A polar bear at SeaWorld San Diego died suddenly after her companion was taken to another zoo for a breeding programme.
Szenja, a 21-year-old female, died on Tuesday. Her keepers had noticed she was not eating properly, but her death remains unexplained.
NBC San Diego reports that Szenja made headlines in March when tens of thousands of people signed a petition asking SeaWorld San Diego not to separate her from her "best friend", another female polar bear named Snowflake. The pair had shared an enclosure since the San Diego park opened in 1997.
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However, Snowflake was transported for a breeding visit to Pittsburgh Zoo, where she remains.
Al Garver, SeaWorld San Diego's vice president of zoological operations, said: ”Szenja was a beloved member of our animal family, so this is a very difficult day for all of us.”
Garver added Szenja had inspired people around the world to want to protect polar bears in the wild.
But the bear’s death provoked an angry response from animal rights activists.
Tracy Remain, executive vice-president of People for the Ethical Treatment of Animals (PETA), said Szenja had died of a "broken heart".
"This should be a wake-up call to SeaWorld: stop breeding and shipping animals around, close the animal exhibits, and retire the animals to sanctuaries," she told NBC San Diego.
SeaWorld San Diego shipped Snowflake to Pittsburgh as part of a national programme to boost the polar bear population, which scientists estimate to be between 22,000 and 31,000 worldwide.
SeaWorld has been the target of criticism in recent years, says the San Diego Union Tribune following the release of the 2013 documentary Blackfish, which raised concerns about the theme parks’ practice of holding killer whales in captivity.
