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

Abstract

The order Eurotiales is diverse and includes species that impact our daily lives in many ways. In the past, its taxonomy was difficult due to morphological similarities, which made accurate identification of species difficult. This situation improved and stabilised with recent taxonomic and nomenclatural revisions that modernised Aspergillus, Penicillium and Talaromyces. This was mainly due to the availability of curated accepted species lists and the publication of comprehensive DNA sequence reference datasets. This has also led to a sharp increase in the number of new species described each year with the accepted species lists in turn also needing regular updates. The focus of this study was to review the 160 species described between the last list of accepted species published in 2020 until 31 December 2022. To review these species, single-gene phylogenies were constructed and GCPSR (Genealogical Concordance Phylogenetic Species Recognition) was applied. Multi-gene phylogenetic analyses were performed to further determine the relationships of the newly introduced species. As a result, we accepted 133 species (37 Aspergillus, two Paecilomyces, 59 Penicillium, two Rasamsonia, 32 Talaromyces and one Xerochrysium), synonymised 22, classified four as doubtful and created a new combination for Paraxerochrysium coryli, which is classified in Xerochrysium. This brings the number of accepted species to 453 for Aspergillus, 12 for Paecilomyces, 535 for Penicillium, 14 for Rasamsonia, 203 for Talaromyces and four for Xerochrysium. We accept the newly introduced section Tenues (in Talaromyces), and series Hainanici (in Aspergillus sect. Cavernicolarum) and Vascosobrinhoana (in Penicillium sect. Citrina). In addition, we validate the invalidly described species Aspergillus annui and A. saccharicola, and series Annuorum (in Aspergillus sect. Flavi), introduce a new combination for Dichlaena lentisci (type of the genus) and place it in a new section in Aspergillus subgenus Circumdati, provide an updated description for Rasamsonia oblata, and list excluded and recently synonymised species that were previously accepted. This study represents an important update of the accepted species lists in Eurotiales.

Taxonomic novelties: New sections: Aspergillus section Dichlaena Visagie, Kocsubé & Houbraken. New series: Aspergillus series Annuorum J.J. Silva, B.T. Iamanaka, Frisvad. New species: Aspergillus annui J.J. Silva, M.H.P. Fungaro, Frisvad, M.H. Taniwaki & B.T. Iamanaka; Aspergillus saccharicola J.J. Silva, Frisvad, M.H.P. Fungaro, M.H. Taniwaki & B.T. Iamanaka. New combinations: Aspergillus lentisci (Durieu & Mont.) Visagie, Malloch, L. Kriegsteiner, Samson & Houbraken; Xerochrysium coryli (Crous & Decock) Visagie & Houbraken.

Citation: Visagie CM, Yilmaz N, Kocsubé S, Frisvad JC, Hubka V, Samson RA, Houbraken J (2024). A review of recently introduced Aspergillus, Penicillium, Talaromyces and other Eurotiales species. Studies in Mycology 107: 1–66. doi: 10.3114/sim.2024.107.01

INTRODUCTION

Eurotiales is one of the most diverse orders of fungi and includes genera such as Aspergillus, Penicillium, Paecilomyces and Talaromyces. Species identification in these speciose genera has been very difficult in the past. Recent taxonomic and nomenclatural studies have modernised the morphology-based classifications to the extent that these genera now have one of the most modern taxonomies of all fungi. The basis and main driving force for this are the so-called ‘accepted species lists’. Nomenclators such as MycoBank (https://www.mycobank.org/) list more than 3 000 names of Eurotiales. However, many of these names belong to other genera, were considered synonyms of accepted species, or remain unrecognisable because old descriptions were insufficient for recognition and/or no material is available. Knowledge of a genus at a given time was traditionally published in monographs that contained descriptions for all species and keys for their identification. Pitt (1980), in his monograph on Penicillium and its associated sexual (teleomorphic) genera Eupenicillium and Talaromyces, published a review of the recognised names and listed synonyms and excluded names that he considered indeterminate. This overview was later extended to the ‘Names in Current Use’ published by Pitt & Samson (1993) for Trichocomaceae and updated by Pitt et al. (2000). These lists were based on the morphological species concept used at the time, and did not include comments on taxonomy. The main aim of these lists was to record information on species that were ‘accepted’ in these genera, and the purpose was not to formally conserve or reject names as allowed for by the International Code of Nomenclature for algae, fungi, and plants (ICNafp; Turland et al. 2018). The authors of these lists were aware that taxonomy may change as species concepts evolve, and when new concepts were adopted, old names may be accepted in the future. This happened when Houbraken & Samson (2011) revised the taxonomy of Trichocomaceae and classified the species into three families: Aspergillaceae (Aspergillus, Hamigera, Leiothecium, Monascus, Penicilliopsis, Penicillium, Phialomyces, Sclerocleista, Warcupiella and Xeromyces), Thermoascaceae (Byssochlamys/Paecilomyces and Thermoascus) and Trichocomaceae (Rasamsonia, Sagenomella, Talaromyces, Thermomyces, and Trichocoma). They proposed the adoption of Aspergillus and Penicillium over their associated sexual genera, pre-empting the move to a single name nomenclature for fungi (McNeill et al. 2012), and reclassified several other sexual and asexual genera (e.g. Chromocleista, Eupenicillium, Eladia, Hemicarpenteles, Torulomyces and Thysanophora were considered synonymous with Penicillium). These changes were mainly based on phylogenetic analyses, which have become standard practise to study the relationships between and within these genera.

In the following years, accepted species lists for Aspergillus, Penicillium and Talaromyces were published (Samson et al. 2014, Visagie et al. 2014, Yilmaz et al. 2014), representing the first modern lists for fungi to incorporate DNA sequence data into decision-making. Several recommendations were made, from how to work with these genera or describe new species to the procedures required to identify strains more precisely, including the use of the recommended DNA barcode markers β-tubulin (BenA; for Penicillium and Talaromyces) or calmodulin (CaM; for Aspergillus). These recommendations were supported by the metadata associated with each name entry, including authority, citation, MycoBank number, type, ex-type, subgeneric classification and GenBank accession numbers for DNA sequences obtained from ex-type cultures. As with previous lists (Pitt & Samson 1993, Pitt et al. 2000), excluded names were not formally rejected. This approach, together with the wealth of reference data released, resulted in a strong backbone for these genera on which taxonomic revisions of specific groups could be built. It also facilitated the description of new species and their comparison with close relatives, leading to the description of many new species. For this reason, Houbraken et al. (2020) updated the accepted species lists of Aspergillus (increased from 339 to 446 species), Penicillium (increased from 354 to 483 species), Talaromyces (increased from 88 to 171 species) and expanded the list to include other Eurotiales (but excluding Elaphomycetaceae). Houbraken et al. (2020) also reintroduced a series classification in Aspergillus and Penicillium. This taxonomic rank provides information on what functional characters the species might have and is useful in phenotype-based identification. In addition, the current series classification makes it even easier than before to compare putative new species and their close relatives. Since 2020, 160 new species were described in Eurotiales. The focus of this study was to review these and provide comments and opinions on them.

MATERIALS AND METHODS

Phylogenetic analyses

Phylogenies were calculated for all species described since Houbraken et al. (2020), with datasets compiled to represent the genera, sections and/or series these belong to. Datasets (see Table 1 & Suppl. Table S1) were assembled using DNA reference sequences obtained from NCBIs GenBank nucleotide database (https://www.ncbi.nlm.nih.gov/genbank/) and included the internal transcribed spacer rDNA region (ITS), 28S large subunit (LSU), beta-tubulin (BenA), calmodulin (CaM), and RNA polymerase II second largest subunit (RPB2).

Table 1.

List of species described since Houbraken et al. (2020), and reviewed in the current study.

