An elephant described as the loneliest in the world by animal rights campaigners has died in Tokyo at the age of 69.

Hanako was a gift from the government of Thailand in 1949 and had lived alone in a small concrete enclosure at Inokashira Park Zoo, in Tokyo, since she was two.

Her plight became a campaign after photographs of her solitary existence circulated among animal-lovers. She is not believed to have seen another elephant since arriving in Japan.

Nearly 470,000 people signed an online petition set up in the name of Elephant Freedom Fighters demanding that Hanako be allowed to leave her "concrete prison" and to see out the rest of her life at a sanctuary in Thailand.

The zoo said a transfer would not be possible because she was too old to endure the journey. Another organisation, the Friends of Asian Elephants, proposed that she at least be allowed to socialise with other animals and have some greenery in her enclosure.