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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 modem land tortoises have probably stayed on land 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 tree of all modem 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 modem 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 re-
emerged 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 driest of deserts.