Date: June 27 2012
He was on Ecuador's bank notes and stamps, an evolutionary remnant, a money-spinning tourist attraction and an icon of international conservation. No one knew if he was gay, impotent, terminally bored or just very shy. And he is thought to have been about 100 years old and in the prime of life when he died on Sunday at the Charles Darwin research centre in the Galapagos. But the giant tortoise known as Lonesome George and commonly called the ''rarest animal on Earth'' could, in fact, have been far older - or younger.
In the 40-odd years that he spent in a field on Santa Cruz island, having been relocated from Pinto Island in 1972, the 90 kilogram, 150-centimetre-long animal showed little interest in either man or other tortoises. He mostly ignored the female company provided to encourage him to breed, kept his 90-centimetre-long scraggy neck down in the long grass, and only responded to his long-time keeper, Fausto Llerena, who runs a tortoise breeding centre.
''The park ranger in charge of looking after the tortoises found Lonesome George, his body was motionless,'' the head of the Galapagos National Park, Edwin Naula, says. ''His life cycle came to an end.''
George was found near a water hole but no one knows how or why he died and evolutionary scientists are still baffled by his life in the volcanic Pacific islands that inspired Darwin's theories on evolution, which are now a global laboratory for conservation.
But history shows that the last known representative of the giant Galapagos tortoise subspecies Chelonoidis nigra abingdonis had every reason to shun humanity. His closest relatives had been systematically exterminated for food or for oil by passing whalers and seal hunters in the 19th century and his habitat on Pinto Island had then been devastated by introduced feral goats. George possibly has relations on neighbouring Isabela Island but it is more likely his whole subspecies is now extinct - the end of what was probably a 10-million-year-old line.
On Monday, scientists who had spent time with George recalled his peculiar ways. ''George was the last of his kind,'' the head vet of the Houston Zoo, Joe Flanagan, said. ''He had a unique personality. His natural tendency was to avoid people. He was very evasive. He had his favourites and his routines but he really only came close to his keeper, Llerena. He represents what we wanted to preserve forever. When he looked at you, you saw time in the eyes.'' Flanagan knew George for more than 20 years.
Scientists' attempts to get him to mate with other giant tortoises from the Galapagos Islands, and to eventually repopulate Pinto, all failed and were often comical. Artificial insemination did not work, nor did a $10,000 reward offered by the Ecuadorean government for a suitable mate. In the 1990s, Sveva Grigioni, a Swiss zoology graduate student smeared herself with female tortoise hormones and, in the cause of science, spent four months trying to manually stimulate him - all to no avail.
Breeding success was thought to be close in 2008 and in 2009, when George unexpectedly mated with one of two female companions he was living with, but although two clutches of eggs were collected and then incubated, they all failed to hatch.
However, Henry Nicholls, author of Lonesome George: The Life and Loves of the World's Most Famous Tortoise, reported that he was irresistibly attracted to the late Lord Devon's wartime helmet, presumably because it resembled the shell of a young tortoise.
Even after being put on a diet, the celibate tortoise with the scraggy neck, who could have been expected to live until he was well over 200, remained obstinately alone.
Conservation scientists on Monday said George was important because he symbolised both the rapid loss of biodiversity now taking place around the world, and provided the inspiration to begin restoring it in places such as the Galapagos Islands.
Richard Knab of the Galapagos Conservancy, which is running giant tortoise breeding programmes with the Ecuadorean government says: ''Because of George's fame, Galapagos tortoises which were down to just a few animals on some islands have recovered their populations. He opened the door to finding new genetic techniques to help them breed and showed the way to restore habitats.''
In 1960, 11 of the Galapagos Islands' original 14 populations of tortoises remained, and most were on the point of extinction. Today, about 20,000 giant tortoises of different subspecies inhabit the islands and most of the wild goats have been eradicated.
But George will be sorely missed for financial reasons, too. As the star of the islands and an icon of global wildlife, he helped attract 180,000 money-spinning visitors a year to the archipelago 1000 kilometres off the Ecuadorean coast. Now, says the national park, he is likely to become a conservation relic and will probably be embalmed and displayed - alone until the end.
■ John Vidal is The Guardian's environment editor
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