vs MN969095: 99.8 %). Aspergillus ferenczii is regarded to be conspecific with Aspergillus sublevisporus Someya et al.



Aspergillus frequens Hubka et al., Mycologia 107: 183. 2015.



Notes: The type strains of A. frequens (NRRL 4578) and A. micronesiensis (CBS 138183) have highly similar BenA, CaM and RPB2 sequences. Based on these data, Arzanlou et al. (2016) reduced A. frequens (Hubka et al. 2015) in synonymy with A. micronesiensis Visagie et al. (Visagie et al. 2014a).



Aspergillus korhogoensis A. Carvajal-Campos et al., Toxins 9, 353: 11. 2017. nom. inval.



Notes: An identifier issued by a recognised repository for A. korhogoensis was not cited in the protologue and this species is therefore invalidly described [Art. 42.1, McNeill et al. (2012)]. Frisvad et al. (2019) studied the taxonomy of Aspergillus sect. Flavi and based on their studied specimens, they introduced Aspergillus cerealis. Based on sequence similarity, Aspergillus korhogoensis is treated as a synonym of A. cerealis Houbraken et al.



Aspergillus latus (Thom & Raper) A.J. Chen et al., Stud. Mycol. 84: 69. 2016.



Notes: Chen et al. (2016a) treated Aspergillus sublatus as a synonym of A. latus. Even though A. nidulans var. latus Thom & Raper (1939) is the oldest name of this species, the new combination/status proposed by Chen et al. (2016a) cannot have priority over A. sublatus (oldest name in the species rank) (Hubka et al. 2016a). The correct name for this species is Aspergillus sublatus Y. Horie.



Aspergillus mangaliensis A. Nováková et al., Mycologia 107: 187. 2015.



Notes: Based on gene concordance and a high sequence homology, Arzanlou et al. (2016) treated Aspergillus mangaliensis as a synonym of A. templicola Visagie et al. Their conclusion is followed here.



Aspergillus occiafricanus P. Singh & P.J. Cotty, Int. J. Food Microbiol. 289: 148. 2018, nom. inval.



Notes: This species was invalidly described (without an identifier, Art. F.5.1). NRRL A-11612 was designated as the type. The BenA (MK119746) and CaM (MK119712) sequences derived from the type strain have high homology (99.8 %) with the type of A. aflatoxiformans (CBS 143679). Aspergillus occiafricanus can therefore be considered an invalidly described synonym of A. aflatoxiformans Frisvad et al.



Aspergillus parafelis Sugui et al., J. Clin. Microbiol. 52: 3709. 2014.



Notes: Hubka et al. (2018a) re-evaluated the species boundaries in the Aspergillus viridinutans species complex using a large set of clinical and environmental strains. In their analyses, they showed that A. parafelis and A. pseudofelis are included in the genetically diverse A. felis lineage. This is in contrary to the results of Sugui et al. (2014); however, only two isolates of each species were included in their analysis and these isolates did not sufficiently cover the genetic diversity within A. felis (Hubka et al. 2018a).



Aspergillus pseudofelis Sugui et al., J. Clin. Microbiol. 52: 3709. 2014.



Notes: This species is a synonym of A. felis Barrs et al.; see A. parafelis.



Aspergillus similis (Y. Horie et al.) Samson et al., Stud. Mycol. 78: 157. 2014.



Notes: Samson et al. (2014) combined Emericella similis in Aspergillus. Chen et al. (2016a) noted that the ascospore morphology of the ex-type culture (CBS 293.93) is identical to that of Aspergillus violaceus. This close relationship is confirmed by sequence data and we follow the conclusion of Chen et al. (2016a) and treat A. similis as a synonym of A. violaceus Fennell & Raper.



Aspergillus texensis P. Singh et al., Toxins 10, 513: 6. 2018.



Notes: Molecular phylogenetic analyses of Aspergillus flavus-like isolates producing small-sized sclerotia (average sclerotium size < 400 μm) collected from across the United States resulted in the discovery of a novel aflatoxin-producing species, named Aspergillus texensis. The phylogenetic position of this species was studied using partial β-tubulin (0.9 kb), calmodulin (1.2 kb), and nitrate reductase (niaD, 2.1 kb) gene sequences (Singh et al. 2018). The former two gene regions are commonly used in Aspergillus taxonomy; however, the sequenced fragments are longer than those recommended by Samson et al. (2014). On the other hand, the nitrate reductase gene is part of the aflatoxin biosynthetic pathway and not commonly used in phylogenetic studies. Based on the results of Singh et al. (2018), A. texensis is closely related to A. minisclerotigenes and a previously reported unnamed lineage designated “Lethal Aflatoxicosis Fungus” (LAF). In this study, we re-evaluated the species boundaries of A. minisclerotigenes using a larger set of strains. Single gene phylogenies using BenA, CaM, and niaD sequences were constructed (using the longer fragments as described by Singh et al. (2018)) (Fig. 20). The five representative A. texensis strains clustered together in all analyses, indicating their close genetic relationship. The two included LAF strains (A1168, A1170) causing lethal aflatoxicosis in Kenya (ex maize), clustered together in the BenA and niaD phylogenies, and were closely related in the CaM phylogram. In the BenA phylogram, all A. minisclerotigenes, A. texensis and LAF strains clustered together with high statistical support and there were four minor deviating lineages present. Three main clades are present in the CaM phylogram. One clade contains the A. texensis strains and A. minisclerotigenes strains DTO 275-E9 and CBS 145094, one clade includes the two LAF strains, CBS 117635^T and four other A. minisclerotigenes strains and one clade with strains only identified as A. minisclerotigenes. There are also three main clades present in the niaD phylogram; however, the clade containing the LAF strains does not have statistical support. The position of A. minisclerotigenes CBS 145094 is basal to the other investigated strains. The A. texensis and the LAF strains clustered together in the combined analysis of the three genes, confirming the results of Singh et al. (2018). However, with the addition of A. minisclerotigenes strains and therefore increasing the genetic diversity within the analysis, it becomes clear that A. texensis and the LAF strains fall within the genetic diversity of A. minisclerotigenes. We therefore consider both as synonyms of A. minisclerotigenes Vaamonde et al.



Fig. 20.

Phylogenetic trees based on single and a combined data set of BenA, CaM and niaD sequences showing the relationship between Aspergillus texensis and related species. The BI posterior probability (pp) values and bootstrap percentages of the maximum likelihood (ML) analysis are presented at the nodes; fully supported branches are thickened. Values less than 70 % bootstrap support (ML) or less than 0.95 posterior probability (Bayesian analysis) are indicated with a hyphen or not shown. The bar indicates the number of substitutions per site. The phylogram is rooted with Aspergillus tamarii NRRL 20818.

Penicillium attenuatum Kirichuk & Pivkin, Mycol. Prog. 16: 21. 2017.



Notes: Kirichuk et al. (2017) introduced Penicillium attenuatum, P. ochotense and P. piltunense as new species closely related to P. antarcticum (sect. Canescentia, ser. Atroveneta) based on DNA sequences of the ITS, BenA and CaM regions. Penicillium sect. Canescentia is currently being reviewed (Visagie et al., unpublished) and the authors found that the published sequences for those species are of poor quality. Alignment of them against reference sequences revealed numerous suspicious regions towards read ends (e.g., gaps within coding regions of BenA and CaM, as well as conserved regions of the 28S rDNA region of ITS). Removing suspected low quality read ends, resulted in phylograms where all three species resolve with Penicillium antarcticum A.D. Hocking & C.F. McRae. Unfortunately, these strains are not available for study and are thus reduced to synonyms.



Penicillium cellarum Strausbaugh & Dugan, Pl. Dis. 101: 1783. 2017.



Notes: Strausbaugh & Dugan (2017) described P. cellarum from sugar beet. Based on BenA sequencing, this species is most closely related to P. aurantiogriseum in Penicillium sect. Fasciculata. Seven unique sequence types were found among the studied P. cellarum strains (F727^T, F759, F761, F769, F776, F785, F790) (Strausbaugh and Dugan, 2017, Strausbaugh, 2018). Those BenA, CaM and RPB2 sequences were re-analysed and compared with a set of nine P. aurantiogriseum strains from the CBS culture collection (Supplementary Fig. S4). The BenA analysis shows that all P. cellarum strains reside in a clade with P. aurantiogriseum with 88 % bootstrap support (no statistical support in the Bayesian analysis). The statistical support of the P. aurantiogriseum clade, including the P. cellarum sequences, is poor in the CaM analysis (<70 % BS, <0.95 pp). Strains F727^T, F769 and F785 cluster together within the P. aurantiogriseum clade (81 % BS, 0.99 pp), while the other P. cellarum strains cluster with strains previously identified as P. aurantiogriseum (Frisvad & Samson 2004b). The phylogenetic analysis of the RPB2 data set resulted in a less well-supported phylogram. The majority of the RPB2 sequences obtained from the CBS strains (including CBS 324.89^T) cluster with F769, F785, F761 and F790. A smaller clade with strains F727^T, F759 and F776 is present in the RPB2 phylogram (full support). The strains within this cluster differ from those in the cluster present in the CaM phylogram, showing that the phylograms are not congruent. These data indicate that P. cellarum is a synonym of P. aurantiogriseum Dierckx. This observation is also supported by the BenA phylogram.



Penicillium imranianum Imran Ali, Pakistan J. Bot. 50: 2055. 2018.



Notes: The deposited ITS sequence in GenBank (KP780293) cannot be reliably aligned in a dataset that contains the ITS barcodes of the accepted species in the families Aspergillaceae, Trichocomaceae and Thermoascaceae. This is probably due to a sequencing error, because also the conserved 5.8S nrDNA region is difficult to align. The taxonomic position of Penicillium imranianum remains unresolved and is considered a doubtful species. Other genes need to be sequenced to get insight in the taxonomic position and status of this species.



Penicillium ochotense Kirichuk & Pivkin, Mycol. Prog. 16: 21. 2017.



Notes: Synonym of Penicillium antarcticum A.D. Hocking & C.F. McRae; see notes for P. attenuatum above.



Penicillium pedernalense Laich & J. Andrade, Int. J. Syst. Evol. Microbiol. 66: 4386. 2016. nom. inval.



Notes: Penicillium pedernalense was invalidly described and validated in 2018. In the original description, two strain numbers (CBS 140770; CECT 20949) were indicated as the type (invalid under Art. 40.7, Melbourne). The name was later validated with the CBS strain designated as holotype (Index Fungorum 361: 1).



Penicillium piltunense Kirichuk & Pivkin, Mycol. Prog. 16: 19 2017.



Notes: Synonym of Penicillium antarcticum A.D. Hocking & C.F. McRae; see notes for P. attenuatum above.



Penicillium wandoense Hyang B. Lee et al., Fungal Diversity 96: 105. 2019.



Notes: Comparison of the BenA, CaM and RPB2 sequences deposited in GenBank shows that Penicillium laevigatum and P. wandoense are conspecific. Penicillium laevigatum was effectively published online on 19 December 2018 (online version of record) whereas the other article was only published on 24 June 2019. Penicillium laevigatum L. Cai et al. has priority over P. wandoense.



Phialosimplex halophilus [as “halophila”] (F.J.H. Beyma) Greiner et al., IMA Fungus 5: 170. 2014.



Notes: Samson et al. (2014) combined Basipetospora halophila (basionym Oospora halophila J.F.H. Beyma) in Aspergillus as Aspergillus baarnensis. In the same year, Greiner et al. (2014) combined this species in the genus Phialosimplex. Even though Phialosimplex species (e.g., Ph. caninus, the generic type) produce reduced conidiophores or solitary phialides (and lack typical aspergillus-type conidiophores), they phylogenetically belong to Aspergillus subgen. Polypaecilum (Fig. 17). We therefore accept A. baarnensis Samson et al., which is in agreement with Tanney et al. (2017).



Phialosimplex salinarum (Greiner et al.) Zalar & Greiner, Extremophiles 21: 762. 2017.



Notes: Martinelli et al. (2017) combined Ph. salinarum in Aspergillus (as Aspergillus salinarus) and Tanney et al. (2017) accepted this combination in their overview paper on Aspergillus subgen. Polypaecilum. Based on the multigene phylogenetic study performed here, we treat Phialosimplex as a synonym of Aspergillus and accept Aspergillus salinarum [as “salinarus”] (Greiner et al.) Zalar & Greiner over Ph. salinarum.



Talaromyces omanensis Halo et al., Phytotaxa 404: 192. 2019.



Notes: Penicillium resedanum belongs to Talaromyces and will be combined in this genus as Talaromyces resedanus elsewhere. Based on sequence and phenotypic similarities, Tal. omanensis as a synonym of Tal. resedanus (pers. communication, A. J. Chen).



Talaromyces rubrifaciens W.W. Gao, Mycologia 108: 775. 2016.



Notes: During a study on new Talaromyces species from indoor environments in China, Chen et al. (2016b) synonymised Tal. rubrifaciens with Tal. albobiverticillius based on GCPSR approach and phenotypic characters. Similarly as with A. parafelis and A. pseudofelis, a limited number of Tal. albobiverticillius sequences were included and the full sequence diversity was not represented in the phylogenetic analyses. We follow Chen et al. (2016b) and treat Tal. rubrifaciens as a synonym of Talaromyces albobiverticillius (H.-M. Hsieh et al.) Samson et al.

List of species in Eurotiales

A list of “Names in Current Use” (NCU) for the family Trichocomaceae was prepared in 1993 and was of great value for anyone working with these fungi (Pitt & Samson 1993). The list was updated in 2000, and this version included the accepted species and their synonyms in this family (Pitt et al. 2000). In total, 29 genera and 617 species were accepted. The genera Penicillium and Aspergillus made up for the majority of the included species. New overviews of accepted species in Aspergillus (Samson et al. 2014), Penicillium (Visagie et al. 2014b) and Talaromyces (Yilmaz et al. 2014) were published. These lists were urgently needed, due to many changes since the list of 2000. The majority of changes were caused by the move from a dual to a single name nomenclature system (McNeill et al. 2012), with the result that many teleomorphic genera were synonymised with Penicillium or Aspergillus (Houbraken and Samson, 2011, Samson et al., 2014). Furthermore, new names were introduced due to descriptions of new species, and old names were resurrected because taxonomic studies using molecular data showed that the phenotype-based species were actually consisting of complexes of multiple species (e.g., sibling species such as Aspergillus niger and A. welwitschiae; Hong et al. (2013). Another advantage of the “2014 Aspergillus/Penicillium/Talaromyces lists” are the inclusion of MycoBank numbers, type and ex-type culture (accession number) information and GenBank accession numbers for BenA, CaM, ITS and RPB2 sequences.

 Compared with the “2014 Aspergillus/Penicillium/Talaromyces lists”, the list of species presented below is expanded with the species described after publication in 2014. In the “2014 Aspergillus/Penicillium/Talaromyces lists”, 339 Aspergillus, 354 Penicillium and 88 Talaromyces species were accepted. These numbers increased significantly, and now includes 446 Aspergillus (32 % increase), 483 Penicillium (36 % increase) and 171 Talaromyces (94 % increase) species. An overview of the number of accepted species in the 2000, 2014 and the current list is given in Table 6. We decided that if no sequence data is available for examination of the type or representative strain, then we only included the species that were described since 1970. Also, species such as A. amazonensis were excluded because it was described in 1904, but combined later (in 1985) (Samson & Seifert 1985). Additional BenA, CaM, ITS and RPB2 sequences were generated for the species lacking this data in GenBank or for those that were linked to poor or short sequences. Sequence data are now available for the majority of species. Of the 446 Aspergillus taxa, 3.1 % (n = 14), 3.1 % (n = 14), 4.5 % (n = 20) and 5.1 % (n = 23) of the species are missing BenA, CaM, ITS or RPB2 sequences, respectively. No sequence data are available for 13 species: Aspergillus argenteus, A. beijingensis, A. collembolorum, A. crassihyphae, A. curviformis, A. ellipsoideus, A. maritimus, A. qizutongii, A. raianus, A. subunguis, A. tapirirae, A. vinosobubalinus, A. wangduanlii, and only an ITS sequence is available for Aspergillus assiutensis. The taxonomic position of these species needs to be confirmed. The list contains 483 Penicillium species and BenA, CaM, ITS and RPB2 sequences are lacking for 1.9 % (n = 8), 2.3 % (n = 11), 2.3 %, (n = 11) and 5.4 % (n = 26) of the taxa, respectively. No sequence data are available Penicillium asymmetricum, P. coniferophilum, P. glaucoalbidum, P. longisporum, P. melanostipe, P. parviverrucosum and P. taiwanense. Besides these species, only P. dravunii is not represented by a BenA sequence (an ITS sequence is known though, which is distinct from all other accepted species). Out of the 171 Talaromyces species, three (1.8 %) taxa do not have a BenA sequence, five (2.9 %) are lacking CaM sequences, one (0.6 %) is lacking an ITS sequence and nine (5.3 %) RPB2 sequences. New in the list is the inclusion of other genera and species belonging to the Eurotiales (e.g., Monascus, Paecilomyces, Rasamsonia), with exception of taxa classified in the Elaphomycetaceae. The list includes 1 187 species, distributed over 27 genera (Table 6). Due to the change to a single name nomenclature system, a name does not reflect the ability of a species to produce a sexual morph anymore. In order to make this information readily available, we also included information on the mode of reproduction. Species could be asexual, homothallic, heterothallic or protoheterothallic. The latter term is used in those species where both idiomorphs are known in the species populations; if only one of the idiomorphs is known, then the species is listed as asexual (Houbraken & Dyer 2015). Information on several type specimens are not available or has been lost for various reasons. Where ex-type strains were available, new dried specimens were prepared and the typifications fixed below.

Table 6.

Genera and number of species in current and earlier lists.

Genus	Number of accepted species			Number of species producing sexual morph
	Pitt et al. (2000)	2014-lists3	This study4
Acidotalaromyces	–	–	1	0
Ascospirella	–	–	1	1
Aspergillago	–	–	1	0
Aspergillus1	184	339	446	139 (incl. 19 heterothallic spp.)
Byssochlamys	4	–	Paecilomyces	n/a
Chaetosartorya	3	–	Aspergillus	n/a
Cristaspora	1	–	Aspergillus	n/a
Dendrosphaera	1	–	1	1
Dichlaena	1	–	2	2
Dichotomomyces	1	–	Aspergillus	n/a
Emericella	27	–	Aspergillus	n/a
Eupenicillium	43	–	Penicillium	n/a
Eurotium	19	–	Aspergillus	n/a
Evansstolkia	–	–	1	1
Fennellia	3	–	Aspergillus	n/a
Geosmithia1	8	–	Hypocreales	n/a
Hamigera	1	–	9	8
Hemicarpenteles	2	–	Penicillium	n/a
Leiothecium	–	–	2	2
Merimbla1	1	–	Hamigera	n/a
Monascus	222	–	9	7
Neosartorya	12	–	Aspergillus	n/a
Paecilomyces1	41	–	10	5 (incl. 1 heterothallic spp.)
Penicillago	–	–	4	0
Penicilliopsis	2	–	4	3
Penicillium1	225	354	483	64 (incl. 2 heterothallic spp.)
Petromyces	2	–	Aspergillus	n/a
Phialomyces	–	–	4	0
Pseudohamigera	–	–	1	1
Pseudopenicillium	–	–	3	3
Raperia1	1	–	Warcupiella	n/a
Rasamsonia	–	–	12	4
Sagenomella	–	–	6	1
Sarophorum1	1	–	Penicilliopsis	n/a
Sclerocleista	2	–	2	2
Stilbodendron1	1	–	Penicilliopsis	n/a
Talaromyces	24	88	171	40 (incl. 3 heterothallic spp.)
Thermoascus	4	–	7	7
Thermomyces	–	–	2	1
Torulomyces1	1	–	Penicillium	n/a
Trichocoma	1	–	1	1
Warcupiella	1	–	1	1
Xerochrysium	–	–	2	0
Xeromyces	–	–	1	1
Total	6172	7813	1 187	295 (incl. 25 heterothallic spp.)

¹Genera originally introduced for strictly asexual reproducing species.

²Species producing a sexual and asexual morph are counted two times in this list due to the at that time dual nomenclature rules.

³Species of genera other than Aspergillus, Penicillium and Talaromyces were not included in the 2014-lists.

⁴If applicable, then the current generic name is provided.

List of accepted species

Acidotalaromyces

Acidotalaromyces lignorum (Stolk) Houbraken, Frisvad & Samson, this study. 2020. [MB832555]. Basionym: Penicillium lignorum Stolk, Antonie van Leeuwenhoek 35: 264. 1969. [MB335743]. — Type: CBS 709.68. Ex-type: CBS 709.68 = ATCC 22051 = FRR 804 = IMI 151899 = UPSC 3184. Reproduction: asexual. ITS barcode: JF910285 (alternative markers: BenA = HQ156946; CaM = JX140680; RPB2 = MN969215).



Ascospirella

Ascospirella lutea (Zukal) Houbraken et al., this study. 2020. [MB832556]. Basionym: Penicillium luteum Zukal, Sitzungsber. Kaiserl. Akad. Wiss. Math.-Naturwiss. Cl., Abt. 1 98: 561. 1890. [MB306716]. — Type: IMI 89305. Ex-type: CBS 348.51 = DTO 165-C7 = CECT 2950 = IFO 31753 = IMI 089305 = LSHB BB228. Reproduction: homothallic. ITS barcode: MN431414 (alternative markers: BenA = MN969437; CaM = MN969356; RPB2 = MN969216).



Aspergillago

Aspergillago clavatoflava (Raper & Fennell) Samson et al., Stud. Mycol. 85: 211. 2016. [MB819187]. Basionym: Aspergillus clavatoflavus Raper & Fennell, Gen. Aspergillus: 378. 1965. [MB326619]. — Type: WB5113. Ex-type: DTO 022-B2 = CBS 473.65 = NRRL 5113 = ATCC 16866 = IMI 124937 = LCP 89.2589. Reproduction: asexual. ITS barcode: EF669713 (alternative markers: BenA = EF669686; CaM = EF669700; RPB2 = EF669668).



Aspergillus

Aspergillus acanthosporus Udagawa & Takada, Bull. Nat. Sci. Mus. Tokyo 14: 503. 1971. [MB309201]. — Type: NHL 22462. Ex-type: CBS 558.71 = NRRL 5293 = ATCC 22931 = IMI 164621 = NHL 2462. Infragen. class: subgen. Fumigati, sect. Clavati, ser. Clavati. Reproduction: homothallic. ITS barcode: EU078625 (alternative markers: BenA = MN969364; CaM = EU078676; RPB2 = EF669779).

Aspergillus acidohumus A.J. Chen et al., Stud. Mycol. 85: 71. 2016. [MB817723]. — Type: CBS H-22730. Ex-type: CBS 141577 = CGMCC 3.18217 = DTO 340-H1 = IBT 34346. Infragen. class: subgen. Fumigati, sect. Cervini, ser. Acidohumorum. Reproduction: asexual. ITS barcode: KX423646 (alternative markers: BenA = KX423623; CaM = KX423634; RPB2 = KX423663).

Aspergillus acrensis Hubka et al., Persoonia 41: 163. 2018. [MB822542]. — Type: IFM 57291H. Ex-type: IFM 57291 = CCF 4670. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Viridinutantes. Reproduction: protoheterothallic; both MAT idiomorphs detected (Hubka et al. 2018a). ITS barcode: n.a. (alternative markers: BenA = LT795980; CaM = LT795981; RPB2 = LT795982).

Aspergillus aculeatinus Noonim et al., Int. J. Syst. Evol. Microbiol. 58: 1733. 2008. [MB505075]. — Type: unknown. Ex-type: CBS 121060 = DTO 202-G5 = IBT 29077. Infragen. class: subgen. Circumdati, sect. Nigri, ser. Japonici. Reproduction: protoheterothallic; unpublished (genome data; Vesth et al. 2018). ITS barcode: EU159211 (alternative markers: BenA = EU159220; CaM = EU159241; RPB2 = HF559233).

Aspergillus aculeatus Iizuka, J. Agric. Chem. Soc. Japan 27: 806. 1953. [MB292831]. — Type: IMI 211388. Ex-type: CBS 172.66 = NRRL 5094 = NRRL 20623 = IMI 211388 = ATCC 16872 = WB 5094. Infragen. class: subgen. Circumdati, sect. Nigri, ser. Japonici. Reproduction: protoheterothallic; MAT1-2-1 detected (de Vries et al. 2017). ITS barcode: EF661221 (alternative markers: BenA = HE577806; CaM = EF661148; RPB2 = EF661046).

Aspergillus aeneus Sappa, Allionia 2: 84. 1954. [MB292832]. — Type: CBS H-6735. Ex-type: CBS 128.54 = NRRL 4769 = ATCC 16803 = IMI 69855 = LSHBBB 355 = MUCL 13570 = QM 1945 = WB 4279 = WB 4769. Infragen. class: subgen. Nidulantes, sect. Aenei, ser. Aenei. Reproduction: asexual. ITS barcode: EF652474 (alternative markers: BenA = EF652298; CaM = EF652386; RPB2 = EF652210).

Aspergillus aerius A.J. Chen et al., Stud. Mycol. 88: 79. 2017. [MB818731]. — Type: CBS H-22823. Ex-type: CBS 141771 = DTO 241-G7 = IBT 34446. Infragen. class: subgen. Aspergillus, sect. Aspergillus, ser. Aspergillus. Reproduction: homothallic. ITS barcode: LT670916 (alternative markers: BenA = LT670990; CaM = LT670991; RPB2 = LT670992).

Aspergillus affinis Davolos et al., Int. J. Syst. Evol. Microbiol. 62: 1014. 2012. [MB517245]. — Type: ATCC MYA-4773. Ex-type: CBS 129190 = DTO 223-C6 = BT 32310 = TCC MYA-4773. Infragen. class: subgen. Circumdati, sect. Circumdati, ser. Circumdati. Reproduction: asexual. ITS barcode: MN431360 (alternative markers: BenA = GU721092; CaM = GU721091; RPB2 = MN969063).

Aspergillus aflatoxiformans Frisvad et al., Stud. Mycol. 93: 32. 2019. [MB823770]. — Type: CBS H-23361. Ex-type: CBS 143679 = DTO 228-G2 = IBT 32085. Infragen. class: subgen. Circumdati, sect. Flavi, ser. Flavi. Reproduction: protoheterothallic; both MAT idiomorphs detected (Carvajal-Campos et al. 2017, referred to as A. parvisclerotigenes). ITS barcode: MG662388 (alternative markers: BenA = MG517706; CaM = MG518076; RPB2 = MG517897).

Aspergillus alabamensis Balajee et al., Eukaryot. Cell 8: 720. 2009. [MB543648]. — Type: UAB20. Ex-type: CBS 125693 = UAB20 = DTO 045-C5. Infragen. class: subgen. Circumdati, sect. Terrei, ser. Terrei. Reproduction: asexual. ITS barcode: KP987071 (alternative markers: BenA = KP987049; CaM = EU147583; RPB2 = KP987018).

Aspergillus allahabadii B.S. Mehrotra & Agnihotri, Mycologia 54: 400. 1963. [MB326609]. — Type: CBS H-6736. Ex-type: CBS 164.63 = NRRL 4539 = ATCC 15055 = IMI 139273 = MUCL 13571 = WB 4539. Infragen. class: subgen. Circumdati, sect. Terrei, ser. Nivei. Reproduction: asexual. ITS barcode: EF669601 (alternative markers: BenA = EF669531; CaM = EF669559; RPB2 = EF669643).

Aspergillus alliaceus Thom & Church, Aspergilli: 163. 1926. [MB256402]. — Type: CBS H-7812. Ex-type: CBS 536.65 = DTO 034-B3 = DTO 046-B1 = ATCC 10060 = DSM 813 = IFO 7538 = IMI 051982 = IMI 051982ii = NRRL 315 = QM 1885 = WB 315. Infragen. class: subgen. Circumdati, sect. Flavi, ser. Alliacei. Reproduction: homothallic (Fennell & Warcup 1959). ITS barcode: EF661551 (alternative markers: BenA = EF661465; CaM = EF661534; RPB2 = MG517825).

Aspergillus amazonicus D. Mares, Curr. Microbiol. 57: 228. 2008. [MB531888]. — Type: E19D. Ex-type: CBS 124228 = DTO 092-D6 = DTO 411-B6. Infragen. class: subgen. Nidulantes, sect. Sparsi, ser. Conjuncti. Reproduction: asexual. ITS barcode: MN431399 (alternative markers: BenA = FJ943939; CaM = FJ943936; RPB2 = KU866979).

Aspergillus ambiguus Sappa, Allionia 2: 254. 1955. [MB292834]. — Type: CBS H-6737. Ex-type: CBS 117.58 = NRRL 4737 = ATCC 16827 = IMI 139274 = QM 8155 = WB 4737. Infragen. class: subgen. Circumdati, sect. Terrei, ser. Ambigui. Reproduction: asexual. ITS barcode: EF669606 (alternative markers: BenA = EF669534; CaM = EF669564; RPB2 = EF669648).

Aspergillus amethystinus F. Sklenář et al., Mycologia 112: 356. 2020. [MB832712]. — Type: PRM 951579. Ex-type: NRRL 4178 = CCF 5261. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Nidulantes. Reproduction: homothallic. ITS barcode: EF652462 (alternative markers: BenA = EF652286; CaM = EF652374; RPB2 = EF652198).

Aspergillus amoenus M. Roberg, Hedwigia 70: 138. 1931. [MB250654]. — Type: Münster i.W., isol. ex Berberis sp. fruit, M. Roberg (type locality, this specimen was not deposited into herbarium). Ex-type: NRRL 4838 = CBS 111.32. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Versicolores. Reproduction: protoheterothallic; MAT 1-1-1 detected . ITS barcode: EF652480 (alternative markers: BenA = JN853946; CaM = JN854035; RPB2 = JN853824).

Aspergillus angustatus A.J. Chen et al., Stud. Mycol. 84: 41. 2016. [MB816090]. — Type: CBS H-22487. Ex-type: CBS 273.65 = DTO 319-H8. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Stellati. Reproduction: homothallic. ITS barcode: EU448283 (alternative markers: BenA = AY339993; CaM = EU443984; RPB2 = KU867013).

Aspergillus anthodesmis Bartoli & Maggi, Trans. Brit. Mycol. Soc. 71: 386. 1979 [1978]. [MB309207]. — Type: RO 103 S. Ex-type: CBS 552.77 = NRRL 22884 = IMI 223070. Infragen. class: subgen. Nidulantes, sect. Sparsi, ser. Conjuncti. Reproduction: asexual. ITS barcode: FJ491662 (alternative markers: BenA = EF661108; CaM = FJ491648; RPB2 = EF661039).

Aspergillus appendiculatus Blaser, Sydowia 28: 38. 1975. [MB309209]. — Type: ZT 8286. Ex-type: CBS 374.75 = DTO 196-H3 = ETH8286 = IMI 278374 = KACC 45268. Infragen. class: subgen. Aspergillus, sect. Aspergillus, ser. Rubri. Reproduction: homothallic. ITS barcode: HE615132 (alternative markers: BenA = HE801333; CaM = HE801318; RPB2 = HE801307).

Aspergillus arachidicola Pildain et al., Int. J. Syst. Evol. Microbiol. 58: 730. 2008. [MB505189]. — Type: Pildain et al. 2008, Int. J. Syst. Evol. Microbiol. 58: p. 731 Fig. 2 (– lectotype designated here, MBT392255; CBS H-24274 [dried culture] – epitype designated here, MBT392256). Ex-epitype: DTO 009-G3 = CBS 117610 = IBT 117610 = IBT 25020. Infragen. class: subgen. Circumdati, sect. Flavi, ser. Flavi. Reproduction: protoheterothallic; both MAT idiomorphs detected (Carvajal-Campos et al. 2017). ITS barcode: MF668184 (alternative markers: BenA = EF203158; CaM = EF202049; RPB2 = MG517802).

Aspergillus arcoverdensis Y. Horie et al., Mycoscience 56: 130. 2015. [MB804028]. — Type: CBM-FA-39845. Ex-type: IFM 61334 = JCM 19878 = CCF 4695 = CBS 139187 = DTO 316-F7. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Viridinutantes. Reproduction: protoheterothallic; both MAT idiomorphs detected (Matsuzawa et al. 2015). ITS barcode: MN431385 (alternative markers: BenA = AB818845; CaM = AB818856; RPB2 = MN969103).

Aspergillus ardalensis A. Nováková et al., Mycologia 107: 179. 2015. [MB808140]. — Type: PRM 923450. Ex-type: CCF 4031 = CCF 4426 = CMF ISB 1688 = CBS 134372 = NRRL 62824. Infragen. class: subgen. Circumdati, sect. Flavipedes, ser. Flavipedes. Reproduction: asexual. ITS barcode: FR733808 (alternative markers: BenA = HG916683; CaM = HG916725; RPB2 = HG916704).

Aspergillus arenarioides Visagie, Hirooka & Samson, Stud. Mycol. 78: 110. 2014. [MB809195]. — Type: CBS H-21812. Ex-type: CBS 138200 = DTO 268-E3 = CCF 4928. Infragen. class: subgen. Circumdati, sect. Petersoniorum, ser. Petersoniorum. Reproduction: asexual. ITS barcode: KJ775562 (alternative markers: BenA = KJ775091; CaM = KJ775390; RPB2 = LN849430).

Aspergillus argenteus J.N. Rai & H.J. Chowdhery, Kavaka 7: 19. 1979. [MB116063]. — Type: MLLU 104. Ex-type: Infragen. class: subgen. Nidulantes, sect.: unknown, ser.: unknown. Reproduction: asexual. ITS barcode: n.a. (alternative markers: BenA = n.a.; CaM = n.a.; RPB2 = n.a.).

Aspergillus arxii (Fort & Guarro) Houbraken, Visagie & Samson, Stud. Mycol. 78: 154. 2014. [MB809575]. Basionym: Cristaspora arxii Fort & Guarro, Mycologia 76: 1115. 1984. [MB106038]. — Type: CBS H-14047. Ex-type: CBS 525.83 = ATCC 52744 = FMR 416. Infragen. class: subgen. Cremei, sect. Cremei, ser. Arxiorum. Reproduction: homothallic; asexual morph unknown. ITS barcode: MN431361 (alternative markers: BenA = MN969365; CaM = MN969223; RPB2 = JN121529).

Aspergillus asclerogenus Jurjević & Hubka, Plant Syst. Evol. 301: 2451. 2015. [MB814441]. — Type: PRM 933843. Ex-type: CCF 4947 = NRRL 58502. Infragen. class: subgen. Circumdati, sect. Petersoniorum, ser. Petersoniorum. Reproduction: asexual. ITS barcode: LN849392 (alternative markers: BenA = LN849406; CaM = LN849421; RPB2 = LN849437).

Aspergillus askiburgiensis A. Nováková et al., Plant Syst. Evol. 302: 1285. 2016. [MB816280]. — Type: PRM 924055. Ex-type: CCF 4716 = CCF 4428 = CBS 134374 = NRRL 62818 = IBT 33114 = IBT 32911. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Speluncei. Reproduction: asexual. ITS barcode: LN873939 (alternative markers: BenA = LN873952; CaM = LN873965; RPB2 = LN873984).

Aspergillus aspearensis Houbraken et al., Stud. Mycol. 93: 32. 2019. [MB823771]. — Type: CBS H-23358. Ex-type: CBS 143672 = DTO 203-D9 = CCTU 758 = IBT 32590 = IBT 34544. Infragen. class: subgen. Circumdati, sect. Flavi, ser. Leporum. Reproduction: asexual. ITS barcode: MG662398 (alternative markers: BenA = MG517669; CaM = MG518040; RPB2 = MG517857).

Aspergillus asper Jurjević & S.W. Peterson, Int. J. Syst. Evol. Microbiol. 66: 2567. 2016. [MB814412]. — Type: BPI-893218. Ex-type: CBS 140842 = NRRL 35910 = CCF 5174. Infragen. class: subgen. Nidulantes, sect. Usti, ser. Calidousti. Reproduction: asexual. ITS barcode: KT698840 (alternative markers: BenA = KT698838; CaM = KT698839; RPB2 = KT698842).

Aspergillus asperescens Stolk, Antonie van Leeuwenhoek 20: 303. 1954. [MB292835]. — Type: IMI 46813. Ex-type: CBS 110.51 = NRRL 2252 = NRRL 4770 = ATCC 11079 = DSM 871 = IMI 46813 = QM 1946 = WB 2252 = WB 4770 = WB 5038. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Speluncei. Reproduction: asexual. ITS barcode: EF652475 (alternative markers: BenA = EF652299; CaM = EF652387; RPB2 = EF652211).

Aspergillus assiutensis Moub. & Soliman, J. Basic Appl. Mycol. 2: 84. 2011. [MB584202]. — Type: AUMC 5748. Ex-type: CBS 132773 = AUMC 5748. Infragen. class: subgen. Circumdati, sect. Nigri, ser. Japonici. Reproduction: asexual. ITS barcode: JN393254 (alternative markers: BenA = n.a.; CaM = n.a.; RPB2 = n.a.).

Aspergillus assulatus (S.B. Hong et al.) Houbraken et al., Stud. Mycol. 78: 154. 2014. [MB809576]. Basionym: Neosartorya assulata S.B. Hong et al., Antonie van Leeuwenhoek 93: 95. 2008. [MB506376]. — Type: KACC 41691. Ex-type: IBT 27911 = DTO 043-E8. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Unilaterales. Reproduction: homothallic. ITS barcode: HF545007 (alternative markers: BenA = DQ114123; CaM = MN969222; RPB2 = HF545311).

Aspergillus astellatus (Fennell & Raper) Houbraken et al., Stud. Mycol. 78: 154. 2014. [MB809577]. Basionym: Aspergillus variecolor var. astellatus Fennell & Raper, Mycologia 47: 81. 1955. [MB346549]. — Type: IMI 061455. Ex-type: CBS 134.55 = CBS 261.93 = NRRL 2396 = ATCC 16817 = IMI 61455 = IMI 61455ii = NRRL A-1634 = QM 1910 = WB 2396. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Stellati. Reproduction: homothallic. ITS barcode: EF652446 (alternative markers: BenA = EF652270; CaM = EF652358; RPB2 = EF652182).

Aspergillus atacamensis Zalar et al., Extremophiles 21:766. 2017. [MB818565]. — Type: CBS H-23062. Ex-type: EXF-6660 = CBS 142046 = DTO 411-B9. Infragen. class: subgen. Polypaecilum, sect. Polypaecilum, ser. Kalimarum. Reproduction: asexual. ITS barcode: KX900619 (alternative markers: BenA = MN969415; CaM = MN969325; RPB2 = MN969192).

Aspergillus aurantiacoflavus Hubka et al., Stud. Mycol. 88: 82. 2017. [MB818732]. — Type: CBS H-22827. Ex-type: CBS 141930 = EMSL No. 2903 = CCF 5393 = DTO 355-I1 = IBT 34485. Infragen. class: subgen. Aspergillus, sect. Aspergillus, ser. Aspergillus. Reproduction: homothallic. ITS barcode: LT670917 (alternative markers: BenA = LT670993; CaM = LT670994; RPB2 = LT670995).

Aspergillus aurantiobrunneus (G.A. Atkins et al.) Raper & Fennell, Gen. Aspergillus: 511. 1965. [MB326612]. Basionym: Emericella nidulans var. aurantiobrunnea G.A. Atkins, Hindson & A.B. Russell, Trans. Brit. Mycol. Soc. 41: 504. 1958 [MB346743]. — Type: IMI 74897. Ex-type: CBS 465.65 = NRRL 4545 = NRRL 2775 = ATCC 16821 = IMI 074897. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Aurantiobrunnei. Reproduction: homothallic. ITS barcode: EF652465 (alternative markers: BenA = EF652289; CaM = EF652377; RPB2 = EF652201).

Aspergillus aurantiopurpureus A.J. Chen et al., Stud. Mycol. 84: 46. 2016. [MB816087]. — Type: CBS H-22488. Ex-type: CBS 140608 = IBT 12601 = DTO 060-A7. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Nidulantes. Reproduction: homothallic. ITS barcode: KU866588 (alternative markers: BenA = KU866824; CaM = KU866711; RPB2 = KU866966).

Aspergillus auratus Warcup, Gen. Aspergillus: 263. 1965. [MB326613]. — Type: IMI 75886. Ex-type: CBS 466.65 = NRRL 4378 = ATCC 16894 = IFO 8783 = IMI 75886 = QM 7861 = WB 4378. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Neoglabri. Reproduction: homothallic. ITS barcode: EF669979 (alternative markers: BenA = EF669835; CaM = EF669905; RPB2 = EF669766).

Aspergillus aureolatus Munt.-Cvetk. & Bata, Bull. Inst. Jard. Bot. Univ. Beograd 1: 196. 1964. [MB326614]. — Type: CBS H-6738. Ex-type: CBS 190.65 = NRRL 5126 = ATCC 16810 = IMI 136527 = IMI 136527ii = WB 5126. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Speluncei. Reproduction: asexual. ITS barcode: EF652501 (alternative markers: BenA = EF652325; CaM = EF652413; RPB2 = EF652237).

Aspergillus aureolus Fennell & Raper, Mycologia 47: 71. 1955. [MB292836]. — Type: CBS 105.55. Ex-type: CBS 105.55 = NRRL 2244 = ATCC 16896 = IFO 8105 = IMI 61451 = IMI 061451ii = MUCL 13579 = QM 1906 = WB 2244 = DTO 052-C8 = DTO 331-G6. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Viridinutantes. Reproduction: homothallic. ITS barcode: EF669950 (alternative markers: BenA = EF669808; CaM = EF669877; RPB2 = EF669738).

Aspergillus aureoterreus Samson et al., Stud. Mycol. 69: 45. 2011. [MB560392]. — Replaced synonym: Aspergillus terreus var. aureus Thom & Raper, A manual of the Aspergilli: 198. 1945. [MB351655]. — Type: Thom & Raper 1945, A Manual of the Aspergilli: p. 199 Fig. 57B (– lectotype designated here, MBT392257; CBS H-24275 [dried culture] – epitype designated here, MBT392258). Ex-epitype: CBS 503.65 = NRRL 1923 = ATCC 16793 = IFO 30536 = IMI 82431 = MUCL 38644 = QM 7472 = VKM F-2035 = WB 1923. Infragen. class: subgen. Circumdati, sect. Terrei, ser. Terrei. Reproduction: asexual. ITS barcode: EF669580 (alternative markers: BenA = EF669524; CaM = EF669538; RPB2 = EF669622).

Aspergillus auricomus (Guég.) Saito, J. Ferment. Technol. 17: 3. 1939. [MB119950]. Basionym: Sterigmatocystis auricoma Guég., Bull. Soc. Mycol. Fr. 15: 186. 1899. [MB209799]. — Type: CBS H-9173. Ex-type: CBS 467.65 = NRRL 391 = IBT 14581 = ATCC 16890 = IMI 172277 = LCP 89.2596 = LSHBA 41 = WB 391. Infragen. class: subgen. Circumdati, sect. Circumdati, ser. Circumdati. Reproduction: asexual. ITS barcode: EF661411 (alternative markers: BenA = EF661320; CaM = EF661379; RPB2 = EF661300).

Aspergillus australiensis [as “australensis”] (Samson et al.) Houbraken et al., Stud. Mycol. 78: 154. 2014. [MB821660]. Basionym: Neosartorya australensis Samson, S.B. Hong & Varga, Stud. Mycol. 59: 174. 2007. [MB492203]. — Type: CBS 112.55. Ex-type: CBS 112.55 = NRRL 2392 = IMI 061450 = DTO 026-H3. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Neoglabri. Reproduction: homothallic. ITS barcode: EF669953 (alternative markers: BenA = EF669811; CaM = EF669880; RPB2 = EF669741).

Aspergillus austroafricanus Jurjević et al., IMA Fungus 3: 67. 2012. [MB800597]. — Type: BPI 880914. Ex-type: CBS 145748 = NRRL 233 = DTO 225-D8. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Versicolores. Reproduction: asexual. ITS barcode: JQ301891 (alternative markers: BenA = JN853963; CaM = JN854025; RPB2 = JN853814).

Aspergillus austwickii Frisvad et al., Stud. Mycol. 93: 38. 2019. [MB823772]. — Type: CBS H-23360. Ex-type: CBS 143677 = DTO 228-F7 = IBT 32590 = IBT 32076. Infragen. class: subgen. Circumdati, sect. Flavi, ser. Flavi. Reproduction: asexual. ITS barcode: MG662391 (alternative markers: BenA = MG517702; CaM = MG518072; RPB2 = MG517893).

Aspergillus avenaceus G. Sm., Trans. Brit. Mycol. Soc. 26: 24. 1943. [MB284296]. — Type: CBS H-6739. Ex-type: CBS 109.46 = NRRL 517 = ATCC 16861 = IMI 16140 = LCP 89.2592 = LSHBBB 155 = QM 6741 = WB 517. Infragen. class: subgen. Circumdati, sect. Flavi, ser. Avenacei. Reproduction: asexual. ITS barcode: AF104446 (alternative markers: BenA = FJ491481; CaM = FJ491496; RPB2 = JN121424).

Aspergillus baarnensis Samson et al., Stud. Mycol. 78: 154. 2014. [MB809579]. Replaced synonym: Oospora halophila J.F.H. Beyma, Zentralbl. Bakteriol. Parasitenk., Abt. 2 88: 134. 1933. [MB266778]. — Type: van Beyma 1933, Zentralbl. Bakteriol. Parasitenk., Abt. 2 88: p 134 Fig. 2 (– lectotype designated here, MBT392360; CBS H-24276 [dried culture] – epitype designated here, MBT392361). Ex-epitype: CBS 232.32 = VKM F-204. Infragen. class: subgen. Polypaecilum, sect. Polypaecilum, ser. Salinarum. Reproduction: asexual. ITS barcode: KY980621 (alternative markers: BenA = KY980549; CaM = KY980585; RPB2 = JN121509).

Aspergillus baeticus A. Nováková & Hubka, Int. J. Syst. Evol. Microbiol. 62: 2783. 2012. [MB564188]. — Type: PRM 860609. Ex-type: NRRL 62501 = CCF 4226 = CMFISB 2153. Infragen. class: subgen. Nidulantes, sect. Usti, ser. Usti. Reproduction: asexual. ITS barcode: HE615086 (alternative markers: BenA = HE615092; CaM = HE615117; RPB2 = HE615124).

Aspergillus beijingensis D.M. Li et al., Mycoscience 39: 299. 1998. [MB446575]. — Type: CBM FD-285. Ex-type: CBM FD-285. Infragen. class: subgen.: unknown, sect.: unknown, ser.: unknown. Reproduction: asexual. ITS barcode: n.a. (alternative markers: BenA = n.a.; CaM = n.a.; RPB2 = n.a.).

Aspergillus bertholletiae [as “bertholletius”] Taniwaki et al., PLoS ONE 7: e42480, 6. 2012. [MB622229]. — Type: CCT 7615. Ex-type: DTO 223-D3 = ITAL 270/06 = IBT 29228. Infragen. class: subgen. Circumdati, sect. Flavi, ser. Bertholletiarum. Reproduction: protoheterothallic; MAT1-1-1 detected (Carvajal-Campos et al. 2017). ITS barcode: JX198673 (alternative markers: BenA = MG517689; CaM = MN969224; RPB2 = MG517880).

Aspergillus bezerrae J.P. Andrade et al., Persoonia 42: 379. 2019. [MB830186]. — Type: HURB 22323 (holotype). Ex-type: CCDCA 11511 = 9EM2. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Viridinutantes. Reproduction: Heterothallic. ITS barcode: n.a. (alternative markers: BenA = MK597913; CaM = MK597915; RPB2 = n.a.).

Aspergillus bicephalus J.P.Z. Siqueira et al., Persoonia 37: 283. 2016. [MB818290]. — Type: CBS H-22807. Ex-type: CBS 142900 = FMR 14918. Infragen. class: subgen. Circumdati, sect. Terrei, ser. Nivei. Reproduction: asexual. ITS barcode: LT601380 (alternative markers: BenA = LT601381; CaM = LT601382; RPB2 = LT601383).

Aspergillus bicolor M. Chr. & States, Mycologia 70: 337. 1978. [MB309212]. — Type: NY RMF 2058. Ex-type: CBS 425.77 = NRRL 6364 = ATCC 36104 = IMI 216612. Infragen. class: subgen. Nidulantes, sect. Aenei, ser. Aenei. Reproduction: homothallic. ITS barcode: EF652511 (alternative markers: BenA = EF652335; CaM = EF652423; RPB2 = EF652247).

Aspergillus biplanus Raper & Fennell, Gen. Aspergillus: 434. 1965. [MB326615]. — Type: IMI 235602. Ex-type: CBS 468.65 = NRRL 5071 = ATCC 16858 = IMI 235602 = QM 8873 = WB 5071. Infragen. class: subgen. Nidulantes, sect. Sparsi, ser. Biplani. Reproduction: asexual. ITS barcode: EF661210 (alternative markers: BenA = EF661116; CaM = EF661130; RPB2 = EF661036).

Aspergillus bisporus Kwon-Chung & Fennell, Mycologia 63: 479. 1971. [MB309213]. — Type: BPI NRRL 3693. Ex-type: CBS 707.71 = NRRL 3693 = ATCC 22527 = IMI 350350 = NRRL A-17271 = QM 9700. Infragen. class: subgen. Nidulantes, sect. Bispori, ser. Bispori. Reproduction: asexual. ITS barcode: EF661208 (alternative markers: BenA = EF661121; CaM = EF661139; RPB2 = EF661077).

Aspergillus botswanensis A.J. Chen et al., Stud. Mycol. 84: 49. 2016. [MB816095]. — Type: CBS H-22494. Ex-type: CBS 314.89 = DTO 047-I4. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Nidulantes. Reproduction: homothallic; asexual morph unknown. ITS barcode: KU866572 (alternative markers: BenA = KU866812; CaM = KU866695; RPB2 = KU866949).

Aspergillus brasiliensis Varga et al., Int. J. Syst. Evol. Microbiol. 57: 57. 2007. [MB510581]. — Type: CBS 101740. Ex-type: CBS 101740 = IMI 381727 = IBT 101740. Infragen. class: subgen. Circumdati, sect. Nigri, ser. Nigri. Reproduction: protoheterothallic; MAT1-2-1 detected (de Vries et al. 2017). ITS barcode: FJ629321 (alternative markers: BenA = FJ629272; CaM = FN594543; RPB2 = EF661063).

Aspergillus brevijanus S.W. Peterson, Mycologia 100: 217. 2008. [MB506751]. Replaced synonym: Aspergillus janus var. brevis Raper & Thom Mycologia 36: 561. 1944. [MB351654]. — Type: IMI 16066. Ex-type: CBS 111.46 = NRRL 1935 = ATCC 16828 = CBS 119.45 = IMI 016066ii = IMI 16066 = NCTC 6971 = QM 7417 = WB 1935. Infragen. class: subgen. Circumdati, sect. Janorum, ser. Janorum. Reproduction: asexual. ITS barcode: EF669582 (alternative markers: BenA = EU014078; CaM = EF669540; RPB2 = EF669624).

Aspergillus brevipes G. Sm., Trans. Brit. Mycol. Soc. 35: 241. 1952. [MB292837]. — Type: IMI 51494. Ex-type: CBS 118.53 = NRRL 2439 = NRRL 4078 = NRRL 4772 = NRRL A-5521 = ATCC 16899 = IFO 5821 = IMI 51494 = LSHBBB 263 = LSHBSm 242 = QM 1948 = WB 4078 = WB 4224 = WB 4772. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Brevipedes. Reproduction: asexual. ITS barcode: EF669954 (alternative markers: BenA = EF669812; CaM = EF669881; RPB2 = EF669742).

Aspergillus brevistipitatus A. Nováková & Hubka, Fungal Diversity 64: 260. 2014. [MB803934]. — Type: PRM 860543. Ex-type: CBS 135454 = CCF 4149 = CMF ISB 2152 = NRRL 62500 = IFM 60858 = DTO 311-F5. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Neoglabri. Reproduction: protoheterothallic (Nováková et al. 2014). ITS barcode: HF937386 (alternative markers: BenA = HF933364; CaM = HF933388; RPB2 = HF937380).

Aspergillus bridgeri M. Chr., Mycologia 74: 210. 1982. [MB110494]. — Type: NY JB 26-1-2. Ex-type: CBS 350.81 = NRRL 13000 = IBT 13380 = ATCC 44562 = IMI 259098. Infragen. class: subgen. Circumdati, sect. Circumdati, ser. Sclerotiorum. Reproduction: asexual. ITS barcode: EF661404 (alternative markers: BenA = EF661335; CaM = EF661358; RPB2 = EF661290).

Aspergillus brunneouniseriatus Suj. Singh & B.K. Bakshi, Trans. Brit. Mycol. Soc. 44: 160. 1961. [MB326616]. — Type: IMI 227677. Ex-type: CBS 127.61 = NRRL 4273 = ATCC 16916 = IFO 6993 = IMI 227677 = QM 6990 = WB 4273. Infragen. class: subgen. Cremei, sect. Cremei, ser. Brunneouniseriati. Reproduction: asexual. ITS barcode: EF652141 (alternative markers: BenA = EF652123; CaM = EF652138; RPB2 = EF652089).

Aspergillus brunneoviolaceus Bat. & H. Maia, Anais Soc. Biol. Pernambuco 13: 91. 1955. [MB292838]. — Type: IMI 312981. Ex-type: CBS 621.78 = NRRL 4912 = IMI 312981 = WB 4912. Infragen. class: subgen. Circumdati, sect. Nigri, ser. Japonici. Reproduction: asexual. ITS barcode: AJ280003 (alternative markers: BenA = EF661105; CaM = EF661147; RPB2 = EF661045).

Aspergillus brunneus Delacr., Bull. Soc. Mycol. Fr. 9: 185. 1893. [MB204832]. — Type: IMI 211378. Ex-type: CBS 112.26 = CBS 524.65 = NRRL 131 = NRRL 134 = ATCC 1021 = IFO 5862 = IMI 211378 = QM 7406 = Thom 4481 = Thom 5633.4 = WB 131. Infragen. class: subgen. Aspergillus, sect. Aspergillus, ser. Aspergillus. Reproduction: homothallic (Delacroix 1893, Chen et al. 2017). ITS barcode: EF652060 (alternative markers: BenA = EF651907; CaM = EF651998; RPB2 = EF651939).

Aspergillus caatingaensis Y. Horie et al., Mycoscience 55: 84. 2014. [MB801323]. — Type: IFM 61335H. Ex-type: IFM 61335 = CBS 137446 = DTO 278-B3 = DTO 316-F8. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Unilaterales. Reproduction: homothallic. ITS barcode: MN431362 (alternative markers: BenA = AB743854; CaM = AB743860; RPB2 = MN969064).

Aspergillus caelatus B.W. Horn, Mycotaxon 61: 186. 1997. [MB436955]. — Type: BPI 737601. Ex-type: DTO 046-A8 = CBS 763.97 = NRRL 25528 = ATCC 201128. Infragen. class: subgen. Circumdati, sect. Flavi, ser. Kitamyces. Reproduction: protoheterothallic; both idiomorphs detected (Ramirez-Prado et al. 2008, unpublished data in GenBank). ITS barcode: AF004930 (alternative markers: BenA = EF661470; CaM = EF661522; RPB2 = EF661436).

Aspergillus caesiellus Saito, J. Coll. Sci. Imp. Univ. Tokyo 18: 49. 1904. [MB205025]. — Type: IMI 172278. Ex-type: CBS 470.65 = NRRL 5061 = ATCC 11905 = IFO 4882 = IMI 172278 = WB 5061. Infragen. class: subgen. Aspergillus, sect. Restricti, ser. Restricti. Reproduction: asexual. ITS barcode: EF652044 (alternative markers: BenA = EF651884; CaM = EF652030; RPB2 = EF651981).

Aspergillus caespitosus Raper & Thom, Mycologia 36: 563. 1944. [MB284298]. — Type: IMI 16034ii. Ex-type: CBS 103.45 = NRRL 1929 = ATCC 11256 = IMI 16034 = MUCL 13587 = NCTC 6972 = NCTC 6973 = QM 7399 = WB 1929. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Stellati. Reproduction: asexual. ITS barcode: EF652428 (alternative markers: BenA = EF652252; CaM = EF652340; RPB2 = EF652164).

Aspergillus calidoustus Varga et al., Eukaryot. Cell 7: 636. 2008. [MB504846]. — Type: Varga et al. 2008, Eukaryot. Cell 7: p. 636 Fig. 3 (– lectotype designated here, MBT392259; CBS H-24277 [dried culture] – epitype designated here, MBT392260). Ex-epitype: CBS 121601. Infragen. class: subgen. Nidulantes, sect. Usti, ser. Calidousti. Reproduction: asexual. ITS barcode: HE616558 (alternative markers: BenA = FJ624456; CaM = HE616559; RPB2 = MN969061).

Aspergillus californicus Frisvad et al., Stud. Mycol. 69: 91. 2011. [MB560400]. — Type: CBS H-20635. Ex-type: CBS 123895 = IBT 16748 = DTO 061-D4. Infragen. class: subgen. Nidulantes, sect. Cavernicolarum, ser. Cavernicolarum. Reproduction: asexual. ITS barcode: FJ531153 (alternative markers: BenA = FJ531180; CaM = FJ531128; RPB2 = MN969065).

Aspergillus campestris M. Chr., Mycologia 74: 212. 1982. [MB110495]. — Type: NY ST 2–3–1. Ex-type: CBS 348.81 = NRRL 13001 = ATCC 44563 = IMI 259099. Infragen. class: subgen. Circumdati, sect. Candidi, ser. Candidi. Reproduction: asexual. ITS barcode: EF669577 (alternative markers: BenA = EU014091; CaM = EF669535; RPB2 = EF669619).

Aspergillus canadensis Visagie et al., Stud. Mycol. 88: 187. 2017. [MB818935]. — Type: DAOM 740109. Ex-type: CCF 5548 = KAS 6194 = DTO 356-H9 = IBT 34520 = IBT 34642 = NRRL 66614. Infragen. class: subgen. Aspergillus, sect. Restricti, ser. Penicillioides. Reproduction: asexual. ITS barcode: KY087667 (alternative markers: BenA = KY117731; CaM = KY068215; RPB2 = KY117909).

Aspergillus candidus Link, Mag. Ges. Naturf. Freunde Berlin 3: 16. 1809. [MB204868]. — Type: CBS 566.65. Ex-type: CBS 566.65 = NRRL 303 = ATCC 1002 = IMI 16264 = IMI 91889 = LSHBA c .27 = NCTC 595 = QM 1995 = Thom 106 = WB 303. Infragen. class: subgen. Circumdati, sect. Candidi, ser. Candidi. Reproduction: asexual. ITS barcode: EF669592 (alternative markers: BenA = EU014089; CaM = EF669550; RPB2 = EF669634).

Aspergillus caninus (Sigler et al.) Houbraken et al., Stud. Mycol. 78: 154. 2014. [MB809580]. Basionym: Phialosimplex caninus Sigler et al., Med. Mycol. 48: 338. 2010. [MB513393]. — Type: UAMH 10337. Ex-type: CBS 128032 = UAMH 10337 = DTO 139-A6. Infragen. class: subgen. Polypaecilum, sect. Polypaecilum, ser. Canini. Reproduction: asexual. ITS barcode: KY980618 (alternative markers: BenA = KY980546; CaM = MN969225; RPB2 = JN121445).

Aspergillus capensis Visagie et al., Stud. Mycol. 78: 105. 2014. [MB809193]. — Type: CBS H-21810. Ex-type: CBS 138188 = DTO 179-E6. Infragen. class: subgen. Circumdati, sect. Flavipedes, ser. Flavipedes. Reproduction: asexual. ITS barcode: KJ775550 (alternative markers: BenA = KJ775072; CaM = KJ775279; RPB2 = KP987020).

Aspergillus caperatus A.J. Chen et al., Stud. Mycol. 88: 85. 2017. [MB818733]. — Type: CBS H-22825. Ex-type: CBS 141774 = DTO 337-E6 = IBT 34451. Infragen. class: subgen. Aspergillus, sect. Aspergillus, ser. Chevalierorum. Reproduction: homothallic. ITS barcode: LT670922 (alternative markers: BenA = LT671008; CaM = LT671009; RPB2 = LT671010).

Aspergillus carbonarius (Bainier) Thom, J. Agric. Res. 7: 12. 1916. [MB100545]. Basionym: Sterigmatocystis carbonaria Bainier, Bull. Soc. Bot. France 27: 27. 1880. [MB195901]. — Type: CBS 556.65. Ex-type: CBS 111.26 = NRRL 369 = ATCC 1025 = IMI 16136 = LSHBA c .11 = NCTC 1325 = NRRL 1987 = QM 331 = Thom 4030.1 = WB 369. Infragen. class: subgen. Circumdati, sect. Nigri, ser. Carbonarii. Reproduction: protoheterothallic; MAT1-2-1 detected (de Vries et al. 2017). ITS barcode: EF661204 (alternative markers: BenA = EF661099; CaM = EF661167; RPB2 = EF661068).

Aspergillus carlsbadensis Frisvad et al., Stud. Mycol. 69: 88. 2011. [MB560399]. — Type: CBS H-30634. Ex-type: CBS 123894 = IBT 14493 = DTO 061-C7. Infragen. class: subgen. Nidulantes, sect. Usti, ser. Calidousti. Reproduction: asexual. ITS barcode: FJ531151 (alternative markers: BenA = FJ531179; CaM = FJ531126; RPB2 = MN969066).

Aspergillus carneus Blochwitz, Ann. Mycol. 31: 81. 1933. [MB259903]. — Type: IMI 1358818. Ex-type: CBS 494.65 = NRRL 527 = ATCC 16798 = IMI 135818 = QM 7401 = Thom 5740.4 = WB 527. Infragen. class: subgen. Circumdati, sect. Terrei, ser. Nivei. Reproduction: asexual. ITS barcode: EF669611 (alternative markers: BenA = EF669529; CaM = EF669569; RPB2 = EF669653).

Aspergillus cavernicola Lörinczi, Contrtii bot. Univ. Babes-Bolyai, Cluj, Grad. bot.: 341. 1969. [MB326617]. — Type: CBS 117.76. Ex-type: CBS 117.76 = NRRL 6327. Infragen. class: subgen. Nidulantes, sect. Cavernicolarum, ser. Cavernicolarum. Reproduction: asexual. ITS barcode: EF652508 (alternative markers: BenA = EF652332; CaM = EF652420; RPB2 = EF652244).

Aspergillus cejpii (Milko) Samson et al., Stud. Mycol. 78: 155. 2014. [MB809582]. Basionym: Talaromyces cejpii Milko, Novosti Sist. Nizsh. Rast. 1: 208. 1964. [MB339918]. — Type: CBS H-7011. Ex-type: CBS 157.66. Infragen. class: subgen. Fumigati, sect. Vargarum, ser. Vargarum. Reproduction: homothallic. ITS barcode: MN431363 (alternative markers: BenA = EU076314; CaM = MN969226; RPB2 = JN121447).

Aspergillus cerealis Houbraken et al., Stud. Mycol. 93: 43. 2019. [MB823773]. — Type: CBS H-23359. Ex-type: CBS 143674 = DTO 228-E7 = IBT 32067. Infragen. class: subgen. Circumdati, sect. Flavi, ser. Flavi. Reproduction: protoheterothallic; both MAT idiomorphs detected (Carvajal-Campos et al. 2017; referred to as A. korhogoensis). ITS barcode: MG662394 (alternative markers: BenA = MG517693; CaM = MG518063; RPB2 = MG517884).

Aspergillus cervinus Massee, Bull. Misc. Inform. Kew 1914: 158. 1914. [MB211549]. — Type: WIS WISC WT 540. Ex-type: CBS 537.65 = NRRL 5025 = ATCC 16915 = IMI 126542 = QM 8875 = WB 5025. Infragen. class: subgen. Fumigati, sect. Cervini, ser. Cervini. Reproduction: asexual. ITS barcode: EF661268 (alternative markers: BenA = EF661251; CaM = EF661261; RPB2 = EF661229).

Aspergillus chaetosartoryae Hubka et al., this study. [MB832557]. Replaced synonym: Chaetosartorya stromatoides B.J. Wiley & E.G. Simmons, Mycologia 65: 935. 1973. [MB310956]. — Type: QM 8944. Ex-type: CBS 265.73 = ATCC 24480 = IMI 171880 = NRRL 5501. Infragen. class: subgen. Cremei, sect. Cremei, ser. Cremei. Reproduction: homothallic. ITS barcode: EF652144 (alternative markers: BenA = EF652117; CaM = EF652129; RPB2 = EF652099).

Aspergillus chevalieri (L. Mangin) Thom & Church, Aspergilli: 111. 1926. [MB292839]. Basionym: Eurotium chevalieri L. Mangin, Ann. Sci. Nat., Bot., ser. 9, 10: 361. 1909. [MB238304]. — Type: IMI 211382. Ex-type: CBS 522.65 = NRRL 78 = ATCC 16443 = IMI 211382 = NRRL A-7803 = Thom 4125.3 = WB 78. Infragen. class: subgen. Aspergillus, sect. Aspergillus, ser. Chevalierorum. Reproduction: homothallic. ITS barcode: EF652068 (alternative markers: BenA = EF651911; CaM = EF652002; RPB2 = EF651954).

Aspergillus chlamydosporus (Gené & Guarro) Houbraken et al., Stud. Mycol. 78: 155. 2014. [MB809584]. Basionym: Sagenomella chlamydospora Gené & Guarro, J. Clin. Microbiol. 41: 1723. 2003. [MB488173]. — Type: IMI 387422. Ex-type: CBS 109945 = IMI 387422 = FMR 7371 = DTO 138-C2 = UAMH 10961. Infragen. class: subgen. Polypaecilum, sect. Polypaecilum, ser. Canini. Reproduction: asexual. ITS barcode: KY980617 (alternative markers: BenA = KY980545; CaM = MN969227; RPB2 = JN121425).

Aspergillus christenseniae A.J. Chen et al., Stud. Mycol. 85: 75. 2016. [MB817724]. — Type: CBS H-9217. Ex-type: CBS 122.56 = DTO 022-C8 = IBT 22043 = IBT 23735 = IMI 343732 = NRRL 4897 = WB 4897. Infragen. class: subgen. Fumigati, sect. Cervini, ser. Cervini. Reproduction: asexual. ITS barcode: FJ491613 (alternative markers: BenA = FJ491639; CaM = FJ491608; RPB2 = EF661235).

Aspergillus chrysellus Kwon-Chung & Fennell, Gen. Aspergillus: 424. 1965. [MB326618]. — Type: IMI 238612. Ex-type: CBS 472.65 = NRRL 5084 = ATCC 16852 = IMI 238612 = IMI 238612ii = QM 8876 = WB 5084. Infragen. class: subgen. Cremei, sect. Cremei, ser. Wentiorum. Reproduction: homothallic. ITS barcode: EF652155 (alternative markers: BenA = EF652109; CaM = EF652136; RPB2 = EF652090).

Aspergillus cibarius S.B. Hong & Samson, J. Microbiol 50: 713. 2012. [MB800861]. — Type: KACC 46346. Ex-type: DTO 197-D3 = KACC 46346. Infragen. class: subgen. Aspergillus, sect. Aspergillus, ser. Aspergillus. Reproduction: homothallic. ITS barcode: JQ918177 (alternative markers: BenA = JQ918180; CaM = JQ918183; RPB2 = JQ918186).

Aspergillus citocrescens Hubka et al., Persoonia 35: 311. 2015. [MB814680]. — Type: PRM 934413. Ex-type: CCF 4001 = CBS 140566 = DTO 376-B3. Infragen. class: subgen. Cremei, sect. Cremei, ser. Brunneouniseriati. Reproduction: asexual. ITS barcode: FR727121 (alternative markers: BenA = FR775317; CaM = LN878969; RPB2 = MN969163).

Aspergillus citrinoterreus J. Guinea et al., J. Clin. Microbiol. 53: 612. 2015. [MB810584]. — Type: CBS H-22005. Ex-type: CBS 138921 = GM 228 = DTO 331-H6. Infragen. class: subgen. Circumdati, sect. Terrei, ser. Terrei. Reproduction: asexual. ITS barcode: KP175260 (alternative markers: BenA = LN680657; CaM = LN680685; RPB2 = MN969155).

Aspergillus clavatonanicus Bat. et al., Anais Fac. Med. Univ. Recife 15: 197. 1955. [MB292840]. — Type: IMI 235352. Ex-type: CBS 474.65 = NRRL 4741 = ATCC 12413 = DMUR 532 = IMI 235352 = JCM 10183 = QM 7059 = WB 4741. Infragen. class: subgen. Fumigati, sect. Clavati, ser. Clavati. Reproduction: asexual. ITS barcode: EF669986 (alternative markers: BenA = EF669842; CaM = EF669912; RPB2 = EF669773).

Aspergillus clavatophorus F. Sklenář et al., Stud. Mycol. 88: 187. 2017. [MB818936]. — Type: PRM 944440. Ex-type: NRRL 25874 = CCF 5454 = IBT 34560 = IBT 34823 = DTO 356-D8. Infragen. class: subgen. Aspergillus, sect. Restricti, ser. Penicillioides. Reproduction: asexual. ITS barcode: KY087772 (alternative markers: BenA = KY117836; CaM = KY068323; RPB2 = KY118014).

Aspergillus clavatus Desm., Ann. Sci. Nat., Bot., ser. 2, 2: 71. 1834. [MB211530]. — Type: IMI 15949. Ex-type: CBS 513.65 = NRRL 1 = ATCC 1007 = ATCC 9598 = ATCC 9602 = CECT2674 = DSM 816 = IMI 15949 = LSHBA c .86 = LSHBA c .95 = MIT213 = NCTC 3887 = NCTC 9 = NCTC 978 = NRRL 1656 = QM 1276 = QM 7404 = Thom 107 = WB 1. Infragen. class: subgen. Fumigati, sect. Clavati, ser. Clavati. Reproduction: Heterothallic (Ojeda-López et al. 2018). ITS barcode: EF669942 (alternative markers: BenA = EF669802; CaM = EF669871; RPB2 = EF669730).

Aspergillus collembolorum Dörfelt & A.R. Schmidt, Mycol. Res. 109: 956, Figs 1–9. 2005. [MB344420]. — Type: Russia: Kaliningrad (Koenigsberg), in succinum Balticum, in exemplare subordines Entomobryomorpha (Collembola), C. & H. W. Hoffeins (coll. Hoffeins, Hamburg, no. 805, holotypus). Ex-type: n.a. Infragen. class: subgen.: unknown, sect.: unknown, ser.: unknown. Reproduction: asexual. ITS barcode: n.a. (alternative markers: BenA = n.a.; CaM = n.a.; RPB2 = n.a.).

Aspergillus collinsii Jurjević & S.W. Peterson, Int. J. Syst. Evol. Microbiol. 66: 2570. 2016. [MB814413]. — Type: BPI 893219. Ex-type: CBS 140843 = NRRL 66196 = CCF 5175. Infragen. class: subgen. Nidulantes, sect. Usti, ser. Deflecti. Reproduction: asexual. ITS barcode: KT698845 (alternative markers: BenA = KT698843; CaM = KT698844; RPB2 = KT698848).

Aspergillus coloradensis F. Sklenář et al., Mycologia 112: 357. 2020. [MB832715]. — Type: PRM 951699. Ex-type: CCF 6118 = EMSL No. 2726 = NRRL 66888. Infragen. class: subgen. Nidulantes, sect. Aenei, ser. Aenei. Reproduction: homothallic. ITS barcode: MK713539 (alternative markers: BenA = MK695646; CaM = MK695657; RPB2 = MK695668).

Aspergillus conicus Blochwitz, Ann. Mycol. 12: 38. 1914. [MB120214]. — Type: IMI 172281. Ex-type: CBS 475.65 = NRRL 149 = ATCC 16908 = IMI 172281 = QM 7405 = Thom 4733.701 = WB 149. Infragen. class: subgen. Aspergillus, sect. Restricti, ser. Restricti. Reproduction: asexual. ITS barcode: EF652039 (alternative markers: BenA = EF651881; CaM = EF652033; RPB2 = EF651975).

Aspergillus conjunctus Kwon-Chung & Fennell, Gen. Aspergillus: 552. 1965. [MB326620]. — Type: IMI 135421. Ex-type: CBS 476.65 = NRRL 5080 = ATCC 16796 = IMI 135421 = QM 8878 = WB 5080. Infragen. class: subgen. Nidulantes, sect. Sparsi, ser. Conjuncti. Reproduction: asexual. ITS barcode: EF661179 (alternative markers: BenA = EF661111; CaM = EF661133; RPB2 = EF661042).

Aspergillus contaminans Hubka et al., Persoonia 39: 285. 2017. [MB821684]. — Type: PRM 944503. Ex-type: CCF 4682 = CBS 142451 = NRRL 66666. Infragen. class: subgen. Nidulantes, sect. Usti, ser. Calidousti. Reproduction: asexual. ITS barcode: LT594451 (alternative markers: BenA = LT594443; CaM = LT594425; RPB2 = LT594434).

Aspergillus conversis Hubka & A. Nováková, Fungal Diversity 64: 262. 2014. [MB803935]. — Type: PRM 860541. Ex-type: CBS 135457 = NRRL 62496 = CCF 4190 = CMF ISB 2151 = IFM 60857. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Neoglabri. Reproduction: protoheterothallic; MAT 1-2-1 detected (Nováková et al. 2014). ITS barcode: HF937385 (alternative markers: BenA = HF933363; CaM = HF933387; RPB2 = HF937379).

Aspergillus coremiiformis Bartoli & Maggi, Trans. Brit. Mycol. Soc. 71: 386. 1979 [1978]. [MB309214]. — Type: RO 102 S. Ex-type: CBS 553.77 = NRRL 13603 = ATCC 38576 = IMI 223069 = NRRL 13756. Infragen. class: subgen. Circumdati, sect. Flavi, ser. Coremiiformes. Reproduction: asexual. ITS barcode: EF661544 (alternative markers: BenA = EU014104; CaM = EU014112; RPB2 = EU021623).

Aspergillus corrugatus Udagawa & Y. Horie, Mycotaxon 4: 535. 1976. [MB309216]. — Type: NHL 2763. Ex-type: CBS 191.77 = NHL 2763 = DTO 047-I9 = CBM-FA-73. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Nidulantes. Reproduction: homothallic. ITS barcode: KU866574 (alternative markers: BenA = KU866814; CaM = MN969228; RPB2 = KU866951).

