LOS ANGELES >> The mountain lion P-22, who lived in the heart of Los Angeles for more than a decade and became the face of an international campaign to save California’s threatened puma population, was “compassionately euthanized” Saturday morning, according to the California Department of Fish & Wildlife’s director, because of the serious injuries he suffered earlier this week.

P-22 was thought to be about 12 years old.

Wildlife biologists with the National Park Service and the state’s wildlife department captured the mountain lion in December after he began to exhibit increasing “signs of distress,” including three attacks on dogs in a month and several near-miss encounters with people walking in Los Feliz and Silver Lake. Scientists tracked the big cat to a Los Feliz back yard, where he may have been laying low after being hit by a car.

A series of health exams showed that P-22 was significantly underweight, with a thinning coat and damage to his right eye, possibly from being hit by a car. A local animal control department had received a call reporting a vehicle collision with a mountain lion, and P-22’s radio collar placed him near the intersection where the crash was reported, officials said.

The mountain lion was not healthy enough to be released back to Griffith Park, state wildlife officials said. Advocates, scientists and residents had harbored hopes that the beloved animal would be healthy enough to retire to a nature preserve.

P-22 surprised the world in 2012 when his fluffy hindquarters and black-tipped tail appeared on a photograph snapped by a motion-sensing camera in Griffith Park. The adolescent cat had made an improbable trek to Griffith Park from his likely birthplace in the Santa Monica Mountains, journeying through the Hollywood Hills and across the 405 and 101 freeways.

P-22 was first introduced to the world in a Los Angeles Times story. The big cat soon became a bona fide celebrity, appearing in a glossy National Geographic feature that showed the mountain lion prowling past the Hollywood sign at night, muscles rippling under his tawny fur.

Scientists assumed the apex predator would move on in search of a mate and more space to roam. Instead, the wayward cat stayed in Los Feliz for more than 10 years, feasting on mule deer and raccoons and occasionally appearing on video doorbell cameras on quiet streets near the park. The cat lived alone and, as far as scientists can tell, never mated.

Catching a glimpse of P-22 on a nighttime prowl became one of the most coveted celebrity sightings in Los Angeles.

Like many cougars, which are sometimes called “ghost cats,” P-22 was shy by nature. For years, he preferred the park’s dark canyons and hillsides — and, sometimes, a darkened city sidewalk — to populated areas. But he had recently started to venture deeper into L.A., wandering as far south as Silver Lake and staying in residential areas for longer stretches of time.

Those forays coincided with an increase in run-ins with humans, including attacking three dogs in the span of a month, and chasing a man and his dog back up a set of steps and into their house in Silver Lake, wildlife officials said.

P-22’s discovery in the park in 2012 led to one of the most unusual elements of his life: the city taking his side, instead of demanding that he be removed. Big cats prowl large swaths of the United States, but few cities would allow a cougar to live in their midst, let alone stay for a decade.

Many Angelenos saw themselves in P-22, an aging bachelor who adjusted to a too-small space in the big city, waiting for a mate who might never arrive. Others identified with his story, crossing borders and freeways in search of a place he could call home.

“Crossing the border, being persecuted in some areas of the country — people feel a connection with that,” said Miguel Ordeñana, the scientist who first discovered P-22, in a 2022 interview.

P-22 went on to achieve the kind of lasting fame of which most Angelenos can only dream. His photogenic face, including dark markings around the eyes that resembled eyeliner, appeared in a documentary and an exhibit at Los Angeles County’s Natural History Museum. The big cat was featured on socks, tattoos and bumper stickers. And by order of the City Council, every Oct. 22 was celebrated as “P-22 Day.”

P-22’s presence in Griffith Park was a reminder that Los Angeles is far wilder than it appears, with one of the highest levels of biological diversity of any big city in North America. The big cat’s isolation in the park, surrounded by freeways, helped him become the poster cat for the conservation campaign called “Save L.A. Cougars.”

The 101 Freeway forms an almost impenetrable barrier for the threatened puma population in the Santa Monica Mountains, cutting them off from a wider gene pool to the north. That has led to inbreeding that has caused genetic abnormalities and could lead to infertility.

Recent scientific modeling has drawn a dire conclusion: Without intervention, pumas in the Santa Monica and Santa Ana mountains could be extinct within 50 years.

California nature activists spent more than a decade raising $77 million in private donations and state funding for a wildlife bridge across a 10-lane stretch of the 101 in Agoura Hills, which they hope will widen the breed’s gene pool. The plight of P-22, alone in a tiny territory hemmed in by freeways, drew support from around the world, including from Leonardo DiCaprio’s charitable foundation.

The state broke ground on the wildlife bridge in April. Its presence, advocates say, may be P-22’s most lasting contribution.

“His story of being isolated and trapped is what really got people to realize why a crossing like that was needed, more than any scientific paper could,” said Beth Pratt, a regional executive director in California for the National Wildlife Federation, who called herself P-22’s agent. “He changed the world for his kind.”

Pumas are so retiring that they’re sometimes called “ghost cats,” and P-22 often went days without being spotted. But there were high-profile high jinks, too.

About two years after arriving in the park, P-22 appeared on trail-camera footage looking gaunt, his tail as thin as a pipe cleaner. The NPS trapped and treated him with topical medications and vitamin K injections, then released him.

Tests later confirmed that P-22 had been exposed to rat poison and was suffering from mange, a parasitic mite. The photo released of the big cat while sick went viral, showing the once-handsome face bedraggled, the eyes sagging.

The image helped spur action in the California Legislature and ultimately led to a 2020 law temporarily banning some kinds of rat poison.

A few months later, a contractor for a home-security company found the cat lounging in a crawl space beneath a house in the hills of Los Feliz. Soon, helicopters were hovering above the street, covering the incident like an FBI raid. One local news station added a caption that screamed: “BREAKING NEWS: P-22 TRAPPED INSIDE HOME.”

When officials finally cleared the area, P-22 slipped back, unseen, to Griffith Park.

In 2016, P-22 became the prime suspect in the death of a 14-year-old koala named Killarney, whose mutilated body was found about 400 yards away from her enclosure at the Los Angeles Zoo. The attack was not recorded, but the zoo’s surveillance cameras placed the puma at the scene. Few animals can easily leap over an 8-foot fence topped with barbed wire.

After the attack, a city councilman suggested that P-22 be moved to a new habitat, saying Griffith Park was “ultimately not suitable for him.” But where the puma could go stymied the discussion. Moving P-22 to any mountainous area already occupied by another mountain lion could be a death sentence, because the big cats kill to protect their territory.

The zoo, however, took P-22’s side. The zoo’s director of animal programs later told a Times reporter: “We’re in Griffith Park, and Griffith Park is his home, and we have to respect that. You can’t hold a mountain lion accountable for being a mountain lion.”