Species	Strain	Status	Subgenus	Section	Series	Country	Substrate	ITS	BenA	CaM	RPB2	LSU	Citation
Aspergillus agricola	A2-A		Circumdati	Flavi	Flavi	USA	Soil (maize field)	-	-	MN987060	-	-	Singh et al. (2020)
	BC09-F		Circumdati	Flavi	Flavi	USA	Soil (maize field)	-	-	MN987065	-	-	Singh et al. (2020)
	C3-J		Circumdati	Flavi	Flavi	USA	Soil (maize field)	-	-	MN987059	-	-	Singh et al. (2020)
	E13-L		Circumdati	Flavi	Flavi	USA	Soil (maize field)	-	-	MN987064	-	-	Singh et al. (2020)
	EC37-C		Circumdati	Flavi	Flavi	USA	Soil (maize field)	-	-	MN987066	-	-	Singh et al. (2020)
	J11-B		Circumdati	Flavi	Flavi	USA	Soil (maize field)	-	-	MN987062	-	-	Singh et al. (2020)
	J11-C		Circumdati	Flavi	Flavi	USA	Soil (maize field)	-	-	MN987063	-	-	Singh et al. (2020)
	J15-H		Circumdati	Flavi	Flavi	USA	Soil (maize field)	-	-	MN987061	-	-	Singh et al. (2020)
	NRRL 66869	T	Circumdati	Flavi	Flavi	USA	Soil (maize field)	-	-	MN987053	-	-	Singh et al. (2020)
	NRRL 66870		Circumdati	Flavi	Flavi	USA	Soil (maize field)	-	-	MN987054	-	-	Singh et al. (2020)
	NRRL 66871		Circumdati	Flavi	Flavi	USA	Soil (maize field)	-	-	MN987055	-	-	Singh et al. (2020)
	NRRL 66872		Circumdati	Flavi	Flavi	USA	Soil (maize field)	-	-	MN987056	-	-	Singh et al. (2020)
	NRRL 66873		Circumdati	Flavi	Flavi	USA	Soil (maize field)	-	-	MN987057	-	-	Singh et al. (2020)
	Sanpatong22		Circumdati	Flavi	Flavi	USA	Soil (maize field)	-	-	MN987068	-	-	Singh et al. (2020)
	Sukhothai 19		Circumdati	Flavi	Flavi	USA	Soil (maize field)	-	-	MN987067	-	-	Singh et al. (2020)
	TXA35-K		Circumdati	Flavi	Flavi	USA	Soil (maize field)	-	-	MN987058	-	-	Singh et al. (2020)
	Ubon3		Circumdati	Flavi	Flavi	USA	Soil (maize field)	-	-	MN987069	-	-	Singh et al. (2020)
Aspergillus alboluteus	CBS 145854		Circumdati	Flavipedes	Spelaei	USA	Air (indoor)	MW448664	MW478498	MW478512	MW478533	-	Sklenář et al. (2021)
	CBS 145855	T	Circumdati	Flavipedes	Spelaei	USA	Air (outdoor)	MW448663	MW478497	MW478511	MW478532	-	Sklenář et al. (2021)
	CBS 145859		Circumdati	Flavipedes	Spelaei	USA	Airconditioner	MW448662	MW478496	MW478510	MW478531	-	Sklenář et al. (2021)
	CBS 147065		Circumdati	Flavipedes	Spelaei	Nigeria	Unknown	MW448666	MW478500	MW478514	MW478535	-	Sklenář et al. (2021)
	CCF 5849		Circumdati	Flavipedes	Spelaei	USA	Storage room	MW448665	MW478499	MW478513	MW478534	-	Sklenář et al. (2021)
	CMW 56637		Circumdati	Flavipedes	Spelaei	Botswana	Soil (bat cave)	MW480881	MW480789	MW480707	MW480791	-	Sklenář et al. (2021)
Aspergillus alboviridis	CBS 142665	T	Circumdati	Flavipedes	Spelaei	Spain	Dung (herbivor)	LT798909	LT798936	LT798937	LT798938	-	Sklenář et al. (2021)
Aspergillus annui	IBT 36122	T	Circumdati	Flavi	Annuorum	Brazil	Sweet paprika	OP691228	ON529842	ON529841	ON529843	-	Silva et al. (2022)
	IBT 36123		Circumdati	Flavi	Annuorum	Brazil	Sweet paprika	-	ON643012	ON643060	ON642964	-	Silva et al. (2022)
	IBT 36124		Circumdati	Flavi	Annuorum	Brazil	Sweet paprika	-	-	-	-	-	Silva et al. (2022)
Aspergillus arizonicus	CCF 5341	T	Fumigati	Fumigati	Neoglabri	USA	Air (hospital)	OK322364	OK334128	OK334127	OK334129	-	Crous et al. (2021b)
Aspergillus banksianus	FRR 6047	T	Fumigati	Fumigati	Brevipedes	Australia	Soil under Banksia integrifolia	MH280013	MT 184780	MT184786	MT184792	-	Crous et al. (2020b)
Aspergillus barbosae	CBS 145863	T	Circumdati	Terrei	Terrei	Brazil	Indoor environment	LR536042	LR031377	LR031392	LR031407	-	Barbosa et al. (2018)
	URM 5870		Circumdati	Terrei	Terrei	Brazil	Industrial castor cake	LR536043	LR031378	LR031393	LR031408	-	Barbosa et al. (2018)
	URM 7011		Circumdati	Terrei	Terrei	Brazil	Soil	LR536041	LR031376	LR031391	LR031406	-	Barbosa et al. (2018)
Aspergillus burnettii	CBS 146237	T	Circumdati	Flavi	Alliacei	Australia	Soil	MK429758	MT211761	MT211762	MT211763	-	Gilchrist et al. (2020)
Aspergillus curvatus	EMCCN2213	T	Circumdati	Circumdati	Steyniorum	Egypt	Water (alkaline lake)	MN006961	-	-	-	-	Al-Bedak (2020a)
Aspergillus gaarensis	AUMC 11046	T	Circumdati	Circumdati	Steyniorum	Egypt	Soil (lake)	MN648408	-	-	-	-	Al-Bedak (2020b)
Aspergillus guangdongensis	CGMCC 3.19704	T	Nidulantes	Ochraceorosei	Funiculosi	China	Soil	MN640760	MN635246	MN635257	MN635269	-	Sun et al. (2022a)
Aspergillus guangxiensis	CGMCC 3.19709	T	Nidulantes	Sparsi	Conjuncti	China	Soil	MN640765	MN635251	MN635262	MN635274	-	Sun et al. (2022a)
	CGMCC 3.19710		Nidulantes	Sparsi	Conjuncti	China	Soil	MN640766	MN635252	MN635263	MN635275	-	Sun et al. (2022a)
Aspergillus hainanicus	CGMCC 3.20888	T	Nidulantes	Cavernicolarum	Hainanici	China	Soil	OM414846	OM475626	OM475630	OM475634	-	Wang & Zhuang (2022b)
Aspergillus hydei	KUMCC 18-0196	T	Circumdati	Nigri	Japonici	China	Air (outdoor)	MT152332	MT161679	MT178247	MT384370	-	Doilom et al. (2020)
Aspergillus inusitatus	CBS 147044	T	Circumdati	Flavipedes	Spelaei	Tunisia	Soil	MW448669	MW478502	MW478517	MW478542	-	Sklenář et al. (2021)
Aspergillus jilinensis	CGMCC 3.18132		Circumdati	Terrei	Terrei	China	Soil	KX443223	KX443161	KX443192	-	-	Huang et al. (2020)
	CGMCC 3.18134	T	Circumdati	Terrei	Terrei	China	Soil	KX443224	KX443162	KX443193	-	-	Huang et al. (2020)
Aspergillus kumbius	FRR 6049	T	Circumdati	Circumdati	Sclerotiorum	Australia	Soil (pasture)	MT179307	MT184782	MT184788	MT184794	-	Crous et al. (2020b)
Aspergillus lannaensis	SDBR-CMUO 6		Nidulantes	Ochraceorosei	Funiculosi	Thailand	Soil	-	MW219782	MW219780	MW219784	-	Boonmee et al. (2021)
	SDBR-CMUO 8	T	Nidulantes	Ochraceorosei	Funiculosi	Thailand	Soil	-	MW219783	MW219781	MW219785	-	Boonmee et al. (2021)
Aspergillus lanuginosus	NRRL4610	T	Circumdati	Flavipedes	Spelaei	Haiti	Soil	EF669604	EU014080	EF669562	EF669646	-	Sklenář et al. (2021)
Aspergillus lebretii	URM 8450		Cremei	Cremei	Wentiorum	Brazil	Air (outdoor)	ON862927	OP672381	OP290539	OP290510	-	Alves et al. (2022b)
	URM 8451	T	Cremei	Cremei	Wentiorum	Brazil	Air (outdoor)	ON862928	OP672382	OP290540	OP290511	-	Alves et al. (2022b)
Aspergillus lentisci	CBS 150189	T	Circumdati	Dichlaena	-	Portugal	Pistacia leaf	OR142402	OR145977	OR145992	OR146003	OR142413	Present study
	DTO 426-F1		Circumdati	Dichlaena	-	Portugal	Pistacia leaf	OR142405	OR145976	OR145991	OR146002	OR142414	Present study
	DTO 426-F2		Circumdati	Dichlaena	-	Portugal	Pistacia leaf	OR142404	OR145975	OR145990	OR146001	OR142415	Present study
	DTO 426-F3		Circumdati	Dichlaena	-	Portugal	Pistacia leaf	OR142409	OR145978	OR145993	OR146004	OR142416	Present study
Aspergillus limoniformis	CGMCC 3.19323	T	Polypaecilum	Polypaecilum	Canini	China	Bat guano	MK329066	MK336093	-	MK335972	-	Zhang et al. (2020)
	LC12610		Polypaecilum	Polypaecilum	Canini	China	Bat guano	MK329067	MK336094	-	MK335973	-	Zhang et al. (2020)
Aspergillus luteorubrus	CBS 146723	T	Fumigati	Fumigati	Fennelliarum	Australia	Soil	MT179305	MT 184781	MT184787	MT184793	-	Crous et al. (2020b)
Aspergillus magnus	UAMH 1324	T	Circumdati	Candidi	Candidi	Canada	Mouse	ON156376	ON164570	ON164619	ON164517	-	Glässnerová et al. (2022)
Aspergillus malvicolor	CBS 146724	T	Circumdati	Circumdati	Sclerotiorum	Australia	Soil under Arachis hypogaea	MT179308	MT184784	MT184790	MT184796	-	Crous et al. (2020b)
Aspergillus marneyi	BRIP 71536a	T	Circumdati	Terrei	Terrei	Australia	Crown of Medicago sativa	OL691080	OL741659	-	OL741656	-	Tan & Shivas (2022)
Aspergillus montoensis	BRIP 71717	T	Circumdati	Terrei	Terrei	Australia	Root of Vigna radiata	OK441076	OK533535	-	OK509073	-	Tan et al. (2021)
Aspergillus nanangensis	CBS 146238	T	Circumdati	Janorum	Janorum	Australia	Soil	MK979278	MT184783	MT184789	MT184795	-	Crous et al. (2020b)
Aspergillus neoterreus	CGMCC 3.20891	T	Circumdati	Terrei	Terrei	China	Soil	OM414849	OM475629	OM475633	OM475637	-	Wang & Zhuang (2022b)
Aspergillus neotritici	CBS 129260		Circumdati	Candidi	Candidi	USA	Soil	ON156397	ON164591	ON164632	ON164541	-	Glässnerová et al. (2022)
	CBS 129307		Circumdati	Candidi	Candidi	Unknown	Soil	ON156398	ON164592	ON164633	ON164542	-	Glässnerová et al. (2022)
	CBS 133055		Circumdati	Candidi	Candidi	Japan	Unknown	ON156395	ON164587	ON164628	ON164537	-	Glässnerová et al. (2022)
	CBS 266.81		Circumdati	Candidi	Candidi	India	Triticum aestivum grains	LT626958	EU076293	EU076305	MN969098	-	Glässnerová et al. (2022)
	CCF 1649		Circumdati	Candidi	Candidi	Czech Republic	Flour	FR733810	LT627024	FR751427	LT627025	-	Glässnerová et al. (2022)
	CCF 3314		Circumdati	Candidi	Candidi	Czech Republic	Air (outdoor)	FR733812	LT627022	FR751426	LT627023	-	Glässnerová et al. (2022)
	CCF 3853	T	Circumdati	Candidi	Candidi	Czech Republic	Human toenail	FR727136	FR775327	HE661598	LT627021	-	Glässnerová et al. (2022)
	CCF 4030		Circumdati	Candidi	Candidi	Czech Republic	Vermicompost	FR733814	LT627018	FR751425	LT627019	-	Glässnerová et al. (2022)
	CCF 4653		Circumdati	Candidi	Candidi	Czech Republic	Human toenail	HG915890	HG916674	HG916677	LT627020	-	Glässnerová et al. (2022)
	CCF 4658		Circumdati	Candidi	Candidi	Czech Republic	Human toenail	HG915891	HG916675	HG916676	LT627026	-	Glässnerová et al. (2022)
	CCF 4914		Circumdati	Candidi	Candidi	USA	Air (hospital)	ON156392	ON164556	ON164605	ON164503	-	Glässnerová et al. (2022)
	CCF 6202		Circumdati	Candidi	Candidi	USA	Air (house)	ON156396	ON164588	ON164629	ON164538	-	Glässnerová et al. (2022)
	CCF 6397		Circumdati	Candidi	Candidi	Czech Republic	Human abdominal cavity	ON156394	ON164589	ON164630	ON164539	-	Glässnerová et al. (2022)
	IBT 12659		Circumdati	Candidi	Candidi	USA	Soil (kangaroo rat burrow)	ON156393	ON164557	ON164606	ON164504	-	Glässnerová et al. (2022)
Aspergillus okavangoensis	CMW 56636	T	Circumdati	Flavipedes	Flavipedes	Botswana	Soil (bat cave)	MW480880	MW480788	MW480706	MW480790	-	Visagie et al. (2021)
Aspergillus oxumiae	CCDCA11546	T	Circumdati	Nigri	Japonici	Brazil	Soil under Agave sisalana	MN431160	-	MN531842	MN521389	-	Crous et al. (2020b)
Aspergillus phialiformis	CGMCC 3.19314	T	Polypaecilum	Polypaecilum	Canini	China	Rock	MK329068	MK336095	-	MK335974	-	Zhang et al. (2020)
	LC12537		Polypaecilum	Polypaecilum	Canini	China	Rock	MK329069	MK336096	-	MK335975	-	Zhang et al. (2020)
Aspergillus phialosimplex	CGMCC 3.19637	T	Polypaecilum	Polypaecilum	Canini	China	Plant debris	MK329070	MK336097	-	MK335976	-	Zhang et al. (2020)
	LC12625		Polypaecilum	Polypaecilum	Canini	China	Animal faeces	MK329071	MK336098	-	MK335977	-	Zhang et al. (2020)
	LC12658		Polypaecilum	Polypaecilum	Canini	China	Plant root	MK329072	MK336099	-	MK335978	-	Zhang et al. (2020)
Aspergillus gilianyuensis	CGMCC 3.20889	T	Nidulantes	Nidulantes	Versicolores	China	Soil	OM414847	OM475627	OM475631	OM475635	-	Wang & Zhuang (2022b)
Aspergillus recifensis	CBS 145864	T	Circumdati	Terrei	Nivei	Brazil	Soil	LR536036	LR031370	LR031385	LR031400	-	Barbosa et al. (2018)
	URM 2803		Circumdati	Terrei	Nivei	Brazil	Bird food	LR536040	LR031375	LR031390	LR031405	-	Barbosa et al. (2018)
	URM 3371		Circumdati	Terrei	Nivei	Brazil	Bird food	-	KR051530	-	-	-	Barbosa et al. (2018)
	URM 3571		Circumdati	Terrei	Nivei	Brazil	Water from tank	LR536039	LR031373	LR031388	LR031403	-	Barbosa et al. (2018)
	URM 5262		Circumdati	Terrei	Nivei	Brazil	Soil (rhizosphere of Croton sp)	LR536037	LR031371	LR031386	LR031401	-	Barbosa et al. (2018)
	URM 5461		Circumdati	Terrei	Nivei	Brazil	Water from pool	LR536038	LR031372	LR031387	LR031402	-	Barbosa et al. (2018)
	URM 6628		Circumdati	Terrei	Nivei	Brazil	Soil	LR536035	LR031369	LR031384	LR031399	-	Barbosa et al. (2018)
Aspergillus rouenensis	CBS 149067	T	Polypaecilum	Polypaecilum	Salinarum	France	Quercus bore dust of Xestobium rufovillosum	ON603782	ON605641	ON653193	ON653194	-	Crous et al. (2022)
	CBS 149068		Polypaecilum	Polypaecilum	Salinarum	France	Quercus bore dust of Xestobium rufovillosum	-	ON605642	-	-	-	Crous et al. (2022)
Aspergillus saccharicola	IBT 36125		Circumdati	Flavi	Flavi	Brazil	Sugarcane	-	ON642978	ON643026	ON642930	-	Silva et al. (2022)
	IBT 36126	T	Circumdati	Flavi	Flavi	Brazil	Sugarcane	OP611470	ON529845	ON529844	ON529846	-	Silva et al. (2022)
	IBT 36127		Circumdati	Flavi	Flavi	Brazil	Sugarcane	-	ON642982	ON643030	ON642934	-	Silva et al. (2022)
Aspergillus sakultaensis	AUMC 13885	T	Circumdati	Flavipedes	Flavipedes	Egypt	Water	MK391495	-	-	-	-	Zhori et al. (2020)
Aspergillus sibiricus	CBS 143307	T	Fumigati	Fumigati	Unilaterales	Russia	Soil (cole mine)	MG587008	MG722970	MG722971	MG710809	-	Iliushin (2022)
Aspergillus sichuanensis	CGMCC 3.19705	T	Nidulantes	Aenei	Aenei	China	Soil	MN640761	MN635247	MN635258	MN635270	-	Sun et al. (2022a)
	CGMCC 3.19706		Nidulantes	Aenei	Aenei	China	Soil	MN640762	MN635248	MN635259	MN635271	-	Sun et al. (2022a)
	CGMCC 3.19708		Nidulantes	Aenei	Aenei	China	Soil	MN640764	MN635250	MN635261	MN635273	-	Sun et al. (2022a)
Aspergillus sigarelli	CBS 141579	T	Nidulantes	Usti	Calidousti	China	Cigarette	MN640758	MN635244	MN635255	MN635267	-	Sun et al. (2020c)
Aspergillus telluris	CGMCC 3.19701	T	Polypaecilum	Polypaecilum	Canini	China	Soil	MN640767	MN635253	MN635264	MN635276	-	Sun et al. (2022a)
	CGMCC 3.19702		Polypaecilum	Polypaecilum	Canini	China	Soil	MN640768	MN635254	MN635265	MN635277	-	Sun et al. (2022a)
	CGMCC 3.19703		Polypaecilum	Polypaecilum	Canini	China	Soil	MN640769	MN635243	MN635266	MN635278	-	Sun et al. (2022a)
Aspergillus tenebricus	CBS 147048	T	Circumdati	Candidi	Candidi	South Africa	Soil	ON156389	ON164584	ON164623	ON164532	-	Glässnerová et al. (2022)
	CBS 147376		Circumdati	Candidi	Candidi	Australia	Soil	ON156390	ON164585	ON164624	ON164533	-	Glässnerová et al. (2022)
	DTO 440-E2		Circumdati	Candidi	Candidi	Australia	Soil	ON156391	ON164586	ON164625	ON164534	-	Glässnerová et al. (2022)
Aspergillus tibetensis	CGMCC 3.19707	T	Nidulantes	Aenei	Aenei	China	Soil	MN640763	MN635249	MN635260	MN635272	-	Sun et al. (2022a)
Aspergillus toxicus	A34-N		Circumdati	Flavi	Flavi	USA	Soil (maize field)	-	-	MN987102	-	-	Singh et al. (2020)
	BG14-F		Circumdati	Flavi	Flavi	USA	Soil (maize field)	-	-	MN987108	-	-	Singh et al. (2020)
	BRG3458A		Circumdati	Flavi	Flavi	USA	Soil (maize field)	-	-	MN987090	-	-	Singh et al. (2020)
	BRG3458H		Circumdati	Flavi	Flavi	USA	Soil (maize field)	-	-	MN987095	-	-	Singh et al. (2020)
	BRG3458J		Circumdati	Flavi	Flavi	USA	Soil (maize field)	-	-	MN987096	-	-	Singh et al. (2020)
	BRG5138J		Circumdati	Flavi	Flavi	USA	Soil (maize field)	-	-	MN987097	-	-	Singh et al. (2020)
	CR10-G		Circumdati	Flavi	Flavi	USA	Soil (maize field)	-	-	MN987105	-	-	Singh et al. (2020)
	CR20-D		Circumdati	Flavi	Flavi	USA	Soil (maize field)	-	-	MN987098	-	-	Singh et al. (2020)
	CR24-F		Circumdati	Flavi	Flavi	USA	Soil (maize field)	-	-	MN987103	-	-	Singh et al. (2020)
	D16-J		Circumdati	Flavi	Flavi	USA	Soil (maize field)	-	-	MN987099	-	-	Singh et al. (2020)
	D25-A-S		Circumdati	Flavi	Flavi	USA	Soil (maize field)	-	-	MN987100	-	-	Singh et al. (2020)
	E21-B		Circumdati	Flavi	Flavi	USA	Soil (maize field)	-	-	MN987101	-	-	Singh et al. (2020)
	EC24-C		Circumdati	Flavi	Flavi	USA	Soil (maize field)	-	-	MN987106	-	-	Singh et al. (2020)
	EC49-L		Circumdati	Flavi	Flavi	USA	Soil (maize field)	-	-	MN987107	-	-	Singh et al. (2020)
	J15-B		Circumdati	Flavi	Flavi	USA	Soil (maize field)	-	-	MN987104	-	-	Singh et al. (2020)
	K44-K		Circumdati	Flavi	Flavi	USA	Soil (maize field)	-	-	MN987085	-	-	Singh et al. (2020)
	K849-B		Circumdati	Flavi	Flavi	USA	Soil (maize field)	-	-	MN987086	-	-	Singh et al. (2020)
	NRRL 66868	T	Circumdati	Flavi	Flavi	USA	Soil (maize field)	-	-	MN987092	-	-	Singh et al. (2020)
	NRRL 66897		Circumdati	Flavi	Flavi	USA	Soil (maize field)	-	-	MN987091	-	-	Singh et al. (2020)
	NRRL 66899		Circumdati	Flavi	Flavi	USA	Soil (maize field)	-	-	MN987093	-	-	Singh et al. (2020)
	NRRL 66900		Circumdati	Flavi	Flavi	USA	Soil (maize field)	-	-	MN987094	-	-	Singh et al. (2020)
	TX04A5-B		Circumdati	Flavi	Flavi	USA	Soil (maize field)	-	-	MN987089	-	-	Singh et al. (2020)
	TX07CB73-I		Circumdati	Flavi	Flavi	USA	Soil (maize field)	-	-	MN987087	-	-	Singh et al. (2020)
	TXLaFeria2-F		Circumdati	Flavi	Flavi	USA	Soil (maize field)	-	-	MN987088	-	-	Singh et al. (2020)
Aspergillus vinaceus	ITAL 47.456	T	Circumdati	Nigri	Nigri	Brazil	Grapes (Vitis labrusca)	MN575692	MN583579	MN583580	MN583581	-	Silva et al. (2020)
Aspergillus xishaensis	CGMCC 3.20890	T	Circumdati	Flavipedes	Flavipedes	China	Soil	OM414848	OM475628	OM475632	OM475636	-	Wang & Zhuang (2022b)
Paecilomyces clematidis	CBS 148466	T	-	-	-	Czech Republic	Root of Clematis	MZ923760	MZ927740	MZ927738	OL332316	-	Spetik et al. (2022)
	MEND-F-0561		-	-	-	Czech Republic	Root of Clematis	MZ923761	MZ927741	MZ927739	OL332317	-	Spetik et al. (2022)
Paecilomyces penicilliformis	CCF 5755	T	-	-	-	USA	Air (pharmacy)	LR679769	LR679768	LR778299	-	-	Crous et al. (2020b)
	CCF 6350		-	-	-	USA	Juice (peach-mango)	LR736038	LR778163	LR778165	-	-	Crous et al. (2020b)
Paraxerochrysium coryli	CBS 148314	T	-	-	-	Belgium	Hazelnut (Corylus avellana)	OK664748	OK651216	-	OK651178	OK663787	Crous et al. (2021b)
Penicillium allaniae	BRIP 74886a	T	Aspergilloides	Exilicaulis	Restrict!	Australia	Soil	OP903476	OP921959	OP921957	OP921958	OP925816	Tan & Shivas (2022)
	BRIP 74899		Aspergilloides	Exilicaulis	Restrict!	Australia	Unknown	OP903475	OP921956	OP921954	OP921955		Tan & Shivas (2022)
Penicillium allsoppiae	CBS 138943	T	Penicillium	Canescentia	Canescentia	South Africa	Soil	JX140830	JX140992	JX157384	KP016895	-	Visagie & Yilmaz (2022)
	CBS 138945		Penicillium	Canescentia	Canescentia	South Africa	Soil	JX140822	JX141004	JX157399	KP016910	-	Visagie & Yilmaz (2022)
	CN 086C6		Penicillium	Canescentia	Canescentia	South Africa	Soil	MW364385	MW357820	MW357831	MW357840	-	Visagie & Yilmaz (2022)
	CN 086C7		Penicillium	Canescentia	Canescentia	South Africa	Soil	MW364386	MW357821	MW357832	MW357841	-	Visagie & Yilmaz (2022)
	CN 086C8		Penicillium	Canescentia	Canescentia	South Africa	Soil	MW364387	MW357822	-	MW357842	-	Visagie & Yilmaz (2022)
Penicillium anthracinoglaciei	EXF-11216		Penicillium	Brevicompacta	Brevicompacta	Greenland	Cryoconite	-	MT080468	MT080527	MT080509	-	Perini et al. (2023)
	EXF-11218		Penicillium	Brevicompacta	Brevicompacta	Greenland	Cryoconite	-	MT080469	MT080528	MT080510	-	Perini et al. (2023)
	EXF-11222		Penicillium	Brevicompacta	Brevicompacta	Greenland	Cryoconite	-	MT080472	MT080531	MT080511	-	Perini et al. (2023)
	EXF-11226		Penicillium	Brevicompacta	Brevicompacta	Greenland	Supraglacial water	-	MT080475	MT080534	MT080512	-	Perini et al. (2023)
	EXF-11230		Penicillium	Brevicompacta	Brevicompacta	Greenland	Supraglacial water	-	MT080479	MT080538	MT080508	-	Perini et al. (2023)
	EXF-11232		Penicillium	Brevicompacta	Brevicompacta	Greenland	Supraglacial water	-	MT080481	MT080540	MT080513	-	Perini et al. (2023)
	EXF-11233		Penicillium	Brevicompacta	Brevicompacta	Greenland	Supraglacial water	-	MT080482	MT080541	MT080514	-	Perini et al. (2023)
	EXF-11237		Penicillium	Brevicompacta	Brevicompacta	Greenland	Cryoconite	-	MT080483	MT080542	MT080515	-	Perini et al. (2023)
	EXF-11240		Penicillium	Brevicompacta	Brevicompacta	Greenland	Cryoconite	-	MT080485	MT080544	MT080516	-	Perini et al. (2023)
	EXF-11241		Penicillium	Brevicompacta	Brevicompacta	Greenland	Clear ice	-	MT080486	MT080545	MT080517	-	Perini et al. (2023)
	EXF-11443	T	Penicillium	Brevicompacta	Brevicompacta	Greenland	Dark ice	-	MT080493	MT080552	MT080519	-	Perini et al. (2023)
	EXF-11444		Penicillium	Brevicompacta	Brevicompacta	Greenland	Dark ice	-	MT080494	MT080553	MT080520	-	Perini et al. (2023)
	EXF-11445		Penicillium	Brevicompacta	Brevicompacta	Greenland	Dark ice	-	MT080495	MT080554	MT080521	-	Perini et al. (2023)
	EXF-11448		Penicillium	Brevicompacta	Brevicompacta	Greenland	Cryoconite	-	MT080498	MT080557	MT080522	-	Perini et al. (2023)
	EXF-11451		Penicillium	Brevicompacta	Brevicompacta	Greenland	Cryoconite	-	MT080501	MT080560	MT080523	-	Perini et al. (2023)
	EXF-11453		Penicillium	Brevicompacta	Brevicompacta	Greenland	Clear ice	-	MT080503	MT080562	MT080524	-	Perini et al. (2023)
	EXF-11454		Penicillium	Brevicompacta	Brevicompacta	Greenland	Clear ice	-	MT080504	MT080563	MT080525	-	Perini et al. (2023)
	EXF-11456		Penicillium	Brevicompacta	Brevicompacta	Greenland	Clear ice	-	MT080506	MT080565	MT080526	-	Perini et al. (2023)
Penicillium aquadulcis	CNUFC JT1301	T	Aspergilloides	Citrina	Westlingiorum	Republic of Korea	Water	OK356194	QK105100	OK105102	-	-	Nguyen et al. (2021b)
	CNUFC JT1302		Aspergilloides	Citrina	Westlingiorum	Republic of Korea	Water	OK356195	OK105101	OK105103	-	-	Nguyen et al. (2021b)
Penicillium archerae	BRIP 72549c	T	Aspergilloides	Exilicaulis	Restricti	Australia	Soil	OP903477	OP921961	-	OP921960	-	Tan & Shivas (2022)
Penicillium aspericonidium	CBS 141832	T	Aspergilloides	Charlesia	Indica	Australia	Soil	MT309657	MT302240	-	MT302224	-	Sun et al. (2021)
Penicillium ausonanum	FMR 16948	T	Aspergilloides	Lanata-Divaricata	Dalearum	Spain	Fluvial sediment	LR655808	LR655809	LR655810	LR655811	-	Torres-Garcia et al. (2022)
Penicillium barbosae	URM 7705	T	Aspergilloides	Sclerotiorum	Adametziorum	Brazil	Soil	MW191494	MG452818	MW183245	LR898886	-	Ramos et al. (2021)
	URM 7824		Aspergilloides	Sclerotiorum	Adametziorum	Brazil	Soil	MW191495	MG452819	MW183246	LR898887	-	Ramos et al. (2021)
Penicillium cerradense	UB23977	T	Aspergilloides	Citrina	Sumatraensia	Brazil	Soil	MT006126	MT416533	MT416534	MT416532	-	Andrade et al. (2021)
	DCFS6b		Aspergilloides	Citrina	Sumatraensia	Brazil	Soil	MT006127	MT416536	MT416537	MT416535	-	Andrade et al. (2021)
Penicillium claroviride	CMW 56197	T	Penicillium	Canescentia	Atroveneta	South Africa	Soil	MT949909	MT957414	MT957456	MT957482	-	Visagie & Yilmaz (2022)
	CMW 56198		Penicillium	Canescentia	Atroveneta	South Africa	Soil	MT949910	MT957415	MT957457	MT957483	-	Visagie & Yilmaz (2022)
Penicillium doidgeae	CBS 138947	T	Penicillium	Canescentia	Atroveneta	South Africa	Mite from Protea repens infructescens	JX140804	JX141006	JX157413	KP016915	-	Visagie & Yilmaz (2022)
	CBS 138948		Penicillium	Canescentia	Atroveneta	South Africa	Mite from Protea repens infructescens	JX140805	JX141007	JX157414	KP016916	-	Visagie & Yilmaz (2022)
Penicillium donggangicum	AS 3.15900	T	Aspergilloides	Lanata-Divaricata	Janthinella	China	Soil	MW946996	MZ004914	MZ004918	MW979253	-	Xu et al. (2022)
Penicillium eickerii	CBS 138939	T	Penicillium	Canescentia	Canescentia	South Africa	Mite from Protea repens infructescens	JX140824	JX140979	JX157365	KP016876	-	Visagie & Yilmaz (2022)
	CBS 138940		Penicillium	Canescentia	Canescentia	South Africa	Bract from Protea repens infructescens	JX140825	JX140980	JX157366	KP016877	-	Visagie & Yilmaz (2022)
Penicillium elizabethiae	NRRL917	T	Penicillium	Canescentia	Canescentia	Scotland	Soil	KP016840	KJ866964	KJ867021	KP016918	-	Visagie & Yilmaz (2022)
Penicillium ezekielii	CBS 149115	T	Aspergilloides	Cinnamopurpurea	Jiangxiensia	Indonesia	Zea mays kernels	ON723772	ON920778	ON920781	ON920784	ON911289	Tan et al. (2022)
	CBS 149114		Aspergilloides	Cinnamopurpurea	Jiangxiensia	Indonesia	Zea mays kernels	ON723771	ON920777	ON920780	ON920783	ON911288	Tan et al. (2022)
	DTO 463-A7		Aspergilloides	Cinnamopurpurea	Jiangxiensia	Nigeria	Oryza sativa kernels	ON723773	ON920779	ON920782	-	ON911290	Tan et al. (2022)
Penicillium ferraniaense	CBS 147594		Aspergilloides	Sclerotiorum	Sclerotiorum	Italy	Compost	MW694952	MW689337	MW689339	MW689341	-	Crous et al. (2021a)
	CBS 147595	T	Aspergilloides	Sclerotiorum	Sclerotiorum	Italy	Compost	MW694951	MW689336	MW689338	MW689340	-	Crous et al. (2021a)
Penicillium fusiforme	CBS 250.66	T	Aspergilloides	Charlesia	Fellutana	The Netherlands	Unknown	MT309668	MT302253	MT302220	MT302236	-	Sun et al. (2021)
Penicillium gercinae	URM 8348	T	Aspergilloides	Ramigena	Georgiensia	Brazil	Soil	MW648591	MW646389	MW646391	MW646393	-	Alves et al. (2022a)
Penicillium guarroi	FMR 17747	T	Aspergilloides	Gracilenta	Estinogena	Spain	Fluvial sediment	LR814139	LR814134	LR814140	LR814145	-	Torres-Garcia et al. (2022)
Penicillium hepuense	AS 3.16039	T	Aspergilloides	Lanata-Divaricata	Oxalica	China	Soil	MW946994	MZ004912	MZ004916	MW979254	-	Xu et al. (2022)
	AS 3.16040		Aspergilloides	Lanata-Divaricata	Oxalica	China	Soil	MW946995	MZ004913	MZ004917	MW979255	-	Xu et al. (2022)
Penicillium irregulare	FMR 17859	T	Penicillium	Canescentia	Canescentia	Spain	Fluvial sediment	LR814181	LR814144	LR814151	LR814182	-	Torres-Garcia et al. (2022)
Penicillium jenningsiae	BRIP 45936a	T	Aspergilloides	Citrina	Sumatraensia	Australia	Compost	-	OL741657	-	OL741660	-	Tan & Shivas (2022)
Penicillium jiaozhouwanicum	AS 3.16027		Aspergilloides	Lanata-Divaricata	Oxalica	China	Soil	OM203537	OM220087	OM220088	OM220089	-	Xu et al. (2022)
	AS 3.16038	T	Aspergilloides	Lanata-Divaricata	Oxalica	China	Soil	MW946993	MZ004911	MZ004915	MW979252	-	Xu et al. (2022)
Penicillium kalander	CMW 56202	T	Aspergilloides	Sclerotiorum	Sclerotiorum	South Africa	Soil	MT949914	MT957421	MT957461	MT957487	-	Visagie & Yilmaz (2022)
	CMW 56203		Aspergilloides	Sclerotiorum	Sclerotiorum	South Africa	Soil	MT949915	MT957422	MT957462	MT957488	-	Visagie & Yilmaz (2022)
	CMW 56204		Aspergilloides	Sclerotiorum	Sclerotiorum	South Africa	Soil	MT949916	MT957423	MT957463	MT957489	-	Visagie & Yilmaz (2022)
	CMW 56205		Aspergilloides	Sclerotiorum	Sclerotiorum	South Africa	Soil	MT949917	MT957424	MT957464	MT957490	-	Visagie & Yilmaz (2022)
	CMW 56390		Aspergilloides	Sclerotiorum	Sclerotiorum	South Africa	Soil	MT949918	MT957425	MT957465	MT957491	-	Visagie & Yilmaz (2022)
Penicillium krskae	CBS 147776	T	Aspergilloides	Exilicaulis	Restricta	Austria	Air (indoor)	MW794123	MW774594	MW774595	MW774593	-	Labuda et al. (2021)
Penicillium limae	URM 7706	T	Aspergilloides	Sclerotiorum	Adametziorum	Brazil	Soil	MW191493	MG452820	MW183244	LR898888	-	Ramos et al. (2021)
Penicillium linzhiense	Z863	T	Penicillium	Canescentia	Canescentia	China	Soil	MT461156	MT 461157	MT461162	-	-	Liang et al. (2021)
Penicillium longiconidiophorum	CBS 141831	T	Aspergilloides	Charlesia	Phoenicea	Madagascar	Soil	MT309669	MT302254	MT302221	MT302237	-	Sun et al. (2021)
	DTO 092-C6		Aspergilloides	Charlesia	Phoenicea	Madagascar	Soil	MT309670	MT302255	MT302222	MT302238	-	Sun et al. (2021)
Penicillium mattheeae	CMW 56195		Aspergilloides	Aspergilloides	Saturniiormia	South Africa	Soil	MT949905	MT957409	MT957452	MT957478	-	Visagie & Yilmaz (2022)
	CMW 56388	T	Aspergilloides	Aspergilloides	Saturniiormia	South Africa	Soil	MT949904	MT957408	MT957451	MT957477	-	Visagie & Yilmaz (2022)
	CMW 56633		Aspergilloides	Aspergilloides	Saturniiormia	South Africa	Soil	MT949906	MT957410	MT957453	MT957479	-	Visagie & Yilmaz (2022)
Penicillium melanosporum	CBS 146938	T	Aspergilloides	Lanata-Divaricata	Janthinella	Spain	Soil	LR655192	LR655196	LR655200	LR655204	-	Rodriguez-Andrade et al. (2021)
Penicillium michoacanense	FMR 17612	T	Aspergilloides	Lanata-Divaricata	Janthinella	Mexico	Soil	LR655194	LR655198	LR655202	LR655206	-	Rodriguez-Andrade et al. (2021)
Penicillium neoherguei	CBS 148692	T	Aspergilloides	Sclerotiorum	Hergueorum	USA	White mushroom sporocarp	MW341222	OL840853	OL840855	MW349119	-	Crous et al. (2022)
Penicillium newtonturnerae	BRIP 74909a	T	Aspergilloides	Lanata-Divaricata	Simplicissima	Australia	Soil	OP903478	OP921964	OP921962	OP921963	OP925817	Tan & Shivas (2022)
Penicillium nordestinense	CBS 564.85		Aspergilloides	Lanata-Divaricata	Janthinella	Brazil	Pollen samples inside nests of (Melipona scutellaris)	OV312015	MH846596	MH846609	MH846584	-	Barbosa et al. (2022)
	URM 8423	T	Aspergilloides	Lanata-Divaricata	Janthinella	Brazil	Pollen samples inside nests of (Melipona scutellaris)	OV265270	OV265324	OV265272	OM927721	-	Barbosa et al. (2022)
	URM 8424		Aspergilloides	Lanata-Divaricata	Janthinella	Brazil	Pollen samples inside nests of (Melipona scutellaris)	OV265271	OV265337	OV265273	OM927721	-	Barbosa et al. (2022)
Penicillium outeniguaense	CMW 56387	T	Aspergilloides	Citrina	Westlingiorum	South Africa	Soil	MT949903	MT957405	MT957450	MT957476	-	Visagie & Yilmaz (2022)
Penicillium poederi	CBS 147622	T	Aspergilloides	Torulomyces	Torulomyces	Iceland	Volcanic soil	MF611757	MF611760	MF611763	MF611766	-	Kirchmair et al. (2022)
	CBS 147623		Aspergilloides	Torulomyces	Torulomyces	Iceland	Volcanic soil	MF611758	MF611761	MF611764	MF611767	-	Kirchmair et al. (2022)
	CBS 147624		Aspergilloides	Torulomyces	Torulomyces	Iceland	Volcanic soil	MF611759	MF611762	MF611765	MF611768	-	Kirchmair et al. (2022)
Penicillium pole-evansii	CBS 138946	T	Penicillium	Canescentia	Atroveneta	South Africa	Bract from Protea repens infructescens	JX140831	JX141005	JX157412	KP016911	-	Visagie & Yilmaz (2022)
Penicillium rotoruae	CBS 145838	T	Aspergilloides	Lanata-Divaricata	Rolfsiorum	New Zealand	Pinus radiata timber on ground	MN315103	MN315104	MN315102	MT240842	-	O’Callahan et al. (2020)
Penicillium saanichanum	DAOMC 251850	T	Aspergilloides	Cinnamopurpurea	Idahoensia	Canada	House dust	KY469059	KY469096	KY469020	MN795070	-	Crous et al. (2020a)
Penicillium sanjayi	NFCCI 5017	T	Aspergilloides	Citrina	Vascosobrinhoana	India	Soil	MZ571358	MZ558484	MZ558492	MZ558482	-	Ashtekar et al. (2022)
	NFCCI 5018		Aspergilloides	Citrina	Vascosobrinhoana	India	Soil	MZ571359	MZ558485	MZ558493	MZ558483	-	Ashtekar et al. (2022)
Penicillium scottii	CBS 138935		Penicillium	Canescentia	Canescentia	South Africa	Air (outdoor)	JX140823	JX140977	JX157351	KP016863	-	Visagie & Yilmaz (2022)
	CBS 138937		Penicillium	Canescentia	Canescentia	South Africa	Soil	JX140826	JX140978	JX157355	KP016867	-	Visagie & Yilmaz (2022)
	CBS 138941		Penicillium	Canescentia	Canescentia	South Africa	Air (outdoor)	JX140827	JX140981	JX157371	KP016882	-	Visagie & Yilmaz (2022)
	CBS 138944		Penicillium	Canescentia	Canescentia	South Africa	Bract from Protea repens infructescens	JX140820	JX141002	JX157396	KP016907	-	Visagie & Yilmaz (2022)
	CBS 138951	T	Penicillium	Canescentia	Canescentia	South Africa	Soil	JX140812	JX140991	JX157383	KP016894	-	Visagie & Yilmaz (2022)
	CV0939		Penicillium	Canescentia	Canescentia	South Africa	Soil	JX140814	JX140994	JX157386	KP016897	-	Visagie & Yilmaz (2022)
	IBT 31903		Penicillium	Canescentia	Canescentia	South Africa	Bract from Protea repens infructescens	JX140821	JX141003	JX157397	KP016908	-	Visagie & Yilmaz (2022)
	IBT 31904		Penicillium	Canescentia	Canescentia	South Africa	Soil	KP016833	JX140995	JX157387	KP016898	-	Visagie & Yilmaz (2022)
	IBT 31906		Penicillium	Canescentia	Canescentia	South Africa	Soil	JX140815	JX140996	JX157388	KP016899	-	Visagie & Yilmaz (2022)
	IBT 31907		Penicillium	Canescentia	Canescentia	South Africa	Soil	KP016832	JX140988	JX157378	KP016889	-	Visagie & Yilmaz (2022)
	IBT 31908		Penicillium	Canescentia	Canescentia	South Africa	Soil	JX140816	JX140997	JX157389	KP016900	-	Visagie & Yilmaz (2022)
	IBT 31909		Penicillium	Canescentia	Canescentia	South Africa	Soil	KP016834	JX140998	JX157390	KP016901	-	Visagie & Yilmaz (2022)
	IBT 31910		Penicillium	Canescentia	Canescentia	South Africa	Soil	JX140817	JX140999	JX157391	KP016902	-	Visagie & Yilmaz (2022)
	IBT 31911		Penicillium	Canescentia	Canescentia	South Africa	Soil	JX140818	JX141000	JX157392	KP016903	-	Visagie & Yilmaz (2022)
	IBT 31912		Penicillium	Canescentia	Canescentia	South Africa	Soil	JX140808	JX140984	JX157374	KP016885	-	Visagie & Yilmaz (2022)
	IBT 31913		Penicillium	Canescentia	Canescentia	South Africa	Soil	JX140811	JX140990	JX157382	KP016893	-	Visagie & Yilmaz (2022)
	IBT 31914		Penicillium	Canescentia	Canescentia	South Africa	Soil	JX140813	JX140993	JX157385	KP016896	-	Visagie & Yilmaz (2022)
	IBT 31915		Penicillium	Canescentia	Canescentia	South Africa	Soil	JX140807	JX140982	JX157372	KP016883	-	Visagie & Yilmaz (2022)
	IBT 31916		Penicillium	Canescentia	Canescentia	South Africa	Soil	JX140809	JX140985	JX157375	KP016886	-	Visagie & Yilmaz (2022)
	IBT 31917		Penicillium	Canescentia	Canescentia	South Africa	Soil	JX140810	JX140987	JX157377	KP016888	-	Visagie & Yilmaz (2022)
	IBT31918		Penicillium	Canescentia	Canescentia	South Africa	Soil	JX140828	JX140986	JX157376	KP016887	-	Visagie & Yilmaz (2022)
	IBT 31919		Penicillium	Canescentia	Canescentia	South Africa	Soil	KP016831	JX140983	JX157373	KP016884	-	Visagie & Yilmaz (2022)
	IBT 31922		Penicillium	Canescentia	Canescentia	South Africa	Mite from Protea repens infructescens	JX140819	JX141001	JX157394	KP016905	-	Visagie & Yilmaz (2022)
	IBT 31953		Penicillium	Canescentia	Canescentia	South Africa	Soil	JX140829	JX140989	JX157379	KP016890	-	Visagie & Yilmaz (2022)
Penicillium setosum	CBS 144865	T	Aspergilloides	Lanata-Divaricata	Janthinella	India	Withania somniiera	KT852579	MF184995	MH105905	-	-	Barbosa et al. (2022)
	CBS 576.70		Aspergilloides	Lanata-Divaricata	Janthinella	Mexico	Soil	-	MH846595	MH846608	MH846583	-	Barbosa et al. (2022)
	DTO 284-F3		Aspergilloides	Lanata-Divaricata	Janthinella	India	Withania somniiera	-	MH846594	MH846607	MH846582	-	Barbosa et al. (2022)
	PPRI 20582		Aspergilloides	Lanata-Divaricata	Janthinella	South Africa	Insect	MK450718	MK451255	MK451649	-	-	Barbosa et al. (2022)
	PPRI 6371		Aspergilloides	Lanata-Divaricata	Janthinella	South Africa	Scarabid larvae	MK450717	MK 451227	MK451648	MK450852	-	Barbosa et al. (2022)
Penicillium sexuale	CBS 146939	T	Aspergilloides	Crypta	Crypta	Spain	Soil	LR655195	LR655199	LR655203	LR655207	-	Rodriguez-Andrade et al. (2021)
Penicillium siccitolerans	FMR 17381	T	Aspergilloides	Lanata-Divaricata	Janthinella	Spain	Soil	LR655193	LR655197	LR655201	LR655205	-	Rodriguez-Andrade et al. (2021)
Penicillium sicoris	FMR 18076	T	Penicillium	Paradoxa	Atramentosa	Spain	Fluvial sediment	LR884497	LR884494	LR884496	LR884495	-	Torres-Garcia et al. (2022)
Penicillium silybi	CBS 147777	T	Aspergilloides	Exilicaulis	Restricta	USA	Milk thistle (Silybum marianum)	KF367458	MW774592	MW774591	AB860248	-	Labuda et al. (2021)
Penicillium soli	KUMCC 18-0202	T	Aspergilloides	Lanata-Divaricata	Janthinella	China	Soil	MT152337	MT161681	MT178249	MT384372	-	Doilom et al. (2020)
Penicillium stangiae	URM 8347	T	Aspergilloides	Lanata-Divaricata	Dalearum	Brazil	Soil	MW648590	MW646388	MW646390	MW646392	-	Alves et al. (2022a)
Penicillium subfuscum	CMW 56196	T	Aspergilloides	Lanata-Divaricata	Simplicissima	South Africa	Soil	MT949907	MT957412	MT957454	MT957480	-	Visagie & Yilmaz (2022)
	CN014A6		Aspergilloides	Lanata-Divaricata	Simplicissima	South Africa	Soil	MW329997	MW340969	MW340970	MW340971	-	Visagie & Yilmaz (2022)
Penicillium submersum	FMR 17140	T	Penicillium	Robsamsonia	Urticicola	Spain	Fluvial sediment	LR814194	LR814187	LR814188	LR814195	-	Torres-Garcia et al. (2022)
Penicillium taurinense	CBS 145672	T	Penicillium	Robsamsonia	Glandicolarum	Italy	Chestnut mill	MF595981	MF595977	MF595979	MT253108	-	Crous et al. (2020b)
	CBS 145673		Penicillium	Robsamsonia	Glandicolarum	Italy	Chestnut mill	MF595982	MF595978	MF595980	-	-	Crous et al. (2020b)
Penicillium tealii	BRIP 72734c	T	Aspergilloides	Cinnamopurpurea	Jiangxiensia	Australia	Dead spider	OP101639	OP039547	-	OP039546	-	Tan et al. (2022)
	BRIP 72735b		Aspergilloides	Cinnamopurpurea	Jiangxiensia	Australia	Dead spider	OP101642	OP039553	-	OP039552	-	Tan et al. (2022)
	BRIP 72742b		Aspergilloides	Cinnamopurpurea	Jiangxiensia	Australia	Dead spider	OP101643	OP039554	-	OP039546	-	Tan et al. (2022)
	BRIP 72731b		Aspergilloides	Cinnamopurpurea	Jiangxiensia	Australia	Dead spider	OP101641	OP039551	-	OP039555	-	Tan et al. (2022)
	BRIP 72732b		Aspergilloides	Cinnamopurpurea	Jiangxiensia	Australia	Dead spider	OP101640	OP039549	-	OP039559	-	Tan et al. (2022)
Penicillium tirolense	CBS 147625	T	Aspergilloides	Torulomyces	Torulomyces	Austria	Sporocarp of Serpula lacrymans	MW145398	MW143069	MW143068	MW143067	-	Kirchmair et al. (2022)
Penicillium tolerans	BRIP 64090a	T	Aspergilloides	Aspergilloides	Sclerotiorum	Australia	Soil	OK639006	OL741658	-	-	-	Tan & Shivas (2022)
Penicillium ucsense	CBS 146492	T	Aspergilloides	Lanata-Divaricata	Oxalica	Brazil	Intestinal tract of Anobium punctatum larva	OM914583	ON024157	ON024158	ON024159	-	Lenz et al. (2022)
Penicillium ulleungdoense	KACC 48990	T	Aspergilloides	Sclerotiorum	Sclerotiorum	Republic of Korea	Root of Phedimus takesimensis	MN640087	MN737487	MN745074	MN756007	-	Choi et al. (2020)
	KACC 48991		Aspergilloides	Sclerotiorum	Sclerotiorum	Republic of Korea	Root of Sedum oryzifolium	MN640088	MN737488	MN745075	MN756008	-	Choi et al. (2020)
	KACC 48992		Aspergilloides	Sclerotiorum	Sclerotiorum	Republic of Korea	Root of Aster spathulifolius	MN640089	MN737489	MN745076	MN756009	-	Choi et al. (2020)
Penicillium umkhoba	CMW 56199		Aspergilloides	Sclerotiorum	Hergueorum	South Africa	Soil	MT949911	MT957416	MT957458	MT957484	-	Visagie & Yilmaz (2022)
	CMW 56200	T	Aspergilloides	Sclerotiorum	Hergueorum	South Africa	Soil	MT949912	MT957417	MT957459	MT957485	-	Visagie & Yilmaz (2022)
	CMW 56201		Aspergilloides	Sclerotiorum	Hergueorum	South Africa	Soil	MT949913	MT957418	MT957460	MT957486	-	Visagie & Yilmaz (2022)
Penicillium uttarakhandense	NFCCI 4808	T	Aspergilloides	Lanata-Divaricata	Simplicissima	India	Soil (garden)	MN967315	MN972443	MN972445	MN972447	-	Crous et al. (2021a)
	NFCCI 4809		Aspergilloides	Lanata-Divaricata	Simplicissima	India	Soil (garden)	MN967316	MN972444	MN972446	MN972448	-	Crous et al. (2021a)
Penicillium vaccaeorum	CBS 110.64		Aspergilloides	Citrina	Roseopurpurea	Turkey	Soil	-	JN606829	JN606533	-	-	Torres-Garcia et al. (2022)
	CBS 118020		Aspergilloides	Citrina	Roseopurpurea	Canada	Ants	-	JN606832	JN606536	-	-	Torres-Garcia et al. (2022)
	CBS 118024		Aspergilloides	Citrina	Roseopurpurea	Canada	Ants	-	JN606833	JN606537	-	-	Torres-Garcia et al. (2022)
	CBS 127029		Aspergilloides	Citrina	Roseopurpurea	Argentina	Soil	-	JN606814	JN606544	-	-	Torres-Garcia et al. (2022)
	CBS 135118		Aspergilloides	Citrina	Roseopurpurea	South Africa	Soil	JX140867	JX141019	JX141510	MN418449	-	Torres-Garcia et al. (2022)
	CBS 135119		Aspergilloides	Citrina	Roseopurpurea	South Africa	Soil	JX140865	JX141020	JX141511	MK461491	-	Torres-Garcia et al. (2022)
	CBS 148.83	T	Aspergilloides	Citrina	Roseopurpurea	Spain	Soil	JN617689	JN606835	JN606543	JN606614	-	Torres-Garcia et al. (2022)
	CBS 300.67		Aspergilloides	Citrina	Roseopurpurea	The Netherlands	Soil	-	JN606841	JN606561	-	-	Torres-Garcia et al. (2022)
	CBS 441.88		Aspergilloides	Citrina	Roseopurpurea	Chile	Soil	-	JN606846	JN606568	-	-	Torres-Garcia et al. (2022)
	CBS 643.73		Aspergilloides	Citrina	Roseopurpurea	Canada	Soil	-	JN606853	JN606576	-	-	Torres-Garcia et al. (2022)
	CBS 644.73		Aspergilloides	Citrina	Roseopurpurea	Canada	Soil	-	JN606854	JN606577	-	-	Torres-Garcia et al. (2022)
	CBS 685.85		Aspergilloides	Citrina	Roseopurpurea	Spain	Soil	JN617711	JN606855	JN606578	-	-	Torres-Garcia et al. (2022)
	CV1865		Aspergilloides	Citrina	Roseopurpurea	South Africa	Soil	JX140866	JX141021	JX157340	MK461547	-	Torres-Garcia et al. (2022)
	FMR 17531		Aspergilloides	Citrina	Roseopurpurea	Spain	Fluvial sediment	LR814213	LR814203	LR814204	-	-	Torres-Garcia et al. (2022)
	FMR 17534		Aspergilloides	Citrina	Roseopurpurea	Spain	Fluvial sediment	OU375272	OU375168	OU375273	-	-	Torres-Garcia et al. (2022)
	FMR 17616		Aspergilloides	Citrina	Roseopurpurea	Spain	Fluvial sediment	LR814217	LR814212	LR814218	-	-	Torres-Garcia et al. (2022)
	FMR 17967		Aspergilloides	Citrina	Roseopurpurea	Spain	Fluvial sediment	LR814235	LR814226	LR814227	-	-	Torres-Garcia et al. (2022)
	FMR 18100		Aspergilloides	Citrina	Roseopurpurea	Spain	Fluvial sediment	LR814241	LR814234	LR814242	-	-	Torres-Garcia et al. (2022)
	FMR 18123		Aspergilloides	Citrina	Roseopurpurea	Spain	Fluvial sediment	LR814273	LR814265	LR814264	-	-	Torres-Garcia et al. (2022)
Penicillium vallebormidaense	CBS 147064	T	Aspergilloides	Exilicaulis	Erubescentia	Italy	Compost	MT316359	MW115862	MW115863	MW115864	-	Crous et al. (2020a)
Penicillium vickeryae	BRIP 72552a	T	Aspergilloides	Lanata-Divaricata	Simplicissima	Australia	Soil	OP903479	OP921966	-	OP921965	-	Tan & Shivas (2022)
Penicillium vietnamense	VTCC 930029	T	Aspergilloides	Charlesia	Indica	Vietnam	Sea water	MT102836	MT230561	ON209438	MT222288	-	Nguyen & Pham (2022)
Penicillium xyleborini	CMW 56800	T	Penicillium	Ramosum	Soppiorum	South Africa	Beetle (Xyleborinus saxesenii)	MW504356	MW480817	MW480823	MW480824	-	Visagie & Yilmaz (2022)
Rasamsonia oblata	IMI 288719	T	-	-	-	Australia	Spoiled baby food	LC546718	LC546729	LC546740	-	-	Yanai et al. (2020)
	CBS 258.87	T	-	-	-	Australia	Spoiled baby food	OR142403	OR145988	OR145994	-	-	Present study
Rasamsonia sabulosa	ATCC 56984	T	-	-	-	Australia	Spoiled fruit juice	LC546720	LC546726	LC546742	-	-	Yanai et al. (2020)
Talaromyces africanus	CBS 147340	T	-	Trachyspermi	-	South Africa	House dust	OK339610	OK338782	OK338808	OK338833	-	Pyrri et al. (2021)
Talaromyces albisclerotius	CBS 141839	T	-	Trachyspermi	-	China	Soil	MN864276	MN863345	MN863322	MN863334	-	Sun et al. (2020b)
Talaromyces aspriconidius	CBS 141835	T	-	Talaromyces	-	China	Soil	MN864274	MN863343	MN863320	MN863332	-	Sun et al. (2020b)
Talaromyces atkinsoniae	BRIP 72528a	T	-	Talaromyces	-	Australia	Gills of Marasmius crinisequi	OP059084	OP087524	-	OP087523	-	Tan et al. (2022)
Talaromyces aureolinus	AS 3.15864		-	Talaromyces	-	China	Soil	MK837954	MK837938	MK837946	MK837962	-	Wei et al. (2021)
	AS 3.15865		-	Talaromyces	-	China	Soil	MK837953	MK837937	MK837945	MK837961	-	Wei et al. (2021)
	NM6-1		-	Talaromyces	-	China	Soil	MN059095	MN059093	MN059094	MN059096	-	Wei et al. (2021)
Talaromyces bannicus	AS 3.15862	T	-	Talaromyces	-	China	Soil	MK837955	MK837939	MK837947	MK837963	-	Wei et al. (2021)
Talaromyces brevis	CBS 141833	T	-	Talaromyces	-	China	Soil	MN864269	MN863338	MN863315	MN863328	-	Sun et al. (2020b)
	CBS 118436		-	Talaromyces	-	Marocco	Soil	MN864271	MN863340	MN863317	MN863330	-	Sun et al. (2020b)
	DTO 307-C1		-	Talaromyces	-	Turkey	Soil	MN864270	MN863339	MN863316	MN863329	-	Sun et al. (2020b)
Talaromyces calidominioluteus	CBS 113167		-	Trachyspermi	-	Unknown	Air (indoor)	OK339611	OK338785	OK338816	OK338836	-	Pyrri et al. (2021)
	CBS 147313	T	-	Trachyspermi	-	The Netherlands	Melon (imported from Brazil)	OK339612	OK338786	OK338817	OK338837	-	Pyrri et al. (2021)
	CBS 147341		-	Trachyspermi	-	Iran	Grapevine	OK339602	OK338788	KU711896	OK338839	-	Pyrri et al. (2021)
	CBS 147342		-	Trachyspermi	-	Thailand	House dust	OK339600	KP330045	OK338815	OK338835	-	Pyrri et al. (2021)
	DTO 265-H8		-	Trachyspermi	-	Iran	Grapevine	OK339601	OK338787	KU711894	OK338838	-	Pyrri et al. (2021)
	DTO 266-A5		-	Trachyspermi	-	Iran	Grapevine	OK339603	OK338789	KU711900	OK338840	-	Pyrri et al. (2021)
	DTO 269-H1		-	Trachyspermi	-	Thailand	House dust	KJ775721	KJ775214	-	-	-	Pyrri et al. (2021)
	DTO 269-H4		-	Trachyspermi	-	Thailand	House dust	KJ775722	KJ775215	-	-	-	Pyrri et al. (2021)
	DTO 270-A1		-	Trachyspermi	-	Thailand	House dust	KJ775728	KJ775221	-	-	-	Pyrri et al. (2021)
	DTO 270-C3		-	Trachyspermi	-	Thailand	House dust	KJ775733	KJ775226	-	-	-	Pyrri et al. (2021)
	DTO 390-E9		-	Trachyspermi	-	Nigeria	Cocoa beans	MN788104	MN787900	MN787896	OK338847	-	Pyrri et al. (2021)
	DTO 390-F1		-	Trachyspermi	-	Nigeria	Cocoa beans	MN788103	MN787901	MN787895	OK338848	-	Pyrri et al. (2021)
	DTO 390-I9		-	Trachyspermi	-	Nigeria	Cocoa beans	MN788115	MN787911	MN787885	OK338849	-	Pyrri et al. (2021)
	DTO 391-A5		-	Trachyspermi	-	Nigeria	Cocoa beans	MN788111	MN787914	MN787883	OK338850	-	Pyrri et al. (2021)
Talaromyces cavernicola	URM 8448	T	-	Talaromyces	-	Brazil	Air (cave)	ON862935	OP672383	OP290543	OP290515	-	Alves et al. (2022b)
	URM 8449		-	Talaromyces	-	Brazil	Air (cave)	ON862936	OP672384	OP290544	OP290516	-	Alves et al. (2022b)
Talaromyces chongqingensis	CBS 270.35		-	Trachyspermi	-	China	Soil	OK339609	OK338781	OK338807	OK338832	-	Zhang et al. (2021b)
	CGMCC 3.20482	T	-	Trachyspermi	-	China	Soil	MZ358001	MZ361343	MZ361350	MZ361357	-	Zhang et al. (2021b)
	CS26-63		-	Trachyspermi	-	China	Soil	MZ358002	MZ361344	MZ361351	MZ361358	-	Zhang et al. (2021b)
	CS26-73		-	Trachyspermi	-	China	Soil	MZ358003	MZ361345	MZ361352	MZ361359	-	Zhang et al. (2021b)
	CS26-75		-	Trachyspermi	-	China	Soil	MZ358004	MZ361346	MZ361353	MZ361360	-	Zhang et al. (2021b)
	NRRL 1064		-	Trachyspermi	-	China	Soil	KM066172	KM066129				Zhang et al. (2021b)
Talaromyces gaditanus	CBS 104.71		-	Trachyspermi	-	The Netherlands	Tulip	OK339614	OK338792	OK338820	OK338852	-	Pyrri et al. (2021)
	CBS 144771		-	Trachyspermi	-	The Netherlands	Sputum of cystic fibroses patient	OK339616	OK338794	OK338822	OK338842	-	Pyrri et al. (2021)
	CBS 169.81	T	-	Trachyspermi	-	Spain	Air	MH861318	OK338775	OK338802	OK338827	-	Pyrri et al. (2021)
	CBS 442.89		-	Trachyspermi	-	Denmark	Soil	OK339615	OK338793	OK338821	OK338853	-	Pyrri et al. (2021)
	CBS 444.89		-	Trachyspermi	-	Denmark	Cranberry (imported from USA)	OK339597	OK338776	OK338803	OK338828	-	Pyrri et al. (2021)
	CBS 996.72		-	Trachyspermi	-	The Netherlands	Jute sugar bag	MH860641	OK338774	OK338813	OK338826	-	Pyrri et al. (2021)
	CBS 138.84		-	Trachyspermi	-	Spain	Apple (Malus sylvestris)	OK339604	OK338791	OK338819	OK338851	-	Pyrri et al. (2021)
Talaromyces germanicus	CBS 147314	T	-	Trachyspermi	-	Germany	Indoor environment	OK339619	OK338799	OK338812	OK338845	-	Pyrri et al. (2021)
Talaromyces ginkgonis	CGMCC 3.20698	T	-	Talaromyces	-	China	Fruit of Ginkgo biloba	OL638158	OL689844	OL689846	OL689848	-	Wang & Zhuang (2022a)
Talaromyces gwangjuensis	CNUFC WT19-1	T	-	Purpurei	-	Republic of Korea	Freshwater	MK766233	MZ318448	-	MK912174	-	Nguyen et al. (2021a)
	CNUFC WT19-2		-	Purpurei	-	Republic of Korea	Freshwater	MK766234	MZ318449	-	MK912175	-	Nguyen et al. (2021a)
Talaromyces haitouensis	AS 3.16101	T	-	Talaromyces	-	China	Soil	MZ045695	MZ054634	MZ054637	MZ054631	-	Han et al. (2021)
Talaromyces koreanus	CNUFC YJW2-13	T	-	Helici	-	Republic of Korea	Freshwater	MZ315100	MZ318450	MZ332529	MZ332533	-	Nguyen et al. (2021a)
	CNUFC YJW2-14		-	Helici	-	Republic of Korea	Freshwater	MZ315101	MZ318451	MZ332530	MZ332534	-	Nguyen et al. (2021a)
Talaromyces nanjingensis	CCTCC-M-2012167	T	-	Talaromyces	-	China	Soil	MW130720	MW147759	MW147760	MW147762	-	Sun et al. (2022b)
Talaromyces peaticola	CGMCC 3.18620	T	-	Trachyspermi	-	China	Soil (peat)	MF135613	MF284705	MF284703	MF284704	-	Tian et al. (2021)
	CGMCC 3.18767		-	Trachyspermi	-	China	Soil (peat)	MF960857	MF960859	MF960861	MF960863	-	Tian et al. (2021)
	CGMCC 3.18768		-	Trachyspermi	-	China	Soil (peat)	MF960858	MF960860	MF960862	MF960864	-	Tian et al. (2021)
Talaromyces penicillioides	AS 3.15822	T	-	Talaromyces	-	China	Soil	MK837956	MK837940	MK837948	MK837964	-	Wei et al. (2021)
Talaromyces pernambucoensis	URM 6894	T	-	Trachyspermi	-	Brazil	Soil	LR535947	LR535945	LR535946	LR535948	-	Crous et al. (2019)
Talaromyces phuphaphetensis	TBRC 16281	T	-	Trachyspermi	-	Thailand	Soil (cave)	ON692803	ON706960	ON706962	ON706964	-	Nuankaew et al. (2022)
Talaromyces pulveris	CBS 146831	T	-	Purpurei	-	France	Bore dust of deathwatch beetle (Xestobium rufovillosum) infesting floorboards (Quercus wood)	MW175345	MW173136	MW173099	MW173115	-	Crous et al. (2020a)
Talaromyces resedanus	CBS 181.71	T	-	Subinflati	-	Seychelles	Soil	MN864280	MN863349	MN863326	MN969214	-	Sun et al. (2020b)
Talaromyces rosarhiza	GUCC 190040.1	T	-	Talaromyces	-	China	Endophyte of Rosa roxburghii	MZ221603	MZ333143	MZ333137	MZ333141	-	Zhang et al. (2021a)
	GUCC 197011.1		-	Talaromyces	-	China	Endophyte of Rosa roxburghii	MZ221604	MZ333144	MZ333138	MZ333142	-	Zhang et al. (2021a)
Talaromyces rufus	CBS 141834	T	-	Talaromyces	-	China	Soil	MN864272	MN863341	MN863318	MN863331	-	Sun et al. (2020b)
	DTO 274-C5		-	Talaromyces	-	Republic of Korea	Soil	MN864273	MN863342	MN863319	-	-	Sun et al. (2020b)
Talaromyces samsonii	CBS 137.84	T	-	Trachyspermi	-	Spain	Apple (Malus sylvestris)	MH861709	OK338798	OK338824	OK338844	-	Pyrri et al. (2021)
	CBS 147356		-	Trachyspermi	-	The Netherlands	Soil	OK339598	OK338777	OK338804	OK338829	-	Pyrri et al. (2021)
	CBS 147357		-	Trachyspermi	-	Greece	Air (indoor)	OK339599	OK338778	OK338805	OK338830	-	Pyrri et al. (2021)
Talaromyces santanderensis	HF05	T	-	Talaromyces	-	Colombia	Soil (cacao field)	OP082331	OP067657	OP067656	OP067655	-	Guerra Sierra et al. (2022)
Talaromyces satunensis	TBRC 16246	T	-	Trachyspermi	-	Thailand	Soil (cave)	ON692804	ON706961	ON706963	-	-	Nuankaew et al. (2022)
Talaromyces saxoxalicus	MUM20.30	T	-	Purpurei	-	Portugal	Limestone walls	MT039882	MT052003	-	MT052004	-	Trovao et al. (2021)
Talaromyces shilinensis	CGMCC 3.20699	T	-	Talaromyces	-	China	Rotten twig	OL638159	OL689845	OL689847	OL689849	-	Wang & Zhuang (2022a)
Talaromyces sparsus	AS 3.15880	T	-	Talaromyces	-	China	Soil	MK837958	MK837942	MK837950	MK837966	-	Wei et al. (2021)
Talaromyces teleomorphus	CNUFC YJW2-5	T	-	Helici	-	Republic of Korea	Freshwater	MZ315102	MZ318452	MZ332531	MZ332535	-	Nguyen et al. (2021a)
	CNUFC YJW2-6		-	Helici	-	Republic of Korea	Freshwater	MZ315103	MZ318453	MZ332532	MZ332536	-	Nguyen et al. (2021a)
Talaromyces tenuis	CBS 141840	T	-	Tenues	-	China	Soil	MN864275	MN863344	MN863321	MN863333	-	Sun et al. (2020b)
Talaromyces wushanicus	CGMCC 3.20481	T	-	Talaromyces	-	China	Soil	MZ356356	MZ361347	MZ361354	MZ361361	-	Zhang et al. (2021b)
	CS17-04		-	Talaromyces	-	China	Soil	MZ356357	MZ361348	MZ361355	MZ361362	-	Zhang et al. (2021b)
	CS17-06		-	Talaromyces	-	China	Soil	MZ356358	MZ361349	MZ361356	MZ361363	-	Zhang et al. (2021b)
Talaromyces yunnanensis	KUMCC 18-0208	T	-	Talaromyces	-	China	Soil	MT152339	MT161683	MT178251	-	-	Doilom et al. (2020)
Talaromyces zhenhaiensis	AS 3.15693		-	Talaromyces	-	China	Soil	KY007094	KY007110	KY007102	KY112592	-	Han et al. (2021)
	AS 3.16102	T	-	Talaromyces	-	China	Soil	MZ045697	MZ054636	MZ054639	MZ054633	-	Han et al. (2021)

