Insights into stem Batomorphii: A new holomorphic ray (Chondrichthyes, Elasmobranchii) from the upper Jurassic of Germany

The Late Jurassic fossil deposits of southern Germany, collectively known as the 'Solnhofen Archipelago', are one of the world's most important sources of Mesozoic vertebrates. Complete skeletons of cartilaginous fishes (Chondrichthyes), whose skeletal remains are rare in the fossil record and therefore all the more valuable, are represented, among others, by exceptionally well-preserved rays (superorder Batomorphii). Despite their potential for research in several areas, including taxonomy, morphology, ecology, and phylogeny, the number of studies on these chondrichthyans is still very limited. Here, we identify a previously unknown ray, †Apolithabatis seioma gen. et sp. nov., which represents the first record of a ray species from the upper Kimmeridgian of Painten, Germany, and thus the oldest Late Jurassic ray taxon from Germany based on skeletal remains. This new batomorph is characterised by a unique body shape and a combination of skeletal features that distinguish it readily from all other known Late Jurassic rays. Two different morphometric approaches confirm differences in body shape and proportions to all known Late Jurassic conspecifics. We thus extend the recent taxonomic revision of these rays and include all described holomorphic specimens in a phylogenetic framework using strict cladistic principles. The phylogenetic analysis reveals all Late Jurassic batomorphs to represent a monophyletic group, for which we introduce the new order Apolithabatiformes, which is sister to all other batomorphs representing a stem group. While the phylogenetic relationships within Apolithabatiformes ord. nov. remain largely unresolved, †Apolithabatis gen. nov. is placed as the sister to †Aellopobatis. This highlights that, despite considerable progress in our understanding of the diversity and phylogeny of early rays, difficulties remain in establishing robust relationships within batomorphs. We therefore emphasise the importance of comprehensive studies of completely preserved fossil cartilaginous fishes to obtain a better understanding of chondrichthyan evolution and their systematics in deep time.