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Filicalean frond segments bearing indusiate sori with gradate development have been discovered in Pennsylvanian deposits of eastern North America. The specimens combine characters of Schizaeaceae, Gleicheniaceae, Cyatheaceae and Hymenophyllaceae, and together with data from other sources, indicate that the Filicales were in the midst of an evolutionary radiation during the late Paleozoic. Evidence from fossil and extant species prompts the hypothesis that filicalean ferns originated near the base of the Carboniferous, and have undergone three major evolutionary radiations.
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Gar W. Rothwell (1987) studied this question.
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