Luke, a lion, has died at the National Zoo in D.C.

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Luke the lion, known for being both gentle and regal, was put down at the National Zoo in Washington on Wednesday.

“Luke was truly the ‘king’ of the big cat exhibit,” said Craig Saffoe, curator of big cats at the Smithsonian’s National Zoo and Conservation Biology Institute in northwest Washington.

Luke was 17 years old, which is the median life expectancy for African lions in captivity. Lesions that may have been cancerous were found in his liver.

Saffoe said Luke is “gentle with his buddies” and “extremely patient,” adding that Luke was also a protective father to the 13 boys he fathered.

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Born in 2005 on a private reserve in South Africa, Luke was sent to the DC Zoo in 2006 after expert breeding advice. Zoo officials said in a statement that Luke was the “most genetically valuable” lion in the North American population at the time because his genes were not represented in US zoos.

He sired all of his surviving cubs with Naba and Shera in four different litters from 2010 to 2014. Some of the cubs went to other zoos. Shera and three of her offspring stayed at the DC Zoo.

In recent years, Luke has had some health issues.

He has had some discomfort and lameness in his right front leg since 2016, zoo officials said. At one point, he didn’t put his full weight — 422 pounds — on his leg. A CT scan showed he had a lesion on his spine.

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However, experts decided against surgery due to concerns about complications. Vets treated him with steroids and anti-inflammatory drugs. They also performed deep tissue laser therapy and acupuncture to try to treat his lameness.

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Last week, his carers noticed that Luke had lost nearly 18 pounds. They drew blood but found nothing of concern. Earlier this week, Luke lost his appetite and his hind legs appeared to have lost muscle mass, experts said. After an examination, the zoo’s veterinarians found multiple cystic masses in his liver and a “moderate to severe progression of his spinal disease” in which several of his cervical vertebrae had fused, zoo officials said.

“Given these results, Luke’s overall long-term quality of life was considered poor and the animal care team decided to humanely euthanize him,” zoo officials said in the statement.

A final pathology report will be prepared in the coming weeks and will likely provide more details on his condition, officials said.

Saffoe said Luke’s health issues caused arthritis in his joints and spine, and that he basically had a “cat’s version of osteoporosis.”

“He was limping for a long time because of a spinal injury,” he said, “and all of this caused him to have an awkward gait.”

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Saffoe said Luke is a unique Leo and known for being “very clear with his mood.” Laughing, Saffoe recalled that Luke didn’t like him, but he did like his female handlers.

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“I was one of the few men he saw consistently, and I think because I’m a man who produces testosterone, it’s a distinctive smell that a lion can pick up on,” Saffoe said. “I think he picked my scent out and he saw me as a raid on his lion pride.”

Luke would see Saffoe and “jump and bark,” so Saffoe kept his distance. But when female zoo keepers sat in front of Luke’s enclosure and cooed at him, “he loved it,” Saffoe said.

“He was very expressive,” Saffoe said. “He was a very open cat about his feelings for you.”

With other cats, Luke is very gentle, Saffoe said. Sometimes when the two female lions “beat him up…even though he was bigger, he backed off,” Saffoe said.

“He was very gentle,” Saffoe added, but he could also be playful. Saffoe recalled how keepers would sometimes see Luke playing in his enclosure, but when Luke realized he was being watched, he stopped, looked up, and had an expression on his face that seemed to say, “‘You didn’t just catch me playing ,'” Saffoe said. ” ‘I’m too royal.’ ”

Luke’s favorite toy was a log, which zookeepers said he enjoyed rolling and chewing on with his legs.

In the wild, lion populations have declined about 30 percent due to disease, hunting and habitat loss, zoo officials said. They are considered an “endangered species” by the International Union for Conservation of Nature, and officials said there are an estimated 20,000 lions in the wild.

Luke’s death comes almost a month after a female African lion named Nababiep – briefly known as Naba – was euthanized. At 18, she had kidney disease, dental problems, and a mass in the intestines. She had been at the DC Zoo for 16 years.

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Speaking of the recent deaths of the two lions at the DC Zoo, Saffoe said, “It’s great when you bring young animals in, but unfortunately they get old too.”

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