Aspergillus costaricensis [as “costaricaensis”] Samson & Frisvad, Stud. Mycol. 50: 52. 2004. [MB369151]. — Type: CBS H-13437. Ex-type: CBS 115574 = IBT 23401 = CECT 20579 = ITEM 7555. Infragen. class: subgen. Circumdati, sect. Nigri, ser. Nigri. Reproduction: protoheterothallic; unpublished (genome data; Vesth et al. 2018). ITS barcode: DQ900602 (alternative markers: BenA = FJ629277; CaM = FN594545; RPB2 = HE984361).

Aspergillus costiformis H.Z. Kong & Z.T. Qi, Acta Mycol. Sin. 14: 10. 1995 [MB363444]. — Type: HMAS 62766. Ex-type: CBS 101749 = AS 3.4664. Infragen. class: subgen. Aspergillus, sect. Aspergillus, ser. Chevalierorum. Reproduction: homothallic. ITS barcode: HE615136 (alternative markers: BenA = HE801338; CaM = HE801320; RPB2 = HE801309).

Aspergillus crassihyphae Wadhwani & N. Mehrotra, Indian Bot. Reporter: 52. 1985. [MB105070]. — Type: unknown. Ex-type: unknown. Infragen. class: subgen. Circumdati, sect. Nigri, ser.: unknown. Reproduction: asexual. ITS barcode: n.a. (alternative markers: BenA = n.a.; CaM = n.a.; RPB2 = n.a.).

Aspergillus creber Jurjević, S.W. Peterson & B.W. Horn, IMA Fungus 3: 69. 2012. [MB800598]. — Type: BPI 800912. Ex-type: CBS 145749 = NRRL 58592 = DTO 225-G7. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Versicolores. Reproduction: asexual. ITS barcode: JQ301889 (alternative markers: BenA = JN853980; CaM = JN854043; RPB2 = JN853832).

Aspergillus cremeus Kwon-Chung & Fennell, Gen. Aspergillus: 418. 1965. [MB326621]. — Type: IMI 123749ii. Ex-type: CBS 477.65 = NRRL 5081 = ATCC 16857 = IMI 123749 = QM 8879 = QM 9191 = WB 5081. Infragen. class: subgen. Cremei, sect. Cremei, ser. Cremei. Reproduction: homothallic. ITS barcode: EF652149 (alternative markers: BenA = EF652120; CaM = EF652125; RPB2 = EF652101).

Aspergillus cretensis Frisvad & Samson, Stud. Mycol. 50: 33. 2004. [MB500002]. — Type: CBS H-13446. Ex-type: CBS 112802 = NRRL 35672 = IBT 17505. Infragen. class: subgen. Circumdati, sect. Circumdati, ser. Circumdati. Reproduction: asexual. ITS barcode: FJ491572 (alternative markers: BenA = EF661332; CaM = FJ491534; RPB2 = EF661311).

Aspergillus cristatus Raper & Fennell, Gen. Aspergillus: 169. 1965. [MB326622]. — Type: IMI 172280. Ex-type: CBS 123.53 = NRRL 4222 = ATCC 16468 = IMI 172280 = MUCL 15644 = WB 4222. Infragen. class: subgen. Aspergillus, sect. Aspergillus, ser. Chevalierorum. Reproduction: homothallic. ITS barcode: EF652078 (alternative markers: BenA = EF651914; CaM = EF652001; RPB2 = EF651957).

Aspergillus croceiaffinis F. Sklenář et al., Mycologia 112: 359. 2020. [MB832713]. — Type: PRM 951576. Ex-type: CCF 6035 = EMSL No. 2282 = NRRL 66887. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Unguium. Reproduction: asexual. ITS barcode: MK713538 (alternative markers: BenA = MK695645; CaM = MK695656; RPB2 = MK695667).

Aspergillus croceus Hubka et al., Plant Syst. Evol. 302: 1291. 2016. [MB816281]. — Type: PRM 924053. Ex-type: CCF 4405 = CBS 134396 = NRRL 62495 = IBT 33602. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Unguium. Reproduction: asexual. ITS barcode: LN873931 (alternative markers: BenA = LN873944; CaM = LN873957; RPB2 = LN873976).

Aspergillus crustosus Raper & Fennell, Gen. Aspergillus: 532. 1965. [MB326623]. — Type: IMI 135819. Ex-type: CBS 478.65 = NRRL 4988 = ATCC 16806 = IMI 135819 = NRRL A-3254 = QM 8910 = WB 4988. Infragen. class: subgen. Nidulantes, sect. Aenei, ser. Aenei. Reproduction: asexual. ITS barcode: EF652489 (alternative markers: BenA = EF652313; CaM = EF652401; RPB2 = EF652225).

Aspergillus cumulatus D.H. Kim & S.B. Hong, J. Microbiol. Biotechnol. 24: 335. 2014. [MB807118]. — Type: KACC 47316. Ex-type: KACC 47316. Infragen. class: subgen. Aspergillus, sect. Aspergillus, ser. Rubri. Reproduction: homothallic. ITS barcode: KF928303 (alternative markers: BenA = KF928297; CaM = KF928300; RPB2 = KF928294).

Aspergillus curviformis H.J. Chowdhery & J.N. Rai, Nova Hedwigia 32: 231. 1980. [MB118396]. — Type: unknown. Ex-type: unknown. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Viridinutantes. Reproduction: asexual. ITS barcode: n.a. (alternative markers: BenA = n.a.; CaM = n.a.; RPB2 = n.a.).

Aspergillus cvjetkovicii Jurjević et al., IMA Fungus 3: 69. 2012. [MB800599]. — Type: BPI 880909. Ex-type: NRRL 227 = CBS 599.65. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Versicolores. Reproduction: asexual. ITS barcode: EF652440 (alternative markers: BenA = EF652264; CaM = EF652352; RPB2 = EF652176).

Aspergillus deflectus Fennell & Raper, Mycologia 47: 83. 1955. [MB292841]. — Type: IMI 61448. Ex-type: CBS 109.55 = NRRL 2206 = ATCC 16807 = IMI 61448 = NRRL A-2700A = QM 1904 = UC4638 = WB 2206. Infragen. class: subgen. Nidulantes, sect. Usti, ser. Deflecti. Reproduction: asexual. ITS barcode: EF652437 (alternative markers: BenA = EF652261; CaM = EF652349; RPB2 = EF652173).

Aspergillus delicatus H.Z. Kong, Mycotaxon 62: 429. 1997. [MB437509]. — Type: HMAS 71159. Ex-type: CBS 101754 = AS 3.4697 = DTO 050-E7. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Thermomutati. Reproduction: homothallic. ITS barcode: MN431364 (alternative markers: BenA = DQ114124; CaM = DQ114132; RPB2 = MN969067).

Aspergillus denticulatus (Samson et al.) Samson et al., Stud. Mycol. 78: 155. 2014. [MB809586]. Basionym: Neosartorya denticulata Samson et al., Antonie van Leeuwenhoek 93: 95. 2008. [MB506375]. — Type: CBS 652.73. Ex-type: CBS 652.73 = KACC 41183 = DTO 050-D8 = DTO 026-G9. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Fennelliarum. Reproduction: homothallic. ITS barcode: MN431365 (alternative markers: BenA = DQ114125; CaM = DQ114133; RPB2 = MN969068).

Aspergillus desertorum (Samson & Mouch.) Samson et al., Stud. Mycol. 78: 155. 2014. [MB809587]. Basionym: Emericella desertorum Samson & Mouch., Antonie van Leeuwenhoek 40: 121. 1974. [MB313502]. — Type: CBS H-7045. Ex-type: CBS 653.73 = NRRL 5921 = IMI 343076. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Nidulantes. Reproduction: homothallic; asexual morph unknown. ITS barcode: EF652505 (alternative markers: BenA = EF652329; CaM = EF652417; RPB2 = EF652241).

Aspergillus destruens Zalar et al., Stud. Mycol. 88: 191. 2017. [MB818930]. — Type: PRM 944428. Ex-type: NRRL 145 = IMI 358691 = CCF 5462 = CBS 593.91 = DTO 079-A8 = IBT 34818. Infragen. class: subgen. Aspergillus, sect. Restricti, ser. Restricti. Reproduction: asexual. ITS barcode: KY087748 (alternative markers: BenA = KY117811; CaM = KY068298; RPB2 = KY117989).

Aspergillus dimorphicus B.S. Mehrotra & R. Prasad, Trans. Brit. Mycol. Soc. 52: 331. 1969. [MB326625]. — Type: IMI 131553. Ex-type: CBS 649.74 = NRRL 3650 = IMI 131553 = QM 9190. Infragen. class: subgen. Cremei, sect. Cremei, ser. Wentiorum. Reproduction: asexual. ITS barcode: EF652154 (alternative markers: BenA = EF652111; CaM = EF652135; RPB2 = EF652096).

Aspergillus dipodomyus F. Sklenář et al., Mycologia 112: 360. 2020. [MB832710]. — Type: PRM 951565. Ex-type: CCF 5265 = NRRL 66273. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Nidulantes. Reproduction: homothallic. ITS barcode: MK713535 (alternative markers: BenA = MK695642; CaM = MK695653; RPB2 = MK695664).

Aspergillus discophorus Samson et al., Mycologia 100: 787. 2008. [MB507360]. — Type: CBS H-19889. Ex-type: CBS 469.88 = IBT 21910 = IMI 328717 = DTO 011-B1. Infragen. class: subgen. Nidulantes, sect. Aenei, ser. Aenei. Reproduction: homothallic. ITS barcode: EU448272 (alternative markers: BenA = AY339999; CaM = EU443970; RPB2 = MN969069).

Aspergillus diversus Raper & Fennell, Gen. Aspergillus: 437. 1965. [MB326626]. — Type: IMI 232882. Ex-type: CBS 480.65 = NRRL 5074 = ATCC 16849 = IMI 232882 = QM 8882 = WB 5074. Infragen. class: subgen. Nidulantes, sect. Sparsi, ser. Biplani. Reproduction: asexual. ITS barcode: EF661213 (alternative markers: BenA = EF661114; CaM = EF661128; RPB2 = EF661034).

Aspergillus dobrogensis A. Nováková et al., Int. J. Syst. Evol. Microbiol. 68: 1004. 2018. [MB821313]. — Type: PRM 935751. Ex-type: CCF 4651 =CCF 4655 = NRRL 62821 = IBT 32697 = CBS 143370. Infragen. class: subgen. Circumdati, sect. Candidi, ser. Candidi. Reproduction: asexual. ITS barcode: LT626959 (alternative markers: BenA = LT627027; CaM = LT558722; RPB2 = LT627028).

Aspergillus domesticus F. Sklenář et al., Stud. Mycol. 88: 194. 2017. [MB818931]. — Type: PRM 944426. Ex-type: DTO 079-F2 = CCF 5464 = NRRL 66616 = IBT 34814. Infragen. class: subgen. Aspergillus, sect. Restricti, ser. Restricti. Reproduction: asexual. ITS barcode: KY087688 (alternative markers: BenA = KY117752; CaM = KY068236; RPB2 = KY117928).

Aspergillus dromiae A.J. Chen et al., Stud. Mycol. 84: 57. 2016. [MB816089]. — Type: CBS H-22489. Ex-type: CBS 140633 = IBT 25166 = DTO 059-H5. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Stellati. Reproduction: homothallic. ITS barcode: KU866580 (alternative markers: BenA = KU866885; CaM = KU866703; RPB2 = KU866958).

Aspergillus duricaulis Raper & Fennell, Gen. Aspergillus: 249. 1965. [MB326627]. — Type: IMI 172282. Ex-type: CBS 481.65 = NRRL 4021 = ATCC 16900 = IMI 172282 = IMI 367413 = NRRL A-5509 = QM 8884 = WB 4021. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Brevipedes. Reproduction: asexual. ITS barcode: EF669971 (alternative markers: BenA = EF669827; CaM = EF669897; RPB2 = EF669758).

Aspergillus eburneocremeus Sappa, Allionia 2: 87. 1954. [MB292842]. — Type: TMI 69856. Ex-type: CBS 130.54 = NRRL 4773 = ATCC 16802 = IMI 69856 = MUCL 13588 = QM 1949 = WB 4773. Infragen. class: subgen. Nidulantes, sect. Aenei, ser. Aenei. Reproduction: asexual. ITS barcode: EF652476 (alternative markers: BenA = EF652300; CaM = EF652388; RPB2 = EF652212).

Aspergillus egyptiacus Moub. & Mustafa, Egypt. J. Bot. 15: 153. 1972. [MB344341]. — Type: IMI 141415. Ex-type: CBS 656.73 = NRRL 5920 = ATCC 32114 = IMI 141415. Infragen. class: subgen. Nidulantes, sect. Cavernicolarum, ser. Egyptiaci. Reproduction: asexual. ITS barcode: EF652504 (alternative markers: BenA = EF652328; CaM = EF652416; RPB2 = EF652240).

Aspergillus elegans Gasperini, Atti Soc. Tosc. Sci. Nat. Pisa Mem. 8: 328. 1887. [MB212852]. — Type: CBS 102.14. Ex-type: CBS 102.14 = CBS 543.65 = NRRL 4850 = IBT 13505 = ATCC 13829 = ATCC 16886 = IFO 4286 = IMI 133962 = QM 8912 = QM 9373 = WB 4850. Infragen. class: subgen. Circumdati, sect. Circumdati, ser. Steyniorum. Reproduction: asexual. ITS barcode: EF661414 (alternative markers: BenA = EF661349; CaM = EF661390; RPB2 = EF661316).

Aspergillus ellipsoideus J.N. Rai & H.J. Chowdhery, Kavaka 7: 17. 1979 [MB116064]. — Type: MLLU 107. Ex-type: unknown. Infragen. class: subgen. Circumdati, sect. Nigri, ser.: unknown. Reproduction: asexual. ITS barcode: n.a. (alternative markers: BenA = n.a.; CaM = n.a.; RPB2 = n.a.).

Aspergillus ellipticus Raper & Fennell, Gen. Aspergillus: 319. 1965. [MB326628]. — Type: CBS 707.79. Ex-type: CBS 482.65 = CBS 707.79 = DTO 035-B7 = NRRL 5120 = ATCC 16876 = IMI 172283 = NRRL 20624 = QM 8886 = WB 5120. Infragen. class: subgen. Circumdati, sect. Nigri, ser. Heteromorphi. Reproduction: asexual. ITS barcode: EF661194 (alternative markers: BenA = AY585530; CaM = EF661170; RPB2 = EF661051).

Aspergillus elongatus J.N. Rai & S.C. Agarwal, Canad. J. Bot. 48: 791. 1970. [MB309217]. — Type: CBS 387.75. Ex-type: CBS 387.75 = NRRL 5176 = QM 9702 = WB 5495. Infragen. class: subgen. Nidulantes, sect. Usti, ser. Deflecti. Reproduction: asexual. ITS barcode: EF652502 (alternative markers: BenA = EF652326; CaM = EF652414; RPB2 = EF652238).

Aspergillus elsenburgensis Visagie, Stud. Mycol., this issue. 2020. [MB834199]. — Type: PREM 62313. Ex-type: PPRI 2994 = CMV 011G4. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Neoglabri. Reproduction: homothallic. ITS barcode: MK450651 (alternative markers: BenA = MK451215; CaM = MK451513; RPB2 = MK450804).

Aspergillus endophyticus Hubka et al., Stud. Mycol. 88: 95. 2017. [MB818734]. — Type: CBS H-22819. Ex-type: CBS 141766 = DTO 354-I2 = CCF 5345 = IBT 34511. Infragen. class: subgen. Aspergillus, sect. Aspergillus, ser. Aspergillus. Reproduction: homothallic. ITS barcode: LT670941 (alternative markers: BenA = LT671067; CaM = LT671068; RPB2 = LT671069).

Aspergillus eucalypticola Varga et al., Stud. Mycol. 69: 9. 2011. [MB560387]. — Type: CBS H-20627. Ex-type: CBS 122712 = IBT 29274. Infragen. class: subgen. Circumdati, sect. Nigri, ser. Nigri. Reproduction: protoheterothallic (genome data, Vesth et al. 2018). ITS barcode: EU482439 (alternative markers: BenA = EU482435; CaM = EU482433; RPB2 = MN969070).

Aspergillus europaeus Hubka et al., Plant Syst. Evol. 302: 645. 2016. [MB815574]. — Type: PRM 933832. Ex-type: CCF 4409 = CBS 134393 = IBT 32228 = NRRL 66252. Infragen. class: subgen. Cremei, sect. Cremei, ser. Wentiorum. Reproduction: asexual. ITS barcode: LN908996 (alternative markers: BenA = LN909006; CaM = LN909007; RPB2 = LT548274).

Aspergillus falconensis Y. Horie et al., Trans. Mycol. Soc. Japan 30: 257. 1989. [MB127891]. — Type: CBM 10001. Ex-type: CBS 271.91 = IFM 4997 = NHL 2999 = ATCC 76117 = DTO 048-A2. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Nidulantes. Reproduction: homothallic. ITS barcode: KU866575 (alternative markers: BenA = KU866815; CaM = KU866697; RPB2 = KU866952).

Aspergillus felis Barrs et al., PLoS ONE 8: e64871, 8. 2013. [MB560382]. — Type: CBS H-21125. Ex-type: CBS 130245 = DTO 131-F4 = CCF 5620. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Viridinutantes. Reproduction: Heterothallic. ITS barcode: MN431358 (alternative markers: BenA = MN969363; CaM = JX021715; RPB2 = MN969062).

Aspergillus fennelliae Kwon-Chung & S.J. Kim, Mycologia 66: 629. 1974. [MB309218]. — Type: IMI 278382. Ex-type: AF4 = CBS 599.74 = NRRL 5535 = ATCC 24326 = KACC 41150 (A); AF5 = CBS 598.74 = DTO 046-E8 = NRRL 5534 = ATCC 24325 = KACC 41125 (a). Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Fennelliarum. Reproduction: Heterothallic. ITS barcode: EF669994 (alternative markers: BenA = AF057320; CaM = EF669920; RPB2 = EF669781).

Aspergillus filifer [as “filifera”] Zalar et al., Mycologia 100: 787. 2008. [MB540309]. — Type: CBS H-19886. Ex-type: CBS 113636 = IBT 23443 = DTO 011-A5. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Stellati. Reproduction: homothallic. ITS barcode: EU448277 (alternative markers: BenA = EF428372; CaM = EU443973; RPB2 = KU866932).

Aspergillus fischeri Wehmer, Zentralbl. Bakteriol. Parasitenk, Abt. 2, 18: 390. 1907. [MB202877]. — Type: IMI 21139ii. Ex-type: CBS 544.65 = NRRL 181 = ATCC 1020 = DSM 3700 = IMI 211391 = QM 1983 = Thom 4651.2 = WB 181. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Fumigati. Reproduction: homothallic. ITS barcode: EF669936 (alternative markers: BenA = EF669796; CaM = EF669865; RPB2 = EF669724).

Aspergillus flaschentraegeri Stolk, Trans. Brit. Mycol. Soc. 47: 123. 1964. [MB326629]. — Type: CBS 108.63. Ex-type: CBS 108.63 = NRRL 5042 = ATCC 15535 = IMI 101651 = QM 8889 = WB 5042. Infragen. class: subgen. Cremei, sect. Cremei, ser. Wentiorum. Reproduction: asexual. ITS barcode: EF652150 (alternative markers: BenA = EF652113; CaM = EF652130; RPB2 = EF652102).

Aspergillus flavipes (Bainier & Sartory) Thom & Church, Aspergilli: 155. 1926. [MB265045]. Basionym: Sterigmatocystis flavipes Bainier & Sartory, Bull. Soc. Mycol. France 27: 90. 1911. [MB452855]. — Type: IMI 171885. Ex-type: NRRL 302 = ATCC 24487 = IMI 171885 = QM 9566 = Thom 4640.474 = WB 302. Infragen. class: subgen. Circumdati, sect. Flavipedes, ser. Flavipedes. Reproduction: asexual; the putative sexual morph of A. flavipes, Fennellia flavipes Wiley & Simmons, represents a different species - A. neoflavipes (Hubka et al. 2015). ITS barcode: EF669591 (alternative markers: BenA = EU014085; CaM = EF669549; RPB2 = EF669633).

Aspergillus flavus Link, Mag. Ges. Naturf. Freunde Berlin 3: 16. 1809. [MB209842]. — Type: IMI 124930. Ex-type: CBS 569.65 = NRRL 1957 = ATCC 16883 = IMI 124930 = QM 9947 = WB 1957. Infragen. class: subgen. Circumdati, sect. Flavi, ser. Flavi. Reproduction: Heterothallic (Horn et al. 2009a). ITS barcode: AF027863 (alternative markers: BenA = EF661485; CaM = EF661508; RPB2 = EF661440).

Aspergillus floccosus (Y.K. Shih) Samson et al., Stud. Mycol. 69: 45. 2011. [MB560393]. Basionym: Aspergillus terreus var. floccosus Y.K. Shih, Lingnan Sci. J. 15: 372. 1936. [MB499550]. — Type: Unknown. Ex-type: CBS 116.37 = CBS H-24278 = IBT 10846 = IBT 22556 = WB 4872 = DTO 067-B7. Infragen. class: subgen. Circumdati, sect. Terrei, ser. Terrei. Reproduction: asexual. ITS barcode: KP987086 (alternative markers: BenA = FJ491714; CaM = KP987066; RPB2 = KP987021).

Aspergillus floridensis Jurjević et al., IMA Fungus 3: 169. 2012. [MB802363]. — Type: BPI 883907. Ex-type: DTO 198-A8 = NRRL 62478 = ITEM 14783. Infragen. class: subgen. Circumdati, sect. Nigri, ser. Japonici. Reproduction: asexual. ITS barcode: MN431366 (alternative markers: BenA = HE984412; CaM = HE984429; RPB2 = HE984376).

Aspergillus foeniculicola Udagawa, Trans. Mycol. Soc. Japan 20: 13. 1979. [MB309220]. — Type: NHL 2777. Ex-type: CBS 156.80 = ATCC 42155 = IMI 334933 = LCP 84.2560 = NHL 2777. Infragen. class: subgen. Nidulantes, sect. Aenei, ser. Aenei. Reproduction: homothallic. ITS barcode: EU448274 (alternative markers: BenA = EU443990; CaM = EU443968; RPB2 = MN969071).

Aspergillus foveolatus Y. Horie, Trans. Mycol. Soc. Japan 19: 313. 1978. [MB309221]. — Type: IFM 4547. Ex-type: CBS 279.81 = IFM 4547 = NHL 2839 = NBRC 30559 = IFO 30559 = DTO 320-D2. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Nidulantes. Reproduction: homothallic. ITS barcode: KX423658 (alternative markers: BenA = KX423622; CaM = MN969229; RPB2 = KU867034).

Aspergillus frankstonensis Barrs et al., PloS ONE 12: e0181660, 8. 2017. [MB819986]. — Type: CBS-H-22969. Ex-type: CBS 142233 = DTO 341-E7. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Viridinutantes. Reproduction: protoheterothallic; MAT1-2-1 detected (Talbot et al. 2017). ITS barcode: KY808756 (alternative markers: BenA = KY808594; CaM = KY808724; RPB2 = KY808948).

Aspergillus fresenii Subram., Hyphomycetes: 552. 1971. [MB309222]. Replaced synonym: Sterigmatocystis sulphurea Fresen., Beitr. Mykol. 3: 83. 1863. [MB231754]. — Type: IMI 211397. Ex-type: CBS 550.65 = NRRL 4077 = ATCC 16893 = IMI 211397 = NRRL A-5355 = NRRL A-5520 = WB 4077. Infragen. class: subgen. Circumdati, sect. Circumdati, ser. Sclerotiorum. Reproduction: asexual. ITS barcode: EF661409 (alternative markers: BenA = EF661341; CaM = EF661382; RPB2 = EF661296).

Aspergillus fructus Jurjević et al., IMA Fungus 3: 70. 2012. [MB800600]. — Type: BPI 880915. Ex-type: NRRL 239 = CBS 584.65. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Versicolores. Reproduction: asexual. ITS barcode: EF652449 (alternative markers: BenA = EF652273; CaM = EF652361; RPB2 = EF652185).

Aspergillus fruticulosus Raper & Fennell, Gen. Aspergillus: 506. 1965. [MB326630]. — Type: IMI 139279. Ex-type: CBS 486.65 = NRRL 4903 = ATCC 16823 = IMI 139279 = O-1077 = QM 8033 = WB 4903. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Nidulantes. Reproduction: homothallic. ITS barcode: EF652483 (alternative markers: BenA = EF652307; CaM = EF652395; RPB2 = EF652219).

Aspergillus fumigatiaffinis S.B. Hong et al., Mycologia 97: 1326. 2006. [MB500296]. — Type: CBS 117186. Ex-type: CBS 117186 = KACC 41148 = IBT 12703 = IFM 55214. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Fumigati. Reproduction: protoheterothallic; both MAT idiomorphs detected (Dudová 2014). ITS barcode: MN431367 (alternative markers: BenA = DQ094885; CaM = DQ094891; RPB2 = MN969072).

Aspergillus fumigatus Fresen., Beitr. Mykol. 3: 81. 1863. [MB211776]. — Type: IMI 16152. Ex-type: CBS 133.61 = NRRL 163 = ATCC 1022 = ATCC 4813 = IMI 16152 = LSHBA c .71 = NCTC 982 = QM 1981 = Thom 118 = WB 163 = DTO 001-D1. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Fumigati. Reproduction: Heterothallic (O’Gorman et al. 2009). ITS barcode: EF669931 (alternative markers: BenA = EF669791; CaM = EF669860; RPB2 = EF669719).

Aspergillus fumisynnematus Y. Horie et al., Trans. Mycol. Soc. Japan 34: 3. 1993. [MB360061]. — Type: CBM FD-0001. Ex-type: DTO 354-A5 = CBS 141446 = IFM 42277. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Fumigati. Reproduction: protoheterothallic (unpublished data). ITS barcode: AB250779 (alternative markers: BenA = AB248076; CaM = AB259968; RPB2 = MN969073).

Aspergillus funiculosus G. Sm., Trans. Brit. Mycol. Soc. 39: 111. 1956. [MB292845]. — Type: IMI 44397. Ex-type: NRRL 4744 = NRRL 2550 = NRRL A-6752. Infragen. class: subgen. Nidulantes, sect. Ochraceorosei, ser. Funiculosi. Reproduction: asexual. ITS barcode: EF661223 (alternative markers: BenA = EF661112; CaM = EF661175; RPB2 = EF661078).

Aspergillus fuscicans S.M. Romero et al., Phytotaxa 343: 69. 2018. [MB823159]. — Type: BAFC 52653. Ex-type: BAFCcult 4564!. Infragen. class: subgen. Nidulantes, sect. Usti, ser. Calidousti. Reproduction: asexual. ITS barcode: n.a. (alternative markers: BenA = KY853416; CaM = KY853415; RPB2 = n.a.).

Aspergillus galapagensis (Frisvad et al.) Samson et al., Stud. Mycol. 78: 155. 2014. [MB809589]. Basionym: Neosartorya galapagensis Frisvad et al., Antonie van Leeuwenhoek 93: 96. 2008. [MB506377]. — Type: CBS 117522. Ex-type: CBS 117522 = IBT 16756 = KACC 41935 = DTO 003-H5 = DTO 022-B6. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Neoglabri. Reproduction: homothallic. ITS barcode: MN431368 (alternative markers: BenA = DQ534145; CaM = DQ534151; RPB2 = MN969074).

Aspergillus germanicus Frisvad et al., Stud. Mycol. 69: 91. 2011. [MB560401]. — Type: CBS H-20636. Ex-type: CBS 123887 = DTO 027-D9. Infragen. class: subgen. Nidulantes, sect. Usti, ser. Calidousti. Reproduction: asexual. ITS barcode: FJ531146 (alternative markers: BenA = FJ531172; CaM = FJ531141; RPB2 = KU866944).

Aspergillus giganteus Wehmer, Mem. Soc. Phys. Genève 33: 85. 1901. [MB206765]. — Type: IMI 227678. Ex-type: CBS 526.65 = NRRL 10 = ATCC 10059 = DSM 1146 = IFO 5818 = IMI 227678 = QM 1970 = Thom 5581.13A = WB 10. Infragen. class: subgen. Fumigati, sect. Clavati, ser. Clavati. Reproduction: asexual. ITS barcode: EF669928 (alternative markers: BenA = EF669789; CaM = EF669857; RPB2 = EF669716).

Aspergillus glabripes F. Sklenář et al., Stud. Mycol. 88: 197. 2017. [MB818934]. — Type: PRM 944436. Ex-type: CCF 5474 = DTO 356-E8 = EMSL No. 2462 = NRRL 66618 = IBT 34820. Infragen. class: subgen. Aspergillus, sect. Restricti, ser. Vitricolarum. Reproduction: asexual. ITS barcode: KY087614 (alternative markers: BenA = KY117683; CaM = KY068166; RPB2 = KY117859).

Aspergillus glaucus (L.) Link, Mag. Ges. Naturf. Freunde Berlin 3: 16. 1809. [MB161735]. Basionym: Mucor glaucus L., Sp. Pl. 2: 1186. 1753. [MB185847]. — Type: IMI 211383. Ex-type: CBS 516.65 = NRRL 116 = ATCC 16469 = IMI 211383 = LCP 64.1859 = Thom 5629.C = WB 116. Infragen. class: subgen. Aspergillus, sect. Aspergillus, ser. Aspergillus. Reproduction: homothallic (Link 1809, Chen et al. 2017). ITS barcode: EF652052 (alternative markers: BenA = EF651887; CaM = EF651989; RPB2 = EF651934).

Aspergillus gorakhpurensis Kamal & Bhargava, Trans. Brit. Mycol. Soc. 52: 338. 1969. [MB326632]. — Type: IMI 130728. Ex-type: CBS 648.74 = NRRL 3649 = IMI 130728 = QM 9187 = WB 5346. Infragen. class: subgen. Cremei, sect. Cremei, ser. Cremei. Reproduction: asexual. ITS barcode: EF652145 (alternative markers: BenA = EF652114; CaM = EF652126; RPB2 = EF652097).

Aspergillus gracilis Bainier, Bull. Soc. Mycol. France 23: 90. 1907. [MB167554]. — Type: IMI 211393. Ex-type: CBS 539.65 = NRRL 4962 = ATCC 16906 = IMI 211393 = QM 8915 = WB 4962. Infragen. class: subgen. Aspergillus, sect. Restricti, ser. Restricti. Reproduction: asexual. ITS barcode: EF652045 (alternative markers: BenA = EF651883; CaM = EF652031; RPB2 = EF651980).

Aspergillus granulosus Raper & Thom, Mycologia 36: 565. 1944. [MB284302]. — Type: IMI 17278ii. Ex-type: NRRL 1932 = ATCC 16837 = IMI 17278 = QM 6846 = WB 1932. Infragen. class: subgen. Nidulantes, sect. Usti, ser. Usti. Reproduction: asexual. ITS barcode: EF652430 (alternative markers: BenA = EF652254; CaM = EF652342; RPB2 = EF652166).

Aspergillus griseoaurantiacus Visagie et al., Stud. Mycol. 78: 112. 2014. [MB809197]. — Type: CBS H-21814. Ex-type: CBS 138191 = DTO 267-D8. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Versicolores. Reproduction: asexual. ITS barcode: KJ775553 (alternative markers: BenA = KJ775086; CaM = KJ775357; RPB2 = KU866988).

Aspergillus haitiensis Varga et al., IMA Fungus 1: 194. 2010. [MB517384]. — Type: CBS H-20503. Ex-type: CBS 464.91. Infragen. class: subgen. Nidulantes, sect. Sparsi, ser. Sparsi. Reproduction: asexual. ITS barcode: FJ491657 (alternative markers: BenA = FJ491670; CaM = FJ491645; RPB2 = KU866943).

Aspergillus halophilicus C.M. Chr. et al., Mycologia 51: 636. 1961. [MB326633]. — Type: BPI 566153. Ex-type: CBS 122.62 = NRRL 2739 = ATCC 16401 = IFO 7054 = IMI 211802 = NRRL 4679 = NRRL A-7206 = QM 8894 = WB 4679. Infragen. class: subgen. Aspergillus, sect. Restricti, ser. Halophilici. Reproduction: homothallic. ITS barcode: EF652088 (alternative markers: BenA = EF651926; CaM = EF652034; RPB2 = EF651982).

Aspergillus hancockii Pitt, PLoS ONE 12: e0170254, 16. 2017. [MB818219]. — Type: FRR 3425. Ex-type: CBS 142004 = DTO 360-G7. Infragen. class: subgen. Circumdati, sect. Flavi, ser. Leporum. Reproduction: asexual. ITS barcode: KX858342 (alternative markers: BenA = MBFL01001228.1:26000-28000; CaM = MBFL01000377.1:5000-7000; RPB2 = MBFL01000137:9000-11000).

Aspergillus heldtiae Visagie, Stud. Mycol., this issue. 2020. [MB834200]. — Type: PREM 50864. Ex-type: PPRI 4229 = CMV 004A2. Infragen. class: subgen. Circumdati, sect. Terrei, ser. Terrei. Reproduction: asexual. ITS barcode: MK450656 (alternative markers: BenA = MK450981; CaM = MK451518; RPB2 = MK450809).

Aspergillus heteromorphus Bat. & H. Maia, Anais Soc. Biol. Pernambuco 15: 200. 1957. [MB292846]. — Type: IMI 172288. Ex-type: CBS 117.55 = NRRL 4747 = ATCC 12064 = IMI 172288 = QM 6954 = WB 4747. Infragen. class: subgen. Circumdati, sect. Nigri, ser. Heteromorphi. Reproduction: protoheterothallic; unpublished (genome data, Vesth et al. 2018). ITS barcode: EU821305 (alternative markers: BenA = EF661103; CaM = EF661169; RPB2 = EF661050).

Aspergillus heterothallicus Kwon-Chung et al., Gen. Aspergillus: 502. 1965. [MB326635]. — Type: CBS 488.65. Ex-type: CBS 488.65 = NRRL 5096 = ATCC 16847 = IMI 139277 = QM 8916 = WB 5096. Infragen. class: subgen. Nidulantes, sect. Usti, ser. Usti. Reproduction: Heterothallic. ITS barcode: EF652499 (alternative markers: BenA = EF652323; CaM = EF652411; RPB2 = EF652235).

Aspergillus heyangensis Z.T. Qi et al., Acta Mycol. Sin. 13: 81. 1994. [MB414654]. — Type: HMAS 58982. Ex-type: CBS 101751 = AS 3.4630 = DTO 026-G6. Infragen. class: subgen. Nidulantes, sect. Aenei, ser. Aenei. Reproduction: asexual. ITS barcode: FJ491520 (alternative markers: BenA = FJ491521; CaM = FJ491522; RPB2 = KX423659).

Aspergillus hiratsukae Udagawa et al., Trans. Mycol. Soc. Japan 32: 23. 1991. [MB354908]. — Type: NHL 3008. Ex-type: CBS 294.93 = NRRL 20820 = IMI 349859 = NHL 3008 = DTO 050-E5. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Unilaterales. Reproduction: homothallic. ITS barcode: MN431369 (alternative markers: BenA = AF057324; CaM = AY870699; RPB2 = MN969075).

Aspergillus homomorphus Steiman et al. ex Samson & Frisvad, Stud. Mycol. 50: 58. 2004. [MB500011]. — Type: CBS H-13440. Ex-type: CBS 101889 = ITEM 7556. Infragen. class: subgen. Circumdati, sect. Nigri, ser. Homomorphi. Reproduction: protoheterothallic; unpublished (genome data, Vesth et al. 2018). ITS barcode: EF166063 (alternative markers: BenA = AY820015; CaM = FN594549; RPB2 = MN969076).

Aspergillus hongkongensis C.C. Tsang et al., Diagnostic Microbiology and Infectious Disease 84: 130. 2016. [MB810279]. — Type: NBRC H-13268. Ex-type: CBS 145671 = HKU49 = NBRC 110693 = NCPF 7870 = BCRC FU30360 = DTO 351-C3. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Versicolores. Reproduction: asexual. ITS barcode: AB987907 (alternative markers: BenA = LC000552; CaM = MN969320; RPB2 = LC000578).

Aspergillus hordei F. Sklenář et al., Stud. Mycol. 88: 207. 2017. [MB818937]. — Type: PRM 944446. Ex-type: NRRL 25825 = CCF 5483 = DTO 356-D3 = IBT 34539. Infragen. class: subgen. Aspergillus, sect. Restricti, ser. Penicillioides. Reproduction: asexual. ITS barcode: KY087759 (alternative markers: BenA = KY117822; CaM = KY068309; RPB2 = KY118000).

Aspergillus hortae [as “hortai”] (Langeron) C.W. Dodge, Medic. Mycol.: 628. 1935. [MB252620]. Basionym: Sterigmatocystis hortae Langeron, Bull. Soc. Pathol. Exot.: 383. 1922. [MB252621]. — Type: Langeron 1922, Bull. Soc. Path. Exot. 15: p. 384, Fig. 1 (– lectotype designated here, MBT392292; CBS H-24279 [dried culture] – epitype designated here, MBT392293). Ex-epitype: CBS 124230 = NRRL 274 = ATCC 10070 = IBT 26384 = DTO 051-D6. Infragen. class: subgen. Circumdati, sect. Terrei, ser. Terrei. Reproduction: asexual. ITS barcode: KP987087 (alternative markers: BenA = FJ491706; CaM = KP987054; RPB2 = KP987022).

Aspergillus huiyaniae Y. Horie et al., Mycoscience 55: 218. 2014. [MB803656]. — Type: IFM 57847H. Ex-type: IFM 57847 = JCM 19448 = CBS 139185 = DTO 316-F5. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Fennelliarum. Reproduction: homothallic. ITS barcode: MN431370 (alternative markers: BenA = AB787219; CaM = AB787564; RPB2 = MN969077).

Aspergillus ibericus R. Serra et al., Mycologia 98: 298. 2006. [MB501326]. — Type: MUM-H 03.49. Ex-type: NRRL 35644. Infragen. class: subgen. Circumdati, sect. Nigri, ser. Carbonarii. Reproduction: protoheterothallic; unpublished (genome data, Vesth et al. 2018). ITS barcode: EF661200 (alternative markers: BenA = EF661102; CaM = EF661163; RPB2 = EF661065).

Aspergillus iizukae Sugiy., J. Fac. Sci. Univ. Tokyo, Sect. 3, Bot. 9: 390. 1967. [MB326636]. — Type: TI 0007. Ex-type: CBS 541.69 = NRRL 3750 = IMI 141552 = QM 9325. Infragen. class: subgen. Circumdati, sect. Flavipedes, ser. Flavipedes. Reproduction: asexual. ITS barcode: EF669597 (alternative markers: BenA = EU014086; CaM = EF669555; RPB2 = EF669639).

Aspergillus implicatus Persiani & Maggi, Mycol. Res. 98: 871. 1994. [MB362533]. — Type: ROHB 110 S. Ex-type: CBS 484.95. Infragen. class: subgen. Nidulantes, sect. Sparsi, ser. Implicati. Reproduction: asexual. ITS barcode: FJ491656 (alternative markers: BenA = FJ491667; CaM = FJ491650; RPB2 = MN969078).

Aspergillus incahuasiensis E. Piontelli et al., Int. J. Syst. Evol. Microbiol. 69: 3354. 2019. [MB828145]. — Type: BPI 910732. Ex-type: NRRL 66825. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Multicolores. Reproduction: asexual. ITS barcode: MH473585 (alternative markers: BenA = MH476273; CaM = MH476276; RPB2 = MH476279).

Aspergillus indologenus Frisvad et al., Stud. Mycol. 69: 9. 2011. [MB560389]. — Type: CBS H-20629. Ex-type: CBS 114.80 = IBT 3679 = ITEM 7038. Infragen. class: subgen. Circumdati, sect. Nigri, ser. Japonici. Reproduction: asexual. ITS barcode: AJ280005 (alternative markers: BenA = AY585539; CaM = AM419750; RPB2 = HE984366).

Aspergillus inflatus (Stolk & Malla) Samson et al., Stud. Mycol. 78: 155. 2014. [MB809590]. Basionym: Penicillium inflatum Stolk & Malla, Persoonia 6: 197. 1971. [MB319276]. — Type: CBS H-7500. Ex-type: CBS 682.70 = FRR 1549 = IMI 191498. Infragen. class: subgen. Cremei, sect. Cremei, ser. Inflati. Reproduction: asexual. ITS barcode: FJ531054 (alternative markers: BenA = FJ531008; CaM = FJ531090; RPB2 = JN406529).

Aspergillus infrequens F. Sklenář et al., Stud. Mycol. 88: 207. 2017. [MB818938]. — Type: PRM 944449. Ex-type: NRRL 25868 = CCF 5486 = DTO 356-D6 = IBT 34524. Infragen. class: subgen. Aspergillus, sect. Restricti, ser. Penicillioides. Reproduction: asexual. ITS barcode: KY087770 (alternative markers: BenA = KY117833; CaM = KY068320; RPB2 = KY118011).

Aspergillus insolitus (G. Sm.) Houbraken et al., Stud. Mycol. 78: 155. 2014. [MB809591]. Basionym: Polypaecilum insolitum G. Sm., Trans. Brit. Mycol. Soc. 44: 437. 1961. [MB337467]. — Type: CBS 384.61. Ex-type: CBS 384.61 = ATCC 18164 = IFO 8788 = IMI 75202 = LSHB BB414 = MUCL 3078 = QM 7961 = DTO 049-I6. Infragen. class: subgen. Polypaecilum, sect. Polypaecilum, ser. Polypaecilum. Reproduction: asexual. ITS barcode: KY980622 (alternative markers: BenA = KY980550; CaM = MN969230; RPB2 = JN121510).

Aspergillus insuetus (Bainier) Thom & Church, Manual of the Aspergilli: 153. 1929. [MB267997]. Basionym: Sterigmatocystis insueta Bainier, Bull. Soc. Mycol. France 24: 85. 1908. [MB218947]. — Type: CBS 107.25. Ex-type: CBS 107.25 = NRRL 279 = NRRL 1726 = ATCC 1033 = IFO 4128. Infragen. class: subgen. Nidulantes, sect. Usti, ser. Calidousti. Reproduction: asexual. ITS barcode: EF652457 (alternative markers: BenA = EF652281; CaM = EF652369; RPB2 = EF652193).

Aspergillus insulicola Montem. & A.R. Santiago, Mycopathol. 55: 130. 1975. [MB309225]. — Type: CBS 382.75. Ex-type: CBS 382.75 = NRRL 6138 = ATCC 26220. Infragen. class: subgen. Circumdati, sect. Circumdati, ser. Steyniorum. Reproduction: asexual. ITS barcode: EF661430 (alternative markers: BenA = EF661353; CaM = EF661396; RPB2 = EF661286).

Aspergillus intermedius Blaser, Sydowia 28: 41. 1976. [MB309226]. — Type: IMI 89278. Ex-type: CBS 523.65 = NRRL 82 = ATCC 16444 = DSM 2830 = IMI 089278ii = IMI 89278 = LSHBBB 107 = LSHTM 107 = QM 7403 = Thom 5612.107 = WB 82. Infragen. class: subgen. Aspergillus, sect. Aspergillus, ser. Chevalierorum. Reproduction: homothallic. ITS barcode: EF652074 (alternative markers: BenA = EF651892; CaM = EF652012; RPB2 = EF651958).

Aspergillus iranicus Arzanlou et al., Mycol. Prog. 15: 1085. 2016. [MB817473]. — Type: CBS H-22338. Ex-type: CCTU 756 = CBS 139561 = IBT 32596 = DTO 203-D7. Infragen. class: subgen. Circumdati, sect. Terrei, ser. Nivei. Reproduction: asexual. ITS barcode: KP987077 (alternative markers: BenA = KP987045; CaM = KP987060; RPB2 = KP987034).

Aspergillus israelensis A.J. Chen et al., Stud. Mycol. 84: 63. 2016. [MB816091]. — Type: CBS H-22491. Ex-type: CBS 140627 = IBT 24293 = DTO 325-E2. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Unguium. Reproduction: asexual. ITS barcode: KU866677 (alternative markers: BenA = KU866915; CaM = KU866797; RPB2 = KU867062).

Aspergillus itaconicus Kinosh., Bot. Mag. (Tokyo) 45: 60. 1931. [MB268225]. — Type: IMI 16119. Ex-type: CBS 115.32 = NRRL 161 = ATCC 10021 = IHEM 4378 = IMI 16119 = LSHBA 48 = MUCL 31306 = QM 1980 = Thom 5344 = Thom 5660.48 = WB 161. Infragen. class: subgen. Cremei, sect. Cremei, ser. Cremei. Reproduction: asexual. ITS barcode: EF652147 (alternative markers: BenA = EF652118; CaM = EF652140; RPB2 = EF652103).

Aspergillus ivoriensis Bartoli & Maggi, Trans. Brit. Mycol. Soc. 71: 383. 1979 [1978]. [MB309228]. — Type: RO 101 S. Ex-type: CBS 551.77 = NRRL 22883. Infragen. class: subgen. Nidulantes, sect. Raperorum, ser. Raperorum. Reproduction: asexual. ITS barcode: EF652441 (alternative markers: BenA = EF652265; CaM = EF652353; RPB2 = EF652177).

Aspergillus jaipurensis Samson et al., Stud. Mycol. 78: 155. 2014. [MB809592]. Replaced synonym: Emericella indica Stchigel & Guarro, Mycol. Res. 103: 1059. 1999. [MB460081]. — Type: IMI 378525. Ex-type: IMI 378525 = DTO 320-A9 = FMR 6232 = CBS 952.97. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Nidulantes. Reproduction: homothallic. ITS barcode: MN431371 (alternative markers: BenA = AY339988; CaM = KU866761; RPB2 = KU867024).

Aspergillus janus Raper & Thom, Mycologia 36: 556. 1944. [MB284303]. — Type: IMI 16065. Ex-type: CBS 118.45 = NRRL 1787 = IMI 16065 = NCTC 6970. Infragen. class: subgen. Circumdati, sect. Janorum, ser. Janorum. Reproduction: asexual. ITS barcode: EF669578 (alternative markers: BenA = EU014076; CaM = EF669536; RPB2 = EF669620).

Aspergillus japonicus Saito, Bot. Mag. (Tokyo) 20: 61. 1906. [MB160656]. — Type: CBS 114.51. Ex-type: CBS 114.51 = ITEM 7034. Infragen. class: subgen. Circumdati, sect. Nigri, ser. Japonici. Reproduction: protoheterothallic; unpublished (genome data, Vesth et al. 2018). ITS barcode: AJ279985 (alternative markers: BenA = HE577804; CaM = FN594551; RPB2 = MN969079).

Aspergillus jensenii Jurjević et al., IMA Fungus 3: 70. 2012. [MB800601]. — Type: BPI 880910. Ex-type: NRRL 58600. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Versicolores. Reproduction: asexual. ITS barcode: JQ301892 (alternative markers: BenA = JN854007; CaM = JN854046; RPB2 = JN853835).

Aspergillus kalimae Tanney et al., Stud. Mycol. 88: 249. 2017. [MB822732]. — Type: DAOM 745800. Ex-type: DAOMC 251762 = UAMH 11837 = CBS 143506 = KAS 8135 = SLOAN 4181 = PN08TH-526. Infragen. class: subgen. Polypaecilum, sect. Polypaecilum, ser. Kalimarum. Reproduction: asexual. ITS barcode: KY980650 (alternative markers: BenA = KY980578; CaM = KY980614; RPB2 = KY980475).

Aspergillus kanagawaensis Nehira, J. Jap. Bot. 26: 109. 1951. [MB292847]. — Type: IMI 126690. Ex-type: CBS 538.65 = NRRL 4774 = NRRL 2854 = NRRL 3156 = NRRL A-13499 = IMI 126690. Infragen. class: subgen. Fumigati, sect. Cervini, ser. Cervini. Reproduction: asexual. ITS barcode: EF661275 (alternative markers: BenA = EF661239; CaM = EF661263; RPB2 = EF661236).

Aspergillus karnatakaensis Varga et al., IMA Fungus 1: 203. 2010. [MB517549]. — Type: CBS H-20502. Ex-type: CBS 102800 = IBT 22153. Infragen. class: subgen. Nidulantes, sect. Aenei, ser. Aenei. Reproduction: asexual. ITS barcode: EU482441 (alternative markers: BenA = EU482438; CaM = EU482431; RPB2 = KU866956).

Aspergillus kassunensis Baghd., Novosti. Sist. Nizsh. Rast., 1968 5: 113. 1968. [MB326639]. — D10 in Universitate Mosquensi (holotype); CBS H-24320 (isotype). Ex-type: CBS 419.69 = NRRL 3752 = IMI 334938. Infragen. class: subgen. Nidulantes, sect. Cavernicolarum, ser. Cavernicolarum. Reproduction: asexual. ITS barcode: EF652461 (alternative markers: BenA = EF652285; CaM = EF652373; RPB2 = EF652197).

Aspergillus keratitidis (W.L. Chen et al.) Zalar & W.L. Chen. Extremophiles 21: 762. 2017. [MB818566]. Basionym: Sagenomella keratitidis W.L. Chen et al., Bot. Stud. 50: 332. 2009. [MB541734]. — Type: BCRC 34221. Ex-type: BCRC 34221 = DTO 198-E8. Infragen. class: subgen. Polypaecilum, sect. Polypaecilum, ser. Noonimiarum. Reproduction: asexual. ITS barcode: KY980616 (alternative markers: BenA = KY980544; CaM = KY980580; RPB2 = KY980443).

Aspergillus keveii Varga et al., Stud. Mycol. 59: 120. 2007. [MB505570]. — Type: CBS 209.92. Ex-type: CBS 209.92 = DTO 013-G8. Infragen. class: subgen. Nidulantes, sect. Usti, ser. Calidousti. Reproduction: asexual. ITS barcode: EU076354 (alternative markers: BenA = EU076376; CaM = EU076365; RPB2 = KU866938).

Aspergillus keveioides L. Wang, Mycosystema 32 (suppl.): 139. 2013. [MB800250]. — Type: HMAS242394. Ex-type: AS 3.15305 = CBS 132737 = DTO 328-D7. Infragen. class: subgen. Nidulantes, sect. Usti, ser. Calidousti. Reproduction: asexual. ITS barcode: JN982704 (alternative markers: BenA = JN982694; CaM = JN982684; RPB2 = MN969151).

Aspergillus koreanus Hyang B. Lee et al., Fungal Diversity 80: 142. 2016. [MB816938]. — Type: NIBR EML-GSNP1-1. Ex-type: JMRC:SF:012334. Infragen. class: subgen. Cremei, sect. Cremei, ser. Inflati. Reproduction: asexual. ITS barcode: KX216525 (alternative markers: BenA = KX216530; CaM = KX216528; RPB2 = KX216531).

Aspergillus krugeri Visagie, Stud. Mycol., this issue. 2020. [MB834203]. — Type: PREM 62309. Ex-type: PPRI 8986 = CMV 006G4. Infragen. class: subgen. Circumdati, sect. Flavi, ser. Flavi. Reproduction: asexual. ITS barcode: MK450655 (alternative markers: BenA = MK451098; CaM = MK451517; RPB2 = MK450808).

Aspergillus labruscus Fungaro et al., Sci. Rep. 7: 6203, 4. 2017. [MB815746]. — Type: CCT 7800. Ex-type: DTO 357-D4 = ITAL 22.223 = IBT 33586. Infragen. class: subgen. Circumdati, sect. Nigri, ser. Japonici. Reproduction: asexual. ITS barcode: KU708544 (alternative markers: BenA = KT986014; CaM = KT986008; RPB2 = MN969196).

Aspergillus laciniosus S.B. Hong et al., Int. J. Syst. Evol. Microbiol. 56: 484. 2006. [MB521269]. — Type: CBS 117721. Ex-type: CBS 117721 = NRRL 35589 = KACC 41657 = DTO 164-I2. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Fumigati. Reproduction: homothalliic. ITS barcode: AB299413 (alternative markers: BenA = AY870756; CaM = AY870716; RPB2 = MN969080).

Aspergillus lanosus Kamal & Bhargava, Trans. Brit. Mycol. Soc. 52: 336. 1969. [MB326640]. — Type: IMI 130727. Ex-type: CBS 650.74 = DTO 034-B7 = NRRL 3648 = IMI 130727 = QM 9183 = WB 5347. Infragen. class: subgen. Circumdati, sect. Flavi, ser. Alliacei. Reproduction: asexual. ITS barcode: EF661553 (alternative markers: BenA = MG517633; CaM = MG518017; RPB2 = EU021642).

Aspergillus latilabiatus A.J. Chen et al., Stud. Mycol. 84: 66. 2016. [MB816093]. — Type: CBS H-22514. Ex-type: CBS 426.93 = IBT 33959 = DTO 320-B2. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Nidulantes. Reproduction: homothallic; asexual morph unknown. ITS barcode: KU866624 (alternative markers: BenA = KU866864; CaM = KU866762; RPB2 = KU867025).

Aspergillus lentulus Balajee & K.A. Marr, Eukaryot. Cell 4: 631. 2005. [MB356679]. — Type: BPI 863540. Ex-type: CBS 117885 = NRRL 35552 = IBT 27201 = KACC 41940 = DTO 004-E9. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Fumigati. Reproduction: Heterothallic. ITS barcode: EF669969 (alternative markers: BenA = EF669825; CaM = EF669895; RPB2 = EF669756).

Aspergillus leporis States & M. Chr., Mycologia 58: 738. 1966. [MB326641]. — Type: NY RMF 99. Ex-type: CBS 151.66 = NRRL 3216 = ATCC 16490 = NRRL A-14256 = NRRL A-15810 = QM 8995 = RMF99 = WB 5188. Infragen. class: subgen. Circumdati, sect. Flavi, ser. Leporum. Reproduction: asexual. ITS barcode: AF104443 (alternative markers: BenA = EF661499; CaM = EF661541; RPB2 = EF661459).

Aspergillus leucocarpus Hadlok & Stolk, Antonie van Leeuwenhoek 35: 9. 1969. [MB326642]. — Type: CBS 353.68. Ex-type: CBS 353.68 = NRRL 3497 = QM 9365 = QM 9707. Infragen. class: subgen. Aspergillus, sect. Aspergillus, ser. Leucocarpi. Reproduction: homothallic. ITS barcode: EF652087 (alternative markers: BenA = EF651925; CaM = EF652023; RPB2 = EF651972).

Aspergillus levisporus Hubka et al., Stud. Mycol. 88: 99. 2017. [MB818735]. — Type: CBS H-22820. Ex-type: CBS 141767 = DTO 355-G4 = EMSL No.3211 = CCF 5378 = IBT 34512. Infragen. class: subgen. Aspergillus, sect. Aspergillus, ser. Aspergillus. Reproduction: homothallic. ITS barcode: LT670950 (alternative markers: BenA = LT671094; CaM = LT671095; RPB2 = LT671096).

Aspergillus longistipitatus F. Sklenář et al., Mycologia 112: 363. 2020. [MB832714]. — Type: PRM 951573. Ex-type: CCF 5788 = EMSL No. 2705 = NRRL 66886. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Unguium. Reproduction: asexual. ITS barcode: MK713534 (alternative markers: BenA = MK695641; CaM = MK695652; RPB2 = MK695663).

Aspergillus longivesica L.H. Huang & Raper, Mycologia 63: 53. 1971. [MB309229]. — Type: WIS Nl l79. Ex-type: CBS 530.71 = NRRL 5215 = ATCC 22434 = IMI 156966 = JCM 10186 = QM 9698. Infragen. class: subgen. Fumigati, sect. Clavati, ser. Clavati. Reproduction: asexual. ITS barcode: EF669991 (alternative markers: BenA = EF669847; CaM = EF669917; RPB2 = EF669778).

Aspergillus loretoensis S. González-Martinez & A. Portillo-López, Extremophiles 23: 562. 2019. [MB830181]. — Type: CM-CNRG 624. Ex-type: BCMEX-UABC 6006. Infragen. class: subgen. Polypaecilum, sect. Polypaecilum, ser. Salinarum. Reproduction: asexual. ITS barcode: KX236325 (alternative markers: BenA = MK309399; CaM = MK309400; RPB2 = MK312162).

Aspergillus luchuensis Inui, J. Coll. Agric. Imp. Univ. Tokyo 13: 469. 1901. [MB151291]. — Type: Inui 1901, J. Coll. Agric. Imp. Univ. Tokyo 13: Tafel XXII, Figs 1–8 (– lectotype designated here, MBT392280; CBS H-24280 [dried culture] – epitype designated here, MBT392281). Ex-epitype: CBS 205.80 = NBRC 4281 = KACC 46772 = IFM 47726 = RIB 2642. Infragen. class: subgen. Circumdati, sect. Nigri, ser. Nigri. Reproduction: protoheterothallic; both MAT idiomorphs detected (Mageswari et al. 2016, Yamada et al. 2016, de Vries et al. 2017). ITS barcode: JX500081 (alternative markers: BenA = JX500062; CaM = JX500071; RPB2 = MN969081).

Aspergillus lucknowensis J.N. Rai et al., Canad. J. Bot. 46: 1483. 1968. [MB326643]. — Type: CBS 449.75. Ex-type: CBS 449.75 = NRRL 3491 = ATCC 18607 = IMI 278379 = PIL623 = QM 9271 = WB 5377. Infragen. class: subgen. Nidulantes, sect. Usti, ser. Deflecti. Reproduction: asexual. ITS barcode: EF652459 (alternative markers: BenA = EF652283; CaM = EF652371; RPB2 = EF652195).

Aspergillus luppiae [as “luppii”] Hubka et al., Mycologia 107: 187. 2015. [MB825372]. — Type: PRM 923447. Ex-type: NRRL 6326 = CBS 653.74 = CCF 4545. Infragen. class: subgen. Circumdati, sect. Flavipedes, ser. Spelaei. Reproduction: asexual. ITS barcode: EF669617 (alternative markers: BenA = EU014079; CaM = EF669575; RPB2 = EF669659).

Aspergillus luteovirescens Blochwitz, Ann. Mycol. 31: 80. 1933. [MB269992]. — Type: CBS H-23401. Ex-type: CBS 620.95 = DTO 010-H1 = CBS 116.32 (dead) = IMI 348034 = NRRL 4858 = WB 4858. Infragen. class: subgen. Circumdati, sect. Flavi, ser. Nomiarum. Reproduction: protoheterothallic; MAT 1-2-1 detected (Ramirez-Prado et al. 2008). ITS barcode: MG662406 (alternative markers: BenA = MG517625; CaM = MG517998; RPB2 = MG517808).

Aspergillus magaliesburgensis Visagie, Stud. Mycol., this issue. 2020. [MB834204]. — Type: PREM 62314. Ex-type: PPRI 6165 = CMV 007A3. Infragen. class: subgen. Circumdati, sect. Flavi, ser. Alliacei. Reproduction: asexual. ITS barcode: MK450649 (alternative markers: BenA = MK451116; CaM = MK451511; RPB2 = MK450802).

Aspergillus magnivesiculatus F. Sklenář et al., Stud. Mycol. 88: 211. 2017. [MB818939]. — Type: PRM 944444. Ex-type: NRRL 25866 = CCF 5488 = IBT 34816. Infragen. class: subgen. Aspergillus, sect. Restricti, ser. Penicillioides. Reproduction: asexual. ITS barcode: KY087768 (alternative markers: BenA = KY117831; CaM = KY068318; RPB2 = KY118009).

Aspergillus mallochii Visagie et al., MycoKeys 19: 16. 2017. [MB819025]. — Type: DAOM 740296. Ex-type: DAOMC 146054 = CBS 141928 = DTO 357-A5 = KAS 7618. Infragen. class: subgen. Aspergillus, sect. Aspergillus, ser. Rubri. Reproduction: homothallic. ITS barcode: KX450907 (alternative markers: BenA = KX450889; CaM = KX450902; RPB2 = KX450894).

Aspergillus maritimus Samson & W. Gams, Adv. Pen. Asp. Syst.: 43. 1986 [1985]. [MB114709]. — Type: CBS 186.77. Ex-type: CBS 186.77. Infragen. class: subgen.: unknown, sect.: unknown, ser.: unknown. Reproduction: homothallic. ITS barcode: n.a. (alternative markers: BenA = n.a.; CaM = n.a.; RPB2 = n.a.).

Aspergillus marvanovae Hubka et al., Int. J. Syst. Evol. Microbiol. 63: 787. 2013. [MB801064]. — Type: PRM 860539. Ex-type: NRRL 62486 = IBT 31279 = CCM 8003 = CCF 4037 = IFM 60873 = DTO 303-A2. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Unilaterales. Reproduction: protoheterothallic; MAT 1-1-1 detected (Hubka et al. 2017). ITS barcode: HE974450 (alternative markers: BenA = HE974387; CaM = HE974389; RPB2 = HE974396).

Aspergillus megasporus Visagie et al., MycoKeys 19: 17. 2017. [MB819028]. — Type: DAOM 741781. Ex-type: DAOMC 250799 = CBS 141929= DTO 356-H7 = KAS 6176. Infragen. class: subgen. Aspergillus, sect. Aspergillus, ser. Aspergillus. Reproduction: homothallic. ITS barcode: KX450910 (alternative markers: BenA = KX450892; CaM = KX450905; RPB2 = KX450897).

Aspergillus melleus Yukawa, J. Coll. Agric. Imp. Univ. Tokyo 1: 358. 1911. [MB164593]. — Type: CBS 546.65. Ex-type: CBS 546.65 = NRRL 5103 = IBT 13510 = IBT 13511 = IBT 13875 = ATCC 16889 = WB 5103. Infragen. class: subgen. Circumdati, sect. Circumdati, ser. Circumdati. Reproduction: asexual. ITS barcode: EF661425 (alternative markers: BenA = EF661326; CaM = EF661391; RPB2 = EF661309).

Aspergillus microcysticus Sappa, Allionia 2: 251. 1955. [MB292848]. — Type: IMI 139275. Ex-type: CBS 120.58 = NRRL 4749 = ATCC 16826 = IMI 139275 = QM 8158 = WB 4749. Infragen. class: subgen. Circumdati, sect. Terrei, ser. Ambigui. Reproduction: asexual. ITS barcode: EF669607 (alternative markers: BenA = EF669515; CaM = EF669565; RPB2 = EF669649).

Aspergillus micronesiensis Visagie et al., Stud. Mycol. 78: 105. 2014. [MB809192]. — Type: CBS H-21809. Ex-type: CBS 138183 = DTO 267-D5. Infragen. class: subgen. Circumdati, sect. Flavipedes, ser. Flavipedes. Reproduction: asexual. ITS barcode: KJ775548 (alternative markers: BenA = KJ775085; CaM = KP987067; RPB2 = KP987023).

Aspergillus microperforatus J.P.Z. Siqueira et al., Med. Mycol. 56: 545. 2018. [MB820080]. — Type: CBS H-22998. Ex-type: UTHSCSA DI16-407 = CBS 142376 = FMR 14071. Infragen. class: subgen. Aspergillus, sect. Aspergillus, ser. Rubri. Reproduction: homothallic. ITS barcode: LT627271 (alternative markers: BenA = LT627296; CaM = LT627321; RPB2 = LT627346).

Aspergillus minisclerotigenes Vaamonde et al., Int. J. Syst. Evol. Microbiol. 58: 733. 2008. [MB505188]. — Type: Pildain et al. 2008, Int. J. Syst. Evol. Microbiol. 58: p. 732, Fig. 3 (– lectotype designated here, MBT392282; CBS H-24281 [dried culture] – epitype designated here, MBT392283). Ex-epitype: CBS 117635 = DTO 009-F7 = DTO 303-C6 = IBT 25032. Infragen. class: subgen. Circumdati, sect. Flavi, ser. Flavi. Reproduction: protoheterothallic; both MAT idiomorphs detected (Soares et al. 2012). ITS barcode: EF409239 (alternative markers: BenA = EF203148; CaM = MG518009; RPB2 = MG517799).

Aspergillus miraensis (L.C. Zhang et al.) Hubka et al., Plant Syst. Evol. 302: 1288. 2016. [MB816283]. Basionym: Emericella miraensis L.C. Zhang et al., Mycotaxon 125: 132. 2013. [MB800444]. — Type: CGMCC 3.14984. Ex-type: CGMCC 3.14984 = DTO 323-B2. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Stellati. Reproduction: homothallic. ITS barcode: KU866642 (alternative markers: BenA = KC342577; CaM = KU866780; RPB2 = KU867045).

Aspergillus monodii (Locq.-Lin.) Varga et al., Stud. Mycol. 69: 91. 2011. [MB560402]. Basionym: Fennellia monodii Locq.-Lin., Mycotaxon 39: 10. 1990. [MB126894]. — Type: LCP 89-3570 (PC). Ex-type: DTO 69-A3 = CBS 435.93 = DTO 026-I4. Infragen. class: subgen. Nidulantes, sect. Usti, ser. Monodiorum. Reproduction: homothallic; asexual morph unknown. ITS barcode: FJ531150 (alternative markers: BenA = FJ531171; CaM = FJ531142; RPB2 = MN969082).

Aspergillus montevidensis Talice & Mackinnon, Compt. Rend. Soc. Biol. Fr. 108: 1007. 1931. [MB309231]. — Type: NRRL 108. Ex-type: CBS 491.65 = NRRL 108 = BPI 884202 = ATCC 10077 = IHEM 3337 = IMI 172290 = NRRL 109 = QM 7423 = Thom 5290 = Thom 5633.24 = WB 108. Infragen. class: subgen. Aspergillus, sect. Aspergillus, ser. Chevalierorum. Reproduction: homothallic. ITS barcode: EF652077 (alternative markers: BenA = EF651898; CaM = EF652020; RPB2 = EF651964).

Aspergillus mottae C. Soares et al., Mycologia 104: 692. 2012. [MB561841]. — Type: MUM-H 10.231. Ex-type: CBS 130016 = DTO 223-C8. Infragen. class: subgen. Circumdati, sect. Flavi, ser. Flavi. Reproduction: protoheterothallic; both MAT idiomorphs detected (Soares et al. 2013). ITS barcode: JF412767 (alternative markers: BenA = HM803086; CaM = MG518058; RPB2 = MG517878).

Aspergillus movilensis A. Nováková et al., Mycologia 107: 190. 2015. [MB808144]. — Type: PRM 923448. Ex-type: PRM 923449 = CCF 4410 = CMF ISB 2614 = NRRL 62819 = CBS 134395 = DTO 316-C6. Infragen. class: subgen. Circumdati, sect. Flavipedes, ser. Spelaei. Reproduction: asexual. ITS barcode: KP987089 (alternative markers: BenA = HG916697; CaM = HG916740; RPB2 = HG916718).

Aspergillus multicolor Sappa, Allionia 2: 87. 1954. [MB292849]. — Type: IMI 69875. Ex-type: CBS 133.54 = NRRL 4775 = ATCC 16804 = IFO 8133 = IMI 69857 = LSHBBB 356 = QM 1952 = WB 4281 = WB 4775. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Multicolores. Reproduction: asexual. ITS barcode: EF652477 (alternative markers: BenA = EF652301; CaM = EF652389; RPB2 = EF652213).

Aspergillus multiplicatus Yaguchi et al., Mycoscience 35: 310. 1994. [MB412530]. — Type: CBM PF-1154. Ex-type: CBS 646.95 = IBT 17517 = DTO 050-E2. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Unilaterales. Reproduction: homothallic. ITS barcode: HE974445 (alternative markers: BenA = DQ114129; CaM = DQ114137; RPB2 = HE974397).

Aspergillus mulundensis Bills & Frisvad, J Antibiot. 69: 143. 2016. [MB813062]. — Type: DSMZ 5745. Ex-type: DTO 316-C9 = DSMZ 5745a = IBT 33104. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Multicolores. Reproduction: asexual. ITS barcode: KP985732 (alternative markers: BenA = KP985735; CaM = KP985734; RPB2 = KU866989).

Aspergillus muricatus Udagawa et al., Mycotaxon 52: 210. 1994. [MB362530]. — Type: CBM BF-42515. Ex-type: CBS 112808 = NRRL 35674 = IBT 19374 = IMI 36852. Infragen. class: subgen. Circumdati, sect. Circumdati, ser. Circumdati. Reproduction: homothallic. ITS barcode: EF661434 (alternative markers: BenA = EF661356; CaM = EF661377; RPB2 = EF661314).

Aspergillus navahoensis M. Chr. & States, Mycologia 74: 226. 1982. [MB110496]. — Type: NY SD-5. Ex-type: CBS 351.81 = NRRL 13002 = ATCC 44663 = IMI 259971. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Nidulantes. Reproduction: homothallic. ITS barcode: EF652424 (alternative markers: BenA = EF652248; CaM = EF652336; RPB2 = EF652160).

Aspergillus neoafricanus Samson et al., Stud. Mycol. 69: 53. 2011. [MB560391]. Replaced synonym: Aspergillus terreus var. africanus Fennell & Raper, Mycologia 47: 86. 1955. [MB351904]. — Type: Fennell & Raper 1955, Mycologia 47: p. 87, Fig. 8 (– lectotype designated here, MBT392284; CBS H-24282 [dried culture] – epitype designated here, MBT392285). Ex-epitype: CBS 130.55 = NRRL 2399 = ATCC 16792 = IHEM 4380 = IMI 61457 = MUCL 31316 = NRRL A-3175 = QM 1913 = VKMF-2037 = WB 2399. Infragen. class: subgen. Circumdati, sect. Terrei, ser. Terrei. Reproduction: asexual. ITS barcode: EF669585 (alternative markers: BenA = EF669516; CaM = EF669543; RPB2 = EF669627).

Aspergillus neoalliaceus A. Nováková et al., Stud. Mycol. 93: 43. 2019. [MB823775]. — Type: CBS H-23363. Ex-type: CBS 143681 = DTO 326-D3 = S765 = CCF 5433 = IBT 33110 = IBT 33353. Infragen. class: subgen. Circumdati, sect. Flavi, ser. Alliacei. Reproduction: asexual. ITS barcode: MH279420 (alternative markers: BenA = MG517763; CaM = MG518133; RPB2 = MG517954).

Aspergillus neobridgeri Frisvad & Samson, Stud. Mycol. 50: 35. 2004. [MB500004]. — Type: CBS 559.82. Ex-type: CBS 559.82 = NRRL 13078 = IBT 14026. Infragen. class: subgen. Circumdati, sect. Circumdati, ser. Sclerotiorum. Reproduction: asexual. ITS barcode: EF661410 (alternative markers: BenA = EF661345; CaM = EF661359; RPB2 = EF661298).

Aspergillus neocarnoyi Kozak., Mycol. Pap. 161: 63. 1989. [MB127756]. Basionym: Eurotium carnoyi Malloch & Cain, Canad. J. Bot. 50: 63. 1972. [MB297362]. — Type: IMI 172279. Ex-type: CBS 471.65 = NRRL 126 = ATCC 16924 = IMI 172279 = LSHTM A32 = QM 7402 = Thom 5612.A32 = WB 126. Infragen. class: subgen. Aspergillus, sect. Aspergillus, ser. Aspergillus. Reproduction: homothallic. ITS barcode: EF652057 (alternative markers: BenA = EF651903; CaM = EF651985; RPB2 = EF651942).

Aspergillus neoflavipes Hubka et al., Mycologia 107: 192. 2015. [MB808147]. Replaced synonym: Fennellia flavipes B.J. Wiley & E.G. Simmons, Mycologia 65: 937. 1973. [MB314109]. — Type: BPI 410858. Ex-type: CBS 260.73 = NNRL 5504 = ATCC 24484 = IMI 171883 = IFM 40894 = CCF 4552. Infragen. class: subgen. Circumdati, sect. Flavipedes, ser. Flavipedes. Reproduction: homothallic. ITS barcode: EF669614 (alternative markers: BenA = EU014084; CaM = EF669572; RPB2 = EF669656).

Aspergillus neoglaber Kozak., Mycol. Pap. 161: 56. 1989. [MB127762]. Replaced synonym: Aspergillus fischeri var. glaber Fennell & Raper, Mycologia 47: 74. 1955. [MB351897]. — Type: IMI 61447. Ex-type: CBS 111.55 = NRRL 2163 = ATCC 16909 = IFO 8789 = IMI 061447ii = IMI 367412 = IMI 61447 = NRRL A-2175 = QM 1903 = WB 2163 = DTO 050-D4 = DTO 164-H7. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Neoglabri. Reproduction: homothallic. ITS barcode: EF669948 (alternative markers: BenA = EU014107; CaM = EU014120; RPB2 = EF669736).

Aspergillus neoindicus Samson et al., Stud. Mycol. 69: 53. 2011. [MB560394]. Replaced synonym: Aspergillus niveus var. indicus [as “indica”] Lal & A.K. Sarbhoy, Indian Phytopathol. 25: 311. 1972 [1973]. [MB252165]. — Type: ITCC 1575 (holotype); CBS H-24283 (isotype). Ex-type: CBS 444.75 = NRRL 6134 = IMI 334935. Infragen. class: subgen. Circumdati, sect. Terrei, ser. Nivei. Reproduction: asexual. ITS barcode: EF669616 (alternative markers: BenA = EF669532; CaM = EF669574; RPB2 = EF669658).

Aspergillus neoniger Varga et al., Stud. Mycol. 69: 16. 2011. [MB560390]. — Type: CBS H-20630. Ex-type: CBS 115656 = NRRL 62634. Infragen. class: subgen. Circumdati, sect. Nigri, ser. Nigri. Reproduction: protoheterothallic; unpublished (genome data; Vesth et al. 2018). ITS barcode: FJ491682 (alternative markers: BenA = FJ491691; CaM = FJ491700; RPB2 = KC796429).

Aspergillus neoniveus Samson et al., Stud. Mycol. 69: 53. 2011. [MB560395]. Replaced synonym: Emericella nivea B.J. Wiley & E.G. Simmons, Mycologia 65: 934. 1973. [MB313507]. — Type: QM 8942. Ex-type: CBS 261.73 = NRRL 5299 = ATCC 24482 = IMI 171878. Infragen. class: subgen. Circumdati, sect. Flavipedes, ser. Neonivei. Reproduction: homothallic. ITS barcode: EF669612 (alternative markers: BenA = EU014098; CaM = EF669570; RPB2 = KP987024).

