LOS ANGELES (AP) — P-22, the celebrated mountain lion that took up residence in the middle of Los Angeles and became a symbol of urban pressures on wildlife, was euthanized Saturday after dangerous changes in his behavior led to examinations that revealed worsening health and injuries likely caused by a car.

Officials with the California Department of Fish and Wildlife said the decision to euthanize the beloved big cat was made after veterinarians determined it had a skull fracture and chronic illnesses including a skin infection and diseases of the kidneys and liver.

“His prognosis was deemed poor,” said the agency’s director, Chuck Bonham, who fought back tears during a news conference announcing the cougar’s death. “This really hurts … it’s been an incredibly difficult several days.”

The animal became the face of the campaign to build a wildlife crossing over a Los Angeles-area freeway to give mountain lions, bobcats, coyotes, deer and other animals a safe path between the nearby Santa Monica Mountains and wildlands to the north.

Seth Riley, wildlife branch chief with the National Park Service, called P-22 “an ambassador for his species,” with the bridge a symbol of his lasting legacy.

State and federal wildlife officials announced earlier this month that they were concerned that P-22 “may be exhibiting signs of distress” due in part to aging, noting the animal needed to be studied to determine what steps to take.

The mountain lion was captured in a residential backyard in LA’s trendy Los Feliz neighborhood on Dec. 12, a month after killing a Chihuahua on a dogwalker’s leash. An anonymous report that indicated P-22 may have been struck by a vehicle was confirmed by a scan that revealed injuries to his head and torso, wildlife officials said.

State authorities determined that the only likely options were euthanasia or confinement in an animal sanctuary — a difficult prospect for a wild lion.

P-22 was believed to be 12 years old, longer-lived than most wild male mountain lions.

His name was his number in a National Park Service study of the challenges the wide-roaming big cats face in habitat fragmented by urban sprawl and hemmed in by massive freeways that are not only dangerous to cross but are also barriers to the local population’s genetic diversity.

The cougar was regularly recorded on security cameras strolling through residential areas near his home in Griffith Park, an oasis of hiking trails and picnic areas in the middle of the city.

P-22 was born in the western Santa Monica Mountains, and he was partly famous for apparently crossing two heavily traveled freeways on his trek east to the 4,200-acre (1,700-hectare) park in the urbanized eastern end of the mountain range.

“P-22’s survival on an island of wilderness in the heart of Los Angeles captivated people around the world and revitalized efforts to protect our diverse native species and ecosystems,” Governor Gavin Newsom said in a statement Saturday.

Ground was broken this year on the wildlife crossing, which will stretch 200 feet (61 meters) over U.S. 101. Construction is expected to be completed by early 2025.

P-22 was outfitted with a tracking collar in 2012. A year later his celebrity was solidified when he appeared in a National Geographic feature with an iconic photo of the big cat on an LA hillside with the Hollywood sign in the background.

In 2017, the cougar became the star of a permanent exhibit at the Natural History Museum called “The Story of P-22, LA’s Most Famous Feline” that documents the lion’s life and times in Southern California. The exhibit will be upgraded next year to pay tribute to the animal’s legacy, officials said Saturday.

“Even in his death, P-22 continues to inspire LA to embrace urban wildlife conservation and the nature that surrounds us,” said Miguel Ordeñana, the museum’s Senior Manager of Community Science.

P-22 usually hunted deer and coyotes, but in November the National Park Service confirmed that the cougar had attacked and killed a Chihuahua mix that was being walked in the narrow streets of the Hollywood Hills.

The cougar also is suspected of attacking another Chihuahua in the Silver Lake neighborhood this month.

Beth Pratt, California Regional Executive Director of the National Wildlife Federation, said she hopes P-22’s life and death will inspire the construction of more wildlife crossings in California and across the nation. The nonprofit was a major advocate for the LA-area bridge.

“He changed the way we look at LA. And his influencer status extended around the world, as he inspired millions of people to see wildlife as their neighbors,” Pratt said.