Early flowers of primuloid Ericales from the Late Cretaceous of Portugal and their ecological and phytogeographic implications

A distinctive feature of the major eudicot diversification that occurred through the Late Cretaceous is the unequivocal presence of Cornales and diverse Ericales. Here we describe well-preserved fossil flowers from the Mira locality in western Portugal, of Campanian-Maastrichtian age, that we assign to a new extinct genus of Ericales with two new species; Miranthus elegans gen. et sp. nov. and Miranthus kvacekii sp. nov. The fossil flowers are pedicellate, structurally bisexual, actinomorphic, pentamerous and isomerous, with five narrowly triangular persistent calyx lobes, a five-lobed corolla, five antepetalous stamens, five staminodes alternating with the petals and a semi-inferior, unilocular ovary. The ovary consists of five carpels and has a raised nectariferous ring with stomata-like openings above the insertion of the perianth, and a long five-angled style. A key feature, which confirms a relationship with Primulaceae s. l., is the free, central dome-shaped placenta that bears numerous, densely spaced ovules. The ovary matures into a capsule containing many, minute, reticulate seeds. Flowers of Miranthus are especially similar to those of extant Samolus, a genus of about twelve species that is sister group to other genera of subfamily Theophrastoideae and that has a disjunct distribution mainly in the Southern Hemisphere. Miranthus also appears to have grown in environments influenced by marine conditions, an ecological preference also seen in Samolus. Miranthus expands the diversity of Ericales known from the Late Cretaceous, and together with previously described fossils provides further evidence that the diversification of Ericales was already underway by the Campanian-Maastrichtian stages of the Late Cretaceous.