A former circus elephant that crushed her keeper to death at a New Zealand zoo in 2012 has died in San Diego.
Mila was retired from the circus in 2009 after years of campaigning by animal rights groups, who say she was psychologically damaged by 30 years spent as a circus elephant.
She was released from the circus only to kill her keeper, who was trying to calm the elephant after something gave Mila a fright.
The average lifespan for an African elephant living in the wild is 70 years. Mila was just 44 years old when she died at San Diego Zoo on Tuesday morning.
Hans Kriek from Save Animals From Exploitation (SAFE) told Newshub life in captivity led to her early death.
"It's quite typical that elephants in captivity live a much shorter life. It's mainly because they don't get enough exercise, so their whole body can't cope," Mr Kriek said.
"It's quite sad. We just hope this will be another reason for people to reflect and think back and realise that keeping these magnificent wild animals in captivity is just no longer acceptable."
A study published in 2008 found elephants in large reserves live twice as long as those kept in zoos. The researchers compared the mean life expectancy of elephants in European zoos with Kenya's large Amboseli National Park over a 45-year time period. They attributed the stark differences in life expectancy to obesity and the stress of zoo life.
Critics at the time said the data failed to account for recent advances in zoo welfare.
Despite Mila's early death, Mr Kriek said her years spent with other elephants at San Diego Zoo would have been the best of her life.
"At least we can take some comfort that in the last years of her life - she had companionship, she had probably the best life she ever had during her entire 44-year life, but she was still in captivity and she still died a premature death."
Video filmed by San Diego Zoo when Mila had recently been moved showed her settling into her new life and socialising with the other elephants.
Animal activists campaigned for two decades to have Mila removed from the circus, a life Mr Kriek says was cruel.
"Elephants live in family groups and she lived by herself for decades and suffered mentally very badly. Anyone could see her stereotypical behaviour. She was chained up, she was swaying from side to side, and that's a clear sign that the animal's not coping with her environment.
"We were very pleased that after decades of campaigning, she was finally released and she then ended up at a local New Zealand zoo, where she was very much loved and very much cared for, but the problem was she was still by herself and she had no companionship."
Mila spent two-and-a-half years at Auckland's Franklin Zoo after her release from the circus. She was due to be shifted to a sanctuary in the United States when she crushed keeper Dr Helen Schofield to death.
The coroner's report, released in 2013, found the elephant had become agitated, probably after being shocked by the electric fence surrounding her enclosure.
Dr Schofield entered the enclosure to help calm the elephant, but Mila moved toward her quickly, the Coroner said.
When Dr Schofield turned to leave the enclosure, she slipped and fell. Before she could get away, Mila reached Dr Schofield and put her trunk around her, in an action described by a witness as "protective".
The coroner found Mila was being "gentle" to Dr Schofield until the end.
The African elephant
- They are the largest land mammals on Earth, bigger than their Asian cousins.
- African elephants have two opposing 'lips' at the end of their nose. They can be used like fingers to grab small items. This is one of the key differences to Asian elephants who have one 'finger'.
- African elephants are threatened by the ivory trade. Many have been killed for their tusks.
- The elephants travel great distances in search of food. They are herbivores and some need to consume more than 100 kilograms of food every day.
- Infant elephants are dependent on their mother's milk for the first two years of their life.