Frenelopsis (Coniferales: Cheirolepidiaceae) and related male organ genera from the Lower Cretaceous of Spain

Vegetative plant remains and microsporangiate cones, related to the fossil genera Frenelopsis and Classostrobus respectively, were studied in three localities from the Lower Cretaceous of the Pyrenees and Iberian Ranges (Spain). Sterile remains belong to three different species: F. rubiesensis Barale, F. ugnaensis sp. nov. and F. turolensis sp. nov. The male cones Classostrobus ugnaensis sp. nov. and C. turolensis sp. nov. are associated with the two latter species respectively, and C turolensis is found in connection with F. turolensis. Intemodes of F. ugnaensis display small hemispherical papillae and long conical hairs covering the entire cuticle surface, while F. turolensis has a strip of hairs near the leaf margin, in addition to more massive outer stomatal papillae and higher stomatal density. The tiny cones of C ugnaensis bear microsporophyll cuticles capped with vermiculate flattened papillae and a fringe of marginal hairs, while C turolensis has a central area of microsporophylls covered by long conical hairs. Sedimentological and taphonomic analyses show that the assemblages studied belong to a wide spectrum of habitats. These indicate that Spanish Lower Cretaceous frenelopsids were xeromorphic plants, able to adapt to helophytic and riparian habitats, and grew in brackish coastal marshes and fluvio-lacustrine freshwater environments.