Datasets were aligned in MAFFT v. 7.490 (Katoh & Standley 2013) using the G-INS-i option. For multigene phylogenies, datasets were concatenated in Geneious Prime v. 2023.0.1 (Biomatters, NZ). Partitionfinder v. 2.1.1 (Lanfear et al. 2017) was used to select the partitioning schemes and nucleotide substitution models for each alignment, with exons, introns, and codon positions treated as independent datasets. Maximum Likelihood phylogenies were computed in IQ-TREE v. 2.1.3 (Minh et al. 2020) with support in nodes calculated using UFBoot (Hoang et al. 2018) ultrafast bootstrapping with 1 000 replicates, as implemented in IQ-TREE. Phylogenies were visualised in TreeViewer v. 2.0.1 (https://treeviewer.org) and further edited for publication in Affinity Publisher v. 2.0.3 [Serif (Europe) Ltd, Nottingham, UK]. Here we use a phylogenetic species concept and apply Genealogical Concordance Phylogenetic Species Recognition (GCPSR; Taylor et al. 2000).

Morphology

Descriptions were made following the recommendations of Samson et al. (2014) and Visagie et al. (2014). Strains were inoculated at three-equidistant points onto Czapek yeast autolysate agar (CYA), CYA with 5 % NaCl (CYAS), CYA with 20 % sucrose (CY20S), malt extract agar (MEA), yeast extract sucrose agar (YES), dichloran 18 % glycerol agar (DG18), oatmeal agar (OA), and Creatine sucrose agar (CREA). These were incubated at 25 °C for 7 d, with additional CYA plates incubated at 20, 30 and 37 °C. A subset of plates were incubated for longer periods to observe possible sexual states. After incubation, the species were characterised based on their growth rates, colony characteristics and microscopic features. Colonies were photographed using a Sony A6400 camera and a Sony SEL50M28 lens (Tokyo, Japan). Microscopic observations were made using a Nikon SMZ25 stereomicroscope (Tokyo, Japan) and Zeiss AXIO Imager.A2 compound microscope (Carl Zeiss CMP, Göttingen, Germany), both equipped with Nikon DS-Ri2 cameras and using Nikon Elements D v. 5.11 software. The photoplates were created in Affinity Photo v. 2.0.3 [Serif (Europe), Nottingham, UK].

RESULTS

Phylogenetic analyses

All alignments, partitioning schemes, model tests, and phylogenetic trees were deposited in the University of Pretoria research data repository hosted on Figshare (doi: 10.25403/UPresearchdata.23723277). The phylogenies largely confirmed the novelty of the newly introduced species, with some exceptions commented on using notes in the species list below (Figs 1–33, Suppl. Figs S1–S30).

Fig. 1.

Phylogenetic tree of Aspergillus section Aenei series Aenei based on a concatenated dataset of BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. The tree was rooted to A. versicolor. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by superscript T. See Suppl. Fig. S1.

Fig. 33.

Phylogenetic tree of Talaromyces section Trachyspermi, based on a concatenated dataset of BenA, CaM and RPB2. Strains of recently described species are shown in bold coloured text. The tree was rooted to T. purpureus. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by superscript T.

TAXONOMY

Here we present new additions and changes to the list of accepted species published in Houbraken et al. (2020) and provide an overview of the 160 species described (51 Aspergillus, three Emericella, one Neosartorya, two Paecilomyces, one Paraxerochrysium, 64 Penicillium, two Rasamsonia, 35 Talaromyces and one Xerochrysium) and the new genus Paraxerochrysium. At the continental level, species were described from Africa (n = 28), Asia (n = 57), Europe (n = 36), North America (n = 15), Oceania (n = 21) and South America (n = 19).

These originated from 42 countries with most coming from China (n = 39), Australia (n = 20), South Africa (n = 16), Brazil (n = 16), Spain (n = 12), and the USA (n = 8). Species were described from a wide range of substrates, mainly soil (n = 91), plant material (n = 15), food and feed (n = 14), air (n = 12) and indoor environments (n = 8). Of the 160, 22 were classified as synonyms and four as doubtful species. Ten were invalidly described. Two of these names are validated below, the remaining eight are considered synonyms or doubtful species. We consider the four combinations introduced by Pitt & Hocking (2022) contradictory to our view on Aspergillus and the sexual (teleomorphic) genera associated with it (Samson et al. 2014, Kocsube et al. 2016). We also document species that were previously accepted but then subsequently reduced as synonyms due to various taxonomic revisions. Finally, species accepted in Houbraken et al. (2020) for which no DNA sequence data are available are now listed as doubtful. Taking these changes into account, we currently accept 453 Aspergillus, 12 Paecilomyces, 535 Penicillium, 14 Rasamsonia, 203 Talaromyces and four Xerochrysium species.

Aspergillus series Annuorum J.J. Silva, Iamanaka & Frisvad ser. nov. MycoBank MB 849339. Fig. 6 & Suppl. Fig. S6.

Fig. 6.

Phylogenetic tree of Aspergillus section Flavi series Alliacei and Flavi based on a concatenated dataset of BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. The tree was midpoint rooted. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by superscript T. Representative strains are indicated by superscript R. See Suppl. Fig. S6.

Synonym: Aspergillus ser. Annuorum [nom. inval. Art. 40.1 (Shenzhen)] J.J. Silva, Fungaro, Frisvad, Taniwaki & Iamanaka, J. Fungi 8 (no 1279): 14. 2022. MycoBank MB 845971.

In: Aspergillus subgen. Circumdati sect. Flavi.

Typus: Aspergillus annui J.J. Silva, Fungaro, Frisvad, Taniwaki & Iamanaka [MB 849336].

Etymology: Named after Aspergillus annui.

Description: See Silva et al. (2022).

Aspergillus annui J.J. Silva, Fungaro, Frisvad, Taniwaki & Iamanaka, sp. nov. MycoBank MB 849336. Fig. 6 & Suppl. Fig. S6.

Synonym: Aspergillus annui [nom. inval. Art. 40.7 & Art. 40.8 (Shenzhen)] J.J. Silva, Fungaro, Frisvad, Taniwaki & Iamanaka, J. Fungi 8 (no 1279): 14. 2022. MycoBank MB 845969.

In: Aspergillus subgen. Circumdati sect. Flavi ser. Annuorum.

DNA barcodes: ITS = OP691228; BenA = ON529842; CaM = ON529841; RPB2 = ON529843.

Etymology: The specific epithet refers to the substrate from which it was isolated, paprika pepper (Capsicum annuum).

Typus: Brazil, São Paulo State, São Paulo City, 23°35’29.7”S 46°40’52.1”W, sweet paprika pepper (Capsicum annuum), 6 Apr. 2017, isol. C.A. Yasumura (holotype IBT 36122 preserved as a metabolically inactive culture, culture ex-type 365-IT-PPK = IBT 36122).

Description: See Silva et al. (2022).

Notes: This species was invalidly described because Silva et al. (2022) cited two holotypes and failed to mention that the holotype was metabolically inactive. This also means that series Annuorum introduced for A. annui is invalid. Here we validate both the species and the series.

Aspergillus saccharicola J.J. Silva, Frisvad, Fungaro, Taniwaki & Iamanaka, sp. nov. MycoBank MB 849338. Fig. 6 & Suppl. Fig. S6.

Synonym: Aspergillus saccharicola [nom. inval. Art. 40.7 & Art. 40.8 (Shenzhen)] J.J. Silva, Frisvad, M.H.P. Fungaro, M.H. Taniwaki & B.T. Iamanaka, J. Fungi 8 (no 1279): 16. 2022. MycoBank MB 845970.

In: Aspergillus subgen. Circumdati sect. Flavi ser. Flavi.

DNA barcodes: ITS = OP611470; BenA = ON529845; CaM = ON529844; RPB2 = ON529846.

Etymology: The specific epithet refers to the substrate from which it was isolated, sugarcane (Saccharum officinarum).

Typus: Brazil, São Paulo State, São Paulo City, 23°35’29.7”S 46°40’52.1”W, sugarcane (Saccharum officinarum), 14 Sep. 2011, isol. B.T. Imanaka (holotype IBT 36126 preserved as a metabolically inactive culture, culture ex-type 117-IT-SGC = IBT 36126).

Description: See Silva et al. (2022).

Notes: This species was invalidly described because Silva et al. (2022) cited two holotypes and failed to mention that the holotype was metabolically inactive. Here we validate the species.

Aspergillus section Dichlaena Visagie, Kocsubé & Houbraken, sect. nov. MycoBank MB 849488. Fig. 14.

Fig. 14.

Phylogenetic tree showing the relationship of Dichlaena within Aspergillus based on a concatenated dataset of BenA, CaM, RPB2, RPB1, ITS and LSU. Strains of recently described species are shown in bold coloured text. The tree was rooted to Trichocoma paradoxa. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by superscript T. Representative strains are indicated by superscript R.

In: Aspergillus subgenus Circumdati.