Aspergillus nidulans (Eidam) G. Winter, Rabenh. Krypt.-Fl., ed. 2, 1: 62. 1884. [MB182069]. Basionym: Sterigmatocystis nidulans Eidam, Beitr. Biol. Pflanzen 3: 393. 1883. [MB221350]. — Type: IMI 86806. Ex-type: CBS 589.65 = NRRL 187 = ATCC 10074 = IHEM 3563 = IMI 126691 = IMI 86806 = QM 1985 = Thom 4640.5 = WB 187. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Nidulantes. Reproduction: homothallic. ITS barcode: EF652427 (alternative markers: BenA = EF652251; CaM = EF652339; RPB2 = EF652163).

Aspergillus niger Tiegh., Ann. Sci. Nat., Bot., ser. 5, 8: 240. 1867.; nom. cons. (Kozakiewicz et al. 1992). [MB284309]. — Type: CBS 554.65. Ex-type: CBS 554.65 = NRRL 326 = ATCC 16888 = IFO 33023 = IHEM 3415 = IMI 050566ii = IMI 50566 = JCM 10254 = QM 9270 = QM 9946 = Thom 2766 = WB 326. Infragen. class: subgen. Circumdati, sect. Nigri, ser. Nigri. Reproduction: protoheterothallic; both MAT idiomorphs detected (Pel et al., 2007, Mageswari et al, 2016). ITS barcode: EF661186 (alternative markers: BenA = EF661089; CaM = EF661154; RPB2 = EF661058).

Aspergillus nishimurae Takada et al., Mycoscience 42: 362. 2001. [MB474712]. — Type: CBM-FA-919 (holotype); PRM 935217 (epitype). Ex-epitype: IFM 54133 = CBM-FA-0910 = CCF 4547 = IBT 29024. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Unilaterales. Reproduction: Heterothallic. ITS barcode: HE974449 (alternative markers: BenA = AB201360; CaM = HE974392; RPB2 = HE974393).

Aspergillus niveoglaucus Thom & Raper, U.S.D.A. Misc. Pub. 426: 35. 1941. [MB120985]. — Type: IMI 32050ii. Ex-type: CBS 114.27 = CBS 517.65 = NRRL 127 = ATCC 10075 = IMI 32050 = LSHBA 16 = NRRL 129 = NRRL 130 = QM 1977 = Thom 5612.A16 = Thom 5633 = Thom 5633.7 = Thom 7053.2 = WB 127 = WB 130. Infragen. class: subgen. Aspergillus, sect. Aspergillus, ser. Aspergillus. Reproduction: homothallic. ITS barcode: EF652058 (alternative markers: BenA = EF651905; CaM = EF651993; RPB2 = EF651943).

Aspergillus niveus Blochwitz, Ann. Mycol. 27: 205. 1929. [MB272402]. — Type: IMI 171878. Ex-type: CBS 115.27 = NRRL 5505. Infragen. class: subgen. Circumdati, sect. Terrei, ser. Nivei. Reproduction: asexual; the putative sexual morph of A. niveus, Emericella nivea Wiley & Simmons, represents a different species - A. neoniveus (Samson et al. 2011a). ITS barcode: EF669615 (alternative markers: BenA = EF669528; CaM = EF669573; RPB2 = EF669657).

Aspergillus nomiae [as “nomius”] Kurtzman et al., Antonie van Leeuwenhoek 53: 151. 1987. [MB634998]. — Type: NRRL 13137. Ex-type: CBS 260.88 = NRRL 13137 = ATCC 15546 = FRR 3339 = IMI 331920 = LCP 89.3558 = NRRL 6108 = NRRL A-13671 = NRRL A-13794. Infragen. class: subgen. Circumdati, sect. Flavi, ser. Nomiarum. Reproduction: Heterothallic (Horn et al. 2011). ITS barcode: AF027860 (alternative markers: BenA = AF255067; CaM = AY017588; RPB2 = EF661456).

Aspergillus noonimiae Tanney et al., Stud. Mycol. 88: 252. 2017. [MB822733]. — Type: DAOM 745797. Ex-type: DAOMC 251754 = UAMH 11836 = CBS 143382 = KAS 8125 = SLOAN 7955 = PN06TH-370. Infragen. class: subgen. Polypaecilum, sect. Polypaecilum, ser. Noonimiarum. Reproduction: asexual. ITS barcode: KY980641 (alternative markers: BenA = KY980569; CaM = KY980605; RPB2 = KY980466).

Aspergillus novofumigatus S.B. Hong et al., Mycologia 97: 1368. 2006. [MB500297]. — Type: CBS 117520. Ex-type: CBS 117520 = IBT 16806 = KACC 41934 = IFM 55215 = CCF 4695 = DTO 022-B7 = DTO 003-H3. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Fumigati. Reproduction: protoheterothallic; both MAT idiomorphs detected (Dudová 2014). ITS barcode: MN431372 (alternative markers: BenA = DQ094886; CaM = DQ094893; RPB2 = MN969083).

Aspergillus novoguineensis A.J. Chen et al., Stud. Mycol. 85: 75. 2016. [MB817725]. — Type: CBS H-22729. Ex-type: CBS 906.96 = DTO 021-G5 = IBT 29312. Infragen. class: subgen. Fumigati, sect. Cervini, ser. Cervini. Reproduction: asexual. ITS barcode: FJ491622 (alternative markers: BenA = FJ491641; CaM = FJ491605; RPB2 = KX423681).

Aspergillus novoparasiticus S.S. Gonçalves et al., Med. Mycol. 50: 158. 2011. [MB516612]. — Type: CBS H-20401. Ex-type: CBS 126849 = DTO 223-C3 = LEMI 250 = FMR 10121. Infragen. class: subgen. Circumdati, sect. Flavi, ser. Flavi. Reproduction: protoheterothallic; MAT1-1-1 detected (Carvajal-Campos et al. 2017). ITS barcode: MG662397 (alternative markers: BenA = MG517684; CaM = MG518055; RPB2 = MG517875).

Aspergillus nutans McLennan & Ducker, Austral. J. Bot. 2: 355. 1954. [MB292850]. — Type: IMI 62874ii. Ex-type: CBS 121.56 = NRRL 575 = NRRL 4364 = NRRL A-6280 = ATCC 16914 = IFO 8134 = IMI 062874ii = IMI 62874 = QM 8159 = WB 4364 = WB 4546 = WB 4776. Infragen. class: subgen. Fumigati, sect. Cervini, ser. Cervini. Reproduction: asexual. ITS barcode: EF661272 (alternative markers: BenA = EF661249; CaM = EF661262; RPB2 = EF661227).

Aspergillus occultus Visagie et al., Stud. Mycol. 78: 32. 2014. [MB809198]. — Type: CBS H-21794. Ex-type: CBS 137330 = IBT 32285 = DTO 231-A7. Infragen. class: subgen. Circumdati, sect. Circumdati, ser. Steyniorum. Reproduction: asexual. ITS barcode: KJ775443 (alternative markers: BenA = KJ775061; CaM = KJ775239; RPB2 = MN969084).

Aspergillus ochraceopetaliformis Bat. & Maia, Anais Soc. Biol. Pernambuco 15: 213. 1957. [MB292851]. — Type: no 270 Instituto de Micologia, Iniversidade do Recife. Ex-type: CBS 123.55 = NRRL 4752 = IBT 14347 = ATCC 12066 = IMI 211804 = QM 6955 = WB 4752. Infragen. class: subgen. Circumdati, sect. Circumdati, ser. Steyniorum. Reproduction: asexual. ITS barcode: EF661429 (alternative markers: BenA = EF661350; CaM = EF661388; RPB2 = EF661283).

Aspergillus ochraceoroseus Bartoli & Maggi, Trans. Brit. Mycol. Soc. 71: 393. 1979 [1978]. [MB309233]. — Type: RO 104 S. Ex-type: CBS 550.77 = NRRL 28622 = ATCC 38873 = SRRC1432. Infragen. class: subgen. Nidulantes, sect. Ochraceorosei, ser. Ochraceorosei. Reproduction: asexual. ITS barcode: EF661224 (alternative markers: BenA = EF661113; CaM = EF661137; RPB2 = EF661074).

Aspergillus ochraceus K. Wilh., Beitr. Kenntn. Aspergillus: 66. 1877. [MB190223]. — Type: IMI 16247iv. Ex-type: CBS 108.08 = NRRL 398 = IBT 11952 = ATCC 1008 = CECT2093 = DSM 824 = HARVARD296 = IMI 16247 = NCTC 3889 = NRRL 1642 = QM 6731 = Thom 112 = WB 398. Infragen. class: subgen. Circumdati, sect. Circumdati, ser. Circumdati. Reproduction: asexual. ITS barcode: EF661419 (alternative markers: BenA = EF661322; CaM = EF661381; RPB2 = EF661302).

Aspergillus oerlinghausenensis Bader & Houbraken, FEMS Microbiol. Letter 363 (3/fnv236): 4. 2016. [MB813868]. — Type: CBS H-22119. Ex-type: CBS 139183 = IBT 33878 = DTO 316-A3. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Fumigati. Reproduction: protoheterothallic; MAT 1-1-1 detected (Houbraken et al. 2016). ITS barcode: KT359601 (alternative markers: BenA = KT359603; CaM = KT359605; RPB2 = MN969162).

Aspergillus olivicola Frisvad et al., Mycologia 100: 781. 2008. [MB507362]. — Type: CBS H-19888. Ex-type: CBS 119.37 = IBT 21903 = DTO 002-I2. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Stellati. Reproduction: homothallic. ITS barcode: EU448268 (alternative markers: BenA = AY339996; CaM = EU443986; RPB2 = KU866923).

Aspergillus olivimuriae S.W. Peterson & S. Crognale, Int. J. Syst. Evol. Microbiol. 69: 2901. 2019. [MB826866]. — Type: BPI 910647. Ex-type: NRRL 66783 = DIBAF 6C2. Infragen. class: subgen. Circumdati, sect. Flavipedes, ser. Olivimuriarum. Reproduction: asexual. ITS barcode: MH298877 (alternative markers: BenA = MH492010; CaM = MH492011; RPB2 = MH492012).

Aspergillus omanensis Y. Horie & Udagawa, Mycoscience 36: 391. 1995. [MB414655]. — Type: CBM FA-700. Ex-type: CBM FA-700 = IFM 54275. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Nidulantes. Reproduction: homothallic. ITS barcode: n.a. (alternative markers: BenA = AB248347; CaM = AB524047; RPB2 = n.a.).

Aspergillus oryzae (Ahlb.) Cohn, Jahresber. Schles. Ges. Vaterl. Cult. 61: 226. 1884. [MB184394]. Basionym: Eurotium oryzae Ahlb., Dingler's Polytechn. J. 230: 330. 1878. [MB225012]. — Type: IMI 16266. Ex-type: CBS 100925 = CBS 102.07 = NRRL 447 = ATCC 1011 = ATCC 12891 = ATCC 4814 = ATCC 7561 = ATCC 9102 = IAM13118 = IFO 4075 = IFO 537 = IFO 5375 = IMI 16266 = IMI 44242 = LSHBA c .19 = NCTC 598 = NRRL 692 = QM 6735 = Thom 113 = WB 447. Infragen. class: subgen. Circumdati, sect. Flavi, ser. Flavi. Reproduction: protoheterothallic; both MAT idiomorphs detected (Wada et al. 2012). ITS barcode: EF661560 (alternative markers: BenA = EF661483; CaM = EF661506; RPB2 = EF661438).

Aspergillus osmophilus Asgari & Zare, Mycoscience 55: 58. 2014. [MB803278]. — Type: IRAN 16110. Ex-type: IRAN 2090C = CBS 134258. Infragen. class: subgen. Aspergillus, sect. Aspergillus, ser. Xerophili. Reproduction: homothallic. ITS barcode: KC473921 (alternative markers: BenA = LT671127; CaM = LT671128; RPB2 = LT671129).

Aspergillus ostianus Wehmer, Bot. Centralbl. 80: 461. 1899. [MB179393]. — Type: IMI 15960. Ex-type: CBS 103.07 = CBS 548.65 = IBT 13386 = NRRL 420 = ATCC 16887 = IMI 015960iii = IMI 15960 = LCP 89.2584 = LSHBA c .35 = NCTC 3788 = QM 7460 = Thom 4724.35 = WB 420. Infragen. class: subgen. Circumdati, sect. Circumdati, ser. Circumdati. Reproduction: asexual. ITS barcode: EF661421 (alternative markers: BenA = EF661324; CaM = EF661385; RPB2 = EF661304).

Aspergillus pachycaulis F. Sklenář et al., Stud. Mycol. 88: 211. 2017. [MB823048]. — Type: PRM 944432. Ex-type: NRRL 25824 = CCF 5492 = DTO 356-D2 = IBT 34521 = IBT 34812. Infragen. class: subgen. Aspergillus, sect. Restricti, ser. Restricti. Reproduction: asexual. ITS barcode: KY087758 (alternative markers: BenA = KY117821; CaM = KY068308; RPB2 = KY117999).

Aspergillus pachycristatus Matsuzawa et al., Mycoscience 53: 439. 2012. [MB580944]. — Type: IFM 55265. Ex-type: IFM 55265 = NBRC 104790. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Nidulantes. Reproduction: homothallic. ITS barcode: n.a. (alternative markers: BenA = AB375875; CaM = AB524062; RPB2 = n.a.).

Aspergillus pallidofulvus Visagie et al., Stud. Mycol. 78: 40. 2014. [MB809199]. — Type: CBS H-21796. Ex-type: CBS 640.78 = NRRL 4789 = IBT 13871 = IFO 4095 = WB 4789. Infragen. class: subgen. Circumdati, sect. Circumdati, ser. Circumdati. Reproduction: asexual. ITS barcode: EF661423 (alternative markers: BenA = EF661328; CaM = EF661389; RPB2 = MN969085).

Aspergillus panamensis Raper & Thom, Mycologia 36: 568. 1944. [MB284311]. — Type: IMI 19393iii. Ex-type: CBS 120.45 = NRRL 1785 = ATCC 16797 = IMI 019393ii = IMI 019393iii = IMI 19393 = LSHBA .61 = NCTC 6974 = QM 6829 = QM 8897 = WB 1785. Infragen. class: subgen. Nidulantes, sect. Sparsi, ser. Conjuncti. Reproduction: asexual. ITS barcode: EF661177 (alternative markers: BenA = EF661109; CaM = EF661135; RPB2 = EF661040).

Aspergillus papuensis (Samson et al.) Samson et al., Stud. Mycol. 78: 155. 2014. [MB809593]. Basionym: Neosartorya papuensis Samson et al., Stud. Mycol. 59: 190. 2007. [MB505571]. — Type: CBS H-6277. Ex-type: CBS 841.96 = IBT 27801 = DTO 050-D1. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Neoglabri. Reproduction: homothallic. ITS barcode: EU220280 (alternative markers: BenA = AY870738; CaM = AY870697; RPB2 = MN969086).

Aspergillus parasiticus Speare, Bull. Hawaiian Sugar Planters Assoc. Exp. Sta. Pathol. Physiol. Ser. 12: 38. 1912. [MB191085]. — Type: IMI 15957ix. Ex-type: CBS 100926 = CBS 103.13 = NRRL 502 = ATCC 1018 = ATCC 6474 = ATCC 7865 = IMI 15957 = IMI 15957ii = IMI 15957iv = IMI 15957ix = IMI 15957vi = IMI 15957vii = LCP 89.2566 = LSHBA c 14 = NCTC 975 = NRRL 1731 = NRRL 3315 = NRRL A-13360 = NRRL A-14693 = Thom 3509 = WB 502. Infragen. class: subgen. Circumdati, sect. Flavi, ser. Flavi. Reproduction: Heterothallic (Horn et al. 2009b). ITS barcode: AY373859 (alternative markers: BenA = EF661481; CaM = AY017584; RPB2 = EF661449).

Aspergillus parvulus G. Sm., Trans. Brit. Mycol. Soc. 44: 45. 1961. [MB121074]. — Type: IMI 86558. Ex-type: CBS 136.61 = NRRL 4753 = ATCC 16911 = IMI 86558 = LSHBBB 405 = NRRL 1846 = QM 7955 = UC4613 = WB 4753. Infragen. class: subgen. Fumigati, sect. Cervini, ser. Cervini. Reproduction: asexual. ITS barcode: EF661269 (alternative markers: BenA = EF661247; CaM = EF661259; RPB2 = EF661233).

Aspergillus penicillioides Speg., Revista Fac. Agron. Univ. Nac. La Plata 2: 246. 1896. [MB309234]. — Type: IMI 211342. Ex-type: CBS 540.65 = NRRL 4548 = ATCC 16910 = IMI 211342 = IMUR540 = QM 9370 = WB 4548. Infragen. class: subgen. Aspergillus, sect. Restricti, ser. Penicillioides. Reproduction: asexual. ITS barcode: EF652036 (alternative markers: BenA = EF651928; CaM = EF652024; RPB2 = EF651930).

Aspergillus pepii Despot et al., Mycol. Prog. 16: 67. 2016 [2017]. [MB817073]. — Type: SZMC 23791. Ex-type: MFBF AV11051B IX = SZMC 22333 = CBS 142028. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Versicolores. Reproduction: asexual. ITS barcode: KU613368 (alternative markers: BenA = KU613371; CaM = KU613365; RPB2 = n.a.).

Aspergillus pernambucoensis Y. Horie et al., Mycoscience 55: 86. 2014. [MB801324]. — Type: IFM 61342H. Ex-type: IFM 61342 = JCM 19244 = CBS 137449 = DTO 316-G1. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Unilaterales. Reproduction: homothallic. ITS barcode: MN431373 (alternative markers: BenA = AB743856; CaM = AB743862; RPB2 = MN969087).

Aspergillus persii A.M. Corte & Zotti, Mycotaxon 83: 276 2002. [MB374215]. — Type: MUCL 41970. Ex-type: CBS 112795 = NRRL 35669 = IBT 22660 = MUCL 41970. Infragen. class: subgen. Circumdati, sect. Circumdati, ser. Sclerotiorum. Reproduction: asexual. ITS barcode: FJ491580 (alternative markers: BenA = AY819988; CaM = FJ491559; RPB2 = EF661295).

Aspergillus petersonii Jurjević & Hubka, Plant Syst. Evol. 301: 2454. 2015. [MB814440]. — Type: PRM 933841. Ex-type: CCF 4999 = NRRL 66216. Infragen. class: subgen. Circumdati, sect. Petersoniorum, ser. Petersoniorum. Reproduction: asexual. ITS barcode: LN849393 (alternative markers: BenA = LN849407; CaM = LN849422; RPB2 = LN849438).

Aspergillus peyronelii Sappa, Allionia 2: 248. 1955. [MB292855]. — Type: Plate 1, subfigures 1–4, in Sappa 1955, Allionia 2: 249 (lectotype); a dried herbarium specimen derived from the culture IMI 139271 (epitype, PRM 933831) (designated in Jurjević et al. 2015). Ex-epitype: IMI 139271 = CCF 4942 = NRRL 4754 = ATCC 16840. Infragen. class: subgen. Circumdati, sect. Petersoniorum, ser. Petersoniorum. Reproduction: asexual. ITS barcode: LN849398 (alternative markers: BenA = LN849412; CaM = LN849428; RPB2 = LN849443).

Aspergillus pipericola Frisvad et al., Stud. Mycol. 93: 46. 2019. [MB823774]. — Type: CBS H-23362. Ex-type: CBS 143680 = DTO 228-H4 = IBT 24628. Infragen. class: subgen. Circumdati, sect. Flavi, ser. Flavi. Reproduction: asexual. ITS barcode: MG662385 (alternative markers: BenA = MG517717; CaM = MG518087; RPB2 = MG517908).

Aspergillus piperis Samson & Frisvad, Stud. Mycol. 50: 57. 2004. [MB500009]. — Type: CBS H-13434. Ex-type: CBS 112811 = IBT 24630 = IBT 26239 = NRRL 62631. Infragen. class: subgen. Circumdati, sect. Nigri, ser. Nigri. Reproduction: protoheterothallic; unpublished (genome data, Vesth et al. 2018). ITS barcode: EU821316 (alternative markers: BenA = FJ629303; CaM = EU163267; RPB2 = KC796427).

Aspergillus pisce [as “pisci”] (A.D. Hocking & Pitt) Houbraken et al., Stud. Mycol. 78: 155. 2014. [MB812441]. Basionym: Polypaecilum pisce A.D. Hocking & Pitt, Mycotaxon 22: 200. 1985. [MB536436]. — Type: FRR 2732. Ex-type: FRR 2732 = ATCC 56982 = IMI 288726. Infragen. class: subgen. Polypaecilum, sect. Polypaecilum, ser. Polypaecilum. Reproduction: asexual. ITS barcode: MF362690 (alternative markers: BenA = MF362691; CaM = MN969231; RPB2 = JN121415).

Aspergillus pluriseminatus (Stchigel & Guarro) Samson et al., Stud. Mycol. 78: 155. 2014. [MB809595]. Basionym: Emericella pluriseminata Stchigel & Guarro, Mycologia 89: 937. 1997. [MB443124]. — Type: FMR 5588. Ex-type: CBS 100523 = FMR 5588 = IMI 370867 = DTO 011-H1. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Multicolores. Reproduction: homothallic; asexual morph unknown. ITS barcode: KU866566 (alternative markers: BenA = AY339989; CaM = EU443988; RPB2 = KU866937).

Aspergillus polyporicola Hubka et al., Mycologia 107: 194. 2015. [MB808145]. — Type: PRM 923452. Ex-type: NRRL 32683 = CCF 4553. Infragen. class: subgen. Circumdati, sect. Flavipedes, ser. Spelaei. Reproduction: asexual. ITS barcode: EF669595 (alternative markers: BenA = EU014088; CaM = EF669553; RPB2 = EF669637).

Aspergillus porosus A.J. Chen et al., Stud. Mycol. 88: 113. 2017. [MB818736]. — Type: CBS H-22822. Ex-type: CBS 141770 = DTO 262-D7 = IBT 34443. Infragen. class: subgen. Aspergillus, sect. Aspergillus, ser. Chevalierorum. Reproduction: homothallic. ITS barcode: LT670961 (alternative markers: BenA = LT671130; CaM = LT671131; RPB2 = LT671132).

Aspergillus porphyreostipitatus Visagie et al., Stud. Mycol. 78: 112. 2014. [MB809196]. — Type: CBS H-21813. Ex-type: CBS 138203 = DTO 266-D9. Infragen. class: subgen. Nidulantes, sect. Usti, ser. Usti. Reproduction: asexual. ITS barcode: KJ775564 (alternative markers: BenA = KJ775080; CaM = KJ775338; RPB2 = KU866987).

Aspergillus posadasensis Y. Marín et al., Int. J. Syst. Evol. Microbiol. 64: 2874. 2014. [MB803514]. — Type: CBS-H 21131. Ex-type: FMR 12168 = CBS 134259 = NBRC 109845. Infragen. class: subgen. Fumigati, sect. Clavati, ser. Clavati. Reproduction: homothallic; asexual morph unknown. ITS barcode: HG529483 (alternative markers: BenA = HG529481; CaM = HG529488; RPB2 = HF954977).

Aspergillus pragensis Hubka et al., Med. Mycol. 52: 570. 2014. [MB800371]. — Type: PRM 922702. Ex-type: CCF 3962 = CBS 135591 = NRRL 62491 = IBT 32274. Infragen. class: subgen. Circumdati, sect. Candidi, ser. Candidi. Reproduction: asexual. ITS barcode: FR727138 (alternative markers: BenA = HE661604; CaM = FR751452; RPB2 = LN849445).

Aspergillus proliferans G. Sm., Trans. Brit. Mycol. Soc. 26: 26. 1943. [MB284312]. — Type: IMI 16105iii. Ex-type: CBS 121.45 = NRRL 1908 = IMI 016105ii = IMI 016105iii = IMI 16105 = LSHB BB.82 = MUCL 15625 = NCTC 6546 = QM 7462 = UC 4303 = WB 1908. Infragen. class: subgen. Aspergillus, sect. Aspergillus, ser. Aspergillus. Reproduction: homothallic (Hubka et al. 2013a). ITS barcode: EF652064 (alternative markers: BenA = EF651891; CaM = EF651988; RPB2 = EF651941).

Aspergillus protuberus Munt.-Cvetk., Mikrobiologia 5: 119. 1968. [MB326650]. — Type: CBS 602.74. Ex-type: CBS 602.74 = NRRL 3505 = ATCC 18990 = QM 9804. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Versicolores. Reproduction: asexual. ITS barcode: EF652460 (alternative markers: BenA = EF652284; CaM = EF652372; RPB2 = EF652196).

Aspergillus pseudocaelatus Varga et al., Stud. Mycol. 69: 63. 2011. [MB560397]. — Type: CBS H-20632. Ex-type: CBS 117616 = DTO 010-H4. Infragen. class: subgen. Circumdati, sect. Flavi, ser. Kitamyces. Reproduction: protoheterothallic; MAT1-1-1 detected (Carvajal-Campos et al. 2017). ITS barcode: EF409242 (alternative markers: BenA = MG517626; CaM = MG517995; RPB2 = MG517809).

Aspergillus pseudodeflectus Samson & Mouch., Antonie van Leeuwenhoek 40: 345. 1975. [MB309236]. — Type: CBS 756.74. Ex-type: CBS 756.74 = NRRL 6135. Infragen. class: subgen. Nidulantes, sect. Usti, ser. Calidousti. Reproduction: asexual. ITS barcode: EF652507 (alternative markers: BenA = EF652331; CaM = EF652419; RPB2 = EF652243).

Aspergillus pseudoelegans Frisvad & Samson, Stud. Mycol. 50: 35. 2004. [MB500005]. — Type: CBS H-13439. Ex-type: CBS 112796 = DTO 077-F5 = NRRL 35670 = IBT 23402. Infragen. class: subgen. Circumdati, sect. Circumdati, ser. Steyniorum. Reproduction: asexual. ITS barcode: FJ491590 (alternative markers: BenA = EU014095; CaM = FJ491552; RPB2 = EF661281).

Aspergillus pseudoglaucus Blochwitz, Ann. Mycol. 27: 207. 1929. [MB275429]. — Type: IMI 016122ii. Ex-type: CBS 123.28 = NRRL 40 = ATCC 10066 = IMI 16122 = IMI 016122ii = LSHBA 19 = MUCL 15624 = QM 7463 = WB 40. Infragen. class: subgen. Aspergillus, sect. Aspergillus, ser. Rubri. Reproduction: homothallic. ITS barcode: EF652050 (alternative markers: BenA = EF651917; CaM = EF652007; RPB2 = EF651952).

Aspergillus pseudogracilis F. Sklenář et al., Stud. Mycol. 88: 216. 2017. [MB818932]. — Type: PRM 944434. Ex-type: CCF 5505 = EMSL No. 2765 = DTO 356-F3 = NRRL 66620 = IBT 34813. Infragen. class: subgen. Aspergillus, sect. Restricti, ser. Restricti. Reproduction: asexual. ITS barcode: KY087634 (alternative markers: BenA = KY117702; CaM = KY068186; RPB2 = KY117879).

Aspergillus pseudonomiae [as “pseudonomius”] Varga et al., Stud. Mycol. 69: 67. 2011. [MB833325]. — Type: CBS H-20633. Ex-type: CBS 119388 = DTO 009-F1 = NRRL 3353 = IBT 27864. Infragen. class: subgen. Circumdati, sect. Flavi, ser. Nomiarum. Reproduction: asexual. ITS barcode: AF338643 (alternative markers: BenA = EF661495; CaM = EF661529; RPB2 = EF661454).

Aspergillus pseudosclerotiorum J.P.Z. Siqueira et al., J. Clin. Microbiol. 55: 950. 2017. [MB818572]. — Type: CBS H-22808. Ex-type: UTHSCSA DI15-13 = FMR 14449 = CBS 141845. Infragen. class: subgen. Circumdati, sect. Circumdati, ser. Sclerotiorum. Reproduction: asexual. ITS barcode: LT574713 (alternative markers: BenA = LT574748; CaM = LT574783; RPB2 = LT574818).

Aspergillus pseudotamarii Yoko Ito et al., Mycol. Res. 105: 237. 2001. [MB466527]. — Type: BPI 746098. Ex-type: CBS 766.97 = DTO 046-C1 = NRRL 25517. Infragen. class: subgen. Circumdati, sect. Flavi, ser. Kitamyces. Reproduction: protoheterothallic; MAT 1-1-1 detected (Carvajal-Campos et al. 2017). ITS barcode: AF272574 (alternative markers: BenA = EF203125; CaM = EF202030; RPB2 = EU021631).

Aspergillus pseudoterreus S.W. Peterson et al., Stud. Mycol. 69: 53. 2011. [MB560396]. — Type: CBS H-20631. Ex-type: CBS 123890 = NRRL 4017. Infragen. class: subgen. Circumdati, sect. Terrei, ser. Terrei. Reproduction: asexual. ITS barcode: EF669598 (alternative markers: BenA = EF669523; CaM = EF669556; RPB2 = EF669640).

Aspergillus pseudoustus Frisvad et al., Stud. Mycol. 69: 91. 2011. [MB560403]. — Type: CBS H-20637. Ex-type: CBS 123904 = NRRL 5856 = IBT 28161 = DTO 083-G3. Infragen. class: subgen. Nidulantes, sect. Usti, ser. Usti. Reproduction: asexual. ITS barcode: FJ531147 (alternative markers: BenA = FJ531168; CaM = FJ531129; RPB2 = KU866978).

Aspergillus pseudoviridinutans Sugui et al., J. Clin. Microbiol. 52: 3709. 2014. [MB808637]. — Type: NRRL 62904. Ex-type: NRRL 62904 = NIH AV1 = CCF 5631 = DTO 304-I5. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Viridinutantes. Reproduction: protoheterothallic; both MAT idiomorphs detected (Hubka et al. 2018a). ITS barcode: MN431384 (alternative markers: BenA = KJ914690; CaM = KJ914708; RPB2 = MN969102).

Aspergillus pulvericola Visagie et al., Stud. Mycol. 78: 43. 2014. [MB809200]. — Type: CBS H-21793. Ex-type: CBS 137327 = DTO 267-C6. Infragen. class: subgen. Circumdati, sect. Circumdati, ser. Steyniorum. Reproduction: asexual. ITS barcode: KJ775440 (alternative markers: BenA = KJ775055; CaM = KJ775236; RPB2 = MN969088).

Aspergillus pulvinus Kwon-Chung & Fennell, Gen. Aspergillus: 455. 1965. [MB326651]. — Type: IMI 139628. Ex-type: CBS 578.65 = NRRL 5078 = ATCC 16842 = IMI 139628 = QM 8937 = WB 5078. Infragen. class: subgen. Cremei, sect. Cremei, ser. Pulvini. Reproduction: asexual. ITS barcode: EF652159 (alternative markers: BenA = EF652121; CaM = EF652139; RPB2 = EF652104).

Aspergillus puniceus Kwon-Chung & Fennell, Gen. Aspergillus: 547. 1965. [MB326652]. — Type: IMI 126692. Ex-type: CBS 495.65 = NRRL 5077 = ATCC 16800 = IMI 126692 = QM 9812 = WB 5077. Infragen. class: subgen. Nidulantes, sect. Usti, ser. Usti. Reproduction: asexual. ITS barcode: EF652498 (alternative markers: BenA = EF652322; CaM = EF652410; RPB2 = EF652234).

Aspergillus purpureocrustaceus Visagie, Stud. Mycol., this issue. 2020. [MB834205]. — Type: PREM 62264. Ex-type: PPRI 3840 = CMV 008B3. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Multicolores. Reproduction: asexual. ITS barcode: MK450653 (alternative markers: BenA = MK451138; CaM = MK451515; RPB2 = MK450806).

Aspergillus purpureus Samson & Mouch., Antonie van Leeuwenhoek 41: 350. 1975. [MB309237]. — Type: CBS 754.74. Ex-type: CBS 754.74 = NRRL 6133 = IMI 334937 = LCP 82.3323. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Aurantiobrunnei. Reproduction: homothallic. ITS barcode: EF652506 (alternative markers: BenA = EF652330; CaM = EF652418; RPB2 = EF652242).

Aspergillus puulaauensis Jurjević et al., IMA Fungus 3: 71. 2012. [MB800602]. — Type: BPI 880911. Ex-type: CBS 145750 = NRRL 35641 = DTO 225-G5. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Versicolores. Reproduction: asexual. ITS barcode: JQ301893 (alternative markers: BenA = JN853979; CaM = JN854034; RPB2 = JN853823).

Aspergillus qinqixianii Y. Horie et al., Mycoscience 41: 183. 2000. [MB464660]. — Type: CBM-FA-0866. Ex-type: IFM 55020 = CBM-FA-0866 = DTO 098-H6. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Stellati. Reproduction: homothallic. ITS barcode: KU866980 (alternative markers: BenA = AB524360; CaM = AB524051; RPB2 = KU866980).

Aspergillus qizutongii D.M. Li et al., Mycoscience 39: 301. 1998. [MB446576]. — Type: CBM FD-284. Ex-type: CBM FD-284. Infragen. class: subgen.: unknown, sect.: unknown, ser.: unknown. Reproduction: asexual. ITS barcode: n.a (alternative markers: BenA = n.a.; CaM = n.a.; RPB2 = n.a.).

Aspergillus quadricinctus E. Yuill, Trans. Brit. Mycol. Soc. 36: 57. 1953. [MB292857]. — Type: IMI 48583ii. Ex-type: CBS 135.52 = NRRL 2154 = ATCC 16897 = IMI 048583ii = IMI 48583 = QM 6874 = WB 2154 = DTO 050-E9 = DTO 164-I5. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Brevipedes. Reproduction: homothallic. ITS barcode: EF669947 (alternative markers: BenA = EF669806; CaM = EF669875; RPB2 = EF669735).

Aspergillus quadrilineatus Thom & Raper, Mycologia 31: 660. 1939. [MB275888]. — Type: IMI 89351. Ex-type: CBS 591.65 = NRRL 201 = ATCC 16816 = IMI 089351ii = IMI 89351 = LSHBA 546 = QM 7465 = Thom 4138.N8 = WB 201 = DTO 009-C7 = DTO 048-A9. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Nidulantes. Reproduction: homothallic. ITS barcode: EF652433 (alternative markers: BenA = EF652257; CaM = EF652345; RPB2 = EF652169).

Aspergillus raianus H.J. Chowdhery, Curr. Sci. 48: 953. 1979. [MB309239]. — Type: MLLU 110. Ex-type: n.a. Infragen. class: subgen.: unknown, sect.: unknown, ser.: unknown. Reproduction: asexual. ITS barcode: n.a. (alternative markers: BenA = n.a.; CaM = n.a.; RPB2 = n.a.).

Aspergillus rambellii Frisvad & Samson, Syst. Appl. Microbiol. 28: 449. 2005. [MB501209]. — Type: CBS 101887. Ex-type: CBS 101887 = ATCC 42001 = IBT 14580. Infragen. class: subgen. Nidulantes, sect. Ochraceorosei, ser. Ochraceorosei. Reproduction: asexual. ITS barcode: AJ874116 (alternative markers: BenA = JN217228; CaM = KU866700; RPB2 = JN121416).

Aspergillus raperi Stolk & J.A. Mey., Trans. Brit. Mycol. Soc. 40: 190. 1957. [MB292858]. — Type: [dried culture from soil] Zaire, Yangambi, Meyer (K). Ex-type: CBS 123.56 = NRRL 2641 = ATCC 16917 = IFO 6416 = IMI 70949 = NRRL 4778 = NRRL A-7462 = QM 1898 = WB 4221 = WB 4778. Infragen. class: subgen. Nidulantes, sect. Raperorum, ser. Raperorum. Reproduction: asexual. ITS barcode: EF652454 (alternative markers: BenA = EF652278; CaM = EF652366; RPB2 = EF652190).

Aspergillus recurvatus Raper & Fennell, Gen. Aspergillus: 529. 1965. [MB326653]. — Type: IMI 36528. Ex-type: CBS 496.65 = NRRL 4902 = ATCC 16809 = IMI 136528 = O-566 = QM 7972 = WB 4902. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Nidulantes. Reproduction: asexual. ITS barcode: EF652482 (alternative markers: BenA = EF652306; CaM = EF652394; RPB2 = EF652218).

Aspergillus restrictus G. Sm., J. Textile Inst. 22: 115. 1931. [MB276290]. — Type: IMI 16267. Ex-type: CBS 117.33 = CBS 541.65 = NRRL 154 = ATCC 16912 = B35855 = CECT2075 = IHEM 3920 = IMI 16267 = LSHBBB 94 = LSHTM 93 = MUCL 31313 = NCTC 6976 = NRRL 4155 = QM 1979 = Thom 5660.93 = UC4312 = VTTD-77065 = WB 154. Infragen. class: subgen. Aspergillus, sect. Restricti, ser. Restricti. Reproduction: asexual. ITS barcode: EF652042 (alternative markers: BenA = EF651880; CaM = EF652029; RPB2 = EF651978).

Aspergillus reticulatus F. Sklenář et al., Stud. Mycol. 88: 219. 2017. [MB818940]. — Type: PRM 944442. Ex-type: NRRL 25852 = CCF 5516 = DTO 356-D4 = IBT 34540. Infragen. class: subgen. Aspergillus, sect. Restricti, ser. Penicillioides. Reproduction: asexual. ITS barcode: KY087765 (alternative markers: BenA = KY117828; CaM = KY068315; RPB2 = KY118006).

Aspergillus rhizopodus J.N. Rai et al., Trans. Brit. Mycol. Soc. 64: 515. 1975. [MB309240]. — Type: Rai et al. 1975, Trans. Brit. Mycol. Soc. 64: p. 516, Fig. 1 (– lectotype designated here, MBT392286; CBS H-24284 [dried culture] – epitype designated here, MBT392287). Ex-epitype: CBS 450.75 = IMI 385057 = WB 5442 = NRRL 6136. Infragen. class: subgen. Fumigati, sect. Clavati, ser. Clavati. Reproduction: asexual. ITS barcode: EU078652 (alternative markers: BenA = EU076327; CaM = EF669926; RPB2 = MN969089).

Aspergillus robustus M. Chr. & Raper, Mycologia 70: 200. 1978. [MB309241]. — Type: NY WB 5286. Ex-type: CBS 428.77 = NRRL 6362 = ATCC 36106 = IMI 216610 = NRRL A-17351 = WB 5286. Infragen. class: subgen. Circumdati, sect. Robusti, ser. Robusti. Reproduction: asexual. ITS barcode: EF661176 (alternative markers: BenA = EU014101; CaM = EF661357; RPB2 = EF661033).

Aspergillus roseoglobulosus Frisvad & Samson, Stud. Mycol. 50: 30. 2004. [MB500001]. — Type: CBS H-13438. Ex-type: CBS 112800 = NRRL 4565 = IBT 14720. Infragen. class: subgen. Circumdati, sect. Circumdati, ser. Sclerotiorum. Reproduction: asexual. ITS barcode: FJ491583 (alternative markers: BenA = AY819984; CaM = FJ491555; RPB2 = EF661299).

Aspergillus ruber (Jos. König et al.) Thom & Church, Aspergillus: 112. 1926. [MB490579]. Basionym: Eurotium rubrum Jos. König et al., Z. Untersuch. Nahr. Genusm.: 726. 1901. [MB219613]. — Type: CBS 530.65. Ex-type: CBS 530.65 = NRRL 52 = ATCC 16441 = IMI 211380 = QM 1973 = Thom 5599B = WB 52. Infragen. class: subgen. Aspergillus, sect. Aspergillus, ser. Rubri. Reproduction: homothallic. ITS barcode: EF652066 (alternative markers: BenA = EF651920; CaM = EF652009; RPB2 = EF651947).

Aspergillus rugulosus Thom & Raper, Mycologia 31: 660. 1939. [MB277104]. — Type: IMI 136775. Ex-type: CBS 133.60 = NRRL 206 = ATCC 16820 = IMI 136775 = QM 1987 = Thom 4138.T11 = WB 206. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Nidulantes. Reproduction: homothallic. ITS barcode: EF652434 (alternative markers: BenA = EF652258; CaM = EF652346; RPB2 = EF652170).

Aspergillus saccharolyticus Sørensen et al., Int. J. Syst. Evol. Microbiol. 61: 3081. 2011. [MB518695]. — Type: Sørenson et al. 2011, Int. J. Syst. Evol. Microbiol. 61: p. 3081, Fig. 2 (– lectotype designated here, MBT392288; CBS H-24285 [dried culture] – epitype designated here, MBT392289). Ex-epitype: CBS 127449 = IBT 28509. Infragen. class: subgen. Circumdati, sect. Nigri, ser. Japonici. Reproduction: asexual. ITS barcode: HM853552 (alternative markers: BenA = HM853553; CaM = HM853554; RPB2 = HF559235).

Aspergillus salinarum [as “salinarus”] (Greiner et al.) Zalar & Greiner, Extremophiles 21: 762. 2017. [MB818567]. Basionym: Phialosimplex salinarum Greiner et al., IMA Fungus 5: 166. 2014. [MB809044]. — Type: CBS H-23061. Ex-type: CBS142047 = EXF-10247. Infragen. class: subgen. Polypaecilum, sect. Polypaecilum, ser. Salinarum. Reproduction: asexual. ITS barcode: KY980619 (alternative markers: BenA = KY980547; CaM = KY980583; RPB2 = KY980445).

Aspergillus salinicola Zalar et al., Stud. Mycol. 88: 221. 2017. [MB818941]. — Type: PRM 944448. Ex-type: EXF-10401 = IBT 34266 = CCF 5526 = NRRL 66621. Infragen. class: subgen. Aspergillus, sect. Restricti, ser. Penicillioides. Reproduction: asexual. ITS barcode: KY087722 (alternative markers: BenA = KY117785; CaM = KY068272; RPB2 = KY117963).

Aspergillus salisburgensis Zalar et al., Extremophiles 21:762. 2017. [MB818564]. — Type: CBS H-23061. Ex-type: EXF-10247 = CBS 142047 = DTO 410-E7. Infragen. class: subgen. Polypaecilum, sect. Polypaecilum, ser. Salinarum. Reproduction: asexual. ITS barcode: KX900623 (alternative markers: BenA = MN969414; CaM = MN969324; RPB2 = MN969191).

Aspergillus salwaensis Visagie et al., Stud. Mycol. 78: 49. 2014. [MB809201]. — Type: QCC F001/14. Ex-type: CBS 138172 = DTO 297-B3. Infragen. class: subgen. Circumdati, sect. Circumdati, ser. Sclerotiorum. Reproduction: asexual. ITS barcode: KJ775447 (alternative markers: BenA = KJ775056; CaM = KJ775244; RPB2 = MN969090).

Aspergillus savannensis A.J. Chen et al., Stud. Mycol. 84: 89. 2016. [MB816096]. — Type: CBS H-22495. Ex-type: CBS 140607 = IBT 23422 = DTO 059-H6. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Nidulantes. Reproduction: homothallic. ITS barcode: KU866581 (alternative markers: BenA = KU866818; CaM = KU866704; RPB2 = KU866959).

Aspergillus sclerotialis (W. Gams & Breton) Houbraken et al., Stud. Mycol. 78: 157. 2014. [MB809596]. Basionym: Sagenomella sclerotialis W. Gams & Breton, Persoonia 10: 109. 1978. [MB323039]. — Type: CBS 366.77. Ex-type: CBS 366.77 = IAM 14794 = DTO 107-E2 = DTO 137-F4. Infragen. class: subgen. Polypaecilum, sect. Polypaecilum, ser. Noonimiarum. Reproduction: asexual. ITS barcode: KF267869 (alternative markers: BenA = KY980579; CaM = MN969232; RPB2 = JN121505).

Aspergillus sclerotiicarbonarius Noonim et al., Int. J. Syst. Evol. Microbiol. 58: 1733. 2008. [MB504407]. — Type: Noonim et al. 2008, Int. J. Syst. Evol. Microbiol. 58: p. 1731, Fig. 2 (– lectotype designated here, MBT392290; CBS H-24286 [dried culture] – epitype designated here, MBT392291). Ex-epitype: CBS 121057 = IBT 121057. Infragen. class: subgen. Circumdati, sect. Nigri, ser. Carbonarii. Reproduction: Heterothallic (Darbyshir et al. 2013). ITS barcode: EU159216 (alternative markers: BenA = EU159229; CaM = EU159235; RPB2 = MN969091).

Aspergillus sclerotioniger Samson & Frisvad, Stud. Mycol. 50: 57. 2004. [MB500010]. — Type: CBS H-13433. Ex-type: CBS 115572 = IBT 22905. Infragen. class: subgen. Circumdati, sect. Nigri, ser. Carbonarii. Reproduction: protoheterothallic; unpublished (genome data, Vesth et al. 2018). ITS barcode: DQ900606 (alternative markers: BenA = FJ629304; CaM = FN594557; RPB2 = HE984369).

Aspergillus sclerotiorum G.A. Huber, Phytopathology 23: 306. 1933. [MB277707]. — Type: IMI 56673. Ex-type: CBS 549.65 = NRRL 415 = IBT 11931 = ATCC 16892 = DSM 870 = IFO 7542 = IMI 56732 = IMI 56673 = LCP 89.2594 = QM 6732 = Thom 5351 = WB 415. Infragen. class: subgen. Circumdati, sect. Circumdati, ser. Sclerotiorum. Reproduction: asexual. ITS barcode: EF661400 (alternative markers: BenA = EF661337; CaM = EF661384; RPB2 = EF661287).

Aspergillus seifertii Visagie, Stud. Mycol., this issue. 2020. [MB834206]. — Type: PREM 49066. Ex-type: PPRI 3211 = CMV 006F5. Infragen. class: subgen. Fumigati, sect. Clavati, ser. Clavati. Reproduction: asexual. ITS barcode: MK450647 (alternative markers: BenA = MK451093; CaM = MK451509; RPB2 = MK450800).

Aspergillus sergii P. Rodrigues, S.W. Peterson, Venâncio & N. Lima, Mycologia 104: 693. 2012. [MB561842]. — Type: MUM-H 10.219. Ex-type: CBS 130017 = DTO 223-C9 = DTO 223-D1. Infragen. class: subgen. Circumdati, sect. Flavi, ser. Flavi. Reproduction: protoheterothallic; MAT 1-2-1 detected (Soares et al. 2012). ITS barcode: JF412769 (alternative markers: BenA = MG517688; CaM = MG518059; RPB2 = HM802985).

Aspergillus serratalhadensis L.F. Oliveira et al., Persoonia 40: 263. 2018. [MB824978]. — Type: URM 91189. Ex-type: URM 7866. Infragen. class: subgen. Circumdati, sect. Nigri, ser. Japonici. Reproduction: asexual. ITS barcode: MH169127 (alternative markers: BenA = LT993222; CaM = LT993223; RPB2 = LT995971).

Aspergillus sesamicola Visagie et al., Stud. Mycol. 78: 52. 2014. [MB809202]. — Type: CBS H-21792. Ex-type: CBS 137324 = IBT 29314 = DTO 148-B4. Infragen. class: subgen. Circumdati, sect. Circumdati, ser. Circumdati. Reproduction: asexual. ITS barcode: KJ775437 (alternative markers: BenA = KJ775063; CaM = KJ775233; RPB2 = MN969092).

Aspergillus shendaweii (Yaguchi et al.) Samson et al., Stud. Mycol. 78: 157. 2014. [MB809597]. Basionym: Neosartorya shendaweii Yaguchi et al., Mycoscience 51: 260. 2010. [MB513151]. — Type: CBM FA-0958. Ex-type: IFM 57611 = CBS 128793 = DTO 148-G9. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Neoglabri. Reproduction: homothallic. ITS barcode: MN431374 (alternative markers: BenA = AB488754; CaM = AB488762; RPB2 = LC367696).

Aspergillus siamensis Manoch & Eamvijarn, Mycoscience 54: 403. 2013. [MB561946]. — Type: IFM 59793. Ex-type: IFM 59793 = KUFC 6349T = CCF 4685 = CBS 137452 = DTO 278-B6. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Viridinutantes. Reproduction: homothallic. ITS barcode: MN431375 (alternative markers: BenA = AB646989; CaM = AB776704; RPB2 = MN969093).

Aspergillus sigurros Visagie, Stud. Mycol., this issue. 2020. [MB834207]. — Type: PREM 62308. Ex-type: PPRI 15889 = CMV 005I4. Infragen. class: subgen. Nidulantes, sect. Usti, ser. Calidousti. Reproduction: asexual. ITS barcode: MK450650 (alternative markers: BenA = MK451066; CaM = MK451512; RPB2 = MK450803).

Aspergillus silvaticus Fennell & Raper, Mycologia 47: 83. 1955. [MB292859]. — Type: IMI 61456. Ex-type: CBS 128.55 = NRRL 2398 = ATCC 16843 = ATCC 46904 = IFO 8173 = IMI 61456 = NRRL A-3107 = QM 1912 = WB 2398. Infragen. class: subgen. Nidulantes, sect. Silvatici, ser. Silvatici. Reproduction: asexual. ITS barcode: EF652448 (alternative markers: BenA = EF652272; CaM = EF652360; RPB2 = EF652184).

Aspergillus similanensis Dethoup, Mycotaxon 131: 9. 2016. [MB810782]. — Type: BCC 75436. Ex-type: KUFA 0012 = KUFA 0013. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Fennelliarum. Reproduction: homothallic. ITS barcode: n.a. (alternative markers: BenA = KM095494; CaM = KC920701; RPB2 = n.a.).

Aspergillus sloanii Visagie et al., Stud. Mycol. 78: 108. 2014. [MB809194]. — Type: CBS H-21811. Ex-type: CBS 138177 = DTO 245A1. Infragen. class: subgen. Aspergillus, sect. Aspergillus, ser. Rubri. Reproduction: homothallic. ITS barcode: KJ775540 (alternative markers: BenA = KJ775074; CaM = KJ775309; RPB2 = KX463365).

Aspergillus sojae Sakag. & K. Yamada ex Murak., Rep. Res. Inst. Brewing: 8. 1971. [MB292860]. — Type: IMI 191300. Ex-type: CBS 100928 = DTO 046-C3 = IMI 191300. Infragen. class: subgen. Circumdati, sect. Flavi, ser. Flavi. Reproduction: protoheterothallic; MAT 1-1-1 detected (Ramirez-Prado et al. 2008). ITS barcode: KJ175434 (alternative markers: BenA = KJ175494; CaM = KJ175550; RPB2 = MG517831).

Aspergillus solicola Samson et al., Stud. Mycol. 78: 157. 2014 [MB809599]. Replaced synonym: Neosartorya warcupii S.W. Peterson et al., Stud. Mycol. 59: 201. 2007. [MB505572]. — Type: NRRL 35723. Ex-type: NRRL 35723 = DTO 047-E8. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Neoglabri. Reproduction: homothallic. ITS barcode: EU220279 (alternative markers: BenA = MN969370; CaM = EU220284; RPB2 = MN969104).

Aspergillus sparsus Raper & Thom, Mycologia 36: 572. 1944. [MB284314]. — Type: IMI 19394. Ex-type: CBS 139.61 = NRRL 1933 = ATCC 16851 = IHEM 4377 = IMI 19394 = IMI 19394ii = MUCL 31314 = NCTC 6975 = QM 7470 = WB 1933. Infragen. class: subgen. Nidulantes, sect. Sparsi, ser. Sparsi. Reproduction: asexual. ITS barcode: EF661181 (alternative markers: BenA = EF661125; CaM = EF661173; RPB2 = EF661071).

Aspergillus spathulatus Takada & Udagawa, Mycotaxon 24: 396. 1985. [MB104019]. — Type: NHL 2947. Ex-type: NRRL 20549 = ATCC 64222 = NHL 2948. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Spathulati. Reproduction: Heterothallic. ITS barcode: EF669943 (alternative markers: BenA = EF669803; CaM = EF669872; RPB2 = EF669731).

Aspergillus spectabilis M. Chr. & Raper, Mycologia 70: 333. 1978. [MB309243]. — Type: NY RMFH 429. Ex-type: CBS 429.77 = NRRL 6363 = ATCC 36105 = IMI 216611 = RMFH429. Infragen. class: subgen. Nidulantes, sect. Aenei, ser. Aenei. Reproduction: homothallic. ITS barcode: EF652510 (alternative markers: BenA = EU482437; CaM = EF652422; RPB2 = EF652246).

Aspergillus spelaeus A. Nováková et al., Mycologia 107: 194. 2015. [MB808146]. — Type: PRM 923462. Ex-type: CCF 4425 = CMF ISB 2615 = CBS 134371 = NRRL 62826. Infragen. class: subgen. Circumdati, sect. Flavipedes, ser. Spelaei. Reproduction: asexual. ITS barcode: HG915905 (alternative markers: BenA = HG916698; CaM = HG916741; RPB2 = HG916719).

Aspergillus spelunceus [as “speluneus”] Raper & Fennell, Gen. Aspergillus: 457. 1965. [MB326656]. — Type: IMI 211389. Ex-type: CBS 497.65 = NRRL 4989 = ATCC 16838 = IMI 211389 = NRRL A-3676 = QM 8898 = WB 4989. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Speluncei. Reproduction: asexual. ITS barcode: EF652490 (alternative markers: BenA = EF652314; CaM = EF652402; RPB2 = EF652226).

Aspergillus spinosus Kozak., Mycol. Pap. 161: 58. 1989. [MB127763]. — Type: IMI 211390. Ex-type: CBS 483.65 = NRRL 5034 = ATCC 16898 = IFO 8782 = IMI 211390 = NRRL A-1148 = QM 8888 = WB 5034 = DTO 050-D9. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Fumigati. Reproduction: homothallic. ITS barcode: EF669988 (alternative markers: BenA = EF669844; CaM = EF669914; RPB2 = EF669775).

Aspergillus spinulosporus Hubka et al., Plant Syst. Evol. 302: 1290. 2016. [MB816282]. Replaced synonym: Aspergillus nidulans var. echinulatus Fennell & Raper, Mycologia 47: 79 1955. [MB346543]. — Type: IMI 061454. Ex-type: CBS 120.55 = NRRL 2395 = ATCC 16825 = IMI 61454 = LCP 84.2557 = QM 1909 = WB 2395. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Nidulantes. Reproduction: homothallic. ITS barcode: EF652445 (alternative markers: BenA = EF652269; CaM = EF652357; RPB2 = EF652181).

Aspergillus stella-maris Zalar et al., Mycologia 100: 789. 2008. [MB507363]. — Type: CBS H-19887. Ex-type: CBS 113638 = IBT 23439 = DTO 011-A2. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Stellati. Reproduction: homothallic. ITS barcode: EU448269 (alternative markers: BenA = KU866886; CaM = EU443978; RPB2 = KU866925).

Aspergillus stellatus Curzi, Atti Reale Accad. Naz. Lincei, Rendiconti Cl. Sci. Fis.19: 428. 1934. [MB254841]. — Type: Bowenpilly near Secundarabad, s. coll., (K). Ex-type: CBS 598.65 = NRRL 1858 = ATCC 16819 = IMI 136778 = QM 6835 = WB 1858. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Stellati. Reproduction: homothallic. ITS barcode: EF652426 (alternative markers: BenA = EF652250; CaM = EF652338; RPB2 = EF652162).

Aspergillus stelliformis F. Sklenář et al., Mycologia 112: 363. 2020. [MB832711]. — Type: PRM 951569. Ex-type: CCF 5375 = EMSL No. 2293 = NRRL 66885. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Stellati. Reproduction: homothallic. ITS barcode: MK713531 (alternative markers: BenA = MK695638; CaM = MK695649; RPB2 = MK695660).

Aspergillus stercorarius A.J. Chen et al., Stud. Mycol. 84: 100. 2016. [MB816094]. — Type: CBS H-22496. Ex-type: CBS 428.93 = IBT 28024 = DTO 320-B3. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Nidulantes. Reproduction: homothallic; asexual morph unknown. ITS barcode: KU866625 (alternative markers: BenA = KU866865; CaM = KU866763; RPB2 = KU867026).

Aspergillus steynii Frisvad & Samson, Stud. Mycol. 50: 39. 2004. [MB500006]. — Type: CBS H-13445. Ex-type: CBS 112812 = NRRL 35675 = IBT 23096. Infragen. class: subgen. Circumdati, sect. Circumdati, ser. Steyniorum. Reproduction: asexual. ITS barcode: EF661416 (alternative markers: BenA = EF661347; CaM = EF661378; RPB2 = JN121428).

Aspergillus stramenius R.O. Novak & Raper, Gen. Aspergillus: 260. 1965. [MB326658]. — Type: IMI 172293. Ex-type: CBS 498.65 = NRRL 4652 = ATCC 16895 = IFO 9611 = IMI 172293 = QM 8900 = WB 4652 = DTO 046-E3. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Neoglabri. Reproduction: homothallic. ITS barcode: EF669984 (alternative markers: BenA = EF669840; CaM = EF669910; RPB2 = EF669771).

Aspergillus striatus J.N. Rai et al., Canad. J. Bot. 42: 1521. 1964. [MB326659]. — Type: IMI 96679. Ex-type: CBS 283.67 = CBS 592.65 = NRRL 4699 = ATCC 16815 = IMI 96679 = QM 8901 = WB 4699. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Nidulantes. Reproduction: homothallic; asexual morph unknown. ITS barcode: EF652470 (alternative markers: BenA = EF652294; CaM = EF652382; RPB2 = EF652206).

Aspergillus stromatoides Raper & Fennell, Gen. Aspergillus: 421. 1965. [MB326660]. — Type: IMI 123750. Ex-type: CBS 500.65 = DTO 059-B3 = DTO 080-G9 = NRRL 4519 = ATCC 16854 = ATCC 24485 = IMI 123750 = QM 8959 = QM 8974 = WB 4519. Infragen. class: subgen. Cremei, sect. Cremei, ser. Cremei. Reproduction: asexual, Chaetosartorya stromatoides B.J. Wiley & E.G. Simmons, a putative sexual morph of A. stromatiodes, is not conspecific with this species as recognised by Peterson (2008). ITS barcode: EF652146 (alternative markers: BenA = FJ531038; CaM = EF652127; RPB2 = EF652098).

Aspergillus subalbidus Visagie et al., Stud. Mycol. 78: 101. 2014. [MB809190]. — Type: CBS H-21807. Ex-type: CBS 567.65 = ATCC 16871 = IMI 230752 = NRRL 312 = DTO 045-D7 = DTO 045-D7. Infragen. class: subgen. Circumdati, sect. Candidi, ser. Candidi. Reproduction: asexual. ITS barcode: KJ866983 (alternative markers: BenA = MN969366; CaM = EF669551; RPB2 = MN969094).

Aspergillus subflavus Hubka et al., Stud. Mycol. 93: 46. 2019. [MB823776]. — Type: CBS H-23364. Ex-type: CBS 143683 = DTO 326-E8 = S778 = CCF 4957 = NRRL 66254 = IBT 34939. Infragen. class: subgen. Circumdati, sect. Flavi, ser. Flavi. Reproduction: asexual. ITS barcode: MH279429 (alternative markers: BenA = MG517773; CaM = MG518143; RPB2 = MG517964).

Aspergillus sublatus Y. Horie, Trans. Mycol. Soc. Japan 20: 481. 1979. [MB118407]. — Type: IFM 4553. Ex-type: CBS 140630 = DTO 338-F7 = IFO 30906 = IMI 334870 = NBRC 30906 = IFM 4553. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Nidulantes. Reproduction: homothallic. ITS barcode: KU866683 (alternative markers: BenA = KU866920; CaM = KU866804; RPB2 = KU867069).

Aspergillus sublevisporus Someya et al., Mycoscience 40: 405. 1999. [MB459822]. — Type: CBM PF-1207. Ex-type: CBS 128796 = IFM 53598 = DTO 148-H3. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Fennelliarum. Reproduction: homothallic. ITS barcode: MN431376 (alternative markers: BenA = AB488759; CaM = AB488767; RPB2 = MN969095).

Aspergillus subnutans A.J. Chen et al., Stud. Mycol. 85: 83. 2016. [MB817726]. — Type: CBS H-22728. Ex-type: CBS 129386 = DTO 202–C2 = WSF 445 = IBT 34352. Infragen. class: subgen. Fumigati, sect. Cervini, ser. Cervini. Reproduction: asexual. ITS barcode: KX528456 (alternative markers: BenA = KX528454; CaM = KX528455; RPB2 = KX528453).

Aspergillus subramanianii Visagie et al., Stud. Mycol. 78: 55. 2014. [MB809203]. — Type: CBS H-21791. Ex-type: CBS 138230 = NRRL 6161 = ATCC 18413. Infragen. class: subgen. Circumdati, sect. Circumdati, ser. Sclerotiorum. Reproduction: asexual. ITS barcode: EF661403 (alternative markers: BenA = EF661339; CaM = EF661397; RPB2 = EF661289).

Aspergillus subsessilis Raper & Fennell, Gen. Aspergillus: 530. 1965. [MB119551]. — Type: IMI 135820. Ex-type: CBS 502.65 = NRRL 4905 = ATCC 16808 = IMI 135820 = O-325 = QM 8035 = WB 4905. Infragen. class: subgen. Nidulantes, sect. Cavernicolarum, ser. Cavernicolarum. Reproduction: asexual. ITS barcode: EF652485 (alternative markers: BenA = EF652309; CaM = EF652397; RPB2 = EF652221).

Aspergillus subunguis Wadhwani et al., Curr. Sci. 53: 444. 1984. [MB105934]. — Type: IMI 254637. Ex-type: IMI 254637. Infragen. class: subgen. Nidulantes, sect.: unknown, ser.: unknown. Reproduction: asexual. ITS barcode: n.a. (alternative markers: BenA = n.a.; CaM = n.a.; RPB2 = n.a.).

Aspergillus subversicolor Jurjević et al., IMA Fungus 3: 72. 2012. [MB800603]. — Type: BPI 880918. Ex-type: CBS 145751 = NRRL 58999 = DTO 225-G9. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Versicolores. Reproduction: asexual. ITS barcode: JQ301894 (alternative markers: BenA = JN853970; CaM = JN854010; RPB2 = JN853799).

Aspergillus sulphureoviridis A.J. Chen et al., Stud. Mycol. 84: 103. 2016. [MB816097]. — Type: CBS H-22497. Ex-type: CBS 140626 = IBT 21868 = DTO 325-D1). Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Nidulantes. Reproduction: asexual. ITS barcode: KU866673 (alternative markers: BenA = KU866911; CaM = KU866793; RPB2 = KU867058).

Aspergillus suttoniae J.P.Z. Siqueira et al., Mycosis 61: 820. 2018. [MB823689]. — Type: CBS H-23243. Ex-type: UTHSCSA DI14-215 = FMR 13523. Infragen. class: subgen. Circumdati, sect. Flavipedes, ser. Flavipedes. Reproduction: asexual. ITS barcode: LT899487 (alternative markers: BenA = LT899536; CaM = LT899589; RPB2 = LT899644).

Aspergillus sydowii (Bainier & Sartory) Thom & Church, Aspergilli: 147. 1926. [MB279636]. Basionym: Sterigmatocystis sydowii Bainier & Sartory, Ann. Mycol. 11: 25. 1913. [MB197979]. — Type: IMI 211384. Ex-type: CBS 593.65 = NRRL 250 = IMI 211384 = NRRL 254. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Versicolores. Reproduction: protoheterothallic; MAT1-2-1 detected (de Vries et al. 2017). ITS barcode: EF652450 (alternative markers: BenA = EF652274; CaM = EF652362; RPB2 = EF652186).

Aspergillus tabacinus Nakaz. et al., J. Agric. Chem. Soc. Japan 10: 177. 1934. [MB539544]. — Type: Unknown. Ex-type: CBS 122718 = CBS H-24287 = NRRL 4791 = IFO 4098 = QM 9766 = WB 4791. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Versicolores. Reproduction: asexual. ITS barcode: EF652478 (alternative markers: BenA = EF652302; CaM = EF652390; RPB2 = EF652214).

Aspergillus taichungensis Yaguchi et al., Mycoscience 36: 421 1995. [MB434473]. — Type: CBM PF-1167. Ex-type: IBT 19404 = DTO 031-C6. Infragen. class: subgen. Circumdati, sect. Candidi, ser. Candidi. Reproduction: asexual. ITS barcode: MN431377 (alternative markers: BenA = MN969367; CaM = EU076310; RPB2 = MN969096).

Aspergillus takadae Y. Horie et al., Mycoscience 60: 358. 2019. [MB827072]. — Type: CBM-FA-929-3H. Ex-type: IFM 62979 = CBM-FA-929-1. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Spathulati. Reproduction: Heterothallic. ITS barcode: n.a. (alternative markers: BenA = LC367646; CaM = LC367657; RPB2 = LC367699).

Aspergillus takakii Y. Horie et al., Mycoscience 42: 91. 2001. [MB467818]. — Type: CBM FA-884. Ex-type: CBM FA-884 = IFM 53599 = CBS 137454 = DTO 278-B8. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Fumigati. Reproduction: homotallic. ITS barcode: MN431378 (alternative markers: BenA = AB787221; CaM = AB787566; RPB2 = MN969097).

Aspergillus tamarii Kita, Centralbl. Bakteriol. 2. Abth. 37: 433. 1913. [MB191425]. — Type: CBS 104.13. Ex-type: CBS 104.13 = NRRL 20818 = QM 9374. Infragen. class: subgen. Circumdati, sect. Flavi, ser. Kitamyces. Reproduction: protoheterothallic; MAT 1-1-1 detected (Ramirez-Prado et al. 2008). ITS barcode: AF004929 (alternative markers: BenA = EF661474; CaM = EF661526; RPB2 = EU021629).

Aspergillus tamarindosoli A.J. Chen et al., Stud. Mycol. 88: 123. 2017. [MB818737]. — Type: CBS H-22826. Ex-type: CBS 141775 = DTO 054-A8 = IBT 34432. Infragen. class: subgen. Aspergillus, sect. Aspergillus, ser. Tamarindosolorum. Reproduction: homothallic. ITS barcode: LT670981 (alternative markers: BenA = LT671191; CaM = LT671192; RPB2 = LT671193).

Aspergillus tanneri Kwon-Chung et al., J. Clin. Microbiol. 50: 3312. 2012. [MB801149]. — Type: BPI 882529. Ex-type: NRRL 62426 = NIH 1005 = ATCC MYA-4905. Infragen. class: subgen. Circumdati, sect. Tannerorum, ser. Tannerorum. Reproduction: asexual. ITS barcode: JN853798 (alternative markers: BenA = JN896582; CaM = JN896583; RPB2 = JN896585).

Aspergillus tapirirae C. Ram & A. Ram, Atti Reale Accad. Sci. Napoli 41: 100. 1972. [MB309245]. — Type: IMUFPe 2175. Ex-type: unknown. Infragen. class: subgen. Aspergillus, sect. Restricti, ser.: unknown. Reproduction: asexual. ITS barcode: n.a. (alternative markers: BenA = n.a.; CaM = n.a.; RPB2 = n.a.).

Aspergillus tardicrescens F. Sklenář et al., Stud. Mycol. 88: 221. 2017. [MB818942]. — Type: PRM 944439. Ex-type: DTO 316-B5 = CCF 5529 = IBT 34558 = NRRL 66623). Infragen. class: subgen. Aspergillus, sect. Restricti, ser. Penicillioides. Reproduction: asexual. ITS barcode: KY087710 (alternative markers: BenA = KY117772; CaM = KY068259; RPB2 = KY117951).

Aspergillus tardus Bissett & Widden, Canad. J. Bot. 62: 2521. 1984. [MB105071]. — Type: DAOM 183872. Ex-type: CBS 433.93 = DAOM 175187 (representative strain). Infragen. class: subgen. Cremei, sect. Cremei, ser. Inflati. Reproduction: asexual. ITS barcode: FJ531045 (alternative markers: BenA = FJ531001; CaM = FJ531084; RPB2 = n.a.).

Aspergillus tasmanicus Hubka et al., Plant Syst. Evol. 303: 801. 2017. [MB819519]. — Type: PRM 933840. Ex-type: CBS 283.66 = KACC 41141 = IBT 3211 = NBRC 8008. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Unilaterales. Reproduction: protoheterothallic; MAT 1-1-1 detected (Hubka et al. 2017). ITS barcode: AB185279 (alternative markers: BenA = AY685180; CaM = AY689367; RPB2 = LN874010).

Aspergillus tatenoi Y. Horie et al., Trans. Mycol. Soc. Japan 33: 395. 1992. [MB358433]. — Type: CBM-FA 0022. Ex-type: CBM-FA 0022 = CBS 407.93 = NRRL 4584 = DTO 046-E4. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Thermomutati. Reproduction: homothallic. ITS barcode: EF669982 (alternative markers: BenA = EF669838; CaM = EF669908; RPB2 = EF669769).

Aspergillus templicola Visagie et al., Stud. Mycol. 78: 103. 2014. [MB809191]. — Type: CBS H-21808. Ex-type: CBS 138181 = DTO 270-C6. Infragen. class: subgen. Circumdati, sect. Flavipedes, ser. Flavipedes. Reproduction: asexual. ITS barcode: KJ775545 (alternative markers: BenA = KJ775092; CaM = KJ775394; RPB2 = KP987017).

Aspergillus tennesseensis Jurjević et al., IMA Fungus 3: 73. 2012. [MB800604]. — Type: BPI 880917. Ex-type: CBS 145752 = NRRL 13150 = DTO 225-F5. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Versicolores. Reproduction: asexual. ITS barcode: JQ301895 (alternative markers: BenA = JN853976; CaM = JN854017; RPB2 = JN853806).

Aspergillus teporis A.J. Chen et al., Stud. Mycol. 88: 123. 2017. [MB818738]. — Type: CBS H-22821. Ex-type: CBS 141768 = DTO 058-E5 = IBT 34513. Infragen. class: subgen. Aspergillus, sect. Aspergillus, ser. Teporium. Reproduction: homothallic. ITS barcode: LT670982 (alternative markers: BenA = LT671194; CaM = LT671195; RPB2 = LT671196).

Aspergillus terreus Thom, Amer. J. Bot. 5: 85. 1918. [MB191719]. — Type: IMI 17294. Ex-type: CBS 601.65 = NRRL 255 = ATCC 10071 = ATCC 1012 = IFO 33026 = IMI 017294ii = IMI 17294 = JCM 10257 = LSHBA c .24 = MUCL 38640 = NCTC 981 = NRRL 543 = QM 1 = QM 1991 = Thom 144 = VKMF-67 = WB 255. Infragen. class: subgen. Circumdati, sect. Terrei, ser. Terrei. Reproduction: Heterothallic (Arabatzis & Velegraki 2013). ITS barcode: EF669586 (alternative markers: BenA = EF669519; CaM = EF669544; RPB2 = EF669628).

Aspergillus thailandensis Tanney et al., Stud. Mycol. 88: 255. 2017. [MB822734]. — Type: DAOM 745798. Ex-type: DAOMC 251755 = UAMH 11840 = CBS 143383 = KAS 8126 = SLOAN 4554 = PN10TH-749. Infragen. class: subgen. Polypaecilum, sect. Polypaecilum, ser. Noonimiarum. Reproduction: asexual. ITS barcode: KY980642 (alternative markers: BenA = KY980570; CaM = KY980606; RPB2 = KY980467).

Aspergillus thermomutatus (Paden) S.W. Peterson, Mycol. Res. 96: 549. 1992. [MB358403]. Basionym: Aspergillus fischeri var. thermomutatus Paden, Mycopathol. Mycol. Appl. 36: 161. 1968. [MB349035]. — Type: BPI 1108305. Ex-type: CBS 208.92 = NRRL 20748 = DTO 051-D7. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Thermomutati. Reproduction: homothallic. ITS barcode: EF669946 (alternative markers: BenA = EF669805; CaM = EF669874; RPB2 = EF669734).

Aspergillus thesauricus Hubka & A. Nováková, Int. J. Syst. Evol. Microbiol. 62: 2784. 2012. [MB564187]. — Type: PRM 860609. Ex-type: NRRL 62487 = CCF 4166 = CMFISB 2155. Infragen. class: subgen. Nidulantes, sect. Usti, ser. Calidousti. Reproduction: asexual. ITS barcode: HE615088 (alternative markers: BenA = HE615095; CaM = HE615120; RPB2 = HE615126).

Aspergillus togoensis (Henn.) Samson & Seifert, Adv. Pen. Asp. Syst.: 419. 1986 [1985]. [MB114720]. Basionym: Stilbothamnium togoense Henn., Bot. Jahrb. Syst. 23: 542. 1897. [MB374610]. — Type: BR B 1009. Ex-type: CBS 205.75 = NRRL 13551 = LCP 67.3456 (CBS 272.89 (representative strain)). Infragen. class: subgen. Circumdati, sect. Flavi, ser. Coremiiformes. Reproduction: asexual. ITS barcode: MN431379 (alternative markers: BenA = FJ491477; CaM = FJ491489; RPB2 = JN121479).

Aspergillus tonophilus Ohtsuki, Bot. Mag. (Tokyo) 75: 438. 1962. [MB326663]. — Type: IMI 108299. Ex-type: CBS 405.65 = NRRL 5124 = ATCC 16440 = ATCC 36504 = IMI 108299 = QM 8599 = WB 5124. Infragen. class: subgen. Aspergillus, sect. Aspergillus, ser. Rubri. Reproduction: homothallic. ITS barcode: EF652081 (alternative markers: BenA = EF651919; CaM = EF652000; RPB2 = EF651969).

Aspergillus transcarpathicus A.J. Chen et al., Stud. Mycol. 85: 83. 2016. [MB817727]. — Type: CBS H-22727. Ex-type: CBS 423.68 = DTO 022-C7 = IBT 22080 = IMI 134108 = VKM F-1331. Infragen. class: subgen. Fumigati, sect. Cervini, ser. Cervini. Reproduction: asexual. ITS barcode: FJ491624 (alternative markers: BenA = FJ491632; CaM = FJ491610; RPB2 = KX423680).

Aspergillus transmontanensis P. Rodrigues et al., Mycologia 104: 694. 2012. [MB561843]. — Type: MUM-H 10.214. Ex-type: DTO 223-C7 = CBS 130015. Infragen. class: subgen. Circumdati, sect. Flavi, ser. Flavi. Reproduction: protoheterothallic; both MAT idiomorphs detected (Soares et al. 2013). ITS barcode: JF412774 (alternative markers: BenA = HM803101; CaM = HM803020; RPB2 = HM802980).

