After making his home in the urban Griffith Park — home of the Hollywood Sign — for the past decade, P-22 became a symbol for California’s endangered mountain lions and their decreasing genetic diversity. The mountain lion’s name comes from being the 22nd puma in a National Park Service study.

The death of the cougar late last year set off a debate between the tribes in the Los Angeles area and wildlife officials over whether scientists could keep samples of the mountain lion’s remains for future testing and research.

Some representatives of the Chumash, Tataviam and Gabrielino (Tongva) peoples argued that samples taken during the necropsy should be buried with the rest of his body in the ancestral lands where he spent his life. Some tribal elders said keeping the specimens for scientific testing would be disrespectful to their traditions. Mountain lions are regarded as relatives and considered teachers in LA’s tribal communities.

Tribal representatives, wildlife officials and others discussed a potential compromise in recent weeks, but a consensus was not reached before P-22 was buried in an unspecified location in the Santa Monica Mountains on Saturday.

“While we have done everything we could to keep the carcass intact, the Tribes and agencies involved are still working toward a conclusion about some of the samples,” the state Department of Fish and Wildlife said in a statement Monday. “What is important to understand is that the Tribes and agencies involved all agreed on moving forward with the burial and it was a moving ceremony. We have come to a better place of understanding and we look forward to continued growth from this place.”

It was not clear whether the unspecified samples might also be buried with the animal in the future or if the tribes have agreed to let scientists keep some specimens for additional testing.

Saturday’s traditional tribal burial included songs, prayers and sage smoke cleansings, according to Alan Salazar, a tribal member of the Fernandeño Tataviam Band of Mission Indians and a descendent of the Chumash tribe.

The Natural History Museum of Los Angeles County, where the cougar’s remains had been kept in a freezer before the burial, called the burial a “historically significant ceremony.”

“The death of P-22 has affected all of us and he will forever be a revered icon and ambassador for wildlife conservation,” the museum said in a statement Monday.

Salazar, who attended the ceremony, said he believes P-22’s legacy will help wildlife officials and scientists realize the importance of being respectful to animals going forward.

Beth Pratt, the California executive director for the National Wildlife Federation who also attended the ceremony, wrote on Facebook that the burial “ helped me achieve some measure of peace” as she grieves the animal’s death.

“I can also imagine P-22 at peace now, with such a powerful and caring send-off to the next place,” she wrote. “As we laid him to rest, a red-tailed hawk flew overhead and called loudly, perhaps there to help him on his journey.”

Los Angeles and Mumbai are the world’s only major cities where large cats have been a regular presence for years — mountain lions in one, leopards in the other — though pumas began roaming the streets of Santiago, Chile, during pandemic lockdowns.

Wildlife officials believe P-22 was born about 12 years ago in the western Santa Monica Mountains but left because of his father’s aggression and his own struggle to find a mate amid a dwindling population. That drove the cougar to cross two heavily traveled freeways and migrate east to Griffith Park, where a wildlife biologist captured him on a trail camera in 2012.

His journey over the freeways inspired a wildlife crossing over a Los Angeles-area highway that will allow big cats and other animals safe passage between the mountains and wildlands to the north. The bridge broke ground in April.

P-22 was captured last December in a residential backyard following dog attacks. Examinations revealed a skull fracture — the result of being hit by a car — and chronic illnesses including a skin infection and diseases of the kidneys and liver. The city’s cherished big cat was euthanized five days later.

Los Angeles celebrated his life last month at the Greek Theater in Griffith Park in a star-studded memorial that featured musical performances, tribal blessings, speeches about the importance of P-22’s life and wildlife conservation, and a video message from Gov. Gavin Newsom.

To honor the place where the animal made his home among the city’s urban sprawl, a boulder from Griffith Park was brought to the gravesite in the Santa Monica Mountains and placed near P-22’s grave, Salazar said.