Etymology: Named after the genus Dichlaena.

Typus: Aspergillus lentisci (Durieu & Mont.) Visagie, Malloch, L. Kriegsteiner, Samson & Houbraken [MB 849485].

Description: We follow the Malloch & Cain (1972) description of Dichlaena and apply this to the section.

Notes: The genus Dichlaena is synonymised with Aspergillus. Aspergillus lentisci (bas. Dichlaena lentisci) strains resolve as a distinct clade, with section Petersoniorum (subg. Circumdati) members being the closest relatives (Fig. 14). Aspergillus lentisci strains are strongly xerophilic, similar to Aspergillus sections Aspergillus and Restricti, while section Petersoniorum species also grow well on low water activity media. We thus introduce this as a new section. The section is based on the single species, Aspergillus lentisci. The sexual state for this xerophilic species has not been observed in culture, but was documented in detail by Malloch & Cain (1972) who observed it from leaves, twigs and dung. Three other species have been described in Dichlaena. Dichlaena bovina [MB 329872] was considered a synonym of Thielavia bovina [MB 240118] by Malloch & Cain (1973), while we consider Dichlaena indica [MB 127024] and Dichlaena pterodontis [MB 312952] doubtful species with no material available for study.

Aspergillus lentisci (Durieu & Mont.) Visagie, Malloch, L. Kriegsteiner, Samson & Houbraken comb. nov. MycoBank MB 849485. Figs 34 & 35.

Fig. 34.

Aspergillus lentisci in nature. A, B. Ascomata on dried leaf material. C, D. Ascomata observed under light microscope. E. Outer cell layer of ascomata. F. Ascospores released from ascomata. G. Ascospores. Scale bars = 10 µm.

Fig. 35.

Aspergillus lentisci in culture. A. Colonies on, from left to right, MY50G, MY20, MY40, DG18 and OA. B–D. Colony overview on DG18 showing sclerotia and conidiophores (D). E, F. Section of sclerotia showing potential early development of asci and ascomata. G. Section of sclerotia showing its wall structure. H. Hülle cells from sclerotia. I. Conidia. J–M. Conidiophores. Scale bars: B = 2 mm; C, D = 500 µm; E = 50 µm; F–H = 20 µm; I–M = 10 µm.

Basionym: Dichlaena lentisci Durieu & Mont., Expl. Sci. l’Algérie 1 (11): 405. 1849. MycoBank MB 249716.

In: Aspergillus subgen. Circumdati sect. Dichlaena

DNA barcodes: ITS = OR142402; BenA = OR145977; CaM = OR145992; RPB2 = OR146003; LSU = OR142413.

Typus: Durieu & Mont., Expl. Sci. l’Algérie 1 (11): 405, tab. 22 bis, fig. 2, lectotype designated here, MBT10014202. Portugal, Aljezur, Pistacia leaf, Apr. 2019, coll. L. Kriegsteiner, isol. R.A. Samson [epitype designated here, CBS H-25292 (dried culture of CBS 150189), MBT10014196, culture ex-epitype CBS 150189 = DTO 426-E9].

Additional strains examined: Portugal, Aljezur, Pistacia leaf, Apr. 2019, coll. L. Kriegsteiner, isol. R.A. Samson, cultures DTO 426-F1, DTO 426-F2, and DTO 426-F3.

Colony diam, 25 °C (7 d, in mm): CYA no growth; MEA no growth; OA no growth; YES no growth; MY20 no growth; MY40 microcolonies–6; MY50G 5–10. After 14 d (in mm): CYA no growth; MEA no growth; OA microcolonies–6; YES no growth; MY20 no growth to microcolonies; MY40 9–17; MY50G 13–20; DG18 5–13; CREA no growth.

Colony characters (25 °C, 14 d): DG18, colonies floccose; mycelial areas white to yellowish; ascocarp bearing stromata light yellow to orange yellow to greyish yellow (4A4–6–B6), globose, sclerotiod, 600–1 200(−1 900) µm; sporulation absent to sparse, becoming moderate with prolonged incubation, white to cream; exudate sometimes present, clear; soluble pigment brown, inconspicuous; reverse yellowish white (4A2) to light brown to yellowish brown to brown (5D7–E8–6E8). MY50G, colonies floccose; mycelial areas white to yellowish; ascocarp bearing stromata light yellow to orange yellow to greyish yellow (4A4–6–B6), globose, sclerotiod, 600–1 200(−1 900) µm; sporulation sparse, white to cream; exudate absent; soluble pigment absent; reverse yellowish white (4A2) to greyish orange (5B6). MY40, colonies floccose; mycelial areas white to yellowish; sporulation absent; exudate absent; soluble pigment absent.

Micromorphology: Conidial heads radiate; conidiophores uniseriate; stipes hyaline, minor proportion pigmented, smooth, 200–830 × 4–8 μm; vesicles subglobose, sometimes clavate, phialides cover 50–75 % of head, 11–22(−32) μm wide; phialides ampulliform, 6.5–13 × 2.5–4(−4.5) μm; conidia globose to subglobose, some tapering towards one end, rough to echinulate, 3.5–5 × 3–4.5 (4.25 ± 0.27 × 3.88 ± 0.27, n = 50) μm, length/width 0.91 ± 0.1. Ascomata not produced in culture, in nature fide Malloch & Cain (1972), ‘arising from coiled or twisted initials at one to three points in the stroma, completely filling the stroma at maturity, irregular in shape, colorless to pale yellow; stroma tissue hard and stony, of two types when young; stromal cells of the outer tissue pale yellow, very thick-walled, pyriform to globose, often nearly open at one end, about 5–13 µm in diameter; stromal cells of the inner tissue hyaline, very thick-walled (but thinner-walled than those of the outer tissue), irregular in shape, usually elongated, often greater than 40 µm length, up to 15 µm broad; ascocarp peridium hyaline to pale yellow, consisting of a filamentous tissue with a membranous “backing”; asci irregularly disposed, ellipsoidal, short-stipitate, eight-spored, evanescent, 9 × 7 µm [fide Von Höhnel (1910)]; ascospores oblate, hyaline, smooth, with an inconspicuous equatorial band, 3.0–4.2 × 2.0–3.5 µm’.

Rasamsonia oblata (Pitt & A.D. Hocking) Yanai & Udagawa, Jap. J. Mycol. 61: 93. 2020. MycoBank MB 836491. Figs 28, 36 & Suppl. Fig. S26.

Fig. 28.

Phylogenetic tree of Rasamsonia, based on a concatenated dataset of BenA, CaM and ITS. Strains of recently described species are shown in bold coloured text. The tree was rooted to Trichocoma paradoxa. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by superscript T. See Suppl. Fig. S26.

Fig. 36.

Rasamsonia oblata. A. Colonies on, from left to right, CYA, MEA, YES, DG18 and OA. B–D. Colony overview on MEA (B) and OA (C, D). E–I. Conidiophores. J. Conidia. Scale bars: B = 2.5 mm; C, D = 1 mm; E–J = 10 µm.

Basionym: Penicillium oblatum Pitt & A.D. Hocking, Mycologia 77: 810. 1985. MycoBank MB 104603.

DNA barcodes: ITS = LC546718; BenA = LC546729; CaM = LC546740.

Typus: Australia, New South Wales, Sydney, from spoiled baby food, 1979, isol. A.D. Hocking (holotype FRR 2234, culture ex-type CBS 258.87 = IMI 288719 = NBRC 33091).

Colony diam, 25 °C (7 d, in mm): CYA 20 °C 8–9; CYA 8–9; CYA 30 °C 10–12; CYA 37 °C 4–5; CYAS no growth; CREA no growth; DG18 14–15; MEA 29–32; OA 22–23; YES 9–10.

Colony characters: CYA 25 °C, 7 d: Colonies low, plane; margins low, narrow, entire; mycelia white; texture funiculose; sporulation very sparse, conidia en masse not determined; soluble pigments absent; exudates absent; reverse pale yellow (3A2). MEA 25 °C, 7 d: Colonies deep, plane; margins subsurface, wide; mycelia white; texture funiculose; sporulation sparse to moderately dense, conidia en masse Brown to Dark Blonde (5D4); soluble pigments absent; exudates absent. YES 25 °C, 7 d: Colonies deep, plane; margins moderately deep, narrow, entire; mycelia white; texture funiculose; sporulation very sparse, conidia en masse not determined; soluble pigments absent; exudates absent; reverse pale yellow (4A2). DG18 25 °C, 7 d: Colonies low, plane; margins low, narrow, entire; mycelia white; texture funiculose; sporulation very sparse, conidia en masse not determined; soluble pigments absent; exudates absent; reverse pale yellow (3A2). OA 25 °C, 7 d: Colonies low, plane; margins subsurface, wide; mycelia white; texture funiculose; sporulation sparse to moderately dense, conidia en masse Brown to Dark Blonde (5D4); soluble pigments absent; exudates absent.

Micromorphology: Conidiophores monoverticillate, small proportion biverticillate; stipes rough walled, some finely roughened, nonvesiculate, 10–28 × 2.5–3 μm; metulae 2–3 per stipe, 9.5–14 × 2.5–3 μm; phialides acerose, in verticils of 3–6, smooth to roughened, 7–8.5(−9.5) × 2–3 μm; conidia smooth, broadly ellipsoid, some subglobose, 2–2.5 × 1.5–2 μm (2.14 ± 0.1 × 1.67 ± 0.1), average width/length = 0.78, n = 50, connected in chains with distinct connectors, connected on short end.

Notes: Pitt & Hocking (1985a) described Penicillium oblatum and referred to it as an interface species, noting that its ‘rough-walled stipes bearing appressed monoverticillate penicilli and its brown, often oblate conidia distinguish it from other Penicillia’, but that its conidial and phialide shape preclude it from Paecilomyces or Geosmithia. Both Samson et al. (2011) and Yilmaz et al. (2014) mentioned that the ex-type strain (CBS 258.87) was contaminated and was not representative of the original description. Recently, Yanai et al. (2020) analysed sequences generated for P. oblatum (ex-type strains IMI 288719 and NBRC 33091) and P. sabulosum (ex-type strain ATCC 56984; CBS 261.87 is contaminated), showed that they belong to Rasamsonia, and provided new combinations as R. oblata and R. sabulosa. They also provided new morphological descriptions in Japanese. We were able to purify CBS 258.87 and provide an English description above based on this strain.

Xerochrysium coryli (Crous & Decock) Visagie & Houbraken, comb. nov. MycoBank MB 849335. Fig. 16.

Fig. 16.

Phylogenetic tree showing the relationship of Xerochrysium coryli (bas. Paraxerochrysium coryli) within Aspergillaceae based on a concatenated dataset of BenA, RPB2, ITS and LSU. Strains of recently described species are shown in bold coloured text. The tree was rooted to Talaromyces amestolkiae. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by superscript T.

Basionym: Paraxerochrysium coryli Crous & Decock, Persoonia 47: 261. 2021. MycoBank MB 841830.

DNA barcodes: ITS = OK664748; BenA = OK651216; RPB2 = OK651178.

Typus: Belgium, Louvain-la-Neuve, from dry hazelnut (Corylus avellana), Feb. 2021, C. Decock (holotype CBS H-24853, culture ex-type CBS 148314 = CPC 41272 = MUCL 58103).

Description: See Crous et al. (2021b).

Notes: A multigene phylogenetic analysis of Aspergillaceae based on BenA, RPB2, ITS and LSU showed that Paraxerochrysium coryli resolves as a unique species inside Xerochrysium (Fig. 16). We thus synonymise Paraxerochrysium with Xerochrysium and provide a new combination here.

Current Eurotiales families, genera and subgeneric classifications (* indicates recently introduced sections or series)

• Aspergillaceae

  • Aspergillago

  • Aspergillus

    • subgenus Aspergillus

      • section Aspergillus

        • series Aspergillus, Chevalierorum, Leucocarpi, Rubri, Tamarindosolorum, Teporium and Xerophili

      • section Restricti

        • series Halophilici, Penicillioides, Restricti and Vitricolarum

    • subgenus Circumdati

      • section Candidi

        • series Candidi

      • section Circumdati

        • series Circumdati, Sclerotiorum and Steyniorum

      • section Dichlaena*

      • section Flavi

        • series Alliacei, Annuorum*, Avenacei, Bertholletiarum, Coremiiformes, Flavi, Kitamyces, Leporum and Nomiarum

      • section Flavipedes

        • series Flavipedes, Neonivei, Olivimuriarum and Spelaei

      • section Janorum

        • series Janorum

      • section Nigri

        • series Carbonarii, Heteromorphi, Homomorphi, Japonici and Nigri

      • section Petersoniorum

        • series Petersoniorum

      • section Robusti

        • series Robusti

      • section Tannerorum

        • series Tannerorum

      • section Terrei

        • series Ambigui, Nivei and Terrei

    • subgenus Cremei

      • section Cremei

        • series Arxiorum, Brunneouniseriati, Cremei, Inflati, Pulvini and Wentiorum

    • subgenus Fumigati

      • section Cervini

        • series Acidohumorum and Cervini

      • section Clavati

        • series Clavati

      • section Fumigati

        • series Brevipedes, Fennelliarum, Fumigati, Neoglabri, Spathulati, Thermomutati, Unilaterales and Viridinutantes

      • section Vargarum

        • series Vargarum

    • subgenus Nidulantes

      • section Aenei

        • series Aenei

      • section Bispori

        • series Bispori

      • section Cavernicolarum

        • series Cavernicolarum, Egyptiaci and Hainanici*

      • section Nidulantes

        • series Aurantiobrunnei, Multicolores, Nidulantes, Speluncei, Stellati, Unguium and Versicolores

      • section Ochraceorosei

        • series Funiculosi and Ochraceorosei

      • section Raperorum

        • series Raperorum

      • section Silvatici

        • series Silvatici

      • section Sparsi

        • series Biplani, Conjuncti, Implicati and Sparsi

      • section Usti

        • series Calidousti, Deflecti, Monodiorum and Usti

    • subgenus Polypaecilum

      • section Polypaecilum

        • series Canini, Kalimarum, Noonimiarum, Polypaecilum, Salinarum and Whitfieldiorum

  • Evansstolkia

  • Hamigera

  • Leiothecium

  • Monascus

  • Paraxerochrysium

  • Penicilliopsis

  • Penicillium

    • subgenus Aspergilloides

      • section Alfrediorum

        • series Alfrediorum

      • section Aspergilloides

        • series Fortuita, Glabra, Hoeksiorum, Improvisa, Kiamaensia, Livida, Longicatenata, Pinetorum, Quercetorum, Saturniformia, Spinulosa, Sublectatica, Thiersiorum, Thomiorum and Verhageniorum

      • section Charlesia

        • series Costaricensia, Fellutana, Indica and Phoenicea

      • section Cinnamopurpurea

        • series Cinnamopurpurea, Idahoensia, Jiangxiensia and Nodula

      • section Citrina

        • series Citrina, Copticolarum, Euglauca, Gallaica, Paxillorum, Roseopurpurea, Sheariorum, Sumatraensia, Vascosobrinhoana* and Westlingiorum

      • section Crypta

        • series Crypta

      • section Eremophila

        • series Eremophila

      • section Exilicaulis

        • series Alutacea, Citreonigra, Corylophila, Erubescentia, Lapidosa and Restricta

      • section Gracilenta

        • series Angustiporcata, Estinogena, Gracilenta and Macrosclerotiorum

      • section Griseola

        • series Griseola

      • section Inusitata

        • series Inusitata

      • section Lanata-Divaricata

        • series Dalearum, Janthinella, Oxalica, Rolfsiorum and Simplicissima

      • section Lasseniorum

        • series Lasseniorum

      • section Ochrosalmonea

        • series Ochrosalmonea

      • section Ramigena

        • series Georgiensia and Ramigena

      • section Sclerotiorum

        • series Adametziorum, Herqueorum and Sclerotiorum

      • section Stolkia

        • series Stolkia

      • section Thysanophora

        • series Thysanophora

      • section Torulomyces

        • series Torulomyces

    • subgenus Penicillium

      • section Brevicompacta

        • series Brevicompacta, Buchwaldiorum, Olsoniorum and Tularensia

      • section Canescentia

        • series Atroveneta and Canescentia

      • section Chrysogena

        • series Aethiopica, Chrysogena, Crustacea, Goetziorum and Persicina

      • section Eladia

        • series Eladia

      • section Fasciculata

        • series Camembertiorum, Corymbifera, Gladioli, Verrucosa and Viridicata

      • section Formosana

        • series Formosana

      • section Osmophila

        • series Osmophila and Samsoniorum

      • section Paradoxa

        • series Atramentosa and Paradoxa

      • section Penicillium

        • series Clavigera, Digitata, Italica, Penicillium and Sclerotigena

      • section Ramosum

        • series Lanosa, Raistrickiorum, Scabrosa, Soppiorum and Virgata

      • section Robsamsonia

        • series Claviformia, Glandicolarum, Robsamsonia and Urticicola

      • section Roquefortorum

        • series Roquefortorum

      • section Turbata

        • series Turbata

  • Phialomyces

  • Pseudohamigera

  • Pseudopenicillium

  • Sclerocleista

  • Warcupiella

  • Xerochrysium

  • Xeromyces

• incertae sedis

  • Dendrosphaera

• Penicillaginaceae

  • Penicillago

• Thermoascaceae

  • Paecilomyces

  • Thermoascus

• Trichocomaceae

  • Acidotalaromyces

  • Ascospirella

  • Rasamsonia

  • Sagenomella

  • Talaromyces

    • section Bacillispori

    • section Helici

    • section Islandici

    • section Purpurei

    • section Subinflati

    • section Talaromyces

    • section Tenues*

    • section Trachyspermi

  • Thermomyces

  • Trichocoma

List of species described since Houbraken et al. (2020)

Aspergillus agricola [nom. inval. Art. 40.8 (Shenzhen)] P. Singh, K.A. Callicott, M.J. Orbach & P.J. Cotty, Front. Microbiol. 11, 1236: 7. 2020. [MB 830377]. — Type: NRRL 66869 (holotype). Ex-type: NRRL 66869 = CR9-G = A2400. Infragen. class.: subgen. Circumdati sect. Flavi ser. Flavi. DNA barcodes: ITS = n.a.; BenA = n.a.; CaM = MN987053; RPB2 = n.a.

Synonym of: Aspergillus flavus Link, Mag. Ges. Naturf. Freunde Berlin 3: 16. 1809. [MB 209842].

Notes: This species was invalidly described. The CaM phylogeny resolved this species within A. flavus (Suppl. Fig. S6). We therefore consider it a synonym of A. flavus and choose not to validate this species.

Aspergillus alboluteus F. Sklenar, Jurjević, Ezekiel, Houbraken & Hubka, Stud. Mycol. 99 (no. 100120): 19. 2021. [MB 839382]. — Type: PRM 952200 (holotype). Ex-type: CBS 145855 = EMSL 2420 = CCF 5695 = IFM 66815. Infragen. class.: subgen. Circumdati sect. Flavipedes ser. Spelaei. DNA barcodes: ITS = MW448663; BenA = MW478497; CaM = MW478511; RPB2 = MW478532.

Aspergillus alboviridis J.P.Z. Siqueira, Gene, F. Sklenar & Hubka, Stud. Mycol. 99 (no. 100120): 19. 2021. [MB 821808]. — Type: CBS H-23128 (holotype). Ex-type: CBS 142665 = FMR 15175 = CCF 6049 = IFM 66819. Infragen. class.: subgen. Circumdati sect. Flavipedes ser. Spelaei. DNA barcodes: ITS = LT798909; BenA = LT798936; CaM = LT798937; RPB2 = LT798938.

Aspergillus annui J.J. Silva, Fungaro, Frisvad, Taniwaki & Iamanaka, published here. [MB 849336]. — Type: IBT 36122 (holotype). Ex-type: 365-IT-PPK = IBT 36122. Infragen. class.: subgen. Circumdati sect. Flavi ser. Annuorum. DNA barcodes: ITS = OP691228; BenA = ON529842; CaM = ON529841; RPB2 = ON529843.

Notes: This species was invalidly described in Silva et al. (2022) and is validated above.

Aspergillus arizonicus Jurjević, Glässnerová, Yaguchi & Hubka, Persoonia 47: 273. 2021. [MB 841359]. — Type: PRM 954611 (holotype). Ex-type: CCF 5341 = CBS 148476 = IFM 66805 = EMSL 2204. Infragen. class.: subgen. Fumigati sect. Fumigati ser. Neoglabri. DNA barcodes: ITS = OK322364; BenA = OK334128; CaM = OK334127; RPB2 = OK334129.

Aspergillus banksianus Pitt, Persoonia 44: 355. 2020. [MB 835223]. — Type: DAR 85042 (holotype). Ex-type: FRR 6047 = MST FP2248. Infragen. class.: subgen. Fumigati sect. Fumigati ser. Brevipedes. DNA barcodes: ITS = MH280013; BenA = MT184780; CaM = MT184786; RPB2 = MT184792.

Aspergillus barbosae A.C.R. Barros-Correia, R.N. Barbosa, Houbraken & Souza-Motta, Mycol. Prog. 19: 892. 2020. [MB 830077]. — Type: URM 93046 (holotype). Ex-type: URM 5930 = CBS 145863. Infragen. class.: subgen. Circumdati sect. Terrei ser. Terrei. DNA barcodes: ITS = LR536042; BenA = LR031377; CaM = LR031392; RPB2 = LR031407.

Aspergillus burnettii Pitt, Fungal Genet. Biol. 143 (no. 103435): 5. 2020. [MB 835453]. — Type: DAR 84902 (holotype). Ex-type: CBS 146237 = FRR 5400 = MST FP2249. Infragen. class.: subgen. Circumdati sect. Flavi ser. Alliacei. DNA barcodes: ITS = MK429758; BenA = MT211761; CaM = MT211762; RPB2 = MT211763.

Synonym of: Aspergillus alliaceus Thom & Church, Aspergilli: 163. 1926. [MB 256402].

Notes: BenA and CaM phylogenies resolved A. burnettii inside A. alliaceus, while RPB2 resolved it on an unsupported distinct branch (Suppl. Fig. S6). As a result, we consider A. burnetti a synonym of A. alliaceus.

Aspergillus chiangmaiensis S. Khuna, N. Suwannarach & S. Lumyong, Front. Microbiol. 12, 705896: 6. 2021. [MB 830887]. — Type: SDBR-CMUI4 (holotype). Ex-type: TBRC 10407. Infragen. class.: subgen. Circumdati sect. Nigri ser. Nigri. DNA barcodes: ITS = MK457198; BenA = MK457200; CaM = MK457199; RPB2 = MK457203.

Synonym of: Aspergillus tubingensis Mosseray, La Cellule 43: 245. 1934. [MB 255209] (Bian et al. 2022).

Notes: Khuna et al. (2021) introduced A. chiangmaiensis based on DNA sequences that are of low quality and the species was subsequently considered a synonym of A. tubingensis by Bian et al. (2022).

Aspergillus curvatus Al-Bedak & Moubasher, Asian J. Mycol. 3: 327. 2020. [MB 557788]. — Type: AUMC 11038 (holotype). Ex-type: EMCCN 2213. Infragen. class.: subgen. Circumdati sect. Circumdati ser. Steyniorum. DNA barcodes: ITS = MN006961; BenA = n.a.; CaM = n.a.; RPB2 = n.a.

Notes: Doubtful species. This species was described with only an ITS sequence. Based on sequence alignments, the ITS sequence is potentially of low-quality. We obtained a culture of the ex-type strain but this unfortunately represented a different species. As we could not confirm the status of A. curvatus, we consider it a doubtful species (Suppl. Fig. S4). Two identifiers were cited in the protologue. IF 557788 (= MB 557788) was listed first and is thus the one that is permanently associated with the name. MycoBank MB 831990 should not be used.

Aspergillus gaarensis [nom. inval. Art. 40.7 (Shenzhen)] Al-Bedak & Moubasher, Stud. Fungi 5: 61. 2020. [MB 833223]. — Type: AUMC 11046 = EMCCN 2221 (holotype). Ex-type: AUMC 11046 = EMCCN 2221. Infragen. class.: subgen. Circumdati sect. Circumdati ser. Steyniorum. DNA barcodes: ITS = MN648408; BenA = n.a.; CaM = n.a.; RPB2 = n.a.

Notes: Doubtful species. This species was invalidly described with only an ITS sequence to support it. However, based on sequence alignments it was potentially a low-quality sequence. We obtained a culture of the ex-type strain but this unfortunately represented a different species. As we could not confirm the status of A. gaarensis, we consider it a doubtful species (Suppl. Fig. S4) and choose not to validate this species.

Aspergillus guangdongensis B.D. Sun, X.Z. Jiang & A.J. Chen, J. Fungi 8 (no 1205): 4. 2022. [MB 837898]. — Type: HMAS 248373 (holotype). Ex-type: CGMCC 3.19704 = MN 014767. Infragen. class.: subgen. Nidulantes sect. Ochraceorosei ser. Funiculosi. DNA barcodes: ITS = MN640760; BenA = MN635246; CaM = MN635257; RPB2 = MN635269.

Synonym of: Aspergillus lannaensis Suwannar., S. Khuna & Lumyong, Fungal Divers. 111: 145. 2021. [MB 838058].

Notes: Phylogenies resolved A. lannaensis and A. guangdongensis in the same clade (Fig. 3 & Suppl. Fig. S3) and we consider them synonyms with A. lannaensis having priority over A. guangdongensis.

Fig. 3.

Phylogenetic tree of Aspergillus sections Cavernicolarum, Ochraceorosei and Sparsi based on a concatenated dataset of BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. The tree was rooted to A. bisporus. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by superscript T. See Suppl. Fig. S3.

Aspergillus guangxiensis B.D. Sun, X.Z. Jiang & A.J. Chen, J. Fungi 8 (no 1205): 9. 2022. [MB 837899]. — Type: HMAS 248372 (holotype). Ex-type: CGMCC 3.19709 = MN 115635. Infragen. class.: subgen. Nidulantes sect. Sparsi ser. Conjuncti. DNA barcodes: ITS = MN640765; BenA = MN635251; CaM = MN635262; RPB2 = MN635274.

Aspergillus hainanicus X.C. Wang & W.Y. Zhuang, J. Fungi 8 (225): 9. 2022. [MB 570967]. — Type: HMAS 247855 (holotype). Ex-type: CGMCC 3.20888. Infragen. class.: subgen. Nidulantes sect. Cavernicolarum ser. Hainanici. DNA barcodes: ITS = OM414846; BenA = OM475626; CaM = OM475630; RPB2 = OM475634.

Aspergillus hydei Doilom, Front. Microbiol. 11 (no. 585215): 4. 2020. [MB 557860]. — Type: MFLU 20-0430 (holotype). Ex-type: KUMCC 18-0196. Infragen. class.: subgen. Circumdati sect. Nigri ser. Japonici. DNA barcodes: ITS = MT152332; BenA = MT161679; CaM = MT178247; RPB2 = MT384370.

Notes: The phylogenetic trees are not well-resolved (Fig. 10 & Suppl. Fig. S10), and further studies including additional strains and/or extended datasets are needed. Here, we give benefit of doubt to the publisher.

Fig. 10.

Phylogenetic tree of Aspergillus section Nigri series Japonici and Nigri based on a concatenated dataset of BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. The tree was rooted to A. candidus. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by superscript T. See Suppl. Fig. S10.

Aspergillus inusitatus F. Sklenar, C. Silva Pereira, Houbraken & Hubka, Stud. Mycol. 99: 20. 2021. [MB 839383]. — Type: PRM 954606 (holotype). Ex-type: CBS 147044 = DTO 121-G5 = CCF 6552. Infragen. class.: subgen. Circumdati sect. Flavipedes ser. Spelaei. DNA barcodes: ITS = MW448669; BenA = MW478502; CaM = MW478517; RPB2 = MW478542.

Aspergillus jilinensis X.Z. Jiang, Mycoscience 61: 207. 2020. [MB 819490]. — Type: HMAS 247010 (holotype). Ex-type: CGMCC 3.18134. Infragen. class.: subgen. Circumdati sect. Terrei ser. Terrei. DNA barcodes: ITS = KX443224; BenA = KX443162; CaM = KX443193; RPB2 = n.a.

Aspergillus kumbius Pitt, Persoonia 44: 359. 2020. [MB 835225]. — Type: DAR 85044 (holotype). Ex-type: FRR 6049 = MST FP2250. Infragen. class.: subgen. Circumdati sect. Circumdati ser. Sclerotiorum. DNA barcodes: ITS = MT179307; BenA = MT184782; CaM = MT184788; RPB2 = MT184794.

Aspergillus lannaensis Suwannar., S. Khuna & Lumyong, Fungal Divers. 111: 145. 2021. [MB 838058]. — Type: SDBR-CMUO 8 (holotype). Ex-type: SDBR-CMUO 8. Infragen. class.: subgen. Nidulantes sect. Ochraceorosei ser. Funiculosi. DNA barcodes: ITS = n.a.; BenA = MW219783; CaM = MW219781; RPB2 = MW219785.

