Lizards and snakes from the earliest Miocene of Saint-Gérand-le-Puy, France: an anatomical and histological approach of some of the oldest Neogene squamates from Europe

Abstract

Background
The earliest Miocene (Aquitanian) represents a crucial time interval in the evolution of European squamates (i.e., lizards and snakes), witnessing a high diversity of taxa, including an array of extinct forms but also representatives of extant genera. We here conduct a taxonomical survey along with a histological/microanatomical approach on new squamate remains from the earliest Miocene of Saint-Gérand-le-Puy, France, an area that has been well known for its fossil discoveries since the nineteenth century.

Results
We document new occurrences of taxa, among which, the lacertid Janosikia and the anguid Ophisaurus holeci, were previously unknown from France. We provide a detailed description of the anatomical structures of the various cranial and postcranial remains of lizards and snakes from Saint-Gérand-le-Puy. By applying micro-CT scanning in the most complete cranial elements of our sample, we decipher previously unknown microanatomical features. We report in detail the subsurface distribution and 3D connectivity of vascular channels in the anguid parietal. The fine meshwork of channels and cavities or sinuses in the parietal of Ophisaurus could indicate some thermoregulatory function, as it has recently been demonstrated for other vertebrate groups, providing implications for the palaeophysiology of this earliest Miocene anguine lizard.

Conclusions
A combination of anatomical and micro-anatomical/histological approach, aided by micro-CT scanning, enabled the documentation of these new earliest Miocene squamate remains. A distinct geographic expansion is provided for the extinct anguine Ophisaurus holeci and the lacertid Janosikia (the closest relative of the extant insular Gallotia from the Canary Islands).