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

Abstract

The Eurotiales is a relatively large order of Ascomycetes with members frequently having positive and negative impact on human activities. Species within this order gain attention from various research fields such as food, indoor and medical mycology and biotechnology. In this article we give an overview of families and genera present in the Eurotiales and introduce an updated subgeneric, sectional and series classification for Aspergillus and Penicillium. Finally, a comprehensive list of accepted species in the Eurotiales is given. The classification of the Eurotiales at family and genus level is traditionally based on phenotypic characters, and this classification has since been challenged using sequence-based approaches. Here, we re-evaluated the relationships between families and genera of the Eurotiales using a nine-gene sequence dataset. Based on this analysis, the new family Penicillaginaceae is introduced and four known families are accepted: Aspergillaceae, Elaphomycetaceae, Thermoascaceae and Trichocomaceae. The Eurotiales includes 28 genera: 15 genera are accommodated in the Aspergillaceae (Aspergillago, Aspergillus, Evansstolkia, Hamigera, Leiothecium, Monascus, Penicilliopsis, Penicillium, Phialomyces, Pseudohamigera, Pseudopenicillium, Sclerocleista, Warcupiella, Xerochrysium and Xeromyces), eight in the Trichocomaceae (Acidotalaromyces, Ascospirella, Dendrosphaera, Rasamsonia, Sagenomella, Talaromyces, Thermomyces, Trichocoma), two in the Thermoascaceae (Paecilomyces, Thermoascus) and one in the Penicillaginaceae (Penicillago). The classification of the Elaphomycetaceae was not part of this study, but according to literature two genera are present in this family (Elaphomyces and Pseudotulostoma). The use of an infrageneric classification system has a long tradition in Aspergillus and Penicillium. Most recent taxonomic studies focused on the sectional level, resulting in a well-established sectional classification in these genera. In contrast, a series classification in Aspergillus and Penicillium is often outdated or lacking, but is still relevant, e.g., the allocation of a species to a series can be highly predictive in what functional characters the species might have and might be useful when using a phenotype-based identification. The majority of the series in Aspergillus and Penicillium are invalidly described and here we introduce a new series classification. Using a phylogenetic approach, often supported by phenotypic, physiologic and/or extrolite data, Aspergillus is subdivided in six subgenera, 27 sections (five new) and 75 series (73 new, one new combination), and Penicillium in two subgenera, 32 sections (seven new) and 89 series (57 new, six new combinations). Correct identification of species belonging to the Eurotiales is difficult, but crucial, as the species name is the linking pin to information. Lists of accepted species are a helpful aid for researchers to obtain a correct identification using the current taxonomic schemes. In the most recent list from 2014, 339 Aspergillus, 354 Penicillium and 88 Talaromyces species were accepted. These numbers increased significantly, and the current list includes 446 Aspergillus (32 % increase), 483 Penicillium (36 % increase) and 171 Talaromyces (94 % increase) species, showing the large diversity and high interest in these genera. We expanded this list with all genera and species belonging to the Eurotiales (except those belonging to Elaphomycetaceae). The list includes 1 187 species, distributed over 27 genera, and contains MycoBank numbers, collection numbers of type and ex-type cultures, subgenus, section and series classification data, information on the mode of reproduction, and GenBank accession numbers of ITS, beta-tubulin (BenA), calmodulin (CaM) and RNA polymerase II second largest subunit (RPB2) gene sequences.

