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The killers littered a South African beach with dozens of bodies and disappeared, leaving a massacre and a mystery.

The culprits were unknown until experts looked more closely at the 63 African penguins found dead Friday morning on Boulders Beach near Cape Town and discovered stingers in their eyes. A veterinarian returned to the scene and spotted what had escaped investigators’ eyes on first pass: many more bodies scattered on the ground – tinier ones this time.

Honey bees. The whodunit had been solved.

The penguins died suddenly between Thursday afternoon and Friday morning, according to a statement from South African National Parks. Veterinarians, penguin experts and government officials rallied to investigate but found no obvious injuries. The bodies of the African penguins, so named because they’re the only species of penguin that breeds on that continent, were taken to the Southern African Foundation for the Conservation of Coastal Birds for post-mortem examination.

The penguins had been stung around their eyes and flippers, areas not protected by feathers, Alison Kock, a marine biologist with the national parks agency, wrote in a statement to The Washington Post.

“The feathers over the penguin’s body are densely packed and it’s unlikely the bees stings could have penetrated through these feathers,” she wrote. “On the other hand, the skin around the eyes and flippers have no feathers and the stings could penetrate in those regions.”

Katta Ludynia, the conservation foundation’s research manager, said they’ve never seen bees attack the African penguin, or any penguins for that matter. She characterized the massacre as unprecedented and “bad luck for the penguins.”

“This is a complete freak accident,” she told NBC News Now.

As part of the post-mortem exams, scientists took samples from the penguins’ bodies for disease and toxicology testing. They were still being tested on Saturday when the national parks agency sent out a news release, but officials say they believe the bees’ nest was disturbed, causing “a mass of bees to flee the nest, swarm and they became defensive and aggressive,” Kock wrote in her email.

“Unfortunately the bees encountered a group of penguins on their flight path,” she added.

Rangers with the conservation foundation will monitor the penguin colony, keeping an eye out for any nests the dead birds may have had. If they find any, the rangers will rescue the eggs or chicks so they can be hand-raised, something the foundation does routinely.

David Roberts, the conservation foundation’s clinical veterinarian, told The Post test results have confirmed the penguins did not have avian influenza, which was “the biggest worry.”

“So we’re happy there,” Roberts said, adding that a bees’ nest was found near where the penguins died and government officials have moved it.

The African penguin is endangered, threatened by oil and gas drilling, mining, hunting, fishing and industrial pollution, according to the International Union for Conservation of Nature’s Red List of Threatened Species. There are about 41,700 mature adults, and their numbers continue to dwindle.

The population has plummeted by 60 percent in the last 28 years, according to the San Diego Zoo. In South Africa alone, according to the conservation foundation, the number of breeding adults has dropped from about 80,000 to 20,600 in the last 20 years.

“The African penguin population is rapidly declining, and it is very sad to see the deaths of so many healthy, most likely breeding adults,” Roberts told NBC News.

“This unusual event is part of what can happen in a normal balanced ecosystem and if the penguins were not in such trouble already, it wouldn’t be such a tragedy.”

The colony at Boulders Beach, which sits on False Bay near one of the most southern points of Africa, has been a refuge for the endangered bird. Two breeding pairs in 1982 have grown in the decades since to as many as 2,200, making the Boulders colony world famous. It’s one of the few places people can see the African penguin up close.

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