Although Aspergillus fumigatus is the major agent of invasive aspergillosis, an increasing number of infections are caused by its cryptic species, especially A. lentulus and the A. viridinutans species complex (AVSC). Their identification is clinically relevant because of antifungal drug resistance and refractory infections. Species boundaries in the AVSC are unresolved since most species have uniform morphology and produce interspecific hybrids in vitro. Clinical and environmental strains from six continents (n = 110) were characterized by DNA sequencing of four to six loci. Biological compatibilities were tested within and between major phylogenetic clades, and ascospore morphology was characterised. Species delimitation methods based on the multispecies coalescent model (MSC) supported recognition of ten species including one new species. Four species are confirmed opportunistic pathogens; A. udagawae followed by A. felis and A. pseudoviridinutans are known from opportunistic human infections, while A. felis followed by A. udagawae and A. wyomingensis are agents of feline sino-orbital aspergillosis. Recently described human-pathogenic species A. parafelis and A. pseudofelis are synonymized with A. felis and an epitype is designated for A. udagawae. Intraspecific mating assay showed that only a few of the heterothallic species can readily generate sexual morphs in vitro. Interspecific mating assays revealed that five different species combinations were biologically compatible. Hybrid ascospores had atypical surface ornamentation and significantly different dimensions compared to parental species. This suggests that species limits in the AVSC are maintained by both pre- and post-zygotic barriers and these species display a great potential for rapid adaptation and modulation of virulence. This study highlights that a sufficient number of strains representing genetic diversity within a species is essential for meaningful species boundaries delimitation in cryptic species complexes. MSC-based delimitation methods are robust and suitable tools for evaluation of boundaries between these species.

1st Faculty of Medicine Charles University Kateřinská 32 121 08 Prague 2 Czech Republic

Department of Biotechnology and Biomedicine Technical University of Denmark Kongens Lyngby Denmark

Department of Botany Faculty of Science Charles University Benátská 2 128 01 Prague 2 Czech Republic

Laboratory of Fungal Genetics and Metabolism Institute of Microbiology of the CAS v v i Vídeňská 1083 142 20 Prague 4 Czech Republic

Medical Mycology Research Center Chiba University 1 8 1 Inohana Chuo ku Chiba 260 8673 Japan

Sydney School of Veterinary Science Faculty of Science and Marie Bashir Institute of Infectious Diseases and Biosecurity University of Sydney Camperdown NSW Australia

University of Nagasaki 1 1 1 Manabino Nagayo cho Nishi Sonogi gun Nagasaki 851 2195 Japan

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