Aspergillus lanuginosus F. Sklenar & Hubka, Stud. Mycol. 99: 20. 2021. [MB 839384]. — Type: PRM 954608 (holotype). Ex-type: NRRL 4610 = IMI 350352 = CCF 4551 = IFM 66818. Infragen. class.: subgen. Circumdati sect. Flavipedes ser. Spelaei. DNA barcodes: ITS = EF669604; BenA = EU014080; CaM = EF669562; RPB2 = EF669646.

Aspergillus lebretii V.C.S. Alves, J.D.P. Bezerra & R.N. Barbosa, Fungal Syst. Evol. 10: 144. 2022. [MB 846119]. — Type: URM 95150 (holotype). Ex-type: URM 8451 = FCCUFG 09. Infragen. class.: subgen. Cremei sect. Cremei ser. Wentiorum. DNA barcodes: ITS = ON862928; BenA = OP672382; CaM = OP290540; RPB2 = OP290511.

Aspergillus lentisci (Durieu & Mont.) Visagie, Malloch, L. Kriegsteiner, Samson & Houbraken, published here. [MB 849485]. Basionym: Dichlaena lentisci Durieu & Mont., Expl. Sci. l’Algérie 1: 405. 1849. [MB 249716]. — Type: Durieu & Mont., Expl. Sci. l’Algérie 1 (11): 405, tab. 22 bis, fig. 2 (lectotype), CBS H-25292 (epitype). Ex-epitype: CBS 150189. Infragen. class.: subgen. Circumdati sect. Dichlaena ser. Dichlaena. DNA barcodes: ITS = OR142402; BenA = OR145977; CaM = OR145992; RPB2 = OR146003.

Aspergillus limoniformis Z.F. Zhang & L. Cai, Fungal Divers. 106: 77. 2021. [MB 556394]. — Type: HMAS 248014 (holotype). Ex-type: CGMCC 3.19323 = LC126098. Infragen. class.: subgen. Polypaecilum sect. Polypaecilum ser. Canini. DNA barcodes: ITS = MK329066; BenA = MK336093; CaM = n.a.; RPB2 = MK335972.

Aspergillus luteorubrus Pitt, Persoonia 44: 361. 2020. [MB 835226]. — Type: DAR 85045 (holotype). Ex-type: CBS 146723 = FRR 5427 = MST FP2246. Infragen. class.: subgen. Fumigati sect. Fumigati ser. Fennelliarum. DNA barcodes: ITS = MT179305; BenA = MT184781; CaM = MT184787; RPB2 = MT184793.

Synonym of: Aspergillus fennelliae Kwon-Chung & S.J. Kim, Mycologia 66: 629. 1974. [MB 309218].

Notes: Even though the multigene phylogeny resolves A. luteorubrus basal to A. fennelliae, both the BenA and RPB2 phylogenies places it intermixed with strains of the latter (Fig. 8 & Suppl. Fig. S8). We thus consider it a synonym.

Fig. 8.

Phylogenetic tree of Aspergillus section Fumigati based on a concatenated dataset of BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. The tree was rooted to A. clavatus. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by superscript T. Representative strains are indicated by superscript R. See Suppl. Fig. S8.

Aspergillus magnus Glässnerová & Hubka, Stud. Mycol. 102: 38. 2022. [MB 844202]. — Type: PRM 956934 (holotype). Ex-type: UAMH 1324 = IBT 14560. Infragen. class.: subgen. Circumdati sect. Candidi ser. Candidi. DNA barcodes: ITS = ON156376; BenA = ON164570; CaM = ON164619; RPB2 = ON164517.

Aspergillus malvicolor A.D. Hocking, Persoonia 44: 363. 2020. [MB 835227]. — Type: DAR 85046 (holotype). Ex-type: CBS 146724 = FRR 2383 = MST FP2244. Infragen. class.: subgen. Circumdati sect. Circumdati ser. Sclerotiorum. DNA barcodes: ITS = MT179308; BenA = MT184784; CaM = MT184790; RPB2 = MT184796.

Aspergillus marneyi Y.P. Tan, Bishop-Hurley, E. Lacey & R.G. Shivas, Index Austral. Fungi 3: 1. 2022. [MB 900143]. — Type: BRIP 71536a (holotype). Ex-type: BRIP 71536a. Infragen. class.: subgen. Circumdati sect. Terrei ser. Terrei. DNA barcodes: ITS = OL691080; BenA = OL741659; CaM = n.a.; RPB2 = OL741656.

Synonym of: Aspergillus alabamensis Balajee, Baddley, Frisvad & Samson, Eukar. Cell 8: 720. 2009. [MB 543648].

Notes: Phylogenies resolve A. marneyi inside the A. alabamensis clade (Fig. 12 & Suppl. Fig. S12). We thus consider it a synonym.

Fig. 12.

Phylogenetic tree of Aspergillus section Terrei series Nivei and Terrei based on a concatenated dataset of BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. The tree was rooted to A. ambiguus. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by superscript T. See Suppl. Fig. S12.

Aspergillus montoensis Y.P. Tan, Bishop-Hurley, S.M. Thompson & R.G. Shivas, Index Fungorum 503: 1. 2021. [MB 558828]. — Type: BRIP 64553a (holotype). Ex-type: BRIP 64553a = CBS 149107. Infragen. class.: subgen. Circumdati sect. Terrei ser. Terrei. DNA barcodes: ITS = OK441076; BenA = OK533535; CaM = n.a.; RPB2 = OK509073.

Notes: Both the ITS and BenA sequences are identical to A. terreus, but RPB2 resolves it distinct from A. terreus as a close relative of A. citrinoterreus (Fig. 12 & Suppl. Fig. S12). Further studies including additional strains and/or extended datasets are needed. Here, we give benefit of doubt to the publisher.

Aspergillus nanangensis Pitt, Persoonia 44: 365. 2020. [MB 836001]. — Type: DAR 84903 (holotype). Ex-type: CBS 146238 = FRR 6048 = MST FP2251. Infragen. class.: subgen. Circumdati sect. Janorum ser. Janorum. DNA barcodes: ITS = MK979278; BenA = MT184783; CaM = MT184789; RPB2 = MT184795.

Aspergillus neoterreus X.C. Wang and W.Y. Zhuang, J. Fungi 8 (3, no. 225): 10. 2022. [MB 570968]. — Type: HMAS 247856 (holotype). Ex-type: CGMCC 3.20891. Infragen. class.: subgen. Nidulantes sect. Terrei ser. Terrei. DNA barcodes: ITS = OM414849; BenA = OM475629; CaM = OM475633; RPB2 = OM475637.

Notes: The phylogenetic trees are not well-resolved (Fig. 12 & Suppl. Fig. S12), and further studies including additional strains and/or extended datasets are needed. Here, we give benefit of doubt to the publisher.

Aspergillus neotritici Glässnerová & Hubka, Stud. Mycol. 102: 38. 2022. [MB 844204]. — Type: PRM 956940 (holotype). Ex-type: CCF 3853 = IBT 32725. Infragen. class.: subgen. Circumdati sect. Candidi ser. Candidi. DNA barcodes: ITS = FR727136; BenA = FR775327; CaM = HE661598; RPB2 = LT627021.

Notes: This new name was introduced for the invalid A. tritici [nom. inval. Art. 8.1, Art. 8.4, Art 40.1, Art. 40.4 (Shenzhen)] by Glässnerová et al. (2022).

Aspergillus okavangoensis Visagie & Nkwe, Fungal Syst. Evol. 8: 85. 2021. [MB 840269]. — Type: PREM 63212 (holotype). Ex-type: CBS 147420 = CMW 56636 = CN073A3 = DN24. Infragen. class.: subgen. Circumdati sect. Flavipedes ser. Flavipedes. DNA barcodes: ITS = MW480880; BenA = MW480788; CaM = MW480706; RPB2 = MW480790.

Aspergillus oxumiae C.N. Figueiredo, L.S. Sales, Y.F. Figueiredo, J.P. Andrade & J.T. De Souza, Persoonia 44: 357. 2020. [MB 832766]. — Type: HURB 22369 (holotype). Ex-type: CCDCA 11546 = UFLA115. Infragen. class.: subgen. Circumdati sect. Nigri ser. Japonici. DNA barcodes: ITS = MN431160; BenA = MN521388; CaM = MN531842; RPB2 = MN521389.

Aspergillus pakistanicus [nom. inval. Art. 40.7 & Art. 40.8 (Shenzhen)] Abrar, Mughal & S. Sarwar, Appl. Ecol. Environm. Res. 18: 3797. 2020. [MB 830877]. — Type: AA100717 (holotype). Ex-type: AA100717. Infragen. class.: subgen. Circumdati sect. Flavi ser. Flavi. DNA barcodes: ITS = n.a.; BenA = n.a.; CaM = n.a.; RPB2 = n.a.

Notes: The species was invalidly described. Only an SSU (18S small subunit nrDNA) sequence is available for this species (GenBank MK371711). Even though it is unique, there are very little SSU data available for Aspergillus, and we therefore decided not to validate the species here.

Aspergillus phialiformis Z.F. Zhang & L. Cai, Fungal Divers. 106: 79. 2021. [MB 556395]. — Type: HMAS 248017 (holotype). Ex-type: CGMCC 3.19314 = LC12536. Infragen. class.: subgen. Polypaecilum sect. Polypaecilum ser. Canini. DNA barcodes: ITS = MK329068; BenA = MK336095; CaM = n.a.; RPB2 = MK335974.

Aspergillus phialosimplex Z.F. Zhang & L. Cai, Fungal Divers. 106: 79. 2021. [MB 556396]. — Type: HMAS 248007 (holotype). Ex-type: CGMCC 3.19637 = LC12578. Infragen. class.: subgen. Polypaecilum sect. Polypaecilum ser. Canini. DNA barcodes: ITS = MK329070; BenA = MK336097; CaM = n.a.; RPB2 = MK335976.

Aspergillus pseudopiperis S. Khuna, N. Suwannarach & S. Lumyong, Front. Microbiol. 12, 705896: 6. 2021. [MB 830888]. — Type: SDBR-CMUI1 (holotype). Ex-type: TBRC 10408. Infragen. class.: subgen. Circumdati sect. Nigri ser. Nigri. DNA barcodes: ITS = n.a.; BenA = MK457194; CaM = MK457193; RPB2 = MK457196.

Synonym of: Aspergillus tubingensis Mosseray, La Cellule 43: 245. 1934. [MB 255209] (Bian et al. 2022).

Notes: Khuna et al. (2021) introduced this species based on DNA sequences that are of low quality and the species was subsequently considered a synonym of A. tubingensis by Bian et al. (2022).

Aspergillus pseudotubingensis S. Khuna, N. Suwannarach & S. Lumyong, Front. Microbiol. 12, 705896: 9. 2021. [MB 830889]. — Type: SDBR-CMUO2 (holotype). Ex-type: TBRC 10409. Infragen. class.: subgen. Circumdati sect. Nigri ser. Nigri. DNA barcodes: ITS = n.a.; BenA = MK457206; CaM = MK457205; RPB2 = MK457208.

Notes: Doubtful species. Khuna et al. (2021) introduced this species based on DNA sequences that are of low quality and its taxonomic position was subsequently considered doubtful by Bian et al. (2022).

Aspergillus qilianyuensis X.C. Wang & W.Y. Zhuang, J. Fungi 8 (225): 12. 2022. [MB 570969]. — Type: HMAS 247857 (holotype). Ex-type: CGMCC 3.20889. Infragen. class.: subgen. Nidulantes sect. Nidulantes ser. Versicolores. DNA barcodes: ITS = OM414847; BenA = OM475627; CaM = OM475631; RPB2 = OM475635.

Aspergillus recifensis A.C.R. Barros-Correia, R.N. Barbosa, Houbraken & Souza-Motta, Mycol. Prog. 19: 894. 2020. [MB 830081]. — Type: URM 93050 (holotype). Ex-type: URM 6605 = CBS 145864. Infragen. class.: subgen. Circumdati sect. Terrei ser. Nivei. DNA barcodes: ITS = LR536036; BenA = LR031370; CaM = LR031385; RPB2 = LR031400.

Aspergillus rouenensis Crous & Decock, Persoonia 48: 289. 2022. [MB 844262]. — Type: CBS H-24987. Ex-type: CBS 149067 = CPC 41585 = MUCL 58110. Infragen. class.: subgen. Polypaecilum sect. Polypaecilum ser. Salinarum. DNA barcodes: ITS = ON603782; BenA = ON605641; CaM = ON653193; RPB2 = ON653194.

Aspergillus saccharicola J.J. Silva, Frisvad, M.H.P. Fungaro, M.H. Taniwaki & B.T. Iamanaka, published here. [MB 849338]. — Type: IBT 36126 (holotype). Ex-type: 117-IT-SGC = IBT 36126. Infragen. class.: subgen. Circumdati sect. Flavi ser. Flavi. DNA barcodes: ITS = OP611470; BenA = ON529845; CaM = ON529844; RPB2 = ON529846.

Notes: This species was invalidly described in Silva et al. (2022) and is validated above.

Aspergillus sakultaensis [nom. inval. Art. 40.8 (Shenzhen)] Al-Bedak, Zohri & Abdel-Kareem, J. Environm. Stud. 20: 23. 2020. [MB 831480]. — Type: UMC 13885 (holotype). Ex-type: AUMC 13885. Infragen. class.: subgen. Circumdati sect. Flavipedes ser. Flavipedes. DNA barcodes: ITS = MK391495; BenA = n.a.; CaM = n.a.; RPB2 = n.a.

Synonym of: Aspergillus templicola Visagie, Hirooka & Samson, Stud. Mycol. 78: 103. 2014. [MB 809191].

Notes: Aspergillus sakultaensis was described with only an ITS sequence to support it, but this is identical to A. templicola (Suppl. Fig. S7). As a result, we consider it a synonym of the latter.

Aspergillus sibiricus V.A. Iliushin, Phytotaxa 531: 68. 2022. [MB 841752]. — Type: LE F-341005 (holotype). Ex-type: CBS 143307. Infragen. class.: subgen. Fumigati sect. Fumigati ser. Unilaterales. DNA barcodes: ITS = MG587008; BenA = MG722970; CaM = MG722971; RPB2 = MG710809.

Aspergillus sichuanensis B.D. Sun, X.Z. Jiang & A.J. Chen, J. Fungi 8 (no 1205): 10. 2022. [MB 837900]. — Type: HMAS 248374 (holotype). Ex-type: CGMCC 3.19706 = MN 18437. Infragen. class.: subgen. Nidulantes sect. Aenei ser. Aenei. DNA barcodes: ITS = MN640762; BenA = MN635248; CaM = MN635259; RPB2 = MN635271.

Aspergillus sigarelli B.D. Sun, Houbraken, A.J. Chen and Samson, Int. J. Syst. Evol. Microbiol. 70: 9. 2020. [MB 833129]. — Type: CBS H-22725 (holotype). Ex-type: CBS 141579 = DTO 348-D4 = CGMCC 3.03936. Infragen. class.: subgen. Nidulantes sect. Usti ser. Calidousti. DNA barcodes: ITS = MN640758; BenA = MN635244; CaM = MN635255; RPB2 = MN635267.

Aspergillus telluris B.D. Sun, X.Z. Jiang & A.J. Chen, Phytotaxa 455: 146. 2020. [MB 834494]. — Type: HMAS 248375 (holotype). Ex-type: CGMCC 3.19701. Infragen. class.: subgen. Polypaecilum sect. Polypaecilum ser. Canini. DNA barcodes: ITS = MN640767; BenA = MN635253; CaM = MN635264; RPB2 = MN635276.

Aspergillus tenebricus Houbraken, Glässnerová & Hubka, Stud. Mycol. 102: 45. 2022. [MB 844203]. — Type: PRM 957108 (holotype). Ex-type: CBS 147048 = DTO 337-H7. Infragen. class.: subgen. Circumdati sect. Candidi ser. Candidi. DNA barcodes: ITS = ON156389; BenA = ON164584; CaM = ON164623; RPB2 = ON164532.

Aspergillus tibetensis B.D. Sun, X.Z. Jiang & A.J. Chen, J. Fungi 8 (no 1205): 12. 2022. [MB 837901]. — Type: HMAS 248371 (holotype). Ex-type: CGMCC 3.19707 = MN 110445. Infragen. class.: subgen. Nidulantes sect. Aenei ser. Aenei. DNA barcodes: ITS = MN640763; BenA = MN635249; CaM = MN635260; RPB2 = MN635272.

Aspergillus toxicus [nom. inval. Art. 40.8 (Shenzhen)] P. Singh, K.A. Callicott, M.J. Orbach & P.J. Cotty, Front. Microbiol. 11, 1236: 10. 2020. [MB 832486]. — Type: NRRL 66898 (holotype). Ex-type: NRRL 66898 = A5-B-S = A2406. Infragen. class.: subgen. Circumdati sect. Flavi ser. Flavi. DNA barcodes: ITS = n.a.; BenA = n.a.; CaM = MN987092; RPB2 = n.a.

Synonym of: Aspergillus minisclerotigenes Vaamonde, Frisvad & Samson, Int. J. Syst. Evol. Microbiol. 58: 733. 2008. [MB 505188].

Notes: This species was invalidly described. The CaM phylogeny resolved this species within the A. minisclerotigenes clade (Suppl. Fig. S6) and we thus consider this species to be a synonym of the latter and, therefore, choose not to validate this species.

Aspergillus vinaceus Ferranti, Iamanaka, Frisvad, O. Puel & J.J. da Silva, J. Fungi 6 (no. 371): 14. 2020. [MB 833399]. — Type: ITAL 47.456 (holotype). Ex-type: ITAL 47.456 = IBT 35556. Infragen. class.: subgen. Circumdati sect. Nigri ser. Nigri. DNA barcodes: ITS = MN575692; BenA = MN583579; CaM = MN583580; RPB2 = MN583581.

Synonym of: Aspergillus niger Tiegh., Ann. Sci. Nat., Bot., ser. 58: 240. 1867. [MB 284309].

Notes: Phylogenies resolved A. vinaceus in the A. niger clade (Fig. 10 & Suppl. Fig. S10), similar to the results found during a recent revision of series Nigri (Bian et al. 2022).

Aspergillus xishaensis X.C. Wang & W.Y. Zhuang, J. Fungi 8 (225): 13. 2022. [MB 570970]. — Type: HMAS 247858 (holotype). Ex-type: CGMCC 3.20890. Infragen. class.: subgen. Circumdati sect. Flavipedes ser. Flavipedes. DNA barcodes: ITS = OM414848; BenA = OM475628; CaM = OM475632; RPB2 = OM475636.

Emericella sydowii (Bainier & Sartory) Pitt & A.D. Hocking, Fungi and Food Spoilage. 4^th Ed.: 611. 2022. [MB 838069]. Basionym: Sterigmatocystis sydowii Bainier & Sartory, Ann. Mycol. 11: 25. 1913. [MB 197979] — Type: IMI 211384.

Synonym of: Aspergillus sydowii (Bainier & Sartory) Thom & Church, The Aspergilli: 147. 1926. [MB 279636]. — Type: IMI 211384. Ex-type: CBS 593.65 = NRRL 250 = IMI 211384 = NRRL 254. Infragen. class.: subgen. Nidulantes sect. Nidulantes ser. Versicolores. DNA barcodes: ITS = EF652450; BenA = EF652274; CaM = EF652362; RPB2 = EF652186.

Notes: Pitt & Hocking (2022) introduced this new combination to accommodate A. sydowii in Emericella. The genus Emericella is considered a synonym of Aspergillus (Samson et al. 2014). We therefore do not accept this new combination and consider it a synonym of Aspergillus sydowii.

Emericella versicolor (Vuill.) Pitt & A.D. Hocking, Fungi and Food Spoilage. 4^th Ed.: 611. 2022. [MB 838068]. Basionym: Sterigmatocystis versicolor Vuill., Erreur Dét. Asp. Paras. Homme: 15. 1903. [MB 233198]. — Type: CBS 583.65 (holotype).

Synonym of: Aspergillus versicolor (Vuill.) Tirab., Ann. Bot. (Roma) 7: 9. 1908. [MB 172159]. — 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. DNA barcodes: ITS = EF652442; BenA = EF652266; CaM = EF652354; RPB2 = EF652178.

Notes: Pitt & Hocking (2022) introduced this new combination to accommodate A. versicolor in Emericella. The genus Emericella is considered a synonym of Aspergillus (Samson et al. 2014). We therefore do not accept this new combination and consider this species a synonym of Aspergillus versicolor.

Emericella usta (Bainier) Pitt & A.D. Hocking, Fungi and Food Spoilage. 4^th Ed.: 611. 2022. [MB 838032]. Basionym: Sterigmatocystis usta Bainier, Bull. Soc. Bot. France 28: 78. 1881. [MB 536545]. — Type: IMI 211805 (holotype).

Synonym of: Aspergillus ustus (Bainier) Thom & Church, The Aspergilli: 152. 1926. [MB 281216]. — 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. DNA barcodes: ITS = EF652455; BenA = EF652279; CaM = EF652367; RPB2 = EF652191.

Notes: Pitt & Hocking (2022) introduced this new combination to accommodate A. ustus in Emericella. The genus Emericella is considered a synonym of Aspergillus (Samson et al. 2014). We therefore do not accept this new combination and consider this species a synonym of Aspergillus ustus.

Neosartorya clavata (Desm.) Pitt & A.D. Hocking, Fungi and Food Spoilage. 4^th Ed.: 611. 2022. [MB 838031]. — Type: IMI 15949 (holotype).

Synonym of: Aspergillus clavatus Desm., Ann. Sci. Nat., Bot., ser. 2: 71. 1834. [MB 211530]. — 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. DNA barcodes: ITS = EF669942; BenA = EF669802; CaM = EF669871; RPB2 = EF669730.

Notes: Pitt & Hocking (2022) introduced this new combination to accommodate A. clavatus in Neosartorya. The genus Neosartorya is considered a synonym of Aspergillus (Samson et al. 2014). We therefore do not accept this new combination and consider it a synonym of Aspergillus clavatus.

Paecilomyces clematidis Spetik, Eichmeier, Gramaje & Berraf-Tebbal, Phytotaxa 559: 242. 2022. [MB 843540]. — Type: BRNU 677844 (holotype). Ex-type: CBS 148466 = MEND-F-0560. DNA barcodes: ITS = MZ923760; BenA = MZ927740; CaM = MZ927738; RPB2 = OL332317.

Paecilomyces penicilliformis Jurjević & Hubka, Persoonia 44: 431. 2020. [MB 834874]. — Type: BPI 911216 (holotype). Ex-type: CCF 5755 = CBS 146003 = EMSL 3392. DNA barcodes: ITS = LR679769; BenA = LR679768; CaM = LR778299; RPB2 = n.a.

Paraxerochrysium Crous & Decock, Persoonia 47: 261. 2021. [MB 841829]

Synonym of: Xerochrysium Pitt, SIM 4 (2): 236. 2013. [MB 807003].

Notes: See notes for Xerochrysium coryli above.

Paraxerochrysium coryli Crous & Decock, Persoonia 47: 261. 2021. [MB 841830]. — Type: CBS H-24853 (holotype). Ex-type: CBS 148314 = CPC 41272 = MUCL 58103. DNA barcodes: ITS = OK664748; BenA = OK651216; CaM = n.a.; RPB2 = OK651178.

Notes: See notes for Xerochrysium coryli above.

Penicillium allaniae Y.P. Tan, Bishop-Hurley, Marney, R.G. Shivas, Index Austral. Fungi 3: 6. 2022. [MB 900139]. — Type: BRIP 74886a (holotype). Ex-type: BRIP 74886a. Infragen. class.: subgen. Aspergilloides sect. Exilicaulis ser. Restricta. DNA barcodes: ITS = OP903476; BenA = OP921959; CaM = OP921957; RPB2 = OP921958.

Notes: Since Houbraken et al. (2020), P. allaniae, P. archerae, P. krskae and P. silybi were introduced bringing the number of accepted species in series Restricta to thirteen. However, the series clearly needs a taxonomic revision (Fig. 23 & Suppl. Fig. S21) as noted by Visagie et al. (2016). In the meantime, we accept the four recently introduced species.

Fig. 23.

Phylogenetic tree of Penicillium sections Exilicaulis and Gracilenta based on a concatenated dataset of BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. The tree was rooted to P. stolkiae. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by superscript T. See Suppl. Fig. S21.

Penicillium allsoppiae Visagie, A. Visagie, Frisvad & K. Jacobs, Persoonia 46: 176. 2021. [MB 834426]. — Type: CBS H-22036 (holotype). Ex-type: CBS 138943 = DAOM 241348 = DTO 182-D5 = CV 931. Infragen. class.: subgen. Penicillium sect. Canescentia ser. Canescentia. DNA barcodes: ITS = JX140830; BenA = JX140992; CaM = JX157384; RPB2 = KP016895.

Penicillium anthracinoglaciei Perini, Frisvad & Zalar, Microbial Ecol. 86: 287. 2022 (2023). [MB 835602]. — Type: EXF11443H (Holotype). Ex-type: EXF-11443 = IBT 34739. Infragen. class.: subgen. Penicillium sect. Brevicompacta ser. Brevicompacta. DNA barcodes: ITS = n.a.; BenA = MT080493; CaM = MT080552; RPB2 = MT080519.

Notes: The phylogenetic relationship between P. anthracinoglaciei and P. bialowiezense is unresolved and needs further study (Fig. 18 & Suppl. Fig. S16).

Fig. 18.

Phylogenetic tree of Penicillium section Brevicompacta based on a concatenated dataset of BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. The tree was rooted to P. tularense. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by superscript T. See Suppl. Fig. S16.

Penicillium aquadulcis Hyang B. Lee & T.T.T. Nguyen, Mycobiology 49: 542. 2021. [MB 556953]. — Type: CNUFC HT19009 (holotype). Ex-type: CNUFC JT1301. Infragen. class.: subgen. Aspergilloides sect. Citrina ser. Westlingiorum. DNA barcodes: ITS = OK356194; BenA = OK105100; CaM = OK105102; RPB2 = n.a.

Penicillium archerae Y.P. Tan, Bishop-Hurley, R.G. Shivas, Index Austral. Fungi 3: 7. 2022. [MB 900140]. — Type: BRIP 72549c (holotype). Ex-type: BRIP 72549c. Infragen. class.: subgen. Aspergilloides sect. Exilicaulis ser. Restricta. DNA barcodes: ITS = OP903477; BenA = OP921961; CaM = n.a.; RPB2 = OP921960.

Notes: Since Houbraken et al. (2020), P. allaniae, P. archerae, P. krskae and P. silybi were introduced bringing the number of accepted species in series Restricta to thirteen. However, the series clearly needs a taxonomic revision (Fig. 23 & Suppl. Fig. S21) as noted by Visagie et al. (2016). In the meantime, we accept the four recently introduced species.

Penicillium aspericonidium B.D. Sun, A.J. Chen & Houbraken, Mycol. Prog. 20: 1387. 2021. [MB 838211]. — Type: CBS H-22830 (holotype). Ex-type: CBS 141832 = DTO 030-C5. Infragen. class.: subgen. Aspergilloides sect. Charlesia ser. Phoenicea. DNA barcodes: ITS = MT309657; BenA = MT302240; CaM = n.a.; RPB2 = MT302224.

Penicillium ausonanum Torres-Garcia, Gené & Dania García, MycoKeys 86: 119. 2022. [MB 840556]. — Type: CBS H-24781 (holotype). Ex-type: CBS 148237 = FMR 16948. Infragen. class.: subgen. Aspergilloides sect. Lanata-Divaricata ser. Dalearum. DNA barcodes: ITS = LR655808; BenA = LR655809; CaM = LR655810; RPB2 = LR655811.

Penicillium barbosae S. Ramos, R. Cruz, R.N. Barbosa & Houbraken, Mycol. Prog. 20: 828. 2021. [MB 837908]. — Type: URM 94474 (holotype). Ex-type: URM 7705. Infragen. class.: subgen. Aspergilloides sect. Sclerotiorum ser. Adametziorum. DNA barcodes: ITS = MW191494; BenA = MG452818; CaM = MW183245; RPB2 = LR898886.

Penicillium cerradense Cruvinel, P.O. Magalh. & Pinho, Sci. Rep. 11 (no. 17861): 2. 2021. [MB 835241]. — Type: UB23977 (holotype). Ex-type: DCFS6a. Infragen. class.: subgen. Aspergilloides sect. Citrina ser. Westlingiorum. DNA barcodes: ITS = MT006126; BenA = MT416533; CaM = MT416534; RPB2 = MT416532.

Synonym of: Penicillium sumatraense [as ‘sumatrense’] Szilvinyi, Archiv. Hydrobiol. 14 Suppl. 6: 535. 1936. [MB 319297].

Notes: Penicillium cerradense belongs to the P. sumatraense clade and is thus considered a synonym (Fig. 21 & Suppl. Fig. S19).

Fig. 21.

Phylogenetic tree of Penicillium section Citrina based on a concatenated dataset of BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. The tree was rooted to P. euglaucum. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by superscript T. See Suppl. Fig. S19.

