Both flagellated and nonflagellated fungi have an extensive fossil record, which is, however, unevenly documented and often difficult to interpret. Recent work on Silurian, Devonian and Carboniferous terrestrial assemblages has provided plausible evidence for all major groups of extant fungi in the Paleozoic. Key events in fungal macroevolution thus probably took place in the early Paleozoic or the late Precambrian, and the likelihood of finding definitive fossil evidence for them is small. The fossil record also provides evidence for morphological conservatism and early establishment of a spectrum of intimate associations between fungi and vascular plants. A model for the origin of terrestrial fungi involving two distinct lines of biotrophs in lichen-like symbioses with algae is proposed.