Abstract

A new marine lizard showing exceptional soft tissue preservation was found in Late Cretaceous deposits of the Apulian Platform (Puglia, Italy). Primitivus manduriensis gen. et sp. nov. is not only the first evidence of the presence of dolichosaurs in a southern Italian Carbonate Platform, filling a palaeogeographic gap in the Mediterranean Tethys, but also extends the range of this group to the upper Campanian–lower Maastrichtian. Our parsimony analysis recovers a monophyletic non-ophidian pythonomorph clade, including Tetrapodophis amplectus at the stem of Mosasauroidea + Dolichosauridae, which together represent the sister group of Ophidia (modern and fossil snakes). Based on Bayesian inference instead, Pythonomorpha is monophyletic, with Ophidia representing the more deeply nested clade, and the new taxon as basal to all other pythonomorphs. Primitivus displays a fairly conservative morphology in terms of both axial elongation of the trunk and limb reduction, and the coexistence of aquatic adaptations with features hinting at the retention of the ability to move on land suggests a semi-aquatic lifestyle. The exceptional preservation of mineralized muscles, portions of the integument, cartilages and gut content provides unique sources of information about this extinct group of lizards. The new specimen may represent local persistence of a relict dolichosaur population until almost the end of the Cretaceous in the Mediterranean Tethys, and demonstrates the incompleteness of our knowledge of dolichosaur temporal and spatial distributions.

Highlights

  • Pythonomorpha is a clade including both extinct and extant squamates

  • The spectra resulting from the Scanning electron microscopy (SEM)/energy dispersive X-ray (EDX) analyses of the soft tissues, bones and sediment are presented in figure 9, and electronic supplementary material, figure S10

  • Both bony and soft tissues are rich in phosphorus (P), and display a very similar composition, while there is no P in the embedding sediment

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Summary

Introduction

Pythonomorpha (mosasauroids, dolichosaurs and snakes) is a clade including both extinct and extant squamates. We present new data from an extremely well-preserved specimen, including soft tissue remains, of the first dolichosaur from the latest Cretaceous of southern Italy (Puglia), recovered from a new Lagerstätte-quality locality. This new finding fills a palaeogeographic gap in the Mediterranean Tethys for this group, being the first record from the Apulian Platform, but it extends the range of dolichosaurs sensu Nopcsa [18] by about 10 million years (from the Santonian to the upper Campanian– lower Maastrichtian). The astonishing preservation of the soft tissues provides an unprecedented source of information to help us better understand the morphology of pythonomorphs and their interrelationships

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