Spore wall ultrastructure in the early lycopsid Leclercqia (Protolepidodendrales) from the Lower Devonian of North America: Evidence for a fundamental division in the lycopsids

Documenting the morphology and ultrastructure of spores from known Silurian-Devonian plants clarifies organization and probable affinities of dispersed spores and contributes to analyses of evolutionary changes and phylogenetic relationships in early plants. In this study of fossil in situ spores from the early protolepidodendralean lycopsid Leclercqia, we identified new characters including an additional synapomorphy of the ligulate lycopsid clade. A detailed light (LM), scanning electron (SEM), and transmission electron microscope (TEM) analysis of spores from two species of Leclercqia from the Lower Devonian (Emsian) of New Brunswick, eastern Canada, L. andrewsii and L. complexa, shows both are homosporous, yielding spores belonging to the dispersed spore form taxon Acinosporites lindlarensis. Important features of wall ultrastructure include the presence of a paraexospore, peculiar exospore-derived, peg-like structures located in the gap between the outer exospore/inner paraexospore, and multilamellate regions in the interradial areas of the proximal surface. Similar interradial multilamellate regions occur in other ligulate lycopsids (fossil and extant). This character is probably a further synapomorphy for the ligulate lycopsid clade, within which heterosporous lycopods form a monophyletic group. These data suggest the ligule and interradial multilamellate region appeared prior to heterospory.