Aspergillus section Aspergillus (formerly the genus Eurotium) includes xerophilic species with uniseriate conidiophores, globose to subglobose vesicles, green conidia and yellow, thin walled eurotium-like ascomata with hyaline, lenticular ascospores. In the present study, a polyphasic approach using morphological characters, extrolites, physiological characters and phylogeny was applied to investigate the taxonomy of this section. Over 500 strains from various culture collections and new isolates obtained from indoor environments and a wide range of substrates all over the world were identified using calmodulin gene sequencing. Of these, 163 isolates were subjected to molecular phylogenetic analyses using sequences of ITS rDNA, partial β-tubulin (BenA), calmodulin (CaM) and RNA polymerase II second largest subunit (RPB2) genes. Colony characteristics were documented on eight cultivation media, growth parameters at three incubation temperatures were recorded and micromorphology was examined using light microscopy as well as scanning electron microscopy to illustrate and characterize each species. Many specific extrolites were extracted and identified from cultures, including echinulins, epiheveadrides, auroglaucins and anthraquinone bisanthrons, and to be consistent in strains of nearly all species. Other extrolites are species-specific, and thus valuable for identification. Several extrolites show antioxidant effects, which may be nutritionally beneficial in food and beverages. Important mycotoxins in the strict sense, such as sterigmatocystin, aflatoxins, ochratoxins, citrinin were not detected despite previous reports on their production in this section. Adopting a polyphasic approach, 31 species are recognized, including nine new species. ITS is highly conserved in this section and does not distinguish species. All species can be differentiated using CaM or RPB2 sequences. For BenA, Aspergillus brunneus and A. niveoglaucus share identical sequences. Ascospores and conidia morphology, growth rates at different temperatures are most useful characters for phenotypic species identification.
Combined alignment: benA caM RPB2
Combined dataset of BenA, CaM and RPB2 sequences for 164 taxa belonging to section Aspergillus (formerly Eurotium). Hamigera avellanea NRRL 1938T represents an outgroup. BenA: positions 1-408; CaM: positions 409-1124; RPB2: positions 1125-2093
benA caM RPB2.fas
Phylogeny of sect. Aspergillus inferred from combined data: BenA, CaM, RPB2
A 50% majority rule Maximum likelihood consensus tree based on combined dataset of BenA, CaM and RPB2 sequences showing the relationship of species within Aspergillus sect. Aspergillus. Dataset contained 164 taxa. Maximum likelihood bootstrap proportion (bs) and Bayesian posterior probability (pp) are appended to nodes; only bs ≥ 70 % and pp ≥ 95% are shown, lower supports are indicated with a hyphen, whereas asterisks indicate full support (100% bs or 1.00 pp); ex-type strains are designated by a superscript T. The tree is rooted with Hamigera avellanea NRRL 1938T.
Fig 1 Eurotium combined phylogeny.pdf
β-tubulin (BenA) alignment
Alignment: β-tubulin (BenA) sequences for 164 taxa belonging to section Aspergillus (formerly Eurotium). Hamigera avellanea NRRL 1938T represents an outgroup.
benA.fas
Aspergillus sect. Aspergillus: Tree based on benA
A 50% majority rule Maximum likelihood consensus tree based on partial β-tubulin (BenA) sequences showing the relationship of species within Aspergillus sect. Aspergillus. Maximum likelihood bootstrap proportion (bs) and Bayesian posterior probability (pp) are appended to nodes; only bs ≥ 70 % and pp ≥ 95% are shown, lower supports are indicated with a hyphen, whereas asterisks indicate full support (100% bs or 1.00 pp); ex-type strains are designated by a superscript T. The tree is rooted with Hamigera avellanea NRRL 1938T.
Fig 2 Eurotium Tree benA.pdf
Calmodulin (caM) alignment
Alignment: calmodulin (caM) sequences for 164 taxa belonging to section Aspergillus (formerly Eurotium). Hamigera avellanea NRRL 1938T represents an outgroup.
caM.fas
Aspergillus sect. Aspergillus: Tree based on CaM
A 50% majority rule Maximum likelihood consensus tree based on partial calmodulin (CaM) sequences showing the relationship of species within Aspergillus sect. Aspergillus. Maximum likelihood bootstrap proportion (bs) and Bayesian posterior probability (pp) are appended to nodes; only bs ≥ 70 % and pp ≥ 95% are shown, lower supports are indicated with a hyphen, whereas asterisks indicate full support (100% bs or 1.00 pp); ex-type strains are designated by a superscript T. The tree is rooted with Hamigera avellanea NRRL 1938T.
Fig 3 Eurotium Tree CaM.pdf
RPB2 alignment
Alignment: RNA polymerase II second largest subunit (RPB2) sequences for 164 taxa belonging to section Aspergillus (formerly Eurotium). Hamigera avellanea NRRL 1938T represents an outgroup.
RPB2.fasta
Aspergillus sect. Aspergillus: Tree based on RPB2
A 50% majority rule Maximum likelihood consensus tree based on partial RNA polymerase II second largest subunit (RPB2) sequences showing the relationship of species within Aspergillus sect. Aspergillus. Maximum likelihood bootstrap proportion (bs) and Bayesian posterior probability (pp) are appended to nodes; only bs ≥ 70 % and pp ≥ 95% are shown, lower supports are indicated with a hyphen, whereas asterisks indicate full support (100% bs or 1.00 pp); ex-type strains are designated by a superscript T. The tree is rooted with Hamigera avellanea NRRL 1938T.
Fig 4 Eurotium Tree RPB2.pdf
ITS rDNA alignmet
Alignment: RNA polymerase II second largest subunit (RPB2) sequences for 164 taxa belonging to section Aspergillus (formerly Eurotium). Hamigera avellanea NRRL 1938T represents an outgroup.
ITS.fas
Aspergillus sect. Aspergillus: Tree based on ITS rDNA
A 50% majority rule Maximum likelihood consensus tree based on ITS rDNA sequences. Maximum likelihood bootstrap proportion (bs) and Bayesian posterior probability (pp) are appended to nodes; only bs ≥ 70 % and pp ≥ 95% are shown, lower supports are indicated with a hyphen, whereas asterisks indicate full support (100% bs or 1.00 pp); ex-type strains are designated by a superscript T. The tree is rooted with Hamigera avellanea NRRL 1938T.
Fig 5 Eurotium Tree ITS.pdf