Penicillium claroviride Visagie & Yilmaz, Mycologia 115: 90. 2022. [MB 844184]. — Type: PREM 63221 (holotype). Ex-type: CMW 56197 = CBS 147458 = CN014D2. Infragen. class.: subgen. Penicillium sect. Canescentia ser. Atroveneta. DNA barcodes: ITS = MT949909; BenA = MT957414; CaM = MT957456; RPB2 = MT957482.

Penicillium doidgeae Visagie, Frisvad & K. Jacobs, Persoonia 46: 176. 2021. [MB 834427]. — Type: CBS H-22038 (holotype). Ex-type: CBS 138947 = IBT 31950 = DAOM 241107 = DTO 183-G7 = CV 2189. Infragen. class.: subgen. Penicillium sect. Canescentia ser. Atroveneta. DNA barcodes: ITS = JX140804; BenA = JX141006; CaM = JX157413; RPB2 = KP016915.

Penicillium donggangicum L. Wang, PeerJ 10 (e13224): 8. 2022. [MB 841518]. — Type: HMAS 350265 (holotype). Ex-type: AS 3.15900. Infragen. class.: subgen. Aspergilloides sect. Lanata-Divaricata ser. Janthinella. DNA barcodes: ITS = MW946996; BenA = MZ004914; CaM = MZ004918; RPB2 = MW979253.

Penicillium eickeri Visagie, Frisvad & K. Jacobs, Persoonia 46: 179. 2021. [MB 834428]. — Type: CBS H-22034 (holotype). Ex-type: CBS 138939 = IBT 31921 = DAOM 241352 = DTO 181-G3 = CV 475. Infragen. class.: subgen. Penicillium sect. Canescentia ser. Canescentia. DNA barcodes: ITS = JX140824; BenA = JX140979; CaM = JX157365; RPB2 = KP016876.

Penicillium elizabethiae Visagie & Frisvad, Persoonia 46: 179. 2021. [MB 834432]. Basionym: Penicillium echinatum E. Dale, Ann. Mycol. 24: 137. 1926. (nom. illegit. Art. 53.1; non Rivolta (1873). [MB 505484]. — Type: CBS H-22052 (holotype). Ex-type: NRRL 917 = MUCL 29170 = IBT 21955 = DTO 189-B8. Infragen. class.: subgen. Penicillium sect. Canescentia ser. Canescentia. DNA barcodes: ITS = KP016840; BenA = KJ866964; CaM = KJ867021; RPB2 = KP016918.

Notes: Penicillium elizabethiae was introduced for the illegitimate P. echinatum E. Dale.

Penicillium ezekielii Houbraken & Oyedele, Persoonia 49: 309. 2022. [MB 844770]. — Type: CBS H-25015 (holotype). Ex-type: CBS 149115 = DTO 065-D2. Infragen. class.: subgen. Aspergilloides sect. Cinnamopurpurea ser. Jiangxiensia. DNA barcodes: ITS = ON723772; BenA = ON920778; CaM = ON920781; RPB2 = ON920784.

Penicillium ferraniaense Houbraken & Di Piazza, Persoonia 46: 491. 2021. [MB 839119]. — Type: CBS H-24757 (holotype). Ex-type: CBS 147595 = DTO 400-D8. Infragen. class.: subgen. Aspergilloides sect. Sclerotiorum ser. Sclerotiorum. DNA barcodes: ITS = MW694951; BenA = MW689336; CaM = MW689338; RPB2 = MW689340.

Penicillium fuscoglaucum Biourge, La Cellule 33: 128. 1923. [MB 265659]. — Type: Belgium, source and collection date unknown, P. Biourge [Biourge, La Cellule 33: Pl. col. I. dr., n°4; Pl. noire II, fig. 9. 1923, lectotype designated here, MBT10014260); CBS H-15446 (dried culture), epitype designated here, MBT10014261]. Ex-epitype: CBS 261.29 = IMI 092259 = LSHB P81 = MUCL 28651 = NRRL 892 = CBS 122423 = DTO 461-D2. Infragen. class.: subgen. Penicillium sect. Fasciculata ser. Camembertiorum. DNA barcodes: ITS = MH855062; BenA = OR206420; CaM = OR206421; RPB2 = OR206422.

Notes: Giraud et al. (2010) and Ropars et al. (2020) considered P. fuscoglaucum as a distinct species, closely related to P. camemberti, and suggested that P. caseifulvum (CBS 101134^T) is a variety, P. camemberti var. ‘caseifulvum’. Following Frisvad & Samson (2004), we disagree with the idea of introducing varieties in Penicillium. We accept P. camemberti, P. caseifulvum and P. commune, as suggested in the taxonomy of Frisvad & Samson (2004), but also tentatively accept P. fuscoglaucum and P. biforme following the taxonomic insights of Ropars et al. (2020). The taxonomies of Frisvad & Samson (2004) and Ropars et al. (2020) are not congruent and a taxonomic revision is therefore needed to resolve relationships and species boundaries within the clade (see Fig. 19 & Suppl. Fig. S17), noting that P. camemberti, and maybe also P. caseifulvum, are domesticated species related to cheese production (Bodinaku et al. 2019, Ropars et al. 2020).

Fig. 19.

Phylogenetic tree of Penicillium section Fasciculata series Camembertiorum based on a concatenated dataset of BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. The tree was rooted to P. expansum. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by superscript T. See Suppl. Fig. S17.

Penicillium fusiforme B.D. Sun, A.J. Chen & Houbraken, Mycol. Prog. 20: 1389. 2021. [MB 838212]. — Type: CBS H-22840 (holotype). Ex-type: CBS 250.66 = DTO 035-D7. Infragen. class.: subgen. Aspergilloides sect. Charlesia ser. Fellutana. DNA barcodes: ITS = MT309668; BenA = MT302253; CaM = MT302220; RPB2 = MT302236.

Penicillium gercinae A.L. Alves, A.C.S. Santos, R.N. Barbosa, C.M. Souza-Motta, R.F.R. Melo, P.V. Tiago, Acta Bot. Brasil 36 (e2022abb0006): 10. 2022. [MB 841260]. — Type: URM 94476 (holotype). Ex-type: URM 8348. Infragen. class.: subgen. Aspergilloides sect. Ramigena ser. Georgiensia. DNA barcodes: ITS = MW648591; BenA = MW646389; CaM = MW646391; RPB2 = MW646393.

Penicillium guarroi Torres-Garcia, Gené & Dania García, MycoKeys 86: 121. 2022. [MB 840567]. — Type: CBS H-24782 (holotype). Ex-type: CBS 148238 = FMR 17747. Infragen. class.: subgen. Aspergilloides sect. Gracilenta ser. Estinogena. DNA barcodes: ITS = LR814139; BenA = LR814134; CaM = LR814140; RPB2 = LR814145.

Penicillium hepuense L. Wang, PeerJ 10 (e13224): 11. 2022. [MB 841525]. — Type: HMAS 350263 (holotype). Ex-type: AS 3.16039 = TT2-4X3. Infragen. class.: subgen. Aspergilloides sect. Lanata-Divaricata ser. Oxalica. DNA barcodes: ITS = MW946994; BenA = MZ004912; CaM = MZ004916; RPB2 = MW979254.

Penicillium irregulare Torres-Garcia, Gené & Dania García, MycoKeys 86: 125. 2022. [MB 840558]. — Type: CBS H-24783 (holotype). Ex-type: CBS 148240 = FMR 17859. Infragen. class.: subgen. Penicillium sect. Canescentia ser. Canescentia. DNA barcodes: ITS = LR814181; BenA = LR814144; CaM = LR814151; RPB2 = LR814182.

Penicillium jenningsiae Y.P. Tan, Bishop-Hurley, E. Lacey & R.G. Shivas, Index Austral. Fungi 3: 8. 2022. [MB 900141]. — Type: BRIP 45936a (holotype). Ex-type: BRIP 45936a. Infragen. class.: subgen. Aspergilloides sect. Citrina ser. Sumatraensia. DNA barcodes: ITS = n.a.; BenA = OL741657; CaM = n.a.; RPB2 = OL741660.

Synonym of: Penicillium sumatraense [as ‘sumatrense’] Szilvinyi, Archiv. Hydrobiol. 14 Suppl. 6: 535. 1936. [MB 319297].

Notes: Penicillium jenningsiae belongs to the P. sumatraense clade and is thus considered a synonym. (Fig. 21 & Suppl. Fig. S19)

Penicillium jiaozhouwanicum L. Wang, PeerJ 10 (e13224): 13. 2022. [MB 841531]. — Type: HMAS 350262 (holotype). Ex-type: AS 3.16038 = 0801H2-2. Infragen. class.: subgen. Aspergilloides sect. Lanata-Divaricata ser. Oxalica. DNA barcodes: ITS = MW946993; BenA = MZ004911; CaM = MZ004915; RPB2 = MW979252.

Penicillium kalander Visagie & Yilmaz, Mycologia 115: 95. 2022. [MB 844185]. — Type: PREM 63223 (holotype). Ex-type: CMW 56202 = CN014E1. Infragen. class.: subgen. Aspergilloides sect. Sclerotiorum ser. Sclerotiorum. DNA barcodes: ITS = MT949914; BenA = MT957421; CaM = MT957461; RPB2 = MT957487.

Penicillium krskae Labuda, Kubátová, C. Schüller & J. Strauss, J. Fungi 7 (7, no. 557): 7. 2021. [MB 839112]. — Type: PRM 955188 (holotype). Ex-type: CBS 147776 = BiMM-F280 = CCF 6561. Infragen. class.: subgen. Aspergilloides sect. Exilicaulis ser. Restricta. DNA barcodes: ITS = MW794123; BenA = MW774594; CaM = MW774595; RPB2 = MW774593.

Notes: Since Houbraken et al. (2020), P. allaniae, P. archerae, P. krskae and P. silybi were introduced bringing the number of accepted species in series Restricta to thirteen. However, the series clearly needs a taxonomic revision (Fig. 23 & Suppl. Fig. S21) as noted by Visagie et al. (2016). In the meantime, we accept the four recently introduced species.

Penicillium limae S. Ramos, R. Cruz, C. Souza-Motta & N. Tinti, Mycol. Prog. 20: 831. 2021. [MB 837909]. — Type: URM 94475 (holotype). Ex-type: URM 7706. Infragen. class.: subgen. Aspergilloides sect. Sclerotiorum ser. Adametziorum. DNA barcodes: ITS = MW191493; BenA = MG452820; CaM = MW183244; RPB2 = LR898888.

Penicillium linzhiense H-K. Wang & R. Jeewon, Front. Cell. Infect. Microbiol. 10 (no 6045044): 4. 2021. [MB 838576]. — Type: CCTCC-M 2019870 (holotype). Ex-type: CCTCC-M 2019870. Infragen. class.: subgen. Penicillium sect. Canescentia ser. Canescentia. DNA barcodes: ITS = MT461156; BenA = MT461157; CaM = MT461162; RPB2 = n.a.

Penicillium longiconidiophorum B.D. Sun, A.J. Chen & Houbraken, Mycol. Prog. 20: 1389. 2021. [MB 838213]. — Type: CBS H-22829 (holotype). Ex-type: CBS 141831 = DTO 088-C1. Infragen. class.: subgen. Aspergilloides sect. Charlesia ser. Phoenicea. DNA barcodes: ITS = MT309669; BenA = MT302254; CaM = MT302221; RPB2 = MT302237.

Penicillium mattheeae Visagie & Yilmaz, Mycologia 115: 97. 2022. [MB 844186]. — Type: PREM 63219 (holotype). Ex-type: CMW 56388 = CBS 147415 = CN014C5. Infragen. class.: subgen. Aspergilloides sect. Aspergilloides ser. Saturniformia. DNA barcodes: ITS = MT949904; BenA = MT957408; CaM = MT957451; RPB2 = MT957477.

Penicillium melanosporum Rodr.-Andr., Cano & Stchigel, J. Fungi 7 (2, no. 126): 5. 2021. [MB 835938]. — Type: CBS H-24465 (holotype). Ex-type: CBS 146938 = FMR 17424. Infragen. class.: subgen. Aspergilloides sect. Lanata-Divaricata ser. Janthinella. DNA barcodes: ITS = LR655192; BenA = LR655196; CaM = LR655200; RPB2 = LR655204.

Penicillium michoacanense Rodr.-Andr., Cano & Stchigel, J. Fungi 7 (2, no. 126): 8. 2021. [MB 835940]. — Type: CBS H-24467 (holotype). Ex-type: FMR 17612. Infragen. class.: subgen. Aspergilloides sect. Lanata-Divaricata ser. Janthinella. DNA barcodes: ITS = LR655194; BenA = LR655198; CaM = LR655202; RPB2 = LR655206.

Penicillium neoherquei Labuda, Kubátová, Nebesářová, Oberlies & Raja, Persoonia 48: 339. 2022. [MB 842267]. — Type: PRM 956035 (holotype). Ex-type: CBS 148692 = CCF 6604. Infragen. class.: subgen. Aspergilloides sect. Sclerotiorum ser. Herqueorum. DNA barcodes: ITS = MW341222; BenA = OL840853; CaM = OL840855; RPB2 = MW349119.

Penicillium newtonturnerae Y.P. Tan, Bishop-Hurley, Marney & R.G. Shivas, Index Austral. Fungi 3: 10. 2022. [MB 900142]. — Type: BRIP 74909a (holotype). Ex-type: BRIP 74909a. Infragen. class.: subgen. Aspergilloides sect. Lanata-Divaricata ser. Simplicissima. DNA barcodes: ITS = OP903478; BenA = OP921964; CaM = OP921962; RPB2 = OP921963.

Penicillium nordestinense J.E.F. Santos & R.N. Barbosa, Acta Bot. Brasil 36 (e2021abb0390): 4. 2022. [MB 845495]. — Type: URM 83558 (holotype). Ex-type: URM 8423. Infragen. class.: subgen. Aspergilloides sect. Lanata-Divaricata ser. Janthinella. DNA barcodes: ITS = OV265270; BenA = OV265324; CaM = OV265272; RPB2 = OM927721.

Penicillium outeniquaense Visagie & Yilmaz, Mycologia 115: 97. 2022. [MB 844187]. — Type: PREM 63218 (holotype). Ex-type: CMW 56387 = CBS 147414 = CN014C2. Infragen. class.: subgen. Aspergilloides sect. Citrina ser. Westlingiorum. DNA barcodes: ITS = MT949903; BenA = MT957405; CaM = MT957450; RPB2 = MT957476.

Penicillium poederi Kirchm. & Neuh., Fungal Syst. Evol. 10: 92. 2022. [MB 845496]. — Type: IBF2017/0007 (holotype). Ex-type: CBS 147622 = SF014017 = Bq214. Infragen. class.: subgen. Aspergilloides sect. Torulomyces ser. Torulomyces. DNA barcodes: ITS = MF611757; BenA = MF611760; CaM = MF611763; RPB2 = MF611766.

Penicillium pole-evansii Visagie, Frisvad & K. Jacobs, Persoonia 46: 179. 2021. [MB 834429]. — Type: CBS H-22037 (holotype). Ex-type: CBS 138946 = IBT 31929 = DAOM 241106 = DTO 183-D5 = CV 1758. Infragen. class.: subgen. Penicillium sect. Canescentia ser. Atroveneta. DNA barcodes: ITS = JX140831; BenA = JX141005; CaM = JX157412; RPB2 = KP016911.

Penicillium rotoruae O’Callahan & Vaidya, Curr. Microbiol. 77: 4131. 2020. [MB 834084]. — Type: NZFS 4797 (holotype). Ex-type: CBS 145838 = NMI V19/026738. Infragen. class.: subgen. Aspergilloides sect. Lanata-Divaricata ser. Rolfsiorum. DNA barcodes: ITS = MN315103; BenA = MN315104; CaM = MN315102; RPB2 = MT240842.

Penicillium saanichanum Visagie, Assabgui & Seifert, Persoonia 45: 373. 2020. [MB 835962]. — Type: DAOM 745787 (holotype). Ex-type: DAOMC 251850 = KAS 6184. Infragen. class.: subgen. Aspergilloides sect. Cinnamopurpurea ser. Idahoensia. DNA barcodes: ITS = KY469059; BenA = KY469096; CaM = KY469020; RPB2 = MN795070.

Penicillium sanjayi Rajeshk., Visagie, N. Ashtekar & Yilmaz, Mycol. Prog. 21 (4, no. 42): 5. 2022. [MB 840642]. — Type: AMH 10349 (holotype). Ex-type: NFCCI 5017. Infragen. class.: subgen. Aspergilloides sect. Citrina ser. Vascosobrinhoana. DNA barcodes: ITS = MZ571358; BenA = MZ558484; CaM = MZ558492; RPB2 = MZ558482.

Notes: The identifier “MB 840643” cited in Ashtekar et al. (2022) is that of the series Vascosobrinhoana introduced in the same publication, but the species is nevertheless validly published (Art. F.5.6 San Juan Chapter F) (May et al. 2019) since “MB 840642” was issued by MycoBank prior to the publication of the name.

Penicillium scottii Visagie, Frisvad & K. Jacobs, Persoonia 46: 182. 2021. [MB 834430]. — Type: CBS H-22040 (holotype). Ex-type: CBS 138951 = IBT 31905 = DTO 185-F8 = CV 930. Infragen. class.: subgen. Penicillium sect. Canescentia ser. Canescentia. DNA barcodes: ITS = JX140812; BenA = JX140991; CaM = JX157383; RPB2 = KP016894.

Penicillium setosum T.K. George, Houbraken, L. Mathew & M.S. Jisha, Acta Bot. Brasil 36 (e2021abb0390): 6. 2022. [MB 842377]. — Type: CBS H-24872 (holotype). Ex-type: CBS 144865 = DTO 455-G4 = WSR 62 = MCC 1370 = NCFT NO 8222.16 = AMH-9974. Infragen. class.: subgen. Aspergilloides sect. Lanata-Divaricata ser. Janthinella. DNA barcodes: ITS = KT852579; BenA = MF184995; CaM = MH105905; RPB2 = n.a.

Notes: In the original description (George et al. 2018), P. setosum was invalidly described because it was not stated that the holotype was preserved as a metabolically inactive culture. The name was subsequently validated by Barbosa et al. (2022).

Penicillium sexuale Rodr.-Andr., Cano & Stchigel, J. Fungi 7 (2, no. 126): 10. 2021. [MB 835941]. — Type: CBS H-24468 (holotype). Ex-type: CBS 146939 = FMR 17380. Infragen. class.: subgen. Aspergilloides sect. Crypta ser. Crypta. DNA barcodes: ITS = LR655195; BenA = LR655199; CaM = LR655203; RPB2 = LR655207.

Penicillium siccitolerans Rodr.-Andr., Cano & Stchigel, J. Fungi 7 (2, no. 126): 7. 2021. [MB 835939]. — Type: CBS H-24466 (holotype). Ex-type: FMR 17381. Infragen. class.: subgen. Aspergilloides sect. Lanata-Divaricata ser. Janthinella. DNA barcodes: ITS = LR655193; BenA = LR655197; CaM = LR655201; RPB2 = LR655205.

Penicillium sicoris Torres-Garcia, Gené & Dania García, MycoKeys 86: 127. 2022. [MB 840559]. — Type: CBS H-24784 (holotype). Ex-type: CBS 148241 = FMR 18076. Infragen. class.: subgen. Penicillium sect. Paradoxa ser. Atramentosa. DNA barcodes: ITS = LR884497; BenA = LR884494; CaM = LR884496; RPB2 = LR884495.

Penicillium silybi Labuda, Kubátová, Raja & Oberlies, J. Fungi 7 (7, no. 557): 9. 2021. [MB 839113]. — Type: PRM 955189 (holotype). Ex-type: CBS 147777 = G85 = CCF 6562. Infragen. class.: subgen. Aspergilloides sect. Exilicaulis ser. Restricta. DNA barcodes: ITS = KF367458; BenA = MW774592; CaM = MW774591; RPB2 = AB860248.

Notes: Since Houbraken et al. (2020), P. allaniae, P. archerae, P. krskae and P. silybi were introduced bringing the number of accepted species in series Restricta to thirteen. However, the series clearly needs a taxonomic revision (Fig. 23 & Suppl. Fig. S21) as noted by Visagie et al. (2016). In the meantime, we accept the four recently introduced species.

Penicillium soli Doilom, C.F. Liao & D. Pem, Front. Microbiol. 11 (no. 585215): 13. 2020. [MB 557862]. — Type: MFLU 20-0432 (holotype). Ex-type: KUMCC 18-0202. Infragen. class.: subgen. Aspergilloides sect. Lanata-Divaricata ser. Janthinella. DNA barcodes: ITS = MT152337; BenA = MT161681; CaM = MT178249; RPB2 = MT384372.

Synonym: Penicillium cluniae [nom. inval. Art. 40.7 (Shenzhen)] Quintan., Avances en Alimentation y Mechora Animal 30: 174. 1990. [MB 130240].

Notes: Phylogenies resolved P. soli in the P. cluniae clade (Fig. 24 & Suppl. Fig. S22). The exception was RPB2. Pairwise comparisons showed that the ex-type sequences of both species were identical for BenA, but that there was some variation between them for CaM and RPB2. This suggests that they belong to the same species. Penicillium cluniae was previously included in the accepted species lists (Visagie et al. 2014, Houbraken et al. 2020), but was invalidly described by Quintanilla (1990) because no collection or herbarium was specified with the “1532” type designation. Consequently, we reduce P. cluniae as a synonym of P. soli.

Fig. 24.

Phylogenetic tree of Penicillium section Lanata-Divaricata series Janthinella and Simplicissima, based on a concatenated dataset of BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. The tree was rooted to P. stolkiae. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by superscript T. See Suppl. Fig. S22.

Penicillium stangiae A.L. Alves, A.C.S. Santos, R.N. Barbosa, C.M. Souza-Motta, R.F.R. Melo, P.V. Tiago, Acta Bot. Brasil 36: e2022abb0006: 10. 2022. [MB 841261]. — Type: URM 94477 (holotype). Ex-type: URM 8347. Infragen. class.: subgen. Aspergilloides sect. Lanata-Divaricata ser. Dalearum. DNA barcodes: ITS = MW648590; BenA = MW646388; CaM = MW646390; RPB2 = MW646392.

Penicillium subfuscum Visagie & Yilmaz, Mycologia 115: 99. 2022. [MB 844188]. — Type: PREM 63220 (holotype). Ex-type: CMW 56196 = CBS 147455 = CN014C9. Infragen. class.: subgen. Aspergilloides sect. Lanata-Divaricata ser. Simplicissima. DNA barcodes: ITS = MT949907; BenA = MT957412; CaM = MT957454; RPB2 = MT957480.

Penicillium submersum Torres-Garcia, Gené & Dania García, MycoKeys 86: 129. 2022. [MB 840560]. — Type: CBS H-24785 (holotype). Ex-type: CBS 148242 = FMR 17140. Infragen. class.: subgen. Penicillium sect. Robsamsonia ser. Urticicola. DNA barcodes: ITS = LR814194; BenA = LR814187; CaM = LR814188; RPB2 = LR814195.

Penicillium taurinense S. Prencipe, Houbraken & D. Spadaro, Persoonia 44: 435. 2020. [MB 834715]. — Type: CBS H-24332 (holotype). Ex-type: CBS 145672 = DTO 333-B8 = CAS16. Infragen. class.: subgen. Penicillium sect. Robsamsonia ser. Glandicolarum. DNA barcodes: ITS = MF595981; BenA = MF595977; CaM = MF595979; RPB2 = MT253108.

Penicillium tealii Y.P. Tan, Bishop-Hurley, Marney & R.G. Shivas, Persoonia 49: 307. 2022. [MB 845000]. — Type: BRIP 72734c (holotype). Ex-type: BRIP 72734c. Infragen. class.: subgen. Aspergilloides sect. Cinnamopurpurea ser. Jiangxiensia. DNA barcodes: ITS = OP101639; BenA = OP039547; CaM = n.a.; RPB2 = OP039546.

Penicillium tirolense Kirchm., Embacher & Neuh., Fungal Syst. Evol. 10: 96. 2022. [MB 845496]. — Type: IBF2019/0162 (holotype). Ex-type: CBS 147625 = SF014017. Infragen. class.: subgen. Aspergilloides sect. Torulomyces ser. Torulomyces. DNA barcodes: ITS = MW145398; BenA = MW143069; CaM = MW143068; RPB2 = MW143067.

Penicillium tolerans Y.P. Tan, Bishop-Hurley, E. Lacey, Grice & R.G. Shivas, Index Austral. Fungi 3: 12. 2022. [MB 900151]. — Type: BRIP 64090a (holotype). Ex-type: BRIP 64090a. Infragen. class.: subgen. Aspergilloides sect. Sclerotiorum ser. Sclerotiorum. DNA barcodes: ITS = OK639006; BenA = OL741658; CaM = n.a.; RPB2 = n.a.

Synonym of: Penicillium sclerotiorum J.F.H. Beyma, Zentralbl. Bakteriol. Parasitenk., Abt. 2 96: 418. 1937. [MB 277708].

Notes: Species classified in Penicillium section Sclerotiorum were reviewed in Rivera & Seifert (2011) and Visagie et al. (2013), both who accepted a broad concept of P. sclerotiorum. Phylogenetically, the strains considered to belong to P. sclerotiorum can be divided into two clades, but without a taxonomic revision of the species, we consider P. tolerans to be a synonym of the former. (Fig. 27 & Suppl. Fig. S25)

Fig. 27.

Phylogenetic tree of Penicillium section Sclerotiorum, based on a concatenated dataset of BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. The tree was rooted to P. glabrum. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by superscript T. See Suppl. Fig. S25.

Penicillium ucsense A. Lenz & Houbraken, Antonie van Leeuwenhoek 115: 9. 2022. [MB 839721]. — Type: CBS H-24331 (holotype). Ex-type: CBS 146492 = DTO 426-B1 = IOC 4717 = 2HH. Infragen. class.: subgen. Aspergilloides sect. Lanata-Divaricata ser. Oxalica. DNA barcodes: ITS = OM914583; BenA = ON024157; CaM = ON024158; RPB2 = ON024159.

Synonym of: Penicillium hepuense L. Wang, PeerJ 10 (e13224): 11. 2022. [MB 841525].

Notes: Phylogenies resolved P. ucsense and P. hepuense in the same clade (Fig. 25 & Suppl. Fig. S23). Both were introduced in 2022, but P. hepuense was published first (2022/05/06 vs 2022/06/09) and thus has priority over P. ucsense.

Fig. 25.

Phylogenetic tree of Penicillium section Lanata-Divaricata series Dalearum, Oxalica and Rolfsiorum, based on a concatenated dataset of BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. The tree was rooted to the series Oxalica clade. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by superscript T. See Suppl. Fig. S23.

Penicillium ulleungdoense D.H. Choi & J.G. Kim, Mycobiology 49: 48. 2021. [MB 835474]. — Type: KACC 48990 (holotype). Ex-type: KACC 48990. Infragen. class.: subgen. Aspergilloides sect. Sclerotiorum ser. Sclerotiorum. DNA barcodes: ITS = MN640087; BenA = MN737487; CaM = MN745074; RPB2 = MN756007.

Penicillium umkhoba Visagie & Yilmaz, Mycologia 115: 101. 2022. [MB 844189]. — Type: PREM 63222 (holotype). Ex-type: CMW 56200 = CBS 147457 = CN014D5. Infragen. class.: subgen. Aspergilloides sect. Sclerotiorum ser. Herqueorum. DNA barcodes: ITS = MT949912; BenA = MT957417; CaM = MT957459; RPB2 = MT957485.

Penicillium uttarakhandense Rajeshk., N. Ashtekar, Visagie, G. Anand & Yilmaz, Persoonia 46: 493. 2021. [MB 834093]. — Type: AMH 10225 (holotype). Ex-type: NFCCI 4808. Infragen. class.: subgen. Aspergilloides sect. Lanata-Divaricata ser. Simplicissima. DNA barcodes: ITS = MN967315; BenA = MN972443; CaM = MN972445; RPB2 = MN972447.

Notes: The phylogenetic relationship between P. brasilianum, P. onobense, P. paraherquei and P. skrjabinii is unresolved (Fig. 24 & Suppl. Fig. S22). Further studies including additional strains and/or extended datasets are needed. Here, we give benefit of doubt to the publisher.

Penicillium vaccaeorum Quintan., Mycopathologia 80: 77. 1982. [MB 109999]. — Type: CBS 148.83 (holotype). Ex-type: CBS 148.83 = DTO 009-E2 = CECT 2753. Infragen. class.: subgen. Aspergilloides sect. Citrina ser. Roseopurpurea. DNA barcodes: ITS = JN617689; BenA = JN606835; CaM = JN606543; RPB2 = JN606614.

Notes: Considered a synonym of P. sanguifluum by Houbraken, Frisvad & Samson (2011), this old name was more recently re-introduced by Torres-Garcia et al. (2022).

Penicillium vallebormidaense R.N. Barbosa & J.D.P. Bezerra, Persoonia 45: 371. 2020. [MB 837659]. — Type: CBS H-24527 (holotype). Ex-type: CBS 147064 = DTO 402-H5. Infragen. class.: subgen. Aspergilloides sect. Exilicaulis ser. Erubescentia. DNA barcodes: ITS = MT316359; BenA = MW115862; CaM = MW115863; RPB2 = MW115864.

