The genus Codinaea is a phialidic, dematiaceous hyphomycete known for its intriguing morphology and turbulent taxonomic history. This polyphasic study represents a new, comprehensive view on the taxonomy, systematics, and biogeography of Codinaea and its relatives. Phylogenetic analyses of three nuclear loci confirmed that Codinaea is polyphyletic. The generic concept was emended; it includes four morphotypes that contribute to its morphological complexity. Ancestral inference showed that the evolution of some traits is correlated and that these traits previously used to delimit taxa at the generic level occur in species that were shown to be congeneric. Five lineages of Codinaea-like fungi were recognized and introduced as new genera: Codinaeella, Nimesporella, Stilbochaeta, Tainosphaeriella, and Xyladelphia. Dual DNA barcoding facilitated identification at the species level. Codinaea and its segregates thrive on decaying plants, rarely occurring as endophytes or plant pathogens. Environmental ITS sequences indicate that they are common in bulk soil. The geographic distribution found using GlobalFungi database was consistent with known data. Most species are distributed in either the Holarctic realm or tropical geographic regions. The ancestral climatic zone was temperate, followed by transitions to the tropics; these fungi evolved primarily in Eurasia and Americas, with subsequent transitions to Africa and Australasia.

CEITEC Central European Institute of Technology Masaryk University 625 00 Brno Czech Republic

Department of Clinical Biochemistry and Diagnostics University Hospital Hradec Králové 500 05 Hradec Králové Czech Republic

Department of Taxonomy Institute of Botany The Czech Academy of Sciences 252 43 Průhonice Czech Republic

Illinois Natural History Survey University of Illinois Urbana Champaign Champaign IL 61820 USA

Laboratory of Fungal Genetics and Metabolism Institute of Microbiology The Czech Academy of Sciences 142 20 Prague Czech Republic

Westerdijk Fungal Biodiversity Institute 3508 AD Utrecht The Netherlands

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