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SUMMARY Lichen‐forming fungi are a large, taxonomically diverse group of nutritional specialists which acquire fixed carbon from a population of minute green algal or cyanobacterial cells. Mycobionts of foliose or fruticose lichens differ from the rest of the fungi by expressing morphologically and anatomically complex symbiotic phenotypes. The extracellularly located photobiont cell population of these macrolichens is housed and controlled by the quantitatively predominant fungal partner which competes for space above ground, secures adequate illumination and facilitates gas exchange. This review summarizes data on the ontogeny, functional morphology, growth patterns and internal thalline differentiation of macrolichens.

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Summary Lichenization is assumed to be a very ancient mode of fungal nutrition, but fossil records are rare. Here we describe two fragments of exceptionally preserved, probably charred, lichen thalli with internal stratification. Cyanolichenomycites devonicus has a cyanobacterial and Chlorolichenomycites salopensis a unicellular, presumably green algal photobiont. Fruiting bodies are missing. Cyanolichenomycites devonicus forms asexual spores in a pycnidium. All specimens were examined with scanning electron microscopy techniques. The fossils were extracted by maceration. Extant lichens and free‐living cyanobacteria were either experimentally charcoalified for comparison or conventionally prepared. Based on their septate hyphal structure, both specimens are tentatively interpreted as representatives of the Pezizomycotina (Ascomycota). Their presence in 415 million yr (Myr) old rocks from the Welsh Borderland predates existing Late Cretaceous records of pycnidial conidiomata by some 325 Myr and Triassic records of lichens with broadly similar organization by some 195 Myr. These fossils represent the oldest known record of lichens with symbionts and anatomy as typically found in morphologically advanced taxa today. The latter does not apply to Winfrenatia reticulata, the enigmatic crustose lichen fossil from the Lower Devonian, nor to presumed lichen‐like organisms such as the Cambrian Farghera robusta or to the Lower Devonian Spongiophyton minutissimum.