THE "SEEDS" ON PADGETTIA READI ARE INSECT GALLS: REASSIGNMENT OF THE PLANT TO ODONTOPTERIS, THE GALL TO OVOFOLIGALLITES N. GEN., AND THE EVOLUTIONARY IMPLICATIONS THEREOF

The Early Permian (Asselian) Euramerican plant Padgettia readi Mamay is reassigned to Odontopteris Brongniart, as O. readi (Mamay) Stull et al. n. comb. Distinctive elongate structures on neuropteroid pinnules of this plant, previously interpreted as fructifications, are herein reinterpreted as foliar histoid galls, structurally analogous to blister or vein galls, and probably induced by an early lineage of hemipterans or mites. These distinctive features are assigned to the new gall ichnogenus Ovofoligallites Labandeira, n. ichnogen. n. ichnosp, as O. padgetti Labandeira. The Early Permian association between an Odontopteris host and Ovofoligallites gallers probably originated during the Middle Pennsylvanian as a similar, antecedent association between Macroneuropteris scheuchzeri (Hoffmann) Cleal, Shute, and Zodrow and the maker of U-shaped surface features long known as a distinctive, unattributed damage type, but now recognizable as a likely gall. The persistence of this association between the galler and certain medullosan pteridosperms into the Permian adds to the morphological richness of the Permian galler insect fauna. The Permian ecological expansion of galling insects resulted in colonization of new host plants, primarily through a shift from the consumption of entire, mostly pteridophyte axial organs during the Pennsylvanian to the partitioning of seed plant tissues in leaves and small branches in the Permian. The Ovofoligallites galler was part of a diverse Permian galler guild involving a variety of plant taxa, organs and tissues that overwhelmingly targeted multiple lineages of seed plants.