Morphology and ultrastructure of cicatricose spores found in an isolated sporangium-bearing structure of a schizaealean fern from the Lower Cretaceous rocks at Casal do Borracho, Torres Vedras, western Portugal

Fossil spores provide important insights into the evolutionary history of ferns, including their morphological changes and adaptations over time. The morphology and ultrastructure of trilete spores with cicatricose sculpture composed of sets of parallel or near-parallel ribs (muri) have been studied. Observations on the spores were performed using light microscopy, scanning electron microscopy, and transmission electron microscopy. The cicatricose spores were extracted from a single sporangium-bearing structure collected from Casal do Borracho mesofossil flora near Torres Vedras (Lusitanian Basin, western Portugal) from sediments belonging to the Almargem Formation, considered to be of late Barremian or early Aptian age (and possible younger than that). The sporangium-bearing structure of a schizaealean fern and in situ cicatricose spores are compared with the family Anemiaceae. On the basis of spore size and shape, laesura length, with about 20 proximal narrow ribs, oriented perpendicular to the spore sides, and three sets of distal ribs forming V-shaped patterns, the cicatricose spores in the present study are clearly assigned to the fossil-taxon Cicatricosisporites venustus. The spore ultrastructure includes a homogeneous exospore of a blechnoid type with a slight difference in the thickness in the proximal, distal, and equatorial regions. The morphology and ultrastructure of the studied spores are corresponded to Anemia type.