Transitional fossils and the origin of turtles

Abstract

The origin of turtles is one of the most contentious issues in systematics with three currently viable hypotheses: turtles as the extant sister to (i) the crocodile-bird clade, (ii) the lizard-tuatara clade, or (iii) Diapsida (a clade composed of (i) and (ii)). We reanalysed a recent dataset that allied turtles with the lizard-tuatara clade and found that the inclusion of the stem turtle Proganochelys quenstedti and the 'parareptile' Eunotosaurus africanus results in a single overriding morphological signal, with turtles outside Diapsida. This result reflects the importance of transitional fossils when long branches separate crown clades, and highlights unexplored issues such as the role of topological congruence when using fossils to calibrate molecular clocks.

Figure 1.

The position of turtles based on molecular (1: e.g. Hugall et al. 2007) and morphological datasets (2: e.g. deBraga & Rieppel 1997; 3: Gauthier et al. 1988). The addition of key fossils eliminates the apparent disagreement among morphological datasets in support of turtles outside Diapsida (3). The Permian ‘parareptile’ Eunotosaurus shares uniquely derived features with turtles that help fill important gaps in the evolutionary origin of the turtle shell. Bootstrap (top) and Bremer (bottom) support values are provided for the Eunotosaurus-turtle clade. Star indicates complete shell.

Figure 2.

Strict consensus cladogram of two trees of 483 steps showing the phylogenetic relationships of ‘parareptiles’ when turtles are included in the analysis. CI = 0.4079, RI = 0.6720. Bootstrap (top) and Bremer (bottom) support values are provided for each node. The asterisk (*) indicates a bootstrap value under 50%.