At Thailand’s infamous Tiger Temple, paying tourists could pet and pose for selfies with the dozens of big cats that called the attraction home. They could walk tigers on leashes and bottle-feed cubs. But the Buddhist monastery turned tourist magnet had for years faced allegations of abuse and, in 2016, a raid by Thai authorities uncovered ghastly sights, including 40 frozen tiger cubs shoved into a refrigerator and a monk attempting to flee with 1,600 tiger parts.

The government soon removed 147 tigers from the compound in the West Thailand town of Kanchanaburi, taking them to two state-run facilities. But in a tragic update on the case, Thai media reported Friday that 86 of the rescued animals have died. A government official attributed the animals’ deaths to a viral disease, saying their immune systems had been compromised by inbreeding.

“There were six tigers [originally at the temple] and later that became 147 or even more, so there were always the risks and also we found that their health were not good to begin with,” Pattarapol Maneeorn, a wildlife veterinarian for the Department of National Parks, Wildlife and Plant Conservation, said during a Monday news conference, according to Reuters. “Their genetics have made their body weak and susceptible to the risk of infection.”

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The revelation came as a blow to conservationists who had long advocated the closure of the Tiger Temple. Opened in 1994 as a forest monastery, the temple took in its first tiger cub five years later, according to the history recounted on its website. Over the years came more cubs, the temple claimed, many after their mothers had been killed by poachers. Soon the tigers began reproducing, and their numbers climbed. Eventually, the temple opened to visitors, charging them to see the tigers.

Although the monks insisted their “hands on approach” resulted in “happy tigers,” critics said the attraction was a front for the illegal tiger trade. It was among the hundreds of so-called “tiger farms” believed to be spread across China, Thailand, Laos and Vietnam. The World Wildlife Fund, which views facilities that breed tigers for commercial purposes as a threat to conservation, was monitoring the temple and similar operations, said Ginette Hemley, the organization’s senior vice president for wildlife conservation.

“To call it a temple absolutely gave the impression that this was a place that was good for tigers,” she told The Washington Post. “And I think all of us were surprised, although we heard rumors, at the scale of the issue.”

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The monastery first ran into controversy in 2001, when authorities discovered it was keeping tigers without a permit. The controversy continued over the years, with activists alleging the facility was exploiting the beloved endangered animal. Numbering about 100,000 a century ago, the tiger population today is estimated at 3,900 worldwide, said John Goodrich, chief scientist and tiger program senior director for Panthera, a nonprofit that focuses on the conservation of wild cat species. They live in some of the most densely populated parts of Asia, he said, and while some South Asian countries are starting to see a rebound, “right in the heart of where these tiger facilities are, we still are facing a crisis.”

For years, the Tiger Temple continued attracting a steady stream of visitors, despite being accused of mistreatment. As tourists clamored for a chance to get up close with the majestic animals, paying between $17 and $140 apiece, ticket sales brought in $5.7 million a year, the New York Times reported in 2016.

Then, in 2016, an Australian nonprofit called Cee4Life (Conservation and Environmental Education for Life) published a report alleging the monastery was involved in commercial breeding and illegal trading. Tiger parts are coveted in Asia for use in traditional medicine, and experts say the illegal industry poses one of the greatest threats to the species.

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In a National Geographic exposé, the nonprofit’s co-founder, Australian wildlife management expert Sybelle Foxcroft, described newborn cubs being torn from their mothers to be cuddled by paying tourists. She also put together a list of 281 tigers that had been born at the Tiger Temple between 1999 and 2015, telling the magazine that the number no longer at the property — 134 — was “too great to be accounted for by deaths alone.” The temple denied any wrongdoing, writing on its Facebook page that volunteers may have “jumped to conclusions.”

Within six months, after fighting the temple in court, the government had raided the temple and removed all of its tigers. Three monks were arrested for trying to smuggle tiger parts off the property, the New York Times reported. The facility has been closed to the public ever since.

After learning that 86 of the confiscated tigers had died, the temple’s caretaker, Athithat Srimanee, singled out the government as responsible, claiming to Reuters that the government kept the animals locked in small cages.

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“When they raided the temple three years ago, they did not say anything about infection, so this is just a blame game,” he said.

Edwin Wiek, founder of the Wildlife Friends Foundation Thailand, told the New York Times that he had urged the Department of National Parks to take precautionary measures to keep the tigers healthy. He said he believed that through steps such as keeping a distance between cages, the deaths might have been prevented.

“It is a very sad story,” he said. “I warned them about it at that time. It was avoidable, but they wouldn’t listen.”

But other conservationists place the blame squarely on the Tiger Temple, with Hemley calling it “more evidence of the horrific downside of keeping large numbers of tigers in captive conditions.” Goodrich said that while the deaths are “very sad and may or may not have been avoidable,” the temple is “on shaky ground” to cast blame.

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Foxcroft, too, said the fault lies only with the monastery, where she said she had noticed illness among the tigers. News of the tigers’ fate, she wrote in a statement on her nonprofit’s website, “is devastating, but it comes as no surprise to me.”

“On hearing about the 86 deaths, it was like being smashed in the chest by a sledgehammer,” Foxcroft wrote. “But, I also know that if the Tiger Temple had continued, and the tigers were not confiscated, they would have still died of the same [illnesses], but the difference would be that the Tiger Temple would have skinned the dead bodies, and used the body parts for sales.”

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