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Vancouver Aquarium sea otters Milo and Tanu gained international fame after being filmed holding paws.

A sea otter who grew to be adored worldwide when he was filmed holding paws with his companion at the Vancouver Aquarium has died of cancer.

Milo died Wednesday, and the aquarium says he was the first otter to be diagnosed with lymphoma and also the first to be treated with chemotherapy.

Veterinarians and other specialists say Milo responded positively to treatment for nearly six months and spent much of its time in a habitat with a companion named Tanu.

"Our team, especially those who worked closely with Milo during the past months to provide specialized care while he was receiving treatment, are saddened by his death but take solace in his peaceful departure," said Clint Wright, the aquarium's senior vice-president.

The aquarium says medical care for the otter since he was diagnosed last August will improve the understanding of sea otter diseases and responses to treatment.

An online video showing Milo holding paws with another otter was posted online four years ago, and 16 million people have seen it.

Milo was born in Portugal's Lisbon Aquarium in 1999 and arrived at the Vancouver Aquarium a year later.

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