Pseudopus pannonicus (Squamata), the largest known anguid lizard—Redescription of the type material and new specimens from the Neogene and Quaternary of Hungary and Poland

Herein, we revise the material of the extinct taxon Pseudopus pannonicus from Central Europe, the largest known anguid lizard and iconic member of herpetofaunas from the Upper Cenozoic of Europe. The geographical position of Polgárdi 2, the type locality of P. pannonicus, as well as several other closely located important localities make Central Europe a valuable area of high interest for studies regarding European Cenozoic palaeoherpetological assemblages. We clarified the nature of the type material of Pseudopus pannonicus, showing that it consisted not only of the five specimens originally figured. Instead, the syntype series also includes a considerable number of specimens from Polgárdi 2 that were only described or figured 12 years after the original description of P. pannonicus. Detailed osteological descriptions are provided for specimens from the type series and Polish specimens, with the aid of high-resolution imaging (SEM and μCT scanning), and intraspecific variability is discussed. The articular surface with the lappet of the parietal overlapping the frontal is discussed as a character potentially relevant for the diagnosis of P. pannonicus. We updated the identification of several specimens of P. pannonicus and discussed the biogeographic implications of such revisions. In Poland, P. pannonicus is an abundant component of Neogene and early Quaternary herpetofaunas, known with certainty from the Middle Miocene of Przeworno (the oldest known occurrence of the species globally), the Early-Late Pliocene of Węże I, the Late Pliocene of Rębielice Królewskie I, the Late Pliocene of Węże II, and the Early Pleistocene of Kadzielnia (one of the youngest occurrences documented globally). An indeterminate anguine with Anguis affinities is newly reported from Rębielice Królewskie II. The taxonomic status of other large anguids from the Neogene of Europe is discussed and we conclude that most are junior synonyms of P. pannonicus. We also show that another purported synonym, that is, Ophisaurus intermedius from the Early Pleistocene of Romania, is instead a nomen nudum. We conducted phylogenetic analyses (18 taxa, 65 characters) to understand the relationship of P. pannonicus relative to other anguid representatives and anguid-related group (i.e., glyptosaurids). A single most parsimonious tree (length: 134 steps) was recovered. The clade Pseudopus is stable, comprising the two distinct sister subclades [Pseudopus laurillardi + Pseudopus ahnikoviensis] and [Pseudopus pannonicus + Pseudopus apodus]. These phylogenetic results are in accordance with previously published works.