Evolution and palaeoecology of early turtles: a review based on recent discoveries in the Middle Jurassic

In recent years, no less than five new species of early turtles have been described worldwide. Among them are three new turtles from Middle Jurassic deposits that partially fill a previous temporal and morphological gap in our knowledge of the early evolution of these shelled amniotes: Heckerochelys romani, Condorchelys antiqua and Eileanchelys waldmani. For the first time, the phylogenetic position of these three new species is tested in the context of the two presently competing cladistic models of turtle evolution. The addition of these taxa to each matrix does not favour or alter any of the two opposed hypotheses. However, it is demonstrated here that, by documenting yet unknown stages in the evolution of several morphological structures, these three species give stronger support to the model of an extended phylogenetic stem for turtles. These new lines of evidence include the structure of the vomer, the position of the aditus canalis stapedio-temporalis and of the posterior opening of the canalis cavernosus, and the morphology of the processus interfenestralis of the opisthotic.Recent discoveries also reinvigorate the debate about the palaeoecology of early turtles. Whereas simple morphological characters (e.g., shell fontanelle, ligamentous bridge, flattened carapace) can be misleading, forelimb proportions and shell bone histology have led to the conclusion that most stem turtles (i.e., Proganochelys quenstedti, Palaeochersis talampayensis, Proterochersis robusta, Kayentachelys aprix and meiolaniids) were terrestrial forms. At least two out of the five recently described early turtles have been convincingly interpreted as having aquatic habits: Odontochelys semitestacea and Eileanchelys waldmani. More investigation is needed, but this will undoubtedly trigger further debate on the primitive ecology of turtles and on the origin of aquatic habits in testudines (i.e., the crown-group), respectively.