Penicillium vickeryae Y.P. Tan & R.G. Shivas, Index Austral. Fungi 3: 10. 2022. [MB 900144]. — Type: BRIP 72552a (holotype). Ex-type: BRIP 72552a. Infragen. class.: subgen. Aspergilloides sect. Lanata-Divaricata ser. Simplicissima. DNA barcodes: ITS = OP903479; BenA = OP921966; CaM = n.a.; RPB2 = OP921965.

Penicillium vietnamense [nom. inval. Arts 40.7 & 40.8 (Shenzhen)] V.D. Nguyen & T.T. Pham, Mycobiology 50: 157. 2022. [MB 840587]. — Type: VTCC 930029 (holotype). Ex-type: VTCC 930029 = NTU DW14M = NITIA DW14M. Infragen. class.: subgen. Aspergilloides sect. Charlesia ser. Indica. DNA barcodes: ITS = MT102836; BenA = MT230561; CaM = ON209438; RPB2 = MT222288.

Synonym of: Penicillium chermesinum Biourge, Cellule 33: 284. 1923. [MB 260472].

Notes: This species was invalidly described. Phylogenies resolved P. vietnamense inside P. chermesinum (Fig. 17 & Suppl. Fig. S15), and even though CaM was not yet available for analysis, we consider P. vietnamense a synonym of P. chermesinum. We therefore choose not to validate this species.

Fig. 17.

Phylogenetic tree of Penicillium sections Aspergilloides, Charlesia, Cinnamopurpurea and Ramigena based on a concatenated dataset of BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. The tree was rooted to P. taxi. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by superscript T. See Suppl. Fig. S15.

Penicillium xyleborini Visagie & W.J. Nel, Persoonia 47: 349. 2021. [MB 840990]. — Type: PREM 63078 (holotype). Ex-type: CMW 56800 = CN001D4. Infragen. class.: subgen. Penicillium sect. Ramosum ser. Soppiorum. DNA barcodes: ITS = MW504356; BenA = MW480817; CaM = MW480823; RPB2 = MW480824.

Rasamsonia oblata (Pitt & A.D. Hocking) Yanai & Udagawa, Jap. J. Mycol. 61: 93. 2020. [MB 836491]. Basionym: Penicillium oblatum Pitt & A.D. Hocking, Mycologia 77: 810. 1985. [MB 104603]. — Type: FRR 2234. Ex-type: CBS 258.87 = IMI 288719 = NBRC 33091. DNA barcodes: ITS = LC546718; BenA = LC546729; CaM = LC546740; RPB2 = n.a.

Notes: Penicillium oblatum was considered a doubtful species by Yilmaz et al. (2014), because sequence data obtained for the ex-type strain CBS 258.87 (= FRR 2234) indicated it as a close relative of T. dendriticus, but its morphology clearly did not match the original description of Pitt & Hocking (1985a). The species was recently revised based on sequences from IMI 288719 and NBRC 33091 (both ex-type strains), which showed that it represents a unique species in Rasamsonia (Yanai et al. 2020). Penicillium oblatum was described as strictly monoverticillate (Pitt & Hocking 1985a), with some subterminal metulae produced. However, Yanai et al. (2020) description of P. oblatum based on IMI 288719 showed both mono- and biverticillate conidiophores.

Rasamsonia sabulosa (Pitt & A.D. Hocking) Yanai & Udagawa, Jap. J. Mycol. 61: 96. 2020. [MB 836492]. Basionym: Penicillium sabulosum Pitt & A.D. Hocking, Mycologia 77: 810. 1985. [MB 104604]. — Type: FRR 2743. Ex-type: ATCC 56984 = FRR 2743 = IMI 288715. DNA barcodes: ITS = LC546720; BenA = LC546726; CaM = LC546742; RPB2 = n.a.

Notes: Penicillium sabulosum and P. corynephorum (CBS 256.87^T) were until recently considered synonyms of P. smithii (CBS 276.83^T) (Visagie et al. 2016). Penicillium sabulosum was reviewed by Yanai et al. (2020) using ATCC 56984 and they showed it is a unique Rasamsonia species, and a new combination was subsequently made. Morphologically, P. sabulosum was described as growing slowly on CYA at 25 °C (3–6 mm) (Pitt & Hocking 1985a), while P. corynephorum grows 32–40 mm (Pitt & Hocking 1985b) and P. smithii 30–35 mm (after 2 wk) (Quintanilla 1982). Additionally, P. sabulosum has roughened phialides which was not reported for the other species. Penicillium smithii (section Exilicaulis) remains an accepted species with P. corynephorum its synonym.

Talaromyces africanus Houbraken, Pyrri & Visagie, J. Fungi 7 (11, no. 993): 10. 2021. [MB 841228]. — Type: CBS H-24874 (holotype). Ex-type: CBS 147340 = DTO 179-C5 = KAS 3859. Infragen. class.: sect. Trachyspermi. DNA barcodes: ITS = OK339610; BenA = OK338782; CaM = OK338808; RPB2 = OK338833.

Talaromyces albisclerotius B.D. Sun, A.J. Chen, Houbraken & Samson, MycoKeys 68: 88. 2020. [MB 833135]. — Type: CBS H-22837 (holotype). Ex-type: CBS 141839 = DTO 340-G5. Infragen. class.: sect. Trachyspermi. DNA barcodes: ITS = MN864276; BenA = MN863345; CaM = MN863322; RPB2 = MN863334.

Talaromyces aspriconidius B.D. Sun, A.J. Chen, Houbraken & Samson, MycoKeys 68: 90. 2020. [MB 833134]. — Type: CBS H-22833 (holotype). Ex-type: CBS 141835 = DTO 340-F8. Infragen. class.: sect. Talaromyces. DNA barcodes: ITS = MN864274; BenA = MN863343; CaM = MN863320; RPB2 = MN863332.

Talaromyces atkinsoniae Y.P. Tan, Bishop-Hurley, Bransgr. & R.G. Shivas, Persoonia 49: 323. 2022. [MB 845020]. — Type: BRIP 72528a (holotype). Ex-type: BRIP 72528a. Infragen. class.: sect. Talaromyces. DNA barcodes: ITS = OP059084; BenA = OP087524; CaM = n.a.; RPB2 = OP087523.

Talaromyces aureolinus L. Wang, Mycologia 113: 495. 2021. [MB 831405]. — Type: HMAS 248136 (holotype). Ex-type: AS 3.15865. Infragen. class.: sect. Talaromyces. DNA barcodes: ITS = MK837953; BenA = MK837937; CaM = MK837945; RPB2 = MK837961.

Talaromyces bannicus L. Wang, Mycologia 113: 498. 2021. [MB 831406]. — Type: HMAS 248133 (holotype). Ex-type: AS 3.15862. Infragen. class.: sect. Talaromyces. DNA barcodes: ITS = MK837955; BenA = MK837939; CaM = MK837947; RPB2 = MK837963.

Talaromyces brevis B.D. Sun, A.J. Chen, Houbraken & Samson, MycoKeys 68: 92. 2020. [MB 833132]. — Type: CBS H-22833 (holotype). Ex-type: CBS 141833. Infragen. class.: sect. Talaromyces. DNA barcodes: ITS = MN864269; BenA = MN863338; CaM = MN863315; RPB2 = MN863328.

Talaromyces calidominioluteus Houbraken & Pyrri, J. Fungi 7 (11, no. 993): 14. 2021. [MB 841229]. — Type: CBS H-24875 (holotype). Ex-type: CBS 147313 = DTO 052-G3. Infragen. class.: sect. Trachyspermi. DNA barcodes: ITS = OK339612; BenA = OK338786; CaM = OK338817; RPB2 = OK338837.

Talaromyces cavernicola V.C.S. Alves, J.D.P. Bezerra & R.N. Barbosa, Fungal Syst. Evol. 10: 157. 2022. [MB 846125]. — Type: URM 95155 (holotype). Ex-type: URM 8448 = FCCUFG 11. Infragen. class.: sect. Talaromyces. DNA barcodes: ITS = ON862935; BenA = OP672383; CaM = OP290543; RPB2 = OP290515.

Talaromyces chongqingensis X.C. Wang & W.Y. Zhuang, Biology 10: 10. 2021. [MB 570851]. — Type: HMAS 247849 (holotype). Ex-type: CGMCC 3.20482. Infragen. class.: sect. Trachyspermi. DNA barcodes: ITS = MZ358001; BenA = MZ361343; CaM = MZ361350; RPB2 = MZ361357.

Talaromyces gaditanus (C. Ramírez & A.T. Martínez) Houbraken & Soccio, J. Fungi 7 (11, no. 993): 17. 2021. [MB 841226]. Basionym: Penicillium gaditanum C. Ramírez & A.T. Martínez, Mycopathologia 74: 165. 1981. [MB 112521]. — Type: IJFM 5146 (holotype). Ex-type: CBS 169.81 = DTO 228-B8 = ATCC 42230 = IMI 253792 = VKM F-2188 = IJFM 5146. Infragen. class.: sect. Trachyspermi. DNA barcodes: ITS = MH861318; BenA = OK338775; CaM = OK338802; RPB2 = OK338827.

Notes: Penicillium gaditanum was considered a synonym of T. minioluteus by Yilmaz et al. (2014). A recent revision of T. minioluteus and related species revealed that P. gaditanum is a distinct species and was therefore combined in Talaromyces as T. gaditanus (Pyrri et al. 2021).

Talaromyces germanicus Houbraken & Pyrri, J. Fungi 7 (11, no. 993): 19. 2021. [MB 841227]. — Type: CBS H-24876 (holotype). Ex-type: CBS 147314 = DTO 055-D1. Infragen. class.: sect. Trachyspermi. DNA barcodes: ITS = OK339619; BenA = OK338799; CaM = OK338812; RPB2 = OK338845.

Talaromyces ginkgonis W.C. Wang & W.Y. Zhuang, J. Fungi 8 (7, no. 647): 7. 2022. [MB 570954]. — Type: HMAS 247853 (holotype). Ex-type: CGMCC 3.20698. Infragen. class.: sect. Trachyspermi. DNA barcodes: ITS = OL638158; BenA = OL689844; CaM = OL689846; RPB2 = OL689848.

Talaromyces gwangjuensis Hyang B. Lee & T.T.T. Nguyen, J. Fungi 7 (9, no. 722): 8. 2021. [MB 554801]. — Type: CNUFC HT19191 (holotype). Ex-type: CNUFC WT19-1. Infragen. class.: sect. Purpurei. DNA barcodes: ITS = MK766233; BenA = MZ318448; CaM = n.a.; RPB2 = MK912174.

Talaromyces haitouensis L. Wang, J. Fungi 8 (1, no. 36): 3. 2021. [MB 570868]. — Type: HMAS 350335 (holotype). Ex-type: AS 3.16101 = HR1-7. Infragen. class.: sect. Talaromyces. DNA barcodes: ITS = MZ045695; BenA = MZ054634; CaM = MZ054637; RPB2 = MZ054631.

Talaromyces koreanus [as ‘koreana’] Hyang B. Lee, J. Fungi 7 (9, no. 722): 9. 2021. [MB 648574]. — Type: CNUFC HT19213 (holotype). Ex-type: CNUFC YJW2-13. Infragen. class.: sect. Helici. DNA barcodes: ITS = MZ315100; BenA = MZ318450; CaM = MZ332529; RPB2 = MZ332533.

Talaromyces nanjingensis X.R. Sun, X.Q. Wu & W. Wei, J. Fungi 8 (2, no. 155): 18. 2022. [MB 837590]. — Type: CCTCC-M 2012167 (holotype). Ex-type: CCTCC-M 2012167. Infragen. class.: sect. Talaromyces. DNA barcodes: ITS = MW130720; BenA = MW147759; CaM = MW147760; RPB2 = MW147762.

Notes: The phylogenies of T. nanjingensis and its close relatives T. brevis and T. liani proved to be problematic (Fig. 31 & Suppl. Fig. S29). BenA resolved T. nanjingensis and T. liani as close relatives, while T. brevis is distantly related. Based on CaM, T. nanjingensis, T. liani and T. brevis are phylogenetically similar species. Finally, T. nanjingensis and T. brevis share RPB2 sequences, and these species are closely related to T. liani. A taxonomic revision including a broader strain sampling is required to assess this species complex, but for the time being, we accept T. nanjingensis as a distinct species.

Fig. 31.

Phylogenetic tree of Talaromyces section Talaromyces, based on a concatenated dataset of BenA, CaM, RPB2, and ITS. Strains of recently described species are shown in bold coloured text. The tree was rooted to T. helicus. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by superscript T. See Suppl. Fig. S29.

Talaromyces peaticola Jian Q. Tian & Jing Z. Sun, Stud. Fungi 6: 396. 2021. [MB 553909]. — Type: HMAS 247296 (holotype). Ex-type: CGMCC 3.18620. Infragen. class.: sect. Trachyspermi. DNA barcodes: ITS = MF135613; BenA = MF284705; CaM = MF284703; RPB2 = MF284704.

Synonym of: Talaromyces diversus (Raper & Fennell) Samson, Yilmaz & Frisvad, Stud. Mycol. 70: 175. 2011. [MB 560649].

Notes: The BenA phylogeny resolved T. peaticola inside T. diversus (Fig. 33 & Suppl. Fig. S30) and we thus consider it a synonym of the latter.

Talaromyces penicillioides L. Wang, Mycologia 113: 501. 2021. [MB 831407]. — Type: HMAS 248132 (holotype). Ex-type: AS 3.15822. Infragen. class.: sect. Talaromyces. DNA barcodes: ITS = MK837956; BenA = MK837940; CaM = MK837948; RPB2 = MK837964.

Talaromyces phuphaphetensis Nuankaew, Chuaseehar. & Somrith., J. Fungi 8 (8: no. 825): 7. 2022. [MB 844613]. — Type: BBH 49306 (holotype). Ex-type: TBRC 16281 = CV00299. Infragen. class.: sect. Trachyspermi. DNA barcodes: ITS = ON692803; BenA = ON706960; CaM = ON706962; RPB2 = ON706964.

Talaromyces pulveris Crous, Persoonia 45: 293. 2020. [MB 837837]. — Type: CBS H-24417 (holotype). Ex-type: CBS 146831 = CPC 38523 = MUCL pd8781 = DTO 432-H1. Infragen. class.: sect. Purpurei. DNA barcodes: ITS = MW175345; BenA = MW173136; CaM = MW173099; RPB2 = MW173115.

Talaromyces resedanus (McLennan & Ducker) A.J. Chen, Houbraken & Samson, MycoKeys 68: 96. 2020. [MB 811695]. Basionym: Penicillium resedanum McLennan & Ducker, Austral. J. Bot. 2: 360. 1954. [MB 302422] — Type: IMI 062877 (holotype). Ex-type: CBS 181.71 = DTO 376-A7 = ATCC 22356 = FRR 578 = IMI 062877 = NRRL 578. Infragen. class.: sect. Subinflati. DNA barcodes: ITS = MN431413; BenA = MN969436; CaM = MN969355; RPB2 = MN969214.

Notes: Although an ITS sequence resolved P. resedanum in Talaromyces, Yilmaz et al. (2014) considered it a doubtful species because the CBS strain was not viable. The strain was subsequently successfully revived and formally combined as Talaromyces resedanus by Sun et al. (2020a). The identifier “MB 302422” cited in Sun et al. (2020a) is that of the basionym (P. resedanum), but the species is still validly published (Art. F.5.6 San Juan Chapter F) (May et al. 2019) because “MB 811695” was issued by MycoBank prior to the publication of the name. Sun et al. (2020a) considered T. omanensis (SQUCC 13153^T) as synonym of T. resedanus based on morphology and DNA sequence data, which we follow here.

Talaromyces rosorhizae [as ‘rosarhiza’] [nom. inval. Art. 40.8 (Shenzhen)] H. Zhang & Y.L. Jiang, Biodivers. Data J. 9 (e70088): 11. 2021. [MB 662132]. — Type: GUCC 190040.1 (holotype). Ex-type: GUCC 190040.1. Infragen. class.: sect. Talaromyces. DNA barcodes: ITS = MZ221603; BenA = MZ333143; CaM = MZ333137; RPB2 = MZ333141.

Synonym of: Talaromyces penicillioides L. Wang, Mycologia 113: 501. 2021. [MB 831407].

Notes: This species was invalidly described. Phylogenies resolved T. rosorhizae and T. penicillioides, both introduced in 2021, in the same clade (Fig. 31 & Suppl. Fig. S29). We therefore choose not to validate this species.

Talaromyces rufus B.D. Sun, A.J. Chen, Houbraken & Samson, MycoKeys 68: 99. 2020. [MB 833133]. — Type: CBS H-22832 (holotype). Ex-type: CBS 141834 = DTO 349-D7 = CGMCC 3.13203. Infragen. class.: sect. Talaromyces. DNA barcodes: ITS = MN864272; BenA = MN863341; CaM = MN863318; RPB2 = MN863331.

Talaromyces samsonii (Quintan.) Houbraken & Pyrri, J. Fungi 7 (11, no. 993): 25. 2021. [MB 841230]. Basionym: Penicillium samsonii Quintan., Mycopathologia 91: 69. 1985. [MB 105614] — Type: CBS H-24877 (holotype). Ex-type: CBS 137.84 = DTO 304-C3 = DTO 169-G6 = CECT 2772 = IMI 282404 = IMI 327872. Infragen. class.: sect. Trachyspermi. DNA barcodes: ITS = MH861709; BenA = OK338798; CaM = OK338824; RPB2 = OK338844.

Notes: Penicillium samsonii was considered a synonym of T. minioluteus by Yilmaz et al. (2014). A recent revision of T. minioluteus and related species showed that P. samsonii is a distinct species and therefore combined in Talaromyces as T. samsonii (Pyrri et al. 2021).

Talaromyces santanderensis [nom. inval. Art. 40.8 (Shenzhen)] B.E. Guerra-Sierra & L.A. Arteaga-Figueroa, J. Fungi 8 (10, no. 1042): 5. 2022. [MB 845323]. — Type: CBUDES:UDES:3068 (holotype). Ex-type: HF05. Infragen. class.: sect. Talaromyces. DNA barcodes: ITS = OP082331; BenA = OP067657; CaM = OP067656; RPB2 = OP067655.

Synonym of: Talaromyces lentulus X.-Z. Jiang & L. Wang, Sci. Rep. 8 (no. 4932): 3. 2018. [MB 824519].

Notes: This species was invalidly described. All phylogenies resolve T. santanderensis inside the T. lentulus clade (Fig. 31 & Suppl. Fig. S29). We consider T. santaderensis a synonym of T. lentulus and therefore choose not to validate this species.

Talaromyces satunensis Nuankaew, Chuaseehar. & Somrith., J. Fungi 8 (8: no. 825): 8. 2022. [MB 844614]. — Type: BBH 49305 (holotype). Ex-type: TBRC 16246 = CV00055. Infragen. class.: sect. Trachyspermi. DNA barcodes: ITS = ON692804; BenA = ON706961; CaM = ON706963; RPB2 = n.a.

Talaromyces saxoxalicus J. Trovão, F. Soares, I. Tiago & A. Portugal, Int. J. Syst. Evol. Microbiol. 71 (12, no. 5175): 4. 2021. [MB 834608]. — Type: MUM-H 20.30 (holotype). Ex-type: MUM 20.30. Infragen. class.: sect. Purpurei. DNA barcodes: ITS = MT039882; BenA = MT052003; CaM = n.a.; RPB2 = MT052004.

Talaromyces shilinensis X.C. Wang & W.Y. Zhuang, J. Fungi 8 (7, no. 647): 10. 2022. [MB 570955]. — Type: HMAS 247854 (holotype). Ex-type: CGMCC 3.20699. Infragen. class.: sect. Talaromyces. DNA barcodes: ITS = OL638159; BenA = OL689845; CaM = OL689847; RPB2 = OL689849.

Talaromyces sparsus L. Wang, Mycologia 113: 501. 2021. [MB 831409]. — Type: HMAS 248135 (holotype). Ex-type: AS 3.16003. Infragen. class.: sect. Talaromyces. DNA barcodes: ITS = MT077182; BenA = MT083924; CaM = MT083925; RPB2 = MT083926.

Talaromyces teleomorphus [as ‚teleomorpha‘] Hyang B. Lee, Frisvad, P.M. Kirk, H.J. Lim & T.T.T. Nguyen, J. Fungi 7 (9, no. 722): 11. 2021. [MB 648560]. — Type: CNUFC HT19251 (holotype). Ex-type: CNUFC YJW2-5. Infragen. class.: sect. Helici. DNA barcodes: ITS = MZ315102; BenA = MZ318452; CaM = MZ332531; RPB2 = MZ332535.

Talaromyces tenuis B.D. Sun, A.J. Chen, Houbraken & Samson, MycoKeys 68: 86. 2020. [MB 833136]. — Type: CBS H-22838 (holotype). Ex-type: CBS 141840 = DTO 340-G9. Infragen. class.: sect. Tenues. DNA barcodes: ITS = MN864275; BenA = MN863344; CaM = MN863321; RPB2 = MN863333.

Talaromyces wushanicus X.C. Wang & W.Y. Zhuang, Biology 10 (8, no. 745): 15. 2021. [MB 570852]. — Type: HMAS 247848 (holotype). Ex-type: CGMCC 3.20481. Infragen. class.: sect. Talaromyces. DNA barcodes: ITS = MZ356356; BenA = MZ361347; CaM = MZ361354; RPB2 = MZ361361.

Talaromyces yunnanensis Doilom & C.F. Liao, Front. Microbiol. 11 (no. 585215): 18. 2020. [MB 557863]. — Type: MFLU 20-0434 (holotype). Ex-type: KUMCC 18-0208. Infragen. class.: sect. Talaromyces. DNA barcodes: ITS = MT152339; BenA = MT161683; CaM = MT178251; RPB2 = n.a.

Talaromyces zhenhaiensis L. Wang, J. Fungi 8 (1, no. 36): 7. 2021. [MB 570869]. — Type: HMAS 350336 (holotype). Ex-type: AS 3.16102 = ZH3-18. Infragen. class.: sect. Talaromyces. DNA barcodes: ITS = MZ045697; BenA = MZ054636; CaM = MZ054639; RPB2 = MZ054633.

Xerochrysium bohemicum Kubátová & Hubka, Persoonia 46: 519. 2021. [MB 839239]. — Type: PRM 954080. Ex-type: CCF 3693 = CBS 147157. DNA barcodes: ITS = MW798184; BenA = n.a.; CaM = n.a.; RPB2 = n.a.

Xerochrysium coryli (Crous & Decock) Visagie & Houbraken, published here. [MB 849335]. Basionym: Paraxerochrysium coryli Crous & Decock, Persoonia 47: 261. 2021. [MB 841830]. — Type: CBS H-24853 (holotype). Ex-type: CBS 148314 = CPC 41272 = MUCL 58103. DNA barcodes: ITS = OK664748; BenA = OK651216; CaM = n.a.; RPB2 = OK651178.

Notes: See notes for Xerochrysium coryli above in the description.

Accepted species published before 2020 not included in Houbraken et al. (2020)

The following species were not included the list published by Houbraken et al. (2020).

Aspergillus coreanus S.B. Hong, Frisvad & Samson, Int. J. Syst. Evol. Microbiol. 56: 485. 2006. [MB 521268]. — Type: CBS 117059 (holotype). Ex-type: CBS 117059 = NRRL 35590 = KACC 41659. Infragen. class.: subgen. Fumigati sect. Fumigati ser. Fumigati. DNA barcodes: ITS = JN943570; BenA = AY870758; CaM = AY870718; RPB2 = n.a.

Species with no or inconclusive sequence data available

The following species were accepted by Houbraken et al. (2020). However, since no sequence data are available for these, we exclude these names until their phylogenetic relationships can be verified.

Aspergillus argenteus [as ‘argentum’] J.N. Rai & H.J. Chowdhery, Kavaka 7: 19. 1980. [MB 116063]. — Type: MLLU 104. Ex-type: unknown.

Aspergillus beijingensis D.M. Li, Y. Horie, Yu X. Wang & R.Y. Li, Mycoscience 39: 299. 1998. [MB 446575]. — Type: CBM FD-285. Ex-type: CBM FD-285.

Aspergillus collembolorum Dörfelt & A.R. Schmidt, Mycol. Res. 109: 956, figs 1–9. 2005. [MB 344420]. — Type: Russia: Kaliningrad (Koenigsberg), in succinum Balticum, in exemplare subordines Entomobryomorpha (Collembola), C. & H. W. Hoffeins (coll. Hoffeins, Hamburg, no. 805, holotypus). Ex-type: unknown.

Aspergillus crassihyphae Wadhwani & N. Mehrotra, Indian Bot. Reporter: 52. 1985. [MB 105070]. — Type: unknown. Ex-type: unknown.

Aspergillus curviformis H.J. Chowdhery & J.N. Rai, Nova Hedwigia 32: 231. 1980. [MB 118396]. — Type: unknown. Ex-type: unknown.

Aspergillus ellipsoideus J.N. Rai & H.J. Chowdhery, Kavaka 7: 17. 1980. [MB 116064]. — Type: MLLU 107. Ex-type: unknown.

Aspergillus maritimus Samson & W. Gams, Adv. Pen. Asp. Syst.: 43. 1986. [MB 114709]. — Type: CBS 186.77. Ex-type: CBS 186.77.

Aspergillus qizutongii D.M. Li, Y. Horie, Yu X. Wang & R.Y. Li, Mycoscience 39: 301. 1998. [MB 446576]. — Type: CBM FD-284. Ex-type: CBM FD-284.

Aspergillus raianus [as ‘raianum’] H.J. Chowdhery, Curr. Sci. 48: 953. 1979. [MB 309239]. — Type: MLLU 110. Ex-type: unknown.

Aspergillus subunguis Wadhwani, Dudeja & M.P. Srivast., Curr. Sci. 53: 443. 1984. [MB 105934]. — Type: IMI 254637. Ex-type: IMI 254637.

Aspergillus tapirirae C. Ram & A. Ram, Atti Reale Accad. Sci. Napoli: 100. 1972. [MB 309245]. — Type: IMUFPe 2175. Ex-type: unknown.

Aspergillus vinosobubalinus Udagawa, Kamiya & Kaori Osada, Trans. Mycol. Soc. Japan 34: 255. 1993. [MB 361186]. — Type: CBM BF-33501. Ex-type: CBM BF-33501.

Aspergillus wangduanlii D.M. Li, Y. Horie, Yu X. Wang & R.Y. Li, Mycoscience 39: 302. 1998. [MB 447107]. — Type: CBM FD-283. Ex-type: CBM FD-283.

Penicilliopsis africana Samson & Seifert, Adv. Pen. Asp. Syst.: 408. 1985. [MB 114759]. — Type: unknown. Ex-type: unknown.

Penicilliopsis pseudocordyceps H.M. Hsieh & Y.M. Ju, Mycologia 9: 541. 2002. [MB 484663]. — Type: HAST (Taiwan) Hsieh & Ju 89112611. Ex-type: BCRC 33730.

Penicillium asymmetricum (Subramanian & Sudha) Houbraken & Samson, Stud. Mycol. 70: 47. 2011. [MB 561963]. Basionym: Thysanophora asymmetrica Subram. & Sudha, Kavaka 13: 88. 1985. [MB 135502]. — Type: unknown. Ex-type: unknown.

Penicillium coniferophilum Houbraken & Samson, Stud. Mycol. 70: 47. 2011. [MB 561968]. Basionym: Thysanophora striatispora G.L. Barron & W.B. Cooke, Mycopathologia et Mycologia Applicata 40 (3-4): 353. 1970. [MB 324607]. — Type: unknown. Ex-type: unknown.

Penicillium glaucoalbidum (Desmazières) Houbraken & Samson, Stud. Mycol. 70: 47. 2011. [MB 561965]. Basionym: Sclerotium glaucoalbidum Desm., Ann. Sci. Nat. Bot. 16: 329. 1851. [MB 212120]. — Type: unknown. Ex-type: unknown.

Penicillium longisporum (W.B. Kendr.) Houbraken & Samson, Stud. Mycol. 70: 47. 2011. [MB 561966]. Basionym: Thysanophora longispora W.B. Kendr., Can. J. Bot. 39 (4): 826. 1961. [MB 340086]. — Type: DAOM 63073. Ex-type: CBS 354.62 = DAOM 63073 = MUCL 4168.

Phialomyces fusiformis G. Delgado & Decock, Mycologia 95: 896. 2003. [MB 489106]. — Type: MUCL 43747. Ex-type: MUCL 43747.

Thermoascus taitungiacus K.Y. Chen & Z.C. Chen, Mycotaxon 60: 226. 1996. [MB 436720]. — Type: TAl-Mycology K-Y Chen 8709-2. Ex-type: unknown.

Species in Houbraken et al. (2020) synonymised afterwards

Aspergillus amoenus M. Roberg, Hedwigia 70: 138. 1931. [MB 250654]. — Type: Munster, 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. DNA barcodes: ITS = EF652480; BenA = JN853946; CaM = JN854035; RPB2 = JN853824.

