An outspoken Peruvian opponent of illegal logging and three other native Ashaninka community leaders were shot and killed in the remote region bordering Brazil where they live, villagers and authorities said on Monday.

The activist, Edwin Chota, had received frequent death threats from illegal loggers, who he had tried for years to expel from the lands for which his community was seeking title.

Illegal loggers were suspected in the killings, said Ashaninka regional leader Reyder Sebastián. Pervasive corruption lets the loggers operate with impunity, stripping the Amazon region's river basins of prized hardwoods, especially mahogany and tropical cedar.

"He threatened to upset the status quo," said David Salisbury, a professor at the University of Richmond who was advising Chota on the title quest and had known him for a decade. "The illegal loggers are on record for wanting Edwin dead."

Chota's region is home to about 80 percent of illegal logging in Peru, which thrives on a web of corruption involving the widespread issuance of counterfeit logging permits.

The wood from a single old-growth mahogany tree can fetch more than $11,000 on the U.S. lumber market, the Environmental Investigation Agency nonprofit said in a 2012 report on Peru's troubled forest concession system.

Chota and the others were apparently killed on Sept. 1, the day they left Saweto, the village he led on the Upper Tamaya river, to hike to a Brazilian Ashaninka community, said the village schoolteacher, María Elena Paredes.

When the men did not arrive at the Brazilian village, worried comrades who had traveled ahead of them returned and found the bodies — apparently killed by shotgun blasts — near some shacks on the Putaya river, Paredes said.

She said by phone that vultures had begun to feed on the bodies, which were found a six-hour walk from the village with 45 inhabitants.

Paredes identified the other slain men as Jorge Ríos, who was Chota's deputy, Leoncio Quinticima and Francisco Pinedo.

She said no villagers had seen the killers.

Peru's main indigenous federation, AIDESEP, expressed outrage at police and the judiciary in a statement for "doing absolutely nothing despite repeated complaints" to protect the slain men.

Peru's deputy minister for intercultural affairs, Patricia Balbuena, said authorities planned to fly by helicopter to Saweto to investigate and retrieve the bodies.

The Ashaninkas are Peru's leading Amazon ethnic group and Sebastián says violence against them has been rising since they began agitating for titles to their territories.

The Associated Press