Wolf 527 was a survivor. She lived through a rival pack's crippling 12-day siege of her den. When another pair of wolves laid down stakes in her territory, she killed the mother and picked off the pups while the invader's mate howled nearby in frustration and fury.
She was not a charmer. But successful wolves are not known for their geniality. She was large and black and wary -- and cruel when she needed to be. As the alpha female of the Cottonwood Creek pack, she also was equipped with a radio collar so wildlife biologists could track her movements, making her one of Yellowstone National Park's best-known wolves.
Then she ventured outside the park boundaries.
Wolf 527 was killed Oct. 3 by a hunter on Buffalo Plateau north of Yellowstone, less than three weeks into Montana's backcountry elk season. Wolves often stalk elk outside the park and are attracted by entrails the hunters leave behind. But this year, the elk season coincided with the opening of the state's first wolf hunt in modern times.
"She was a genius wolf in her tactics," said Laurie Lyman, a former San Diego County teacher who has spent the last five years tracking the recovery of the endangered gray wolves that were reintroduced into Yellowstone in 1995. "Her strategies were just unbelievable. She knew how to survive anything, but she didn't know how to survive a man with a gun."
Park officials believe four of the Cottonwood pack's 10 wolves -- including 527's mate, the alpha male, and her daughter -- died during those first weeks, in effect ending research into one of the park's most important study groups.
"Whether the pack exists anymore or not, to us the pack is gone," said Doug Smith, the biologist in charge of the Yellowstone reintroduction program that helped bring wolves back from the brink of extinction in the Northern Rockies. Cottonwood "was a key pack on the northern range," he said, giving researchers a window into the existence of animals that had little or no interaction with humans.
State wildlife officials, caught off guard by the ease with which the wolves were cut down, called off the backcountry hunt along a section of Yellowstone's northern boundary for the rest of the year.
But the general wolf hunting season opens today throughout much of the rest of Montana, including other areas bordering the 3,468-square-mile park. Wildlife advocates have sought, so far unsuccessfully, a buffer zone to protect Yellowstone's storied wolf packs.
With more than 1,600 wolves now in Montana, Idaho and Wyoming, state officials are allowing hunters this year to take up to 75 in Montana and 220 in Idaho. Federal protections remain in Wyoming.
"We've got quite a number of other border packs. So people need to decide how hunting's going to occur on the park boundaries," Smith said. "Whose wolves are they? Are they national wolves? Montana wolves? And we have to decide what is the value of our research on wolf populations that are not affected by people."
Conservationists fear that allowing the wolves to be targeted just four months after they were removed from the endangered species list could damage the recovery process. They have argued for delaying a hunt until at least 2,000 wolves have gained a foothold in the region.
Yet residents who have seen livestock and elk plundered say the quota of 12 wolves in a small area immediately north of the national park is too modest to control a predator that under federal protection has broadened its territory and become shockingly adept at killing.
"Those wolves they're talking about in Yellowstone are all over the place out here. They're traveling everywhere," said Ryan Counts, a hunting guide and team-rope rodeo rider from the town of Pray, Mont. He shot Wolf 527.
"They're just decimating our elk herd and everything else. They're bothering cows all the time," Counts said. "Twelve ain't going to do any good at all, you know."
In Dillon, Mont., 180 miles northwest of Yellowstone, a rancher in late August found the carcasses of 122 purebred adult sheep strewn in bloody heaps in his pasture. It was the worst livestock predation in memory -- an example of the ability of wolves to kill for the pure pleasure of it -- and wildlife officials authorized the killing of the entire pack.
Here in the Paradise Valley, which winds through snow-dusted peaks on either side of the Yellowstone River, many blame wolves for the destruction of the northern Yellowstone elk herd, whose numbers are down 60% since the predators were reintroduced to the park from Canada.
Federal biologists say bears, drought and hunters are partly to blame for the decline, but it's hard to find anyone in these small towns who doesn't blame wolves.
"We're starting to see the wildlife just disappear," said Randy Petrich, a rancher and big-game outfitter.
"I had a couple clients out goat hunting. We were back in country that's traditionally just beautiful elk country, and we never even saw an elk track. But it was just full of wolf tracks," he said. "If all these people who are for the wolves only knew what was out here now. It's dead. It's beautiful country, but there's nothing living in it. No deer. No elk. I think it's going to be a brutal winter."
Given the stunning speed with which the small group of wolves reintroduced into Yellowstone managed to expand, few people argued that hunting should not be allowed at some point to control their numbers.
But equally few expected Yellowstone's rock-star wolves to be among the first hit.
Wolf 527 and her daughter, 716, originated from two of the best-known packs in Yellowstone's Lamar Valley, the scene of numerous documentaries for National Geographic, PBS and the BBC. For years, the movements of the Lamar packs have been monitored in careful detail by biologists equipped with radio tracking devices and powerful spotting telescopes.
