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The evolution and interrelationships of carnivorous squamates (mosasaurs, snakes, monitor lizards, Gila Monsters) are a contentious part of reptile systematics and go to the heart of conflict between morphological and molecular data in inferring evolutionary history. One of the best-preserved fossils in this motley grouping is “Saniwa” feisti Stritzke, 1983, represented by complete skeletons from the early-middle Eocene of Messel, Germany. We re-describe it on the basis of superficial examination, stereoradiography, and high-resolution X-ray computed tomography of new and published specimens. The scalation of the lizard is unique, consisting of small, keeled scales on the head (including a row of enlarged medial supraorbitals) and large, rhomboidal, keeled scales (invested by osteoderms) that covered the rest of the body. Two paired longitudinal rows of enlarged scales ran down the neck. The head was laterally compressed and box-shaped due to the presence of a strong canthal-temporal ridge; the limbs and tail were very long. Notable osteological features include: a toothed, strap-like vomer; septomaxilla with a long posterior process; palpebral with a long posterolateral process; a lacrimal boss and a single lacrimal foramen; a well-developed cultriform process of the parabasisphenoid; two hypoglossal (XII) foramina in addition to the vagus; a lack of resorption pits for replacement teeth; and possibly the presence of more than one wave of developing replacement teeth per locus. There are no osteological modifications suggestive of an intramandibular hinge, but postmortem displacement of the angular-prearticular-surangular complex in multiple specimens suggests that there might have been some degree of mobility in the lower jaw based on soft-tissue modifications. Using phylogenetic analyses on a data-set comprising 473 morphological characters and 46 DNA loci, we infer that a monophyletic Palaeovaranidae Georgalis, 2017, including Eosaniwa Haubold, 1977, lies on the stem of Varanidae Merrem, 1820, basal to various Cretaceous Mongolian taxa. We transfer feisti to the new genus Paranecrosaurus n. gen. Analysis of gut contents reveals only the second known specimen of the cryptozoic lizard Cryptolacerta hassiaca Müller, Hipsley, Head, Kardjilov, Hilger, Wuttke & Reisz, 2011, confirming a diet that was at least partly carnivorous; the preservation of the teeth of C. hassiaca suggests that the gastric physiology of Paranecrosaurus feisti (Stritzke, 1983) n. comb. had high acidity but low enzyme activity. Based on the foregoing and linear discriminant function analysis, we reconstruct P. feisti n. comb., as a powerful, widely roaming, faunivorous-carnivorous stem monitor lizard with a sensitive snout. If the molecular phylogeny of anguimorphs is correct, then many of the features shared by Helodermatidae Gray, 1837 and Varanidae must have arisen convergently, partly associated with diet. In that case, a reconciliation of morphological and molecular data would require the discovery of equally primitive fossils on the helodermatid stem.
Our knowledge of early evolution of snakes is improving, but all that we can infer about the evolution of modern clades of snakes such as boas (Booidea) is still based on isolated bones. Here, we resolve the phylogenetic relationships of Eoconstrictor fischeri comb. nov. and other booids from the early-middle Eocene of Messel (Germany), the best-known fossil snake assemblage yet discovered. Our combined analyses demonstrate an affinity of Eoconstrictor with Neotropical boas, thus entailing a South America-to-Europe dispersal event. Other booid species from Messel are related to different New World clades, reinforcing the cosmopolitan nature of the Messel booid fauna. Our analyses indicate that Eoconstrictor was a terrestrial, medium- to large-bodied snake that bore labial pit organs in the upper jaw, the earliest evidence that the visual system in snakes incorporated the infrared spectrum. Evaluation of the known palaeobiology of Eoconstrictor provides no evidence that pit organs played a role in the predator–prey relations of this stem boid. At the same time, the morphological diversity of Messel booids reflects the occupation of several terrestrial macrohabitats, and even in the earliest booid community the relation between pit organs and body size is similar to that seen in booids today.