Synonym of: Aspergillus versicolor (Vuill.) Tirab., Ann. Bot. (Roma) 7: 9. 1908. [MB 172159] (Sklenář et al. 2022).

Aspergillus austroafricanus Ž. Jurjević, S.W. Peterson & B.W. Horn, SIM 3: 67. 2012. [MB 800597]. — Type: BPI 880914. Ex-type: CBS 145748 = NRRL 233 = DTO 225-D8. Infragen. class.: subgen. Nidulantes sect. Nidulantes ser. Versicolores. DNA barcodes: ITS = JQ301891; BenA = JN853963; CaM = JN854025; RPB2 = JN853814.

Synonym of: Aspergillus versicolor (Vuill.) Tirab., Ann. Bot. (Roma) 7: 9. 1908. [MB 172159] (Sklenář et al. 2022).

Aspergillus capensis Visagie, Hirooka & Samson, Stud. Mycol. 78: 105. 2014. [MB 809193]. — Type: CBS H-21810. Ex-type: CBS 138188 = DTO 179-E6. Infragen. class.: subgen. Circumdati sect. Flavipedes ser. Flavipedes. DNA barcodes: ITS = KJ775550; BenA = KJ775072; CaM = KJ775279; RPB2 = KP987020.

Synonym of: Aspergillus iizukae Sugiy., J. Fac. Sci. Univ. Tokyo, Sect. 3 9: 390. 1967. [MB 326636] (Sklenář et al. 2021).

Aspergillus costaricensis [as ‘costaricaensis’] Samson & Frisvad, Stud. Mycol. 50: 52. 2004. [MB 369151]. — Type: CBS H-13437. Ex-type: CBS 115574 = IBT 23401 = CECT 20579 = ITEM 7555. Infragen. class.: subgen. Circumdati sect. Nigri ser. Nigri. DNA barcodes: ITS = DQ900602; BenA = FJ629277; CaM = FN594545; RPB2 = HE984361.

Synonym of: Aspergillus tubingensis Mosseray, La Cellule 43: 245. 1934. [MB 255209] (Bian et al. 2022).

Aspergillus cvjetkovicii Jurjevic, S.W. Peterson & B.W. Horn, SIM 3: 69. 2012. [MB 800599]. — Type: BPI 880909. Ex-type: NRRL 227 = CBS 599.65. Infragen. class.: subgen. Nidulantes sect. Nidulantes ser. Versicolores. DNA barcodes: ITS = EF652440; BenA = EF652264; CaM = EF652352; RPB2 = EF652176.

Synonym of: Aspergillus creber Jurjevic, S.W. Peterson & B.W. Horn, SIM 3: 69. 2012. [MB 800598] (Sklenář et al. 2022).

Aspergillus fructus Jurjevic, S.W. Peterson & B.W. Horn, SIM 3: 70. 2012. [MB 800600]. — Type: BPI 880915. Ex-type: NRRL 239 = CBS 584.65. Infragen. class.: subgen. Nidulantes sect. Nidulantes ser. Versicolores. DNA barcodes: ITS = EF652449; BenA = EF652273; CaM = EF652361; RPB2 = EF652185.

Synonym of: Aspergillus versicolor (Vuill.) Tirab., Ann. Bot. (Roma) 7: 9. 1908. [MB 172159] (Sklenář et al. 2022).

Aspergillus griseoaurantiacus Visagie, Hirooka & Samson, Stud. Mycol. 78: 112. 2014. [MB 809197]. — Type: CBS H-21814. Ex-type: CBS 138191 = DTO 267D8. Infragen. class.: subgen. Nidulantes sect. Nidulantes ser. Versicolores. DNA barcodes: ITS = KJ775553; BenA = KJ775086; CaM = KJ775357; RPB2 = KU866988.

Synonym of: Aspergillus versicolor (Vuill.) Tirab., Ann. Bot. (Roma) 7: 9. 1908. [MB 172159] (Sklenář et al. 2022).

Aspergillus hongkongensis C.C. Tsang, T.W.S. Hui, K.C. Lee, J.H.K. Chen, A.H.Y. Ngan, E.W.T. Tam, J.F.W. Chan, A.L. Wu, M. Cheung, B.P.H. Tse, A.K.L. Wu, C.K.C. Lai, D.N.C. Tsang, T.L. Que, C.W. Lam, K.Y. Yuen, S.K.P. Lau & P.C.Y. Woo, Diagn. Microbiol. Infect. Dis. 84: 130. 2016. [MB 810279]. — 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. DNA barcodes: ITS = AB987907; BenA = LC000552; CaM = MN969320; RPB2 = LC000578.

Synonym of: Aspergillus versicolor (Vuill.) Tirab., Ann. Bot. (Roma) 7: 9. 1908. [MB 172159] (Sklenář et al. 2022).

Aspergillus jensenii Jurjevic, S.W. Peterson & B.W. Horn, SIM 3: 70. 2012. [MB 800601]. — Type: BPI 880910. Ex-type: NRRL 58600. Infragen. class.: subgen. Nidulantes sect. Nidulantes ser. Versicolores. DNA barcodes: ITS = JQ301892; BenA = JN854007; CaM = JN854046; RPB2 = JN853835.

Synonym of: Aspergillus creber Jurjevic, S.W. Peterson & B.W. Horn, SIM 3: 69. 2012. [MB 800598] (Sklenář et al. 2022).

Aspergillus neoniger Varga, Frisvad & Samson, Stud. Mycol. 69: 16. 2011. [MB 560390]. — Type: CBS H-20630. Ex-type: CBS 115656 = NRRL 62634. Infragen. class.: subgen. Circumdati sect. Nigri ser. Nigri. DNA barcodes: ITS = FJ491682; BenA = FJ491691; CaM = FJ491700; RPB2 = KC796429.

Synonym of: Aspergillus tubingensis Mosseray, La Cellule 43: 245. 1934. [MB 255209] (Bian et al. 2022).

Aspergillus pepii Despot, Kocsubé, Varga & Klarić, Mycol. Prog. 16: 67. 2017. [MB 817073]. — Type: SZMC 23791 (holotype). Ex-type: CBS 142028 = MFBF AV11051B IX = SZMC 22333. Infragen. class.: subgen. Nidulantes sect. Nidulantes ser. Versicolores. DNA barcodes: ITS = KU613368; BenA = KU613371; CaM = KU613365; RPB2 = n.a.

Synonym of: Aspergillus versicolor (Vuill.) Tirab., Ann. Bot. (Roma) 7: 9. 1908. [MB 172159] (Sklenář et al. 2022).

Aspergillus piperis Samson & Frisvad, Stud. Mycol. 50: 57. 2004. [MB 500009]. — Type: CBS H-13434. Ex-type: CBS 112811 = IBT 24630 = IBT 26239 = NRRL 62631. Infragen. class.: subgen. Circumdati sect. Nigri ser. Nigri. DNA barcodes: ITS = EU821316; BenA = FJ629303; CaM = EU163267; RPB2 = KC796427.

Synonym of: Aspergillus luchuensis Inui, Journal of the Faculty of Science, Imperial University of Tokyo 15: 469. 1901. [MB 151291] (Bian et al. 2022).

Aspergillus protuberus Munt.-Cvetk., Mikrobiologia 5: 119. 1968. [MB 326650]. Basionym: Aspergillus versicolor var. protuberus (Munt.-Cvetk.) Kozak., Mycological Papers 161: 139. 1989. [MB 127752]. — Type: CBS 602.74. Ex-type: CBS 602.74 = NRRL 3505 = ATCC 18990 = QM 9804. Infragen. class.: subgen. Nidulantes sect. Nidulantes ser. Versicolores. DNA barcodes: ITS = EF652460; BenA = EF652284; CaM = EF652372; RPB2 = EF652196.

Synonym of: Aspergillus versicolor (Vuill.) Tirab., Ann. Bot. (Roma) 7: 9. 1908. [MB 172159] (Sklenář et al. 2022).

Aspergillus puulaauensis Jurjevic, S.W. Peterson & B.W. Horn, SIM 3: 71. 2012. [MB 800602]. — Type: BPI 880911. Ex-type: CBS 145750 = NRRL 35641 = DTO 225-G5. Infragen. class.: subgen. Nidulantes sect. Nidulantes ser. Versicolores. DNA barcodes: ITS = JQ301893; BenA = JN853979; CaM = JN854034; RPB2 = JN853823.

Synonym of: Aspergillus creber Jurjevic, S.W. Peterson & B.W. Horn, SIM 3: 69. 2012. [MB 800598] (Sklenář et al. 2022).

Aspergillus tabacinus Nakaz., Y. Takeda, Simo & A. Watan., J. Agric. Chem. Soc. Japan 10: 177. 1934. [MB 539544]. — Type: CBS H-24287. Ex-type: CBS 122718 = NRRL 4791 = IFO 4098 = QM 9766 = WB 4791. Infragen. class.: subgen. Nidulantes sect. Nidulantes ser. Versicolores. DNA barcodes: ITS = EF652478; BenA = EF652302; CaM = EF652390; RPB2 = EF652214.

Synonym of: Aspergillus versicolor (Vuill.) Tirab., Ann. Bot. (Roma) 7: 9. 1908. [MB 172159] (Sklenář et al. 2022).

Aspergillus tennesseensis Jurjevic, S.W. Peterson & B.W. Horn, SIM 3: 73. 2012. [MB 800604]. — Type: BPI 880917. Ex-type: CBS 145752 = NRRL 13150 = DTO 225-F5. Infragen. class.: subgen. Nidulantes sect. Nidulantes ser. Versicolores. DNA barcodes: ITS = JQ301895; BenA = JN853976; CaM = JN854017; RPB2 = JN853806. (Sklenář et al. 2022)

Synonym of: Aspergillus creber Jurjevic, S.W. Peterson & B.W. Horn, SIM 3: 69. 2012. [MB 800598] (Sklenář et al. 2022).

Aspergillus venenatus Jurjevic, S.W. Peterson & B.W. Horn, SIM 3: 73. 2012. [MB 800605]. — Type: BPI 880916. Ex-type: CBS 145753 = NRRL 13147 = DTO 225-F4. Infragen. class.: subgen. Nidulantes sect. Nidulantes ser. Versicolores. DNA barcodes: ITS = JQ301896; BenA = JN854003; CaM = JN854014; RPB2 = JN853803.

Synonym of: Aspergillus creber Jurjevic, S.W. Peterson & B.W. Horn, SIM 3: 69. 2012. [MB 800598] (Sklenář et al. 2022).

Aspergillus welwitschiae (Bres.) Henn. apud Wehmer, Centralbl. Bakteriol. Parasitenk., 2. Abth. 18: 294. 1907. [MB 490584]. Basionym: Ustilago welwitschiae Bres., Bol. Soc. Brot. 11: 68. 1893. [MB 176748]. — Type: CBS 139.54. Ex-type: CBS 139.54. Infragen. class.: subgen. Circumdati sect. Nigri ser. Nigri. DNA barcodes: ITS = FJ629340; BenA = MN969369; CaM = KC480196; RPB2 = MN969100.

Synonym of: Aspergillus niger Tiegh., Ann. Sci. Nat., Bot. ser. 5 8: 240. 1867. [MB 284309] (Bian et al. 2022).

Penicillium cluniae [nom. inval. Arts 40.7 (Shenzhen)] Quintan., Av. Aliment. Mejora Anim. 30: 174. 1990. [MB 130240].

Synonym of: Penicillium soli Doilom, C.F. Liao & D. Pem, Front. Microbiol. 11 (no. 585215): 13. 2020. [MB 557862] (this paper).

Talaromyces omanensis Halo, Maharachch., Al-Yahyai & Al-Sadi, Phytotaxa 404: 192. 2019. [MB 830302]. — Type: SQU H-106 (holotype). Ex-type: SQUCC 13153. Infragen. class.: sect. Subinflati. DNA barcodes: ITS = MH784402; BenA = MH794502; CaM = MH794503; RPB2 = n.a.

Synonym of: Talaromyces resedanus (McLennan & Ducker) A.J. Chen, Houbraken & Samson, MycoKeys 68: 96. 2020. [MB 811695] (Sun et al. 2020a).

DISCUSSION

In this study, we review Eurotiales species described since the previous accepted species list by Houbraken et al. (2020) up until 31 December 2022. Species were revised based on a phylogenetic species concept and the application of Genealogical Concordance Phylogenetic Species Recognition (GCPSR; Taylor et al. 2000), which is the most widely used species concept in Eurotiales. However, we believe that the polyphasic approach (or consilient concept of species) provides a holistic view of species and therefore consider it important to include morphology and, where possible, extrolite (secondary metabolite) as well as whole-genome sequence data, which will play an important role in the near future. Taxonomists dealing with Eurotiales have historically kept classifications practical and useful to the end user, an approach that has served the community dealing with these important fungi well. In case of doubt about the delimitation of species during this review, we have given preference to the publisher and accepted the name. However, we have classified newly published species as synonyms when the currently available data clearly do not support their novelty, while in cases where species were published without or with poor quality sequence data, we considered them doubtful because their future identification would not be possible. This follows Charles Thom’s idea that a species is only useful if others can identify it (Thom 1954). During this study, several trends were noted where descriptions can go wrong. We discuss these trends in more detail below and provide recommendations to avoid them in the future.

The ICNafp (Turland et al. 2018) governs the naming of fungi and sets out the rules for the valid and legitimate publication of names. Of the 160 new Eurotiales names introduced, 10 were invalid because they either did not comply with Art. 40.7 (the single herbarium, collection or institution in which the type is conserved must be specified), Art. 40.8 (the protologue must include a statement that the culture is preserved in a metabolically inactive state), or in some cases both. In addition to ICNafp, there are other helpful resources to guide valid publication of names. These include ‘The Code Decoded’ (Turland 2019) and several publications on behalf of the International Commission on the Taxonomy of Fungi (ICTF) outlining the best practises for describing a new species (Sigler & Hawksworth 1987, Seifert & Rossman 2010, Aime et al. 2021).

DNA sequence comparisons have become the most important tool for the classification and identification of fungal species. A reliable and complete reference database is crucial for this approach. The openly available and mostly complete ex-type DNA sequence reference sets (Samson et al. 2014, Visagie et al. 2014, Yilmaz et al. 2014, Houbraken et al. 2020) for ITS, BenA, CaM and RPB2 are a great resource. However, as critical as they may be, the ex-type sequences only serve as anchor points for the species, while additional sequences are needed to capture infraspecies variation and delimit species. Bian et al. (2022), Glässnerová et al. (2022) and Sklenář et al. (2022) emphasise the importance of these sequences, which should preferably come from strains isolated from different substrates and (ecological) regions. We therefore recommend not to compare putative new species only with ex-type strains in phylogenies where more extensive data is available as this does not provide information on species boundaries and can lead to incorrect conclusions. An example where this was a problem was the description of Penicillium tolerans as a close relative of P. sclerotiorum (Tan & Shivas 2022). The latter was subject to several taxonomic revisions with phylogenies that placed P. sclerotiorum strains into two closely related clades. However, both Rivera & Seifert (2011) and Visagie et al. (2013) concluded that from a morphological and ecological point of view there is no reason to consider these clades as separate species. In the case of P. tolerans, the ex-type strain (BRIP 64090a) resolves into the clade without the P. sclerotiorum ex-type strain (CBS 287.36) (Fig. 27). The phylogenies presented by Tan & Shivas (2022) did not include a comparison with all closely related sequences and left unmentioned their conclusions about these other ‘P. sclerotiorum’ strains and the reasons why the clade with P. tolerans should be considered distinct. In view of this, we consider P. tolerans a synonym of P. sclerotiorum.

Homology searches are a popular tool to quickly identify strains at the species level and generally works well if you get hits with well curated sequences. If a search does not find highly similar sequences, this may be due to several reasons. The sequence in question could belong to an unsequenced described species, represent an unsequenced genotype of an existing species or indicate a new species. The list of accepted species is very informative because, at least for Eurotiales, we can practically ignore the names not accepted and we have reference sequences of all known species. Dissimilarity higher than 4 %, especially in CaM and RPB2, is a relatively safe indication of new species in Aspergillus (Bian et al. 2022). Nevertheless, the results of homology searches should be interpreted with caution and should not be confused with phylogenetic relatedness. We recommend to always confirm new species or genotypes by phylogenetic analyses and not to rely only on homology search results.

Examples of low-quality sequence data used for species descriptions were noted for several recently described species. This has already been discussed for fungi (Nilsson et al. 2017) and remains problematic, even though careful contig assembly and proper trimming and base-calling should resolve this issue. In our analyses, the ITS alignments generated for Aspergillus sect. Circumdati revealed suspect sequences for both A. curvatus and A. gaarensis. Similarly, the sequences available for the recently published A. pseudopiperis and A. pseudotubingensis were considered to be of low quality (Bian et al. 2022), while Visagie et al. (2021) illustrated the same problem in the descriptions of P. attenuatum, P. ochotense and P. piltunense (Kirichuk, Pivkin & Matveeva 2016). Aspergillus pseudopiperis were considered a synonym of A. tubingensis (Bian et al. 2022), while P. attenuatum, P. ochotense and P. piltunense was considered synonyms of P. antarcticum (Visagie et al. 2021). In most cases, however, low-quality sequences make classification and future identification impossible, so that the species must be considered doubtful.

As mentioned above, the phylogenetic species concept and the application of GCPSR is important for the taxonomy of Eurotiales within our preferred consilient concept of species. GCPSR has largely proven to be a consistent approach to defining species boundaries. In principle, a species is delimited in the narrowest sense at the most recent node that forms consistently between different gene phylogenies (Taylor et al. 2000). In Eurotiales, the four most commonly used gene regions are ITS, BenA, CaM and RPB2, and we recommend that these are at least sequenced when describing new species. Recently, several Eurotiales species were introduced that lacked an ITS sequence, the official DNA barcode for fungi (Schoch et al. 2012). Admittedly, ITS is generally not informative at the species level, but we consider it good practice to sequence ITS at least for the ex-type strain of a new species. Several species have also been introduced without sequence data for all recommended genes. We cannot stress enough how important this is for the taxonomy of Eurotiales at a time when mycologists are struggling with the implications of the ‘Nagoya Protocol on Access to Genetic Resources and the Fair and Equitable Sharing of Benefits Arising from their utilisation under the Convention on Biological Diversity’ and the interpretation by governments that restricts and, in many cases, prohibits the exchange of strains between mycologists from different countries. The ICNafp (Rec. 8B.1) recommends that strains of new species should be deposited in two recognised public culture or genetic resource collections, preferably in different countries. It is not inconceivable that strain sharing will become even more complicated in the future, making taxonomic revisions more difficult. This is a complex issue, but we recommend looking for ways to enable sharing of strains. Nevertheless, we believe it is critical that new species are described with as much data as possible, not only to prove conclusively that the proposed species are indeed new, but also to avoid a future in which species might be classified as doubtful because no material is available for study or a lack of data on the species renders it unrecognisable, as was the case with the many doubtful species described in the late 1800s, what Pitt (1980) called the ‘dark ages’.

The introduction of species with a single isolate is not ideal, but it is also not considered a problem, provided it is clearly unique based on all available data, which at least includes BenA, CaM and RPB2. Of course, there is always the risk that a species is not new when future data are collected. Currently, 467 of the 1 279 recognised Eurotiales species are known from only the ex-type strain and 219 species for which one additional reference sequence is available in GenBank. For Aspergillus, 143 of 453 accepted species are represented by the ex-type strain only, while an additional strain was sequenced for 93; for Penicillium, 175 of 535 accepted species are represented by the ex-type strain only, while an additional strain was sequenced for 75; and for Talaromyces, 88 of 203 accepted species are represented by the ex-type strain only, while an additional strain was sequenced for 34. Thus, for a considerable proportion of accepted species, we have little knowledge about infraspecific variability and thus also species boundaries, and much remains to be done to capture them. We therefore recommend that when isolating these poorly sequenced species, BenA, CaM and RPB2 sequences be generated and the strains deposited in recognised culture collections or, if this is not possible, the strains be shared with taxonomists working on these genera. Because of the uncertainty that a lack of DNA reference data can cause, we also recommend that workers who are uncertain about the identity of a species ask a taxonomist specialising in these fungi for a second opinion or advice.

Throughout the history of these genera, the way we describe species has evolved, bringing with it many taxonomic changes. The one thing that has remained is the community of collaborators who have adhered to best practises and worked together to create a standard for Eurotiales that serves everyone and provides a solid taxonomic basis for the study of these species. We believe that our recommendations will help to expand this community and provide further stability to the taxonomy of Eurotiales.

Fig. 2.

Phylogenetic tree of Aspergillus section Candidi based on a concatenated dataset of BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. The tree was rooted to A. neotritici. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by superscript T. See Suppl. Fig. S2.

Fig. 4.

Phylogenetic tree of Aspergillus section Circumdati series Sclerotiorum based on a concatenated dataset of BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. The tree was rooted to A. ochraceus. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by superscript T. See Suppl. Fig. S4.

Fig. 5.

Phylogenetic tree of Aspergillus section Cremei series Wentiorum based on a concatenated dataset of BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. The tree was rooted to A. cremeus. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by superscript T. See Suppl. Fig. S5.

Fig. 7.

Phylogenetic tree of Aspergillus sections Flavipedes and Janorum based on a concatenated dataset of BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. The tree was rooted to A. terreus. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by superscript T. See Suppl. Fig. S7.

Fig. 9.

Phylogenetic tree of Aspergillus section Nidulantes series Versicolores based on a concatenated dataset of BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. The tree was rooted to A. aeneus. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by superscript T. See Suppl. Fig. S9.

Fig. 11.

Phylogenetic tree of Aspergillus section Polypaecilum based on a concatenated dataset of BenA, CaM, RPB2, and ITS. Strains of recently described species are shown in bold coloured text. The tree was rooted to A. cremeus. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by superscript T. See Suppl. Fig. S11.

Fig. 13.

Phylogenetic tree of Aspergillus section Usti series Calidousti based on a concatenated dataset of BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. The tree was rooted to A. ustus. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by superscript T. See Suppl. Fig. S13.

Fig. 15.

Phylogenetic tree of Paecilomyces based on a concatenated dataset of BenA, CaM, RPB2 and ITS. Strains of recently described species are shown in bold coloured text. The tree was rooted to Thermoascus crustaceus. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by superscript T. See Suppl. Fig. S14.

Fig. 20.

Phylogenetic tree of Penicillium section Canescentia based on a concatenated dataset of BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. The tree was midpoint rooted. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by superscript T. See Suppl. Fig. S18.

Fig. 22.

Phylogenetic tree of Penicillium sections Crypta and Torulomyces based on a concatenated dataset of BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. The tree was rooted to P. alfredii. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by superscript T. See Suppl. Fig. S20.

Fig. 26.

Phylogenetic tree of Penicillium sections Paradoxa, Ramosum and Robsamsonia, based on a concatenated dataset of BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. The tree was midpoint rooted. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by superscript T. See Suppl. Fig. S24.

Fig. 29.

Phylogenetic tree of Talaromyces section Helici, based on a concatenated dataset of BenA, CaM, RPB2, and ITS. Strains of recently described species are shown in bold coloured text. The tree was rooted to T. purpureogenus. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by superscript T. See Suppl. Fig. S27.

Fig. 30.

Phylogenetic tree of Talaromyces sections Purpurei and Subinflati, based on a concatenated dataset of BenA, CaM, RPB2, and ITS. Strains of recently described species are shown in bold coloured text. The tree was midpoint rooted. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by superscript T. See Suppl. Fig. S28.

Fig. 32.

Phylogenetic tree of Talaromyces showing the phylogenetic relationship of the recently introduced section Tenues, based on a concatenated dataset of BenA, CaM, RPB2, and ITS. Strains of recently described species are shown in bold coloured text. The tree was rooted to Trichocoma paradoxa. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by superscript T. Representative strains are indicated by superscript R. See Suppl. Fig. S30.

Supplementary Material: https://studiesinmycology.org/

Fig. S1

Phylogenetic trees of Aspergillus section Aenei series Aenei based on BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. Trees were rooted to A. versicolor. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by T.

Fig. S2

Phylogenetic trees of Aspergillus section Candidi series Candidi based on BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. Trees were rooted to A. neotritici. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by T.

Fig. S3

Phylogenetic trees of Aspergillus sections Cavernicolarum, Ochraceorosei and Sparsi based on BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. Trees were rooted to A. bisporus. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by T.

Fig. S4

Phylogenetic trees of Aspergillus section Circumdati series Sclerotiorum based on BenA, CaM, and RPB2, and series Steynii based on ITS. Strains of recently described species are shown in bold coloured text. Trees were rooted to A. ochraceus. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by T.

Fig. S5

Phylogenetic trees of Aspergillus section Cremei series Wentiorum based on BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. Trees were rooted to A. cremeus. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by T.

Fig. S6

Phylogenetic trees of Aspergillus section Flavi series Alliacei and Flavi based on BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. The tree was rooted to A. avenaceus. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by T. Representative strains are indicated by R.

Fig. S7

Phylogenetic trees of Aspergillus section Flavipedes based on BenA, CaM, RPB2 and ITS. Strains of recently described species are shown in bold coloured text. Trees were rooted to A. terreus. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by T.

Fig. S8

Phylogenetic trees of Aspergillus section Fumigati based on BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. Trees were rooted to A. clavatus. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by T.

Fig. S9

Phylogenetic trees of Aspergillus section Nidulantes series Versicolores based on BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. Trees were rooted to A. aeneus. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by T.

Fig. S10

Phylogenetic trees of Aspergillus section Nigri series Japonici and Nigri based on BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. Trees were rooted to A. candidus. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by T.

Fig. S11

Phylogenetic trees of Aspergillus section Polypaecilum based on BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. Trees were rooted to A. cremeus. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by T.

Fig. S12

Phylogenetic trees of Aspergillus section Terrei series Nivei and Terrei based on BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. Trees were rooted to A. ambiguus. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by T.

Fig. S13

Phylogenetic trees of Aspergillus section Usti series Calidousti based on BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. Trees were rooted to A. ustus. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by T.

Fig. S14

Phylogenetic trees of Paecilomyces based on BenA, CaM and RPB2. Strains of recently described species are shown in bold coloured text. Trees were rooted to Thermoascus crustaceus. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by T.

Fig. S15

Phylogenetic trees of Penicillium sections Aspergilloides, Charlesia, Cinnamopurpurea and Ramigena based on BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. Trees were rooted to P. taxi. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by T.

Fig. S16

Phylogenetic trees of Penicillium section Brevicompacta based on BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. Trees were rooted to P. tularense (except RPB2 that was midpoint rooted). UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by T.

Fig. S17

Phylogenetic trees of Penicillium section Fasciculata series Camembertiorum based on BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. Trees were rooted to P. expansum. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by T.

Fig. S18

Phylogenetic trees of Penicillium section Canescentia based on BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. The tree was midpoint rooted. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by T.

Fig. S19

Phylogenetic trees of Penicillium section Citrina based on BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. Trees were rooted to P. euglaucum. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by T.

Fig. S20

Phylogenetic trees of Penicillium sections Crypta and Torulomyces based on BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. Trees were rooted to P. alfredii. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by T.

Fig. S21

Phylogenetic trees of Penicillium sections Exilicaulis and Gracilenta based on BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. Trees were rooted to P. stolkiae. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by T.

Fig. S22

Phylogenetic trees of Penicillium section Lanata-Divaricata series Janthinella and Simplicissima, based on BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. Trees were rooted to P. stolkiae. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by T.

Fig. S23

Phylogenetic trees of Penicillium section Lanata-Divaricata series Dalearum, Oxalica and Rolfsiorum, based on BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. Trees were rooted to the series Oxalica clade. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by T.

Fig. S24

Phylogenetic trees of Penicillium sections Paradoxa, Ramosum and Robsamsonia, based on BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. The tree was midpoint rooted. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by T.

Fig. S25

Phylogenetic trees of Penicillium section Sclerotiorum, based on BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. Trees were rooted to P. glabrum. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by T.

Fig. S26

Phylogenetic trees of Rasamsonia, based on BenA, CaM and ITS. Strains of recently described species are shown in bold coloured text. The tree was rooted to Trichocoma paradoxa. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by superscript T. Representative strains are indicated by R.

Fig. S27

Phylogenetic trees of Talaromyces section Helici, based on BenA, CaM and RPB2. Strains of recently described species are shown in bold coloured text. Trees were rooted to T. purpureogenus. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by T.

Fig. S28

Phylogenetic trees of Talaromyces section Purpurei and Subinflati, based on BenA, CaM, and RPB2. Strains of recently described species are shown in bold coloured text. Trees were midpoint rooted. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by T.

Fig. S29

Phylogenetic trees of Talaromyces section Talaromyces, based on BenA, CaM and RPB2. Strains of recently described species are shown in bold coloured text. Trees were rooted to T. helicus. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by T.

Fig. S30

Phylogenetic trees of Talaromyces section Trachyspermi, based on BenA, CaM and RPB2. Strains of recently described species are shown in bold coloured text. Trees were rooted to T. purpureus. UltraFast Bootstrap support values higher than 95 % are shown at relevant branches. Ex-type strains are indicated by T.

Table S1

List of reference species used for comparisons in this study.