Trichoderma: systematics, the sexual state, and ecology.

ABSTRACT A chronology is presented that charts the development of a genus and species concept in Trichoderma. Eighty-nine species of Trichoderma have been named, and several species of Hypocrea have been linked to unnamed Trichoderma anamorphs. Eighty-three taxa of Trichoderma and their teleomorphs, Hypocrea spp., have been included in phylogenetic analyses, including 11 species of Hypocrea with unnamed Trichoderma anamorphs. Phylogenetic analyses show that Trichoderma and Hypocrea are congeneric. Trichoderma species not linked to Hypocrea teleomorphs are derived from among species that are linked to teleomorphs, indicating sexual and asexual lineages are not independent of each other. Many more species remain to be discovered and described. Molecular phylogenetic analyses have revealed the existence of more species than have been recognized on the basis of morphology alone. A suggestion is made to modify the International Code of Botanical Nomenclature to enable adoption of a single generic name for Trichoderma/Hypocrea, with Trichoderma being the older and more utilitarian name. As increasing numbers of species are studied, the few morphological characters of anamorph and teleomorph have reached their limit for defining species. DNA-based characters have assumed an indispensable role. Exploration of new niches, such as within tree trunks and new geographic locations, have resulted in a substantial increase in the number of species of Trichoderma. Trichoderma is usually considered a genus of free-living soil fungi but evidence suggests that Trichoderma species may be opportunistic, avirulent plant symbionts as well as parasites of other fungi. Members of the genus Trichoderma are universally present in soils, although individual species may be either cosmopolitan (e.g., T. harzianum) or limited (e.g., T. viride) in their geographic distribution. To facilitate identification of species, a list of correctly identified strains of Trichoderma and their GenBank numbers for sequences of translation-elongation factor EF-1alpha and internal transcribed spacer rDNA is provided.