The end came in furious fashion. In a clearing high atop Specimen Ridge in Yellowstone National Park, a bloody battle over territory -- and perhaps even revenge in a long-standing rivalry -- ended this weekend in the death of an important matriarch, a wolf who helped write the opening chapter in the park's wolf recovery program.
Wolf No. 42, as she was known, was the only wolf still surviving in Yellowstone who was part of the 28 wolves brought to the park during the reintroduction effort in 1995 and 1996.
Her sister, No. 41, is now the only member left from that original group. She lives outside of the park in the Sunlight Basin north of Cody, Wyo.
''She was truly part of history," said Doug Smith, leader of the Yellowstone Wolf Project. ''We're looking at one of the wolves that had a significant impact on wolves in Yellowstone."
The life and death of No. 42 had elements of a Shakespearean drama: revenge, power struggles, violence, family bonds, rivalries and tests of will and survival.
Ultimately, No. 42, the alpha female of the Druid Peak pack, appears to have been killed by Mollie's pack, a neighboring group of wolves that were once displaced by the Druids and have been poking around the old territory ever since.
''This is a rivalry that goes back to 1996," Smith said.
Mollie's pack, which was known then as the Crystal Creek pack, lost its alpha male in a battle that year when the Druids pushed them out of the Lamar Valley.
Sometime late Saturday or early Sunday, the two packs probably clashed again. Likely, No. 42 and the Druid pack's alpha male, No. 21, were targets in the attack.
''It's not uncommon in wolf warfare ... to go for the alphas," Smith said. ''It's like taking out a general as opposed to a corporal or a private."
On Sunday, No. 42's carcass was spotted on Specimen Ridge by aerial crews. Her radio collar was putting out a ''mortality signal," which is activated when a wolf stops moving.
On Tuesday, Smith and a group of biologists trekked about four miles through the snow to get to No. 42's body. By then, eagles, ravens and coyotes were already feeding.
Park officials removed No. 42's head and left the body to be eaten. Smith said he's hoping No. 42's skull can eventually be displayed for visitors.
''For Yellowstone, which of all national parks in the United States is steeped in history, we had to bring back that skull," Smith said.
No. 42's ability to live to the ripe age of 8 -- several years beyond average -- and remain the alpha female of one of Yellowstone' s most powerful packs might be seen as a testament to both her toughness and her intelligence.
''We find that most wolves are dead by 5 or 6 but there are a few that stick out that are very good survivors," Smith said. Often, ''it's a combination of being tough and sharp."
No. 42 was born in April 1995 in British Columbia, Canada. Earlier that year, 14 wolves from Alberta had been brought to Yellowstone as part of an ambitious and controversial plan to reintroduce gray wolves to the northern Rocky Mountains.
In 1996, 17 wolves were brought to the park from British Columbia, including No. 42, her mother, No. 39, and sisters Nos. 40 and 41.
While they were in a holding pen, the group hooked up with an alpha male, who completed the nucleus for the Druid Peak pack.
Aside from the Lamar Valley skirmishes with the Crystal Creek pack, the Druids coped with internal drama too.
No. 39 left the pack and No. 40, a dominant and aggressive wolf, became the alpha female When No. 39 tried to return to the group a year later, she was not welcomed by No. 40. Later that year, No. 39 and her daughter, No. 41, left the pack for good.
A month later, the pack's alpha male was illegally shot and a new alpha male moved in. Suddenly, the only original members of the Druid pack were the sisters, Nos. 40 and 42.
Things didn't go well.
No. 40 was assertive and had a penchant for reinforcing her dominance through violence. She was seen attacking No. 42 and not relenting until her sister completely submitted, rolling on her back, tucking her tail and licking No. 40's face.
Although the two females bred with the same male, they kept separate dens. Biologists believe that No. 40 may have once raided her sister's den and killed the pups.
The relationship remained tense and finally came to a head in May 2000.
Sometime during the night, No. 40 was attacked. The speculation is that No. 42, and perhaps others in the pack, had had enough of the alpha female' s domineering. The next day, No. 40 was found hurt and bloody in a roadside culvert. She died later that day.
No. 42 soon assumed the role of alpha female, taking up with the head male, No. 21.
Over the years, the pair ruled the Druids as they hunted, bred and defended their territories, often in front of scores of wolf watchers perched on the side of the road. No. 42 took a slightly less violent approach in reigning over the pack.
''She dominated her subordinates but she just didn't flaunt it," Smith said.
These days, there about 170 wolves in Yellowstone. As the population has grown, established packs have had to work to guard the territory from new or growing packs.
For the Druids, that has often meant dealing with Mollie's pack. The two packs have battled periodically over the years and even had a howl-filled standoff across the Lamar River last year.
On Saturday, members of Mollie's pack were seen on Specimen Ridge. They were also seen there on Tuesday.
Smith said it seems likely that the two packs fought again this weekend.
''We're 100 percent sure she got killed by other wolves and I'm 80 to 90 percent sure it was the Mollie's," he said.
No. 42's death sent a buzz through the cadre of wolf watchers who regularly train their spotting scopes on Lamar Valley.
It also appears to be affecting No. 42's longtime mate, No. 21.
Ever since the weekend fight between packs, there have been reports that No. 21 has been periodically keeping his distance from the rest of the Druids. More noticeable, Smith said, is that he's howling.
''I've heard from people that he's acting different and that he's howling and howling and howling," Smith said. ''Wailing his guts out."
But, there are also reports that he's already courting new females. Survival for a wolf pack means overcoming challenges every day -- including the death of a leader.
''This is not going to affect their future as a pack or their ability to defend their territory," Smith said.