Two new ootaxa from the late Jurassic: The oldest record of crocodylomorph eggs, from the Lourinhã Formation, Portugal

The Late Jurassic Lourinha Formation is known for its abundant remains of dinosaurs, crocodylomorphs and other vertebrates. Among this record are nine localities that have produced either dinosaur embryos, eggs or eggshell fragments. Herein, we describe and identify the first crocodiloid morphotype eggs and eggshells from the Lourinha Formation, from five occurrences. One clutch from Cambelas, composed of 13 eggs, eggshell fragments from Casal da Rola and Peralta, one crushed egg and eggshells from Paimogo North, and four crushed eggs as well as eggshell fragments from Paimogo South. We observed and confirmed diagnostic morphological characters for crocodiloid eggshells and which are consistent with a crocodylomorph affinity, such as the ellipsoidal shape, wedgeshaped shell units, triangular extinction under cross-polarized light, and tabular ultrastructure. This material is distinctive enough to propose two new ootaxa within the oofamily Krokolithidae, Suchoolithus portucalensis, oogen. and oosp. nov., for the material from Cambelas, the most complete clutch known for crocodiloid eggs, and Krokolithes dinophilus, oosp. nov., for the remaining material. These are the oldest crocodylomorph eggs known, extending the fossil record for this group to the Late Jurassic. Furthermore, except for the clutch from Cambelas, the material was found with theropod eggs and nests, in the other four occurrences, which seem to suggest some form of biological relationship, still unclear at this point.