Reinterpretation of Paleoazolla: a heterosporous water fern from the Late Cretaceous of Patagonia, Argentina.

PREMISE: Undoubtedly, fossils are critical for understanding evolutionary transformations in deep time. Here, we reinvestigate the microspores and megaspores of Paleoazolla patagonica, a water fern found in Late Cretaceous sediments of the Chubut Province, Patagonia, Argentina, which provides novel evidence on the past history of the water fern clade. The study was based on recently collected specimens and additional observations of the original material. METHODS: Most specimens analyzed herein were obtained from new palynological samples collected at the Cerro Bosta and Cañadón del Irupé localities, La Colonia Formation. Samples were mechanically disaggregated and treated with hydrofluoric and hydrochloric acid. Spores were studied using standard light microscopy and scanning electron microscopy. We also reexamined the original materials. RESULTS: The newly described characters of Paleoazolla include the presence of heterosporangiate sori composed of one ellipsoidal megasporangium surrounded by three to four oval microsporangia, megasporangium containing one hairy massula that encloses two trilete megaspores (rarely one or three), and microsporangia containing numerous microspore massulae with non-septate multibarbed glochidia and one trilete microspore per massula. CONCLUSIONS: The reinterpretation has revealed a novel set of characters for understanding the evolution of heterosporous water ferns. The presence of two megaspores in the megasporangium of Paleoazolla exposes serious gaps in the current knowledge on the evolution of monomegaspory in heterosporous water ferns, a fact that emphasizes the need of including fossils within phylogenies to elucidate patterns of character acquisition among water ferns.