Aspergillus trinidadensis Jurjević et al., IMA Fungus 3: 170. 2012. [MB802364]. — Type: BPI 883908. Ex-type: DTO 198-D1 = NRRL 62479 = ITEM 14821. Infragen. class: subgen. Circumdati, sect. Nigri, ser. Japonici. Reproduction: asexual. ITS barcode: MN431380 (alternative markers: BenA = HE984420; CaM = HE984434; RPB2 = HE984379).

Aspergillus trisporus S.C. Souza et al., Curr. Res. Environ. & Appl. Mycol. 9: 179. 2019. [MB822378]. — Type: CCDCA FI15. Ex-type: CML 3603. Infragen. class: subgen. Circumdati, sect. Janorum, ser. Janorum. Reproduction: asexual. ITS barcode: MF616388 (alternative markers: BenA = MF616387; CaM = MN013146; RPB2 = MF616389).

Aspergillus tritici [as “triticus”] B.S. Mehrotra & M. Basu, Nova Hedwigia 27: 599. 1976. [MB309248]. — Type: Mehrotra & Basu 1976, Nova Hedwigia 27: p. 603 Fig. 8 (– lectotype designated here, MBT392358; CBS H-24289 [dried culture] – epitype designated here, MBT392359). Ex-epitype: CBS 266.81 = DTO 031-F4. Infragen. class: subgen. Circumdati, sect. Candidi, ser. Candidi. Reproduction: asexual. ITS barcode: MN431381 (alternative markers: BenA = MN969368; CaM = MN969233; RPB2 = MN969098).

Aspergillus tsunodae (Yaguchi et al.) Samson et al., Stud. Mycol. 78: 157. 2014. [MB809600]. Basionym: Neosartorya tsunodae Yaguchi et al., Mycoscience 51: 261. 2010. [MB513152]. — Type: CBM FA-0950. Ex-type: IFM 57609 = NBRC 106416 = CBS 128794 = DTO 148-H1. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Unilaterales. Reproduction: homothallic. ITS barcode: HE974447 (alternative markers: BenA = AB488755; CaM = AB488763; RPB2 = HE974400).

Aspergillus tsurutae Y. Horie, Mycoscience 44: 399. 2003. [MB489534]. — Type: CBM FA-933. Ex-type: CBM FA-933 = CBS 137455 = IFM 56811 = DTO 279-D5. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Brevipedes. Reproduction: homothallic. ITS barcode: MN431382 (alternative markers: BenA = AB488760; CaM = AB488768; RPB2 = MN969099).

Aspergillus tubingensis Mosseray, La Cellule 43: 245. 1934. [MB255209]. — Type: Mosseray 1934, La Cellule 43: Pl. III Fig. 58 (– lectotype designated here, MBT392362; CBS H-24288 [dried culture] – epitype designated here, MBT392363). Ex-epitype: NRRL 4875 = QM 8904 = WB 4875 = CBS 133056. Infragen. class: subgen. Circumdati, sect. Nigri, ser. Nigri. Reproduction: Heterothallic (Horn et al. 2013). ITS barcode: EF661193 (alternative markers: BenA = EF661086; CaM = EF661151; RPB2 = EF661055).

Aspergillus tumidus J.P.Z. Siqueira et al., Persoonia 40: 261. 2018. [MB823690]. — Type: CBS H-23244. Ex-type: FMR 15743 = CBS 143587. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Multicolores. Reproduction: asexual. ITS barcode: LT903691 (alternative markers: BenA = LT903682; CaM = LT903685; RPB2 = LT903688).

Aspergillus turcosus S.B. Hong et al., Antonie van Leeuwenhoek 93: 97. 2008. [MB506378]. — Type: KACC 42091. Ex-type: KACC 42091 = DTO 035-E7. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Unilaterales. Reproduction: Heterothallic (Hubka et al. 2017). ITS barcode: MN431383 (alternative markers: BenA = DQ534143; CaM = DQ534148; RPB2 = HF545310).

Aspergillus turkensis Varga et al., Stud. Mycol. 69: 91. 2011. [MB560404]. — Type: CBS H-20638. Ex-type: CBS 504.65 = NRRL A-3261 = NRRL 4993 = ATCC 16799 = IMI 135420. Infragen. class: subgen. Nidulantes, sect. Usti, ser. Deflecti. Reproduction: asexual. ITS barcode: FJ531160 (alternative markers: BenA = FJ531191; CaM = FJ531145; RPB2 = EF652230).

Aspergillus udagawae Y. Horie et al., Mycoscience 36: 199. 1995. [MB412533]. — Type: CBM-FA-0711 (holotype); PRM 945579 (epitype, Hubka et al. 2018a). Ex-type: IFM 46972 = CBS 114217 = DTO 157-D7 = CBM-FA 0702 = KACC 41155 = CCF 4558. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Viridinutantes. Reproduction: Heterothallic. ITS barcode: AB185265 (alternative markers: BenA = LT796063; CaM = LT796064; RPB2 = LT796065).

Aspergillus undulatus H.Z. Kong & Z.T. Qi, Acta Mycol. Sin. 4: 211. 1985 [MB129004]. — Type: HMAS 47644. Ex-type: CBS 261.88 = DTO 011-A1. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Stellati. Reproduction: homothallic. ITS barcode: EU448275 (alternative markers: BenA = EF428363; CaM = EU443989; RPB2 = KU866928).

Aspergillus unguis (Émile-Weill & L. Gaudin) C.W. Dodge, Med. Mycol.: 637. 1935. [MB255264]. Basionym: Sterigmatocystis unguis Émile-Weill & L. Gaudin, Arch. Med. Exp. Anat. Pathol. 28: 463. 1918. [MB452891]. — Type: IMI 136526. Ex-type: CBS 132.55 = NRRL 2393 = ATCC 16812 = IMI 136526 = NRRL A-2391 = NRRL A-445 = QM 25B = WB 2393. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Unguium. Reproduction: asexual (?); contradictory (unpublished) data (Fennell & Raper 1955, Kakkar & Mehrotra 1971, Hubka et al. 2016a). ITS barcode: EF652443 (alternative markers: BenA = EF652267; CaM = EF652355; RPB2 = EF652179).

Aspergillus unilateralis Thrower, Austral. J. Bot. 2: 355. 1954. [MB292862]. — Type: IMI 62876. Ex-type: CBS 126.56 = NRRL 577 = ATCC 16902 = IFO 8136 = IMI 62876 = QM 8163 = WB 4366 = WB 4779 = DTO 001-E6 = DTO 050-F4. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Unilaterales. Reproduction: protoheterothallic; MAT 1-1-1 detected (Hubka et al. 2017). ITS barcode: EF669997 (alternative markers: BenA = EF669852; CaM = EF669923; RPB2 = EF669784).

Aspergillus urmiensis Arzanlou et al., Mycol. Prog. 15: 1089. 2016. [MB817474]. — Type: CBS H-22671. Ex-type: CCTU 742 = C B S 139558 = IBT 32593 = DTO 203-C2. Infragen. class: subgen. Circumdati, sect. Flavipedes, ser. Flavipedes. Reproduction: asexual. ITS barcode: KP987073 (alternative markers: BenA = KP987041; CaM = KP987056; RPB2 = KP987030).

Aspergillus ustus (Bainier) Thom & Church, Aspergilli: 152. 1926. [MB281216]. Basionym: Sterigmatocystis usta Bainier, Bull. Soc. Bot. France 28: 78. 1881. [MB536545]. — Type: IMI 211805. Ex-type: CBS 261.67 = NRRL 275 = ATCC 1041 = ATCC 16818 = IMI 211805 = QM 7477 = WB 275. Infragen. class: subgen. Nidulantes, sect. Usti, ser. Usti. Reproduction: asexual. ITS barcode: EF652455 (alternative markers: BenA = EF652279; CaM = EF652367; RPB2 = EF652191).

Aspergillus uvarum G. Perrone et al., Int. J. Syst. Evol. Microbiol. 58: 1036. 2008. [MB510962]. — Type: IMI 388523. Ex-type: CBS 121591 = IBT 26606 = IMI 388523 = ITEM 4834. Infragen. class: subgen. Circumdati, sect. Nigri, ser. Japonici. Reproduction: asexual. ITS barcode: AM745757 (alternative markers: BenA = AM745751; CaM = AM745755; RPB2 = HE984370).

Aspergillus vadensis R.P. de Vries et al., Antonie van Leeuwenhoek 87: 201. 2005. [MB340234]. — Type: CBS 113365. Ex-type: CBS 113365 = CECT20584 = IMI 313493. Infragen. class: subgen. Circumdati, sect. Nigri, ser. Nigri. Reproduction: protoheterothallic (genome data, Vesth et al. 2018). ITS barcode: AY585549 (alternative markers: BenA = AY585531; CaM = FN594560; RPB2 = HE984371).

Aspergillus vandermerwei Frisvad et al., Stud. Mycol. 93: 46. 2019. [MB823777]. — Type: CBS H-23381. Ex-type: CBS 612.78 = DTO 069-D2 = DTO 034-B5 = NRRL 5108 = CCF 5683 = IBT 13876. Infragen. class: subgen. Circumdati, sect. Flavi, ser. Alliacei. Reproduction: asexual. ITS barcode: EF661567 (alternative markers: BenA = EF661469; CaM = EF661540; RPB2 = MG517838).

Aspergillus varians Wehmer, Bot. Centralbl. 80: 460. 1899. [MB172782]. — Type: IMI 172297. Ex-type: CBS 505.65 = NRRL 4793 = ATCC 16836 = IFO 4114 = IMI 172297 = WB 4793. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Speluncei. Reproduction: asexual. ITS barcode: EF652479 (alternative markers: BenA = EF652303; CaM = EF652391; RPB2 = EF652215).

Aspergillus venenatus Jurjević et al., IMA Fungus 3: 73. 2012. [MB800605]. — Type: BPI 880916. Ex-type: CBS 145753 = NRRL 13147 = DTO 225-F4. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Versicolores. Reproduction: asexual. ITS barcode: JQ301896 (alternative markers: BenA = JN854003; CaM = JN854014; RPB2 = JN853803).

Aspergillus venezuelensis Frisvad & Samson, Syst. Appl. Microbiol. 27: 678. 2004. [MB368544]. — Type: CBS 868.97. Ex-type: CBS 868.97 = IBT 20956 = DTO 011-A4. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Stellati. Reproduction: homothallic. ITS barcode: AJ874119 (alternative markers: BenA = AY339998; CaM = EU443977; RPB2 = KU866931).

Aspergillus versicolor (Vuill.) Tirab., Ann. Bot. (Roma) 7: 9. 1908 [MB172159]. Basionym: Sterigmatocystis versicolor Vuill., Erreur Dét. Asp. Paras. Homme: 15. 1903. [MB233198]. — Type: CBS 583.65. Ex-type: CBS 583.65 = NRRL 238 = ATCC 9577 = IFO 33027 = IMI 229970 = JCM 10258 = QM 7478 = Thom 5519.57 = WB 238. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Versicolores. Reproduction: protoheterothallic; MAT1-2-1 detected (de Vries et al. 2017). ITS barcode: EF652442 (alternative markers: BenA = EF652266; CaM = EF652354; RPB2 = EF652178).

Aspergillus villosus F. Sklenář et al., Stud. Mycol. 88: 224. 2017. [MB818933]. — Type: PRM 944430. Ex-type: NRRL 25813 = CCF 5531 = DTO 356-C9 = IBT 34822. Infragen. class: subgen. Aspergillus, sect. Restricti, ser. Restricti. Reproduction: asexual. ITS barcode: KY087752 (alternative markers: BenA = KY117815; CaM = KY068302; RPB2 = KY117993).

Aspergillus vinosobubalinus Udagawa et al., Trans. Mycol. Soc. Japan 34: 255. 1993. [MB361186]. — Type: CBM BF-33501. Ex-type: CBM BF-33501. Infragen. class: subgen.: unknown, sect.: unknown, ser.: unknown. Reproduction: asexual. ITS barcode: n.a (alternative markers: BenA = n.a.; CaM = n.a.; RPB2 = n.a.).

Aspergillus violaceus Fennell & Raper, Mycologia 47: 75. 1955. [MB292863]. — Type: IMI 61449. Ex-type: CBS 138.55 = NRRL 2240 = ATCC 16813 = CECT2587 = IFO 8106 = IMI 061449ii = IMI 61449 = LCP 82.3318 = NRRL A-3156 = QM 1905 = UC4511 = WB 2240. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Nidulantes. Reproduction: homothallic. ITS barcode: EF652438 (alternative markers: BenA = EF652262; CaM = EF652350; RPB2 = EF652174).

Aspergillus viridicatenatus A.J. Chen et al., Stud. Mycol. 84: 112. 2016. [MB816088]. — Type: CBS H-22498. Ex-type: CBS 140629 = IBT 31492 = DTO 325-F4. Infragen. class: subgen. Nidulantes, sect. Nidulantes, ser. Speluncei. Reproduction: asexual. ITS barcode: KU866682 (alternative markers: BenA = KX423621; CaM = KU866802; RPB2 = KU867067).

Aspergillus viridinutans Ducker & Thrower, Austral. J. Bot. 2: 355. 1954. [MB292864]. — Type: IMI 62875. Ex-type: CBS 127.56 = NRRL 4365 = NRRL 4782 = NRRL 576 = NRRL A-16083 = NRRL A-6281 = ATCC 16901 = IMI 367415 = IMI 62875 = WB 4081 = WB 4365 = WB 4782 = DTO 050-F1. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Viridinutantes. Reproduction: protoheterothallic; MAT 1-1-1 detected (Nováková et al. 2014). ITS barcode: EF669978 (alternative markers: BenA = EF661252; CaM = DQ534162; RPB2 = EF669765).

Aspergillus vitricola [as “vitricolae”] Ohtsuki, Bot. Mag. (Tokyo) 75: 436. 1962. [MB326665]. — Type: No. 16 (Gi-4) (Herb. Nagao Institute). Ex-type: DTO 356-F7 = CBS H-24290 = CBS 146239 = NRRL 5125 = ATCC 16905 = ATCC 36505 = IMI 108298 = WB 5125. Infragen. class: subgen. Aspergillus, sect. Restricti, ser. Vitricolarum. Reproduction: asexual. ITS barcode: EF652046 (alternative markers: BenA = EF651927; CaM = EF652035; RPB2 = EF651973).

Aspergillus waksmanii Hubka et al., Int. J. Syst. Evol. Microbiol. 63: 786. 2013. [MB801063]. — Type: PRM 860537. Ex-type: NRRL 179 = CCF 4266 = Thom 4138.H52 = IBT 31900 = DTO 239-D8. Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Unilaterales. Reproduction: homothallic. ITS barcode: EF669934 (alternative markers: BenA = EF669794; CaM = EF669863; RPB2 = EF669722).

Aspergillus wangduanlii D.M. Li et al., Mycoscience 39: 302. 1998. [MB447107]. — Type: CBM FD-283. Ex-type: CBM FD-283 = CMMB 2309. Infragen. class: subgen.: unknown, sect.: unknown, ser.: unknown. Reproduction: asexual. ITS barcode: n.a. (alternative markers: BenA = n.a.; CaM = n.a.; RPB2 = n.a.).

Aspergillus waynelawii Tanney et al., Stud. Mycol. 88: 255. 2017. [MB822735]. — Type: DAOM 745796. Ex-type: DAOMC 251751 = UAMH 11926 = CBS 143384 = KAS 8123 = SLOAN 7951a = WL03MI-231. Infragen. class: subgen. Polypaecilum, sect. Polypaecilum, ser. Noonimiarum. Reproduction: asexual. ITS barcode: KY980639 (alternative markers: BenA = KY980567; CaM = KY980603; RPB2 = KY980464).

Aspergillus welwitschiae (Bres.) Henn., in Wehmer, Centralbl. Bakteriol. Parasitenk., 2. Abth. 18: 394. 1907. [MB490584]. Basionym: Ustilago welwitschiae Bres., in Saccardo, Bol. Soc. Brot. 11: 68. 1893. [MB176748]. — Type: CBS 139.54. Ex-type: CBS 139.54. Infragen. class: subgen. Circumdati, sect. Nigri, ser. Nigri. Reproduction: protoheterothallic; both MAT idiomorphs detected (Mageswari et al. 2016). ITS barcode: FJ629340 (alternative markers: BenA = MN969369; CaM = KC480196; RPB2 = MN969100).

Aspergillus wentii Wehmer, Centralbl. Bakteriol. Parasitenk., 2. Abth., 2: 149. 1896. [MB172623]. — Type: IMI 17295. Ex-type: CBS 104.07 = NRRL 375 = ATCC 1023 = IMI 17295 = NCTC 597 = NRRL 1269 = QM 7479 = Thom 116 = WB 375. Infragen. class: subgen. Cremei, sect. Cremei, ser. Wentiorum. Reproduction: protoheterothallic; MAT1-2-1 detected (de Vries et al. 2017). ITS barcode: EF652151 (alternative markers: BenA = EF652106; CaM = EF652131; RPB2 = EF652092).

Aspergillus westerdijkiae Frisvad & Samson, Stud. Mycol. 50: 30. 2004. [MB500000]. — Type: CBS H-13444. Ex-type: CBS 112803 = NRRL 3174 = IBT 10738 = ATCC 22947 = IBT 10738 = MUCL 39539. Infragen. class: subgen. Circumdati, sect. Circumdati, ser. Circumdati. Reproduction: asexual. ITS barcode: EF661427 (alternative markers: BenA = EF661329; CaM = EF661360; RPB2 = EF661307).

Aspergillus westlandensis Visagie et al., Stud. Mycol. 78: 59. 2014. [MB809204]. — Type: CBS H-21795. Ex-type: CBS 137321 = IBT 32139 = DTO 231-A9. Infragen. class: subgen. Circumdati, sect. Circumdati, ser. Circumdati. Reproduction: asexual. ITS barcode: KJ775434 (alternative markers: BenA = KJ775066; CaM = KJ775230; RPB2 = MN969101).

Aspergillus whitfieldii Tanney et al., Stud. Mycol. 88: 258. 2017. [MB822736]. — Type: DAOM 745799. Ex-type: DAOMC 251760 = UAMH 11842 = CBS 143385 = KAS 8129 = SLOAN 4178 = PN08TH-523. Infragen. class: subgen. Polypaecilum, sect. Polypaecilum, ser. Whitfieldiorum. Reproduction: asexual. ITS barcode: KY980645 (alternative markers: BenA = KY980573; CaM = KY980609; RPB2 = KY980470).

Aspergillus wisconsinensis A.J. Chen et al., Stud. Mycol. 85: 86. 2016. [MB817728]. — Type: CBS H-9203. Ex-type: CBS 413.64 = DTO 022–B1 = NRRL 5027 = IBT 22042 = IBT 22082 = WSF 380 = DTO 070-A5 = WB 5027. Infragen. class: subgen. Fumigati, sect. Cervini, ser. Cervini. Reproduction: asexual. ITS barcode: FJ491618 (alternative markers: BenA = FJ491638; CaM = FJ491609; RPB2 = KX423671).

Aspergillus wyomingensis A. Nováková et al., Fungal Diversity 64: 270. 2014. [MB803936]. — Type: PRM 861504. Ex-type: CCF 4417 = CMF ISB 2494 = CBS 135456 = DTO 332-B1 (= purified culture of DTO 311-F7). Infragen. class: subgen. Fumigati, sect. Fumigati, ser. Viridinutantes. Reproduction: Heterothallic. ITS barcode: HG324081 (alternative markers: BenA = HF933359; CaM = HF933397; RPB2 = HF937378).

Aspergillus xerophilus Samson & Mouch., Antonie van Leeuwenhoek 41: 348. 1975. [MB309251]. — Type: CBS 938.73. Ex-type: CBS 938.73 = NRRL 6131. Infragen. class: subgen. Aspergillus, sect. Aspergillus, ser. Xerophili. Reproduction: homothallic. ITS barcode: EF652085 (alternative markers: BenA = EF651923; CaM = EF651983; RPB2 = EF651970).

Aspergillus yunnanensis W.J. Cai et al., Mycoscience 61: 72. 2020. [MB831500]. — Type: HMAS 248248. Ex-type: CGMCC 3.19711. Infragen. class: subgen. Circumdati, sect. Janorum, ser. Janorum. Reproduction: asexual. ITS barcode: MN066373 (alternative markers: BenA = MN072909; CaM = MN072911; RPB2 = MN072913).

Aspergillus zutongqii A.J. Chen et al., Stud. Mycol. 88: 129. 2017. [MB818739]. — Type: CBS H-22824. Ex-type: CBS 141773 = CGMCC 3.13917 = DTO 349-E1 = IBT 34450. Infragen. class: subgen. Aspergillus, sect. Aspergillus, ser. Rubri. Reproduction: homothallic. ITS barcode: LT670986 (alternative markers: BenA = LT671206; CaM = LT671207; RPB2 = LT671208).



Dendrosphaera

Dendrosphaera eberhardtii Pat., Bull. Soc. Mycol. France 23: 69. 1907. [MB183425]. — Type: Patouillard 1907, Bull. Soc. Mycol. France 23: Pl. VIII, Fig. 1. (– lectotype designated here, MBT392299). Ex-type: n.a. Reproduction: homothallic. ITS barcode: n.a. (alternative markers: BenA = n.a.; CaM = n.a.; RPB2 = n.a.).



Dichlaena

Dichlaena indica A.B. Pawar et al., Geobios New Rep. 4: 66. 1985. [MB127024]. — Type: HCIO 32780. Ex-type: n.a. Reproduction: homothallic. ITS barcode: n.a. (alternative markers: BenA = n.a.; CaM = n.a.; RPB2 = n.a.).

Dichlaena lentisci Durieu & Mont., Exploration scientifique de l'Algérie 1: 405. 1849. [MB249716]. — Type: von Höhnel (FH). Ex-type: n.a. Reproduction: homothallic. ITS barcode: n.a. (alternative markers: BenA = n.a.; CaM = n.a.; RPB2 = n.a.).



Evansstolkia

Evansstolkia leycettana (H.C. Evans & Stolk) Houbraken et al., this study. 2020. [MB832558]. Basionym: Talaromyces leycettanus H.C. Evans & Stolk, Trans. Brit. Mycol. Soc. 56: 45. 1971. [MB324419]. — Type: CBS 398.68. Ex-type: CBS 398.68 = ATCC 22469 = IMI 178525 = JCM 12814 = NRRL 5178. Reproduction: homothallic. ITS barcode: AF454080 (alternative markers: BenA = GU092791; CaM = GU092837; RPB2 = EU021654).



Hamigera

Hamigera avellanea (Thom & Turesson) Stolk & Samson, Persoonia 6: 345. 1971. [MB314868]. Basionym: Penicillium avellaneum Thom & Turesson, Mycologia 7: 284. 1915. [MB248029]. — Type: CBS 295.48. Ex-type: CBS 295.48 = ATCC 10414 = CECT 2265 = DSM 2208 = IMI 040230 = NRRL 1938. Reproduction: homothallic. ITS barcode: AF454075 (alternative markers: BenA = EU021664; CaM = EU021682; RPB2 = EU021627).

Hamigera brevicompacta (H.Z. Kong) Samson et al., this study. 2020. [MB832579]. Basionym: Talaromyces brevicompactus H.Z. Kong, Mycosystema 18: 9. 1999. [MB460109]. — Type: HMAS 62770. Ex-type: CBS 102661 = AS 3.4676. Reproduction: homothallic. ITS barcode: MN431402 (alternative markers: BenA = MN969421; CaM = MN969342; RPB2 = MN969203).

Hamigera fusca S.W. Peterson et al., Mycologia 102: 857. 2010. [MB516020]. — Type: BPI 879307. Ex-type: DTO 194-D6 = CBS 132829 = NRRL 35601. Reproduction: homothallic. ITS barcode: GU092938 (alternative markers: BenA = GU092780; CaM = GU092813; RPB2 = GU111755).

Hamigera inflata S.W. Peterson et al., Mycologia 102: 854. 2010. [MB516017]. — Type: BPI 879308. Ex-type: NRRL 58014. Reproduction: homothallic. ITS barcode: GU092949 (alternative markers: BenA = GU092793; CaM = GU092823; RPB2 = GU092908).

Hamigera ingelheimensis (J.F.H. Beyma) S.W. Peterson, Mycology 5: 105. 2014. [MB807715]. Basionym: Penicillium ingelheimense J.F.H. Beyma, Antonie van Leeuwenhoek 8: 109. 1942. [MB289090]. — Type: IMI 234977. Ex-type: CBS 163.42 = DTO 027-G9 = FRR 2110 = IMI 234977 = NRRL 2110. Reproduction: asexual. ITS barcode: MN431403 (alternative markers: BenA = GU092756; CaM = GU092829; RPB2 = GU092912).

Hamigera insecticola S.W. Peterson et al., Mycologia 102: 852. 2010. [MB516016]. — Type: BPI 879309. Ex-type: DTO 194-D5 = CBS 132828 = NRRL 35386. Reproduction: homothallic. ITS barcode: EF634410 (alternative markers: BenA = GU092773; CaM = GU092816; RPB2 = GU111754).

Hamigera pallida S.W. Peterson et al., Mycologia 102: 856. 2010. [MB516019]. — Type: BPI 879310. Ex-type: DTO 194-D7 = CBS 132830 = NRRL 35718. Reproduction: homothallic. ITS barcode: GU092950 (alternative markers: BenA = GU092786; CaM = GU092824; RPB2 = GU111758).

Hamigera paravellanea S.W. Peterson et al., Mycologia 102: 852. 2010. [MB516015]. — Type: BPI 879311. Ex-type: DTO 194-D8 = CBS 132831 = NRRL 35720. Reproduction: homothallic. ITS barcode: GU092952 (alternative markers: BenA = GU092788; CaM = GU092826; RPB2 = GU092919).

Hamigera terricola S.W. Peterson et al., Mycologia 102: 855. 2010. [MB516018]. — Type: BPI 879312. Ex-type: DTO 194-D4 = CBS 132827 = NRRL 29055. Reproduction: homothallic. ITS barcode: GU092946 (alternative markers: BenA = GU092759; CaM = GU092811; RPB2 = GU111751).



Leiothecium

Leiothecium cristatum Y. Marín et al., Int. J. Syst. Evol. Microbiol. 64: 2873. 2014. [MB803513]. — Type: CBS-H 21130. Ex-type: FMR 11998 = CBS 134260 = NBRC 109843. Reproduction: homothallic. ITS barcode: KF732838 (alternative markers: BenA = n.a.; CaM = n.a.; RPB2 = HF954976).

Leiothecium ellipsoideum Samson & Mouch., Canad. J. Bot. 53: 1634. 1975. [MB316445]. — Type: CBS 607.74. Ex-type: CBS 607.74 = ATCC 32453. Reproduction: homothallic. ITS barcode: KF732839 (alternative markers: BenA = KY709178; CaM = KY611939; RPB2 = JN121541).



Monascus

Monascus argentinensis Stchigel & Guarro, Stud. Mycol. 50: 301. 2004. [MB500076]. — Type: FMR 6778. Ex-type: CBS 109402 = FMR 6778. Infragen. class: sect. Floridani. Reproduction: homothallic. ITS barcode: JF922046 (alternative markers: BenA = KY709174; CaM = KY611935; RPB2 = JN121423).

Monascus flavipigmentosus R.N. Barbosa et al., Stud. Mycol. 86: 43. 2017. [MB820072]. — Type: URM 90064. Ex-type: URM 7536 = CBS 142366 = DTO 353-A2. Infragen. class: sect. Floridani. Reproduction: homothallic. ITS barcode: KY511751 (alternative markers: BenA = KY709168; CaM = KY611929; RPB2 = MN969201).

Monascus floridanus P.F. Cannon & E.L. Barnard, Mycologia 79: 480. 1987. [MB132123]. — Type: IMI 282587. Ex-type: FLAS F54662 = CBS 142228 = CGMCC 3.5843 = BCRC 33310 = UAMH 4180. Infragen. class: sect. Floridani. Reproduction: homothallic. ITS barcode: KY635848 (alternative markers: BenA = KY709172; CaM = KY611933; RPB2 = KY611972).

Monascus lunisporas Udagawa & H. Baba, Cryptog. Mycol. 19: 270. 1998. [MB446999]. — Type: SUM 3116. Ex-type: CBS 142230 = CGMCC 3.7951 = ATCC 204397 = NBRC 33241 = BCRC 33640. Infragen. class: sect. Floridani. Reproduction: homothallic. ITS barcode: KY635847 (alternative markers: BenA = KY709171; CaM = KY611932; RPB2 = KY611971).

Monascus mellicola R.N. Barbosa et al., Stud. Mycol. 86: 44. 2017. [MB820073]. — Type: URM 90065. Ex-type: URM 7510 = CBS 142364 = DTO 350-E6. Infragen. class: sect. Floridani. Reproduction: asexual. ITS barcode: KY511726 (alternative markers: BenA = KY709143; CaM = KY611904; RPB2 = KY611943).

Monascus pallens P.F. Cannon et al., Mycol. Res. 99: 659. 1995. [MB413476]. — Type: IMI 356820. Ex-type: BSRA 10266 = CBS 142229 = CGMCC 3.5844 = ATCC 200612 = BCRC 33641. Infragen. class: sect. Floridani. Reproduction: homothallic. ITS barcode: KY635849 (alternative markers: BenA = KY709173; CaM = KY611934; RPB2 = KY611973).

Monascus purpureus Went, Ann. Sci. Nat., Bot., Sér. 8, 1: 1. 1895. [MB235390]. — Type: IMI 210765. Ex-type: CBS 109.07 = IF0 45 13 = ATCC 16426 = NRRL 1596 = FRR 1596. Infragen. class: sect. Rubri. Reproduction: homothallic. ITS barcode: KY635851 (alternative markers: BenA = KY709176; CaM = KY611937; RPB2 = JN121422).

Monascus recifensis R.N. Barbosa et al., Stud. Mycol. 86: 47. 2017. [MB820074]. — Type: URM 90066. Ex-type: URM 7524 = CBS 142365 = DTO 350-G6. Infragen. class: sect. Floridani. Reproduction: asexual. ITS barcode: KY511740 (alternative markers: BenA = KY709157; CaM = KY611918; RPB2 = KY611957).

Monascus ruber Tiegh., Bull. Soc. Mycol. France 31: 227. 1884. [MB234876]. — Type: IMI 81596. Ex-type: CBS 135.60 = IFO 8451 = ATCC 15670. Infragen. class: sect. Rubri. Reproduction: homothallic. ITS barcode: KY635850 (alternative markers: BenA = KY709175; CaM = KY611936; RPB2 = KY611974).



Paecilomyces

Paecilomyces brunneolus (N. Inagaki) Samson & Houbraken, Persoonia 22: 21. 2009. [MB512559]. Basionym: Paecilomyces variotii var. brunneolus N. Inagaki, Trans. Mycol. Soc. Japan 4: 3. 1962. [MB353669]. — Type: unknown. Ex-type: CBS 370.70 = DTO 093-D7 = IFO 7563. Reproduction: asexual. ITS barcode: EU037050 (alternative markers: BenA = EU037068; CaM = EU037033; RPB2 = MN969152).

Paecilomyces dactylethromorphus Bat. & H. Maia, Anais Soc. Biol. Pernambuco 15: 152. 1957. [MB302183]. — Type: IMUR 235. Ex-type: DTO 280-D1 = CBS 251.55 = ATCC 11971 = IMI 065752 = MUCL 9649. Reproduction: asexual. ITS barcode: FJ389951 (alternative markers: BenA = FJ390002; CaM = FJ389960; RPB2 = n.a.).

Paecilomyces divaricatus (Thom) Samson et al., Persoonia 22: 21. 2009. [MB512561]. Basionym: Penicillium divaricatum Thom, U.S.D.A. Bur. Ani. Ind. Bull. 118: 72. 1910. [MB170004]. — Type: Thom 1910, U.S.D.A. Bur. Animal Industr. Bull. 118: p. 73, Fig. 29. (– lectotype designated here, MBT392294; CBS 284.48 [metabolically inactive] – epitype designated here, MBT392295). Ex-epitype: DTO 093-D8 = CBS 284.48 = ATCC 10121 = ATCC 18502 = DSM 1961 = IAM 5001 = IMI 040025 = NBRC 100534 = NRRL 1115 = QM 6764 = VTT D-83214 = Thom 34. Reproduction: asexual. ITS barcode: FJ389931 (alternative markers: BenA = FJ389992; CaM = FJ389953; RPB2 = n.a.).

Paecilomyces formosus Sakag. et al. ex Houbraken & Samson, Persoonia 22: 21. 2009, nom. inval. [MB512562]. Basionym: Monilia formosa Sakag. et al., Zentralbl. Bakteriol., Abt. 2 100: 302. 1939 (nom. inval.) [MB252219]. — Type: CBS 990.73B. Ex-type: DTO 093-D2 = CBS 990.73B = ATCC 10865 = IMI 058427 = LSHB Pa31 = LSHB X26 = NRRL 1282. Reproduction: protoheterothallic (Heidarian et al. 2018). ITS barcode: FJ389929 (alternative markers: BenA = FJ389993; CaM = FJ389978; RPB2 = MN969154).

Paecilomyces fulvus Stolk & Samson, Persoonia 6: 354. 1971. [MB319107]. — Type: CBS 132.33. Ex-type: CBS 132.33 = 1MI 58.421. Reproduction: homothallic. ITS barcode: FJ389939 (alternative markers: BenA = FJ389988; CaM = FJ389957; RPB2 = n.a.).

Paecilomyces lagunculariae (C. Ram) Houbraken et al., this study. 2020. [MB832559]. Basionym: Byssochlamys nivea var. lagunculariae C. Ram, Nova Hedwigia 16: 311. 1968. [MB349108]. — Type: IMUFPe 2195. Ex-type: CBS 373.70. Reproduction: homothallic. ITS barcode: FJ389944 (alternative markers: BenA = FJ389995; CaM = FJ389965; RPB2 = MN969204).

Paecilomyces niveus Stolk & Samson, Persoonia 6: 351. 1971. [MB319117]. — Type: CBS 100.11. Ex-type: CBS 100.11 = ATCC 22260. Reproduction: homothallic. ITS barcode: FJ389934 (alternative markers: BenA = FJ389999; CaM = FJ389956; RPB2 = JF417414).

Paecilomyces tabacinus Jurjević et al., Persoonia 36: 409. 2016. [MB816870]. — Type: BPI 910044. Ex-type: CBS 141098 = DTO 412-B7 = CCF 5290. Reproduction: asexual. ITS barcode: LT548280 (alternative markers: BenA = MN969434; CaM = LT548288; RPB2 = MN969210).

Paecilomyces variotii Bainier, Bull. Soc. Mycol. France 23: 27. 1907. [MB248517]. — Type: unknown. Ex-type: DTO 032-I8 = DTO 280-D5 = CBS 102.74 = CECT 2803 = NRRL 1116. Reproduction: Heterothallic (Houbraken et al. 2008). ITS barcode: EU037055 (alternative markers: BenA = EU037073; CaM = EU037038; RPB2 = MN969153).

Paecilomyces zollerniae Stolk & Samson, Persoonia 6: 356. 1971. [MB319129]. — Type: CBS 374.70. Ex-type: CBS 374.70 = JCM 12808. Reproduction: homothallic. ITS barcode: FJ389933 (alternative markers: BenA = FJ390008; CaM = FJ389966; RPB2 = n.a.).



Penicillago

Penicillago kabunica (Baghd.) Houbraken et al., this study. 2020. [MB832560]. Basionym: Penicillium kabunicum Baghd., Novosti Sist. Nizsh. Rast. 5: 98. 1968. [MB335738]. — Type: CBS 409.68 (neotype). Ex-type: CBS 575.90 = CBS 409.69 (dead) = DTO 105-H9 = FRR 513 = IMI 140341 = VKM F-1072. Reproduction: asexual. ITS barcode: MN431415 (alternative markers: BenA = MN969438; CaM = MN969357; RPB2 = MN969217).

Penicillago mirabilis (Beliakova & Milko) Houbraken et al., this study. 2020. [MB832561]. Basionym: Penicillium mirabile Beliakova & Milko, Mikol. Fitopatol. 6: 145. 1972. [MB319286]. — Type: BKM F-1328. Ex-type: CBS 624.72 = DTO 304-C2 = CCRC 31665 = FRR 1959 = IMI 167383 = LCP 72.2193 = MUCL 31206 = VKM F-1328. Reproduction: asexual. ITS barcode: MN431416 (alternative markers: BenA = MN969439; CaM = MN969358; RPB2 = MN969218).

Penicillago moldavica (Milko & Beliakova) Houbraken et al., this study. 2020. [MB832562]. Basionym: Penicillium moldavicum Milko & Beliakova, Novosti Sist. Nizsh. Rast. 4: 255. 1967. [MB335751]. — Type: IMI 129966. Ex-type: CBS 574.90 = CBS 627.67 (dead) = DTO 041-H9 = ATCC 18355 = FRR 665 = IMI 129966 = VKM F-922. Reproduction: asexual. ITS barcode: MN431417 (alternative markers: BenA = MN969440; CaM = MN969359; RPB2 = MN969219).

Penicillago nodositata (Valla) Guevara-Suarez et al., Fungal Syst. Evol. 5: 64. 2020. [MB822074]. Basionym: Penicillium nodositatum Valla, Plant and Soil 114: 146. 1989. [MB126535]. — Type: Figs 1–4 in Valla et al. (1989) (lectotype, designated in Guevara-Suarez et al. 2020, MBT388228). Ex-epitype: CBS 333.90 = DTO 252-C7. Reproduction: asexual. ITS barcode: KC790403 (alternative markers: BenA = KC790399; CaM = MN969361; RPB2 = MN969220).



Penicilliopsis

Penicilliopsis africana Samson & Seifert, Adv. Pen. Asp. Syst.: 408. 1986 [1985]. [MB114759]. — Type: Metiquette Louis 6275 (BR). Ex-type: n.a. Reproduction: homothallic. ITS barcode: n.a. (alternative markers: BenA = n.a.; CaM = n.a.; RPB2 = n.a.).

Penicilliopsis clavariiformis Solms, Ann. Jard. Bot. Buitenzorg 6: 53. 1886. [MB120178]. — Type: Bot. Garden Bogor, Solms-Laubach in herb. Hauman (BR). Ex-type: n.a. Reproduction: homothallic. ITS barcode: MN431401 (alternative markers: BenA = MN969420; CaM = n.a.; RPB2 = EF669667).

Penicilliopsis pseudocordyceps H.M. Hsieh & Y.M. Ju, Mycologia 9: 541. 2002. [MB484663]. — Type: HAST (Taiwan) Hsieh & Ju 89112611. Ex-type: BCRC 33730. Reproduction: homothallic. ITS barcode: n.a. (alternative markers: BenA = n.a.; CaM = n.a.; RPB2 = n.a.).

Penicilliopsis zonata (Kwon-Chung & Fennell) Samson et al., Stud. Mycol. 85: 211. 2016. [MB819185]. Basionym: Aspergillus zonatus Kwon-Chung & Fennell, Gen. Aspergillus: 377. 1965. [MB326666]. — Type: WB 5079. Ex-type: DTO 022-B4 = CBS 506.65 = NRRL 5079 = ATCC 16867 = IFO 8817 = IMI 124936 = LCP 89.2588 = WB 5079. Reproduction: asexual. ITS barcode: EF669712 (alternative markers: BenA = EF669679; CaM = EF669701; RPB2 = EF669665).



Penicillium

Penicillium abidjanum Stolk, Antonie van Leeuwenhoek 34: 49. 1968. [MB335705]. — Type: CBS 246.67. Ex-type: CBS 246.67 = DTO 101-B4 = ATCC 18385 = FRR 1156 = IMI 136244. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Dalearum. Reproduction: homothallic. ITS barcode: GU981582 (alternative markers: BenA = GU981650; CaM = MN969234; RPB2 = JN121469).

Penicillium acidum Hyang B. Lee et al., Fungal Diversity 89: 173. 2018. [MB822167]. — Type: CNUFC-DLW4-1. Ex-type: JMRC SF:013659 = CNUFC-DLW4-1. Infragen. class: subgen. Aspergilloides, sect. Sclerotiorum, ser. Sclerotiorum. Reproduction: asexual. ITS barcode: KY587441 (alternative markers: BenA = KY587439; CaM = KY587442; RPB2 = KY587446).

Penicillium adametzii K.W. Zaleski, Bull. Int. Acad. Polon. Sci., Sér. B., Sci. Nat., 1927: 507. 1927. [MB119777]. — Type: IMI 39751. Ex-type: CBS 209.28 = ATCC 10407 = IMI 039751 = MUCL 29106 = NRRL 737. Infragen. class: subgen. Aspergilloides, sect. Sclerotiorum, ser. Adametziorum. Reproduction: asexual. ITS barcode: JN714929 (alternative markers: BenA = JN625957; CaM = KC773796; RPB2 = JN121455).

Penicillium adametzioides S. Abe ex G. Sm., Trans. Brit. Mycol. Soc. 46: 335. 1963. [MB302372]. — Type: IMI 068227. Ex-type: CBS 313.59 = ATCC 18306 = FAT1302 = IFO 6055 = IMI 068227 = NRRL 3405 = QM 7312. Infragen. class: subgen. Aspergilloides, sect. Sclerotiorum, ser. Adametziorum. Reproduction: asexual. ITS barcode: JN686433 (alternative markers: BenA = JN799642; CaM = JN686387; RPB2 = JN406578).

Penicillium aeris Visagie & Samson, Persoonia 36: 139. 2016. [MB808262]. — Type: CBS H-21608. Ex-type: CBS 135897 = DTO 207-D4. Infragen. class: subgen. Aspergilloides, sect. Torulomyces, ser. Torulomyces. Reproduction: asexual. ITS barcode: KF303654 (alternative markers: BenA = KF303614; CaM = KF303627; RPB2 = KF303681).

Penicillium alagoense L.O. Ferro et al., Persoonia 42: 447. 2019. [MB830760]. — Type: URM 93058. Ex-type: URM 8086. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Simplicissima. Reproduction: asexual. ITS barcode: MK804503 (alternative markers: BenA = MK802333; CaM = MK802336; RPB2 = MK802338).

Penicillium albocoremium (Frisvad) Frisvad, Int. Mod. Tax. Meth. Pen. Asp. Clas.: 275. 2000. [MB459817]. Basionym: Penicillium hirsutum var. albocoremium Frisvad, Mycologia 81: 856. 1990. [MB126411]. — Type: IMI 285511. Ex-type: CBS 472.84 = FRR 2931 = IBT 10682 = IBT 21502 = IMI 285511. Infragen. class: subgen. Penicillium, sect. Fasciculata, ser. Corymbifera. Reproduction: asexual. ITS barcode: AJ004819 (alternative markers: BenA = KU896812; CaM = KU896819; RPB2 = KU904344).

Penicillium alexiae Visagie et al., Persoonia 31: 59. 2013. [MB803785]. — Type: CBS H-21142. Ex-type: CBS 134558. Infragen. class: subgen. Aspergilloides, sect. Sclerotiorum, ser. Adametziorum. Reproduction: asexual. ITS barcode: KC790400 (alternative markers: BenA = KC773778; CaM = KC773803; RPB2 = KX961291).

Penicillium alfredii Visagie et al., Stud. Mycol. 78: 116. 2014. [MB809180]. — Type: CBS H-21800. Ex-type: CBS 138224 = DTO 269-A4. Infragen. class: subgen. Aspergilloides, sect. Alfrediorum, ser. Alfrediorum. Reproduction: asexual. ITS barcode: KJ775684 (alternative markers: BenA = KJ775177; CaM = KJ775411; RPB2 = KJ834520).

Penicillium allii Vincent & Pitt, Mycologia 81: 300. 1989. [MB125498]. — Type: MU Vincent 114. Ex-type: CBS 131.89 = IMI 321505 = NRRL 13630 = ATCC 64636 = IMI 321506 = IBT 6610. Infragen. class: subgen. Penicillium, sect. Fasciculata, ser. Corymbifera. Reproduction: asexual. ITS barcode: AJ005484 (alternative markers: BenA = AY674331; CaM = KU896820; RPB2 = KU904345).

Penicillium allii-sativi Frisvad et al., Persoonia 29: 89. 2012. [MB801873]. — Type: CBS H-21058. Ex-type: DTO 149-A8 = CBS 132074 = IBT 26507 = LJC 206. Infragen. class: subgen. Penicillium, sect. Chrysogena, ser. Chrysogena. Reproduction: asexual. ITS barcode: JX997021 (alternative markers: BenA = JX996891; CaM = JX996232; RPB2 = JX996627).

Penicillium alogum Visagie et al., Persoonia 36: 263. 2016. [MB815772]. — Type: DAOM 695759. Ex-type: DAOMC 250543 = CBS 140996 = DTO 410-E1 = IBT 23947= KAS 2475. Infragen. class: subgen. Aspergilloides, sect. Stolkia, ser. Stolkia. Reproduction: asexual. ITS barcode: KT887869 (alternative markers: BenA = KT887830; CaM = KT887791; RPB2 = MN969172).

Penicillium alutaceum D.B. Scott, Mycopathol. Mycol. Appl. 36: 17. 1968. [MB335708]. — Type: CBS 317.67. Ex-type: CBS 317.67 = ATCC 18542 = FRR 1158 = IFO 31728 = IMI 136243. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Alutacea. Reproduction: homothallic. ITS barcode: AF033454 (alternative markers: BenA = KJ834430; CaM = KP016768; RPB2 = JN121489).

Penicillium amaliae Visagie et al., Persoonia 31: 52. 2013. [MB803784]. — Type: CBS H-21141. Ex-type: CBS 134209 = CV 1875 = DTO 183F3 = DAOM 241034. Infragen. class: subgen. Aspergilloides, sect. Sclerotiorum, ser. Adametziorum. Reproduction: asexual. ITS barcode: JX091443 (alternative markers: BenA = JX091563; CaM = JX141557; RPB2 = KX961292).

Penicillium americanum Jurjević et al., Persoonia 42: 443. 2019. [MB830667]. — Type: BPI 910642. Ex-type: NRRL 66819 = EMSL1473 = ITEM 17520. Infragen. class: subgen. Penicillium, sect. Ramosum, ser. Soppiorum. Reproduction: asexual. ITS barcode: MK791278 (alternative markers: BenA = MK803427; CaM = MK803428; RPB2 = n.a.).

Penicillium amphipolaria Visagie et al., Persoonia 36: 269. 2016. [MB815777]. — Type: DAOM 695760. Ex-type: DAOMC 250551 = CBS 140997 = DTO 410-E2 = W 284 = KAS 2555. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Dalearum. Reproduction: asexual. ITS barcode: KT887872 (alternative markers: BenA = KT887833; CaM = KT887794; RPB2 = MN969177).

Penicillium anatolicum Stolk, Antonie van Leenwenhoek 34: 46. 1968. [MB335710]. — Type: CBS 479.66. Ex-type: CBS 479.66 = IBT 30764. Infragen. class: subgen. Aspergilloides, sect. Citrina, ser. Euglauca. Reproduction: homothallic. ITS barcode: AF033425 (alternative markers: BenA = JN606849; CaM = JN606571; RPB2 = JN606593).

Penicillium angulare S.W. Peterson et al., Mycologia 96: 1289. 2004. [MB487891]. — Type: BPI 842268. Ex-type: CBS 130293 = IBT 27051 = NRRL 28157. Infragen. class: subgen. Aspergilloides, sect. Sclerotiorum, ser. Adametziorum. Reproduction: asexual. ITS barcode: AF125937 (alternative markers: BenA = KC773779; CaM = KC773804; RPB2 = JN406554).

Penicillium angustiporcatum Takada & Udagawa, Trans. Mycol. Soc. Japan 24: 143. 1983. [MB108322]. — Type: NHL 6481. Ex-type: CBS 202.84. Infragen. class: subgen. Aspergilloides, sect. Gracilenta, ser. Angustiporcata. Reproduction: homothallic. ITS barcode: KC411690 (alternative markers: BenA = KJ834431; CaM = MN969235; RPB2 = JN406617).

Penicillium annulatum Visagie & K. Jacobs, Mycol. Prog. 14 (no. 96): 14. 2015. [MB809817]. — Type: CBS H-21333. Ex-type: CBS 135126 = CV 0037 = DTO 180-G7. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Rolfsiorum. Reproduction: asexual. ITS barcode: JX091426 (alternative markers: BenA = JX091514; CaM = JX141545; RPB2 = KF296410).

Penicillium antarcticum A.D. Hocking & C.F. McRae, Polar Biol. 21: 103. 1999. [MB482749]. — Type: DAR 72813. Ex-type: CBS 100492 = FRR 4989 = DTO 187-B3. Infragen. class: subgen. Penicillium, sect. Canescentia, ser. Atroveneta. Reproduction: asexual. ITS barcode: KJ834503 (alternative markers: BenA = MN969371; CaM = MN969236; RPB2 = JN406653).

Penicillium aotearoae Visagie & Seifert, Persoonia 36: 265. 2016. [MB815774]. — Type: PDD 107543. Ex-type: DAOMC 250538 = CBS 140999 = DTO 410-E4 = KAS 3088. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Lapidosa. Reproduction: asexual. ITS barcode: KT887874 (alternative markers: BenA = KT887835; CaM = KT887796; RPB2 = MN969174).

Penicillium apimei R.N. Barbosa et al., Antonie van Leeuwenhoek 111: 1891. 2018. [MB822208]. — Type: URM 90489. Ex-type: CBS 142502 = URM 7591. Infragen. class: subgen. Aspergilloides, sect. Gracilenta, ser. Macrosclerotiorum. Reproduction: asexual. ITS barcode: MF278310 (alternative markers: BenA = LT854641; CaM = LT882717; RPB2 = LT854650).

Penicillium aquaticum Hyang B. Lee et al., Fungal Diversity 89: 173. 2018. [MB822166]. — Type: CNUFC-YSW8-1. Ex-type: JMRC SF:013660. Infragen. class: subgen. Aspergilloides, sect. Gracilenta, ser. Macrosclerotiorum. Reproduction: asexual. ITS barcode: KY587453 (alternative markers: BenA = KY587450; CaM = KY587447; RPB2 = KY587449).

Penicillium arabicum Baghd., Novosti Sist. Nizsh. Rast. 5: 105. 1968. [MB335711]. — Type: T16 in Universitate Mosquensi (holotype); CBS H-7471 (isotype). Ex-type: CBS 414.69 = ATCC 22347 = DSM 2205 = FRR 507 = IMI 140335 = VKMF-1077. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Restricta. Reproduction: asexual. ITS barcode: KC411758 (alternative markers: BenA = KP016750; CaM = KP016770; RPB2 = KP064574).

Penicillium araracuaraense Houbraken, et al., Int. J. Syst. Evol. Microbiol. 61: 1469. 2011. [MB518025]. — Type: HUA 170334. Ex-type: CBS 113149 = DTO 056-D5 = DTO 297-H3 = IBT 23247. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Simplicissima. Reproduction: asexual. ITS barcode: GU981597 (alternative markers: BenA = GU981642; CaM = MN969237; RPB2 = KF296414).

Penicillium ardesiacum Novobr., Novosti Sist. Nizsh. Rast. 11: 228. 1974. [MB319257]. — Type: IMI 174719. Ex-type: CBS 497.73 = ATCC 24719 = FRR 1479 = IFO 30540 = IMI 174719 = VKMF-1749. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Pinetorum. Reproduction: asexual. ITS barcode: KM189565 (alternative markers: BenA = KM088805; CaM = KM089190; RPB2 = KM089577).

Penicillium argentinense Houbraken et al., Stud. Mycol. 70: 78. 2011. [MB563185]. — Type: CBS H-20461. Ex-type: CBS 130371 = IBT 30761. Infragen. class: subgen. Aspergilloides, sect. Citrina, ser. Euglauca. Reproduction: homothallic. ITS barcode: JN831361 (alternative markers: BenA = JN606815; CaM = JN606549; RPB2 = MN969105).

Penicillium arianeae Visagie et al., Persoonia 31: 59. 2013. [MB803786]. — Type: CBS H-21143. Ex-type: CBS 134559. Infragen. class: subgen. Aspergilloides, sect. Sclerotiorum, ser. Adametziorum. Reproduction: asexual. ITS barcode: KC773833 (alternative markers: BenA = KC773784; CaM = KC773811; RPB2 = KX961294).

Penicillium arizonense Frisvad et al., Sci. Rep. 6: 35112, 8. 2016. [MB817128]. — Type: C-F-101845. Ex-type: IBT 12289 = CBS 141311 = DTO 193-G8. Infragen. class: subgen. Penicillium, sect. Canescentia, ser. Canescentia. Reproduction: asexual. ITS barcode: MH492021 (alternative markers: BenA = MH492019; CaM = MH492020; RPB2 = MH492022).

Penicillium armarii Houbraken et al., Stud. Mycol. 78: 410. 2014. [MB809955]. — Type: CBS H-21870. Ex-type: CBS 138171 = DTO 235-F1. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Glabra. Reproduction: asexual. ITS barcode: KM189758 (alternative markers: BenA = KM089007; CaM = KM089394; RPB2 = KM089781).

Penicillium astrolabium R. Serra & S.W. Peterson, Mycologia 99: 80. 2007. [MB504766]. — Type: BPI 872160. Ex-type: CBS 122427 = NRRL 35611 = MUM 06.161. Infragen. class: subgen. Penicillium, sect. Brevicompacta, ser. Olsoniorum. Reproduction: asexual. ITS barcode: DQ645804 (alternative markers: BenA = DQ645793; CaM = DQ645808; RPB2 = JN406634).

Penicillium asymmetricum (Subram. & Sudha) Houbraken & Samson, Stud. Mycol. 70: 47. 2011. [MB561963]. Basionym: Thysanophora asymmetrica Subram. & Sudha, Kavaka 13: 88. 1987. [MB135502]. — Type: unknown. Ex-type: n.a. Infragen. class: subgen. Aspergilloides, sect. Thysanophora, ser. Thysanophora. Reproduction: asexual. ITS barcode: n.a. (alternative markers: BenA = n.a.; CaM = n.a.; RPB2 = n.a.).

Penicillium athertonense Houbraken, Stud. Mycol. 78: 412. 2014. [MB809956]. — Type: CBS H-21874. Ex-type: CBS 138161 = DTO 030-C2. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Pinetorum. Reproduction: asexual. ITS barcode: KM189462 (alternative markers: BenA = KM088690; CaM = KM089075; RPB2 = KM089462).

Penicillium atramentosum Thom, U.S.D.A. Bur. Animal Industr. Bull. 118: 65. 1910. [MB237291]. — Type: IMI 39752. Ex-type: CBS 291.48 = ATCC 10104 = FRR 795 = IBT 6616 = IFO 8137 = IMI 039752 = IMI 039752ii = LSHBP 1 = MUCL 29071 = MUCL 29126 = NRRL 795 = QM 7483. Infragen. class: subgen. Penicillium, sect. Paradoxa, ser. Atramentosa. Reproduction: asexual. ITS barcode: AF033483 (alternative markers: BenA = AY674402; CaM = KU896821; RPB2 = JN406584).

Penicillium atrofulvum Houbraken et al., Stud. Mycol. 70: 80. 2011. [MB563183]. — Type: CBS H-20650. Ex-type: CBS 109.66 = DTO 031-B2 = FRR 799 = IBT 30032 = IBT 29667. Infragen. class: subgen. Aspergilloides, sect. Citrina, ser. Westlingiorum. Reproduction: asexual. ITS barcode: JN617663 (alternative markers: BenA = JN606677; CaM = JN606387; RPB2 = JN606620).

Penicillium atrolazulinum Visagie & K. Jacobs, IMA Fungus 7: 91. 2016. [MB811001]. — Type: CBS H-22043. Ex-type: CBS 139136 = DAOMC 241083 = DTO 180-H4 = CV 55. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Corylophila. Reproduction: asexual. ITS barcode: JX140913 (alternative markers: BenA = JX141077; CaM = JX157416; RPB2 = KP064575).

Penicillium atrosanguineum B.X. Dong, Ceská Mycol. 27: 174. 1973. [MB319260]. — Type: PRC 1397 (holotype); CBS H-15524 (isotype). Ex-type: CBS 380.75 = FRR 1726 = IMI 197488. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Lapidosa. Reproduction: asexual. ITS barcode: JN617706 (alternative markers: BenA = KJ834435; CaM = KP016771; RPB2 = JN406557).

Penicillium atrovenetum G. Sm., Trans. Brit. Mycol. Soc. 39: 112. 1956. [MB302377]. — Type: IMI 061837. Ex-type: CBS 241.56 = ATCC 13352 = FRR 2571 = IFO 8138 = IMI 061837 = LSHBSm683 = QM 6963. Infragen. class: subgen. Penicillium, sect. Canescentia, ser. Atroveneta. Reproduction: asexual. ITS barcode: AF033492 (alternative markers: BenA = JX140944; CaM = KJ867004; RPB2 = JN121467).

Penicillium aurantiacobrunneum Houbraken et al., Stud. Mycol. 70: 80. 2011. [MB563206]. — Type: CBS H-20662. Ex-type: CBS 126228 = DTO 078-G2 = IBT 18753. Infragen. class: subgen. Aspergilloides, sect. Citrina, ser. Westlingiorum. Reproduction: asexual. ITS barcode: JN617670 (alternative markers: BenA = JN606702; CaM = MN969238; RPB2 = MN969106).

Penicillium aurantiogriseum Dierckx, Ann. Soc. Sci. Bruxelles 25: 88. 1901. [MB247956]. — Type: IMI 195050. Ex-type: CBS 324.89 = ATCC 48920 = FRR 971 = IBT 14016 = IMI 195050 = MUCL 29090 = NRRL 971. Infragen. class: subgen. Penicillium, sect. Fasciculata, ser. Viridicata. Reproduction: asexual. ITS barcode: AF033476 (alternative markers: BenA = MN969372; CaM = KU896822; RPB2 = JN406573).

Penicillium aurantioviolaceum Biourge, Cellule 33: 282. 1923. [MB257885]. — Type: CBS H-21954. Ex-type: CBS 137777 = NRRL 762 = ATCC 14974. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Thomiorum. Reproduction: asexual. ITS barcode: KM189756 (alternative markers: BenA = KM089005; CaM = KM089392; RPB2 = KM089779).

Penicillium austricola Visagie & K. Jacobs, Persoonia 36: 139. 2016. [MB805184]. — Type: CBS H-21605. Ex-type: CBS 135900 = CV 1842 = DTO 183-E6 = DAOMC 241066. Infragen. class: subgen. Aspergilloides, sect. Torulomyces, ser. Torulomyces. Reproduction: asexual. ITS barcode: JX091466 (alternative markers: BenA = JX091579; CaM = JX141600; RPB2 = KF303705).

Penicillium austroafricanum Houbraken & Visagie, Stud. Mycol. 78: 412. 2014. [MB809957]. — Type: CBS H-21864. Ex-type: CBS 137773 = DTO 133-G5. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Thomiorum. Reproduction: asexual. ITS barcode: KM189610 (alternative markers: BenA = KM088854; CaM = KM089241; RPB2 = KM089628).

Penicillium austrosinense L. Cai et al., Cladistics 35: 525. 2018 [2019]. [MB818164]. — Type: HMAS 247725. Ex-type: CGMCC 3.18797 = NN072318. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Dalearum. Reproduction: asexual. ITS barcode: KY495007 (alternative markers: BenA = KY495116; CaM = MN969328; RPB2 = KY495061).

Penicillium austrosinicum X.C. Wang & W.Y. Zhuang, Sci. Rep. 7: 8233, 3. 2017. [MB570338]. — Type: HMAS 248734. Ex-type: CGMCC 3.18410. Infragen. class: subgen. Aspergilloides, sect. Sclerotiorum, ser. Sclerotiorum. Reproduction: asexual. ITS barcode: KX885061 (alternative markers: BenA = KX885041; CaM = KX885051; RPB2 = KX885032).

Penicillium balearicum Guevara-Suarez et al., Fungal Syst. Evol. 5: 54. 2019 [2020]. [MB822061]. — Type: CBS H-23215. Ex-type: CBS 143044 = FMR 15191. Infragen. class: subgen. Penicillium, sect. Paradoxa, ser. Atramentosa. Reproduction: asexual. ITS barcode: LT899762 (alternative markers: BenA = LT898227; CaM = LT899758; RPB2 = LT899760).

Penicillium beceitense Guevara-Suarez et al., Fungal Syst. Evol. 5: 55. 2019 [2020] [MB822063]. — Type: CBS H-23183. Ex-type: CBS 142989 = FMR 15038. Infragen. class: subgen. Penicillium, sect. Ramosum, ser. Lanosa. Reproduction: asexual. ITS barcode: LT899780 (alternative markers: BenA = LT898229; CaM = LT899764; RPB2 = LT899798).

Penicillium bialowiezense K.W. Zaleski, Bull. Int. Acad. Polon. Sci., Sér. B., Sci. Nat. 1927: 450. 1927. [MB258429]. — Type: IMI 092237. Ex-type: CBS 227.28 = IBT 23044 = IMI 092237 = LSHBP 71. Infragen. class: subgen. Penicillium, sect. Brevicompacta, ser. Brevicompacta. Reproduction: asexual. ITS barcode: EU587315 (alternative markers: BenA = AY674439; CaM = AY484828; RPB2 = JN406604).

Penicillium biforme Thom, U.S.D.A. Bur. Animal Industr. Bull. 118: 54. 1910. [MB240764]. — Type: unknown. Ex-type: DTO 060-F9 = DTO 060-F8 = CBS 297.48 = ATCC 10416 = FRR 885 = IFO 7722 = IMI 039820 = LSHB P72 = MUCL 29165 = NRRL 885 = QM 7492. Infragen. class: subgen. Penicillium, sect. Fasciculata, ser. Camembertiorum. Reproduction: asexual. ITS barcode: KC411731 (alternative markers: BenA = MN969373; CaM = KU896823; RPB2 = KU904346).

Penicillium bilaiae Chalab., Bot. Mater. Otd. Sporov. Rast. 6: 165. 1950. [MB302379]. — Type: IMI 113677. Ex-type: CBS 221.66 = ATCC 22348 = ATCC 48731 = CCRC 31675 = FRR 3391 = IJFM 5025 = IMI 113677 = MUCL 31187 = VKMF-854. Infragen. class: subgen. Aspergilloides, sect. Sclerotiorum, ser. Adametziorum. Reproduction: asexual. ITS barcode: JN714937 (alternative markers: BenA = JN625966; CaM = JN626009; RPB2 = JN406610).

Penicillium bissettii Visagie & Seifert, Persoonia 36: 269. 2016. [MB815778]. — Type: DAOM 695761. Ex-type: DAOMC 167011 = CBS 140972 = KAS 1951. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Rolfsiorum. Reproduction: asexual. ITS barcode: KT887845 (alternative markers: BenA = KT887806; CaM = KT887767; RPB2 = MN969178).

Penicillium boreae S.W. Peterson & Sigler, Mycol. Res. 106: 1112. 2002. [MB483980]. — Type: BPI 841395. Ex-type: CBS 111717 = NRRL 31002 = UAMH 3896. Infragen. class: subgen. Aspergilloides, sect. Stolkia, ser. Stolkia. Reproduction: asexual. ITS barcode: AF481122 (alternative markers: BenA = JN617715; CaM = AF481138; RPB2 = MN969107).

Penicillium bovifimosum (Tuthill & Frisvad) Houbraken & Samson, Stud. Mycol. 70: 47. 2011. [MB561957]. Basionym: Eupenicillium bovifimosum Tuthill & Frisvad, Mycologia 94: 241. 2002. [MB456124]. — Type: WY RMF 82071. Ex-type: CBS 102825 = RMF 9598. Infragen. class: subgen. Penicillium, sect. Turbata, ser. Turbata. Reproduction: homothallic. ITS barcode: AF263347 (alternative markers: BenA = KJ834436; CaM = FJ530989; RPB2 = JN406649).

Penicillium brasilianum Bat., Anais Soc. Biol. Pernambuco 15: 162. 1957. [MB302381]. — Type: URM IMUR 56. Ex-type: CBS 253.55 = DTO 015-D3 = DTO 095-C4 = ATCC 12072 = FRR 3466 = QM 6947. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Simplicissima. Reproduction: asexual. ITS barcode: GU981577 (alternative markers: BenA = GU981629; CaM = MN969239; RPB2 = KF296420).

Penicillium brefeldianum B.O. Dodge, Mycologia 25: 92. 1933. [MB258851]. — Type: IMI 216896. Ex-type: CBS 235.81 = NRRL 710 = FRR 710 = IFO 31731 = IMI 216896 = LCP 89.2573 = LCP 89.2578 = MUCL 38762 = QM 1872 = Thom 5296. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Janthinella. Reproduction: homothallic. ITS barcode: AF033435 (alternative markers: BenA = GU981623; CaM = EU021683; RPB2 = KF296421).

Penicillium brevicompactum Dierckx, Ann. Soc. Sci. Bruxelles 25: 88. 1901. [MB149773]. — Type: IMI 40225. Ex-type: CBS 257.29 = ATCC 10418 = ATCC 9056 = DSM3825 = FRR 862 = IBT 23045 = IMI 040225 = LSHBP 75 = MUCL 28647 = MUCL 28813 = MUCL 28935 = MUCL 30240 = MUCL 30241 = MUCL 30256 = MUCL 30257 = NRRL 2011 = NRRL 862 = NRRL 864 = QM 7496. Infragen. class: subgen. Penicillium, sect. Brevicompacta, ser. Brevicompacta. Reproduction: asexual. ITS barcode: AY484912 (alternative markers: BenA = AY674437; CaM = AY484813; RPB2 = JN406594).

Penicillium brevistipitatum L. Wang & W.Y. Zhuang, Mycotaxon 93: 234. 2005. [MB356064]. — Type: HMAS 130354-1-4. Ex-type: DTO 105-I7 = CBS 122277 = AS 3.6887. Infragen. class: subgen. Penicillium, sect. Robsamsonia, ser. Robsamsonia. Reproduction: asexual. ITS barcode: DQ221696 (alternative markers: BenA = DQ221695; CaM = KU896824; RPB2 = JN406528).

Penicillium brocae S.W. Peterson et al., Mycologia 95: 143. 2003. [MB373658]. — Type: BPI 841763. Ex-type: CBS 116113 = IBT 26293 = NRRL 31472. Infragen. class: subgen. Aspergilloides, sect. Sclerotiorum, ser. Adametziorum. Reproduction: asexual. ITS barcode: AF484398 (alternative markers: BenA = KC773787; CaM = KC773814; RPB2 = JN406639).

Penicillium brunneoconidiatum Visagie et al., Stud. Mycol. 78: 415. 2014. [MB809958]. — Type: CBS H-21873. Ex-type: CBS 137732 = DTO 182-E4 = CV 949 = DAOM 241359. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Pinetorum. Reproduction: asexual. ITS barcode: KM189666 (alternative markers: BenA = KM088911; CaM = KM089298; RPB2 = KM089685).

Penicillium buchwaldii Frisvad & Samson, FEMS Microbiol. Lett. 339: 86. 2013. [MB800966]. — Type: IMI 304286. Ex-type: CBS 117181 = IBT 6005 = IMI 304286. Infragen. class: subgen. Penicillium, sect. Brevicompacta, ser. Buchwaldiorum. Reproduction: asexual. ITS barcode: JX313164 (alternative markers: BenA = MN969374; CaM = JX313148; RPB2 = JN406637).

Penicillium burgense Quintan. ex Visagie, IMA Fungus 7: 94. 2016. [MB816641]. — Type: CBS H-22567. Ex-type: CBS 325.89. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Lapidosa. Reproduction: asexual. ITS barcode: KC411736 (alternative markers: BenA = KJ834437; CaM = KP016772; RPB2 = JN406572).

Penicillium bussumense Houbraken, Stud. Mycol. 78: 415. 2014. [MB809959]. — Type: CBS H-21869. Ex-type: CBS 138160 = DTO 018-B2. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Glabra. Reproduction: asexual. ITS barcode: KM189458 (alternative markers: BenA = KM088685; CaM = KM089070; RPB2 = KM089457).

Penicillium cainii K.G. Rivera et al., Stud. Mycol. 70: 147. 2011. [MB563159]. — Type: DAOM 239914. Ex-type: CCFC 239914 = DTO 328-C1. Infragen. class: subgen. Aspergilloides, sect. Sclerotiorum, ser. Sclerotiorum. Reproduction: asexual. ITS barcode: JN686435 (alternative markers: BenA = JN686366; CaM = JN686389; RPB2 = MT156346).

Penicillium cairnsense Houbraken et al., Stud. Mycol. 70: 83. 2011. [MB563184]. — Type: CBS H-20686. Ex-type: CBS 124325 = DTO 030-E6 = IBT 29042. Infragen. class: subgen. Aspergilloides, sect. Citrina, ser. Westlingiorum. Reproduction: asexual. ITS barcode: JN617669 (alternative markers: BenA = JN606693; CaM = MN969240; RPB2 = MN969108).

Penicillium camemberti Thom, U.S.D.A. Bur. Animal Industr. Bull. 82: 33. 1906. [MB175171]. — Type: IMI 27831. Ex-type: DTO 246-F1 = CBS 299.48 = ATCC 1105 = ATCC 4845 = FRR 878 = IBT 21508 = IMI 027831 = IMI 092200 = LCP 66.584 = LSHBP 11 = MUCL 29790 = NCTC 582 = NRRL 877 = NRRL 878. Infragen. class: subgen. Penicillium, sect. Fasciculata, ser. Camembertiorum. Reproduction: asexual. ITS barcode: AB479314 (alternative markers: BenA = FJ930956; CaM = KU896825; RPB2 = MN969109).

Penicillium camponotum Visagie et al., Persoonia 36: 271. 2016. [MB815779]. — Type: DAOM 695762. Ex-type: DAOMC 250557 = CBS 140982 = NBBR-2-1 = W 471 = KAS 2177. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Rolfsiorum. Reproduction: asexual. ITS barcode: KT887855 (alternative markers: BenA = KT887816; CaM = KT887777; RPB2 = MN969179).

Penicillium canariense S.W. Peterson & Sigler, Mycol. Res. 106: 1113. 2002. [MB483981]. — Type: BPI 841396. Ex-type: CBS 111720 = NRRL 31003 = IJFM 536 = UAMH 6403. Infragen. class: subgen. Aspergilloides, sect. Stolkia, ser. Stolkia. Reproduction: asexual. ITS barcode: AF481121 (alternative markers: BenA = JN617714; CaM = AF481137; RPB2 = MN969110).

Penicillium canescens Sopp, Skr. Vidensk.-Selsk. Christiana Math.-Nat. Kl. 11: 181. 1912. [MB153765]. — Type: IMI 28260. Ex-type: CBS 300.48 = ATCC 10419 = DSM1215 = FRR 910 = IMI 028260 = MUCL 29169 = NCTC 6607 = NRRL 910 = QM 7550 = VKMF-1148. Infragen. class: subgen. Penicillium, sect. Canescentia, ser. Canescentia. Reproduction: asexual. ITS barcode: AF033493 (alternative markers: BenA = JX140946; CaM = MN969241; RPB2 = JN121485).

Penicillium canis S.W. Peterson, J. Clin. Microbiol. 52: 2450. 2014. [MB807056]. — Type: BPI 892763. Ex-type: NRRL 62798. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Erubescentia. Reproduction: asexual. ITS barcode: KJ511291 (alternative markers: BenA = KF900167; CaM = KF900177; RPB2 = KF900196).

Penicillium cantabricum Visagie & Samson, Persoonia 36: 142. 2016. [MB808263]. — Type: CBS H-21612. Ex-type: CBS 120415 = DTO 076-I9 = FMR 9121. Infragen. class: subgen. Aspergilloides, sect. Torulomyces, ser. Torulomyces. Reproduction: asexual. ITS barcode: KF303655 (alternative markers: BenA = KF303615; CaM = KF303646; RPB2 = KF303682).

Penicillium caperatum Udagawa & Y. Horie, Trans. Mycol. Soc. Japan 14: 371. 1973. [MB319262]. — Type: NHL 6454. Ex-type: CBS 443.75 = DTO 101-B2 = ATCC 28046 = DSM2209 = NHL 6465. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Janthinella. Reproduction: homothallic. ITS barcode: KC411761 (alternative markers: BenA = GU981660; CaM = MN969242; RPB2 = KF296422).

Penicillium caprifimosum Guevara-Suarez et al., Fungal Syst. Evol. 5: 55. 2019 [2020]. [MB822064]. — Type: CBS H-23184. Ex-type: CBS 142990 = FMR 15041. Infragen. class: subgen. Penicillium, sect. Turbata, ser. Turbata. Reproduction: asexual. ITS barcode: LT899781 (alternative markers: BenA = LT898238; CaM = LT899765; RPB2 = LT899799).

Penicillium capsulatum Raper & Fennell, Mycologia 40: 528. 1948. [MB289079]. — Type: IMI 40576. Ex-type: CBS 301.48 = ATCC 10420 = DSM2210 = FRR 2056 = IJFM 5120 = IMI 040576 = NRRL 2056 = QM 4869 = VKMF-445. Infragen. class: subgen. Aspergilloides, sect. Ramigena, ser. Ramigena. Reproduction: asexual. ITS barcode: AF033429 (alternative markers: BenA = MN969375; CaM = KP735539; RPB2 = JN406582).

Penicillium carneum (Frisvad) Frisvad, Microbiology 142: 546. 1996. [MB415652]. Basionym: Penicillium roqueforti var. carneum Frisvad, Mycologia 81: 858. 1990. [MB126415]. — Type: IMI 293204. Ex-type: CBS 112297 = IBT 6884 = IBT 18419 = IMI 293204. Infragen. class: subgen. Penicillium, sect. Roquefortorum, ser. Roquefortorum. Reproduction: asexual. ITS barcode: HQ442338 (alternative markers: BenA = AY674386; CaM = HQ442322; RPB2 = JN406642).

Penicillium cartierense Houbraken, Stud. Mycol. 78: 415. 2014. [MB809960]. — Type: CBS H-21861. Ex-type: CBS 137956 = DTO 092-H9. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Thomiorum. Reproduction: asexual. ITS barcode: KM189564 (alternative markers: BenA = KM088804; CaM = KM089189; RPB2 = KM089576).

