It’s mating season for wolves in Yellowstone, and the alpha male of the Druid Peak pack sits alone on a snowy ridge, howling mournfully. His mate, whose only name was the number 42, is dead. One of Yellowstone’s oldest wolves at eight, 42 was killed by a rival pack the previous night.

She was also known as the "Cinderella wolf," because of her overnight transformation from submissive female to dominant female, and she was featured in two National Geographic TV specials. She was one of the original 31 Canadian wolves transplanted to Yellowstone to kick off the wolf restoration effort in the Northern Rockies. Much of the park’s spectacular wolf recovery can be attributed to her breeding success: At least three of her daughters have gone on to form their own packs. And not only was she the alpha female of the largest wolf pack ever recorded — the Druid pack numbered 37 wolves in 2000 — but she also contributed mightily to our knowledge of wolf behavior and pack dynamics.

Wolf-watchers loved her. Her beautiful charcoal coat and underdog role endeared her to thousands of visitors to Yellowstone. In her celebrity status, she even made it onto a personalized license plate, WOLF 42F. When wolf biologist Rick McIntyre announced her death recently to a small crowd of wolf enthusiasts in the park, sobs broke out.

Her life was nothing if not dramatic. For her first four years in Yellowstone, 42 suffered under the ruthless domination of her sister, 40, the alpha female at the time, and an especially aggressive wolf. Forty had ousted her own mother as leader and then lost no opportunity to beat up on her sisters, 41 and 42, and their offspring.

Forty-one soon tired of the floggings and left the pack, while 42 stuck it out. Perhaps her bond with 21, the alpha male, enabled her to endure her sister’s assaults. Then in 1999, 21 mated with both 40 and 42, the latter going off to den by herself. One day 40 came over and gave her sister a thrashing, perhaps even killing her pups. The next year, after 42 again mated with 21, she chose a den site far from her sister’s. But when her pups were almost weaned, 40 again came to visit.

Doug Smith, the Wolf Project biologist for Yellowstone, says, "None of the other wolves liked 40 so they would hang out with 42 instead. In fact, the only wolf to visit 40's den was 21." When the aggressive 40 threatened her sister again, Smith said, "This time 42 said, ‘forget it’ and attacked 40, defending her pups. At least two other wolves joined in and left 40 a bloody mess."

The next day 42 moved her pups clear across the Lamar Valley, took over 40's den and raised her sister’s pups along with her own. She quickly assumed the alpha role, which she held until her untimely death this winter.

Upon examining her carcass, Doug Smith marveled at her excellent condition for an eight-year-old wolf. "None of her canines were broken. I don’t think I’ve ever seen a wolf older than five that didn’t have canines broken from trying to bring down running prey."

Smith regarded 42 as an excellent hunter, but in her later years she let the younger wolves do much of the initial chasing and hunting, assuming a supervisory role. On one particular hunt, he noticed that she led the pack to a herd of elk, and as the wolves fanned out across the hillside, she went up to each wolf as if whispering instructions. She even seemed to reposition several of the wolves to more strategic locations.

Forty-two was the quintessential mother. She’d put unruly adolescents in their place and she raised her daughters’ young as well as her own. The year she took over as alpha female, the Druid pack successfully reared 20 out of 21 pups born.

In the past few years, 21 and 42 were nearly inseparable. They cut a distinctive picture as they led their offspring through the Lamar Valley — 21, a large, bulky male, part black, part gray, usually trailing the lithe and darker 42.

After 42 died, 21 spent two days off by himself, howling. Seasoned observers reported it was more howling than they had ever heard the wolf do in his entire life.

"Do wolves mourn?" I asked Smith.

"I’ll leave that up to you," he replied.

Greg Gordon is a contributor to Writers on the Range, a service of High Country News (hcn.org). He writes in Gardiner, Montana.