Morphological, developmental, and cytochemical examinations were made with light and electron microscopy on the apical apparatuses of Morchella esculenta, Helvella crispa, and Rhizina undulata, all species with large, stipitate apothecia. Ascus tips in R. undulata were markedly thinner walled than the rest of the ascus, while those in M. esculenta and H. crispa were slightly thinner than the rest of the ascus wall. Lipid bodies were detected in developing asci of H. crispa and M. esculenta. Their unique occurrence in asci of members of the Morchellaceae and Helvellaceae may have taxonomic significance. With the electron microscope, opercula were distinguished cytochemically in all three species. In H. crispa and M. esculenta dehiscent zones were found to be restricted in the inner layer of the ascal wall. Characters of the apical apparatuses of H. crispa and M. esculenta suggest greater taxonomic relatedness between these species than with any other operculate group. The apical apparatus of R. undulata differed notably from the other species.The present series of studies has demonstrated distinct variability of the operculate ascus and its apical apparatus in morphology, cytochemistry, and development. Several major forms of the apical structures were observed. These examinations support the chemotaxonomic and cytological investigations on representatives of the Pezizales made previously by Arpin and Berthet. Outside of the Thelebolaceae, members of the Pezizales are chiefly characterized by the operculate dehiscence of their asci. Members of the Thelebolaceae eject their spores through a variety of dehiscent mechanisms. Present examinations of those representatives of the Thelebolaceae with functionally operculate apparatuses, i.e., Lasiobolus and Coprotus, support their transferrance to the Pyronemaceae. Taxa which form nonoperative opercula, i.e.,Ascozonus and Trichobolus, also show closer affinities with the Pyronemaceae than with the nonoperculate representatives of the Thelebolaceae. The nonoperculate members of the Thelebolaceae apparently do not belong in the operculate Discomycetes. The operculate ascus wall is structurally compared with the pored and bitunicate ascus walls. The terms 'bitunicate' and 'unitunicate' are redefined.
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