Penicillium caseifulvum Lund et al., J. Food Mycol. 1: 97. 1998. [MB446013]. — Type: C 24999. Ex-type: DTO 145-B8 = CBS 101134 = IBT 18282 = IBT 21510. Infragen. class: subgen. Penicillium, sect. Fasciculata, ser. Camembertiorum. Reproduction: asexual. ITS barcode: KJ834504 (alternative markers: BenA = AY674372; CaM = KU896826; RPB2 = KU904347).

Penicillium catalonicum Visagie & Samson, Persoonia 36: 142. 2016. [MB808265]. — Type: CBS H-21610. Ex-type: CBS 110532 = DTO 078-H5. Infragen. class: subgen. Aspergilloides, sect. Torulomyces, ser. Torulomyces. Reproduction: asexual. ITS barcode: KF303650 (alternative markers: BenA = KF303609; CaM = KF303644; RPB2 = KF303683).

Penicillium cataractarum Visagie et al., Persoonia 36: 271. 2016. [MB819777]. — Type: DAOM 695763. Ex-type: DAOMC 250534 = CBS 140974 = DTO 410-D4 = W 4 = KAS 2145. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Simplicissima. Reproduction: asexual. ITS barcode: KT887847 (alternative markers: BenA = KT887808; CaM = KT887769; RPB2 = MN969180).

Penicillium catenatum D.B. Scott, Mycopathol. Mycol. Appl. 36: 24. 1968. [MB335719]. — Type: CBS 352.67. Ex-type: CBS 352.67 = ATCC 18543 = CSIR 1097 = IFO 31774 = IMI 136241. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Erubescentia. Reproduction: homothallic. ITS barcode: KC411754 (alternative markers: BenA = KJ834438; CaM = KP016774; RPB2 = JN121504).

Penicillium cavernicola Frisvad & Samson, Stud. Mycol. 49: 31. 2004. [MB370976]. — Type: CBS H-13441. Ex-type: DTO 047-C1 = CBS 100540 = IBT 14499. Infragen. class: subgen. Penicillium, sect. Fasciculata, ser. Camembertiorum. Reproduction: asexual. ITS barcode: KJ834505 (alternative markers: BenA = KJ834439; CaM = KU896827; RPB2 = KU904348).

Penicillium chalabudae Visagie, IMA Fungus 7: 94. 2016. [MB816642]. — Type: CBS H-15439. Ex-type: CBS 219.66 = ATCC 18322 = ATCC 18329 = FRR 3393 = VKM F-1037. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Restricta. Reproduction: asexual. ITS barcode: KP016811 (alternative markers: BenA = KP016748; CaM = KP016767; RPB2 = KP064572).

Penicillium charlesii G. Sm., Trans. Brit. Mycol. Soc. 18: 90. 1933. [MB260433]. — Type: NRRL 778. Ex-type: CBS 304.48 = ATCC 8730 = CBS 342.51 = CECT 2277 = FRR 778 = IMI 040232 = LSHBBB127 = LSHBP 146 = NRRL 1887 = NRRL 778 = QM 6338 = QM 6838. Infragen. class: subgen. Aspergilloides, sect. Charlesia, ser. Fellutana. Reproduction: asexual. ITS barcode: AF033400 (alternative markers: BenA = JX091508; CaM = AY741727; RPB2 = JN121486).

Penicillium chermesinum Biourge, Cellule 33: 284. 1923. [MB260472]. — Type: IMI 191730. Ex-type: CBS 231.81 = FRR 2048 = IFO 31745 = IMI 191730 = NRRL 2048. Infragen. class: subgen. Aspergilloides, sect. Charlesia, ser. Indica. Reproduction: asexual. ITS barcode: AY742693 (alternative markers: BenA = KJ834441; CaM = AY741728; RPB2 = MN969111).

Penicillium choerospondiatis X.C. Wang & W.Y. Zhuang, Sci. Rep. 7: 8233, 5. 2017. [MB570333]. — Type: HMAS 248813. Ex-type: CGMCC 3.18411. Infragen. class: subgen. Aspergilloides, sect. Sclerotiorum, ser. Herqueorum. Reproduction: asexual. ITS barcode: KX885063 (alternative markers: BenA = KX885043; CaM = KX885053; RPB2 = KX885034).

Penicillium christenseniae Houbraken et al., Stud. Mycol. 70: 85. 2011. [MB563187]. — Type: CBS H-20656. Ex-type: CBS 126236 = DTO 076-C3 = IBT 23355. Infragen. class: subgen. Aspergilloides, sect. Citrina, ser. Westlingiorum. Reproduction: asexual. ITS barcode: JN617674 (alternative markers: BenA = JN606680; CaM = MN969243; RPB2 = JN606624).

Penicillium chroogomphum F. Xu et al., Mycoscience 57: 82. 2016. [MB813567]. — Type: JZBHM 002. Ex-type: CBS 136204 = DTO 351-H3 = KCTC 46041 = JZB 2120005. Infragen. class: subgen. Penicillium, sect. Ramosum, ser. Soppiorum. Reproduction: asexual. ITS barcode: KC594043 (alternative markers: BenA = KP684056; CaM = KP684057; RPB2 = MN969167).

Penicillium chrysogenum Thom, U.S.D.A. Bur. Animal Industr. Bull. 118: 58. 1910. [MB165757]. — Type: IMI 24314. Ex-type: DTO 012-I1 = CBS 306.48 = ATCC 10106 = ATHUM2889 = CCRC 30564 = FRR 807 = IBT 5233 = IMI 024314 = IMI 092208 = LSHBAd 3 = LSHBP 19 = MUCL 29079 = MUCL 29145 = NCTC 589 = NRRL 807 = NRRL 810 = QM 7500. Infragen. class: subgen. Penicillium, sect. Chrysogena, ser. Chrysogena. Reproduction: protoheterothallic (Henk et al. 2011); sexual reproduction described by Böhm et al. 2013, reidentified as P. rubens (Houbraken et al. 2014a). ITS barcode: AF033465 (alternative markers: BenA = JF909955; CaM = JX996273; RPB2 = JN121487).

Penicillium chrzaszczii K.W. Zaleski, Bull. Int. Acad. Polon. Sci., Sér. B., Sci. Nat. 1927: 464. 1927. [MB260609]. — Type: CBS 217.28 (lectotype, Houbraken et al. 2011b). Ex-type: CBS 217.28 = DTO 022-E4 = FRR 903 = MUCL 29167 = NRRL 1741 = NRRL 903. Infragen. class: subgen. Aspergilloides, sect. Citrina, ser. Westlingiorum. Reproduction: asexual. ITS barcode: GU944603 (alternative markers: BenA = JN606758; CaM = MN969244; RPB2 = JN606628).

Penicillium cinerascens Biourge, Cellule 33: 308, 1923. [MB260785]. — Type: IMI 92234. Ex-type: DTO 189-A9 = NRRL 748 = ATCC 48693 = BIOURGE 90 = FRR 748 = IMI 92234 = QM 7555 = Thom 4733.34. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Citreonigra. Reproduction: asexual. ITS barcode: AF033455 (alternative markers: BenA = JX141041; CaM = JX157405; RPB2 = MN969112).

Penicillium cinereoatrum Chalab., Bot. Mater. Otd. Sporov. Rast. 6: 167. 1950. [MB302385]. — Type: CBS H-7469. Ex-type: CBS 222.66 = ATCC 22350 = FRR 3390 = IJFM 5024 = IMI 113676 = VKMF-856. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Restricta. Reproduction: asexual. ITS barcode: KC411700 (alternative markers: BenA = KJ834442; CaM = KP125335; RPB2 = JN406608).

Penicillium cinnamopurpureum Abe ex Udagawa, J. Agric. Food Sci. 5: 1. 1959. [MB302386]. — Type: unknown. Ex-type: CBS 429.65 = CBS 847.68 = NRRL 162 = ATCC 18489 = CSIR 936 = FAT 362 = IAM 7016 = IFO 6032 = NHL 6359 = QM 7888. Infragen. class: subgen. Aspergilloides, sect. Cinnamopurpurea, ser. Cinnamopurpurea. Reproduction: homothallic (Scott & Stolk 1967). ITS barcode: EF626950 (alternative markers: BenA = EF626948; CaM = EF626949; RPB2 = JN406533).

Penicillium circulare Hyang B. Lee et al., Fungal Diversity 96: 97. 2019. [MB555413]. — Type: CNUFC-GEU220-1. Ex-type: CNUFC-GEU220-1. Infragen. class: subgen. Aspergilloides, sect. Sclerotiorum, ser. Sclerotiorum. Reproduction: asexual. ITS barcode: n.a. (alternative markers: BenA = MK481057; CaM = MK481061; RPB2 = MK481053).

Penicillium citreonigrum Dierckx, Ann. Soc. Sci. Bruxelles 25: 86. 1901. [MB165197]. — Type: IMI 92209i. Ex-type: CBS 258.29 = ATCC 48736 = FRR 761 = IMI 092209 = LSHBP 20 = LSHBP 98 = MUCL 28648 = MUCL 29062 = MUCL 29116 = NRRL 761. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Citreonigra. Reproduction: asexual. ITS barcode: AF033456 (alternative markers: BenA = EF198621; CaM = EF198628; RPB2 = JN121474).

Penicillium citreosulfuratum Biourge, Cellule 33: 285, 1923. [MB260947]. — Type: France: source unknown; in Biourge, Cellule 33: Fig. 86, no. 21 (lectotype, designated in Visagie et al. 2016c, MBT203135); IMI 92228 (epitype). Ex-epitype: IMI 92228 = DTO 290-I4. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Citreonigra. Reproduction: asexual. ITS barcode: KP016814 (alternative markers: BenA = KP016753; CaM = KP016777; RPB2 = KP064615).

Penicillium citrinum Thom, U.S.D.A. Bur. Animal Industr. Bull. 118: 61. 1910. [MB165293]. — Type: IMI 92196ii. Ex-type: CBS 139.45 = DTO 022-F3 = ATCC 1109 = ATCC 36382 = CECT 2269 = FRR 1841 = IMI 091961 = IMI 092196 = LSHBAd 95 = LSHBP 25 = LSHBP 6 = MUCL 29781 = NRRL 1841 = NRRL 1842. Infragen. class: subgen. Aspergilloides, sect. Citrina, ser. Citrina. Reproduction: asexual. ITS barcode: AF033422 (alternative markers: BenA = GU944545; CaM = MN969245; RPB2 = JF417416).

Penicillium clavigerum Demelius, Verh. Zool.-Bot. Ges. Wien 72: 74. 1923. [MB261069]. — Type: IMI 39807. Ex-type: DTO 248-F6 = CBS 310.48 = ATCC 10427 = CBS 255.94 = FRR 1003 = IMI 039807 = IMI 039807ii = MUCL 15623 = NRRL 1003 = QM 1918. Infragen. class: subgen. Penicillium, sect. Penicillium, ser. Clavigera. Reproduction: asexual. ITS barcode: DQ339555 (alternative markers: BenA = AY674427; CaM = KU896828; RPB2 = KU904349).

Penicillium clavistipitatum Visagie et al., Stud. Mycol. 78: 419. 2014. [MB809961]. — Type: CBS H-21882. Ex-type: CBS 138650 = DTO 182-E5 = CV 336 = KAS 4112 = DAOM 241092. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Pinetorum. Reproduction: asexual. ITS barcode: KM189667 (alternative markers: BenA = KM088912; CaM = KM089299; RPB2 = KM089686).

Penicillium cluniae Quintan. nom. inval. (Art. 40.7), Av. Aliment. Mejora Anim. 30: 174. 1990. [MB130240]. — Type: n.a.; typification needs correction. Ex-type: CBS 326.89 = DTO 265-A8. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Janthinella. Reproduction: asexual. ITS barcode: MN431386 (alternative markers: BenA = MN969376; CaM = MN969246; RPB2 = KF296424).

Penicillium coccotrypicola Holdom et al., Persoonia 33: 285. [MB810327]. — Type: BRIP 59608. Ex-type: BRIP 59608 = DTO 354-F5. Infragen. class: subgen. Penicillium, sect. Penicillium, ser. Clavigera. Reproduction: asexual. ITS barcode: KM605436 (alternative markers: BenA = KM605437; CaM = MN969321; RPB2 = n.a.).

Penicillium coeruleum Sopp in Biourge, Cellule 33: 102. 1923. [MB446014]. — Type: unknown. Ex-type: CBS 141.45 = DTO 035-H5 = NCTC 6595. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Janthinella. Reproduction: asexual. ITS barcode: GU981606 (alternative markers: BenA = GU981655; CaM = MN969247; RPB2 = KF296425).

Penicillium coffeae S.W. Peterson et al., Mycologia 97: 662. 2005. [MB340281]. — Type: BPI 863480. Ex-type: CBS 119387 = IBT 27866 = NRRL 35363. Infragen. class: subgen. Aspergilloides, sect. Charlesia, ser. Phoenicea. Reproduction: asexual. ITS barcode: AY742702 (alternative markers: BenA = KJ834443; CaM = AY741747; RPB2 = JN121436).

Penicillium colei S.W. Peterson et al., PLoS ONE 10: 0121987, 10. 2015. [MB807368]. — Type: BPI 881281. Ex-type: NRRL 13013 = IBT 29696. Infragen. class: subgen. Aspergilloides, sect. Cinnamopurpurea, ser. Idahoensia. Reproduction: asexual. ITS barcode: KF932958 (alternative markers: BenA = KF932926; CaM = KF932942; RPB2 = KF932996).

Penicillium commune Thom, U.S.D.A. Bur. Animal Industr. Bull. 118: 56. 1910. [MB164241]. — Type: IMI 39812. Ex-type: DTO 052-F2 = CBS 311.48 = ATCC 10428 = ATCC 1111 = CCRC 31554 = DSM2211 = IBT 6200 = IFO 5763 = IMI 039812ii = IMI 039812iii = NRRL 890 = QM 1269 = VKMF-3233. Infragen. class: subgen. Penicillium, sect. Fasciculata, ser. Camembertiorum. Reproduction: asexual. ITS barcode: AY213672 (alternative markers: BenA = MN969377; CaM = KU896829; RPB2 = KU904350).

Penicillium compactum L. Wang & Houbraken, Persoonia 36: 309. 2016. [MB810216]. — Type: HMAS 245701. Ex-type: AS 3.15411 = DTO 316-B8 = CBS 138918 = IBT 33393. Infragen. class: subgen. Penicillium, sect. Robsamsonia, ser. Robsamsonia. Reproduction: asexual. ITS barcode: KM973207 (alternative markers: BenA = KM973203; CaM = KM973200; RPB2 = KT698909).

Penicillium concentricum Samson et al., Stud. Mycol. 11: 17. 1976. [MB319263]. — Type: CBS 477.75. Ex-type: CBS 477.75 = IBT 14571 = IBT 6577. Infragen. class: subgen. Penicillium, sect. Robsamsonia, ser. Robsamsonia. Reproduction: asexual. ITS barcode: KC411763 (alternative markers: BenA = AY674413; CaM = DQ911131; RPB2 = KT900575).

Penicillium confertum (Frisvad et al.) Frisvad, Mycologia 81: 852. 1990. [MB126404]. Basionym: Penicillium glandicola var. confertum Frisvad et al., Canad. J. Bot. 65: 769. 1987. [MB131769]. — Type: IMI 296930. Ex-type: CBS 171.87 = IBT 21515 = IBT 3098 = IBT 5672 = IMI 296930 = NRRL 13488 = NRRL A-26904. Infragen. class: subgen. Penicillium, sect. Chrysogena, ser. Chrysogena. Reproduction: asexual. ITS barcode: JX997081 (alternative markers: BenA = AY674373; CaM = JX996963; RPB2 = JX996708).

Penicillium coniferophilum Houbraken & Samson, Stud. Mycol. 70: 47. 2011. [MB561968]. Replaced synonym: Thysanophora striatispora G.L. Barron & W.B. Cooke, Mycopathol. Mycol. Appl. 40 (3–4): 353. 1970. [MB324607]. — Type: unknown. Ex-type: n.a. Infragen. class: subgen. Aspergilloides, sect. Thysanophora, ser. Thysanophora. Reproduction: asexual. ITS barcode: n.a. (alternative markers: BenA = n.a.; CaM = n.a.; RPB2 = n.a.).

Penicillium consobrinum Visagie & K. Jacobs, IMA Fungus 7: 96. 2016. [MB811002]. — Type: CBS H-22045. Ex-type: CBS 139144 = DAOMC 241072 = DTO 181-H9 = CV547. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Corylophila. Reproduction: asexual. ITS barcode: JX140888 (alternative markers: BenA = JX141135; CaM = JX157453; RPB2 = KP064619).

Penicillium contaminatum Houbraken, Stud. Mycol. 78: 419. 2014. [MB809962]. — Type: CBS H-21866. Ex-type: CBS 345.52 = DTO 091-A3 = IMI 049057. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Thomiorum. Reproduction: asexual. ITS barcode: KM189554 (alternative markers: BenA = KM088793; CaM = KM089178; RPB2 = KM089565).

Penicillium coprobium Frisvad, Mycologia 81: 853. 1990. [MB126405]. — Type: IMI 293209. Ex-type: CBS 561.90 = ATCC 58615 = IBT 21516 = IBT 4583 = IBT 6932. Infragen. class: subgen. Penicillium, sect. Robsamsonia, ser. Robsamsonia. Reproduction: asexual. ITS barcode: DQ339559 (alternative markers: BenA = AY674425; CaM = KU896830; RPB2 = KT900576).

Penicillium coprophilum (Berk. & M.A. Curtis) Seifert & Samson, Adv. Pen. Asp. Syst.: 145. 1986 [1985]. [MB114760]. Basionym: Coremium coprophilum Berk. & M.A. Curtis, J. Linn. Soc., Bot., 10: 363. 1868. [MB150510]. — Type: K(M), Wright 666; CBS 111760 (epitype, in Frisvad & Samson 2004b). Ex-epitype: CBS 110760 = IBT 5551 = IBT 3064 = NRRL 13627. Infragen. class: subgen. Penicillium, sect. Robsamsonia, ser. Robsamsonia. Reproduction: asexual. ITS barcode: AF033469 (alternative markers: BenA = AY674421; CaM = KU896831; RPB2 = JN406645).

Penicillium copticola Houbraken et al., Stud. Mycol. 70: 88. 2011. [MB563205]. — Type: CBS H-20643. Ex-type: CBS 127355 = IBT 30771. Infragen. class: subgen. Aspergilloides, sect. Citrina, ser. Copticolarum. Reproduction: asexual. ITS barcode: JN617685 (alternative markers: BenA = JN606817; CaM = JN606553; RPB2 = JN606599).

Penicillium coralligerum Nicot & Pionnat, Bull. Soc. Mycol. France 78: 245. 1963 [1962]. [MB335721]. — Type: IMI 99159. Ex-type: CBS 123.65 = ATCC 16968 = FRR 3465 = IFO 9578 = IHEM 4511 = IMI 099159 = LCP 58.1674 = NRRL 3465 = DTO 104-D9. Infragen. class: subgen. Penicillium, sect. Canescentia, ser. Atroveneta. Reproduction: asexual. ITS barcode: JN617667 (alternative markers: BenA = MN969378; CaM = MN969248; RPB2 = JN406632).

Penicillium corvianum Visagie & Seifert, Persoonia 36: 259. 2016. [MB815770]. — Type: DAOM 695764. Ex-type: DAOMC 250517 = CBS 141000 = DTO 412-B3 = KAS 3618 = IT-2008-4-D. Infragen. class: subgen. Penicillium, sect. Canescentia, ser. Canescentia. Reproduction: asexual. ITS barcode: KT887875 (alternative markers: BenA = KT887836; CaM = KT887797; RPB2 = MN969170).

Penicillium corylophilum Dierckx, Ann. Soc. Sci. Bruxelles 25: 86. 1901. [MB178294]. — Type: IMI 39754. Ex-type: CBS 312.48 = TCC9784 = ATHUM2890 = CECT 2270 = FRR 802 = IMI 039754 = MUCL 28671 = MUCL 29073 = MUCL 29131 = NRRL 802 = QM 7510. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Corylophila. Reproduction: asexual. ITS barcode: AF033450 (alternative markers: BenA = JX141042; CaM = KP016780; RPB2 = KP064631).

Penicillium cosmopolitanum Houbraken et al., Stud. Mycol. 70: 91. 2011. [MB563188]. — Type: CBS H-20665. Ex-type: CBS 126995 = DTO 092-E8 = IBT 30681. Infragen. class: subgen. Aspergilloides, sect. Citrina, ser. Westlingiorum. Reproduction: asexual. ITS barcode: JN617691 (alternative markers: BenA = JN606733; CaM = MN969249; RPB2 = MN969113).

Penicillium costaricense Visagie et al., Persoonia 36: 263. 2016. [MB815773]. — Type: DAOM 695765. Ex-type: DAOMC 250520 = CBS 140998 = DTO 410-E3 = KAS 2597 = 01-RGTHC-294. Infragen. class: subgen. Aspergilloides, sect. Charlesia, ser. Costaricensia. Reproduction: asexual. ITS barcode: MN431396 (alternative markers: BenA = KT887834; CaM = KT887795; RPB2 = MN969173).

Penicillium cravenianum Visagie & K. Jacobs, IMA Fungus 7: 96. 2016. [MB811003]. — Type: CBS H-22044. Ex-type: CBS 139138 = DAOMC 241082 = DTO 180-I5 = CV 92. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Corylophila. Reproduction: asexual. ITS barcode: JX140900 (alternative markers: BenA = JX141076; CaM = JX157418; RPB2 = KP064636).

Penicillium cremeogriseum Chalab., Bot. Mater. Otd. Sporov. Rast. 6: 168. 1950. [MB302390]. — Type: CBS 223.66. Ex-type: CBS 223.66 = DTO 097-B1 = ATCC 18320 = ATCC 18323 = FRR 1734 = IJFM 5011 = IMI 197492 = NRRL 3389 = VKMF-1034. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Janthinella. Reproduction: asexual. ITS barcode: GU981586 (alternative markers: BenA = GU981624; CaM = MN969250; RPB2 = KF296426).

Penicillium crocicola W. Yamam., Sci. Rep. Hyogo Univ. Agric. 2: 28. 1956. [MB302391]. — Type: CBS H-7528. Ex-type: CBS 745.70 = NRRL 6175 = ATCC 18313 = QM 7778. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Thomiorum. Reproduction: asexual. ITS barcode: KM189581 (alternative markers: BenA = KJ834445; CaM = KM089210; RPB2 = JN406535).

Penicillium crustosum Thom, The Penicillia: 399. 1930. [MB262401]. — Type: IMI 91917. Ex-type: CBS 115503 = ATCC 52044 = FRR 1669 = IBT 5528 = IBT 6175 = IMI 091917 = NCTC 4002. Infragen. class: subgen. Penicillium, sect. Fasciculata, ser. Camembertiorum. Reproduction: asexual. ITS barcode: AF033472 (alternative markers: BenA = MN969379; CaM = DQ911132; RPB2 = MN969114).

Penicillium cryptum Goch., Mycotaxon 26: 349. 1986. [MB103648]. — Type: NY 769. Ex-type: CBS 271.89 = ATCC 60138 = IMI 296794 = NRRL 13460 = DTO 122-C9. Infragen. class: subgen. Aspergilloides, sect. Crypta, ser. Crypta. Reproduction: homothallic. ITS barcode: KF303647 (alternative markers: BenA = KF303608; CaM = KF303628; RPB2 = JN121478).

Penicillium crystallinum (Kwon-Chung & Fennell) Samson et al., Stud. Mycol. 78: 355. 2014. [MB809315]. Basionym: Aspergillus crystallinus Kwon-Chung & Fennell, Gen. Aspergillus: 471. 1965. [MB326624]. — Type: IMI 139270. Ex-type: CBS 479.65 = NRRL 5082 = ATCC 16833 = IMI 139270. Infragen. class: subgen. Penicillium, sect. Paradoxa, ser. Paradoxa. Reproduction: asexual. ITS barcode: AF033486 (alternative markers: BenA = EF669682; CaM = FJ530973; RPB2 = EF669669).

Penicillium cuddlyae Visagie & I.H. Rong, Persoonia 43: 38. 2019 [MB832433]. — Type: PREM 623302. Ex-type: PPRI 26355 = CMV016A6. Infragen. class: subgen. Aspergilloides, sect. Charlesia, ser. Indica. Reproduction: asexual. ITS barcode: MK951942 (alternative markers: BenA = MK951835; CaM = MK951908; RPB2 = MN418450).

Penicillium curticaule Visagie & K. Jacobs, Mycol. Prog. 14 (no. 96): 16. 2015. [MB809818]. — Type: CBS H-21334. Ex-type: CBS 135127 = CV 2842 = CV 0188 = DTO 180-D3 = DAOM 241159. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Janthinella. Reproduction: asexual. ITS barcode: FJ231021 (alternative markers: BenA = JX091526; CaM = JX141536; RPB2 = KF296417).

Penicillium cvjetkovicii S.W. Peterson et al., PLoS ONE 10: 0121987, 12. 2015. [MB807369]. — Type: BPI 881283. Ex-type: NRRL 35841 = IBT 29714. Infragen. class: subgen. Aspergilloides, sect. Cinnamopurpurea, ser. Idahoensia. Reproduction: asexual. ITS barcode: KF932963 (alternative markers: BenA = KF932931; CaM = KF932948; RPB2 = KF933002).

Penicillium cyaneum (Bainier & Sartory) Biourge, Cellule 33: 102. 1923. [MB251712]. Basionym: Citromyces cyaneus Bainier & Sartory, Bull. Soc. Mycol. France 29: 157. 1913. [MB178850]. — Type: IMI 39744. Ex-type: CBS 315.48 = ATCC 10432 = FRR 775 = IFO 5337 = IMI 039744 = NRRL 775 = QM 7516. Infragen. class: subgen. Aspergilloides, sect. Ramigena, ser. Ramigena. Reproduction: asexual. ITS barcode: AF033427 (alternative markers: BenA = JX091552; CaM = KP735540; RPB2 = JN406575).

Penicillium cyclopium Westling, Ark. Bot. 11: 90. 1911. [MB156739]. — Type: IMI 089372. Ex-type: DTO 163-I1 = CBS 144.45 = ATCC 8731 = ATHUM2888 = CECT 2264 = DSM1250 = IBT 5130 = IMI 089372 = LSHBP 123 = MUCL 15613 = NRRL 1888 = QM 6839 = VKMF-265. Infragen. class: subgen. Penicillium, sect. Fasciculata, ser. Viridicata. Reproduction: asexual. ITS barcode: JN097811 (alternative markers: BenA = MN969380; CaM = KU896832; RPB2 = JN985388).

Penicillium daejeonium S.H. Yu & H.K. Sang, J. Microbiol. 51: 537. 2013. [MB561572]. — Type: KACC 46609. Ex-type: KACC 46609. Infragen. class: subgen. Aspergilloides, sect. Sclerotiorum, ser. Sclerotiorum. Reproduction: asexual. ITS barcode: JX436489 (alternative markers: BenA = JX436493; CaM = JX436491; RPB2 = n.a.).

Penicillium daleae K.W. Zaleski, Bull. Int. Acad. Polon. Sci., Sér. B., Sci. Nat. 1927: 495. 1927. [MB262773]. — Type: IMI 89338. Ex-type: CBS 211.28 = DTO 105-F2 = ATCC 10435 = DSM 2449 = FRR 2025 = IFO 6087 = IFO 9072 = IMI 034910 = MUCL 29234 = NRRL 2025. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Dalearum. Reproduction: asexual. ITS barcode: GU981583 (alternative markers: BenA = GU981649; CaM = MN969251; RPB2 = KF296427).

Penicillium decaturense S.W. Peterson et al., Mycologia 96: 1290. 2004. [MB487890]. — Type: BPI 842267. Ex-type: CBS 117509 = DTO 003-F7 = NRRL 28152 = IBT 27117. Infragen. class: subgen. Aspergilloides, sect. Citrina, ser. Westlingiorum. Reproduction: asexual. ITS barcode: GU944604 (alternative markers: BenA = JN606685; CaM = MN969252; RPB2 = JN606621).

Penicillium decumbens Thom, U.S.D.A. Bur. Animal Industr. Bull. 118: 71. 1910. [MB156582]. — Type: IMI 190875. Ex-type: CBS 230.81 = FRR 741 = IMI 190875 = MUCL 29107 = NRRL 741. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Alutacea. Reproduction: asexual. ITS barcode: AY157490 (alternative markers: BenA = KJ834446; CaM = KP016782; RPB2 = JN406601).

Penicillium desertorum Frisvad et al., Persoonia 29: 90. 2012. [MB801874]. — Type: CBS H-21056. Ex-type: DTO 148-I6 = CBS 131543 = IBT 16321. Infragen. class: subgen. Penicillium, sect. Chrysogena, ser. Chrysogena. Reproduction: asexual. ITS barcode: JX997011 (alternative markers: BenA = JX996818; CaM = JX996937; RPB2 = JX996682).

Penicillium diabolicalicense Visagie & Seifert, Persoonia 36: 265. 2016. [MB815775]. — Type: PDD 107542. Ex-type: DAOMC 250542 = CBS 140967 = KAS 1726. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Lapidosa. Reproduction: asexual. ITS barcode: KT887840 (alternative markers: BenA = KT887801; CaM = KT887762; RPB2 = MN969175).

Penicillium diatomitis Kubátová et al., Mycol. Prog. 18 (1-2): 223. 2018. [MB824352]. — Type: PRM 861476. Ex-type: CCF 3904 = MH 53 = CBS 140107 = IBT 30728. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Oxalica. Reproduction: asexual. ITS barcode: FJ430748 (alternative markers: BenA = HE651133; CaM = LT970912; RPB2 = LT797560).

Penicillium digitatum (Pers.) Sacc., Fung. Ital. Autogr. Delin.: tab. 894. 1881. Basionym: Aspergillus digitatus Pers., Disp. meth. Fung.: 41. 1794 ≡ Monilia digitata Pers., Syn. Meth. Fung.: 693. 1801. [nom. sanct., Fr., Syst. Mycol. 3: 411. 1832]. [MB169502]. — Neotype: Saccardo, Fung. Ital. Autogr. Delin.: tab. 894. 1881 (lectotype); CBS 112082 (epitype, in Frisvad & Samson 2004b). Ex-type: CBS 112082 = IBT 13068. Infragen. class: subgen. Penicillium, sect. Penicillium, ser. Digitata. Reproduction: asexual. ITS barcode: KJ834506 (alternative markers: BenA = KJ834447; CaM = KU896833; RPB2 = JN121426).

Penicillium dimorphosporum H.J. Swart, Trans. Brit. Mycol. Soc. 55: 310. 1970. [MB120334]. — Type: CBS 456.70. Ex-type: CBS 456.70 = NRRL 5207 = ATCC 22783 = ATCC 52501 = FRR 1120 = IMI 149680. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Erubescentia. Reproduction: asexual. ITS barcode: AF081804 (alternative markers: BenA = KJ834448; CaM = KP016783; RPB2 = JN121517).

Penicillium dipodomyicola (Frisvad et al.) Frisvad, Int. Mod. Meth. Pen. Asp. Clas.: 275. 2000. [MB459818]. Basionym: Penicillium griseofulvum var. dipodomyicola Frisvad et al., Canad. J. Bot. 65: 767. 1987. [MB131771]. — Type: IMI 296935. Ex-type: DTO 202-F7 = CBS 173.87 = IBT 21521 = IMI 296935 = ATCC 64187 = NRRL 13487. Infragen. class: subgen. Penicillium, sect. Robsamsonia, ser. Urticicola. Reproduction: asexual. ITS barcode: MN431387 (alternative markers: BenA = AY674409; CaM = KT900573; RPB2 = KT900577).

Penicillium dipodomyus [as “dipodomyis”] (Frisvad et al.) Banke et al., Int. Mod. Meth. Pen. Asp. Clas.: 271. 2000. [MB459815]. Basionym: Penicillium chrysogenum var. dipodomyus Frisvad et al., Canad. J. Bot. 65: 766. 1987. [MB635036]. — Type: IMI 296926. Ex-type: IBT 5333 = CBS 110412 = DTO 072-B6 = NRRL 13485 = NRRL A-26836 = IMI 296926. Infragen. class: subgen. Penicillium, sect. Chrysogena, ser. Chrysogena. Reproduction: protoheterothallic (Henk et al. 2011). ITS barcode: MN431359 (alternative markers: BenA = AY495991; CaM = JX996950; RPB2 = JF909932).

Penicillium discolor Frisvad & Samson, Antonie van Leeuwenhoek 72: 120. 1997. [MB442902]. — Type: IMI 285513. Ex-type: DTO 046-H9 = CBS 474.84 = IBT 21523 = IBT 5738 = IBT 14440 = IMI 285513 = FRR 2933. Infragen. class: subgen. Penicillium, sect. Fasciculata, ser. Camembertiorum. Reproduction: asexual. ITS barcode: AJ004816 (alternative markers: BenA = AY674348; CaM = KU896834; RPB2 = KU904351).

Penicillium dokdoense Hyang B. Lee & T.T.T. Nguyen, Fungal Diversity 95: 95. 2019. [MB554459]. — Type: CNUFC-DDS11-1. Ex-type: JMRC:SF:013606. Infragen. class: subgen. Aspergilloides, sect. Citrina, ser. Copticolarum. Reproduction: asexual. ITS barcode: MG906868 (alternative markers: BenA = MH243037; CaM = MH243031; RPB2 = n.a.).

Penicillium donkii Stolk, Persoonia 7: 333. 1973. [MB319267]. — Type: CBS 188.72. Ex-type: CBS 188.72 = NRRL 5562 = ATCC 48439 = CCRC 31694 = FRR 1738 = IFO 31746 = IMI 197489 = MUCL 31188. Infragen. class: subgen. Aspergilloides, sect. Stolkia, ser. Stolkia. Reproduction: asexual. ITS barcode: AF033445 (alternative markers: BenA = JN617718; CaM = AF481136; RPB2 = MN969115).

Penicillium dravuni Janso, Mycologia 97: 445. 2005. [MB501442]. — Type: BPI 844248. Ex-type: F01V25. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Erubescentia. Reproduction: asexual. ITS barcode: AY494856 (alternative markers: BenA = n.a.; CaM = n.a.; RPB2 = n.a.).

Penicillium dunedinense Visagie et al., Stud. Mycol. 78: 121. 2014. [MB809183]. — Type: CBS H-21803. Ex-type: CBS 138218 = DTO 244-G1. Infragen. class: subgen. Penicillium, sect. Canescentia, ser. Canescentia. Reproduction: asexual. ITS barcode: KJ775678 (alternative markers: BenA = KJ775171; CaM = KJ775405; RPB2 = MN969116).

Penicillium echinulatum Raper & Thom ex Fassat., Acta Univ. Carol., Biol. 1974: 326. 1977. [MB319269]. — Type: PRM 778523. Ex-type: DTO 047-A4 = CBS 317.48 = IBT 6294 = IMI 040028 = ATCC 10434 = NRRL 1151 = FRR 1151 = IFO 7760 = MUCL 15615 = QM 7519. Infragen. class: subgen. Penicillium, sect. Fasciculata, ser. Camembertiorum. Reproduction: asexual. ITS barcode: AF033473 (alternative markers: BenA = AY674341; CaM = DQ911133; RPB2 = KU904352).

Penicillium echinulonalgiovense S. Abe ex Houbraken & R.N. Barbosa Antonie van Leeuwenhoek 111: 1895. 2018. [MB822213]. — Type: CBS H-23172. Ex-type: DTO 014-H5 = CBS 328.59 = ATCC 18314 = FAT 907 = FRR 638 = IFO 6229 = IMI 068213 = QM 7301. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Simplicissima. Reproduction: asexual. ITS barcode: GU981587 (alternative markers: BenA = GU981631; CaM = KX961269; RPB2 = KX961301).

Penicillium egyptiacum J.F.H. Beyma, Zentralbl. Bakteriol. Parasitenk., Abt. 2 88: 137. 1933 [MB263790]. — Type: IMI 040580. Ex-type: CBS 244.32 = ATCC 10441 = CSIR 707 = FRR 2090 = IBT 14684 = IFO 6094 = IFO 8141 = IFO 8847 = IMI 040580 = NRRL 2090 = QM 1875. Infragen. class: subgen. Penicillium, sect. Chrysogena, ser. Crustacea. Reproduction: homothallic. ITS barcode: AF033467 (alternative markers: BenA = KU896810; CaM = JX996969; RPB2 = JN406598).

Penicillium ehrlichii Kleb., Ber. Deutsch. Bot. Ges. 48: 374. 1930. [MB319270]. — Type: IMI 039737. Ex-type: CBS 324.48 = DTO 097-D7 = ATCC 10442 = IMI 039737 = IMI 039737ii = NRRL 708 = QM 1874 = VKMF-273. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Janthinella. Reproduction: homothallic. ITS barcode: GU981578 (alternative markers: BenA = GU981652; CaM = MN969253; RPB2 = KF296428).

Penicillium elleniae Houbraken et al., Int. J. Syst. Evol. Microbiol. 61: 1470. 2011. [MB518028]. — Type: HUA 170339. Ex-type: CBS 118135 = DTO 057-I9 = IBT 23229. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Janthinella. Reproduction: homothallic. ITS barcode: GU981612 (alternative markers: BenA = GU981663; CaM = MN969254; RPB2 = KF296429).

Penicillium ellipsoideosporum L. Wang & W.Y. Kong, Mycosystema 19: 463. 2000. [MB467721]. — Type: HMAS 71768. Ex-type: CBS 112493 = AS 3.5688. Infragen. class: subgen. Aspergilloides, sect. Cinnamopurpurea, ser. Idahoensia. Reproduction: asexual. ITS barcode: JX012224 (alternative markers: BenA = JQ965104; CaM = AY678559; RPB2 = JN121427).

Penicillium eremophilum (A.D. Hocking & Pitt) Houbraken et al., Stud. Mycol. 86: 47. 2017. [MB820075]. Basionym: Monascus eremophilus A.D. Hocking & Pitt, Mycologia 80: 84. 1988. [MB132383]. — Type: FRR 3338. Ex-type: IMI 313774 = CBS 123361 = ATCC 62925. Infragen. class: subgen. Aspergilloides, sect. Eremophila, ser. Eremophila. Reproduction: asexual. ITS barcode: GU733341 (alternative markers: BenA = KY709170; CaM = KY611931; RPB2 = KY611970).

Penicillium erubescens D.B. Scott, Mycopathol. Mycol. Appl. 36: 14. 1968. [MB335726]. — Type: CBS 318.67. Ex-type: CBS 318.67 = ATCC 18544 = CSIR 1040 = FRR 814 = IFO 31734 = IMI 136204 = NRRL 6223. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Erubescentia. Reproduction: homothallic. ITS barcode: AF033464 (alternative markers: BenA = HQ646566; CaM = EU427281; RPB2 = JN121490).

Penicillium estinogenum A. Komatsu & S. Abe ex G. Sm., Trans. Brit. Mycol. Soc. 46: 335. 1963. [MB302397]. — Type: IMI 68241. Ex-type: CBS 329.59 = DTO 360-C8 = ATCC 18310 = CCRC 31557 = FAT1196 = FRR 3428 = IFO 6230 = IMI 068241 = QM 8149 = VKMF-274. Infragen. class: subgen. Aspergilloides, sect. Gracilenta, ser. Estinogena. Reproduction: asexual. ITS barcode: MN431388 (alternative markers: BenA = MN969381; CaM = MN969255; RPB2 = n.a.).

Penicillium euglaucum J.F.H. Beyma, Antonie van Leeuwenhoek 6: 269. 1940. [MB289081]. — Type: CBS 323.71. Ex-type: CBS 323.71 = IBT 30767. Infragen. class: subgen. Aspergilloides, sect. Citrina, ser. Euglauca. Reproduction: homothallic. ITS barcode: JN617699 (alternative markers: BenA = JN606856; CaM = JN606564; RPB2 = JN606594).

Penicillium excelsum Taniwaki et al., PLoS ONE 10: e0143189, 8. 2015. [MB811066]. — Type: CCT 7772. Ex-type: DTO 357-D7 = ITAL 7572 = IBT 31516. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Rolfsiorum. Reproduction: asexual. ITS barcode: KR815341 (alternative markers: BenA = KP691061; CaM = KR815342; RPB2 = MN969166).

Penicillium expansum Link, Mag. Ges. Naturf. Freunde Berlin 3: 16. 1809. [MB159382]. — Type: CBS H-7082. Ex-type: DTO 141-D5 = CBS 325.48 = ATCC 7861 = ATHUM2891 = CCRC 30566 = FRR 976 = IBT 3486 = IBT 5101 = IMI 039761 = IMI 039761ii = MUCL 29192 = NRRL 976 = VKMF-275. Infragen. class: subgen. Penicillium, sect. Penicillium, ser. Penicillium. Reproduction: asexual. ITS barcode: AY373912 (alternative markers: BenA = AY674400; CaM = DQ911134; RPB2 = JF417427).

Penicillium exsudans X.C. Wang & W.Y. Zhuang, Sci. Rep. 7: 8233, 7. 2017. [MB570336]. — Type: HMAS 248735. Ex-type: CGMCC 3.18412. Infragen. class: subgen. Aspergilloides, sect. Sclerotiorum, ser. Sclerotiorum. Reproduction: asexual. ITS barcode: KX885062 (alternative markers: BenA = KX885042; CaM = KX885052; RPB2 = KX885033).

Penicillium fagi C. Ramírez & A.T. Martínez, Mycopathologia 63: 57. 1978. [MB283595]. — Type: IJFM 3049. Ex-type: CBS 689.77 = CCMF-696 = IJFM 3049 = IMI 253806 = VKMF-2178. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Corylophila. Reproduction: asexual. ITS barcode: AF481124 (alternative markers: BenA = KJ834449; CaM = KP016784; RPB2 = JN406540).

Penicillium fellutanum Biourge, Cellule 33: 262. 1923. [MB264748]. — Type: IMI 39734. Ex-type: CBS 229.81 = CBS 326.48 = ATCC 10443 = FRR 746 = IFO 5761 = IMI 039734 = IMI 039734iii = NRRL 746 = QM 7554. Infragen. class: subgen. Aspergilloides, sect. Charlesia, ser. Fellutana. Reproduction: asexual. ITS barcode: AF033399 (alternative markers: BenA = KJ834450; CaM = AY741753; RPB2 = JN121460).

Penicillium fennelliae Stolk, Antonie van Leeuwenhoek 35: 261. 1969. [MB335728]. — Type: CBS 711.68. Ex-type: CBS 711.68 = ATCC 22050 = ATCC 52492 = FRR 521 = IHEM 4389 = IMI 151747 = MUCL 31322. Infragen. class: subgen. Penicillium, sect. Brevicompacta, ser. Brevicompacta. Reproduction: asexual. ITS barcode: JX313169 (alternative markers: BenA = MN969382; CaM = JX313151; RPB2 = JN406536).

Penicillium fernandesiae R.N. Barbosa et al., Antonie van Leeuwenhoek 111: 1895. 2018. [MB822209]. — Type: URM 90490. Ex-type: CBS 142500 = URM 7600. Infragen. class: subgen. Aspergilloides, sect. Sclerotiorum, ser. Sclerotiorum. Reproduction: asexual. ITS barcode: MF278314 (alternative markers: BenA = MN969416; CaM = LT854649; RPB2 = LT854654).

Penicillium fimorum Frisvad & Houbraken, Persoonia 36: 309. 2016. [MB815871]. — Type: CBS H-22342. Ex-type: CBS 140575 = IBT 29495 = DTO 149-B8 = DTO 159-F1. Infragen. class: subgen. Penicillium, sect. Robsamsonia, ser. Robsamsonia. Reproduction: asexual. ITS barcode: KU904343 (alternative markers: BenA = KT698889; CaM = KT698898; RPB2 = KT698908).

Penicillium fimosum Guevara-Suarez et al., Fungal Syst. Evol. 5: 58. 2019 [2020]. [MB822069]. — Type: CBS H-23185. Ex-type: CBS 142991 = FMR 15104. Infragen. class: subgen. Penicillium, sect. Paradoxa, ser. Atramentosa. Reproduction: asexual. ITS barcode: LT970836 (alternative markers: BenA = LT898273; CaM = LT970837; RPB2 = n.a.).

Penicillium flavigenum Frisvad & Samson, Mycol. Res. 101: 620. 1997. [MB437441]. — Type: CBS 419.89. Ex-type: CBS 419.89 = BT21526 = IBT 3091 = IBT V1035 = IMI 293207. Infragen. class: subgen. Penicillium, sect. Chrysogena, ser. Chrysogena. Reproduction: asexual. ITS barcode: JX997105 (alternative markers: BenA = AY495993; CaM = JX996281; RPB2 = JN406551).

Penicillium flaviroseum L. Cai & X.Z. Jiang, Cladistics 35: 528. 2018 [2019]. [MB818159]. — Type: HMAS 247727. Ex-type: CGMCC 3.18805 = NN072483 = CBS 144479. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Rolfsiorum. Reproduction: asexual. ITS barcode: KY495032 (alternative markers: BenA = KY495141; CaM = MN969329; RPB2 = KY495083).

Penicillium flavisclerotiatum Visagie et al., Houbraken & K. Jacobs, Stud. Mycol. 78: 419. 2014 [MB809963]. — Type: CBS H-21879. Ex-type: CBS 137750 = DTO 180-I8 = CV 100 = DAOM 241157. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Pinetorum. Reproduction: asexual. ITS barcode: KM189644 (alternative markers: BenA = KM088888; CaM = KM089275; RPB2 = KM089662).

Penicillium fluviserpens S.W. Peterson et al., PLoS ONE 10: 0121987, 14. 2015. [MB807370]. — Type: BPI 881284. Ex-type: NRRL 35838 = IBT 29686. Infragen. class: subgen. Aspergilloides, sect. Cinnamopurpurea, ser. Idahoensia. Reproduction: asexual. ITS barcode: KF932961 (alternative markers: BenA = KF932929; CaM = KF932946; RPB2 = KF933000).

Penicillium formosanum H.M. Hsieh et al., Trans. Mycol. Soc. Rep. China 2: 159. 1987. [MB126488]. — Type: PPEH 10001. Ex-type: DTO 206-F3 = CBS 211.92 = IBT 19748 = IBT 21527. Infragen. class: subgen. Penicillium, sect. Formosana, ser. Formosana. Reproduction: asexual. ITS barcode: KC411696 (alternative markers: BenA = AY674426; CaM = KU896835; RPB2 = JN406615).

Penicillium fortuitum Visagie & Seifert, Persoonia 41: 387. 2018. [MB827860]. — Type: DAOM 745786. Ex-type: DTO 313-A3 = DAOMC 251497. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Fortuita. Reproduction: asexual. ITS barcode: MF803942 (alternative markers: BenA = MF803836; CaM = MF803932; RPB2 = MN969206).

Penicillium fractum Udagawa, Trans. Mycol. Soc. Japan 9: 51. 1968. [MB335729]. — Type: CBS H-7086. Ex-type: CBS 124.68 = ATCC 18567 = FRR 3448 = IMI 136701 = NHL 6104 = NRRL 3448. Infragen. class: subgen. Aspergilloides, sect. Inusitata, ser. Inusitata. Reproduction: homothallic. ITS barcode: KC411674 (alternative markers: BenA = KJ834452; CaM = MN969256; RPB2 = JN121441).

Penicillium freii Frisvad & Samson, Stud. Mycol. 49: 28. 2004. [MB369274]. — Type: IMI 285513. Ex-type: DTO 158-D2 = CBS 476.84 = IBT 5137 = IMI 285513. Infragen. class: subgen. Penicillium, sect. Fasciculata, ser. Viridicata. Reproduction: asexual. ITS barcode: MN431389 (alternative markers: BenA = KU896813; CaM = KU896836; RPB2 = KU904353).

Penicillium frequentans Westling, Ark. Bot. 11: 133. 1911. [MB152118]. — Type: CBS 105.11. Ex-type: CBS 105.11. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Glabra. Reproduction: asexual. ITS barcode: KM189525 (alternative markers: BenA = KM088762; CaM = KM089147; RPB2 = KM089534).

Penicillium fructuariae-cellae Lorenzini et al., Phytopathol. Medit. 58: 337. 2019. [MB831228]. — Type: ITEM 18276. Ex-type: CBS 145110. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Rolfsiorum. Reproduction: asexual. ITS barcode: MK039434 (alternative markers: BenA = KU554679; CaM = MK045337; RPB2 = n.a.).

Penicillium fundyense Visagie et al., Persoonia 36: 265. 2016. [MB815776]. — Type: DAOM 695767. Ex-type: DAOMC 250519 = CBS 140980 = DTO 410-D5 = NBBR-2-3 = W466 = KAS 2174. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Citreonigra. Reproduction: asexual. ITS barcode: KT887853 (alternative markers: BenA = KT887814; CaM = KT887775; RPB2 = MN969176).

Penicillium fuscum (Sopp) Biourge, Cellule 33: 103. 1923. [MB289082]. Basionym: Citromyces fuscus Sopp, Skr. Vidensk.-Selsk. Christiana Math.-Nat. Kl. 11: 120. 1912. [MB178643]. — Type: WIS WSF 15-C. Ex-type: CBS 295.62 = ATCC 14770 = CCRC 31517 = DSM2438 = IFO 7743 = IMI 094209 = MUCL 31196 = NRRL 3008 = WSF15c. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Pinetorum. Reproduction: homothallic (Stolk and Samson, 1983, Houbraken et al., 2014b). ITS barcode: AF033411 (alternative markers: BenA = GQ367513; CaM = GQ367539; RPB2 = JN121483).

Penicillium fusisporum L. Wang, PLoS ONE 9: e101454, 2. 2014. [MB806119]. — Type: HMAS 244961. Ex-type: CBS 137463 = NRRL 62805 = AS 3.15338. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Thomiorum. Reproduction: asexual. ITS barcode: KF769424 (alternative markers: BenA = KF769400; CaM = KF769413; RPB2 = MN969117).

Penicillium gallaicum C. Ramírez et al., Mycopathologia 72: 30. 1980. [MB113021]. — Type: CBS H-7464. Ex-type: CBS 167.81 = ATCC 42232 = IJFM 5597. Infragen. class: subgen. Aspergilloides, sect. Citrina, ser. Gallaica. Reproduction: asexual. ITS barcode: JN617690 (alternative markers: BenA = JN606837; CaM = JN606548; RPB2 = JN606609).

Penicillium georgiense S.W. Peterson & B.W. Horn, Mycologia 101: 79. 2009. [MB509290]. — Type: BPI 877332. Ex-type: CBS 132826 = NRRL 35509 = DTO 194-D3. Infragen. class: subgen. Aspergilloides, sect. Ramigena, ser. Georgiensia. Reproduction: asexual. ITS barcode: EF422852 (alternative markers: BenA = EF506223; CaM = EF506239; RPB2 = KM089734).

Penicillium geumsanense Hyang B. Lee et al., Fungal Diversity 96: 101. 2019. [MB555412]. — Type: CNUFC-GEU2229-1. Ex-type: CNUFC-GEU2229-1. Infragen. class: subgen. Penicillium, sect. Robsamsonia, ser. Glandicolarum. Reproduction: asexual. ITS barcode: n.a. (alternative markers: BenA = MK481059; CaM = MK481062; RPB2 = MK481055).

Penicillium glabrum (Wehmer) Westling, Ark. Bot. 11: 131. 1911. [MB120545]. Basionym: Citromyces glaber Wehmer, Beitr. Einh. Pilze 1: 24. 1893. [MB178959]. — Type: IMI 91944. Ex-type: CBS 125543 = IBT 22658 = IMI 91944. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Glabra. Reproduction: asexual. ITS barcode: GU981567 (alternative markers: BenA = GU981619; CaM = KM089152; RPB2 = JF417447).

Penicillium gladioli L. McCulloch & Thom, Science 67: 217. 1928. [MB266048]. — Type: IMI 34911. Ex-type: CBS 332.48 = ATCC 10448 = FRR 939 = IBT 14772 = IMI 034911 = IMI 034911ii = LCP 89.202 = MUCL 29174 = NRRL 939 = QM 1955 = VKMF-2088. Infragen. class: subgen. Penicillium, sect. Fasciculata, ser. Gladioli. Reproduction: asexual. ITS barcode: AF033480 (alternative markers: BenA = AY674287; CaM = KU896837; RPB2 = MN969118).

Penicillium glandicola (Oudem.) Seifert & Samson, Adv. Pen. Asp. Syst.: 147. 1986 [1985]. [MB114761]. Basionym: Coremium glandicola Oudem., Ned. Kruidk. Arch. 2: 918. 1903. [MB240065]. — Type: Netherlands, Valkenburg, Jul 1901, Rick in herb. Oudemans (L); CBS 498.75 (epitype). Ex-epitype: CBS 498.75 = IBT 21529 = IMI 154241. Infragen. class: subgen. Penicillium, sect. Robsamsonia, ser. Glandicolarum. Reproduction: asexual. ITS barcode: AB479308 (alternative markers: BenA = KU896814; CaM = KU896838; RPB2 = KU904354).

Penicillium glaucoalbidum (Desmazières) Houbraken & Samson, Stud. Mycol. 70: 47. 2011. [MB561965]. Basionym: Sclerotium glaucoalbidum Desm., Ann. Sci. Nat., Bot., sér. 3, 16: 329. 1851. [MB212120]. — Type: unknown. Ex-type: n.a. Infragen. class: subgen. Aspergilloides, sect. Thysanophora, ser. Thysanophora. Reproduction: asexual. ITS barcode: n.a. (alternative markers: BenA = n.a.; CaM = n.a.; RPB2 = n.a.).

Penicillium glaucoroseum Demelius, Verh. Zool.-Bot. Ges. Wien 72: 72. 1923. [MB158423]. — Type: Fig. 3. (Demelius, Verh. Zool.-Bot. Ges. Wien 72: 73, 1923 (1922) (lectotype, Visagie et al. 2015); CBS H-22050 (epitype). Ex-epitype: DTO 225-E8 = CBS 138908 = NRRL 908 (authentic acc. Raper & Thom 1949). Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Janthinella. Reproduction: asexual. ITS barcode: MN431390 (alternative markers: BenA = MN969383; CaM = MN969257; RPB2 = MN969119).

Penicillium globosum L. Cai et al., Cladistics 35: 529. 2018 [2019]. [MB818149]. — Type: HMAS 247726. Ex-type: CGMCC 3.18800 = NN072354 = CBS 144639. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Simplicissima. Reproduction: asexual. ITS barcode: KY495014 (alternative markers: BenA = KY495123; CaM = MN969330; RPB2 = KY495067).

Penicillium godlewskii K.W. Zaleski, Bull. Int. Acad. Polon. Sci., Sér. B., Sci. Nat. 1927: 466. 1927. [MB266206]. — Type: CBS 215.28 (lectotype, Houbraken et al. 2011a). Ex-type: CBS 215.28 = DTO 022-E2 = ATCC 10449 = ATCC 48714 = FRR 2111 = IFO 7724 = IMI 040591 = MUCL 29243 = NRRL 2111 = QM 7566 = VKMF-1826. Infragen. class: subgen. Aspergilloides, sect. Citrina, ser. Westlingiorum. Reproduction: asexual. ITS barcode: JN617692 (alternative markers: BenA = JN606768; CaM = MN969258; RPB2 = JN606626).

Penicillium goetzii J. Rogers et al., Persoonia 29: 92. 2012. [MB801876]. — Type: CBS H-21061. Ex-type: DTO 088-G6 = CBS 285.73 = IBT 30199. Infragen. class: subgen. Penicillium, sect. Chrysogena, ser. Goetziorum. Reproduction: asexual. ITS barcode: JX997091 (alternative markers: BenA = KU896815; CaM = JX996971; RPB2 = JX996716).

Penicillium gorlenkoanum Baghd., Novosti Sist. Nizsh. Rast. 5: 97. 1968. [MB335731]. — Type: CBS H-7490. Ex-type: CBS 408.69 = DTO 023-A5 = DTO 034-E3 = FRR 511 = IMI 140339 = VKMF-1079. Infragen. class: subgen. Aspergilloides, sect. Citrina, ser. Citrina. Reproduction: asexual. ITS barcode: GU944581 (alternative markers: BenA = GU944520; CaM = MN969259; RPB2 = JN606601).

Penicillium gracilentum Udagawa & Y. Horie, Trans. Mycol. Soc. Japan 14: 373. 1973. [MB319272]. — Type: NHL 6452. Ex-type: CBS 599.73 = ATCC 28047 = ATCC 48258 = FRR 1557 = IMI 216900 = NHL 6452 = DTO 095-D8. Infragen. class: subgen. Aspergilloides, sect. Gracilenta, ser. Gracilenta. Reproduction: homothallic. ITS barcode: KC411768 (alternative markers: BenA = KJ834453; CaM = MN969260; RPB2 = JN121537).

Penicillium grancanariae C. Ramírez et al., Mycopathologia 66: 79. 1978. [MB319273]. — Type: IJFM 3745. Ex-type: CBS 687.77 = IJFM 3745 = IMI 253783. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Spinulosa. Reproduction: asexual. ITS barcode: KM189529 (alternative markers: BenA = KM088766; CaM = KM089151; RPB2 = KM089538).

Penicillium granulatum Bainier, Bull. Soc. Mycol. Fr. 21: 126. 1905. [MB174620]. — Type: CBS 333.48. Ex-type: CBS 333.48 = DTO 246-F5 = ATCC 10450 = FRR 2036 = IBT 6592 = IMI 040220 = MUCL 15621 = NRRL 2036 = QM 6868. Infragen. class: subgen. Penicillium, sect. Robsamsonia, ser. Glandicolarum. Reproduction: asexual. ITS barcode: DQ339565 (alternative markers: BenA = MT478038.; CaM = MT478036; RPB2 = MT478035).

Penicillium gravinicasei S.W. Peterson et al., Int. J. Food Microbiol. 282: 67. 2018. [MB823510]. — Type: BPI 910534. Ex-type: ITEM 17411 = NRRL 66733. Infragen. class: subgen. Aspergilloides, sect. Cinnamopurpurea, ser. Cinnamopurpurea. Reproduction: asexual. ITS barcode: MG600580 (alternative markers: BenA = MG600565; CaM = MG600570; RPB2 = MG600575).

Penicillium grevilleicola Houbraken & Quaedvlieg, Stud. Mycol. 78: 423. 2014. [MB809964]. — Type: CBS H-21871. Ex-type: CBS 137775 = DTO 174-E6. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Thomiorum. Reproduction: asexual. ITS barcode: KM189630 (alternative markers: BenA = KM088874; CaM = KM089261; RPB2 = KM089648).

Penicillium griseoazureum C. Moreau & M. Moreau ex C. Ramírez, Manual and Atlas of the Penicillia: 61, 1982; Penicillium griseo-azureum C. Moreau & M. Moreau, Revue Mycol. 6: 59, 1941 (nom. inval., Art. 36.1). [MB115800]. — type: CBS 162.42 (holotype). ex-type: CBS 162.42 = FRR 1361. Infragen. class: subgen. Penicillium, sect. Canescentia, ser. Canescentia. Reproduction: asexual. ITS barcode: KC411679 (alternative markers: BenA = KP016919; CaM = KP016823; RPB2 = KP016852).

Penicillium griseoflavum L. Cai & X.Z. Jiang, Cladistics 35: 531. 2018 [2019]. [MB818151]. — Type: HMAS 247729. Ex-type: CGMCC 3.18799 = NN072331. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Simplicissima. Reproduction: asexual. ITS barcode: KY495011 (alternative markers: BenA = KY495120; CaM = MN969331; RPB2 = KY495064).

Penicillium griseofulvum Dierckx, Ann. Soc. Sci. Bruxelles 25: 88. 1901. [MB120566]. — Type: IMI 75832. Ex-type: DTO 072-A5 = CBS 185.27 = ATCC 11885 = ATHUM2893 = CECT 2605 = DSM896 = IBT 6740 = IFO 7640 = IFO 7641 = IMI 075832 = IMI 075832ii = LCP 79.3245 = LSHBP 68 = MUCL 28643 = NRRL 2152 = NRRL 2300 = QM 6902 = VKMF-286. Infragen. class: subgen. Penicillium, sect. Robsamsonia, ser. Urticicola. Reproduction: asexual. ITS barcode: AF033468 (alternative markers: BenA = JF909942; CaM = KT900574; RPB2 = JN121449).

Penicillium griseolum G. Sm., Trans. Brit. Mycol. Soc. 40: 485. 1957. [MB302401]. — Type: IMI 071626. Ex-type: CBS 277.58 = ATCC 18239 = FRR 2671 = IFO 8175 = IMI 071626 = LSHBB323 = NRRL 2671 = QM 7523. Infragen. class: subgen. Aspergilloides, sect. Griseola, ser. Griseola. Reproduction: asexual. ITS barcode: EF422848 (alternative markers: BenA = EF506213; CaM = EF506232; RPB2 = JN121480).

Penicillium griseopurpureum G. Sm., Trans. Brit. Mycol. Soc. 48: 275. 1965. [MB335732]. — Type: IMI 96157. Ex-type: CBS 406.65 = DTO 383-E5 = ATCC 22353 = FRR 3429 = IFO 9147 = IMI 096157. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Dalearum. Reproduction: asexual. ITS barcode: KF296408 (alternative markers: BenA = KF296467; CaM = MN969261; RPB2 = KF296431).

Penicillium guaibinense J.P. Andrade et al., Persoonia 41: 389. 2018. [MB827182]. — Type: HURB 18573. Ex-type: CCDCA 11512 = 23EM8. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Dalearum. Reproduction: asexual. ITS barcode: MH674389 (alternative markers: BenA = MH674391; CaM = MH674393; RPB2 = n.a.).

Penicillium guanacastense K.G. Rivera et al., Mycotaxon 119: 324. 2011. [MB563044]. — Type: DAOM 239912. Ex-type: CCFC 239912. Infragen. class: subgen. Aspergilloides, sect. Sclerotiorum, ser. Sclerotiorum. Reproduction: asexual. ITS barcode: JN626098 (alternative markers: BenA = JN625967; CaM = JN626010; RPB2 = KX961295).

Penicillium guangxiense L. Cai & X.Z. Jiang, Cladistics 35: 533. 2018 [2019]. [MB818150]. — Type: HMAS 247737. Ex-type: CGMCC 3.18793 = NN044175 = CBS 144526. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Simplicissima. Reproduction: asexual. ITS barcode: KY494986 (alternative markers: BenA = KY495095; CaM = MN969332; RPB2 = KY495045).

Penicillium guttulosum J.C. Gilman & E.V. Abbott, Iowa St. Coll. J. Sci. 1: 298. 1927. [MB266689]. — Type: Iowa St. Coll. J. Sci. 1: 298, 1927 Fig. 33 (lectotype), Visagie et al. 2016c; CBS H-22566 (epitype). Ex-epitype: NRRL 907 = ATCC 48734 = FRR 907 = Thom 4894.16. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Erubescentia. Reproduction: asexual. ITS barcode: HQ646592 (alternative markers: BenA = HQ646576; CaM = HQ646587; RPB2 = MG386247).

Penicillium hainanense L. Cai & X.Z. Jiang, Cladistics 35: 534. 2018 [2019]. [MB818156]. — Type: HMAS 247730. Ex-type: CGMCC 3.18798 = NN072329 = CBS 144527. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Rolfsiorum. Reproduction: asexual. ITS barcode: KY495009 (alternative markers: BenA = KY495118; CaM = MN969333; RPB2 = KY495062).

Penicillium halotolerans Frisvad et al., Persoonia 29: 92. 2012. [MB801875]. — Type: CBS H-21060. Ex-type: DTO 148-H9 = CBS 131537 = IBT 4315. Infragen. class: subgen. Penicillium, sect. Chrysogena, ser. Chrysogena. Reproduction: asexual. ITS barcode: JX997005 (alternative markers: BenA = JX996816; CaM = JX996935; RPB2 = JX996680).

Penicillium hemitrachum Visagie & K. Jacobs, IMA Fungus 7: 99. 2016. [MB811004]. — Type: CBS H-22042. Ex-type: CBS 139134 = DAOMC 241098 = DTO 180-D8 = CV 2845. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Lapidosa. Reproduction: asexual. ITS barcode: FJ231003 (alternative markers: BenA = JX141048; CaM = JX157526; RPB2 = KP064642).

Penicillium hennebertii Houbraken & Samson, Stud. Mycol. 70: 47. 2011. [MB561964]. Replaced synonym: Thysanophora canadensis Stolk & Hennebert, Persoonia 5: 189. 1968. [MB340084]. — Type: CBS H-7854. Ex-type: CBS 334.68 = ATCC 18741 = IMI 137644 = MUCL 21216 = VKMF-2999. Infragen. class: subgen. Aspergilloides, sect. Thysanophora, ser. Thysanophora. Reproduction: asexual. ITS barcode: KJ834507 (alternative markers: BenA = KJ834454; CaM = MN969262; RPB2 = JN121493).

Penicillium hermansii Houbraken et al., Mycol. Prog. 18: 232. 2018. [MB823949]. — Type: CBS H-21028. Ex-type: CBS 124296 = DTO 079-D5. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Erubescentia. Reproduction: asexual. ITS barcode: MG333472 (alternative markers: BenA = MG386214; CaM = MG386229; RPB2 = MG386242).

Penicillium herquei Bainier & Sartory, Bull. Soc. Mycol. France 28: 121. 1912. [MB536431]. — Type: IMI 28809. Ex-type: CBS 336.48 = NRRL 1040 = ATCC 10118 = BIOURGE 452 = FRR 1040 = IFO 31747 = IMI 28809 = MUCL 29213 = NCTC 1721 = QM 1926 = Thom 4640.447. Infragen. class: subgen. Aspergilloides, sect. Sclerotiorum, ser. Herqueorum. Reproduction: asexual. ITS barcode: JN626101 (alternative markers: BenA = JN625970; CaM = JN626013; RPB2 = JN121494).

Penicillium heteromorphum H.Z. Kong & Z.T. Qi, Mycosystema 1: 107. 1988. [MB135444]. — Type: CBS 226.89. Ex-type: CBS 226.89. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Restricta. Reproduction: asexual. ITS barcode: KC411702 (alternative markers: BenA = KJ834455; CaM = KP016786; RPB2 = JN406605).

Penicillium hetheringtonii Houbraken et al., Fungal Diversity 44: 125. 2010. [MB518292]. — Type: CBS 122392. Ex -type: CBS 122392 = DTO 005-H9 = IBT 29057. Infragen. class: subgen. Aspergilloides, sect. Citrina, ser. Citrina. Reproduction: asexual. ITS barcode: GU944558 (alternative markers: BenA = GU944538; CaM = MN969263; RPB2 = JN606606).

Penicillium hirayamae Udagawa, J. Agric. Soc. Tokyo 5: 6. 1959. [MB302402]. — Type: IMI 78255. Ex-type: CBS 229.60 = ATCC 18312 = IFO 6435 = IMI 078255 = IMI 078255ii = NHL 6046 = NRRL 143 = QM 7885. Infragen. class: subgen. Aspergilloides, sect. Sclerotiorum, ser. Sclerotiorum. Reproduction: homothallic (Scott & Stolk 1967). ITS barcode: JN626095 (alternative markers: BenA = JN625955; CaM = JN626003; RPB2 = JN121459).

Penicillium hirsutum Dierckx, Ann. Soc. Sci. Bruxelles 25: 89. 1901. [MB152720]. — Type: IMI 40213. Ex-type: CBS 135.41 = ATCC 10429 = FRR 2032 = IBT 21531 = IFO 6092 = IMI 040213 = MUCL 15622 = NRRL 2032. Infragen. class: subgen. Penicillium, sect. Fasciculata, ser. Corymbifera. Reproduction: asexual. ITS barcode: AY373918 (alternative markers: BenA = MN969384; CaM = KU896840; RPB2 = JN406629).

Penicillium hispanicum C. Ramírez et al., Mycopathologia 66: 77. 1978. [MB319274]. — Type: IJFM 3223. Ex-type: CBS 691.77 = ATCC 38667 = DSM2416 = IJFM 3223 = IMI 253785 = VKMF-2179. Infragen. class: subgen. Aspergilloides, sect. Ramigena, ser. Ramigena. Reproduction: asexual. ITS barcode: JX841247 (alternative markers: BenA = KJ834456; CaM = MN969264; RPB2 = JN406539).

Penicillium hoeksii Houbraken, Stud. Mycol. 78: 423. 2014. [MB809965]. — Type: CBS H-21860. Ex-type: CBS 137776 = DTO 192-H4. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Hoeksiorum. Reproduction: asexual. ITS barcode: KM189707 (alternative markers: BenA = KM088954; CaM = KM089341; RPB2 = KM089728).

Penicillium hordei Stolk, Antonie van Leeuwenhoek 35: 270. 1969. [MB335734]. — Type: CBS 701.68. Ex-type: CBS 701.68 = DTO 303-B6 = ATCC 22053 = CECT 2290 = FRR 815 = IBT 17804 = IBT 6980 = IMI 151748 = MUCL 39559. Infragen. class: subgen. Penicillium, sect. Fasciculata, ser. Corymbifera. Reproduction: asexual. ITS barcode: MN431391 (alternative markers: BenA = MN969385; CaM = KU896841; RPB2 = KU904355).

Penicillium ibericum Guevara-Suarez et al., Fungal Syst. Evol. 5: 58. 2019 [2020]. [MB822070]. — Type: CBS H-23186. Ex-type: CBS 142992 = FMR 15040. Infragen. class: subgen. Penicillium, sect. Paradoxa, ser. Atramentosa. Reproduction: asexual. ITS barcode: LT899782 (alternative markers: BenA = LT898285; CaM = LT899766; RPB2 = LT899800).

Penicillium idahoense Paden, Mycopathol. Mycol. Appl. 43: 259. 1971. [MB319275]. — Type: UVIC JWP 66-32. Ex-type: CBS 341.68 = NRRL 5274 = ATCC 22055 = FRR 881 = IMI 148393. Infragen. class: subgen. Aspergilloides, sect. Cinnamopurpurea, ser. Idahoensia. Reproduction: homothallic. ITS barcode: KC411747 (alternative markers: BenA = EF626953; CaM = EF626954; RPB2 = JN121499).

Penicillium improvisum Visagie et al., Persoonia 36: 256. 2016. [MB815769]. — Type: DAOM 695768. Ex-type: DAOMC 250547 = CBS 140994 = DTO 410-D9 = KAS 2386 = W 156. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Improvisa. Reproduction: asexual. ITS barcode: KT887867 (alternative markers: BenA = KT887828; CaM = KT887789; RPB2 = MN969169).

Penicillium incoloratum L.Q. Huang & Z.T. Qi, Acta Mycol. Sin. 13: 264. 1994. [MB363421]. — Type: HMAS 65949. Ex-type: CBS 101753 = AS 3.4672. Infragen. class: subgen. Aspergilloides, sect. Cinnamopurpurea, ser. Nodula. Reproduction: asexual. ITS barcode: KJ834508 (alternative markers: BenA = KJ834457; CaM = KJ866984; RPB2 = JN406651).

Penicillium indicum D.K. Sandhu & R.S. Sandhu, Canad. J. Bot. 41: 1273. 1963. [MB335735]. — Type: New Delhi-6, No. Pe 1602; CBS H-7476 (isotype). Ex-type: CBS 115.63 = NRRL 3387 = ATCC 18324 = FRR 3387 = IFO 31744 = IMI 166620. Infragen. class: subgen. Aspergilloides, sect. Charlesia, ser. Indica. Reproduction: asexual. ITS barcode: AY742699 (alternative markers: BenA = EU427263; CaM = AY741744; RPB2 = JN406640).

Penicillium infra-aurantiacum Visagie et al., Stud. Mycol. 78: 426. 2014 [MB809966]. — Type: CBS H-21880. Ex-type: CBS 137747 = DTO 183-C3 = CV 1518 = DAOM 241145. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Sublectatica. Reproduction: asexual. ITS barcode: KM189684 (alternative markers: BenA = KM088930; CaM = KM089317; RPB2 = KM089704).

Penicillium infrabuccalum Visagie et al., Persoonia 36: 275. 2016. [MB815782]. — Type: DAOM 695769. Ex-type: DAOMC 250537 = CBS 140983 = DTO 410-D6 = NBSM-6-2 = W 475 = KAS 2181. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Simplicissima. Reproduction: asexual. ITS barcode: KT887856 (alternative markers: BenA = KT887817; CaM = KT887778; RPB2 = MN969181).

Penicillium infrapurpureum Visagie et al., Stud. Mycol. 78: 116. 2014. [MB809181]. — Type: CBS H-21801. Ex-type: CBS 138219 = DTO 235-F6. Infragen. class: subgen. Aspergilloides, sect. Cinnamopurpurea, ser. Idahoensia. Reproduction: asexual. ITS barcode: KJ775679 (alternative markers: BenA = KJ775172; CaM = KJ775406; RPB2 = MN969120).

Penicillium inusitatum D.B. Scott, Mycopathol. Mycol. Appl. 36: 20. 1968. [MB335736]. — Type: CBS 351.67. Ex-type: CBS 351.67 = ATCC 18622 = CSIR 1096 = FRR 1163 = IMI 136214 = NRRL 5810. Infragen. class: subgen. Aspergilloides, sect. Inusitata, ser. Inusitata. Reproduction: homothallic. ITS barcode: AF033431 (alternative markers: BenA = KJ834458; CaM = MN969265; RPB2 = JN121503).

Penicillium isariiforme Stolk & J.A. Mey., Trans. Brit. Mycol. Soc. 40: 187. 1957. [MB302403]. — Type: IMI 60371. Ex-type: CBS 247.56 = ATCC 18425 = CCRC 31699 = IFO 6393 = IHEM 4376 = IMI 060371 = LSHBBB308 = MUCL 31191 = MUCL 31323 = NRRL 2638 = QM 1897. Infragen. class: subgen. Aspergilloides, sect. Ochrosalmonea, ser. Ochrosalmonea. Reproduction: asexual. ITS barcode: AF454077 (alternative markers: BenA = KJ834459; CaM = MN969266; RPB2 = JN121470).

Penicillium italicum Wehmer, Hedwigia 33: 211. 1894. [MB162660]. — Type: CBS 339.48. Ex-type: CBS 339.48 = ATCC 10454 = DSM 2754 = FRR 983 = IBT 23029 = IMI 039760 = MUCL 15608 = NRRL 983 = QM 7572. Infragen. class: subgen. Penicillium, sect. Penicillium, ser. Italica. Reproduction: asexual. ITS barcode: KJ834509 (alternative markers: BenA = AY674398; CaM = DQ911135; RPB2 = JN121496).

