Origin and evolutionary relationships of giant Galápagos tortoises

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Abstract
Perhaps the most enduring debate in reptile systematics has involved the giant Galapagos tortoises (Geochelone nigra), whose origins and systematic relationships captivated Charles Darwin and remain unresolved to this day. Here we report a phylogenetic reconstruction based on mitochondrial DNA sequences from Galapagos tortoises and Geochelone from mainland South America and Africa. The closest living relative to the Galapagos tortoise is not among the larger-bodied tortoises of South America but is the relatively small-bodied Geochelone chilensis, or Chaco tortoise. The split between G. chilensis and the Galapagos lineage probably occurred 6 to 12 million years ago, before the origin of the oldest extant Galapagos island. Our data suggest that the four named southern subspecies on the largest island, Isabela, are not distinct genetic units, whereas a genetically distinct northernmost Isabela subspecies is probably the result of a separate colonization. Most unexpectedly, the lone survivor of the abingdoni subspecies from Pinta Island ("Lonesome George") is very closely related to tortoises from San Cristobal and Espanola, the islands farthest from the island of Pinta, To rule out a possible recent transplant of Lonesome George, we sequenced DNA from three tortoises collected on Pinta in 1906, They have sequences identical to Lonesome George, consistent with his being the last survivor of his subspecies. This finding may provide guidance in finding a mate for Lonesome George, who so far has failed to reproduce.