Walter Joyce and Jacques Gauthier, at Yale University, obtained three measurements in these particular bones of 71 species of living turtles and tortoises. They used a kind of triangular graph paper to plot the three measurements against one another. All the land tortoise species formed a tight cluster of points in the upper part of the triangle; all the water turtles cluster in the lower part of the triangular graph. There was no overlap, except when they added some species that spend time both in water and on land. Sure enough, these amphibious species show up on the triangular graph approximately half way between the ‘wet cluster’ of sea turtles and the ‘dry cluster’ of land tortoises. The next step was to determine where the fossils fell. The bones of P. quenstedti and P. talampayensis leave us in no doubt. Their points on the graph are right in the thick of the dry cluster. Both these fossils were dry-land tortoises. They come from the era before our turtles returned to the water.

　　You might think, therefore, that modern land tortoises have probably stayed on and ever since those early terrestrial times, as most mammals did after a few of them went back to the sea. But apparently not. If you draw out the family three of all modern turtles and tortoises, nearly all the branches are aquatic. Today’s land tortoises constitute a single branch, deeply nested among branches consisting of aquatic turtles. This suggests that modern land tortoises have not stayed on land continuously since the time of P. quenstedti and P. talampayensis. Rather, their ancestors were among those who went back to the water, and they then reemerged back onto the land in (relatively) more recent times.

　　Tortoises therefore represent a remarkable double return. In common with all mammals, reptiles and birds, their remote ancestors were marine fish and before that various more or less worm-like creatures stretching back, still in the sea, to the primeval bacteria. Later ancestors lived on land and stayed there for a very large number of generations. Later ancestors still evolved back into the water and became sea turtles. And finally they returned yet again to the land as tortoises, some of which now live in the direst of deserts.