Penicillium jacksonii K.G. Rivera et al., Stud. Mycol. 70: 151. 2011. [MB563160]. — Type: DAOM 239937. Ex-type: CCFC 239937. Infragen. class: subgen. Aspergilloides, sect. Sclerotiorum, ser. Sclerotiorum. Reproduction: asexual. ITS barcode: JN686437 (alternative markers: BenA = JN686368; CaM = JN686391; RPB2 = n.a.).

Penicillium jamesonlandense Frisvad & Overy, Int. J. Syst. Evol. Microbiol. 56: 1435. 2006. [MB521421]. — Type: DAOM 234087. Ex-type: CBS 102888 = DAOM 234087 = IBT 21984 = IBT 24411. Infragen. class: subgen. Penicillium, sect. Ramosum, ser. Lanosa. Reproduction: asexual. ITS barcode: DQ267912 (alternative markers: BenA = DQ309448; CaM = KJ866985; RPB2 = MN969121).

Penicillium janczewskii K.W. Zaleski, Bull. Int. Acad. Polon. Sci., Sér. B., Sci. Nat. 1927: 488. 1927. [MB120703]. — Type: IMI 191499. Ex-type: CBS 221.28 = FRR 919 = IMI 191499 = NRRL 919. Infragen. class: subgen. Penicillium, sect. Canescentia, ser. Canescentia. Reproduction: asexual. ITS barcode: AY157487 (alternative markers: BenA = MN969386; CaM = MN969267; RPB2 = JN406612).

Penicillium janthinellum Biourge, Cellule 33: 258. 1923. [MB119134]. — Type: IMI 40238. Ex-type: CBS 340.48 = DTO 095-C3 = ATCC 10455 = IMI 040238 = NRRL 2016 = QM 6865. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Janthinella. Reproduction: asexual. ITS barcode: GU981585 (alternative markers: BenA = GU981625; CaM = MN969268; RPB2 = JN121497).

Penicillium javanicum J.F.H. Beyma, Verh. Kon. Ned. Akad. Wetensch., Afd. Natuurk. 26: 17. 1929. [MB268394]. — Type: IMI 39733. Ex-type: CBS 341.48 = DTO 097-F9 = ATCC 9099 = CSIR 831 = FRR 707 = IFO 31735 = IMI 039733 = MUCL 29099 = NRRL 707 = QM 1876. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Janthinella. Reproduction: homothallic. ITS barcode: GU981613 (alternative markers: BenA = GU981657; CaM = MN969269; RPB2 = JN121498).

Penicillium jejuense M.S. Park & Y.W. Lim, Mycologia 107: 212. 2015. [MB808392]. — Type: SFC 20140101-M756. Ex-type: CBS 138646 = KCTC 46212. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Thomiorum. Reproduction: asexual. ITS barcode: KF818464 (alternative markers: BenA = KF818461; CaM = KF818470; RPB2 = KF818467).

Penicillium jensenii K.W. Zaleski, Bull. Int. Acad. Polon. Sci., Sér. B., Sci. Nat. 1927: 494. 1927. [MB120708]. — Type: IMI 39768. Ex-type: CBS 327.59 = ATCC 18317 = FRR 909 = IFO 5764 = IMI 039768 = LCP 89.1389 = NRRL 909 = QM 7587. Infragen. class: subgen. Penicillium, sect. Canescentia, ser. Canescentia. Reproduction: asexual. ITS barcode: AY443470 (alternative markers: BenA = JX140954; CaM = AY443490; RPB2 = JN406614).

Penicillium jianfenglingense L. Cai & X.Z. Jiang, Cladistics 35: 535. 2018 [2019]. [MB818161]. — Type: HMAS 247731. Ex-type: CGMCC 3.18802 = NN072384 = CBS 144640. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Dalearum. Reproduction: asexual. ITS barcode: KY495016 (alternative markers: BenA = KY495125; CaM = MN969334; RPB2 = KY495069).

Penicillium jiangxiense H.Z. Kong & Z.Q. Liang, Mycosystema 22: 4. 2003. [MB489161]. — Type: HMAS 82540. Ex-type: AS 3.6521 = DTO 309-A7. Infragen. class: subgen. Aspergilloides, sect. Cinnamopurpurea, ser. Jiangxiensia. Reproduction: asexual. ITS barcode: KJ890411 (alternative markers: BenA = KJ890409; CaM = KJ890407; RPB2 = MN969122).

Penicillium johnkrugii K.G. Rivera et al., Stud. Mycol. 70: 151. 2011. [MB563161]. — Type: DAOM 239943. Ex-type: CCFC 239943. Infragen. class: subgen. Aspergilloides, sect. Sclerotiorum, ser. Sclerotiorum. Reproduction: asexual. ITS barcode: JN686447 (alternative markers: BenA = JN686378; CaM = JN686401; RPB2 = n.a.).

Penicillium jugoslavicum C. Ramírez & Munt.-Cvetk., Mycopathologia 88: 65. 1984. [MB124173]. — Type: CBS 192.87. Ex-type: CBS 192.87 = IJFM 7785 = IMI 314508. Infragen. class: subgen. Aspergilloides, sect. Sclerotiorum, ser. Adametziorum. Reproduction: asexual. ITS barcode: KC773836 (alternative markers: BenA = KC773789; CaM = KC773815; RPB2 = JN406618).

Penicillium kananaskense Seifert et al., Canad. J. Bot. 72: 20. 1994. [MB362160]. — Type: DAOM 216105. Ex-type: CBS 530.93 = ATCC 90282 = DAOM 216105 = IBT 11775 = IMI 356791. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Livida. Reproduction: asexual. ITS barcode: KM189780 (alternative markers: BenA = KM089030; CaM = KM089417; RPB2 = KM089804).

Penicillium katangense Stolk, Antonie van Leeuwenhoek 34: 42. 1968. [MB120725]. — Type: CBS 247.67. Ex-type: CBS 247.67 = ATCC 18388 = IMI 136206 = NRRL 5182. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Restricta. Reproduction: homothallic. ITS barcode: AF033458 (alternative markers: BenA = KP016757; CaM = KP016788; RPB2 = KP064646).

Penicillium kewense G. Sm., Trans. Brit. Mycol. Soc. 44: 42. 1961. [MB335740]. — Type: L.S.H.T.M. BB 400 (holotype); CBS H-7077 (isotype). Ex-type: CBS 344.61 = ATCC 18240 = FRR 3441 = IFO 8113 = IMI 086561 = LSHBBB400 = MUCL 2685 = NRRL 3332 = QM 7958. Infragen. class: subgen. Penicillium, sect. Chrysogena, ser. Crustacea. Reproduction: homothallic. ITS barcode: AF033466 (alternative markers: BenA = KU896816; CaM = JX996973; RPB2 = JF417428).

Penicillium kiamaense Houbraken & Pitt, Stud. Mycol. 78: 426. 2014. [MB809967]. — Type: CBS H-21857. Ex-type: CBS 137947 = FRR 6087 = DTO 056-I6. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Kiamaensia. Reproduction: asexual. ITS barcode: KM189506 (alternative markers: BenA = KM088743; CaM = KM089128; RPB2 = KM089515).

Penicillium kojigenum G. Sm., Trans. Brit. Mycol. Soc. 44: 43. 1961. [MB335741]. — Type: L.S.H.T.M. BB 39. Ex-type: CBS 345.61 = ATCC 18227 = CCRC 31515 = FRR 3442 = IFO 9581 = IMI 086562 = LSHBBB394 = MUCL 2457 = NRRL 3442 = QM 7957. Infragen. class: subgen. Penicillium, sect. Ramosum, ser. Lanosa. Reproduction: asexual. ITS barcode: AF033489 (alternative markers: BenA = KJ834463; CaM = KJ867011; RPB2 = JN406564).

Penicillium kongii L. Wang, Mycologia 105: 1549. 2013. [MB803185]. — Type: HMAS 244382. Ex-type: AS 3.15329. Infragen. class: subgen. Penicillium, sect. Brevicompacta, ser. Brevicompacta. Reproduction: asexual. ITS barcode: KC427191 (alternative markers: BenA = KC427171; CaM = KC427151; RPB2 = n.a.).

Penicillium koreense S.B. Hong et al., J. Microbiol. Biotechnol 24: 1607. 2014. [MB808759]. — Type: KACC 47721. Ex-type: CBS 141338 = KACC 47721 = DTO 347-C1. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Janthinella. Reproduction: asexual. ITS barcode: KJ801939 (alternative markers: BenA = KM000846; CaM = MN969317; RPB2 = MN969159).

Penicillium kurssanovii Chalab., Bot. Mater. Otd. Sporov. Rast. 6: 168. 1950. [MB274327]. — Type: unknown. Ex-type: CBS 625.67 = ATCC 18387 = FRR 3381 = IJFM 5045 = IMI 129965 = NRRL 3381 = VKMF-1244. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Restricta. Reproduction: asexual. ITS barcode: EF422849 (alternative markers: BenA = KP016758; CaM = KP016789; RPB2 = KP064647).

Penicillium labradorum Gibas et al., Med. Mycol., doi.org/10.1093/mmy/myaa016. 2020. [MB831086]. — Type: CBS H-24321. Ex-type UTHSCSA DI19-20 = CBS 145775. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Erubescentia. Reproduction: asexual. ITS barcode: MK881918 (alternative markers: BenA = MK887898; CaM = MK887899; RPB2 = MK887900).

Penicillium laeve (K. Ando & Manoch) Houbraken & Samson, Stud. Mycol. 70: 47. 2011. [MB561960]. Basionym: Torulomyces laevis K. Ando & Manoch, Mycoscience 39: 317. 1998. [MB447110]. — Type: TNS-F-238517. Ex-type: CBS 136665 = KY 12727 = NBRC 109724 = DTO 270-G8. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Erubescentia. Reproduction: homothallic. ITS barcode: KF667369 (alternative markers: BenA = KF667365; CaM = KF667367; RPB2 = KF667371).

Penicillium laevigatum L. Cai et al., Cladistics 35: 537. 2018 [2019]. [MB818154]. — Type: HMAS 247728. Ex-type: CGMCC 3.18801 = NN072364. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Simplicissima. Reproduction: asexual. ITS barcode: KY495015 (alternative markers: BenA = KY495124; CaM = MN969335; RPB2 = KY495068).

Penicillium lagena (Delitsch) Stolk & Samson, Stud. Mycol. 23: 100. 1983. [MB109162]. Basionym: Torulomyces lagena Delitsch, Systematik der Schimmelpilze: 9. 1943. [MB340152]. — Type: Fig. 233 (Delitsch, Ergebnisse der theoretischen und angewandten Mikrobiologie: Band I: Systematik der Schimmelpilze. J. Neumann, Neudamm, Germany, Tafel 30, 1943, Visagie et al. 2016a, MBT203020 (lectotype); CBS 185.65 (epitype). Ex-type: CBS 185.65 = MUCL 8221 = JCM10149 = OAC10034 = DTO 077-I8. Infragen. class: subgen. Aspergilloides, sect. Torulomyces, ser. Torulomyces. Reproduction: asexual. ITS barcode: KF303665 (alternative markers: BenA = KF303619; CaM = KF303634; RPB2 = JN121450).

Penicillium lanosocoeruleum Thom, Penicillia: 322. 1930. [MB268949]. — Type: NRRL 888. Ex-type: CBS 215.30 = CBS 334.48 = ATCC 10459 = IFO 7761 = IMI 039818 = NRRL 888 = QM 6755 = VKMF-3089. Infragen. class: subgen. Penicillium, sect. Chrysogena, ser. Aethiopica. Reproduction: asexual. ITS barcode: JX997110 (alternative markers: BenA = KU896817; CaM = JX996967; RPB2 = JX996723).

Penicillium lanosum Westling, Ark. Bot. 11: 97. 1911. [MB178497]. — Type: IMI 40224. Ex-type: DTO 060-F7 = CBS 106.11 = ATCC 10458 = FRR 2009 = IFO 5851 = IFO 6099 = IMI 040224 = LSHBP 86 = MUCL 29232 = NRRL 2009 = QM 7591. Infragen. class: subgen. Penicillium, sect. Ramosum, ser. Lanosa. Reproduction: asexual. ITS barcode: DQ304540 (alternative markers: BenA = DQ285627; CaM = FJ530974; RPB2 = KU904356).

Penicillium lapidosum Raper & Fennell, Mycologia 40: 524, 1948. [MB289094]. — Type: IMI 39743. Ex-type: CBS 343.48 = ATCC 10462 = CCT4477 = IFO 6100 = IMI 039743 = NRRL 718 = QM 1928. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Lapidosa. Reproduction: homothallic (Scott & Stolk 1967). ITS barcode: MN431392 (alternative markers: BenA = KJ834465; CaM = FJ530984; RPB2 = JN121500).

Penicillium lassenii Paden, Mycopathol. Mycol. Appl. 43: 266. 1971. [MB319281]. — Type: UVIC JWP 69-26. Ex-type: CBS 277.70 = NRRL 5272 = ATCC 22054 = FRR 858 = IMI 148395 = DTO 095-D6. Infragen. class: subgen. Aspergilloides, sect. Lasseniorum, ser. Lasseniorum. Reproduction: homothallic. ITS barcode: KF303648 (alternative markers: BenA = KF303607; CaM = KF303629; RPB2 = JN121481).

Penicillium lemhiflumine S.W. Peterson et al., PLoS ONE 10: 0121987, 4. 2015. [MB807371]. — Type: BPI 881287. Ex-type: NRRL 35843 = IBT 29684. Infragen. class: subgen. Aspergilloides, sect. Cinnamopurpurea, ser. Idahoensia. Reproduction: asexual. ITS barcode: KF932964 (alternative markers: BenA = KF932932; CaM = KF932949; RPB2 = KF933003).

Penicillium lenticrescens Visagie et al., Stud. Mycol. 78: 123. 2014. [MB809184]. — Type: CBS H-21804. Ex-type: CBS 138215 = DTO 129-A8. Infragen. class: subgen. Penicillium, sect. Ramosum, ser. Soppiorum. Reproduction: asexual. ITS barcode: KJ775675 (alternative markers: BenA = KJ775168; CaM = KJ775404; RPB2 = MN969123).

Penicillium levitum Raper & Fennell, Mycologia 40: 511. 1948. [MB289096]. — Type: IMI 039735. Ex-type: CBS 345.48 = DTO 096-I7 = ATCC 10464 = IFO 6101 = IFO 8849 = IMI 039735 = NRRL 705 = QM 1877. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Janthinella. Reproduction: homothallic. ITS barcode: GU981607 (alternative markers: BenA = GU981654; CaM = MN969270; RPB2 = KF296432).

Penicillium lilacinoechinulatum S. Abe ex G. Sm., Trans. Brit. Mycol. Soc. 46: 335. 1963. [MB120793]. — Type: IMI 068211. Ex-type: CBS 454.93 = ATCC 18309 = FAT 84 = FRR 3451 = IFO 6231 = IMI 068211 = QM 7289. Infragen. class: subgen. Aspergilloides, sect. Sclerotiorum, ser. Adametziorum. Reproduction: asexual. ITS barcode: AY157489 (alternative markers: BenA = KC773790; CaM = KC773816; RPB2 = KX961293).

Penicillium limosum S. Ueda, Mycoscience 36: 451. 1995. [MB415136]. — Type: CBM NEI-5220. Ex-type: CBS 339.97 = DTO 096-H8 = NEI5220. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Janthinella. Reproduction: homothallic. ITS barcode: GU981568 (alternative markers: BenA = GU981621; CaM = MN969271; RPB2 = KF296433).

Penicillium lineolatum Udagawa & Y. Horie, Mycotaxon 5: 493. 1977. [MB319283]. — Type: NHL 2776. Ex-type: CBS 188.77 = DTO 097-E1 = NHL 2776. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Janthinella. Reproduction: homothallic. ITS barcode: GU981579 (alternative markers: BenA = GU981620; CaM = MN969272; RPB2 = KF296434).

Penicillium lividum Westling, Ark. Bot. 11: 134. 1911. [MB178817]. — Type: IMI 39736. Ex-type: CBS 347.48 = ATCC 10102 = CCRC 31286 = DSM1180 = IFO 6102 = IMI 039736 = NRRL 754 = QM 1930 = VKMF-303. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Livida. Reproduction: asexual. ITS barcode: KM189582 (alternative markers: BenA = KM088825; CaM = KM089211; RPB2 = KM089598).

Penicillium longicatenatum Visagie et al., Stud. Mycol. 78: 429. 2014. [MB809968]. — Type: CBS H-21875. Ex-type: CBS 137735 = DTO 180-D9 = CV 2847 = DAOM 241119. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Longicatenata. Reproduction: asexual. ITS barcode: KM189636 (alternative markers: BenA = KM088880; CaM = KM089267; RPB2 = KM089654).

Penicillium longisporum (W.B. Kend.) Houbraken & Samson, Stud. Mycol. 70: 47. 2011. [MB561966]. Basionym: Thysanophora longispora W.B. Kend., Canad. J. Bot. 39: 826. 1961. [MB340086]. — Type: DAOM 63073. Ex-type: CBS 354.62 = DAOM 63073 = MUCL 4168. Infragen. class: subgen. Aspergilloides, sect. Thysanophora, ser. Thysanophora. Reproduction: asexual. ITS barcode: n.a. (alternative markers: BenA = n.a.; CaM = n.a.; RPB2 = n.a.).

Penicillium ludwigii Udagawa, Trans. Mycol. Soc. Japan 10: 2. 1969. [MB335744]. — Type: NHL 6118. Ex-type: CBS 417.68 = DTO 094-D8 = DTO 264-I9 = FRR 559. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Janthinella. Reproduction: homothallic. ITS barcode: KF296409 (alternative markers: BenA = KF296468; CaM = MN969273; RPB2 = KF296435).

Penicillium lunae Visagie & N. Yilmaz, Persoonia 42: 449. 2019. [MB830682]. — Type: PREM 62233. Ex-type: PPRI 25881 = CMV006E6. Infragen. class: subgen. Aspergilloides, sect. Charlesia, ser. Indica. Reproduction: asexual. ITS barcode: MK450725 (alternative markers: BenA = MK451088; CaM = MK451660; RPB2 = MK450863).

Penicillium lusitanum Gonçalves & Alves, Int. J. Syst. Evol. Microbiol. 69: 3020. 2019. [MB830331]. — Type: MUM-H 18.49. Ex-type: MUM 18.49 = CMG8. Infragen. class: subgen. Penicillium, sect. Ramosum, ser. Soppiorum. Reproduction: asexual. ITS barcode: MK702084 (alternative markers: BenA = MK702085; CaM = n.a.; RPB2 = n.a.).

Penicillium maclennaniae H.Y. Yip, Trans. Brit. Mycol. Soc. 77: 202. 1981. [MB112523]. — Type: DAR 35238. Ex-type: CBS 198.81 = DAR 35238. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Lapidosa. Reproduction: asexual. ITS barcode: KC411689 (alternative markers: BenA = KJ834468; CaM = KP016791; RPB2 = KP064648).

Penicillium macrosclerotiorum L. Wang et al., Mycol. Res. 111: 1244. 2007. [MB492622]. — Type: HMAS 133177-1-4. Ex-type: CBS 116871 = AS 3.6581. Infragen. class: subgen. Aspergilloides, sect. Gracilenta, ser. Macrosclerotiorum. Reproduction: asexual. ITS barcode: KJ834511 (alternative markers: BenA = KJ834469; CaM = DQ911123; RPB2 = JN121432).

Penicillium madriti G. Sm., Trans. Brit. Mycol. Soc. 44: 44. 1961. [MB335747]. — Type: IMI 86563. Ex-type: CBS 347.61 = ATCC 18233 = CCRC 31672 = FRR 3452 = IFO 9148 = IMI 086563 = LSHBBB389 = MUCL 2456 = MUCL 31193 = NRRL 3452 = QM 7959. Infragen. class: subgen. Penicillium, sect. Turbata, ser. Turbata. Reproduction: asexual. ITS barcode: AF033482 (alternative markers: BenA = KJ834470; CaM = EU644076; RPB2 = JN406561).

Penicillium magnielliptisporum Visagie et al., Stud. Mycol. 78: 127. 2014. [MB809186]. — Type: CBS H-21806. Ex-type: CBS 138225 = DTO 128-H8. Infragen. class: subgen. Penicillium, sect. Paradoxa, ser. Atramentosa. Reproduction: asexual. ITS barcode: KJ775686 (alternative markers: BenA = KJ775179; CaM = KJ775413; RPB2 = MN969124).

Penicillium malacaense C. Ramírez & A.T. Martínez, Mycopathologia 72: 186. 1980. [MB113025]. — Type: IJFM 7093. Ex-type: CBS 160.81 = NRRL 35754 = ATCC 42241 = IJFM 7093 = IMI 253801 = VKMF-2197. Infragen. class: subgen. Aspergilloides, sect. Cinnamopurpurea, ser. Idahoensia. Reproduction: asexual. ITS barcode: EU427300 (alternative markers: BenA = EU427268; CaM = KJ866997; RPB2 = JN406626).

Penicillium malachiteum (Yaguchi & Udagawa) Houbraken & Samson, Stud. Mycol. 70: 47. 2011. [MB561971]. Basionym: Chromocleista malachitea Yaguchi & Udagawa, Trans. Mycol. Soc. Japan 34: 102. 1993. [MB360067]. — Type: CBS 647.95. Ex-type: CBS 647.95 = IBT 17515. Infragen. class: subgen. Aspergilloides, sect. Sclerotiorum, ser. Herqueorum. Reproduction: homothallic. ITS barcode: KC773838 (alternative markers: BenA = KC773794; CaM = KC773820; RPB2 = MN969125).

Penicillium malacosphaerulum Visagie & K. Jacobs, Mycol. Prog. 14: 96, 16. 2015. [MB809819]. — Type: CBS H-21332. Ex-type: CBS 135120 = CV 2855 = CV 0311 = DTO 180-E6 = DAOM 241161. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Janthinella. Reproduction: homothallic. ITS barcode: FJ231026 (alternative markers: BenA = JX091524; CaM = JX141542; RPB2 = KF296438).

Penicillium mali-pumilae Hyang B. Lee et al., Fungal Diversity 96: 101. 2019. [MB555410]. — Type: CBS H-22503. Ex-type: CBS 140671 = DTO 327-D1= EML-MP6080-1 = IBT 33672. Infragen. class: subgen. Penicillium, sect. Fasciculata, ser. Corymbifera. Reproduction: asexual. ITS barcode: KP900991 (alternative markers: BenA = MT425584; CaM = KP900992; RPB2 = MN969213).

Penicillium mallochii K.G. Rivera et al., Mycotaxon 119: 322. 2012. [MB563043]. — Type: DAOM 239917. Ex-type: CCFC 239917. Infragen. class: subgen. Aspergilloides, sect. Sclerotiorum, ser. Sclerotiorum. Reproduction: asexual. ITS barcode: JN626104 (alternative markers: BenA = JN625973; CaM = JN626016; RPB2 = KX961296).

Penicillium malmesburiense Visagie et al., Stud. Mycol. 78: 429. 2014. [MB809969]. — Type: CBS H-21872. Ex-type: CBS 137744 = DTO 182-H5 = CV 1180 = DAOM 241144. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Sublectatica. Reproduction: asexual. ITS barcode: KM189676 (alternative markers: BenA = KM088921; CaM = KM089308; RPB2 = KM089695).

Penicillium malodoratum (Kwon-Chung & Fennell) Samson et al., Stud. Mycol. 78: 355. 2014. [MB809316]. Basionym: Aspergillus malodoratus Kwon-Chung & Fennell, Gen. Aspergillus: 468. 1965. [MB326644]. — Type: IMI 172289. Ex-type: CBS 490.65 = NRRL 5083 = IMI 172289 = ATCC 16834. Infragen. class: subgen. Penicillium, sect. Paradoxa, ser. Paradoxa. Reproduction: asexual. ITS barcode: AF033485 (alternative markers: BenA = EF669681; CaM = FJ530972; RPB2 = EF669672).

Penicillium manginii Duché & R. Heim, Trav. Cryptog.: 450. 1931. [MB270490]. — Type: CBS 253.31. Ex-type: CBS 253.31 = DTO 022-E9 = NRRL 2134. Infragen. class: subgen. Aspergilloides, sect. Citrina, ser. Westlingiorum. Reproduction: asexual. ITS barcode: GU944599 (alternative markers: BenA = JN606651; CaM = MN969274; RPB2 = JN606618).

Penicillium mariae-crucis Quintan., Av. Aliment. Mejora Anim. 23: 334. 1982. [MB114171]. — Type: CBS 270.83. Ex-type: CBS 271.83 = IMI 256075. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Simplicissima. Reproduction: asexual. ITS barcode: GU981593 (alternative markers: BenA = GU981630; CaM = MN969275; RPB2 = KF296439).

Penicillium marinum Frisvad & Samson, Stud. Mycol. 49: 20. 2004. [MB370974]. — Type: CBS 109550. Ex-type: DTO 141-E5 = CBS 109550 = IBT 14360. Infragen. class: subgen. Penicillium, sect. Penicillium, ser. Penicillium. Reproduction: asexual. ITS barcode: KJ834512 (alternative markers: BenA = AY674392; CaM = KU896842; RPB2 = KU904357).

Penicillium marthae-christenseniae Visagie & Samson, Persoonia 36: 145. 2016. [MB808267]. — Type: CBS H-21613. Ex-type: CBS 129213 = DTO 201-B5. Infragen. class: subgen. Aspergilloides, sect. Torulomyces, ser. Torulomyces. Reproduction: asexual. ITS barcode: KF303651 (alternative markers: BenA = KF303613; CaM = KF303645; RPB2 = KF303711).

Penicillium maximae Visagie et al., Persoonia 31: 52. 2013. [MB803783]. — Type: CBS H-21144. Ex-type: CBS 134565 = NRRL 2060. Infragen. class: subgen. Aspergilloides, sect. Sclerotiorum, ser. Sclerotiorum. Reproduction: asexual. ITS barcode: EU427298 (alternative markers: BenA = KC773795; CaM = KC773821; RPB2 = MN969126).

Penicillium mediterraneum Guevara-Suarez et al., Fungal Syst. Evol. 5: 61. 2019 [2020]. [MB822071]. — Type: CBS H-23143. Ex-type: CBS 142754 = FMR 15188. Infragen. class: subgen. Penicillium, sect. Roquefortorum, ser. Roquefortorum. Reproduction: asexual. ITS barcode: LT899784 (alternative markers: BenA = LT898291; CaM = LT899768; RPB2 = LT899802).

Penicillium melanoconidium (Frisvad) Frisvad & Samson, Stud. Mycol. 49: 28. 2004. [MB368219]. Basionym: Penicillium aurantiogriseum var. melanoconidium Frisvad, Mycologia 81: 849. 1989. [MB126407]. — Type: IMI 321503. Ex-type: DTO 158-D1 = CBS 115506 = IBT 3444 = IMI 321503. Infragen. class: subgen. Penicillium, sect. Fasciculata, ser. Viridicata. Reproduction: asexual. ITS barcode: MN431393 (alternative markers: BenA = MN969387; CaM = KU896843; RPB2 = KU904358).

Penicillium melanostipe Houbraken & Samson, Stud. Mycol. 70: 47. 2011. [MB561970]. Replaced synonym: Thysanophora verrucosa Mercado et al., Mycotaxon 67: 419. 1998. [MB443755]. — Type: HAC (M) 9165. Ex-type: n.a. Infragen. class: subgen. Aspergilloides, sect. Thysanophora, ser. Thysanophora. Reproduction: asexual. ITS barcode: n.a. (alternative markers: BenA = n.a.; CaM = n.a.; RPB2 = n.a.).

Penicillium melinii Thom, Penicillia: 273. 1930. [MB270876]. — Type: IMI 40216. Ex-type: CBS 218.30 = ATCC 10469 = FRR 2041 = IFO 7675 = IMI 040216 = MUCL 29235 = NRRL 2041 = QM 7599. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Lapidosa. Reproduction: asexual. ITS barcode: AF033449 (alternative markers: BenA = KJ834471; CaM = KP016792; RPB2 = JN406613).

Penicillium meliponae R.N. Barbosa et al., Antonie van Leeuwenhoek 111: 1897. 2018. [MB822210]. — Type: URM 90491. Ex-type: CBS 142495 = URM 7602. Infragen. class: subgen. Aspergilloides, sect. Sclerotiorum, ser. Sclerotiorum. Reproduction: asexual. ITS barcode: MF278315 (alternative markers: BenA = MN969418; CaM = LT854648; RPB2 = LT854653).

Penicillium mellis R.N. Barbosa et al., Antonie van Leeuwenhoek 111: 1900. 2018. [MB822211]. — Type: URM 90492. Ex-type: CBS 142499 = URM 7605. Infragen. class: subgen. Aspergilloides, sect. Sclerotiorum, ser. Adametziorum. Reproduction: asexual. ITS barcode: MN431398 (alternative markers: BenA = MN969417; CaM = MN969327; RPB2 = LT854652).

Penicillium meloforme Udagawa & Y. Horie, Trans. Mycol. Soc. Japan 14: 376. 1973. [MB120882]. — Type: NHL 6468. Ex-type: CBS 445.74 = DTO 101-B3 = ATCC 28049 = IMI 216903 = NHL 6468. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Janthinella. Reproduction: homothallic. ITS barcode: KC411762 (alternative markers: BenA = GU981656; CaM = MN969276; RPB2 = KF296440).

Penicillium menonorum S.W. Peterson, IMA Fungus 2: 122. 2011. [MB519297]. — Type: BPI 881018. Ex-type: NRRL 50410. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Erubescentia. Reproduction: asexual. ITS barcode: HQ646591 (alternative markers: BenA = HQ646573; CaM = HQ646584; RPB2 = KF900194).

Penicillium meridianum D.B. Scott, Mycopathol. Mycol. Appl. 36: 12. 1968. [MB335750]. — Type: CBS 314.67. Ex-type: CBS 314.67 = ATCC 18545 = CSIR 1052 = IMI 136209. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Restricta. Reproduction: homothallic. ITS barcode: AF033451 (alternative markers: BenA = KJ834472; CaM = KP016794; RPB2 = JN406576).

Penicillium mexicanum Visagie et al., Stud. Mycol. 78: 125. 2014. [MB809185]. — Type: CBS H-21805. Ex-type: CBS 138227 = DTO 270-F1. Infragen. class: subgen. Penicillium, sect. Paradoxa, ser. Atramentosa. Reproduction: asexual. ITS barcode: KJ775685 (alternative markers: BenA = KJ775178; CaM = KJ775412; RPB2 = MN969127).

Penicillium miczynskii K.W. Zaleski, Bull. Int. Acad. Polon. Sci., Sér. B., Sci. Nat. 1927: 482. 1927. [MB271171]. — Type: IMI 40030. Ex-type: CBS 220.28 = DTO 022-E5 = ATCC 10470 = DSM2437 = FRR 1077 = IFO 7730 = IMI 040030 = MUCL 29228 = NRRL 1077 = QM 1957. Infragen. class: subgen. Aspergilloides, sect. Citrina, ser. Westlingiorum. Reproduction: asexual. ITS barcode: GU944600 (alternative markers: BenA = JN606706; CaM = MN969277; RPB2 = JN606623).

Penicillium minnesotense Jurjević et al., Persoonia 42: 445. 2019. [MB830666]. — Type: BPI 910934. Ex-type: NRRL 66823 = ITEM 17524 = EMSL 1719. Infragen. class: subgen. Aspergilloides, sect. Cinnamopurpurea, ser. Idahoensia. Reproduction: asexual. ITS barcode: MK791277 (alternative markers: BenA = MK803429; CaM = MK803430; RPB2 = MK796158).

Penicillium momoii Visagie & K. Jacobs, IMA Fungus 7: 99. 2016. [MB811007]. — Type: CBS H-22046. Ex-type: CBS 139157 = DAOMC 241077 = DTO 182-G4 = CV 1015. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Corylophila. Reproduction: asexual. ITS barcode: JX140895 (alternative markers: BenA = JX141073; CaM = JX157479; RPB2 = KP064673).

Penicillium mononematosum (Frisvad et al.) Frisvad, Mycologia 81: 857. 1990. [MB126406]. Basionym: Penicillium glandicola var. mononematosum Frisvad, Filt. & Wicklow, Canad. J. Bot. 65: 767. 1987. [MB131770]. — Type: IMI 296925. Ex-type: CBS 172.87 = IBT 3072 = IBT 5518 = IBT 21535 = IMI 296925 = NRRL 13482 = NRRL A-26709. Infragen. class: subgen. Penicillium, sect. Chrysogena, ser. Chrysogena. Reproduction: asexual. ITS barcode: JX997082 (alternative markers: BenA = AY495997; CaM = JX996964; RPB2 = JX996709).

Penicillium monsgalena S.W. Peterson et al., PLoS ONE 10: 0121987, 17. 2015. [MB807372]. — Type: BPI 881282. Ex-type: NRRL 22302 = IBT 29713. Infragen. class: subgen. Aspergilloides, sect. Cinnamopurpurea, ser. Idahoensia. Reproduction: asexual. ITS barcode: KF932959 (alternative markers: BenA = KF932927; CaM = KF932943; RPB2 = KF932997).

Penicillium monsserratidens S.W. Peterson et al., PLoS ONE 10: 0121987, 19. 2015. [MB807373]. — Type: BPI 881285. Ex-type: NRRL 35884 = IBT 29695. Infragen. class: subgen. Aspergilloides, sect. Cinnamopurpurea, ser. Idahoensia. Reproduction: asexual. ITS barcode: KF932962 (alternative markers: BenA = KF932930; CaM = KF932947; RPB2 = KF933001).

Penicillium montanense M. Chr. & Backus, Mycologia 54: 574. 1962. [MB335752]. — Type: WIS Cryptogamic Herb. No. GW1-6. Ex-type: CBS 310.63 = ATCC 14941 = FRR 3407 = IFO 7740 = IHEM 4375 = IMI 099468 = MUCL 31326 = NRRL 3407. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Pinetorum. Reproduction: asexual. ITS barcode: KM189551 (alternative markers: BenA = KM088789; CaM = KM089174; RPB2 = KM089561).

Penicillium murcianum C. Ramírez & A.T. Martínez, Mycopathologia 74: 37. 1981 [MB112524]. — Type: IJFM 7031. Ex-type: DTO 036-A2 = CBS 161.81 = ATCC 42239 = IJFM 7031 = IMI 253800 = VKMF-2196. Infragen. class: subgen. Penicillium, sect. Canescentia, ser. Canescentia. Reproduction: asexual. ITS barcode: MN431400 (alternative markers: BenA = MN969419; CaM = MN969341; RPB2 = MN969202).

Penicillium nalgiovense Laxa, Zentralbl. Bakteriol. Parasitenk., Abt. 2 86: 160. 1932. [MB114239]. — Type: CBS 352.48. Ex-type: CBS 352.48 = ATCC 10472 = IBT 21536 = IMI 039804 = MUCL 31194 = NRRL 911. Infragen. class: subgen. Penicillium, sect. Chrysogena, ser. Chrysogena. Reproduction: asexual. ITS barcode: AY371617 (alternative markers: BenA = KU896811; CaM = JX996974; RPB2 = JX996719).

Penicillium namyslowskii K.W. Zaleski, Bull. Int. Aead. Polonc. Sci., Cl. Sci. Math., Sér. B, Sci. Nat. 1927: 479. 1927. [MB272006]. — Type: CBS 353.48. Ex-type: CBS 353.48 = ATCC 11127 = IMI 040033 = MUCL 29226 = NRRL 1070. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Lapidosa. Reproduction: asexual. ITS barcode: AF033463 (alternative markers: BenA = JX141067; CaM = KP016795; RPB2 = JF417430).

Penicillium neocrassum R. Serra & S.W. Peterson, Mycologia 99: 81. 2007. [MB504767]. — Type: BPI 872161. Ex-type: CBS 122428 = NRRL 35639 = MUM 06.160. Infragen. class: subgen. Penicillium, sect. Brevicompacta, ser. Brevicompacta. Reproduction: asexual. ITS barcode: DQ645805 (alternative markers: BenA = DQ645794; CaM = DQ645809; RPB2 = JN406633).

Penicillium neoechinulatum (Frisvad et al.) Frisvad & Samson, Stud. Mycol. 49: 28. 2004. [MB368218]. Basionym: Penicillium aurantiogriseum var. neoechinulatum Frisvad et al., Canad. J. Bot. 65: 767. 1987. [MB131767]. — Type: IMI 296937. Ex-type: CBS 169.87 = CBS 101135 = IBT 3493 = IBT 21537 = IMI 296937 = NRRL 13486. Infragen. class: subgen. Penicillium, sect. Fasciculata, ser. Viridicata. Reproduction: asexual. ITS barcode: JN942722 (alternative markers: BenA = MN969388; CaM = KU896844; RPB2 = JN985406).

Penicillium neomiczynskii A.L.J. Cole et al., Stud. Mycol. 70: 105. 2011. [MB563192]. — Type: CBS H-20661. Ex-type: CBS 126231 = DTO 078-C2 = IBT 23560. Infragen. class: subgen. Aspergilloides, sect. Citrina, ser. Westlingiorum. Reproduction: asexual. ITS barcode: JN617671 (alternative markers: BenA = JN606705; CaM = MN969278; RPB2 = MN969128).

Penicillium nepalense Takada & Udagawa, Trans. Mycol. Soc. Japan 24: 146. 1983. [MB108327]. — Type: NHL 6482. Ex-type: CBS 203.84 = NHL 6482. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Erubescentia. Reproduction: homothallic. ITS barcode: KC411692 (alternative markers: BenA = KJ834474; CaM = KP016796; RPB2 = JN121453).

Penicillium nigricans Bainier ex Thom, Penicillia: 351. 1930. [MB119303]. — Type: CBS H-22051. Ex-type: CBS 354.48 = ATCC 10115 = IFO 6103 = IMI 039767 = NRRL 915 = QM 1933 = VKMF-313. Infragen. class: subgen. Penicillium, sect. Canescentia, ser. Canescentia. Reproduction: asexual. ITS barcode: KC411755 (alternative markers: BenA = KJ866965; CaM = KJ867012; RPB2 = KP016857).

Penicillium nodulum H.Z. Kong & Z.T. Qi, Mycosystema 1: 108. 1988. [MB135445]. — Type: CBS 227.89. Ex-type: CBS 227.89. Infragen. class: subgen. Aspergilloides, sect. Cinnamopurpurea, ser. Nodula. Reproduction: asexual. ITS barcode: KC411703 (alternative markers: BenA = KJ834475; CaM = KJ867003; RPB2 = JN406603).

Penicillium nordicum Dragoni & Cantoni ex C. Ramírez, Adv. Pen. Asp. Syst: 139. 1986 [1985]. [MB114762]. — Type: ATCC 44219. Ex-type: DTO 098-F7 = ATCC 44219 = IBT 13307. Infragen. class: subgen. Penicillium, sect. Fasciculata, ser. Verrucosa. Reproduction: asexual. ITS barcode: KJ834513 (alternative markers: BenA = MN969389; CaM = KU896845; RPB2 = KU904359).

Penicillium nothofagi Houbraken et al., Stud. Mycol. 70: 105. 2011. [MB563189]. — Type: CBS H-20655. Ex-type: CBS 130383 = DTO 076-C2 = IBT 23018 = DTO 076-C2. Infragen. class: subgen. Aspergilloides, sect. Citrina, ser. Westlingiorum. Reproduction: asexual. ITS barcode: JN617712 (alternative markers: BenA = JN606732; CaM = JN606507; RPB2 = MN969129).

Penicillium novae-zeelandiae J.F.H. Beyma, Antonie van Leeuwenhoek 6: 275. 1940. [MB522253]. — Type: IMI 40584ii. Ex-type: CBS 137.41 = ATCC 10473 = IFO 31748 = IMI 040584ii = NRRL 2128 = QM 1934 = VKMF-2886 = DTO 035-D8. Infragen. class: subgen. Penicillium, sect. Canescentia, ser. Atroveneta. Reproduction: asexual. ITS barcode: JN617688 (alternative markers: BenA = MN969390; CaM = MN969279; RPB2 = JN406628).

Penicillium nucicola Visagie et al., Persoonia 36: 259. 2016. [MB815771]. — Type: DAOM 695770. Ex-type: DAOMC 250522 = CBS 140987 = DTO 410-D7 = W 59 = KAS 2203. Infragen. class: subgen. Penicillium, sect. Canescentia, ser. Atroveneta. Reproduction: asexual. ITS barcode: KT887860 (alternative markers: BenA = KT887821; CaM = KT887782; RPB2 = MN969171).

Penicillium ochrochloron Biourge, Cellule 33: 269. 1923. [MB272701]. — Type: IMI 39806. Ex-type: CBS 357.48 = DTO 097-G2 = ATCC 10540 = IMI 039806 = NRRL 926 = QM 7604. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Rolfsiorum. Reproduction: asexual. ITS barcode: GU981604 (alternative markers: BenA = GU981672; CaM = MN969280; RPB2 = KF296445).

Penicillium ochrosalmoneum Udagawa, J. Agric. Sci. Tokyo Nogyo Daig. 5: 10. 1959. [MB302409]. — Type: NHL 6048. Ex-type: CBS 489.66 = ATCC 18338 = CSIR 145 = IMI 116248ii = NRRL 35499. Infragen. class: subgen. Aspergilloides, sect. Ochrosalmonea, ser. Ochrosalmonea. Reproduction: homothallic (Scott & Stolk 1967). ITS barcode: EF626961 (alternative markers: BenA = EF506212; CaM = EF506237; RPB2 = JN121524).

Penicillium odoratum M. Chr. & Backus, Mycologia 53: 459. 1961. [MB335755]. — Type: WSF 2000. Ex-type: CBS 294.62 = CBS 296.62 = ATCC 14769 = DSM2419 = IFO 7741 = IMI 094208ii = NRRL 3007 = WSF2000. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Livida. Reproduction: asexual. ITS barcode: KC411730 (alternative markers: BenA = KJ834478; CaM = KM089363; RPB2 = JN406583).

Penicillium olsonii Bainier & Sartory, Ann. Mycol. 10: 398. 1912. [MB121021]. — Type: IMI 192502. Ex-type: CBS 232.60 = IBT 23473 = IMI 192502 = NRRL 13058 = NRRL 13716. Infragen. class: subgen. Penicillium, sect. Brevicompacta, ser. Olsoniorum. Reproduction: asexual. ITS barcode: EU587341 (alternative markers: BenA = AY674445; CaM = DQ658165; RPB2 = JN121464).

Penicillium onobense C. Ramírez & A.T. Martínez, Mycopathologia 74: 44. 1981. [MB112525]. — Type: CBS 174.81. Ex-type: CBS 174.81 = DTO 036-A8 = ATCC 42225 = IJFM 3026 = VKMF-2183. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Simplicissima. Reproduction: asexual. ITS barcode: GU981575 (alternative markers: BenA = GU981627; CaM = MN969281; RPB2 = KF296447).

Penicillium oregonense Visagie & Samson, Persoonia 36: 145. 2016. [MB808268]. — Type: CBS H-21607. Ex-type: CBS 129775 = DTO 208-A5. Infragen. class: subgen. Aspergilloides, sect. Torulomyces, ser. Torulomyces. Reproduction: asexual. ITS barcode: KF303668 (alternative markers: BenA = KF303623; CaM = KF303640; RPB2 = KF303710).

Penicillium ornatum Udagawa, Trans. Mycol. Soc. Japan 9: 49. 1968. [MB335756]. — Type: NHL 6101. Ex-type: CBS 190.68 = ATCC 18608 = IFO 31739 = IMI 137977 = NHL 6101 = NRRL 3471. Infragen. class: subgen. Aspergilloides, sect. Ramigena, ser. Ramigena. Reproduction: homothallic. ITS barcode: KC411687 (alternative markers: BenA = KJ834479; CaM = MN969282; RPB2 = JN121451).

Penicillium ortum Visagie & K. Jacobs, Mycol. Prog. 14 (no. 96): 18. 2015. [MB809820]. — Type: CBS H-21602. Ex-type: CBS 135669 = CV 0102 = DTO 180-I9. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Janthinella. Reproduction: asexual. ITS barcode: JX091427 (alternative markers: BenA = JX091520; CaM = JX141551; RPB2 = KF296443).

Penicillium osmophilum Stolk & Veenb.-Rijks, Antonie van Leeuwenhoek 40: 1. 1974. [MB319288]. — Type: CBS 462.72. Ex-type: DTO 092-C5 = CBS 462.72 = CBS 439.73 = IBT 14678 = NRRL 5922. Infragen. class: subgen. Penicillium, sect. Osmophila, ser. Osmophila. Reproduction: homothallic. ITS barcode: EU427295 (alternative markers: BenA = MN969391; CaM = KU896846; RPB2 = JN121518).

Penicillium ovatum (K. Ando & Nawawi) Houbraken & Samson, Stud. Mycol. 70: 48. 2011. [MB561961]. Basionym: Torulomyces ovatus K. Ando & Nawawi, Mycoscience 39: 317. 1998. [MB447111]. — Type: TNS-F-238518. Ex-type: CBS 136664 = KY 12726 = DTO 270-G7. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Erubescentia. Reproduction: asexual. ITS barcode: KF667370 (alternative markers: BenA = KF667366; CaM = KF667368; RPB2 = KF667372).

Penicillium oxalicum Currie & Thom, J. Biol. Chem. 22: 289. 1915. [MB121033]. — Type: IMI 192332. Ex-type: CBS 219.30 = ATCC 1126 = FRR 787 = IMI 192332 = MUCL 29047 = NRRL 787 = QM 7606. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Oxalica. Reproduction: asexual. ITS barcode: AF033438 (alternative markers: BenA = KF296462; CaM = MN969283; RPB2 = JN121456).

Penicillium pagulum Visagie & K. Jacobs, IMA Fungus 7: 102. 2016. [MB811005]. — Type: CBS H-22049. Ex-type: CBS 139166 = DAOMC 241069 = DTO 183-H2 = CV 2224. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Corylophila. Reproduction: asexual. ITS barcode: JX140898 (alternative markers: BenA = JX141070; CaM = JX157519; RPB2 = KP064655).

Penicillium palitans Westling, Ark Bot. 11: 83. 1911. [MB203604]. — Type: CBS H-7531. Ex-type: DTO 206-F6 = CBS 107.11 = ATCC 10477 = IBT 23034 = IMI 040215 = NRRL 2033 = VKMF-3088. Infragen. class: subgen. Penicillium, sect. Fasciculata, ser. Camembertiorum. Reproduction: asexual. ITS barcode: KJ834514 (alternative markers: BenA = KJ834480; CaM = KU896847; RPB2 = KU904360).

Penicillium palmense C. Ramírez & A.T. Martínez, Mycopathologia 66: 80. 1978. [MB319289]. — Type: CBS 336.79. Ex-type: CBS 336.79 = ATCC 38669 = IJFM 3840. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Spinulosa. Reproduction: asexual. ITS barcode: KJ834515 (alternative markers: BenA = GQ367508; CaM = GQ367534; RPB2 = JN406566).

Penicillium pancosmium Houbraken et al., Stud. Mycol. 70: 108. 2011. [MB563191]. — Type: CBS H-20651. Ex-type: CBS 276.75 = DTO 031-B4 = DAOM 147467 = IBT 29991. Infragen. class: subgen. Aspergilloides, sect. Citrina, ser. Westlingiorum. Reproduction: asexual. ITS barcode: JN617660 (alternative markers: BenA = JN606790; CaM = MN969284; RPB2 = MN969130).

Penicillium paneum Frisvad, Microbiology 142: 546. 1996. [MB415570]. — Type: C 25000. Ex-type: CBS 101032 = IBT 21541 = IBT 12407. Infragen. class: subgen. Penicillium, sect. Roquefortorum, ser. Roquefortorum. Reproduction: asexual. ITS barcode: HQ442346 (alternative markers: BenA = AY674387; CaM = HQ442331; RPB2 = KU904361).

Penicillium panissanguineum Visagie et al., Persoonia 36: 275. 2016. [MB815783]. — Type: DAOM 695771. Ex-type: DAOMC 250562 = CBS 140989 = DTO 410-D8 = W 93 = KAS 2209. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Simplicissima. Reproduction: asexual. ITS barcode: KT887862 (alternative markers: BenA = KT887823; CaM = KT887784; RPB2 = MN969182).

Penicillium paradoxum (Fennell & Raper) Samson et al., Stud. Mycol. 78: 352. 2014. [MB547045]. Basionym: Aspergillus paradoxus Fennell & Raper, Mycologia 47: 69. 1955. [MB292853]. — Type: IMI 061446. Ex-type: CBS 527.65 = NRRL 2162 = ATCC 16918 = IMI 061446. Infragen. class: subgen. Penicillium, sect. Paradoxa, ser. Paradoxa. Reproduction: homothallic. ITS barcode: EF669707 (alternative markers: BenA = EF669683; CaM = EF669692; RPB2 = EF669670).

Penicillium paraherquei S. Abe ex G. Sm., Trans. Brit. Mycol. Soc. 46: 335. 1963. [MB302412]. — Type: IMI 68220. Ex-type: CBS 338.59 = DTO 015-D4 = DTO 097-F3 = ATCC 22354 = ATCC 46903 = FAT964 = FRR 3454 = IFO 6234 = IMI 068220 = NRRL 3454. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Simplicissima. Reproduction: asexual. ITS barcode: AF178511 (alternative markers: BenA = KF296465; CaM = MN969285; RPB2 = KF296449).

Penicillium parviverrucosum (K. Ando & Pitt) Houbraken & Samson, Stud. Mycol. 70: 48. 2011. [MB561962]. Basionym: Torulomyces parviverrucosus K. Ando & Pitt, Mycoscience 39: 317. 1998. [MB447109]. — Type: TNS-F-238516. Ex-type: KY 12720. Infragen. class: subgen. Aspergilloides, sect. Torulomyces, ser. Torulomyces. Reproduction: asexual. ITS barcode: n.a. (alternative markers: BenA = n.a.; CaM = n.a.; RPB2 = n.a.).

Penicillium parvofructum Guevara-Suarez et al., Persoonia 38: 353. 2017. [MB819947]. — Type: CBS H-22733. Ex-type: FMR 15047 = CBS 141690 = DTO 410-E6. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Erubescentia. Reproduction: asexual. ITS barcode: LT559091 (alternative markers: BenA = LT627645; CaM = LT627646; RPB2 = MN969197).

Penicillium parvulum S.W. Peterson & B.W. Horn, Mycologia 101: 75. 2009. [MB509289]. — Type: BPI 877331. Ex-type: CBS 132825 = NRRL 35504. Infragen. class: subgen. Aspergilloides, sect. Cinnamopurpurea, ser. Cinnamopurpurea. Reproduction: asexual. ITS barcode: EF422845 (alternative markers: BenA = EF506218; CaM = EF506225; RPB2 = MN969131).

Penicillium parvum Raper & Fennell, Mycologia 40: 508. 1948. [MB289101]. — Type: CBS 359.48. Ex-type: CBS 359.48 = ATCC 10479 = IFO 7732 = IMI 040587 = NRRL 2095 = QM 1878. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Erubescentia. Reproduction: homothallic. ITS barcode: AF033460 (alternative markers: BenA = HQ646568; CaM = KF900173; RPB2 = JN406559).

Penicillium pasqualense Houbraken et al., Stud. Mycol. 70: 108. 2011. [MB563190]. — Type: CBS H-20663. Ex-type: CBS 126330 = DTO 080-D5 = IBT 14235. Infragen. class: subgen. Aspergilloides, sect. Citrina, ser. Westlingiorum. Reproduction: asexual. ITS barcode: JN617676 (alternative markers: BenA = JN606673; CaM = MN969286; RPB2 = JN606617).

Penicillium paxilli Bainier, Bull. Soc. Mycol. France 23: 95. 1907. [MB203838]. — Type: IMI 40226. Ex-type: CBS 360.48 = ATCC 10480 = FRR 2008 = IMI 040226 = NRRL 2008 = QM 725. Infragen. class: subgen. Aspergilloides, sect. Citrina, ser. Paxillorum. Reproduction: asexual. ITS barcode: GU944577 (alternative markers: BenA = JN606844; CaM = JN606566; RPB2 = JN606610).

Penicillium pedernalense Laich & J. Andrade, Index Fungorum 361: 1. 2018. [MB554533]. — Type: CBS 140770. Ex-type: CBS 140770 = CECT 20949 = DTO 366-A3. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Simplicissima. Reproduction: asexual. ITS barcode: KU255398 (alternative markers: BenA = KU255396; CaM = MN969322; RPB2 = MN969184).

Penicillium penarojense Houbraken et al., Int. J. Syst. Evol. Microbiol. 61: 1471. 2011. [MB518024]. — Type: HUA 170335. Ex-type: CBS 113178 = DTO 056-D1 = DTO 297-H9 = IBT 23262. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Dalearum. Reproduction: asexual. ITS barcode: GU981570 (alternative markers: BenA = GU981646; CaM = MN969287; RPB2 = KF296450).

Penicillium persicinum L. Wang et al., Antonie van Leeuwenhoek 86: 177. 2004. [MB500259]. — Type: HMAS 80638-1-4. Ex-type: CBS 111235 = AS 3.5891 = IBT 24565. Infragen. class: subgen. Penicillium, sect. Chrysogena, ser. Persicina. Reproduction: asexual. ITS barcode: JX997072 (alternative markers: BenA = JF909951; CaM = JX996954; RPB2 = JN406644).

Penicillium philippinense Udagawa & Y. Horie, J. Jap. Bot. 47: 341. 1972. [MB319291]. — Type: NHL 6130. Ex-type: CBS 623.72 = FRR 1532 = NHL 6130. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Restricta. Reproduction: homothallic. ITS barcode: KC411770 (alternative markers: BenA = KJ834482; CaM = KP016799; RPB2 = JN406543).

Penicillium phoeniceum J.F.H. Beyma, Zentralbl. Bakteriol. Parasitenk., Abt. 2 88: 136. 1933. [MB274284]. — Type: IMI 40585. Ex-type: CBS 249.32 = ATCC 10481 = IJFM 5122 = IMI 040585 = NRRL 2070 = QM 7608 = VKMF-321. Infragen. class: subgen. Aspergilloides, sect. Charlesia, ser. Phoenicea. Reproduction: asexual. ITS barcode: KC411711 (alternative markers: BenA = KJ834483; CaM = AY741729; RPB2 = JN406597).

Penicillium pimiteouiense S.W. Peterson, Mycologia 91: 271. 1999. [MB460126]. — Type: BPI 806262. Ex-type: CBS 102479 = NRRL 25542. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Erubescentia. Reproduction: asexual. ITS barcode: AF037431 (alternative markers: BenA = HQ646569; CaM = HQ646580; RPB2 = JN406650).

Penicillium piscarium Westling, Ark. Bot. 11: 86. 1911. [MB211321]. — Type: IMI 40032. Ex-type: CBS 362.48 = DTO 014-G9 = DTO 100-C1 = ATCC 10482 = FRR 1075 = IFO 8111 = IMI 040032 = NRRL 1075 = VKMF-1823. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Rolfsiorum. Reproduction: asexual. ITS barcode: GU981600 (alternative markers: BenA = GU981668; CaM = MN969288; RPB2 = KF296451).

Penicillium polonicum K.W. Zaleski, Bull. Int. Acad. Polon. Sci., Sér. B., Sci. Nat. 1927: 445. 1927. [MB274889]. — Type: CBS 222.28. Ex-type: CBS 222.28 = IBT 12821 = IMI 291194 = MUCL 29204 = NRRL 995. Infragen. class: subgen. Penicillium, sect. Fasciculata, ser. Viridicata. Reproduction: asexual. ITS barcode: AF033475 (alternative markers: BenA = MN969392; CaM = KU896848; RPB2 = JN406609).

Penicillium porphyreum Houbraken & Samson, Stud. Mycol. 70: 48. 2011. [MB561959]. Replaced synonym: Monocillium humicola var. brunneum M. Chr. & Backus, Mycologia 56: 498.1964. [MB353642]. — Type: NY 00985491. Ex-type: CBS 382.64 = KY 12723 = DTO 078-G7. Infragen. class: subgen. Aspergilloides, sect. Torulomyces, ser. Torulomyces. Reproduction: asexual. ITS barcode: KF303666 (alternative markers: BenA = KF303621; CaM = KF303636; RPB2 = KF303677).

Penicillium psychrosexuale [as “psychrosexualis”] Houbraken & Samson, IMA Fungus 1: 174. 2010. [MB834590]. — Type: CBS H-20501. Ex-type: DTO 070-G9 = CBS 128137 = IBT 29551. Infragen. class: subgen. Penicillium, sect. Roquefortorum, ser. Roquefortorum. Reproduction: homothallic. ITS barcode: HQ442345 (alternative markers: BenA = HQ442356; CaM = HQ442330; RPB2 = KU904362).

Penicillium psychrotrophicum Hyang B. Lee et al., Fungal Diversity 96: 103. 2019. [MB555409]. — Type: CBS H-22504. Ex-type: CBS 140670 = DTO 327-C9 = EML-COD3 = IBT 33673. Infragen. class: subgen. Penicillium, sect. Fasciculata, ser. Corymbifera. Reproduction: asexual. ITS barcode: KP941754 (alternative markers: BenA = KP900995; CaM = KP900994; RPB2 = MN969212).

Penicillium pullum S.W. Peterson & Sigler, Mycol. Res. 106: 1115. 2002. [MB483982]. — Type: BPI 841398. Ex-type: CBS 331.48 = ATCC 10447 = NRRL 721 = FRR 721 = IFO 6097 = IMI 39747 = QM 1925 = Thom 5179.4. Infragen. class: subgen. Aspergilloides, sect. Stolkia, ser. Stolkia. Reproduction: asexual. ITS barcode: AF033443 (alternative markers: BenA = JN617719; CaM = AF481134; RPB2 = MN969132).

Penicillium pulvillorum Turfitt, Trans. Brit. Mycol. Soc. 23: 186. 1939. [MB275682]. — Type: CBS 280.39. Ex-type: CBS 280.39 = DTO 014-G7 = DTO 094-D5 = IFO 7763 = NRRL 2026. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Rolfsiorum. Reproduction: asexual. ITS barcode: AF178517 (alternative markers: BenA = GU981670; CaM = MN969289; RPB2 = KF296452).

Penicillium pulvis Houbraken et al., Stud. Mycol. 78: 429. 2014. [MB809970]. — Type: CBS H-21878. Ex-type: CBS 138432 = DTO 180-B7. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Glabra. Reproduction: asexual. ITS barcode: KM189632 (alternative markers: BenA = KM088876; CaM = KM089263; RPB2 = KM089650).

Penicillium punicae Hyang B. Lee et al., Fungal Diversity 83: 103. 2017. [MB818233]. — Type: CNUFC-FP2-1. Ex-type: JMRC:SF:12421. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Corylophila. Reproduction: asexual. ITS barcode: n.a. (alternative markers: BenA = KX839673; CaM = KX839671; RPB2 = KX839675).

Penicillium purpurescens [as “purpurascens”] (Sopp) Biourge, La Cellule 33: 105. 1923. [MB335761]. Basionym: Citromyces purpurrescens Sopp, Skr. Vidensk.-Selsk. Christiana, Math.-Naturvidensk. Kl. 11: 117. 1912. [MB568761]. — Type: IMI 39745. Ex-type: CBS 366.48 = NRRL 720 = FRR 720 = ATCC 10485 = IMI 39745. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Glabra. Reproduction: asexual. ITS barcode: KM189561 (alternative markers: BenA = KM088801; CaM = KM089186; RPB2 = KM089573).

Penicillium pusillum G. Sm., Trans. Brit. Mycol. Soc. 22 (3–4): 254. 1939. [MB275810]. — Type: unknown. Ex-type: CBS 312.63 = FRR 1541 = IMI 089286 = LSHBB147 = NRRL 2498. Infragen. class: subgen. Aspergilloides, sect. Cinnamopurpurea, ser. Jiangxiensia. Reproduction: asexual. ITS barcode: EF626951 (alternative markers: BenA = KF932925; CaM = KF932941; RPB2 = KF932995).

Penicillium quebecense Houbraken et al., Stud. Mycol. 70: 111. 2011. [MB563202]. — Type: CBS H-20666. Ex-type: CBS 101623 = DTO 009-B8 = IBT 29050. Infragen. class: subgen. Aspergilloides, sect. Citrina, ser. Westlingiorum. Reproduction: asexual. ITS barcode: JN617661 (alternative markers: BenA = JN606700; CaM = JN606509; RPB2 = JN606622).

Penicillium quercetorum Baghd., Novosti Sist. Nizsh. Rast. 5: 110. 1968. [MB335762]. — Type: CBS H-7527. Ex-type: CBS 417.69 = NRRL 3758 = ATCC 48727 = CCRC 31668 = FRR 516 = IFO 31749 = IMI 140342 = MUCL 31203 = VKMF-1074. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Quercetorum. Reproduction: asexual. ITS barcode: KM189556 (alternative markers: BenA = KM088795; CaM = KM089180; RPB2 = KM089567).

Penicillium raciborskii K.W. Zaleski, Bull. Int. Acad. Polon. Sci., Sér. B., Sci. Nat. 1927: 454. 1927. [MB276002]. — Type: IMI 40568. Ex-type: CBS 224.28 = ATCC 10488 = DSM2422 = FRR 2150 = IFO 7676 = IMI 040568 = LSHBP 92 = MUCL 29246 = NRRL 2150. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Lapidosa. Reproduction: asexual. ITS barcode: AF033447 (alternative markers: BenA = JX141069; CaM = KP016800; RPB2 = JN406607).

Penicillium radiatolobatum Lörinczi, Publ. Soc. Nat. Rom. Pent. Stiinta Sol. 10B: 435. 1972. [MB114326]. — Type: CBS H-7530. Ex-type: CBS 340.79. Infragen. class: subgen. Penicillium, sect. Canescentia, ser. Canescentia. Reproduction: asexual. ITS barcode: KC411745 (alternative markers: BenA = MN969413; CaM = MT066183; RPB2 = MN969168).

Penicillium radicicola Overy & Frisvad, Syst. Appl. Microbiol. 26: 633. 2003. [MB488233]. — Type: C 60161. Ex-type: CBS 112430 = IBT 10696 = DTO 051-E1. Infragen. class: subgen. Penicillium, sect. Fasciculata, ser. Corymbifera. Reproduction: asexual. ITS barcode: KJ834516 (alternative markers: BenA = MN969393; CaM = MN969290; RPB2 = MN969133).

Penicillium raistrickii G. Sm., Trans. Brit. Mycol. Soc. 18: 90. 1933. [MB276069]. — Type: IMI 40221. Ex-type: CBS 261.33 = ATCC 10490 = FRR 1044 = IFO 6104 = IMI 040221 = LSHBB100 = NRRL 1044 = NRRL 2039 = QM 1936 = VKMF-337. Infragen. class: subgen. Penicillium, sect. Ramosum, ser. Raistrickiorum. Reproduction: asexual. ITS barcode: AY373927 (alternative markers: BenA = KJ834485; CaM = KJ867006; RPB2 = JN406592).

Penicillium ramusculum Bat. & H. Maia, Anais Soc. Biol. Pernambuco 13: 27. 1955. [MB302419]. — Type: unknown. Ex-type: CBS 251.56 = ATCC 12292 = FRR 3459 = IMI 063546 = IMUR478 = LSHBBB324 = NRRL 3459 = QM 7057. Infragen. class: subgen. Aspergilloides, sect. Ramigena, ser. Ramigena. Reproduction: asexual. ITS barcode: EF433765 (alternative markers: BenA = EU427269; CaM = EU427278; RPB2 = JN121472).

Penicillium ranomafanaense Houbraken & Hagen, Stud. Mycol. 78: 433. 2014. [MB809971]. — Type: CBS H-21862. Ex-type: CBS 137953 = DTO 085-A5. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Verhageniorum. Reproduction: asexual. ITS barcode: KM189541 (alternative markers: BenA = KM088779; CaM = KM089164; RPB2 = KM089551).

Penicillium raperi G. Sm., Trans. Brit. Mycol. Soc. 40: 486. 1957. [MB302421]. — Type: IMI 71625. Ex-type: CBS 281.58 = DTO 097-F6 = DTO 014-H4 = ATCC 22355 = IFO 8179 = IMI 071625 = LSHBBB338 = NRRL 2674 = QM 7527. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Janthinella. Reproduction: asexual. ITS barcode: AF033433 (alternative markers: BenA = GU981622; CaM = MN969291; RPB2 = KF296453).

Penicillium raphiae Houbraken et al., Stud. Mycol. 70: 114. 2011. [MB563203]. — Type: CBS H-20660. Ex-type: CBS 126234 = DTO 078-B8 = IBT 22407. Infragen. class: subgen. Aspergilloides, sect. Citrina, ser. Westlingiorum. Reproduction: asexual. ITS barcode: JN617673 (alternative markers: BenA = JN606657; CaM = MN969292; RPB2 = JN606619).

Penicillium reconvexovelosoi J.P. Andrade et al., Persoonia 43: 383, 2019. [MB832747]. — Type: HURB 18575. Ex-type: CCDCA 11500. Infragen. class: subgen. Aspergilloides, sect. Sclerotiorum, ser. Adametziorum. Reproduction: asexual. ITS barcode: n.a. (alternative markers: BenA = MN497417; CaM = MN497418; RPB2 = n.a.).

Penicillium repensicola Visagie & K. Jacobs, IMA Fungus 7: 102. 2016. [MB811006]. — Type: CBS H-22047. Ex-type: CBS 139160 = DAOMC 241080 = DTO 183-B8 = CV 1495. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Corylophila. Reproduction: asexual. ITS barcode: JX140893 (alternative markers: BenA = JX141150; CaM = JX157490; RPB2 = KP064660).

Penicillium restingae J.P. Andrade et al., Persoonia 32: 293. 2014. [MB807051]. — Type: CMR H-12. Ex-type: CBS 140379 = URM 7075 = DTO 331-H7. Infragen. class: subgen. Aspergilloides, sect. Sclerotiorum, ser. Adametziorum. Reproduction: asexual. ITS barcode: KF803355 (alternative markers: BenA = KF803349; CaM = KF803352; RPB2 = MN969134).

Penicillium restrictum J.C. Gilman & E.V. Abbott, Iowa St. Coll. J. Sci. 1: 297. 1927. [MB276289]. — Type: IMI 40228. Ex-type: CBS 367.48 = ATCC 11257 = FRR 1748 = IMI 040228 = NRRL 1748 = QM 1962. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Restricta. Reproduction: asexual. ITS barcode: AF033457 (alternative markers: BenA = KJ834486; CaM = KP016803; RPB2 = JN121506).

Penicillium reticulisporum Udagawa, Trans. Mycol. Soc. Japan 9: 52. 1968. [MB335763]. — Type: NHL 6105. Ex-type: CBS 122.68 = ATCC 18566 = IFO 9024 = IMI 136700 = NHL 6105 = NRRL 3447 = DTO 097-C4. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Janthinella. Reproduction: homothallic. ITS barcode: AF033437 (alternative markers: BenA = MN969394; CaM = MN969293; RPB2 = KF296454).

Penicillium ribium Frisvad & Overy, Int. J. Syst. Evol. Microbiol. 56: 1436. 2006. [MB501061]. — Type: DAOM 234091. Ex-type: CBS 127809 = DAOM 234091 = IBT 16537 = IBT 24431. Infragen. class: subgen. Penicillium, sect. Ramosum, ser. Lanosa. Reproduction: asexual. ITS barcode: DQ267916 (alternative markers: BenA = MN969395; CaM = KJ866995; RPB2 = JN406631).

Penicillium riverlandense Visagie & K. Jacobs, Persoonia 36: 149. 2016. [MB808269]. — Type: CBS H-21606. Ex-type: CBS 135896 = CV 0979 = DTO 182-F6 = DAOMC 241060. Infragen. class: subgen. Aspergilloides, sect. Torulomyces, ser. Torulomyces. Reproduction: asexual. ITS barcode: JX091457 (alternative markers: BenA = JX091580; CaM = JX141593; RPB2 = KF303685).

Penicillium robsamsonii Frisvad & Houbraken, Persoonia 36: 313. 2016. [MB815872]. — Type: CBS H-22341. Ex-type: CBS 140573 = IBT 29466 = DTO 149-B6. Infragen. class: subgen. Penicillium, sect. Robsamsonia, ser. Robsamsonia. Reproduction: asexual. ITS barcode: KU904339 (alternative markers: BenA = KT698885; CaM = KT698894; RPB2 = KT698904).

Penicillium rolfsii Thom, Penicillia: 489. 1930. [MB276674]. — Type: IMI 40029. Ex-type: CBS 368.48 = DTO 106-H8 = ATCC 10491 = FRR 1078 = IFO 7735 = IMI 040029 = MUCL 29229 = NRRL 1078 = QM 1961. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Rolfsiorum. Reproduction: asexual. ITS barcode: JN617705 (alternative markers: BenA = GU981667; CaM = MN969294; RPB2 = KF296455).

Penicillium roqueforti Thom, U.S.D.A. Bur. Animal Industr. Bull. 82: 35. 1906. [MB213525]. — Type: IMI 24313. Ex-type: CBS 221.30 = ATCC 10110 = ATCC 1129 = CECT 2905 = IBT 6754 = IFO 5459 = IMI 024313 = LSHBP 93 = NCTC 588 = NRRL 849 = QM 1937. Infragen. class: subgen. Penicillium, sect. Roquefortorum, ser. Roquefortorum. Reproduction: Heterothallic (Ropars et al. 2014). ITS barcode: HQ442347 (alternative markers: BenA = MN969396; CaM = HQ442332; RPB2 = JN406611).

Penicillium roseomaculatum Biourge, Cellule 33: 301. 1923. [MB276785]. — Type: unknown. Ex-type: CBS 137962 = IMI 189696 = NRRL 728 = FRR 728. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Spinulosa. Reproduction: asexual. ITS barcode: KM189755 (alternative markers: BenA = KM089004; CaM = KM089391; RPB2 = KM089778).

Penicillium roseopurpureum Dierckx, Ann. Soc. Sci. Bruxelles 25: 86. 1901. [MB213447]. — Type: IMI 40573. Ex-type: CBS 226.29 = ATCC 10492 = ATHUM2895 = FRR 2064 = IMI 040573 = MUCL 28654 = MUCL 29237 = NRRL 2064 = NRRL 2064A. Infragen. class: subgen. Aspergilloides, sect. Citrina, ser. Roseopurpurea. Reproduction: asexual. ITS barcode: GU944605 (alternative markers: BenA = JN606838; CaM = JN606556; RPB2 = JN606613).

Penicillium roseoviride Stapp & Bortels, Zentralbl. Bakteriol. Parasitenk., Abt. 2 93: 51. 1935. [MB492646]. — Type: unknown. Ex-type: CBS 267.35 = ATCC 10412 = IFO 6089 = IMI 039740ii = NRRL 760 = QM 7485. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Thomiorum. Reproduction: asexual. ITS barcode: KM189549 (alternative markers: BenA = KM088787; CaM = KM089172; RPB2 = KM089559).

Penicillium rubefaciens Quintan., Mycopathologia 80: 73. 1982. [MB109998]. — Type: CBS 145.83. Ex-type: CBS 145.83 = CECT 2752. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Corylophila. Reproduction: asexual. ITS barcode: KC411677 (alternative markers: BenA = KJ834487; CaM = KP016804; RPB2 = JN406627).

Penicillium rubens Biourge, Cellule 33: 265. 1923. [MB276884]. — Type: CBS H-20595. Ex-type: DTO 098-E8 = CBS 129667 = NRRL 792 = IBT 30129 = ATCC 9783. Infragen. class: subgen. Penicillium, sect. Chrysogena, ser. Chrysogena. Reproduction: Heterothallic (Böhm et al. 2013; reported as P. chrysogenum, Houbraken et al. 2014a). ITS barcode: JX997057 (alternative markers: BenA = JF909949; CaM = JX996263; RPB2 = JX996658).

Penicillium rubidurum Udagawa & Y. Horie, Trans. Mycol. Soc. Japan 14: 381. 1973. [MB319295]. — Type: NHL 6460. Ex-type: CBS 609.73 = NRRL 6033 = ATCC 28051 = ATCC 48238 = FRR 1558 = IMI 228551 = NHL 6460. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Erubescentia. Reproduction: homothallic. ITS barcode: AF033462 (alternative markers: BenA = HQ646574; CaM = HQ646585; RPB2 = JN406545).

Penicillium rubriannulatum L. Cai et al., Cladistics 35: 539. 2018 [2019]. [MB818162]. — Type: HMAS 247732. Ex-type: CGMCC 3.18804 = NN072456 = CBS 144641. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Dalearum. Reproduction: asexual. ITS barcode: KY495029 (alternative markers: BenA = KY495138; CaM = MN969336; RPB2 = KY495080).

Penicillium rudallense Houbraken et al., Stud. Mycol. 78: 433. 2014. [MB809972]. — Type: CBS H-21867. Ex-type: CBS 138162 = FRR 6085 = DTO 056-I4. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Glabra. Reproduction: asexual. ITS barcode: KM189504 (alternative markers: BenA = KM088741; CaM = KM089126; RPB2 = KM089513).

Penicillium sacculum E. Dale, Ann. Mycol. 24: 137. 1926. [MB277209]. — Type: CBS 231.61. Ex-type: CBS 231.61 = ATCC 18350 = IFO 8114 = IFO 9454 = IMI 051498 = LSHBBB298 = UC4505. Infragen. class: subgen. Penicillium, sect. Eladia, ser. Eladia. Reproduction: asexual. ITS barcode: KC411707 (alternative markers: BenA = KJ834488; CaM = KU896849; RPB2 = JN121462).

Penicillium sajarovii Quintan., Av. Aliment. Majora Anim. 22: 539. 1981. [MB114172]. — Type: CBS 277.83. Ex-type: CBS 277.83 = DTO 334-D4 = CECT 2751 = IMI 259992. Infragen. class: subgen. Penicillium, sect. Ramosum, ser. Raistrickiorum. Reproduction: asexual. ITS barcode: KC411724 (alternative markers: BenA = MN969397; CaM = MN969295; RPB2 = JN406588).

