As the last of his kind, his life was a lonely one and his death brought the extinction of a lineage of animals that stretched back hundreds of thousands of years.
Now Lonesome George, the last giant tortoise from Pinta Island in the Galapagos Islands who died just over a month ago, may be able to achieve in death what he could not in life and produce an heir.
Scientists have cryogenically frozen tissue taken from the five foot long reptile just after his death in the hope that they may be able to resurrect the subspecies of Galapagos giant tortoise.
By using the same cloning techniques that created Dolly the Sheep, they believe it may be possible to one day bring the now extinct Pinta Island tortoises, or Chelonoidis nigra abingdonii as they are known scientifically, back to life.
It is a fitting legacy for an animal that had become a national icon in Ecuador, featuring on the countrys bank notes. His plight became a symbol for the efforts to conserve threatened species around the world, and attempts to find him a mate were followed by a global audience.
As the last of Pinta Island giant tortoises, his death appeared to take the hopes of saving the animals with him.
Dr Oliver Ryder, director of genetics at San Diego Zoos Institute for Conservation Research, said he had hoped to take a tissue sample from Lonesome George while he was alive to store it with other samples from endangered species in the institutes Frozen Zoo.
Genetic material and cells from endangered animals are kept cryogenically frozen in the collection, with the aim of studying their biology and eventually being able to use cloning to save them from extinction.
Dr Ryder said it was a cruel irony that George had died before his cells were collected, but it is hoped that the cells were frozen with enough time to ensure they could be used in the future.
We always thought viable cell cultures should be saved from this animal, he said.
This may be the only chance well have to preserve, document and study this tortoise subspecies," he added in an interview with Discovery News.
Scientists from the Frozen Zoo at San Diego Zoo flew to Ecuador, which owns the Galapagos archipelago, as soon as news of Georges death emerged, in a bid to ensure cells from the 100 year old tortoise could be preserved.
Using canisters of frozen nitrogen, the cells were kept in a cryogenic chamber normally used for storing cells for artificially inseminating cattle while arrangements were made transfer them to a long term storage facility at Frozen Zoo.
During his life Lonesome George found worldwide fame after he was discovered on Pinta Island in 1971.
The giant tortoises there had already been deemed extinct after goats introduced by human settlers destroyed much of the vegetation the reptiles relied upon.
But with no female Pinta Island tortoises left alive, it was apparent he was doomed to spend the rest of his long life plodding slowly around his home alone until subspecies finally went extinct.
The discovery sparked a huge effort to find him a mate so the animals might continue to survive. As the descendant of giant tortoises that arrived on Pinta Island around 300,000 years ago, it was unclear whether evolution during that time had left George unable to mate with related tortoises on nearby islands.
Indeed, it was the very differences between the populations of the giant tortoises that had developed over generations of living isolated on the islands of the Galapagos that had helped lead Charles Darwin form his theory of evolution.
Attempts to get George to mate with tortoises from neighbouring islands failed but last year hopes were raised when genetic testing revealed that his closest relatives were the giant tortoises of Espanola Island.
But before much more could be done, Lonesome George was found dead by his keepers near a water hole. A post-mortem found he died of natural causes.
His death brought nationwide mourning in Ecuador and the countrys President Rafael Correa even paid tribute to the reptile in an address to the nation.
In the UK, Sir David Attenborough, who had filmed with the animal shortly before its death, described the news as very poignant and sad.
After finding George dead, vets and keepers moved fast to ensure his body could be preserved, moving his 200lb body to a freezer.
Marilyn Cruz, a veterinary surgeon who is coordinator of Agrocalidad in the Galapagos, the government agency that oversees agriculture and biosecurity on the islands, took a sample of skin during the post mortem so it could be cryogenically frozen.
Liquid nitrogen also had to be shipped back and forth from the Ecuadorean mainland to help prevent the cells from thawing.
The tissue samples taken from Lonesome George will now be kept frozen until scientists have conducted the necessary research to attempt to clone him. The rest of his body is to be embalmed and put on display at the research centre where he lived in Santa Cruz.
Research in to the cloning of endangered species is still at an early stage as most cloning to date has been achieved with animals whose biology is well understood such as cats, dogs and domestic farm animals like sheep and cows.
Recently, endangered black-footed cats were successfully cloned, while a wild ox called a gaur, wild cattle called banteng and a type of mountain goat called the Pyrenean ibex have been also been cloned, raising hopes for other endangered animals.
Among the challenges researchers face will be finding a suitable tortoise species to supply eggs for the cloning process.
DNA from one of Georges cells would be used to replace that from the donor egg, but this does not necessarily work with any combination of species as they can be incompatible.
Provided the resulting egg can be kick started into developing into an embryo, it would then need to be implanted into a suitable female tortoise to carry it before it can lay the fully developed eggs containing the clone.
Linda Cayot, scientific adviser of the conservation group Galapagos Conservancy, said: It has been a difficult time not just because we lost Lonesome George as an individual but also lost his species forever.
We are trying to save some of Lonesome Georges tissues. Cloning in reptiles has not been developed at all, as most of the success has been in mammals. The technology and methodology is still decades away.
Hopefully they will be able to maintain cells in cryogenics until that can happen.