Keywords: Acidotalaromyces Houbraken, Frisvad & Samson; Acidotalaromyces lignorum (Stolk) Houbraken, Frisvad & Samson; Ascospirella Houbraken, Frisvad & Samson; Ascospirella lutea (Zukal) Houbraken, Frisvad & Samson; Aspergillus chaetosartoryae Hubka, Kocsubé & Houbraken; Classification; Evansstolkia Houbraken, Frisvad & Samson; Evansstolkia leycettana (H.C. Evans & Stolk) Houbraken, Frisvad & Samson; Hamigera brevicompacta (H.Z. Kong) Houbraken, Frisvad & Samson; Infrageneric classification; New combinations, series; New combinations, species; New genera; New names; New sections; New series; New taxa; Nomenclature; Paecilomyces lagunculariae (C. Ram) Houbraken, Frisvad & Samson; Penicillaginaceae Houbraken, Frisvad & Samson; Penicillago kabunica (Baghd.) Houbraken, Frisvad & Samson; Penicillago mirabilis (Beliakova & Milko) Houbraken, Frisvad & Samson; Penicillago moldavica (Milko & Beliakova) Houbraken, Frisvad & Samson; Phialomyces arenicola (Chalab.) Houbraken, Frisvad & Samson; Phialomyces humicoloides (Bills & Heredia) Houbraken, Frisvad & Samson; Phylogeny; Polythetic classes; Pseudohamigera Houbraken, Frisvad & Samson; Pseudohamigera striata (Raper & Fennell) Houbraken, Frisvad & Samson; Talaromyces resinae (Z.T. Qi & H.Z. Kong) Houbraken & X.C. Wang; Talaromyces striatoconidius Houbraken, Frisvad & Samson; Taxonomic novelties: New family; Thermoascus verrucosus (Samson & Tansey) Houbraken, Frisvad & Samson; Thermoascus yaguchii Houbraken, Frisvad & Samson; in Aspergillus: sect. Bispori S.W. Peterson, Varga, Frisvad, Samson ex Houbraken; in Aspergillus: ser. Acidohumorum Houbraken & Frisvad; in Aspergillus: ser. Inflati (Stolk & Samson) Houbraken & Frisvad; in Penicillium: sect. Alfrediorum Houbraken & Frisvad; in Penicillium: ser. Adametziorum Houbraken & Frisvad; in Penicillium: ser. Alutacea (Pitt) Houbraken & Frisvad; sect. Crypta Houbraken & Frisvad; sect. Eremophila Houbraken & Frisvad; sect. Formosana Houbraken & Frisvad; sect. Griseola Houbraken & Frisvad; sect. Inusitata Houbraken & Frisvad; sect. Lasseniorum Houbraken & Frisvad; sect. Polypaecilum Houbraken & Frisvad; sect. Raperorum S.W. Peterson, Varga, Frisvad, Samson ex Houbraken; sect. Silvatici S.W. Peterson, Varga, Frisvad, Samson ex Houbraken; sect. Vargarum Houbraken & Frisvad; ser. Alliacei Houbraken & Frisvad; ser. Ambigui Houbraken & Frisvad; ser. Angustiporcata Houbraken & Frisvad; ser. Arxiorum Houbraken & Frisvad; ser. Atramentosa Houbraken & Frisvad; ser. Aurantiobrunnei Houbraken & Frisvad; ser. Avenacei Houbraken & Frisvad; ser. Bertholletiarum Houbraken & Frisvad; ser. Biplani Houbraken & Frisvad; ser. Brevicompacta Houbraken & Frisvad; ser. Brevipedes Houbraken & Frisvad; ser. Brunneouniseriati Houbraken & Frisvad; ser. Buchwaldiorum Houbraken & Frisvad; ser. Calidousti Houbraken & Frisvad; ser. Canini Houbraken & Frisvad; ser. Carbonarii Houbraken & Frisvad; ser. Cavernicolarum Houbraken & Frisvad; ser. Cervini Houbraken & Frisvad; ser. Chevalierorum Houbraken & Frisvad; ser. Cinnamopurpurea Houbraken & Frisvad; ser. Circumdati Houbraken & Frisvad; ser. Clavigera Houbraken & Frisvad; ser. Conjuncti Houbraken & Frisvad; ser. Copticolarum Houbraken & Frisvad; ser. Coremiiformes Houbraken & Frisvad; ser. Corylophila Houbraken & Frisvad; ser. Costaricensia Houbraken & Frisvad; ser. Cremei Houbraken & Frisvad; ser. Crustacea (Pitt) Houbraken & Frisvad; ser. Dalearum Houbraken & Frisvad; ser. Deflecti Houbraken & Frisvad; ser. Egyptiaci Houbraken & Frisvad; ser. Erubescentia (Pitt) Houbraken & Frisvad; ser. Estinogena Houbraken & Frisvad; ser. Euglauca Houbraken & Frisvad; ser. Fennelliarum Houbraken & Frisvad; ser. Flavi Houbraken & Frisvad; ser. Flavipedes Houbraken & Frisvad; ser. Fortuita Houbraken & Frisvad; ser. Fumigati Houbraken & Frisvad; ser. Funiculosi Houbraken & Frisvad; ser. Gallaica Houbraken & Frisvad; ser. Georgiensia Houbraken & Frisvad; ser. Goetziorum Houbraken & Frisvad; ser. Gracilenta Houbraken & Frisvad; ser. Halophilici Houbraken & Frisvad; ser. Herqueorum Houbraken & Frisvad; ser. Heteromorphi Houbraken & Frisvad; ser. Hoeksiorum Houbraken & Frisvad; ser. Homomorphi Houbraken & Frisvad; ser. Idahoensia Houbraken & Frisvad; ser. Implicati Houbraken & Frisvad; ser. Improvisa Houbraken & Frisvad; ser. Indica Houbraken & Frisvad; ser. Japonici Houbraken & Frisvad; ser. Jiangxiensia Houbraken & Frisvad; ser. Kalimarum Houbraken & Frisvad; ser. Kiamaensia Houbraken & Frisvad; ser. Kitamyces Houbraken & Frisvad; ser. Lapidosa (Pitt) Houbraken & Frisvad; ser. Leporum Houbraken & Frisvad; ser. Leucocarpi Houbraken & Frisvad; ser. Livida Houbraken & Frisvad; ser. Longicatenata Houbraken & Frisvad; ser. Macrosclerotiorum Houbraken & Frisvad; ser. Monodiorum Houbraken & Frisvad; ser. Multicolores Houbraken & Frisvad; ser. Neoglabri Houbraken & Frisvad; ser. Neonivei Houbraken & Frisvad; ser. Nidulantes Houbraken & Frisvad; ser. Nigri Houbraken & Frisvad; ser. Nivei Houbraken & Frisvad; ser. Nodula Houbraken & Frisvad; ser. Nomiarum Houbraken & Frisvad; ser. Noonimiarum Houbraken & Frisvad; ser. Ochraceorosei Houbraken & Frisvad; ser. Olivimuriarum Houbraken & Frisvad; ser. Osmophila Houbraken & Frisvad; ser. Paradoxa Houbraken & Frisvad; ser. Paxillorum Houbraken & Frisvad; ser. Penicillioides Houbraken & Frisvad; ser. Phoenicea Houbraken & Frisvad; ser. Pinetorum (Pitt) Houbraken & Frisvad; ser. Polypaecilum Houbraken & Frisvad; ser. Pulvini Houbraken & Frisvad; ser. Quercetorum Houbraken & Frisvad; ser. Raistrickiorum Houbraken & Frisvad; ser. Ramigena Houbraken & Frisvad; ser. Restricti Houbraken & Frisvad; ser. Robsamsonia Houbraken & Frisvad; ser. Rolfsiorum Houbraken & Frisvad; ser. Roseopurpurea Houbraken & Frisvad; ser. Rubri Houbraken & Frisvad; ser. Salinarum Houbraken & Frisvad; ser. Samsoniorum Houbraken & Frisvad; ser. Saturniformia Houbraken & Frisvad; ser. Scabrosa Houbraken & Frisvad; ser. Sclerotigena Houbraken & Frisvad; ser. Sclerotiorum Houbraken & Frisvad; ser. Sheariorum Houbraken & Frisvad; ser. Simplicissima Houbraken & Frisvad; ser. Soppiorum Houbraken & Frisvad; ser. Sparsi Houbraken & Frisvad; ser. Spathulati Houbraken & Frisvad; ser. Spelaei Houbraken & Frisvad; ser. Speluncei Houbraken & Frisvad; ser. Spinulosa Houbraken & Frisvad; ser. Stellati Houbraken & Frisvad; ser. Steyniorum Houbraken & Frisvad; ser. Sublectatica Houbraken & Frisvad; ser. Sumatraensia Houbraken & Frisvad; ser. Tamarindosolorum Houbraken & Frisvad; ser. Teporium Houbraken & Frisvad; ser. Terrei Houbraken & Frisvad; ser. Thermomutati Houbraken & Frisvad; ser. Thiersiorum Houbraken & Frisvad; ser. Thomiorum Houbraken & Frisvad; ser. Unguium Houbraken & Frisvad; ser. Unilaterales Houbraken & Frisvad; ser. Usti Houbraken & Frisvad; ser. Verhageniorum Houbraken & Frisvad; ser. Versicolores Houbraken & Frisvad; ser. Virgata Houbraken & Frisvad; ser. Viridinutantes Houbraken & Frisvad; ser. Vitricolarum Houbraken & Frisvad; ser. Wentiorum Houbraken & Frisvad; ser. Westlingiorum Houbraken & Frisvad; ser. Whitfieldiorum Houbraken & Frisvad; ser. Xerophili Houbraken & Frisvad; series Tularensia (Pitt) Houbraken & Frisvad.

Fig. 1

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

Fig. 2

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

Fig. 2

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

Fig. 3

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

Fig. 4

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

Fig. 5

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

Fig. 6

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

Fig. 7

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

Fig. 8

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

Fig. 9

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

Fig. 10

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

Fig. 11

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

Fig. 12

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

Fig. 12

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

Fig. 13

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

Fig. 14

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

Fig. 15

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

Fig. 15

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

Fig. 16

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

Fig. 16

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

Fig. 16

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

Fig. 17

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

Fig. 18

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

Fig. 18

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

Fig. 18

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

Fig. 18

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

Fig. 18

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

Fig. 18

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

Fig. 19

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

Fig. 19

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

Fig. 19

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

Fig. 20

Phylogenetic trees based on single and a combined data set of BenA, CaM and niaD sequences showing the relationship between Aspergillus texensis and related species. The BI posterior probability (pp) values and bootstrap percentages of the maximum likelihood (ML) analysis are presented at the nodes; fully supported branches are thickened. Values less than 70 % bootstrap support (ML) or less than 0.95 posterior probability (Bayesian analysis) are indicated with a hyphen or not shown. The bar indicates the number of substitutions per site. The phylogram is rooted with Aspergillus tamarii NRRL 20818.