Penicillium salamii G. Perrone et al., Int. J. Food Microb. 193: 93. 2014. [MB809645]. — Type: CBS H-21341. Ex-type: CBS 135391 = DTO 198-E1 = ITEM 15291. Infragen. class: subgen. Penicillium, sect. Brevicompacta, ser. Olsoniorum. Reproduction: asexual. ITS barcode: HG514431 (alternative markers: BenA = HG514437; CaM = HG514432; RPB2 = MN969160).

Penicillium salmoniflumine S.W. Peterson et al., PLoS ONE 10: 0121987, 21. 2015. [MB807374]. — Type: BPI 881286. Ex-type: NRRL 35837 = IBT 29673. Infragen. class: subgen. Aspergilloides, sect. Cinnamopurpurea, ser. Idahoensia. Reproduction: asexual. ITS barcode: KF932960 (alternative markers: BenA = KF932928; CaM = KF932945; RPB2 = KF932999).

Penicillium samsonianum L. Wang et al., Persoonia 36: 313. 2016. [MB815873]. — Type: HMAS 245107. Ex-type: AS3.15403 = CBS 138919 = IBT 33392 = DTO 316-B7. Infragen. class: subgen. Penicillium, sect. Osmophila, ser. Samsoniorum. Reproduction: asexual. ITS barcode: KJ668590 (alternative markers: BenA = KJ668582; CaM = KJ668586; RPB2 = KT698899).

Penicillium sanguifluum (Sopp) Biourge, Cellule 33: 105. 1923. [MB356682]. Basionym: Citromyces sanguifluus Sopp, Skr. Vidensk.-Selsk. Christiana Math.-Nat. Kl. 11: 115. 1912. [MB491120]. — Type: CBS H-20645. Ex-type: CBS 127032 = IBT 29041 = DTO 020-B7. Infragen. class: subgen. Aspergilloides, sect. Citrina, ser. Roseopurpurea. Reproduction: asexual. ITS barcode: JN617681 (alternative markers: BenA = JN606819; CaM = JN606555; RPB2 = MN969135).

Penicillium sanshaense X.C. Wang & W.Y. Zhuang, Sci. Rep. 7: 8233, 9. 2017. [MB570337]. — Type: HMAS 248820. Ex-type: CGMCC 3.18413. Infragen. class: subgen. Aspergilloides, sect. Sclerotiorum, ser. Herqueorum. Reproduction: asexual. ITS barcode: KX885070 (alternative markers: BenA = KX885050; CaM = KX885060; RPB2 = n.a.).

Penicillium saturniforme (L. Wang & W.Y. Zhuang) Houbraken & Samson, Stud. Mycol. 70: 48. 2011. [MB561958]. Basionym: Eupenicillium saturniforme L. Wang & W.Y. Zhuang Mycopathologia 167: 300. 2009. [MB541663]. — Type: AS 3.6886. Ex-type: CBS 122276 = AS 3.6886. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Saturniformia. Reproduction: homothallic. ITS barcode: EU644081 (alternative markers: BenA = EU644080; CaM = EU644062; RPB2 = JN121439).

Penicillium scabrosum Frisvad et al., Persoonia 14: 177. 1990. [MB136735]. — Type: IMI 285533. Ex-type: CBS 683.89 = FRR 2950 = IBT 3736 = IMI 285533 = DAOM 214786. Infragen. class: subgen. Penicillium, sect. Ramosum, ser. Scabrosa. Reproduction: asexual. ITS barcode: DQ267906 (alternative markers: BenA = DQ285610; CaM = FJ530987; RPB2 = JN406541).

Penicillium sclerotigenum W. Yamam., Sci. Rep. Hyogo Univ Agric. 1: 69. 1955. [MB302424]. — Type: IMI 68616. Ex-type: DTO 128-D7 = CBS 101033 = CBS 343.59 = ATCC 18488 = IBT 14346 = IFO 6167 = IMI 068616 = NRRL 3461 = QM 7779. Infragen. class: subgen. Penicillium, sect. Penicillium, ser. Sclerotigena. Reproduction: asexual. ITS barcode: AF033470 (alternative markers: BenA = AY674393; CaM = KU896850; RPB2 = JN406652).

Penicillium sclerotiorum J.F.H. Beyma, Zentralbl. Bakteriol. Parasitenk., Abt. 2 96: 418. 1937. [MB277708]. — Type: IMI 40569. Ex-type: CBS 287.36 = ATCC 10494 = IFO 6105 = IMI 040569 = NRRL 2074 = QM 1938 = VKMF-353. Infragen. class: subgen. Aspergilloides, sect. Sclerotiorum, ser. Sclerotiorum. Reproduction: asexual. ITS barcode: JN626132 (alternative markers: BenA = JN626001; CaM = JN626044; RPB2 = JN406585).

Penicillium senticosum D.B. Scott, Mycopathol. Mycol. Appl. 36: 5. 1968. [MB335764]. — Type: CBS 316.67. Ex-type: CBS 316.67 = ATCC 18623 = CSIR 1042 = IMI 136211 = IMI 216905. Infragen. class: subgen. Penicillium, sect. Eladia, ser. Eladia. Reproduction: homothallic. ITS barcode: KC411733 (alternative markers: BenA = KJ834490; CaM = MN969296; RPB2 = MN969136).

Penicillium setosum Tijith et al., Mycology 10: 55. 2018. [MB818581]. — Type: WSR 62. Ex-type: CBS 144865 = MCC 1370 = NCFT NO 8222.16 = AMH-9974. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Janthinella. Reproduction: asexual. ITS barcode: KT852579 (alternative markers: BenA = MF184995; CaM = MH105905; RPB2 = MH016196).

Penicillium shearii Stolk & D.B. Scott, Persoonia 4: 396. 1967. [MB335765]. — Type: CBS 290.48. Ex-type: CBS 290.48 = ATCC 10410 = IFO 6088 = IMI 039739 = IMI 039739iv = NRRL 715 = QM 1870. Infragen. class: subgen. Aspergilloides, sect. Citrina, ser. Sheariorum. Reproduction: homothallic. ITS barcode: GU944606 (alternative markers: BenA = JN606840; CaM = EU644068; RPB2 = JN121482).

Penicillium shennongjianum [as “shennonghianum”] H.Z. Kong & Z.T. Qi, Mycosystema 1: 110. 1988. [MB587562]. — Type: CBS 228.89. Ex-type: CBS 228.89. Infragen. class: subgen. Aspergilloides, sect. Cinnamopurpurea, ser. Nodula. Reproduction: asexual. ITS barcode: KC411705 (alternative markers: BenA = KJ834491; CaM = AY678561; RPB2 = JN121458).

Penicillium simile Davolos et al., Int. J. Syst. Evol. Microbiol. 62: 457. 2012. [MB509645]. — Type: ATCC MYA-4591. Ex-type: CBS 129191 = ATCC MYA-4591 = DTO 159-F7. Infragen. class: subgen. Penicillium, sect. Ramosum, ser. Raistrickiorum. Reproduction: asexual. ITS barcode: FJ376592 (alternative markers: BenA = FJ376595; CaM = GQ979710; RPB2 = MN969137).

Penicillium simplicissimum (Oudem.) Thom, Penicillia: 335. 1930. [MB278201]. Basionym: Spicaria simplicissima Oudem., Ned. Kruidk. Arch. 2: 763. 1902. [MB245011]. — Type: CUP Jensen. 1912: No. 5921 (CUP). Ex-type: CBS 372.48 = DTO 014-H2 = ATCC 10495 = FRR 902 = IFO 5762 = IMI 039816 = QM 1939. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Simplicissima. Reproduction: asexual. ITS barcode: GU981588 (alternative markers: BenA = GU981632; CaM = MN969297; RPB2 = JN121507).

Penicillium sinaicum Udagawa & S. Ueda, Mycotaxon 14: 266. 1982. [MB110862]. — Type: NHL 2894. Ex-type: CBS 279.82 = NHL 2894. Infragen. class: subgen. Penicillium, sect. Chrysogena, ser. Crustacea. Reproduction: homothallic. ITS barcode: JX997090 (alternative markers: BenA = KU896818; CaM = JX996970; RPB2 = JN406587).

Penicillium singorense Visagie et al., Stud. Mycol. 78: 119. 2014. [MB809182]. — Type: CBS H-21802. Ex-type: CBS 138214 = DTO 133-C6. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Dalearum. Reproduction: asexual. ITS barcode: KJ775674 (alternative markers: BenA = KJ775167; CaM = KJ775403; RPB2 = MN969138).

Penicillium sizovae Baghd., Novosti Sist. Nizsh. Rast. 1968: 103. 1968. [MB335767]. — Type: CBS 413.69. Ex-type: CBS 413.69 = DTO 023-A7 = FRR 518 = IMI 140344 = VKMF-1073. Infragen. class: subgen. Aspergilloides, sect. Citrina, ser. Citrina. Reproduction: asexual. ITS barcode: GU944588 (alternative markers: BenA = GU944535; CaM = MN969298; RPB2 = JN606603).

Penicillium skrjabinii Schmotina & Golovleva, Mikol. Fitopatol. 8: 530. 1974. [MB319296]. — Type: IMI 196528. Ex-type: CBS 439.75 = DTO 095-C8 = NRRL 13055 = FRR 1945 = IMI 196528 = VKMF-1940. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Simplicissima. Reproduction: asexual. ITS barcode: GU981576 (alternative markers: BenA = GU981626; CaM = MN969299; RPB2 = EU427252).

Penicillium smithii Quintan., Av. Aliment. Majora Anim. 23: 340. 1982. [MB114173]. — Type: CBS 276.83. Ex-type: CBS 276.83 = CECT 2744 = IMI 259693. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Lapidosa. Reproduction: asexual. ITS barcode: KC411723 (alternative markers: BenA = KJ834492; CaM = KP016806; RPB2 = JN406589).

Penicillium soliforme L. Cai et al., Cladistics 35: 540. 2018 [2019]. [MB818158]. — Type: HMAS 247733. Ex-type: CGMCC 3.18806 = NN072519 = CBS 144482. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Rolfsiorum. Reproduction: asexual. ITS barcode: KY495038 (alternative markers: BenA = KY495147; CaM = MN969337; RPB2 = KY495047).

Penicillium solitum Westling, Ark. Bot. 11: 65. 1911. [MB206172]. — Type: CBS 424.89. Ex-type: DTO 248-E4 = DTO 047-B2 = CBS 424.89 = ATCC 9923 = CBS 288.36 = FRR 937 = IBT 3948 = IFO 7765 = IMI 039810 = IMI 092225 = LSHBP 52 = MUCL 28668 = MUCL 29173 = NRRL 937. Infragen. class: subgen. Penicillium, sect. Fasciculata, ser. Camembertiorum. Reproduction: asexual. ITS barcode: AY373932 (alternative markers: BenA = MN969398; CaM = KU896851; RPB2 = KU904363).

Penicillium soosanum Kubátová et al., Mycol. Prog. 18 (1-2): 223. 2018. [MB824353]. — Type: PRM 861478. Ex-type: CCF 3778 = MH 344 = CBS 140106 = IBT 30727. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Oxalica. Reproduction: asexual. ITS barcode: FJ430745 (alternative markers: BenA = FM865811; CaM = LT970913; RPB2 = LT797561).

Penicillium soppii K.W. Zaleski, Bull. Int. Acad. Polon. Sci., Cl. Sci. Math., Sér. B., Sci. Nat. 1927: 476. 1927. [MB121424]. — Type: IMI 40217. Ex-type: CBS 226.28 = ATCC 10496 = FRR 2023 = IFO 7766 = IMI 040217 = MUCL 29233 = NRRL 2023 = QM 1964 = IBT 18220. Infragen. class: subgen. Penicillium, sect. Ramosum, ser. Soppiorum. Reproduction: asexual. ITS barcode: AF033488 (alternative markers: BenA = MN969399; CaM = KJ867002; RPB2 = JN406606).

Penicillium spathulatum Frisvad & Samson, FEMS Microbiol. Lett. 339: 88. 2013. [MB492650]. — Type: CBS 117192. Ex-type: DTO 187-D8 = CBS 117192 = IBT 22220. Infragen. class: subgen. Penicillium, sect. Brevicompacta, ser. Buchwaldiorum. Reproduction: asexual. ITS barcode: JX313165 (alternative markers: BenA = MN969400; CaM = JX313149; RPB2 = JN406636).

Penicillium speluncae Visagie & N. Yilmaz, Fungal Syst. Evol. 5: 10. 2020. [MB828614]. — Type: DAOM 745788. Ex-type: DAOMC 251701 = KAS 7512 = P06201. Infragen. class: subgen. Penicillium, sect. Fasciculata, ser. Camembertiorum. Reproduction: asexual. ITS barcode: MG490869 (alternative markers: BenA = MG490889; CaM = MG490959; RPB2 = MN170741).

Penicillium spinuliferum L. Cai & X.Z. Jiang, Cladistics 35: 542. 2018 [2019]. [MB818153]. — Type: HMAS 247734. Ex-type: CGMCC 3.18807 = NN072545 = CBS 144483. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Simplicissima. Reproduction: asexual. ITS barcode: KY495040 (alternative markers: BenA = KY495149; CaM = MN969338; RPB2 = KY495090).

Penicillium spinulosum Thom, U.S.D.A. Bur. Animal Industr. Bull. 118: 76. 1910. [MB215401]. — Type: IMI 24316i. Ex-type: CBS 374.48 = ATCC 10498 = FRR 1750 = IMI 024316 = LSHBAd 29 = MUCL 13910 = MUCL 13911 = NCTC 591 = NRRL 1750 = QM 7654. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Spinulosa. Reproduction: asexual. ITS barcode: AF033410 (alternative markers: BenA = KJ834493; CaM = GQ367524; RPB2 = JN406558).

Penicillium steckii K.W. Zaleski, Bull. Int. Acad. Polon. Sci., Sér. B., Sci. Nat. 1927: 469. 1927. [MB278769]. — Type: IMI 40583. Ex-type: CBS 260.55 = DTO 022-G5 = ATCC 10499 = CECT 2268 = DSM1252 = IMI 040583 = NRRL 2140 = QM 6413. Infragen. class: subgen. Aspergilloides, sect. Citrina, ser. Citrina. Reproduction: asexual. ITS barcode: GU944597 (alternative markers: BenA = GU944522; CaM = MN969300; RPB2 = JN606602).

Penicillium sterculiniicola Houbraken, Stud. Mycol. 78: 436. 2014 [MB809973]. — Type: CBS H-21877. Ex-type: CBS 122426 = DTO 031-A4. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Spinulosa. Reproduction: asexual. ITS barcode: KM189464 (alternative markers: BenA = KM088693; CaM = KM089078; RPB2 = KM089465).

Penicillium stolkiae D.B. Scott, Mycopathol. Mycol. Appl. 36: 8. 1968. [MB335768]. — Type: CBS 315.67. Ex-type: CBS 315.67 = ATCC 18546 = CSIR 1041 = FRR 534 = IMI 136210 = NRRL 5816. Infragen. class: subgen. Aspergilloides, sect. Stolkia, ser. Stolkia. Reproduction: homothallic. ITS barcode: AF033444 (alternative markers: BenA = JN617717; CaM = AF481135; RPB2 = JN121488).

Penicillium striatisporum Stolk, Antonie van Leeuwenhoek 35: 268. 1969. [MB335769]. — Type: CBS 705.68. Ex-type: CBS 705.68 = ATCC 22052 = CCRC 31679 = FRR 827 = IMI 151749 = MUCL 31202. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Erubescentia. Reproduction: asexual. ITS barcode: AF038938 (alternative markers: BenA = MN969401; CaM = KP016807; RPB2 = JN406538).

Penicillium subarcticum S.W. Peterson & Sigler, Mycol. Res. 106: 1116. 2002. [MB483983]. — Type: BPI 841397. Ex-type: CBS 111719 = NRRL 31108 = UAMH 3897. Infragen. class: subgen. Aspergilloides, sect. Stolkia, ser. Stolkia. Reproduction: asexual. ITS barcode: AF481120 (alternative markers: BenA = JN617716; CaM = AF481141; RPB2 = MN969139).

Penicillium sublectaticum Houbraken et al., Stud. Mycol. 78: 436. 2014. [MB809974]. — Type: CBS H-21955. Ex-type: CBS 138217= DTO 244-G2. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Sublectatica. Reproduction: asexual. ITS barcode: KM189761 (alternative markers: BenA = KM089010; CaM = KM089397; RPB2 = KM089784).

Penicillium subrubescens Houbraken et al., Antonie van Leeuwenhoek 103: 1354. 2013. [MB801306]. — Type: CBS H-21029. Ex-type: CBS 132785 = DTO 188-D6 = FBCC 1632 = IBT 31985. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Rolfsiorum. Reproduction: asexual. ITS barcode: KC346350 (alternative markers: BenA = KC346327; CaM = KC346330; RPB2 = KC346306).

Penicillium subspinulosum Houbraken, Stud. Mycol. 78: 436. 2014. [MB809975]. — Type: CBS H-21856. Ex-type: CBS 137946 = DTO 041-F2. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Spinulosa. Reproduction: asexual. ITS barcode: KM189483 (alternative markers: BenA = KM088719; CaM = KM089104; RPB2 = KM089491).

Penicillium subturcoseum Visagie & K. Jacobs, IMA Fungus 7: 105. 2016. [MB811008]. — Type: CBS H-22041. Ex-type: CBS 139132 = DAOMC 241096 = DTO 180-C9 = CV 2835. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Corylophila. Reproduction: asexual. ITS barcode: FJ231006 (alternative markers: BenA = JX141161; CaM = JX157532; RPB2 = KP064674).

Penicillium sucrivorum Visagie & K. Jacobs, Mycologia 106: 546. 2014. [MB804723]. — Type: CBS H-21331. Ex-type: CBS 135116 = DAOM 241042 = DTO 183-E5. Infragen. class: subgen. Aspergilloides, sect. Citrina, ser. Westlingiorum. Reproduction: asexual. ITS barcode: JX140872 (alternative markers: BenA = JX141015; CaM = JX141506; RPB2 = MN969140).

Penicillium sumatraense [as “sumatrense”] Szilvinyi, Archiv. Hydrobiol.14 Suppl. 6: 535. 1936. [MB319297]. — Type: CBS 281.36. Ex-type: CBS 281.36 = DTO 022-F1 = NRRL 779 = FRR 779. Infragen. class: subgen. Aspergilloides, sect. Citrina, ser. Sumatraensia. Reproduction: asexual. ITS barcode: GU944578 (alternative markers: BenA = JN606639; CaM = MN969301; RPB2 = EF198541).

Penicillium svalbardense Frisvad et al., Antonie van Leeuwenhoek 92: 48. 2007. [MB529943]. — Type: EX-F 1307. Ex-type: CBS 122416 = IBT 23856 = DTO 048-D5 = EXF-1307. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Rolfsiorum. Reproduction: asexual. ITS barcode: GU981603 (alternative markers: BenA = DQ486644; CaM = KC346338; RPB2 = KF296457).

Penicillium swiecickii K.W. Zaleski, Bull. Int. Acad. Polon. Sci., Sér. B., Sci. Nat. 1927: 474. 1927. [MB534781]. — Type: unknown. Ex-type: CBS 119391 = FRR 918 = IBT 27865 = IMI 191500 = NRRL 918. Infragen. class: subgen. Penicillium, sect. Ramosum, ser. Lanosa. Reproduction: asexual. ITS barcode: AF033490 (alternative markers: BenA = KJ834494; CaM = KJ866993; RPB2 = JN406635).

Penicillium synnematicola Guevara-Suarez et al., Fungal Syst. Evol. 5: 62. 2020. [MB822072]. — Type: CBS H-23132. Ex-type: CBS 142669 = FMR 15192. Infragen. class: subgen. Penicillium, sect. Robsamsonia, ser. Glandicolarum. Reproduction: asexual. ITS barcode: LT898167 (alternative markers: BenA = LT898172; CaM = LT898137; RPB2 = LT898142).

Penicillium taiwanense (Matsushima) Houbraken & Samson, Stud. Mycol. 70: 48. 2011. [MB561969]. Basionym: Phialomyces taiwanensis Matsush., Matsushima Mycological Memoirs 4: 12. 1985. [MB105680]. — Type: unknown. Ex-type: n.a. Infragen. class: subgen. Aspergilloides, sect. Thysanophora, ser. Thysanophora. Reproduction: asexual. ITS barcode: n.a. (alternative markers: BenA = n.a.; CaM = n.a.; RPB2 = n.a.).

Penicillium tanzanicum Visagie et al., Persoonia 36: 278. 2016. [MB815781]. — Type: DAOM 695766. Ex-type: DAOMC 250514 = CBS 140968 = DTO 410-D3 = 50.118 = KAS 1946. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Simplicissima. Reproduction: asexual. ITS barcode: KT887841 (alternative markers: BenA = KT887802; CaM = KT887763; RPB2 = MN969183).

Penicillium tardochrysogenum Frisvad et al., Persoonia 29: 93. 2012. [MB801877]. — Type: CBS H-21057. Ex-type: DTO 149-B9 = CBS 132200 = IBT 30075. Infragen. class: subgen. Penicillium, sect. Chrysogena, ser. Chrysogena. Reproduction: asexual. ITS barcode: JX997028 (alternative markers: BenA = JX996898; CaM = JX996239; RPB2 = JX996634).

Penicillium taxi R. Schneid., Zentralbl. Bakteriol. Parasitenk., Abt. 2 110: 43. 1956. [MB282799]. — Type: unknown. Ex-type: CBS 206.57 = ATCC 18484 = BBA 7480 = MUCL 11402 = QM 8153. Infragen. class: subgen. Aspergilloides, sect. Thysanophora, ser. Thysanophora. Reproduction: asexual. ITS barcode: KJ834517 (alternative markers: BenA = KJ834495; CaM = MN969302; RPB2 = JN121454).

Penicillium terrarumae Houbraken et al., Phytotaxa 273: 170. 2016. [MB801431]. — Type: HGUPd2020. Ex-type: HGUP2025 = CBS 131811 = DTO 174-H2. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Rolfsiorum. Reproduction: asexual. ITS barcode: MN431397 (alternative markers: BenA = KX650295; CaM = MN969323; RPB2 = MN969185).

Penicillium terrenum D.B. Scott, Mycopathol. Mycol. Appl. 36: 1. 1968. [MB335771]. — Type: CBS 313.67. Ex-type: CBS 313.67 = ATCC 18547 = CSIR 1022 = IMI 136208. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Lapidosa. Reproduction: homothallic. ITS barcode: AM992111 (alternative markers: BenA = KJ834496; CaM = KP016808; RPB2 = JN406577).

Penicillium terrigenum Seifert et al., Stud. Mycol. 70: 125. 2011. [MB563204]. — Type: CBS H-20667. Ex-type: CBS 127354 = IBT 30769. Infragen. class: subgen. Aspergilloides, sect. Citrina, ser. Copticolarum. Reproduction: asexual. ITS barcode: JN617684 (alternative markers: BenA = JN606810; CaM = JN606583; RPB2 = JN606600).

Penicillium thiersii S.W. Peterson et al., Mycologia 96: 1283. 2004. [MB487738]. — Type: BPI 842269. Ex-type: CBS 117503 = IBT 27050 = NRRL 28162. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Thiersiorum. Reproduction: homothallic (Houbraken et al. 2014b). ITS barcode: AF125936 (alternative markers: BenA = KJ834497; CaM = AY741726; RPB2 = JN121434).

Penicillium thomii Maire, Bull. Soc. Hist. Nat. Afrique N. 8: 189. 1917. [MB202819]. — Type: IMI 189694. Ex-type: CBS 225.81 = IMI 189694 = NRRL 2077. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Thomiorum. Reproduction: asexual. ITS barcode: KM189560 (alternative markers: BenA = KM088799; CaM = KM089184; RPB2 = KM089571).

Penicillium thymicola Frisvad & Samson, Stud. Mycol. 49: 29. 2004. [MB370969]. — Type: CBS 111225. Ex-type: CBS 111225 = IBT 5891. Infragen. class: subgen. Penicillium, sect. Fasciculata, ser. Verrucosa. Reproduction: asexual. ITS barcode: KJ834518 (alternative markers: BenA = MN969402; CaM = FJ530990; RPB2 = KU904364).

Penicillium tricolor Frisvad et al., Canad. J. Bot. 72: 937. 1994. [MB541710]. — Type: DAOM 216240. Ex-type: DTO 157-A4 = CBS 635.93 = IBT 12493 = DAOM 216240. Infragen. class: subgen. Penicillium, sect. Fasciculata, ser. Viridicata. Reproduction: asexual. ITS barcode: JN942704 (alternative markers: BenA = MN969403; CaM = KU896852; RPB2 = JN985422).

Penicillium tropicoides Houbraken et al., Fungal Divers. 44: 127. 2010. [MB518293]. — Type: CBS 122410. Ex-type: CBS 122410 = DTO 010-C4 = IBT 29043. Infragen. class: subgen. Aspergilloides, sect. Citrina, ser. Citrina. Reproduction: homothallic. ITS barcode: GU944584 (alternative markers: BenA = GU944531; CaM = MN969303; RPB2 = JN606608).

Penicillium tropicum Houbraken et al., Fungal Divers. 44: 129. 2010. [MB518294]. — Type: SC42-1. Ex-type: CBS 112584 = DTO 031-B1 = IBT 24580. Infragen. class: subgen. Aspergilloides, sect. Citrina, ser. Citrina. Reproduction: homothallic. ITS barcode: GU944582 (alternative markers: BenA = GU944532; CaM = MN969304; RPB2 = JN606607).

Penicillium trzebinskii K.W. Zaleski, Bull. Int. Acad. Polon. Sci., Sér. B., Sci. Nat. 1927: 498. 1927. [MB280795]. — Type: unknown. Ex-type: CBS 382.48 = ATCC 10507 = FRR 731 = IFO 6110 = IMI 039749 = MUCL 29102 = NRRL 731 = QM 7678. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Spinulosa. Reproduction: asexual. ITS barcode: KM189784 (alternative markers: BenA = KM089034; CaM = KM089421; RPB2 = KM089808).

Penicillium tsitsikammaense Houbraken, Stud. Mycol. 78: 440. 2014. [MB809976]. — Type: CBS H-21881. Ex-type: CBS 328.71 = DTO 006-I3 = CSIR 1092. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Pinetorum. Reproduction: homothallic (Stolk & Samson 1983:127, CBS 328.71). ITS barcode: KM189451 (alternative markers: BenA = KM088675; CaM = KM089060; RPB2 = KM089447).

Penicillium tubakianum Visagie & Samson, Persoonia 36: 151. 2016. [MB808270]. — Type: CBS H-21604. Ex-type: CBS 287.66 = DTO 138-D9 = MUCL 8519 = IFO 8315. Infragen. class: subgen. Aspergilloides, sect. Torulomyces, ser. Torulomyces. Reproduction: asexual. ITS barcode: KF303652 (alternative markers: BenA = KF303611; CaM = KF303637; RPB2 = KF303712).

Penicillium tularense Paden, Mycopathol. Mycol. Appl. 43: 264. 1971. [MB319298]. — Type: UVIC JWP 68-31. Ex-type: CBS 430.69 = ATCC 22056 = FRR 899 = IFO 31740 = IMI 148394 = NRRL 5273 = AS 3.14006. Infragen. class: subgen. Penicillium, sect. Brevicompacta, ser. Tularensia. Reproduction: homothallic. ITS barcode: AF033487 (alternative markers: BenA = KC427175; CaM = JX313135; RPB2 = JN121516).

Penicillium tulipae Overy & Frisvad, Syst. Appl. Microbiol. 26: 634. 2003. [MB488954]. — Type: C 60162. Ex-type: CBS 109555 = CBS 187.88 = IBT 3458. Infragen. class: subgen. Penicillium, sect. Fasciculata, ser. Corymbifera. Reproduction: asexual. ITS barcode: KJ834519 (alternative markers: BenA = MN969404; CaM = MN969305; RPB2 = MN969141).

Penicillium tunisiense S. Ouhibi et al., Int. J. Syst. Evol. Microbiol. 68: 3224. 2018. [MB823626]. — Type: MUM-H 17.62. Ex-type: MUM 17.62 = ITEM 17445. Infragen. class: subgen. Penicillium, sect. Ramosum, ser. Soppiorum. Reproduction: asexual. ITS barcode: MG586956 (alternative markers: BenA = MG586970; CaM = MG586974; RPB2 = n.a.).

Penicillium turbatum Westling, Ark. Bot. 11: 128. 1911. [MB202895]. — Type: IMI 39738. Ex-type: CBS 383.48 = CBS 237.60 = ATCC 9782 = DSM2426 = FRR 757 = IFO 7767 = IMI 039738 = MUCL 29115 = NRRL 757 = NRRL 758 = QM 1941. Infragen. class: subgen. Penicillium, sect. Turbata, ser. Turbata. Reproduction: homothallic (Scott and Stolk 1967; as E. baarnense). ITS barcode: AF034454 (alternative markers: BenA = KJ834499; CaM = KU896853; RPB2 = JN406556).

Penicillium turcosoconidiatum Visagie et al., Stud. Mycol. 78: 440. 2014. [MB809977]. — Type: CBS H-21876. Ex-type: CBS 138557 = DTO 181-A3 = CV 110 = DAOM 241130. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Pinetorum. Reproduction: asexual. ITS barcode: KM189645 (alternative markers: BenA = KM088889; CaM = KM089276; RPB2 = KM089663).

Penicillium ubiquetum Houbraken et al., Stud. Mycol. 70: 127. 2011. [MB563201]. — Type: CBS H-20659. Ex-type: CBS 126437 = DTO 078-B5 = IBT 22226. Infragen. class: subgen. Aspergilloides, sect. Citrina, ser. Westlingiorum. Reproduction: asexual. ITS barcode: JN617680 (alternative markers: BenA = JN606800; CaM = MN969306; RPB2 = MN969142).

Penicillium ulaiense H.M. Hsieh et al., Trans. Mycol. Soc. Rep. China 2: 161. 1987. [MB126489]. — Type: PPEH 29001.87. Ex-type: CBS 210.92 = CBS 261.94 = CCRC 32655 = IBT 18387 = IBT 23037. Infragen. class: subgen. Penicillium, sect. Penicillium, ser. Italica. Reproduction: asexual. ITS barcode: KC411695 (alternative markers: BenA = AY674408; CaM = KU896854; RPB2 = KU904365).

Penicillium uruguayense Guevara-Suarez et al., Persoonia 39: 323. 2017. [MB822920]. — Type: FMR H-14490. Ex-type: CBS 143247 = FMR 14490 = DTO 410-E9. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Janthinella. Reproduction: homothallic. ITS barcode: LT904729 (alternative markers: BenA = LT904699; CaM = LT904698; RPB2 = MN969200).

Penicillium vagum Houbraken et al., Stud. Mycol. 78: 443. 2014. [MB809978]. — Type: CBS H-21926. Ex-type: CBS 137728 = DTO 180-G3 = CV 25 = DAOM 241357. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Longicatenata. Reproduction: asexual. ITS barcode: KM189642 (alternative markers: BenA = KM088886; CaM = KM089273; RPB2 = KM089660).

Penicillium valentinum C. Ramírez & A.T. Martínez, Mycopathologia 72: 183. 1980. [MB113027]. — Type: IJFM 5071. Ex-type: CBS 172.81 = ATCC 42227 = IJFM 5071. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Thomiorum. Reproduction: asexual. ITS barcode: KM189550 (alternative markers: BenA = KM088788; CaM = KM089173; RPB2 = KM089560).

Penicillium vancouverense Houbraken et al., Stud. Mycol. 70: 131. 2011. [MB563207]. — Type: CBS H-20646. Ex-type: CBS 126323 = DTO 082-B8 = IBT 20700. Infragen. class: subgen. Aspergilloides, sect. Citrina, ser. Westlingiorum. Reproduction: asexual. ITS barcode: JN617675 (alternative markers: BenA = JN606663; CaM = MN969307; RPB2 = MN969143).

Penicillium vanderhammenii Houbraken et al., Int. J. Syst. Evol. Microbiol. 61: 1473. 2011. [MB518027]. — Type: HUA 170337. Ex-type: CBS 126216 = DTO 097-A3 = DTO 297-I2 = IBT 23203. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Dalearum. Reproduction: homothallic. ITS barcode: GU981574 (alternative markers: BenA = GU981647; CaM = MN969308; RPB2 = KF296458).

Penicillium vanluykii Frisvad et al., Persoonia 29: 97. 2012. [MB801878]. — Type: CBS H-21059. Ex-type: DTO 148-I2 = CBS 131539 = IBT 14505. Infragen. class: subgen. Penicillium, sect. Chrysogena, ser. Chrysogena. Reproduction: asexual. ITS barcode: JX997007 (alternative markers: BenA = JX996879; CaM = JX996220; RPB2 = JX996615).

Penicillium vanoranjei Visagie et al., Persoonia 31: 46. 2013. [MB803782]. — Type: CBS H-21145. Ex-type: CBS 134406. Infragen. class: subgen. Aspergilloides, sect. Sclerotiorum, ser. Sclerotiorum. Reproduction: asexual. ITS barcode: KC695696 (alternative markers: BenA = KC695686; CaM = KC695691; RPB2 = n.a.).

Penicillium variratense Visagie & Samson, Persoonia 36: 151. 2016. [MB808271]. — Type: CBS H-21611. Ex-type: CBS 337.97 = DTO 137-C8. Infragen. class: subgen. Aspergilloides, sect. Torulomyces, ser. Torulomyces. Reproduction: asexual. ITS barcode: KF303649 (alternative markers: BenA = KF303610; CaM = KF303630; RPB2 = KF303675).

Penicillium vasconiae C. Ramírez & A.T. Martínez, Mycopathologia 72: 189. 1980. [MB113028]. — Type: CBS 339.79. Ex-type: CBS 339.79 = DTO 076-H1 = ATCC 42224 = IJFM 3008. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Rolfsiorum. Reproduction: asexual. ITS barcode: GU981599 (alternative markers: BenA = GU981653; CaM = MN969309; RPB2 = MN969144).

Penicillium vascosobrinhous R.N. Barbosa & J.D.P. Bezerra, Acta Bot. Bras. 2020. [MB833816]. — Type: URM 94140. Ex-type: URM 8193. Infragen. class: subgen. Aspergilloides, sect. Citrina, ser. Euglauca. Reproduction: asexual. ITS barcode: LR744067 (alternative markers: BenA = LR744069; CaM = LR744063; RPB2 = LR744065).

Penicillium velutinum J.F.H. Beyma, Zentralbl. Bakteriol. Parasitenk., Abt. 2 91: 353. 1935. [MB283175]. — Type: IMI 40571. Ex-type: CBS 250.32 = ATCC 10510 = CECT 2318 = IJFM 5108 = IMI 040571 = NRRL 2069 = QM 7686 = VKMF-379. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Lapidosa. Reproduction: asexual. ITS barcode: AF033448 (alternative markers: BenA = JX141170; CaM = MT478037; RPB2 = KP064682).

Penicillium venetum (Frisvad) Frisvad, Int. Mod. Meth. Pen. Asp. Clas.: 275. 2000. [MB459816]. Basionym: Penicillium hirsutum var. venetum Frisvad, Mycologia 81: 856. 1990. [MB126414]. — Type: IMI 321520. Ex-type: IBT 10661 = IMI 321520. Infragen. class: subgen. Penicillium, sect. Fasciculata, ser. Corymbifera. Reproduction: asexual. ITS barcode: AJ005485 (alternative markers: BenA = AY674335; CaM = KU896855; RPB2 = KU904366).

Penicillium verhagenii Houbraken, Stud. Mycol. 78: 443. 2014. [MB809979]. — Type: CBS H-21865. Ex-type: CBS 137959 = DTO 193-A1. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Verhageniorum. Reproduction: asexual. ITS barcode: KM189708 (alternative markers: BenA = KM088955; CaM = KM089342; RPB2 = KM089729).

Penicillium verrucisporum X.C. Wang & W.Y. Zhuang, Sci. Rep. 7: 8233, 10. 2017. [MB570339]. — Type: HMAS 248819. Ex-type: CGMCC 3.18415. Infragen. class: subgen. Aspergilloides, sect. Sclerotiorum, ser. Herqueorum. Reproduction: asexual. ITS barcode: KX885069 (alternative markers: BenA = KX885049; CaM = KX885059; RPB2 = KX885040).

Penicillium verrucosum Dierckx, Ann. Soc. Sci. Bruxelles 25: 88. 1901. [MB212252]. — Type: IMI 200310. Ex-type: CBS 603.74 = ATCC 48957 = ATHUM 2897 = CECT 2906 = FRR 965 = IBT 12809 = IBT 4733 = IMI 200310 = IMI 200310ii = MUCL 28674 = MUCL 29089 = MUCL 29186 = NRRL 965. Infragen. class: subgen. Penicillium, sect. Fasciculata, ser. Verrucosa. Reproduction: asexual. ITS barcode: AY373938 (alternative markers: BenA = MN969405; CaM = DQ911138; RPB2 = JN121539).

Penicillium vinaceum J.C. Gilman & E.V. Abbott, Iowa St. Coll. J. Sci. 1: 299. 1927. [MB281754]. — Type: IMI 29189. Ex-type: CBS 389.48 = ATCC 10514 = FRR 739 = IMI 029189 = NRRL 739 = QM 6746. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Erubescentia. Reproduction: asexual. ITS barcode: AF033461 (alternative markers: BenA = HQ646575; CaM = HQ646586; RPB2 = JN406555).

Penicillium virgatum Nirenberg & Kwasna, Mycol. Res. 109: 977. 2005. [MB341488]. — Type: BBA 65745. Ex-type: CBS 114838 = BBA 65745. Infragen. class: subgen. Penicillium, sect. Ramosum, ser. Virgata. Reproduction: asexual. ITS barcode: AJ748692 (alternative markers: BenA = KJ834500; CaM = KJ866992; RPB2 = JN406641).

Penicillium viridicatum Westling, Ark. Bot. 11: 88. 1911. [MB163349]. — Type: IMI 39758ii. Ex-type: DTO 005-C9 = CBS 390.48 = ATCC 10515 = IBT 23041 = IFO 7736 = IMI 039758 = IMI 039758ii = NRRL 963 = QM 7683. Infragen. class: subgen. Penicillium, sect. Fasciculata, ser. Viridicata. Reproduction: asexual. ITS barcode: AY373939 (alternative markers: BenA = MN969406; CaM = KU896856; RPB2 = JN121511).

Penicillium viridissimum L. Cai & X.Z. Jiang, Cladistics 35: 543. 2018 [2019]. [MB818160]. — Type: HMAS 247735. Ex-type: CGMCC 3.18796 = NN072081 = CBS 144484. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Dalearum. Reproduction: asexual. ITS barcode: KY495004 (alternative markers: BenA = KY495113; CaM = MN969339; RPB2 = KY495059).

Penicillium viticola Nonaka & Masuma, Mycoscience 52: 339. 2011. [MB516048]. — Type: TNS-F38702. Ex-type: JCM 17636 = FKI-4410. Infragen. class: subgen. Aspergilloides, sect. Sclerotiorum, ser. Sclerotiorum. Reproduction: asexual. ITS barcode: AB606414 (alternative markers: BenA = AB540174; CaM = n.a.; RPB2 = n.a.).

Penicillium vulpinum (Cooke & Massee) Seifert & Samson, Adv. Pen. Asp. Syst.: 144. 1986 [1985]. [MB114763]. Basionym: Coremium vulpinum Cooke & Massee, Grevillea 16: 81. 1888. [MB183683]. — Type: "on dung", s. coll., in herb. Cooke (K). Ex-type: CBS 126.23 = ATCC 10426 = IMI 040237 = NRRL 2031 = VKMF-257. Infragen. class: subgen. Penicillium, sect. Robsamsonia, ser. Claviformia. Reproduction: asexual. ITS barcode: AF506012 (alternative markers: BenA = KJ834501; CaM = KU896857; RPB2 = KU904367).

Penicillium waksmanii K.W. Zaleski, Bull. Int. Acad. Polon. Sci., Sér. B., Sci. Nat.: 468. 1927. [MB121677]. — Type: IMI 39746i. Ex-type: CBS 230.28 = DTO 022-E6 = ATCC 10516 = FRR 777 = IFO 7737 = IMI 039746 = IMI 039746i = MUCL 29120 = NRRL 777 = QM 7681. Infragen. class: subgen. Aspergilloides, sect. Citrina, ser. Westlingiorum. Reproduction: asexual. ITS barcode: GU944602 (alternative markers: BenA = JN606779; CaM = MN969310; RPB2 = JN606627).

Penicillium wellingtonense A.L.J. Cole et al., Stud. Mycol. 70: 133. 2011. [MB563208]. — Type: CBS H-20657. Ex-type: CBS 130375 = DTO 076-C6 = IBT 23557 = DTO 76C6. Infragen. class: subgen. Aspergilloides, sect. Citrina, ser. Westlingiorum. Reproduction: asexual. ITS barcode: JN617713 (alternative markers: BenA = JN606670; CaM = MN969311; RPB2 = JN606616).

Penicillium westlingii K.W. Zaleski, Bull. Int. Acad. Polon. Sci., Sér. B., Sci. Nat. 1927: 473. 1927. [MB282076]. — Type: IMI 92272. Ex-type: CBS 231.28 = DTO 022-E7 = IMI 092272. Infragen. class: subgen. Aspergilloides, sect. Citrina, ser. Westlingiorum. Reproduction: asexual. ITS barcode: GU944601 (alternative markers: BenA = JN606718; CaM = MN969312; RPB2 = JN606625).

Penicillium williamettense Visagie & Samson, Persoonia 36: 151. 2016. [MB808272]. — Type: CBS H-21609. Ex-type: CBS 129774 = DTO 208-A4. Infragen. class: subgen. Aspergilloides, sect. Torulomyces, ser. Torulomyces. Reproduction: asexual. ITS barcode: KF303667 (alternative markers: BenA = KF303622; CaM = KF303639; RPB2 = KF303709).

Penicillium wisconsinense Visagie & Samson, Persoonia 36: 151. 2016. [MB808273]. — Type: CBS H-21614. Ex-type: CBS 128279 = DTO 198-H7 = WSF 3132. Infragen. class: subgen. Aspergilloides, sect. Torulomyces, ser. Torulomyces. Reproduction: asexual. ITS barcode: KF303670 (alternative markers: BenA = KF303624; CaM = KF303641; RPB2 = KF303706).

Penicillium wollemiicola Visagie et al., Persoonia 36: 153. 2016. [MB808274]. — Type: DAOM 675862. Ex-type: CBS 137177 = DTO 297-E3. Infragen. class: subgen. Aspergilloides, sect. Torulomyces, ser. Torulomyces. Reproduction: homothallic. ITS barcode: KJ174314 (alternative markers: BenA = KJ174315; CaM = KJ174316; RPB2 = KJ174313).

Penicillium wotroi Houbraken et al., Int. J. Syst. Evol. Microbiol. 61: 1474. 2011. [MB518026]. — Type: HUA 170336. Ex-type: CBS 118171 = DTO 056-E5 = DTO 297-I3 = IBT 23253. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Simplicissima. Reproduction: asexual. ITS barcode: GU981591 (alternative markers: BenA = GU981637; CaM = MN969313; RPB2 = KF296460).

Penicillium xanthomelinii Visagie & K. Jacobs, IMA Fungus 7: 105. 2016. [MB811009]. — Type: CBS H-22048. Ex-type: CBS 139163 = DAOMC 241104 = DTO 183-C7 = CV 1677. Infragen. class: subgen. Aspergilloides, sect. Exilicaulis, ser. Lapidosa. Reproduction: asexual. ITS barcode: JX140921 (alternative markers: BenA = JX141120; CaM = JX157495; RPB2 = KP064683).

Penicillium yarmokense Baghd., Novosti Sist. Nizsh. Rast. 5: 99. 1968. [MB335774]. — Type: CBS H-7536. Ex-type: CBS 410.69 = FRR 520 = IMI 140346 = VKMF-1076. Infragen. class: subgen. Penicillium, sect. Canescentia, ser. Canescentia. Reproduction: asexual. ITS barcode: KC411757 (alternative markers: BenA = MN969407; CaM = MN969314; RPB2 = JN406553).

Penicillium yezoense Hanzawa ex Houbraken, Stud. Mycol. 78: 443. 2014. [MB809980]. — Type: CBS H-21863. Ex-type: CBS 350.59 = ATCC 18333 = FRR 3395 = IFO 5362 = IMI 068615. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Thomiorum. Reproduction: asexual. ITS barcode: KM189553 (alternative markers: BenA = KM088792; CaM = KM089177; RPB2 = KM089564).

Penicillium yunnanense L. Cai & X.Z. Jiang, Cladistics 35: 545. 2018 [2019]. [MB818163]. — Type: HMAS 247736. Ex-type: CGMCC 3.18794 = NN051336 = CBS 144485. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Janthinella. Reproduction: asexual. ITS barcode: KY494990 (alternative markers: BenA = KY495099; CaM = MN969340; RPB2 = KY495048).

Penicillium zhuangii L. Wang, PLoS ONE 9: e101454, 4. 2014. [MB805945]. — Type: HMAS 244922. Ex-type: CBS 137464 = NRRL 62806 = AS 3.15341. Infragen. class: subgen. Aspergilloides, sect. Aspergilloides, ser. Hoeksiorum. Reproduction: asexual. ITS barcode: KF769435 (alternative markers: BenA = KF769411; CaM = KF769422; RPB2 = MN969145).

Penicillium zonatum Hodges & J.J. Perry, Mycologia 65: 697. 1973. [MB319303]. — Type: BPI FSL 525. Ex-type: CBS 992.72 = DTO 096-I3 = ATCC 24353. Infragen. class: subgen. Aspergilloides, sect. Lanata-Divaricata, ser. Dalearum. Reproduction: homothallic. ITS barcode: GU981581 (alternative markers: BenA = GU981651; CaM = MN969315; RPB2 = KF296461).



Phialomyces

Phialomyces arenicola (Chalab.) Houbraken et al., this study. [MB832563]. Basionym: Penicillium arenicola Chalab., Not. Syst. Crypt. Inst. bot. Acad. Sci. USSR: 161-167. 1950. [MB302375]. — Type: IMI 117658. Ex-type: CBS 220.66 = ATCC 18321 = ATCC 18330 = DSM 2435 = FRR 3392 = IMI 117658 = NRRL 3392 = VKM F-1035. Reproduction: asexual. ITS barcode: GU092964 (alternative markers: BenA = GU092771; CaM = GU092801; RPB2 = GU092935).

Phialomyces fusiformis G. Delgado & Decock, Mycologia 95: 896. 2003. [MB489106]. — Type: MUCL 43747. Ex-type: MUCL 43747. Reproduction: asexual. ITS barcode: n.a. (alternative markers: BenA = n.a.; CaM = n.a.; RPB2 = n.a.).

Phialomyces humicoloides (Bills & Heredia) Houbraken et al., this study. [MB832564]. Basionym: Merimbla humicoloides Bills & Heredia, Mycol. Res. 105: 1276. 2001. [MB474487]. — Type: BPI 748244. Ex-type: CBS 102854 = NRRL 35712. Reproduction: asexual. ITS barcode: GU092965 (alternative markers: BenA = GU092782; CaM = GU092804; RPB2 = GU092937).

Phialomyces macrosporus P.C. Misra & P.H.B. Talbot, Canad. J. Bot. 42: 1287. 1964. [MB336291]. — Type: Waite Instit. 15645. Ex-type: CBS 430.64 = ATCC 16661= IMI 110130 = MUCL 9776. Reproduction: asexual. ITS barcode: MN431404 (alternative markers: BenA = MN969422; CaM = MN969343; RPB2 = JN121515).



Pseudohamigera

Pseudohamigera striata (Raper & Fennell) Houbraken et al., this study. [MB832565]. Basionym: Penicillium striatum Raper & Fennell, Mycologia 40: 521. 1948. [MB289109]. — Type: IMI 39741. Ex-type: CBS 377.48 = ATCC 10501 = IFO 6106 = IMI 039741 = NRRL 717 = QM 1857 = VKM F-2044. Reproduction: homothallic. ITS barcode: AF454073 (alternative markers: BenA = GU092799; CaM = GU092841; RPB2 = GU092928).



Pseudopenicillium

Pseudopenicillium cervifimosum Guevara-Suarez et al., Fungal Syst. Evol. 5: 66. 2020. [MB822079]. — Type: CBS H-23133. Ex-type: CBS 142670 = FMR 15299. Reproduction: asexual. ITS barcode: LT899789 (alternative markers: BenA = LT898315; CaM = n.a.; RPB2 = LT899807).

Pseudopenicillium giganteum (R.Y. Roy & G.N. Singh) Guevara-Suarez et al., Fungal Syst. Evol. 5: 68. 2020. [MB822077]. Basionym: Penicillium giganteum R.Y. Roy & G.N. Singh, Trans. Brit. Mycol. Soc. 51: 805. 1968. [MB335730]. — Type: IMI 132774. Ex-type: DTO 036-H2 = CBS 144.69 = ATCC 48996 = FRR 535 = IMI 132774 = NRRL 3553. Reproduction: asexual. ITS barcode: GU092966 (alternative markers: BenA = GU092779; CaM = GU092847; RPB2 = GU092923).

Pseudopenicillium megasporum (Orpurt & Fennell) Guevara-Suarez et al., Fungal Syst. Evol. 5: 68. 2020. [MB822078]. Basionym: Penicillium megasporum Orpurt & Fennell, Mycologia 47: 233. 1955. [MB302408]. — Type: IMI 216904. Ex-type: CBS 256.55 = ATCC 12322 = FRR 2232 = IMI 216904 = NRRL 2232 = QM 6879 = WB 2232. Reproduction: asexual. ITS barcode: AF033494 (alternative markers: BenA = GU092757; CaM = GU092846; RPB2 = GU092921).



Rasamsonia

Rasamsonia aegroticola Houbraken et al., J. Clin. Microbiol. 51: 25. 2013. [MB801150]. — Type: CBS H-21031. Ex-type: DTO 137-A8 = CBS 132819 = IHEM 22641. Reproduction: asexual. ITS barcode: JX272988 (alternative markers: BenA = JX273020; CaM = JX272956; RPB2 = MN969193).

Rasamsonia argillacea (Stolk et al.) Houbraken & Frisvad, Antonie van Leeuwenhoek 101: 412. 2012. [MB519878]. Basionym: Penicillium argillaceum Stolk, H.C. Evans & T. Nilsson, Trans. Brit. Mycol. Soc. 53: 307. 1969. [MB335712]. — Type: CBS 101.69. Ex-type: DTO 097-E4 = CBS 101.69 = IBT 31199. Reproduction: asexual. ITS barcode: JF417491 (alternative markers: BenA = JF417456; CaM = JF417501; RPB2 = JF417415).

Rasamsonia brevistipitata Houbraken & Frisvad, Antonie van Leeuwenhoek 101: 413. 2012. [MB519870]. — Type: CBS H-20546. Ex-type: DTO 025-H2 = CBS 128785 = IBT 31187. Reproduction: asexual. ITS barcode: JF417488 (alternative markers: BenA = JF417454; CaM = JF417499; RPB2 = JN406530).

Rasamsonia byssochlamydoides (Stolk & Samson) Houbraken & Frisvad, Antonie van Leeuwenhoek 101: 415. 2012. [MB519877]. Basionym: Talaromyces byssochlamydoides Stolk & Samson, Stud. Mycol. 2: 45. 1972. [MB324415]. — Type: CBS 413.71. Ex-type: DTO 149-D6 = DTO 108-B4 = CBS 413.71 = IMI 178524 = JCM 12813 = NRRL 3658. Reproduction: homothallic. ITS barcode: JF417476 (alternative markers: BenA = JF417460; CaM = JF417512; RPB2 = JF417437).

Rasamsonia columbiensis Jurjević et al., Persoonia 36: 405. 2016. [MB816869]. — Type: BPI 910043. Ex-type: CBS 141097 = CCF 5289. Reproduction: asexual. ITS barcode: LT548281 (alternative markers: BenA = LT548285; CaM = MN969326; RPB2 = MN969195).

Rasamsonia composticola Y.Y. Su & L. Cai, Mycol. Prog. 12: 217. 2013. [MB800249]. — Type: HMAS 242447. Ex-type: CGMCC 3.13669. Reproduction: homothallic. ITS barcode: JF970184 (alternative markers: BenA = JF970183; CaM = JQ729688; RPB2 = JQ729684).

Rasamsonia cylindrospora (G. Sm.) Houbraken & Frisvad, Antonie van Leeuwenhoek 101: 415. 2012. [MB519876]. Basionym: Penicillium cylindrosporum G. Sm., Trans. Brit. Mycol. Soc. 40: 483. 1957. [MB302392]. — Type: IMI 71623. Ex-type: DTO 138-F8 = CBS 275.58 = IBT 31202 = IMI 071623. Reproduction: asexual. ITS barcode: JF417470 (alternative markers: BenA = JF417448; CaM = JF417493; RPB2 = JF417423).

Rasamsonia eburnea (Yaguchi et al.) Houbraken & Frisvad, Antonie van Leeuwenhoek 101: 416. 2012. [MB519875]. Basionym: Talaromyces eburneus Yaguchi et al., Mycoscience 35: 249. 1994. [MB362928]. — Type: PF 1151. Ex-type: DTO 105-D6 = CBS 100538 = IBT 17519. Reproduction: homothallic; maybe heterothallic (De Ravin et al. 2011). ITS barcode: JF417483 (alternative markers: BenA = JF417462; CaM = JF417494; RPB2 = JN406532).

Rasamsonia emersonii (Stolk) Houbraken & Frisvad, Antonie van Leeuwenhoek 101: 417. 2012. [MB519874]. Basionym: Talaromyces emersonii Stolk, Antonie van Leeuwenhoek 31: 262. 1965. [MB339920]. — Type: CBS 393.64. Ex-type: DTO 164-E9 = DTO 048-I1 = CBS 393.64 = ATCC 16479 = CECT 2607 = IFO 31232 = IMI 116815 = IMI 116815ii. Reproduction: homothallic. ITS barcode: JF417478 (alternative markers: BenA = JF417463; CaM = JF417510; RPB2 = XM013471581).

Rasamsonia frigidotolerans Rodr.-Andr. et al., Microorganisms 2020, 8, 12: 13. 2020 [MB830608]. — Type: CBS H-23373. Ex-type: CBS 143845 = FMR 16675. Reproduction: asexual. ITS barcode: LT985886 (alternative markers: BenA = LT985895; CaM = LT985897; RPB2 = n.a.).

Rasamsonia piperina Houbraken et al., J. Clin. Microbiol. 51: 26. 2013. [MB801151]. — Type: CBS H-21030. Ex-type: DTO 138-G3 = CBS 408.73 = IJFM 1326. Reproduction: asexual. ITS barcode: JX272968 (alternative markers: BenA = JX273000; CaM = JX272936; RPB2 = MN969194).

Rasamsonia pulvericola Tanney & Seifert, IMA Fungus 4: 207. 2013. [MB804677]. — Type: DAOM 242435. Ex-type: DAOM 242435. Reproduction: asexual. ITS barcode: KF242514 (alternative markers: BenA = KF242520; CaM = KF242522; RPB2 = KF242518).



Sagenomella

Sagenomella diversispora (J.F.H. Beyma) W. Gams, Persoonia 10: 102. 1978. [MB323034]. Basionym: Scopulariopsis diversispora J.F.H. Beyma, Zentralbl. Bakteriol. Parasitenk., Abt. 2 96: 430. 1937. [MB263394]. — Type: van Beyma 1937, Zentralbl. Bakteriol. Parasitenk., Abt. 2 96: p. 431 Abb. 1 (– lectotype designated here, MBT392364). Ex-type: CBS 354.36 = IAM 14790 = MUCL 9029. Reproduction: asexual. ITS barcode: MN431407 (alternative markers: BenA = MN969427; CaM = MN969348; RPB2 = MN969207).

Sagenomella griseoviridis (Onions & G.L. Barron) W. Gams, Persoonia 10: 102. 1978. [MB323035]. Basionym: Paecilomyces griseoviridis Onions & G.L. Barron, Mycol. Pap. 107: 22. 1967. [MB335525]. — Type: CBS 426.67. Ex-type: CBS 426.67 = ATCC 18505 = IMI 113160. Reproduction: asexual. ITS barcode: MN431406 (alternative markers: BenA = MN969426; CaM = MN969347; RPB2 = JF417438).

Sagenomella humicola (Onions & G.L. Barron) W. Gams, Persoonia 10: 102. 1978. [MB323036]. Basionym: Paecilomyces humicola Onions & G.L. Barron, Mycol. Pap. 107: 20. 1967. [MB335526]. — Type: IMI 113166. Ex-type: CBS 427.67 = ATCC 18506 = IAM 14793 = IMI 113166. Reproduction: asexual. ITS barcode: MH859021 (alternative markers: BenA = MN969428; CaM = MN969349; RPB2 = JF417439).

Sagenomella ocotl (Bills & Heredia) Samson et al., Stud. Mycol. 70: 179. 2011. [MB560681]. Basionym: Talaromyces ocotl Bills & Heredia, Mycologia 93: 533. 2001. [MB467796]. — Type: BPI GB6125. Ex-type: CBS 102855 = BPI GB6125. Reproduction: homothallic. ITS barcode: AF285113 (alternative markers: BenA = MN969431; CaM = MN969352; RPB2 = n.a.).

Sagenomella striatispora (Onions & G.L. Barron) W. Gams, Persoonia 10: 102. 1978. [MB323040]. Basionym: Paecilomyces striatisporus Onions & G.L. Barron, Mycol. Pap. 107: 19. 1967. [MB335536]. — Type: IMI 113163. Ex-type: CBS 429.67 = DTO 107-B2 = ATCC 18510 = IAM 14795 = IMI 113163. Reproduction: asexual. ITS barcode: MN431408 (alternative markers: BenA = MN969429; CaM = MN969350; RPB2 = JF417440).

Sagenomella verticillata W. Gams & B.E. Söderstr., Persoonia 10: 107. 1978. [MB323041]. — Type: CBS 414.78. Ex-type: CBS 414.78 = IAM 14697. Reproduction: asexual. ITS barcode: MN431409 (alternative markers: BenA = MN969430; CaM = MN969351; RPB2 = MN969208).



Sclerocleista

Sclerocleista ornata (Raper et al.) Subram., Curr. Sci. 41: 757. 1972. [MB323241]. Basionym: Aspergillus ornatus Raper et al., Mycologia 45: 678. 1953. [MB292852]. — Type: IMI 55295. Ex-type: CBS 124.53 = ATCC 16921 = IMI 055295 = LSHB BB311 = MUCL 15643 = NRRL 2256 = QM 1951 = UC 4518 = WB 2256. Reproduction: homothallic. ITS barcode: EF669704 (alternative markers: BenA = EF669676; CaM = EF669690; RPB2 = EF669663).

Sclerocleista thaxteri Subram., Curr. Sci. 41: 757. 1972. [MB323242]. — Type: On caterpillar dung, "Kittery Point", coll. R. Thaxter ex Farlow Herbarium. Harvard University. Ex-type: CBS 105.25 = IFO 4042 = IFO 8130 = IMI 055296 = NRRL 2292 = WB 2292. Reproduction: homothallic. ITS barcode: EU021599 (alternative markers: BenA = EU021668; CaM = EU021689; RPB2 = EU021630).



Talaromyces

Talaromyces acaricola Visagie et al., Persoonia 36: 49. 2016. [MB810899]. — Type: CBS H-21632. Ex-type: CBS 137386 = DTO 183-B3 = DAOM 241025 = IBT 32387. Infragen. class: sect. Islandici. Reproduction: asexual. ITS barcode: JX091476 (alternative markers: BenA = JX091610; CaM = JX140729; RPB2 = KF984956).

Talaromyces aculeatus (Raper & Fennell) Samson et al., Stud. Mycol. 71: 174. 2011. [MB560639]. Basionym: Penicillium aculeatum Raper & Fennell, Mycologia 40: 535. 1948. [MB289073]. — Type: IMI 040588. Ex-type: CBS 289.48 = ATCC 10409 = IMI 040588 = NRRL 2129 = NRRL A-1474. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: KF741995 (alternative markers: BenA = KF741929; CaM = KF741975; RPB2 = MH793099).

Talaromyces adpressus A.J. Chen et al., Stud. Mycol. 84:124. 2016 [MB817397]. — Type: CBS H-22507. Ex-type: CBS 140620 = CGMCC3.18211 = DTO 317-G4. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: KU866657 (alternative markers: BenA = KU866844; CaM = KU866741; RPB2 = KU867001).

Talaromyces aerius A.J. Chen et al., Stud. Mycol. 84:124. 2016 [MB817398]. — Type: CBS H-22506. Ex-type: CBS 140611 = CGMCC3.18197 = DTO 317-C7. Infragen. class: sect. Trachyspermi. Reproduction: asexual. ITS barcode: KU866647 (alternative markers: BenA = KU866835; CaM = KU866731; RPB2 = KU866991).

Talaromyces aerugineus (Samson) N. Yilmaz et al., Stud. Mycol. 78: 210. 2014. [MB809553]. Basionym: Paecilomyces aerugineus Samson, Stud. Mycol. 6: 20. 1974. [MB319096]. — Type: CBS H-7448. Ex-type: CBS 350.66 = BDUN 276 = IMI 105412. Infragen. class: sect. Helici. Reproduction: asexual. ITS barcode: AY753346 (alternative markers: BenA = KJ865736; CaM = KJ885285; RPB2 = JN121502).

Talaromyces affinitatimellis Rodr.-Andr. et al., IMA Fungus 10: 20. 2019. [MB823591]. — Type: CBS H-23370. Ex-type: FMR 15690 = CBS 143840. Infragen. class: sect. Trachyspermi. Reproduction: asexual. ITS barcode: LT906543 (alternative markers: BenA = LT906552; CaM = LT906549; RPB2 = LT906546).

Talaromyces albobiverticillius (H.-M. Hsieh et al.) Samson et al., Stud. Mycol. 71: 174. 2011. [MB560683]. Basionym: Penicillium albobiverticillium H.M. Hsieh, Y.M. Ju & S.Y. Hsieh, Fungal Science 25: 26. 2010. [MB519193]. — Type: BCRC 34774. Ex-type: CBS 133440 = DTO 166-E5 = YMJ 1292. Infragen. class: sect. Trachyspermi. Reproduction: asexual. ITS barcode: HQ605705 (alternative markers: BenA = KF114778; CaM = KJ885258; RPB2 = KM023310).

Talaromyces allahabadensis (B.S. Mehrotra & D. Kumar) Samson et al., Stud. Mycol. 71: 174. 2011. [MB560640]. Basionym: Penicillium allahabadense B.S. Mehrotra & D. Kumar, Canad. J. Bot. 40: 1399. 1962. [MB335707]. — Type: University of Allahabad P-26. Ex-type: CBS 453.93 = ATCC 15067 = CBS 304.63. Infragen. class: sect. Islandici. Reproduction: asexual. ITS barcode: KF984873 (alternative markers: BenA = KF984614; CaM = KF984768; RPB2 = KF985006).

Talaromyces alveolaris Guevara-Suarez et al., Mycoses 60: 656. 2017. [MB820459]. — Type: CBS H-22999. Ex-type: CBS 142379 = UTHSC DI16-147 = FMR 13963. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: LT558969 (alternative markers: BenA = LT559086; CaM = LT795596; RPB2 = LT795597).

Talaromyces amazonensis N. Yilmaz et al., Mycol. Prog. 15: 1052. 2016. [MB816230]. — Type: HUA 197223. Ex-type: CBS 140373 = IBT 23215 = DTO 093-F9. Infragen. class: sect. Talaromyces. Reproduction: homothallic. ITS barcode: KX011509 (alternative markers: BenA = KX011490; CaM = KX011502; RPB2 = MN969186).

Talaromyces amestolkiae N. Yilmaz et al., Persoonia 29: 48. 2012. [MB801358]. — Type: CBS H-21050. Ex-type: CBS 132696 = DTO 179-F5. Infragen. class: sect. Talaromyces. Reproduction: Heterothallic (Yilmaz et al. 2016). ITS barcode: JX315660 (alternative markers: BenA = JX315623; CaM = KF741937; RPB2 = JX315698).

Talaromyces amyrossmaniae Rajeshkumar et al., MycoKeys 45: 47. 2019. [MB518601]. — Type: AMH 9330. Ex-type: NFCCI 1919 = KAS 3038. Infragen. class: sect. Trachyspermi. Reproduction: asexual. ITS barcode: MH909062 (alternative markers: BenA = MH909064; CaM = MH909068; RPB2 = MH909066).

Talaromyces angelicae S.H. Yu et al., J. Microbiol. 51: 707. 2013. [MB818651]. — Type: KACC 46611. Ex-type: KACC 46611. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: KF183638 (alternative markers: BenA = KF183640; CaM = KJ885259; RPB2 = KX961275).

Talaromyces annesophieae Houbraken, Persoonia 39: 461. 2017. [MB823027]. — Type: CBS H-23216. Ex-type: CBS 142939 = DTO 377-F3 = JW9011. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: MF574592 (alternative markers: BenA = MF590098; CaM = MF590104; RPB2 = MN969199).

Talaromyces apiculatus Samson et al., Stud. Mycol. 71: 174. 2011. [MB560641]. — Type: CBS H-20755. Ex-type: CBS 312.59 = ATCC 18315 = FRR 635 = IMI 068239. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: JN899375 (alternative markers: BenA = KF741916; CaM = KF741950; RPB2 = KM023287).

Talaromyces argentinensis Jurjević & S.W. Peterson, Fungal Biol. 123: 751. 2019. [MB827826]. — Type: BPI 910716. Ex-type: NRRL 28750. Infragen. class: sect. Talaromyces. Reproduction: homothallic. ITS barcode: MH793045 (alternative markers: BenA = MH792917; CaM = MH792981; RPB2 = MH793108).

Talaromyces assiutensis Samson & Abdel-Fattah, Persoonia 9: 501. 1978. [MB324414]. — Type: CBS 147.78. Ex-type: CBS 147.78. Infragen. class: sect. Trachyspermi. Reproduction: homothallic. ITS barcode: JN899323 (alternative markers: BenA = KJ865720; CaM = KJ885260; RPB2 = KM023305).

Talaromyces atricola S.W. Peterson & Jurjević, PLoS ONE 8: e78084, 8. 2013. [MB804733]. — Type: unknown. Ex-type: CBS 255.31 = NRRL 1052 = FRR 1052 = Thom 4640.439 = ATCC 52257. Infragen. class: sect. Islandici. Reproduction: asexual. ITS barcode: KF984859 (alternative markers: BenA = KF984566; CaM = KF984719; RPB2 = KF984948).

Talaromyces atroroseus N. Yilmaz et al., PLoS ONE 8: e84102, 8. 2013. [MB804901]. — Type: CBS H-21790. Ex-type: CBS 133442 = IBT 32470 = DTO 178-A4. Infragen. class: sect. Trachyspermi. Reproduction: asexual. ITS barcode: KF114747 (alternative markers: BenA = KF114789; CaM = KJ775418; RPB2 = KM023288).

Talaromyces aurantiacus (J.H. Mill. et al.) Samson et al., Stud. Mycol. 71: 175. 2011. [MB560642]. Basionym: Penicillium aurantiacum J.H. Mill. et al., Mycologia 49: 797. 1957. [MB302378]. — Type: No. 1736 (A.A. Foster). Ex-type: CBS 314.59 = ATCC 13216 = IMI 099722 = NRRL 3398. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: JN899380 (alternative markers: BenA = KF741917; CaM = KF741951; RPB2 = KX961285).

Talaromyces australis Visagie et al., Mycoscience 56: 492 2015. [MB808236]. — Type: CBS H-21598. Ex-type: CBS 137102 = DTO 273-F5 = IBT 14256 = FRR 2005. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: KF741991 (alternative markers: BenA = KF741922; CaM = KF741971; RPB2 = KX961284).

Talaromyces austrocalifornicus Yaguchi & Udagawa, Trans. Mycol. Soc. Japan 34: 245. 1993. [MB361182]. — Type: CBM-PF 1117. Ex-type: CBS 644.95 = IBT 17522. Infragen. class: sect. Trachyspermi. Reproduction: homothallic. ITS barcode: JN899357 (alternative markers: BenA = KJ865732; CaM = KJ885261; RPB2 = MN969147).

Talaromyces bacillisporus [as “bacillosporus”] (Swift) C.R. Benj., Mycologia 47: 682. 1955. [MB118745]. Basionym: Penicillium bacillisporum Swift, Bull. Torrey Bot. Club: 221. 1932. [MB119961]. — Type: CBS H-7813 (Isotype). Ex-type: CBS 296.48 = ATCC 10126 = IMI 040045 = NRRL 1025. Infragen. class: sect. Bacillispori. Reproduction: homothallic. ITS barcode: KM066182 (alternative markers: BenA = AY753368; CaM = KJ885262; RPB2 = JF417425).

Talaromyces basipetosporus Stchigel et al., IMA Fungus 10: 20, 17. 2019. [MB823589]. — Type: CBS H-23365. Ex-type: FMR 9720 = CBS 143836. Infragen. class: sect. Trachyspermi. Reproduction: asexual. ITS barcode: LT906542 (alternative markers: BenA = LT906563; CaM = n.a.; RPB2 = LT906545).

Talaromyces beijingensis A.J. Chen et al., Stud. Mycol. 84:125. 2016 [MB817395]. — Type: CBS H-22508. Ex-type: CBS 140617 = CGMCC3.18200 = DTO 317-D8. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: KU866649 (alternative markers: BenA = KU866837; CaM = KU866733; RPB2 = KU866993).

Talaromyces bohemicus (Fassat. & Pěčková) N. Yilmaz et al., Stud. Mycol. 78: 227. 2014 [MB809554]. Basionym: Sagenomella bohemica Fassat. & Pecková, Ceská Mykologie 44: 240. 1990. [MB361859]. — Type: unknown. Ex-type: CBS 545.86 = CCF 2330 = IAM 14789. Infragen. class: sect. Helici. Reproduction: asexual. ITS barcode: JN899400 (alternative markers: BenA = KJ865719; CaM = KJ885286; RPB2 = JN121532).

Talaromyces boninensis (Yaguchi & Udagawa) Samson et al., Stud. Mycol. 71: 175. 2011. [MB560643]. Basionym: Talaromyces helicus var. boninensis Yaguchi & Udagawa, Trans. Mycol. Soc. Japan 33: 511. 1992. [MB359708]. — Type: CBM PF-1103. Ex-type: CBS 650.95 = IBT 17516 . Infragen. class: sect. Helici. Reproduction: homothallic. ITS barcode: JN899356 (alternative markers: BenA = KJ865721; CaM = KJ885263; RPB2 = KM023276).

Talaromyces borbonicus Houbraken, Mycologia 110: 318. 2018. [MB821643]. — Type: CBS H-22672. Ex-type: CBS 141340 = DTO 351-D3. Infragen. class: sect. Helici. Reproduction: asexual. ITS barcode: MG827091 (alternative markers: BenA = MG855687; CaM = MG855688; RPB2 = MG855689).

Talaromyces brasiliensis R.N. Barbosa et al., Antonie van Leeuwenhoek 111: 1902. 2018 [MB822214]. — Type: URM 90494. Ex-type: CBS 142493 = URM 7618. Infragen. class: sect. Trachyspermi. Reproduction: asexual. ITS barcode: MF278323 (alternative markers: BenA = LT855560; CaM = LT855563; RPB2 = MN969198).

Talaromyces brunneosporus Rodr.-Andr. et al., IMA Fungus 10: 20, 19. 2019. [MB823590]. — Type: CBS H-23375. Ex-type: FMR 16566 = CBS 144320. Infragen. class: sect. Purpurei (tentative, see Supplementary Fig. S3). Reproduction: asexual. ITS barcode: LT962487 (alternative markers: BenA = LT962483; CaM = LT962488; RPB2 = LT962485).

Talaromyces brunneus (Udagawa) Samson et al., Stud. Mycol. 71: 175. 2011. [MB560644]. Basionym: Penicillium brunneum Udagawa, J. Agric. Sci. Tokyo Nogyo Daig. 5: 16. 1959. [MB302383]. — Type: NHL 6054. Ex-type: CBS 227.60 = ATCC 18229 = FRR 646 = IFO 6438 = IHEM 3907 = IMI 078259 = MUCL 31318. Infragen. class: sect. Islandici. Reproduction: asexual. ITS barcode: JN899365 (alternative markers: BenA = KJ865722; CaM = KJ885264; RPB2 = KM023272).

Talaromyces calidicanius (J.L. Chen) Samson et al., Stud. Mycol. 71: 175. 2011. [MB560645]. Basionym: Penicillium calidicanium J.L. Chen, Mycologia 94: 870. 2002. [MB483963]. — Type: CFC-7 (isotype TNM F12246). Ex-type: CBS 112002. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: JN899319 (alternative markers: BenA = HQ156944; CaM = KF741934; RPB2 = KM023311).

Talaromyces californicus Jurjević & S.W. Peterson, Fungal Biol. 123: 752. 2019. [MB827827]. — Type: BPI 910714. Ex-type: NRRL 58168. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: MH793056 (alternative markers: BenA = MH792928; CaM = MH792992; RPB2 = MH793119).

Talaromyces catalonicus Guevara-Suarez et al., Fungal Syst. Evol. 5: 68. 2020. [MB822080]. — Type: CBS H-23212. Ex-type: CBS 143039 = FMR 16441. Infragen. class: sect. Trachyspermi. Reproduction: asexual. ITS barcode: LT899793 (alternative markers: BenA = LT898318; CaM = LT899775; RPB2 = LT899811).

Talaromyces cecidicola (Seifert et al.) Samson et al., Stud. Mycol. 71: 175. 2011. [MB560646]. Basionym: Penicillium cecidicola Seifert, Hoekstra & Frisvad, Stud. Mycol. 50: 520. 2004. [MB500150]. — Type: DAOM 233329. Ex-type: CBS 101419 = DAOM233329. Infragen. class: sect. Purpurei. Reproduction: asexual. ITS barcode: AY787844 (alternative markers: BenA = FJ753295; CaM = KJ885287; RPB2 = KM023309).

Talaromyces cerinus A.J. Chen et al., Stud. Mycol. 84:125. 2016 [MB817393]. — Type: CBS H-22513. Ex-type: CBS 140622 = CGMCC3.18212 = DTO 318-A2. Infragen. class: sect. Islandici. Reproduction: asexual. ITS barcode: KU866658 (alternative markers: BenA = KU866845; CaM = KU866742; RPB2 = KU867002).

