She had been searching for a mate, or another pack, officials said, and had wandered at least 8,712 miles in her hunt. It’s not yet known how the animal died, but state officials say they are investigating. The shooting death of another collared wolf — OR-59 — is unsolved, and the U.S. Fish and Wildlife Service announced a $2,500 reward in January for information in that case.

Researchers say OR-54 was 3 or 4 years old when she died. Her tagging in 2017 was a happy find. “At long last!” the Oregon Fish and Wildlife Office wrote on Facebook.

State and federal wildlife officials had long been trying to collar a wolf belonging to Oregon’s “Rogue Pack.” The tracking collar on the famous OR-7, the founding member of that pack, had stopped working years prior, and officials had not been able to place a new tracking device on any of the protected animals in the area for several years, the Oregonian reported at the time OR-54 was collared.

Then they snagged OR-54, the 54th wolf collared in that state, and OR-7′s daughter.

The Oregon-born OR-7 was famously the first wild gray wolf documented in California since the species was widely eradicated in the 20th century. His presence helped push California’s Fish and Game Commission to vote in 2014 to establish state protections for gray wolves under the California Endangered Species Act. He traveled widely before returning to Oregon, finding a mate and starting the Rogue Pack.

In 2018, OR-54 followed her father’s pawsteps to the Golden State.

She crossed from Oregon into California on Jan. 24, 2018, and mostly stayed, first traversing eastern Siskiyou County before walking through Butte, Lassen, Modoc, Nevada, Plumas, Shasta, Sierra and Tehama counties in search of a mate, according to the Sacramento Bee. She twice returned to Oregon. She briefly sneaked into Nevada. She traveled an average of 13 miles a day — a lengthy journey that seemed to cover a lot of the same ground her father crossed years before.

Amaroq Weiss, a wolf advocate at the Center for Biological Diversity, said in a news release that like her father, “OR-54 was a beacon of hope who showed that wolves can return and flourish here.”

She told The Washington Post the wolf’s death sets back the “trajectory we hoped for for wolf recovery.”

“Being a wolf in the wild is very fragile,” Weiss said. “It’s surprising to most. But a lone wolf usually doesn’t live past 4 or 5 years.”

She said lone wolves may be killed by wolf packs, fatally kicked by the elk they’re chasing or killed by humans.

Except for a few run-ins, OR-54′s journey appeared mostly solitary.

“I think the fact that she traveled so far, if nothing else, is an indication that we don’t have a lot of lone wolves for her to have met up with,” Weiss said.

Fewer than a dozen known wolves live in California. In July, tracking devices found that OR-54 crept toward the territory of the only known wolf pack in the state, according to the Sacramento Bee. But she didn’t stay for long. She was also suspected in at least five livestock attacks.

Wolves have long been despised by ranchers and farmers, an industry whose influence wildlife advocates in part blame for reluctance at the state and federal level to develop recovery plans for the controversial predators.

Last March, the Trump administration proposed stripping federal protections for gray wolves in the Lower 48 states, saying the species had successfully recovered. Still, some states are taking wolf recovery efforts into their own hands: In Colorado, for example, voters will decide via a ballot measure in November whether to reintroduce the endangered species.

Weiss said the loss of numerous wolves underlines “how important it is that we have protections in place for wolves.”

“We’d never have wolves coming back to California, coming to Oregon, if they hadn’t been listed for federal protections,” she said.

She added: “We hope every wolf has a long, flourishing life in which they can live out their own intrinsic existence, and at the same time contribute to a successful recovery of this species.”

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