The closest living relative (though not a direct ancestor) of the Galápagos giant tortoise is the Chaco tortoise (Chelonoidis chilensis), a much smaller species from South America. The divergence between C. chilensis and C. nigra probably occurred 6–12 million years ago, an evolutionary event preceding the volcanic formation of the oldest modern Galápagos Islands 5 million years ago.[56] Mitochondrial DNA analysis indicates that the oldest existing islands (Española and San Cristóbal) were colonised first, and that these populations seeded the younger islands via dispersal in a "stepping stone" fashion via local currents.[57][58] Restricted gene flow between isolated islands then resulted in the independent evolution of the populations into the divergent forms observed in the modern species. The evolutionary relationships between the species thus echo the volcanic history of the islands

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