Shell bone histology of solemydid turtles (stem Testudines): palaeoecological implications

Lately, solemydid turtles have been repeatedly recovered as stem Testudines, indicating that they belong to neither one of the two major branches of crown turtles, the Pancryptodira and Panpleurodira. Despite their wide temporal (Late Jurassic to Late Cretaceous) and spatial (North America and Europe) distributions, solemydid turtles are not particularly well known, as exemplified by the fact that only a single skull has been described for the whole group so far. Furthermore, the palaeoecology of solemydid turtles is still contested with hypotheses ranging from semi-aquatic to terrestrial lifestyles. However, the habitat preference of stem Testudines, such as solemydids, is important to understand the evolution and early radiation of the turtle crown, which is primitively aquatic. Here we describe the shell bone microanatomy and histological microstructures of solemydid turtles using a broad sample of taxa of different ages and localities, as well as review previous histological accounts, to elucidate the palaeoecology of the group independent of the geological setting and gross anatomy of the fossil finds. Our results indicate that Solemydidae share unique histological features pertaining to their strongly ornamented shell bones, which a) in cases allow taxonomic identification of even small shell fragments and b) unambiguously corroborate a terrestrial lifestyle of its members. The latter further supports a terrestrial lifestyle preference of most representatives of the turtle stem