The chronic conditions and need for surgery and long-term medical care, combined with P-22’s age, left the puma with “no hope for a positive outcome,” the California Department of Fish and Wildlife said in a statement Saturday, five days after P-22 was captured for a health assessment.

He was euthanized at 9 a.m. Saturday at the San Diego Zoo Safari Park. The death was announced by state wildlife officials in a tearful Zoom news conference.

“It’s been an incredibly difficult several days,” Charlton H. Bonham, the director of the department, said while crying. “I made the decision that the right thing to do was to bring peace now, rather than have P-22 continue through what would not have been acceptable, from a compassionate level, in my mind.”

The big cat’s life contributed to scientific research and inspired a bridge crossing for wildlife over busy Highway 101 in the Santa Monica Mountains. He was a beloved cultural figure, a symbol of survival and an ambassador for wildlife protection. His death, like his life, spoke to the plight of animals in urban environments.

P-22 was believed to be about 12 years old, having been estimated to be two when biologists first found him a decade ago, according the National Park Service. He had surpassed the life expectancy for wild mountain lions, who generally live up to 10 years. Captive ones can live up to 21 years, according to the National Wildlife Foundation.

He was too unwell to live out the rest of his life in an animal sanctuary, said Beth Pratt, the National Wildlife Federation’s regional executive director in California, who attended a briefing by the puma’s medical team. In a statement, she said keeping him alive with medical intervention would have prolonged his suffering.

“I sat near him, looking into his eyes for a few minutes, and told him he was a good boy,” wrote Pratt, who said goodbye to P-22 before he was euthanized. “I told him how much I loved him. How much the world loved him.”

Famous Los Angeles cougar, P-22, captured for medical exam

P-22’s extraordinary journey began when he crossed two multilane freeways to get from the Santa Monica Mountains to Griffith Park in 2012, where he was found by biologist Miguel Ordeñana, according to the Natural History Museum of Los Angeles County. He then found fame thanks to a National Geographic article on big cats, which included an image by photographer Steve Winter of P-22 prowling beneath the Hollywood sign.

On Saturday, the news of his death prompted an outpouring of posts on social media. More than 200 people had commented on the state’s Facebook announcement of his death, many sharing memories of the feline or saying they were crying.

“P-22 was an icon,” tweeted California Gov. Gavin Newsom (D). “His incredible journey helped inspire a new era of urban conservation, including the world’s largest wildlife crossing in CA.”

The big cat was such a celebrity that his obituary was pre-written by the Los Angeles Times, reporter Laura J. Nelson said on Twitter, a practice reserved for notable figures.

“I won’t rest until P-22 has a bronze statue in Griffith Park and maybe a star on the Hollywood Walk of Fame,” tweeted Laura Friedman (D), a state legislator.

With black markings around his eyes and framing his white muzzle, the tawny, sleek puma was often caught on wildlife cameras mid-prowl. For locals in Los Angeles, seeing P-22 was like a celebrity spotting. He sometimes roamed around town, once even cozying up in a home’s crawl space, and showed up on residents’ doorbell cameras.

“Whenever I hiked to the Hollywood sign, or strolled down a street in Beachwood Canyon to pick up a sandwich at The Oaks, or walked to my car after a concert at the Greek Theater, the wondrous knowledge that I could encounter P-22 always propelled me into a joyous kind of awe,” said Pratt. “We may never see another mountain lion stroll down Sunset Boulevard or surprise customers outside the Los Feliz Trader Joe’s.”

This spring, construction began on the Wallis Annenberg Wildlife Crossing, which Pratt said “would not have been possible” without P-22’s inspiration. Construction on the massive bridge, which will allow wildlife to cross the freeway, is projected to be completed in 2025, according to the Annenberg Foundation.

Because his territory was limited to the park’s island of wilderness, P-22 never found a mate. He existed in the smallest home ever known for an adult male mountain lion, officials have said. He ate deer, coyotes and other prey. When he ventured into the Hollywood Hills, it was mostly late at night.