"They sold this wolf hunt in Montana and Idaho as controlling the predation on cattle and whatnot. Well, these wolves aren't touching cattle. They're feeding on elk. They're doing what they're supposed to do," said Tom Murphy, a wildlife photographer who has been documenting the Yellowstone wolves.
"This is the home ground of all of them, the nursery, the definition of what a healthy ecosystem looks like," he said. "And it drives me crazy that [hunters are] standing on the boundary of the park . . . and killing the ones with radio collars, that people watch every day."
The demise of 716, often known as Dark Female, was reported Sept. 29 in a blog posting from Lyman. Five days later, she followed it up with another item, this time about 527. "It is with a heavy heart that I write yet another obituary for a wolf that was part of our lives for seven years," Lyman said.
Wolf 527 was born in 2002 into what had been Yellowstone's best-known pack, Druid Peak, made famous by the documentary "In the Valley of the Wolves."
Younger wolves often light out on their own, and 527 defected early on to the rival Slough Creek pack, which one of her sisters had joined. She rose to become the second-ranking female and bore pups, including 716.
The pack was ravaged by distemper, as were many of the wolves in the park during 2005. A year later, Slough's long multi-pocketed den was beset by a siege unlike any ever witnessed by park biologists.
For 12 days, 527, 716 (now a yearling) and probably four of the pack's other females and their pups were pinned inside the den by a group of marauding wolves.
Slough Creek's males, apparently sensing a hopeless fight, had abandoned their mates and moved farther into the Lamar Valley, taking up with a young female.
But the mothers inside the den were able to hold the intruders back and finally escape. "They went to find those males," Lyman said. "The next morning when we got up there, there was no cute little thing with them anymore."
The pups were gone -- they had probably either starved or been killed, biologists said.
The reunited pack returned to the den and drove the invaders off. Not long after, 527 left the Sloughs, joined up with a male wolf and formed the new Cottonwood pack -- which included 716. They inhabited an area along Hellroaring Creek, so sparse in elk that other wolves had never been able to stay there long, and they recently had moved up to the open, grassy expanse of the Buffalo Plateau.
In April, they encountered a young wolf couple that had moved in too close for comfort.
"The black male ran away to try to lead [the Cottonwood wolves] away from the den, and then they backed off and he sat there howling," trying to warn his mate, Lyman said.
"Then who shows up but 527? And at that point, the males stepped back, and 527 and 716 went in and killed that female in her den and carried out her pups and ate them.
"I've seen a lot out here," Lyman said. "But I understood why 527 was doing what she did. Because she couldn't afford to have a pack that close to her territory."
Counts, the hunting guide, was on Buffalo Plateau, about 3 1/2 miles outside the park, when he encountered her.
"We just went in there looking for them. It's in the backcountry," he said in a brief interview. "We just figured if they'll let us hunt wolves, we will."
The wolf did not attempt to retreat before he fired, Counts said.
"They aren't showing any fear," he said. "But they will, I'm sure."
Several days before 527 was killed, 716 and another wolf also had approached a group of hunters without fear, said Warren Johnson, a local guide.
"The first [male] wolf saw him, started to come toward him more, and the guy shot him," Johnson said. "He was no more afraid of [the hunter] than the man in the moon."
Johnson and his wife, Susan, shot 716. "I heard it howling. It looked at me, it just watched me coming for a hundred yards, within shooting distance, and I got it," he said.
Since then, Johnson said, he's had another client come in from out of state looking to hunt wolves, with no luck.
"They progressively became harder and harder. Whenever they saw us, they were running. There was no more of that sneaking up on them. . . . They are so much smarter right now," he said.
Wolf 716 is being mounted for display at Johnson's hunting and trail-riding lodge.
"In general, I don't think anybody's out to eliminate the wolves again," said Jim Klyap, owner of Dome Mountain Outfitters in the Paradise Valley, who bought a wolf tag for hunting but didn't kill one.
"People do appreciate them being here. But like any other big-game predator -- mountain lions, bears -- they have to be successfully managed in some way by a quota system. . . . No matter how much we'd like to believe that there's this endless wilderness out there, there isn't. You can only go so far. We're in it now, and it's our job to take care of it."
Montana's wolf program coordinator, Carolyn Sime, said the state might consider imposing additional restrictions around Yellowstone next year.
"It surprised us that the hunters were as successful as they were" in the backcountry, she said, when what officials had hoped was that problem wolves preying on livestock in the valley would be targeted.
"You hear, 'Kill the wolves. . . . They don't belong here. My grandparents killed these wolves for a reason,' " Sime said. But the real danger, she said, is that conservationists' lawsuit to shut down the hunt will succeed, and the wolves will be returned to the endangered species list.
"With that, you risk people's willingness to live with the wolves, and history is replete with what happens to the wolf when people are unwilling to share the landscape with them," she said.
Residents, she predicted, would simply take matters into their own hands.
"They'll kill 'em," she said. "Or they'll put such intense pressure on us to kill them that . . . the program won't survive. It will crumble under the pressure to kill wolves."