Talaromyces chlamydosporus A.J. Chen et al., Stud. Mycol. 84:136. 2016 [MB817392]. — Type: CBS H-22509. Ex-type: CBS 140635 = CGMCC 3.18199 = DTO 317-D5. Infragen. class: sect. Islandici. Reproduction: asexual. ITS barcode: KU866648 (alternative markers: BenA = KU866836; CaM = KU866732; RPB2 = KU866992).

Talaromyces chlorolomus [as “chloroloma”] Visagie & K. Jacobs, Persoonia 28: 18. 2012. [MB585178]. — Type: PREM 60033. Ex-type: DAOM 241016 = CV 2802. Infragen. class: sect. Purpurei. Reproduction: asexual. ITS barcode: FJ160273 (alternative markers: BenA = GU385736; CaM = KJ885265; RPB2 = KM023304).

Talaromyces cinnabarinus (S.C. Jong & E.E. Davis) N. Yilmaz et al., Stud. Mycol. 78: 234. 2014. [MB809557]. Basionym: Paecilomyces cinnabarinus S.C. Jong & E.E. Davis, Mycologia 67: 1144. 1976. [MB319103]. — Type: CBS H-6686. Ex-type: CBS 267.72 = ATCC 26215 = NHL 2673. Infragen. class: sect. Helici. Reproduction: homothallic. ITS barcode: JN899376 (alternative markers: BenA = AY753377; CaM = KJ885256; RPB2 = JN121477).

Talaromyces clemensii Viisagie & N. Yilmaz, Persoonia 43: 407. 2019. [MB832488]. — Type: PREM 62301. Ex-type: PPRI 26753 = CMV016A4. Infragen. class: sect. Trachyspermi. Reproduction: asexual. ITS barcode: MK951940 (alternative markers: BenA = MK951833; CaM = MK951906; RPB2 = MN418451).

Talaromyces cnidii S.H. Yu et al., J. Microbiol. 51: 707. 2013. [MB804809]. — Type: KACC 46617. Ex-type: KACC 46617. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: KF183639 (alternative markers: BenA = KF183641; CaM = KJ885266; RPB2 = KM023299).

Talaromyces coalescens (Quintan.) Samson et al., Stud. Mycol. 71: 175. 2011. [MB560647]. Basionym: Penicillium coalescens Quintan., Mycopathologia 84 (2–3): 115. 1984. [MB107101]. — Type: CBS 103.83. Ex-type: CBS 103.83. Infragen. class: sect. Purpurei. Reproduction: asexual. ITS barcode: JN899366 (alternative markers: BenA = JX091390; CaM = KJ885267; RPB2 = KM023277).

Talaromyces columbiensis N. Yilmaz et al., Mycol. Prog. 15: 1053. 2016. [MB816231]. — Type: HUA 197225. Ex-type: CBS 113151 = IBT 23206 = DTO 058-F3. Infragen. class: sect. Bacillispori. Reproduction: homothallic. ITS barcode: KX011503 (alternative markers: BenA = KX011488; CaM = KX011499; RPB2 = MN969187).

Talaromyces columbinus S.W. Peterson & Jurjević, PLoS ONE 8: e78084, 6. 2013. [MB804732]. — Type: BPI 892668. Ex-type: NRRL 58811. Infragen. class: sect. Islandici. Reproduction: asexual. ITS barcode: KJ865739 (alternative markers: BenA = KF196843; CaM = KJ885288; RPB2 = KM023270).

Talaromyces convolutus Udawaga, Mycotaxon 48: 141. 1993. [MB360474]. — Type: CBM SUM-3018. Ex-type: CBS 100537 = IBT 14989. Infragen. class: sect. Trachyspermi. Reproduction: homothallic. ITS barcode: JN899330 (alternative markers: BenA = KF114773; CaM = MN969316; RPB2 = JN121414).

Talaromyces coprophilus Guevara-Suarez et al., Fungal Syst. Evol. 5: 68. 2020. [MB822088]. — Type: CBS H-23144. Ex-type: CBS 142756 = FMR 15199. Infragen. class: sect. Talaromyces. Reproduction: homothallic. ITS barcode: LT899794 (alternative markers: BenA = LT898319; CaM = LT899776; RPB2 = LT899812).

Talaromyces crassus Visagie et al., Persoonia 36: 49. 2016. [MB810900]. — Type: CBS H-21631. Ex-type: CBS 137381 = DTO 181-C5 = DAOM 241027 = IBT 32814. Infragen. class: sect. Islandici. Reproduction: asexual. ITS barcode: JX091472 (alternative markers: BenA = JX091608; CaM = JX140727; RPB2 = KF984914).

Talaromyces cucurbitiradicus L. Su & Y.C. Niu, Mycologia 110: 380. 2018. [MB820559]. — Type: HMAS 247175. Ex-type: ACCC 39155. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: KY053254 (alternative markers: BenA = KY053228; CaM = KY053246; RPB2 = n.a.).

Talaromyces delawarensis Jurjević & S.W. Peterson, Mycologia 109: 541. 2017. [MB817998]. — Type: BPI-910148. Ex-type: NRRL 58874. Infragen. class: sect. Islandici. Reproduction: asexual. ITS barcode: KX657324 (alternative markers: BenA = KX657055; CaM = KX657158; RPB2 = KX657490).

Talaromyces dendriticus (Pitt) Samson et al., Stud. Mycol. 71: 175. 2011. [MB560648]. Basionym: Penicillium dendriticum Pitt, Gen. Penicillium: 413. 1980. [MB116318]. — Type: IMI 216897. Ex-type: CBS 660.80 = IMI 216897. Infragen. class: sect. Purpurei. Reproduction: asexual. ITS barcode: JN899339 (alternative markers: BenA = JX091391; CaM = KF741965; RPB2 = KM023286).

Talaromyces derxii Takada & Udagawa, Mycotaxon 31: 418. 1988. [MB133755]. — Type: NHL 2980. Ex-type: CBS 412.89 = NHL 2981. Infragen. class: sect. Talaromyces. Reproduction: Heterothallic. ITS barcode: JN899327 (alternative markers: BenA = JX494306; CaM = KF741959; RPB2 = KM023282).

Talaromyces dimorphus X.Z. Jiang & L. Wang, Sci. Rep. 8: 4932, 2. 2018. [MB570521]. — Type: HMAS 247023. Ex-type: AS3.15692 = NN072337. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: KY007095 (alternative markers: BenA = KY007111; CaM = KY007103; RPB2 = KY112593).

Talaromyces diversiformis A.J. Chen et al., Stud. Mycol. 84:136. 2016 [MB818696]. — Type: CBS H-22841. Ex-type: CBS 141931 = CGMCC 3.18204 = DTO 317-E3. Infragen. class: sect. Helici. Reproduction: asexual. ITS barcode: KX961215 (alternative markers: BenA = KX961216; CaM = KX961259; RPB2 = KX961274).

Talaromyces diversus (Raper & Fennell) Samson et al., Stud. Mycol. 71: 175. 2011. [MB560649]. Basionym: Penicillium diversum Raper & Fennell, Mycologia 40: 539. 1948. [MB289080]. — Type: IMI 040579. Ex-type: CBS 320.48 = ATCC 10437 = DSM 2212 = IMI 040579 = IMI 040579ii = NRRL 2121. Infragen. class: sect. Trachyspermi. Reproduction: asexual. ITS barcode: KJ865740 (alternative markers: BenA = KJ865723; CaM = KJ885268; RPB2 = KM023285).

Talaromyces domesticus Jurjević & S.W. Peterson, Fungal Biol. 123: 754. 2019. [MB827828]. — Type: BPI 910711. Ex-type: NRRL 58121. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: MH793055 (alternative markers: BenA = MH792927; CaM = MH792991; RPB2 = MH793118).

Talaromyces duclauxii (Delacr.) Samson et al., Stud. Mycol. 71: 175. 2011. [MB560650]. Basionym: Penicillium duclauxii Delacr., Bull. Soc. Mycol. France 7: 107. 1891. [MB169645]. — Type: IMI 24312. Ex-type: CBS 322.48 = ATCC 10439 = IMI 040044 = MUCL 28672 = MUCL 29094 = MUCL 29212 = NRRL 1030. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: JN899342 (alternative markers: BenA = JX091384; CaM = KF741955; RPB2 = JN121491).

Talaromyces emodensis Udagawa, Mycotaxon 48: 146. 1993. [MB360476]. — Type: CBM SUM-3025. Ex-type: CBS 100536 = IBT 14990. Infragen. class: sect. Bacillispori. Reproduction: homothallic. ITS barcode: JN899337 (alternative markers: BenA = KJ865724; CaM = KJ885269; RPB2 = JF417445).

Talaromyces endophyticus L. Su & Y.C. Niu, Mycologia 110: 380. 2018. [MB817808]. — Type: HMAS 254730. Ex-type: ACCC 39141. Infragen. class: sect. Islandici. Reproduction: asexual. ITS barcode: KX639168 (alternative markers: BenA = KX639174; CaM = KX639165; RPB2 = n.a.).

Talaromyces erythromellis (A.D. Hocking) Samson et al., Stud. Mycol. 71: 175. 2011. [MB560652]. Basionym: Penicillium erythromellis A.D. Hocking, The Genus Penicillium: 459. 1980 [1979]. [MB116314]. — Type: IMI 216899. Ex-type: CBS 644.80 = FRR 1868 = IMI 216899. Infragen. class: sect. Trachyspermi. Reproduction: protoheterothallic, MAT1-1-1 detected (López-Villavicencio et al. 2010). ITS barcode: JN899383 (alternative markers: BenA = HQ156945; CaM = KJ885270; RPB2 = KM023290).

Talaromyces euchlorocarpius Yaguchi et al., Mycoscience 40: 133. 1999. [MB460481]. — Type: CBM-PF 1203. Ex-type: PF 1203 = DTO 176-I3 = DTO 176-I4. Infragen. class: sect. Talaromyces. Reproduction: homothallic. ITS barcode: AB176617 (alternative markers: BenA = KJ865733; CaM = KJ885271; RPB2 = KM023303).

Talaromyces flavovirens (Durieu & Mont.) Visagie et al., Mycotaxon 122: 404. 2012. [MB800438]. Basionym: Lasioderma flavovirens Durieu & Mont., Ann. Sci. Nat. Bot. 4: 364. 1845. [MB120473]. — Type: PC 0088796 (lectotype); BCC 473 (epitype) = BCN 473. Ex-type: CBS 102801 = IBT 27044. Infragen. class: sect. Talaromyces. Reproduction: Heterothallic. ITS barcode: JN899392 (alternative markers: BenA = JX091376; CaM = KF741933; RPB2 = KX961283).

Talaromyces flavus (Klöcker) Stolk & Samson, Stud. Mycol. 2: 10. 1972. [MB324416]. Basionym: Gymnoascus flavus Klöcker, Hedwigia 41: 80. 1902. [MB225997]. — Type: CBS H-7820. Ex-type: CBS 310.38 = IMI 197477 = NRRL 2098. Infragen. class: sect. Talaromyces. Reproduction: homothallic. ITS barcode: JN899360 (alternative markers: BenA = JX494302; CaM = KF741949; RPB2 = JF417426).

Talaromyces francoae N. Yilmaz et al., Mycol. Prog. 15: 1054. 2016. [MB816232]. — Type: HUA 197224. Ex-type: CBS 113134 = IBT 23221 = DTO 056-D9. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: KX011510 (alternative markers: BenA = KX011489; CaM = KX011501; RPB2 = MN969188).

Talaromyces funiculosus (Thom) Samson et al., Stud. Mycol. 71: 176. 2011. [MB560653]. Basionym: Penicillium funiculosum Thom, U.S.D.A. Bur. Ani. Ind. Bull. 118: 69. 1910. [MB152047]. — Type: IMI 193019. Ex-type: CBS 272.86 = IMI 193019. Infragen. class: sect. Talaromyces. Reproduction: protoheterothallic, both mating types detected (López-Villavicencio et al. 2010). ITS barcode: JN899377 (alternative markers: BenA = MN969408; CaM = KF741945; RPB2 = KM023293).

Talaromyces fuscoviridis Visagie et al., Mycoscience 56: 492. 2015. [MB808234]. — Type: CBS H-21603. Ex-type: CBS 193.69 = DTO 258-I7 = IBT 14846 = IBT 32646. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: KF741979 (alternative markers: BenA = KF741912; CaM = KF741942; RPB2 = MN969156).

Talaromyces fusiformis A.J. Chen et al., Stud. Mycol. 84:139. 2016 [MB817396]. — Type: CBS H-22510. Ex-type: CBS 140637 = CGMCC3.18210 = DTO 317-F4. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: KU866656 (alternative markers: BenA = KU866843; CaM = KU866740; RPB2 = KU867000).

Talaromyces galapagensis Samson & Mahoney, Trans. Brit. Mycol. Soc. 69: 158. 1977. [MB324417]. — Type: CBS H-7489. Ex-type: CBS 751.74 = IFO 31796. Infragen. class: sect. Talaromyces. Reproduction: homothallic. ITS barcode: JN899358 (alternative markers: BenA = JX091388; CaM = KF741966; RPB2 = KX961280).

Talaromyces georgiensis M. Guevara-Suarez et al., Mycoses 60: 656. 2017. [MB820460]. — Type: CBS H-23000. Ex-type: UTHSC DI16-145 = CBS 142380 = DTO 410-E8 = FMR 14270. Infragen. class: sect. Helici. Reproduction: asexual. ITS barcode: LT558967 (alternative markers: BenA = LT559084; CaM = n.a.; RPB2 = LT795606).

Talaromyces guatemalensis A. Nováková et al., Persoonia 43: 409. 2019. [MB832313]. — Type: PRM 952195. Ex-type: CCF 6215 = GUA2˗1. Infragen. class: sect. Trachyspermi. Reproduction: asexual. ITS barcode: MN322789 (alternative markers: BenA = MN329687; CaM = MN329688; RPB2 = MN329689).

Talaromyces hachijoensis Yaguchi et al., Mycoscience 37: 157. 1996. [MB416016]. — Type: PF 1174. Ex-type: CBM-PF 1174 = IFM 53624. Infragen. class: sect. Bacillispori. Reproduction: homothallic. ITS barcode: AB176620 (alternative markers: BenA = n.a.; CaM = n.a.; RPB2 = n.a.).

Talaromyces halophytorum Y.H. You & S.B. Hong, Mycobiology, accepted. 2020. [MB830295]. — Type: KACC 48127. Ex-type: KACC 48127 = NIBRFGC 000501933. Infragen. class: sect. Trachyspermi. Reproduction: asexual. ITS barcode: MH725786 (alternative markers: BenA = MH729367; CaM = MK111426; RPB2 = MK111427).

Talaromyces heiheensis X.C. Wang & W.Y. Zhuang, Mycol. Prog. 16: 75. 2016. [MB570286]. — Type: HMAS 248789. Ex-type: CGMCC 3.18012. Infragen. class: sect. Trachyspermi. Reproduction: asexual. ITS barcode: KX447526 (alternative markers: BenA = KX447525; CaM = KX447532; RPB2 = KX447529).

Talaromyces helicus (Raper & Fennel) C.R. Benj., Mycologia 47: 684. 1955. [MB306715]. Basionym: Penicillium helicum Raper & Fennell, Mycologia 40: 515. 1948. [MB289086]. — Type: IMI 040593. Ex-type: CBS 335.48 = ATCC 10451 = DSM 3705 = IMI 040593 = NRRL 2106. Infragen. class: sect. Helici. Reproduction: homothallic. ITS barcode: JN899359 (alternative markers: BenA = KJ865725; CaM = KJ885289; RPB2 = KM023273).

Talaromyces herodensis Jurjević & S.W. Peterson, Mycologia 109: 546. 2017. [MB818001]. — Type: BPI-910152. Ex-type: NRRL 62467. Infragen. class: sect. Islandici. Reproduction: asexual. ITS barcode: KX657338 (alternative markers: BenA = KX657061; CaM = KX657182; RPB2 = KX657524).

Talaromyces indigoticus Takada & Udagawa, Mycotaxon 46: 129. 1993. [MB359290]. — Type: CBM SUM-3010. Ex-type: CBS 100534 = IBT 17590. Infragen. class: sect. Talaromyces. Reproduction: homothallic. ITS barcode: JN899331 (alternative markers: BenA = JX494308; CaM = KF741931; RPB2 = KX961278).

Talaromyces infraolivaceus Visagie et al., Persoonia 36: 52. 2016. [MB810901]. — Type: CBS H-21633. Ex-type: CBS 137385 = DTO 182-I2 = DAOM 241024 = IBT 32487. Infragen. class: sect. Islandici. Reproduction: asexual. ITS barcode: JX091481 (alternative markers: BenA = JX091615; CaM = JX140734; RPB2 = KF984949).

Talaromyces intermedius (Apinis) Stolk & Samson, Stud. Mycol. 2: 21. 1972. [MB324418]. Basionym: Arachniotus intermedius Apinis, Mycol. Pap. 96: 45. 1964. [MB326426]. — Type: CBS H-7828. Ex-type: CBS 152.65 = BDUN 267 = IFO 31752 = IMI 100874. Infragen. class: sect. Talaromyces. Reproduction: homothallic. ITS barcode: JN899332 (alternative markers: BenA = JX091387; CaM = KJ885290; RPB2 = KX961282).

Talaromyces iowaense Jurjević et al., Persoonia 41: 407. 2018. [MB828092]. — Type: BPI 910643. Ex-type: NRRL 66822 = ITEM 17527 = EMSL 2233. Infragen. class: sect. Purpurei. Reproduction: asexual. ITS barcode: MH281565 (alternative markers: BenA = MH282578; CaM = MH282579; RPB2 = MH282577).

Talaromyces islandicus (Sopp) Samson et al., Stud. Mycol. 71: 176. 2011. [MB560654]. Basionym: Penicillium islandicum Sopp, Skr. VidenskSelsk. Christiania, Kl. I, Math.-Natur. (no. 11): 161. 1912. [MB162841]. — Type: IMI 040042. Ex-type: CBS 338.48 = ATCC 10127 = IMI 040042 = MUCL 31324 = NRRL 1036. Infragen. class: sect. Islandici. Reproduction: asexual. ITS barcode: KF984885 (alternative markers: BenA = KF984655; CaM = KF984780; RPB2 = KF985018).

Talaromyces juglandicola Jurjević & S.W. Peterson, Mycologia 109: 551. 2017. [MB818006]. — Type: BPI-910144. Ex-type: NRRL 32382. Infragen. class: sect. Islandici. Reproduction: asexual. ITS barcode: KX657330 (alternative markers: BenA = KX657122; CaM = KX657184; RPB2 = KX657573).

Talaromyces kabodanensis Houbraken et al., Persoonia 37: 253. 2016. [MB819001]. — Type: CBS H-22857. Ex-type: CBS 139564 = DTO 204-F2 = CCTU 850. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: KP851981 (alternative markers: BenA = KP851986; CaM = KP851995; RPB2 = MN969190).

Talaromyces kendrickii Visagie et al., Mycoscience 56: 493. 2015. [MB808235]. — Type: CBS H-21599. Ex-type: CBS 136666 = DTO 273-F4 = IBT 14128. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: KF741987 (alternative markers: BenA = KF741921; CaM = KF741967; RPB2 = MN969158).

Talaromyces kilbournensis Jurjević & S.W. Peterson, Mycologia 109: 545. 2017. [MB818000]. — Type: BPI-910153. Ex-type: NRRL 62700. Infragen. class: sect. Islandici. Reproduction: asexual. ITS barcode: KX657344 (alternative markers: BenA = KX657068; CaM = KX657183; RPB2 = KX657545).

Talaromyces lentulus X.Z. Jiang & L. Wang, Sci. Rep. 8: 4932, 3. 2018. [MB570522]. — Type: HMAS 247024. Ex-type: AS3.15689 = NN071323. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: KY007088 (alternative markers: BenA = KY007104; CaM = KY007096; RPB2 = KY112586).

Talaromyces liani (Kamyschko) N. Yilmaz et al., Stud. Mycol. 78: 266. 2014. [MB809555]. Basionym: Penicillium liani Kamyschko, Not. Syst. Crypt. Inst. bot. Acad. Sci. USSR: 86. 1962. [MB335742]. — Type: unknown. Ex-type: CBS 225.66 = ATCC 18325 = ATCC 18331 = IMI 098480 = NRRL 3380 = VKM F-301. Infragen. class: sect. Talaromyces. Reproduction: homothallic. ITS barcode: JN899395 (alternative markers: BenA = JX091380; CaM = KJ885257; RPB2 = KX961277).

Talaromyces loliensis (Pitt) Samson et al., Stud. Mycol. 71: 176. 2011. [MB560655]. Basionym: Penicillium loliense Pitt, Gen. Penicillium: 450. 1980. [MB116312]. — Type: IMI 216901. Ex-type: CBS 643.80 = ATCC 52252 = FRR 1798 = IMI 216901 = MUCL 31325. Infragen. class: sect. Islandici. Reproduction: asexual. ITS barcode: KF984888 (alternative markers: BenA = KF984658; CaM = KF984783; RPB2 = KF985021).

Talaromyces louisianensis Jurjević & S.W. Peterson, Fungal Biol. 123: 755. 2019. [MB827829]. — Type: BPI 910715. Ex-type: NRRL 35823. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: MH793052 (alternative markers: BenA = MH792924; CaM = MH792988; RPB2 = MH793115).

Talaromyces macrosporus (Stolk & Samson) Frisvad et al., Antonie van Leeuwenhoek 57: 186. 1990. [MB126704]. Basionym: Talaromyces flavus var. macrosporus Stolk & Samson, Stud. Mycol 2: 15. 1972. [MB348888]. — Type: CBS H-7822. Ex-type: CBS 317.63 = FRR 404 = IMI 197478. Infragen. class: sect. Talaromyces. Reproduction: homothallic. ITS barcode: JN899333 (alternative markers: BenA = JX091382; CaM = KF741952; RPB2 = KM023292).

Talaromyces mae X.-Z. Jiang & L. Wang, Sci. Rep. 8: 4932, 4. 2018. [MB570523]. — Type: HMAS 247025. Ex-type: AS3.15690 = NN071328. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: KY007090 (alternative markers: BenA = KY007106; CaM = KY007098; RPB2 = KY112588).

Talaromyces malicola Jurjević & S.W. Peterson, Fungal Biol. 123: 756. 2019. [MB827830]. — Type: BPI 910712. Ex-type: NRRL 3724. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: MH909513 (alternative markers: BenA = MH909406; CaM = MH909459; RPB2 = MH909567).

Talaromyces mangshanicus X.C. Wang & W.Y. Zhuang, Mycol. Prog. 16:77. 2016. [MB570288]. — Type: HMAS 248733. Ex-type: CGMCC 3.18013. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: KX447531 (alternative markers: BenA = KX447530; CaM = KX447528; RPB2 = KX447527).

Talaromyces marneffei (Segretain et al.) Samson et al., Stud. Mycol. 71: 176. 2011. [MB560656]. Basionym: Penicillium marneffei Segretain et al., Bull. Soc. Mycol. France 75: 416. 1959. [MB335749]. — Type: IMI 68794iii. Ex-type: CBS 388.87 = ATCC 18224 = CBS 334.59 = IMI 068794ii = IMI 068794iii. Infragen. class: sect. Talaromyces. Reproduction: protoheterothallic (Woo et al. 2006). ITS barcode: JN899344 (alternative markers: BenA = JX091389; CaM = KF741958; RPB2 = KM023283).

Talaromyces mimosinus A.D. Hocking, Gen. Penicillium: 507. 1980. [MB116382]. — Type: IMI 223991. Ex-type: CBS 659.80 = FRR 1875 = IMI 223991. Infragen. class: sect. Bacillispori. Reproduction: homothallic. ITS barcode: JN899338 (alternative markers: BenA = KJ865726; CaM = KJ885272; RPB2 = MN969149).

Talaromyces minioluteus (Dierckx) Samson et al., Stud. Mycol. 71: 176. 2011. [MB560657]. Basionym: Penicillium minioluteum Dierckx, Ann. Soc. Sci. Bruxelles. 25: 87. 1901. [MB157378]. — Type: CBS 642.68. Ex-type: CBS 642.68 = DTO 304-C4 = IMI 089377 = MUCL 28666. Infragen. class: sect. Trachyspermi. Reproduction: asexual. ITS barcode: JN899346 (alternative markers: BenA = MN969409; CaM = KJ885273; RPB2 = JF417443).

Talaromyces minnesotensis Guevara-Suarez et al., Mycoses 60: 657. 2017. [MB820463]. — Type: CBS H-23001. Ex-type: UTHSC DI16-144 = CBS 142381 = FMR 14265. Infragen. class: sect. Trachyspermi. Reproduction: asexual. ITS barcode: LT558966 (alternative markers: BenA = LT559083; CaM = LT795604; RPB2 = LT795605).

Talaromyces muroii Yaguchi et al., Mycoscience 35: 252. 1994. [MB362930]. — Type: CBM PF-1153. Ex-type: CBS 756.96 = PF 1153. Infragen. class: sect. Talaromyces. Reproduction: homothallic. ITS barcode: MN431394 (alternative markers: BenA = KJ865727; CaM = KJ885274; RPB2 = KX961276).

Talaromyces musae Houbraken et al., Persoonia 39: 341. 2017. [MB821051]. — Type: CBS H-23138. Ex-type: CBS 142504 = DTO 366-C5. Infragen. class: sect. Islandici. Reproduction: asexual. ITS barcode: MF072316 (alternative markers: BenA = MF093729; CaM = MF093728; RPB2 = MF093727).

Talaromyces mycothecae R.N. Barbosa et al., Antonie van Leeuwenhoek 111: 1902. 2018 [MB822215]. — Type: URM 90495. Ex-type: CBS 142494 = URM 7622. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: MF278326 (alternative markers: BenA = LT855561; CaM = LT855564; RPB2 = LT855567).

Talaromyces neofusisporus L. Wang, Sci. Rep. 6: 18622, 2. 2016. [MB811447]. — Type: HMAS 246033. Ex-type: CBS 139516 = DTO 410-D2 = AS 3.15415. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: KP765385 (alternative markers: BenA = KP765381; CaM = KP765383; RPB2 = MN969165).

Talaromyces neorugulosus A.J. Chen et al., Stud. Mycol. 84:139. 2016 [MB817394]. — Type: CBS H-22511. Ex-type: CBS 140623 = CGMCC3.18215 = DTO 318-A8. Infragen. class: sect. Islandici. Reproduction: asexual. ITS barcode: KU866659 (alternative markers: BenA = KU866846; CaM = KU866743; RPB2 = KU867003).

Talaromyces novojersensis Jurjević & S.W. Peterson, Mycologia 109: 544. 2017. [MB817999]. — Type: BPI-910146. Ex-type: NRRL 35858. Infragen. class: sect. Islandici. Reproduction: asexual. ITS barcode: KX657319 (alternative markers: BenA = KX657050; CaM = KX657151; RPB2 = KX657503).

Talaromyces oumae-annae Visagie et al., Stud. Mycol. 78: 130. 2014. [MB809187]. — Type: CBS H-21797. Ex-type: CBS 138208 = DTO 269-E8. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: KJ775720 (alternative markers: BenA = KJ775213; CaM = KJ775425; RPB2 = KX961281).

Talaromyces palmae (Samson et al.) Samson et al., Stud. Mycol. 71: 176. 2011. [MB560658]. Basionym: Penicillium palmae Samson, Stolk & Frisvad, Stud. Mycol 31: 135. 1989. [MB125456]. — Type: CBS 442.88. Ex-type: CBS 442.88 = IMI 343640. Infragen. class: sect. Subinflati. Reproduction: asexual. ITS barcode: JN899396 (alternative markers: BenA = HQ156947; CaM = KJ885291; RPB2 = KM023300).

Talaromyces panamensis (Samson et al.) Samson et al., Stud. Mycol. 71: 176. 2011. [MB560659]. Basionym: Penicillium panamense Samson, Stolk & Frisvad, Stud. Mycol. 31: 136. 1989. [MB125457]. — Type: CBS 128.89. Ex-type: CBS 128.89 = IMI 297546. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: JN899362 (alternative markers: BenA = HQ156948; CaM = KF741936; RPB2 = KM023284).

Talaromyces paucisporus (Yaguchi et al.) Samson & Houbraken, Stud. Mycol. 71: 176. 2011. [MB560684]. Basionym: Erythrogymnotheca paucispora Yaguchi et al., Mycoscience 35: 219. 1994. [MB362926]. — Type: CBM-PF 1150. Ex-type: PF 1150 = IFM 53616. Infragen. class: sect. Talaromyces. Reproduction: homothallic. ITS barcode: AB176603 (alternative markers: BenA = n.a.; CaM = n.a.; RPB2 = n.a.).

Talaromyces pernambucoensis R. Cruz et al., Persoonia 42: 467. 2019. [MB830189]. — Type: URM 93054. Ex-type: URM 6894. Infragen. class: sect. Trachyspermi. Reproduction: asexual. ITS barcode: LR535947 (alternative markers: BenA = LR535945; CaM = LR535946; RPB2 = LR535948).

Talaromyces piceus (Raper & Fennell) Samson et al., Stud. Mycol. 71: 176. 2011. [MB560661]. Basionym: Penicillium piceum Raper & Fennell, Mycologia 40: 533. 1948. [MB289103]. — Type: IMI 040038. Ex-type: CBS 361.48 = ATCC 10519 = IMI 040038 = NRRL 1051. Infragen. class: sect. Islandici. Reproduction: asexual. ITS barcode: KF984792 (alternative markers: BenA = KF984668; CaM = KF984680; RPB2 = KF984899).

Talaromyces pigmentosus R.N. Barbosa et al., Antonie van Leeuwenhoek 111: 1905. 2018 [MB822216]. — Type: URM 90496. Ex-type: CBS 142805 = URM 7624. Infragen. class: sect. Helici. Reproduction: asexual. ITS barcode: MF278330 (alternative markers: BenA = LT855562; CaM = LT855565; RPB2 = LT855568).

Talaromyces pinophilus (Hedgc.) Samson et al., Stud. Mycol. 71: 176. 2011. [MB560662]. Basionym: Penicillium pinophilum Hedgc., U.S.D.A. Bur. Ani. Ind. Bull. 118: 37. 1910. [MB211142]. — Type: IMI 114933. Ex-type: CBS 631.66 = ATCC 36839 = CECT 2809 = DSM 1944 = IAM 7013 =IMI 114933. Infragen. class: sect. Talaromyces. Reproduction: protoheterothallic, both mating types detected (López-Villavicencio et al. 2010). ITS barcode: JN899382 (alternative markers: BenA = JX091381; CaM = KF741964; RPB2 = KM023291).

Talaromyces pittii (Quintan.) Samson et al., Stud. Mycol. 71: 176. 2011. [MB560663]. Basionym: Penicillium pittii Quintan., Mycopathologia 91: 69. 1985. [MB105612]. — Type: CBS 139.84. Ex-type: CBS 139.84 = IMI 327871. Infragen. class: sect. Purpurei. Reproduction: asexual. ITS barcode: JN899325 (alternative markers: BenA = KJ865728; CaM = KJ885275; RPB2 = KM023297).

Talaromyces pratensis Jurjević & S.W. Peterson, Fungal Biol. 123: 756. 2019. [MB827831]. — Type: BPI 910710. Ex-type: NRRL 62170. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: MH793075 (alternative markers: BenA = MH792948; CaM = MH793012; RPB2 = MH793139).

Talaromyces primulinus (Pitt) Samson et al., Stud. Mycol. 71: 176. 2011. [MB560664]. Basionym: Penicillium primulinum Pitt, The Genus Penicillium: 455. 1980 [1979]. [MB116330]. — Type: IMI 040031. Ex-type: CBS 321.48 = ATCC 10438 = CBS 439.88 = FRR 1074 = IMI 040031 = MUCL 31321 = MUCL 31330 = NRRL 1074. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: JN899317 (alternative markers: BenA = JX494305; CaM = KF741954; RPB2 = KM023294).

Talaromyces proteolyticus (Kamyschko) Samson et al., Stud. Mycol. 71: 176. 2011. [MB560665]. Basionym: Penicillium proteolyticum Kamyschko, Nov. Sist. niz. Rast.: 228. 1961. [MB335760]. — Type: CBS 303.67. Ex-type: CBS 303.67 = ATCC 18326 = NRRL 3378. Infragen. class: sect. Bacillispori. Reproduction: asexual. ITS barcode: JN899387 (alternative markers: BenA = KJ865729; CaM = KJ885276; RPB2 = KM023301).

Talaromyces pseudofuniculosus Guevara-Suarez et al., Fungal Syst. Evol. 5: 69. 2020. [MB822090]. — Type: CBS H-23214. Ex-type: CBS 143041 = FMR 15307. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: LT899796 (alternative markers: BenA = LT898323; CaM = LT899778; RPB2 = LT899814).

Talaromyces pseudostromaticus (Hodges et al.) Samson et al., Stud. Mycol. 71: 176. 2011. [MB560666]. Basionym: Penicillium pseudostromaticum Hodges et al., Mycologia 62: 1106. 1970. [MB319292]. — Type: Warner 18 (NY). Ex-type: CBS 470.70 = ATCC 18919 = FRR 2039. Infragen. class: sect. Purpurei. Reproduction: asexual. ITS barcode: JN899371 (alternative markers: BenA = HQ156950; CaM = KJ885277; RPB2 = KM023298).

Talaromyces ptychoconidius [as “ptychoconidium”] Visagie & K. Jacobs, Persoonia 28: 18. 2012. [MB626028]. — Type: PREM 60041. Ex-type: DAOM 241017 = CV 2808 = DTO 180-E7. Infragen. class: sect. Purpurei. Reproduction: asexual. ITS barcode: FJ160266 (alternative markers: BenA = GU385733; CaM = JX140701; RPB2 = KM023278).

Talaromyces purgamentorum N. Yilmaz et al., Mycol. Prog. 15: 1054. 2016. [MB816233]. — Type: HUA 197222. Ex-type: CBS 113145 = IBT 23220 = DTO 056-E1. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: KX011504 (alternative markers: BenA = KX011487; CaM = KX011500; RPB2 = MN969189).

Talaromyces purpureogenus [as “purpurogenus”] (Stoll) Samson et al., Stud. Mycol. 71: 177. 2011. [MB585068]. Basionym: Penicillium purpureogenum Stoll, Beitr. Charakt. Penicillium-Arten: 32. 1904. [MB585067]. — Type: IMI 091926. Ex-type: CBS 286.36 = IMI 091926. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: JN899372 (alternative markers: BenA = JX315639; CaM = KF741947; RPB2 = JX315709).

Talaromyces purpureus (E. Müll. & Pacha-Aue) Stolk & Samson, Stud. Mycol. 2: 57. 1972. [MB324420]. Basionym: Arachniotus purpureus E. Müll. & Pacha-Aue, Nova Hedwigia 15: 552. 1968. [MB326429]. — Type: CBS H-7832 (isotype). Ex-type: CBS 475.71 = ATCC 24069 = ATCC 52513 = FRR 1731 = IMI 181546. Infragen. class: sect. Purpurei. Reproduction: homothallic. ITS barcode: JN899328 (alternative markers: BenA = GU385739; CaM = KJ885292; RPB2 = JN121522).

Talaromyces qii L. Wang, Sci. Rep. 6: 18622, 2. 2016. [MB811448]. — Type: HMAS 246032. Ex-type: CBS 139515 = DTO 410-D1 = AS 3.15414. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: KP765384 (alternative markers: BenA = KP765380; CaM = KP765382; RPB2 = MN969164).

Talaromyces rademirici (Quintan.) Samson et al., Stud. Mycol. 71: 177. 2011. [MB560668]. Basionym: Penicillium rademirici Quintan., Mycopathologia 91: 69. 1985. [MB105613]. — Type: CBS 140.84. Ex-type: CBS 140.84 = CECT 2771 = IMI 282406 = IMI 327870. Infragen. class: sect. Purpurei. Reproduction: asexual. ITS barcode: JN899386 (alternative markers: BenA = KJ865734; CaM = n.a.; RPB2 = KM023302).

Talaromyces radicus (A.D. Hocking & Whitelaw) Samson et al., Stud. Mycol. 71: 177. 2011. [MB560669]. Basionym: Penicillium radicum A.D. Hocking & Whitelaw, Mycol. Res. 102: 802. 1998. [MB445182]. — Type: DAR 72374. Ex-type: CBS 100489 = FRR 4718. Infragen. class: sect. Islandici. Reproduction: asexual. ITS barcode: KF984878 (alternative markers: BenA = KF984599; CaM = KF984773; RPB2 = KF985013).

Talaromyces ramulosus (Visagie & K. Jacobs) Samson et al., Stud. Mycol. 71: 177. 2011. [MB560670]. Basionym: Penicillium ramulosum Visagie & K. Jacobs, Mycologia 101: 890. 2008. [MB512023]. — Type: PREM 59947. Ex-type: DAOM 241660 = CV 2837 = DTO 184-B8. Infragen. class: sect. Purpurei. Reproduction: asexual. ITS barcode: EU795706 (alternative markers: BenA = FJ753290; CaM = JX140711; RPB2 = KM023281).

Talaromyces rapidus Guevara-Suarez et al., Mycoses 60: 658. 2017. [MB820464]. — Type: CBS H-23002. Ex-type: UTHSC DI16-148 = CBS 142382 = FMR 14293. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: LT558970 (alternative markers: BenA = LT559087; CaM = LT795600; RPB2 = LT795601).

Talaromyces resedanus (McLennan & Ducker) A.J. Chen et al., under review. [MB811695]. Basionym: Penicillium resedanum McLennan & Ducker, Australian J. Bot: 360. 1954. [MB302422]. — Type: IMI 62877. Ex-type: CBS 181.71 = DTO 376-A7 = ATCC 22356 = FRR 578 = IMI 062877 = NRRL 578. Infragen. class: sect. Subinflati. Reproduction: asexual. ITS barcode: MN431413 (alternative markers: BenA = MN969436; CaM = MN969355; RPB2 = MN969214).

Talaromyces resinae (Z.T. Qi & H.Z. Kong) Houbraken & X.C. Wang, this study. 2020. [MB833989]. Basionym: Penicillium resinae Z.T. Qi & H.Z. Kong, Acta Mycol. Sin.: 103. 1982. [MB110236]. — Type: HMAS 42799. Ex-type: AS 3.4387 = CBS 324.83 = DTO 027-G5. Infragen. class: sect. Trachyspermi. Reproduction: asexual. ITS barcode: MT079858 (alternative markers: BenA = MN969442; CaM = MT066184; RPB2 = MN969221).

Talaromyces reverso-olivaceus A.J. Chen et al., Stud. Mycol. 84:141. 2016 [MB817391]. — Type: CBS H-22512. Ex-type: CBS 140672 = CGMCC 3.18195 = DTO 317-C3. Infragen. class: sect. Helici. Reproduction: asexual. ITS barcode: KU866646 (alternative markers: BenA = KU866834; CaM = KU866730; RPB2 = KU866990).

Talaromyces ricevillensis Jurjević & S.W. Peterson, Mycologia 109: 547. 2017. [MB818003]. — Type: BPI-910151. Ex-type: NRRL 62296. Infragen. class: sect. Islandici. Reproduction: asexual. ITS barcode: KX657343 (alternative markers: BenA = KX657056; CaM = KX657249; RPB2 = KX657582).

Talaromyces rogersiae Jurjević & S.W. Peterson, Mycologia 109: 550. 2017. [MB818005]. — Type: BPI-910149. Ex-type: NRRL 62223. Infragen. class: sect. Islandici. Reproduction: asexual. ITS barcode: KX657332 (alternative markers: BenA = KX657125; CaM = KF196891; RPB2 = KX657581).

Talaromyces rotundus (Raper & Fennell) C.R. Benj., Mycologia 47: 683. 1955. [MB306719]. Basionym: Penicillium rotundum Raper & Fennell, Mycologia 40: 518. 1948. [MB289106]. — Type: IMI 040589. Ex-type: CBS 369.48 = ATCC 10493 = IMI 040589 = NRRL 2107. Infragen. class: sect. Islandici. Reproduction: homothallic. ITS barcode: JN899353 (alternative markers: BenA = KJ865730; CaM = KJ885278; RPB2 = KM023275).

Talaromyces ruber (Stoll) N. Yilmaz et al., Persoonia 29: 48. 2012. [MB801360]. Basionym: Penicillium rubrum Stoll, Beitr. Charakt. Penicillium-Arten: 35. 1904. [MB205727]. — Type: CBS H-21052. Ex-type: CBS 132704 = DTO 193-H6 = IBT 10703 = CBS 113137. Infragen. class: sect. Talaromyces. Reproduction: protoheterothallic, MAT1-1-1 detected (López-Villavicencio et al. 2010). ITS barcode: JX315662 (alternative markers: BenA = JX315629; CaM = KF741938; RPB2 = JX315700).

Talaromyces rubicundus (J.H. Mill. et al.) Samson et al., Stud. Mycol. 71: 177. 2011. [MB560671]. Basionym: Penicillium rubicundum J.H. Mill., Giddens & A.A. Foster, Mycologia 49: 797. 1957. [MB302423]. — Type: No. 2531 (A.A. Foster). Ex-type: CBS 342.59 = ATCC 13217 = IMI 099723 = NRRL 3400. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: JN899384 (alternative markers: BenA = JX494309; CaM = KF741956; RPB2 = KM023296).

Talaromyces rugulosus (Thom) Samson et al., Stud. Mycol. 71: 177. 2011. [MB560672]. Basionym: Penicillium rugulosum Thom, U.S.D.A. Bur. Ani. Ind. Bull. 118: 60. 1910. [MB210907]. — Type: IMI 040041. Ex-type: CBS 371.48 = ATCC 10128 = IMI 040041 = MUCL 31201 = NRRL 1045. Infragen. class: sect. Islandici. Reproduction: asexual. ITS barcode: KF984834 (alternative markers: BenA = KF984575; CaM = KF984702; RPB2 = KF984925).

Talaromyces ryukyuensis (S. Ueda & Udagawa) Arx, Persoonia 13: 282. 1987. [MB132096]. Basionym: Sagenoma ryukyuense S. Ueda & Udagawa, Mycotaxon 20: 499. 1984. [MB124389]. — Type: unknown. Ex-type: NHL 2917 = DTO 176-I6. Infragen. class: sect. Helici. Reproduction: homothallic. ITS barcode: AB176628 (alternative markers: BenA = n.a.; CaM = n.a.; RPB2 = n.a.).

Talaromyces sayulitensis Visagie et al., Stud. Mycol. 78: 132. 2014. [MB809188]. — Type: CBS H-21798. Ex-type: CBS 138204 = DTO 245-H1. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: KJ775713 (alternative markers: BenA = KJ775206; CaM = KJ775422; RPB2 = MN969146).

Talaromyces scorteus (Nakazawa et al.) S.W. Peterson & Jurjević, PLoS ONE 8: e78084, 8. 2013. [MB804734]. Basionym: Penicillium scorteum Nakazawa et al., J. Agric. Chem. Soc. Japan: 103. 1934. [MB492647]. — Type: unknown. Ex-type: CBS 340.34 = NRRL 1129 = FRR 1129. Infragen. class: sect. Islandici. Reproduction: asexual. ITS barcode: KF984892 (alternative markers: BenA = KF984565; CaM = KF984684; RPB2 = KF984916).

Talaromyces siamensis (Manoch & C. Ramírez) Samson et al., Stud. Mycol. 71: 177. 2011. [MB560674]. Basionym: Penicillium siamense Manoch & C. Ramírez, Mycopathologia 101: 32. 1988. [MB133414]. — Type: CBS 475.88. Ex-type: CBS 475.88 = IMI 323204. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: JN899385 (alternative markers: BenA = JX091379; CaM = KF741960; RPB2 = KM023279).

Talaromyces siglerae S.W. Peterson & Jurjević, Mycologia 109: 540. 2017. [MB818002]. — Type: BPI-910143. Ex-type: NRRL 28620. Infragen. class: sect. Islandici. Reproduction: asexual. ITS barcode: KX657351 (alternative markers: BenA = KX657135; CaM = KX657236; RPB2 = KX657497).

Talaromyces soli Jurjević & S.W. Peterson, Fungal Biol. 123: 757. 2019. [MB827832]. — Type: BPI 910709. Ex-type: NRRL 62165. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: MH793074 (alternative markers: BenA = MH792947; CaM = MH793011; RPB2 = MH793138).

Talaromyces solicola Visagie & K. Jacobs, Persoonia 28: 20. 2012. [MB564328]. — Type: PREM 60037. Ex-type: DAOM 241015 = CV 2800 = DTO 180-D4. Infragen. class: sect. Trachyspermi. Reproduction: asexual. ITS barcode: FJ160264 (alternative markers: BenA = GU385731; CaM = KJ885279; RPB2 = KM023295).

Talaromyces speluncarum Rodr.-Andr. et al., Microorganisms 2020, 8, 12: 10. 2020 [MB830606]. — Type: CBS H-23372. Ex-type: CBS 143844 = FMR 16671. Infragen. class: sect. Trachyspermi. Reproduction: asexual. ITS barcode: LT985890 (alternative markers: BenA = LT985901; CaM = LT985906; RPB2 = LT985911).

Talaromyces stellenboschensis [as “stellenboschiensis”] Visagie & K. Jacobs, Mycoscience 56:497. 2015. [MB834764]. — Type: CBS H-21601. Ex-type: CBS 135665 = DTO 181-A2 = DAOM 241021 = IBT 32631. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: JX091471 (alternative markers: BenA = JX091605; CaM = JX140683; RPB2 = MN969157).

Talaromyces stipitatus (Thom) C.R. Benj., Mycologia 47: 684. 1955. [MB306722]. Basionym: Penicillium stipitatum Thom, Mycologia 27: 138. 1935. [MB119544]. — Type: CBS H-7835. Ex-type: CBS 375.48 = ATCC 10500 = NRRL 1006 = IMI 39805. Infragen. class: sect. Talaromyces. Reproduction: homothallic. ITS barcode: JN899348 (alternative markers: BenA = KM111288; CaM = KF741957; RPB2 = KM023280).

Talaromyces stollii N. Yilmaz et al., Persoonia 29: 52. 2012. [MB801359]. — Type: CBS H-21053. Ex-type: CBS 408.93. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: JX315674 (alternative markers: BenA = JX315633; CaM = JX315646; RPB2 = JX315712).

Talaromyces striatoconidius (R.F. Castañeda & W. Gams) Houbraken et al., this study. 2020. [MB832566]. Replaced synonym: Phialomyces striatus R.F. Castañeda & W. Gams, Mycotaxon 42: 239. 1991. [MB358862]. — Type: INIFAT C89/170. Ex-type: CBS 550.89 = DTO 418-H4. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: MN431418 (alternative markers: BenA = MN969441; CaM = MN969360; RPB2 = MT156347).

Talaromyces subaurantiacus Visagie et al., Persoonia 36: 52. 2016. [MB810902]. — Type: CBS H-21630. Ex-type: CBS 137383 = DTO 181-I2 = DAOM 241020 = IBT 32383. Infragen. class: sect. Islandici. Reproduction: asexual. ITS barcode: JX091475 (alternative markers: BenA = JX091609; CaM = JX140728; RPB2 = KF984960).

Talaromyces subericola Rodr.-Andr. et al., Microorganisms 2020, 8, 12: 12. 2020 [MB830607]. — Type: CBS H-23366. Ex-type: CBS 144322 = FMR 15656. Infragen. class: sect. Trachyspermi. Reproduction: asexual. ITS barcode: LT985888 (alternative markers: BenA = LT985899; CaM = LT985904; RPB2 = LT985909).

Talaromyces subinflatus Yaguchi & Udagawa, Trans. Mycol. Soc. Japan 34: 249. 1993. [MB361184]. — Type: CBM PF-1113. Ex-type: CBS 652.95 = DTO 105-D5 = IBT 17520. Infragen. class: sect. Subinflati. Reproduction: homothallic. ITS barcode: JN899397 (alternative markers: BenA = MK450890; CaM = KJ885280; RPB2 = KM023308).

Talaromyces subtropicalis Jurjević & S.W. Peterson, Mycologia 109: 548. 2017. [MB818004]. — Type: BPI-910147. Ex-type: NRRL 58084. Infragen. class: sect. Islandici. Reproduction: asexual. ITS barcode: KX657337 (alternative markers: BenA = KX657060; CaM = KX657250; RPB2 = KX657531).

Talaromyces systylus S.M. Romero et al., Nova Hedwigia 102: 244. 2016. [MB812381]. — Type: BAFC 52367. Ex-type: BAFCcult 3419. Infragen. class: sect. Trachyspermi. Reproduction: asexual. ITS barcode: KP026917 (alternative markers: BenA = KR233838; CaM = KR233837; RPB2 = n.a.).

Talaromyces tabacinus Jurjević et al., Persoonia 40: 323. 2018. [MB823318]. — Type: BPI 910533. Ex-type: NRRL 66727 = EMSL 2174. Infragen. class: sect. Helici. Reproduction: asexual. ITS barcode: MG182613 (alternative markers: BenA = MG182627; CaM = MG182606; RPB2 = MG182620).

Talaromyces tardifaciens Udagawa, Mycotaxon 48: 150. 1993. [MB360478]. — Type: CBM SUM 3017. Ex-type: CBS 250.94. Infragen. class: sect. Islandici. Reproduction: homothallic. ITS barcode: JN899361 (alternative markers: BenA = KF984560; CaM = KF984682; RPB2 = KF984908).

Talaromyces thailandensis Manoch et al., Mycoscience 54: 339. 2013. [MB801737]. — Type: CBS H-21075. Ex-type: CBS 133147 = KUFC 3399. Infragen. class: sect. Talaromyces. Reproduction: homothallic. ITS barcode: JX898041 (alternative markers: BenA = JX494294; CaM = KF741940; RPB2 = KM023307).

Talaromyces tiftonensis Jurjević & S.W. Peterson, Mycologia 109: 552. 2017. [MB818008]. — Type: BPI-910150. Ex-type: NRRL 62264. Infragen. class: sect. Islandici. Reproduction: asexual. ITS barcode: KX657353 (alternative markers: BenA = KX657129; CaM = KX657163; RPB2 = KX657602).

Talaromyces trachyspermus (Shear) Stolk & Samson, Stud. Mycol. 2: 32. 1972. [MB324421]. Basionym: Arachniotus trachyspermus Shear, Science New York 16: 138. 1902. [MB164396]. — Type: IMI 040043. Ex-type: CBS 373.48 = ATCC 10497 = IMI 040043 = NRRL 1028. Infragen. class: sect. Trachyspermi. Reproduction: homothallic. ITS barcode: JN899354 (alternative markers: BenA = KF114803; CaM = KJ885281; RPB2 = JF417432).

Talaromyces tratensis Manoch et al., Mycoscience 54: 337. 2013. [MB801738]. — Type: CBS H-21074. Ex-type: CBS 133146 = KUFC 3383. Infragen. class: sect. Islandici. Reproduction: homothallic. ITS barcode: KF984891 (alternative markers: BenA = KF984559; CaM = KF984690; RPB2 = KF984911).

Talaromyces tumuli Jurjević & S.W. Peterson, Fungal Biol. 123: 758. 2019. [MB827833]. — Type: BPI 910713. Ex-type: NRRL 62151. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: MH793071 (alternative markers: BenA = MH792944; CaM = MH793008; RPB2 = MH793135).

Talaromyces tzapotlensis Jurjević & S.W. Peterson, Mycologia 109: 553. 2017. [MB818009]. — Type: BPI-910145. Ex-type: NRRL 35203. Infragen. class: sect. Subinflati. Reproduction: asexual. ITS barcode: KX946902 (alternative markers: BenA = KX946884; CaM = KX946893; RPB2 = KX946922).

Talaromyces ucrainicus (Panas.) Udagawa, Trans. Mycol. Soc. Japan 7: 94. 1966. [MB449587]. Basionym: Penicillium ucrainicum Panas., Mycologia 56: 59. 1964. [MB335773]. — Type: unknown. Ex-type: CBS 162.67 = ATCC 22344 = FRR 3462 = NHL 6086. Infragen. class: sect. Trachyspermi. Reproduction: homothallic. ITS barcode: JN899394 (alternative markers: BenA = KF114771; CaM = KJ885282; RPB2 = KM023289).

Talaromyces udagawae Stolk & Samson, Stud. Mycol. 2: 36. 1972. [MB324424]. — Type: CBS H-7841. Ex-type: CBS 579.72 = FRR 1727 = IMI 197482. Infragen. class: sect. Trachyspermi. Reproduction: homothallic. ITS barcode: JN899350 (alternative markers: BenA = KF114796; CaM = KX961260; RPB2 = MN969148).

Talaromyces unicus Tzean et al., Mycologia 84: 739. 1992. [MB360172]. — Type: PPH 16 Nat. Taiwan Univ. Ex-type: CBS 100535 = CCRC 32703 = IBT 18385. Infragen. class: sect. Bacillispori. Reproduction: homothallic. ITS barcode: JN899336 (alternative markers: BenA = KJ865735; CaM = KJ885283; RPB2 = MN969150).

Talaromyces varians (G. Sm.) Samson et al., Stud. Mycol. 71: 177. 2011. [MB560677]. Basionym: Penicillium varians G. Sm., Trans. Brit. Mycol. Soc. 18: 89. 1933. [MB289112]. — Type: IMI 040586. Ex-type: CBS 386.48 = ATCC 10509 = IMI 040586 = NRRL 2096. Infragen. class: sect. Helici. Reproduction: asexual. ITS barcode: JN899368 (alternative markers: BenA = KJ865731; CaM = KJ885284; RPB2 = KM023274).

Talaromyces veerkampii Visagie et al., Mycoscience 56: 497. 2015. [MB808233]. — Type: CBS H-21600. Ex-type: CBS 500.78; DTO 258-I8; IBT 14845; IBT 32648. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: KF741984 (alternative markers: BenA = KF741918; CaM = KF741961; RPB2 = KX961279).

Talaromyces verruculosus (Peyronel) Samson et al., Stud. Mycol. 71: 177. 2011. [MB560678]. Basionym: Penicillium verruculosum Peyronel, I germi astmosferici dei fungi con micelio, Diss. (Padova): 22. 1913. [MB166576]. — Type: IMI 040039. Ex-type: DTO 264-I8 = CBS 136671 = ATCC 10513 = DSM 2263 = IMI 040039 = NRRL 1050. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: KF741994 (alternative markers: BenA = KF741928; CaM = KF741944; RPB2 = KM023306).

Talaromyces versatilis Bridge & Buddie, Index Fungorum 26: 1. 2013. [MB550190]. — Type: IMI 134755. Ex-type: DTO 326-B7 = IMI 134755. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: MN431395 (alternative markers: BenA = MN969412; CaM = MN969319; RPB2 = MN969161).

Talaromyces viridis (Stolk & G.F. Orr) Arx, Persoonia 13: 2821. 1987. [MB132097]. Basionym: Sagenoma viride Stolk & G.F. Orr, Mycologia 66: 677. 1974. [MB323033]. — Type: CBS H-7732 (isotype), CBS H-7733 (isotype), CBS H-7734 (isotype). Ex-type: CBS 114.72 = ATCC 22467 = NRRL 5575. Infragen. class: sect. Talaromyces. Reproduction: homothallic. ITS barcode: AF285782 (alternative markers: BenA = JX494310; CaM = KF741935; RPB2 = JN121430).

Talaromyces viridulus Samson et al., Stud. Mycol. 71: 177. 2011. [MB560679]. — Type: FRR 1863. Ex-type: CBS 252.87 = FRR 1863 = IMI 288716. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: JN899314 (alternative markers: BenA = JX091385; CaM = KF741943; RPB2 = JF417422).

Talaromyces wortmannii (Klöcker) C.R. Benj., Mycologia 47: 683. 1955. [MB344294]. Basionym: Penicillium wortmannii Klöcker, Compt. Rend. Trav. Lab. Carlsberg: serie Physiologique 6: 100. 1903. [MB119648]. — Type: IMI 040047. Ex-type: CBS 391.48 = ATCC 10517 = IMI 040047 = NRRL 1017. Infragen. class: sect. Islandici. Reproduction: homothallic. ITS barcode: KF984829 (alternative markers: BenA = KF984648; CaM = KF984756; RPB2 = KF984977).

Talaromyces xishaensis X.C. Wang et al., Phytotaxa 267: 193. 2016. [MB815751]. — Type: HMAS 248732. Ex-type: CGMCC 3.17995. Infragen. class: sect. Talaromyces. Reproduction: asexual. ITS barcode: KU644580 (alternative markers: BenA = KU644581; CaM = KU644582; RPB2 = n.a.).

Talaromyces yelensis Visagie et al., Stud. Mycol. 78: 134. 2014. [MB809189]. — Type: CBS H-21799. Ex-type: CBS 138209 = DTO 268-E5. Infragen. class: sect. Islandici. Reproduction: asexual. ITS barcode: KJ775717 (alternative markers: BenA = KJ775210; CaM = KP119161; RPB2 = KP119163).



Thermoascus

Thermoascus aegyptiacus S. Ueda & Udagawa, Trans. Mycol. Soc. Japan 24: 135. 1983. [MB108584]. — Type: NHL 2914. Ex-type: DTO 424-H2 = CBS 146342 = NHL 2914 = IFM 61569. Reproduction: homothallic. ITS barcode: MT068197 (alternative markers: BenA = MN969411; CaM = MN969318; RPB2 = n.a.).

Thermoascus aurantiacus Miehe, Die Selbsterhitzung des Heus, Jena: 70. 1907. [MB122028]. — Type: UC M206516. Ex-type: n.a. Reproduction: homothallic. ITS barcode: EU021617 (alternative markers: BenA = EU021667; CaM = n.a.; RPB2 = EU021657).

Thermoascus crustaceus (Apinis & Chesters) Stolk, Antonie van Leeuwenhoek 31: 272. 1965. [MB340045]. Basionym: Dactylomyces crustaceus Apinis & Chesters, Trans. Brit. Mycol. Soc. 47: 428. 1964. [MB329643]. — Type: IMI 102470. Ex-type: IMI 102470 = BDUN 378. Reproduction: homothallic. ITS barcode: MN431405 (alternative markers: BenA = MN969423; CaM = MN969344; RPB2 = MN969205).

Thermoascus taitungiacus K.Y. Chen & Z.C. Chen, Mycotaxon 60: 226. 1996. [MB436720]. — Type: TAl-Mycology K-Y Chen 8709-2. Ex-type: n.a. Reproduction: homothallic. ITS barcode: n.a. (alternative markers: BenA = n.a.; CaM = n.a.; RPB2 = n.a.).

Thermoascus thermophilus (Sopp) Arx, The genera of fungi sporulating in pure culture: 94. 1974. [MB324541]. Basionym: Dactylomyces thermophilus Sopp, Skr. VidenskSelsk. Christiania, Kl. I, Math.-Natur. (no. 11): 35. 1912. [MB224271]. — Type: CBS 528.71. Ex-type: CBS 528.71 = BDUN 394 = IMI 123298 = NRRL 5208. Reproduction: homothallic. ITS barcode: MH860254 (alternative markers: BenA = MN969424; CaM = MN969345; RPB2 = JF417442).

Thermoascus verrucosus (Samson & Tansey) Houbraken et al., this study. 2020. [MB832567]. Basionym: Byssochlamys verrucosa Samson & Tansey, Trans. Brit. Mycol. Soc. 65: 512. 1975. [MB310015]. — Type: CBS 605.74. Ex-type: CBS 605.74 = ATCC 34163. Reproduction: homothallic. ITS barcode: DQ073329 (alternative markers: BenA = MN969425; CaM = MN969346; RPB2 = JN121540).

Thermoascus yaguchii (Yaguchi et al.) Houbraken et al., this study. 2020. [MB833988]. Replaced synonym: Thermoascus crustaceus var. verrucosus Yaguchi et al., Mycoscience 36: 151. 1995. [MB414278]. — Type: PF-1160. Ex-type: DTO 424-H3 = CBS 146343 = PF-1160 = IFM 66000. Reproduction: homothallic. ITS barcode: MT068196 (alternative markers: BenA = MT070995; CaM = MN969362; RPB2 = n.a.).



Thermomyces

Thermomyces dupontii (Griffon & Maubl.) Houbraken & Samson, Adv. Appl. Microbiol. 86: 218. 2014. [MB805186]. Basionym: Penicillium dupontii Griffon & Maubl., Bull. Soc. Mycol. France 27: 73. 1911. [MB120357]. — Type: CBS 236.58. Ex-type: CBS 236.58 = ATCC 10518 = ATCC 16461 = ATCC 52514 = IFO 31798 = IMI 048593 = NRRL 2155. Reproduction: homothallic (fide Emerson, in Raper & Thom 1949). ITS barcode: MN431410 (alternative markers: BenA = MN969432; CaM = MT066185; RPB2 = JF417420).

Thermomyces lanuginosus Tsikl., Ann. Inst. Pasteur 13: 500. 1899. [MB239786]. — Type: IMI 84400 (Pugh et al.). Ex-type: CBS 632.91 = DTO 024-B3 = ATCC 16455 = DAOM 232588 (representative). Reproduction: asexual. ITS barcode: MN431411 (alternative markers: BenA = MN969433; CaM = MN969353; RPB2 = MN969209).



Trichocoma

Trichocoma paradoxa Jungh., Praemissa in floram cryptogamicam Javae insulae: 9. 1838. [MB161024]. — Type: "Trichocoma paradoxa, Junghuhn (BO)". Ex-type: CBS 247.57, CBS 103.73, CBS 788.83 (representative strains). Reproduction: homothallic. ITS barcode: MH860643 (alternative markers: BenA = JF417469; CaM = JF417506; RPB2 = JN121417).



Warcupiella

Warcupiella spinulosa (Warcup) Subram., Curr. Sci. 41: 757. 1972. [MB325542]. Basionym: Aspergillus spinulosus Warcup, Gen. Aspergillus: 204. 1965. [MB326657]. — Type: IMI 75885. Ex-type: CBS 512.65 = DTO 021-H3 = ATCC 16919 = IFO 31800 = IMI 075885 = IMI 075885ii = IMI 075885iii = IMI 075885iv = NRRL 4376 = WB 4376. Reproduction: homothallic. ITS barcode: EF669706 (alternative markers: BenA = MN969410; CaM = EF669698; RPB2 = EF669666).



Xerochrysium

Xerochrysium dermatitidis (A. Agostini) Pitt, IMA Fungus 4: 237. 2013. [MB807005]. Basionym: Glenosporella dermatitidis A. Agostini, Atti Ist. Bot. R. Univ. Pavia, 3 Sér. 4: 98. 1931. [MB536220]. — Type: CBS 132.31 (Pitt et al. 2013). Ex-type: DTO 375-C6 = IMI 96729 = UAMH 802 = FRR 2376. Reproduction: asexual. ITS barcode: KY635853 (alternative markers: BenA = MT070993; CaM = MT066186; RPB2 = JN121443).

Xerochrysium xerophilum (Pitt) Pitt, IMA Fungus 4: 238. 2013. [MB807006]. Basionym: Chrysosporium xerophilum Pitt, Trans. Brit. Mycol. Soc. 49: 468. 1966. [MB328223]. — Type: UAMH 2368. Ex-type: CBS 153.67 = FRR 503 = ATCC 18053 = IMI 126287. Reproduction: asexual. ITS barcode: GU733339 (alternative markers: BenA = n.a.; CaM = n.a.; RPB2 = n.a.).



Xeromyces

Xeromyces bisporus L.R. Fraser, Proc. Linn. Soc. New South Wales 78: 245. 1953. [MB307870]. — Type: unknown. Ex-type: CBS 236.71 = IMI 063718 = VKM F-1978 = FRR 525. Reproduction: homothallic. ITS barcode: KY635854 (alternative markers: BenA = KY709179; CaM = Genome sequence; CCCX01000040.1:6800–8000; RPB2 = JN121466).

General discussion

Infrageneric classification

Dierckx (1901) proposed the first infrageneric classification of Penicillium. Various other monographs used an infrageneric classification to sort Aspergillus and Penicillium species in groups. The use of subgenera and sections are therefore well-established for these genera; however, a series classification based on sequence data was lacking. This was considered problematic as both Aspergillus and Penicillium have grown significantly in species numbers the past few years. To make working with these genera and sections easier, various taxonomic studies often refer to a “clade” or a “species complex”. Here we show that series are often the equivalents of these “clades” and “species complexes”. The use of a formal series classification makes working with large speciose genera easier and stabilises its taxonomy. As mentioned in the introduction, the allocation of a species to a series can be highly predictive in what functional characters the species might have. On the other hand, these functional characters are not known in various series (e.g., in Aspergillus sect. Fumigati) and future studies might reveal those. Furthermore, the application of a series classification might be useful when using a phenotype-based identification. Phenotypic identification of strains on species level can be difficult and identification on series rank can be a suitable alternative: a series name can be used for a phylogenetically defined complex of species that are morphological highly similar. An example is ser. Versicolores in Aspergillus, which includes commonly occurring species that are morphologically highly similar and usually have a similar impact or function. Species in Penicillium and Aspergillus series are in many cases phenotypically recognizable, for example by colony obverse and reverse colours, colony texture (for example by image analysis; (Dörge et al. 2000), ecophysiological features, nutritional features (Ahmad & Malloch 1999), growth on alternative nitrogen sources and enzyme profiles (Barrett et al. 2020). Until now, profiles of CAZymes appear to be very promising to be used for classification of species into sections, as these are congruent with a phylogenetic cladification (Barrett et al. 2020); it is likely that these methods will be applicable on series as well. While most series are phylogenetically recognizable, there were some examples of a more provisional designation of series (e.g., series in Aspergillus sect. Fumigati; series Claviformia, Glandicolarum and Longicatenata in Penicillium). Genome sequence data will help circumscribing such series more accurately (Vesth et al., 2018, Kjærbølling et al., 2020), preferably using a comprehensive phenotypic characterization on top of the phylogenetic characterization.

It has been an experience that species in filamentous fungi are clear cut with no intergrading strains. Only a few hybrids have ever been reported in fungal taxonomy. Of course, individual characters may be overlapping, such as the size, form and ornamentation of conidia or growth and conidiation as related to temperature, but a large number of characters are non-overlapping. This is especially true concerning small molecule extrolites characters (Frisvad et al. 2008). Species are thus stable for many years (punctuated equilibria), and may then speciate under certain circumstances (Gould and Eldredge, 1977, Gould and Eldredge, 1993, Zander, 2010). Thus, selection can not only work at the individual level but also at the species level (Gould 1998). However, there is one problem with the use of clear non-overlapping characters: these characters are not necessarily present in all isolates of a species or all species in a series, section, subgenus or genus. This phenomenon may be caused by horizontal transfer of genes or gene clusters, epigenetical factors or mutation-based loss of certain genes or cluster. For this reason, the concept of polythetic classes is very valuable, i.e. that any individual character is present in most, but not all members of the class and that most members of the class also possess any individual character. Polythetic classes can be found by a character analysis (cluster analysis) of several operational taxonomic units (OTUs) using several measured features, but not by a cladistic analysis. Thus, while cladistic analysis appear to be optimal for DNA sequence data, cluster analysis appears to be most effective for phenotypic characters.

In some cases, small molecule extrolites (secondary metabolites) support the species relationship in a series, in other cases this is not supported, at least not with currently available data. Genome sequencing may help in determining if cryptic gene clusters for secondary metabolites will support the species series as proposed by household gene sequence data, apart from the extrolites that are actually produced on agar media. For example, citreoviridin production is present among the two species in Penicillium series Ochrosalmonea: while one species is slow-growing and produces no synnemata (P. ochrosalmoneum), the other species grows fast and produces synnemata (P. isariiforme). Citreoviridin is also produced by species in other series, and therefore ideally several secondary metabolite families need to be characterised for each series. More data on physiology such as water activity relations, temperature, pH and exoenzyme profiles may help to circumscribe the series phenotypically in a more accurate way and help discriminate between series. Examples of series that are both phylogenetically and phenotypically consistent are series Viridicata and Roquefortorum, and in sect. Sclerotiorum, series Adametziorum, Herqueorum and Sclerotiorum where each have unique extrolites typical of the series (sclerotiorins, herqueinones and dithiodiketopiperazines, respectively).

The fundamental species level

The species is the fundamental unit of taxonomy of fungi (Kirk et al. 2008) reflecting both evolution and ecology (Rollo, 1994, Sneath, 1995, Westoby et al., 1995, West-Eberhard, 2005). For this reason, as many unweighted characters as possible should be used in classification and for discovering species according to Sneath and Sokal, 1962, Sneath and Sokal, 1973 and Sneath (1995). This taxometric concept has been used for many years, even though the weighting of the characters has been discussed, like many characters, for example, chemotaxonomic characters based on the primary metabolism, that is shared with most other species and are constantly and quantitatively in flux, are hardly useful at all in a numerical taxonomy. For this reason it was proposed to use a weighted omnispective (Blackwelder 1964), polyphasic (Vandamme et al. 1996) or consilient (as consolidated) (Quaedvlieg et al. 2014) approach in discovering species based on different kinds of distinctive characters, especially morphological, nutritional, physiological, chemical and molecular (sequence-based) features. Following Hennig (1966), many scientists have emphasised synapomorphic features only, but using this approach in a strict sense will result in many important features not being considered in taxonomy (Sneath 1995).

For many years sequencing of household and other genes has been used solely to develop a cladification of isolates or species using the DNA sequence-based phylogenetic species recognition approach based on concordance of multiple gene genealogies (Taylor et al. 2000), but this approach does not take into account anagenesis (Mayr, 1981, Mayr, 1995), the functional phenotype (Rollo 1994) or epigenetics (Zander 2010). Thus, a polyphasic approach to taxonomy using symplesiomorphies, synapomorphies, and autapomorphies in conjunction with quantitative ecophysiological characters and sequence-based analyses appears to be an effective approach to discover species. The removal of autapomorphies in cladistic analyses has been criticised by Yeates (1992) and those autapomorphies are clear species-specific features. The autapomorphies may also help explain anagenetic evolution. Recently genome sequencing has added a lot of potentially new features, provided these features can be annotated, but this genome-based approach does not directly take into account epigenetics and phenotypic evolution (Rollo, 1994, Schlichting and Pigliucci, 1998, West-Eberhard, 2003, West-Eberhard, 2005). However, genomics, in conjunction with transcriptomics and exometabolomics, will undoubtedly be extremely valuable in taxonomy and cladonomy (Uilenberg et al., 2004, Zhi et al., 2012, Thompson et al., 2015). The revision of the species, series and sections in Aspergillus and Penicillium was guided by sequences of several primary metabolism “house-hold” genes. The results are often strongly supported by phenotypic features, showing that a combination of phenomics and genomics (a polyphasic approach) is needed for a predictive cladification and classification. Often series suggested by early phenotypic studies were confirmed by phylogenetic data, but in other cases, surprising relationships were encountered that were not expected from pure morphological data (for example, see Frisvad & Samson 2004b). It is crucial to take all important phenotypic features into account, as it is rarely the case that one type of phenotypic features can be used for classification that is in agreement with a sequence-based cladification. Until now the most predictive classification was based on profiles of function / family annotated CAZymes (Barrett et al. 2020), but in-depth comparative studies on the genomes and phenomes of Aspergillus, Penicillium and Talaromyces species will help suggesting a highly predictive taxonomy and phylogeny in the future.

Dedication and acknowledgments

This work is dedicated to Keith A. Seifert for his invaluable contributions to the taxonomy of the Eurotiales on the occasion of his retirement from Ottawa Research and Development Centre, Agriculture and Agri-Food Canada (Biodiversity: Mycology and Microbiology). Jens Frisvad thanks Agilent for a Thought leader award from Agilent (# 2871) and support from the Danish National Research Foundation for the center of excellence CeMiSt (DNRF137). We are grateful to David Malloch for his valuable suggestions to improve the manuscript and together with Lothar Kriegsteiner for making specimens of Dichlaena lentisci available. This work was partially supported by the Hungarian Research Fund OTKA K115690 via Sándor Kocsubé. We thank Takashi Yaguchi and the CBS culture collection for supplying cultures for our study and Shaun Pennycook for his nomenclatural advice. Cony Decock is acknowledged for supplying literature of P. Biourge.

Appendix A. Supplementary data

The following are the Supplementary data to this article:

Supplementary Fig. S1

Phylogenetic relationship within the Eurotiales based on a combined ML phylogeny using nine loci (RPB1, RPB2, Cct8, Tsr1, CaM, BenA, SSU, LSU, ITS) and a partition containing a binary matrix of indels present in the Tsr1, CaM, BenA and ITS dataset. Bootstrap values are above or below branches. The phylogram is based on 263 species belonging to the order Eurotiales and 16 species from the order Onygenales (used an outgroup).

Supplementary Fig. S2

Combined phylogeny for BenA, CaM and RPB2 data sets showing the phylogenetic relation of species within the genus Hamigera. The new combination Hamigara brevicompacta is shown in bold font. The BI posterior probability (pp) values and bootstrap percentages of the maximum likelihood (ML) analysis are presented at the nodes; fully supported branches are thickened. Values less than 70 % bootstrap support (ML) or less than 0.95 posterior probability (Bayesian analysis) are indicated with a hyphen or not shown. The bar indicates the number of substitutions per site. The phylogram is rooted with Aspergillus glaucus.

Supplementary Fig. S3

Combined phylogeny for BenA, CaM and RPB2 data sets showing the phylogenetic relation of species within the genus Talaromyces. The new combinations Tal. resinae and Tal. striatoconidium are shown in bold font. The BI posterior probability (pp) values and bootstrap percentages of the maximum likelihood (ML) analysis are presented at the nodes; fully supported branches are thickened. Values less than 70 % bootstrap support (ML) or less than 0.95 posterior probability (Bayesian analysis) are indicated with a hyphen or not shown. The bar indicates the number of substitutions per site. The phylogram is rooted with Ascospirella lutea.

Supplementary Fig. S4

Phylogenetic trees based on BenA, CaM and RPB2 sequence data sets showing the relationship between Penicillium cellarum and P. aurantiogriseum. The data of species mentioned in the phylogram can be found in the “accepted species” list in this article. The BI posterior probability (pp) values and bootstrap percentages of the maximum likelihood (ML) analysis are presented at the nodes; fully supported branches are thickened. Values less than 70 % bootstrap support (ML) or less than 0.95 posterior probability (Bayesian analysis) are indicated with a hyphen or not shown. The bar indicates the number of substitutions per site. The phylogram is rooted with Penicillium chrysogenum NRRL 20818.