Tracked by the National Park Service since March 2012, P-22 had become one of the oldest mountain lions in a long-term research study on the animals. He was a “critical part” of the study and one of its most interesting participants, the Santa Monica Mountains National Recreation Area said in a statement.

About 100 mountain lions live in the area. Los Angeles and Mumbai are the only two of the world’s largest cities with big cats.

He leaves behind a scientific legacy, the park said, in addition to his legacy as a living demonstration of the hardships and possibilities for urban wildlife.

“We’re really going to miss following him,” Seth Riley, the National Park Service branch chief for the Santa Monica Mountains, said at the news briefing, choking up. “Already this week, we don’t get to go and look and see his points on the website and see what he’s doing.”

In the Santa Monica Mountains, the long-term survival of a stable mountain lion population is threatened by development, the National Park Service has said. Roads break up habitat, prevent animals from roaming to breed and lead to car collisions.

For P-22, it eventually led to suffering. As he got older, “the challenges associated with living on an island of habitat” seemed to increase, state officials said last week.

He had lived to be “remarkably old” for a cat in the wild, they said. Recent changes in his behavior indicated he might have been in distress.

In March, P-22 left Griffith Park and went farther into an urban area than he ever had, eventually returning to the park, according to the Los Angeles County Natural History Museum. He killed a pet Chihuahua in November, and had been venturing into residential areas. On camera, he looked like he was growing thin and his coat didn’t look healthy, said Deana Clifford, a state wildlife veterinarian.

State wildlife officials began planning to capture P-22 so they could give him a checkup, she said. Then, on Dec. 7, they received an anonymous report that P-22 might have been hit by a car. Biologists found him in a woman’s backyard on Monday, where wildlife officers tranquilized him and took him in for a medical examination.

P-22’s weakened health may have hindered his ability to evade cars, but if such a collision led to his death, that still represented a failure by humans to better protect wildlife, Pratt said. State officials said they would not investigate the car incident.

“This situation is not the fault of P-22, nor of a driver who may have hit him,” the wildlife agency said. “Rather, it is an eventuality that arises from habitat loss and fragmentation, and it underscores the need for thoughtful construction of wildlife crossings and well-planned spaces that provide wild animals room to roam.”

Six veterinarians from the San Diego Zoo and four veterinary specialists were involved in the assessment, the state wildlife department said. They found that the car collision had torn P-22’s diaphragm and pushed his liver into his chest cavity, damaging abdominal organs, said Hendrik Nollens, San Diego Zoo Wildlife Alliance’s wildlife health vice president.

Among P-22’s other ailments were kidney disease with Stage 2 failure, liver disease, a skin infection and arthritis. He had lost at least 20 percent of his weight, Nollens said at the news conference, where he showed a CT scan of P-22’s body. The damage to his internal organs from the likely car collision would have required invasive surgery.

P-22 will have a post-mortem examination, said Clifford. His body will then be given to the county’s Natural History Museum, where it will be received by Ordeñana, the biologist who discovered him.

Bonham, the state wildlife director, said he had gone through moments of hope for the cat all week, followed by successive bad diagnoses. Officials had hoped to place P-22 in a sanctuary if he couldn’t return to the wild, the director indicated.

Bonham gave a tearful apology, saying he hoped people would “move through the pain” and find ways to honor P-22. His 11-year-old son, he said, planned to take a hike at Griffith Park. He urged the public to support making urban environments safer for wildlife.

“The animal brought us together,” he said. “Let’s make a difference, so the rest of the large animals out there have a future that’s brighter.”

Pratt said she had attended a tearful meeting with the veterinarians and state workers who had examined P-22, where they presented their findings and passed boxes of tissues around the room.

“He changed the way we look at L.A. And his influencer status extended around the world, as he inspired millions of people to see wildlife as their neighbors. He made us more human, made us connect more to that wild place in ourselves,” Pratt said. “His legacy to us, and to his kind will never fade.”