Osteology and phylogeny of Zalmoxes (n. g.), an unusual Euornithopod dinosaur from the latest Cretaceous of Romania

Synopsis

The dinosaurs of the Hateg Basin of Transylvania (late Maastrichtian; western Romania) include Theropoda, Sauropoda, Ornithopoda and Ankylosauria. Of these, one of the most enigmatic taxa is the ornithopod that Franz Baron Nopcsa originally described as Mochlodon suessi and M. robustus in 1902. These two species have come to be regarded as a single species of Rhabdodon, R. robustus, which is distinct from R. priscus from the Late Cretaceous of southern France and northern Spain.

This study provides a detailed anatomical revision of the Rhabdodon material that was described originally by Nopcsa during the early decades of the 20th century. It also adds information on material discovered in the Hateg area of Romania since the 1930s. A phylogenetic analysis of basal euornithopods indicates that the non‐hadrosaurid material from Hateg comprises two distinct, but congeneric, species. These two species can be distinguished unambiguously from R. priscus. A new genus Zalmoxes is established for the Romanian ornithopod, comprising Z. robustus comb. nov. (the type‐species of the genus) and Z. shqiperorum sp. nov.

Phylogenetic analysis indicates that the two species of Zalmoxes and R. priscus are united in the monophyletic clade Rhabdodontidae (nov.). Rhabdodontidae constitutes the sibling clade to Iguanodontia. R. septimanicus, M. suessi, and the Villeveyrac Rhabdodon also appear to be members of Rhabdodontidae. The evolutionary implications of this phylogenetic analysis include the recognition of a ghost lineage, extending from the most recent common ancestor of Rhabdodontidae and Iguanodontia, which extends for 73 million years. This extraordinarily long ghost lineage duration may reflect considerable gaps in the history of this group or the geographical isolation of Rhabdodontidae in Europe during much of the Cretaceous period. The area of origin of the Rhabdodontidae + Iguanodontia clade may be North America, while the common ancestor of Rhabdodontidae dispersed to Europe, at that time a marine‐dominated region with tectonically‐active terrestrial habitats. Adult individuals of Z. robustus are smaller than either of its two closest relatives, Z. shqiperorum and R. priscus, within the Rhabdodontidae, or with many species of Iguanodontia and, therefore, is considered a possible paedomorphic dwarf.

Key words:

• Ornithopoda
• Cladistics
• Systematics
• Biogeography
• Heterochrony
• Transylvania