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galapagos tortoise chelonoidis nigra warm tone galápagos giant niger species large genus lifespans wild 100 years one longest-lived vertebrates adult captive individual shell saddleback conservation efforts juveniles exceeded threatened insular radiations gigantic evolutionary history breeding pair bony dull brown grey plates ribs integral skeleton characteristic scute segment withdraw head neck fore limbs protection stumpy gigantism size larger gigantothermy fossil biogeographic spectrum carapace morphology ranging upward arching edge resembling largest toitoise earth live 180 old found islands