For decades, the wolves of the storied East Fork pack were beloved by researchers and tourists alike at Alaska's Denali National Park. They frequented the park's entrance and roads and became the stars of hundreds of thousands of family vacation photos.
Since the 1930s, scientists have documented every detail of the pack's lives: their hunting ranges, mating rituals, even the content of their droppings. They traced family lineage through dozens of generations, giving individual wolves names like "The Dandy," "Grandpa" and "Robber Mask."
Now the researchers must record one final detail in the wolves' long history: They may all be dead.
The last radio-collared male was found shot dead near a hunting camp in May. Now, park officials can't find the last three pack members: a mother wolf without a collar and her two pups.
It's impossible to know for sure what happened to them, officials said, but it's unlikely that the mother and her pups will survive without the support and protection of a pack. The family's den is empty and overgrown with weeds. Porcupines have taken it over since June 28, when the group was last seen.
The wolf pack is the most recent fatality of a controversial Alaska policy that allows hunters to kill wolves and other large predators in the state's national wildlife refuges, wildlife advocates say.
Park officials estimated 49 wolves lived in Denali National Park this spring, only three more than the park's all-time low of 46 in 1986 and a significant decline from the early 2000s when it was common to count more than 100. In 2015, only 5 per cent of Denali visitors reported seeing a wolf - down from 45 per cent in 2010.
The East Fork pack's decline was fast and drastic. In 2013, the nine-member East Fork pack was one of the largest of the nine monitored groups. By the fall of 2014 the pack's numbers had grown to 17, according to Park Service data. Then, the numbers steadily dropped.
The causes of their deaths vary. Many are shot and killed (legally and illegally) by hunters. One died of blood loss after becoming trapped in a snare. Some become untraceable and others die of natural causes.
But one pattern emerges: About 75 per cent of deaths in the East Fork pack in the past year were caused by human trapping and hunting, park biologist Bridget Borg told Alaska Public Media.
By May, only the mother wolf and her two cubs remained. Now, they are gone as well.
In a July report that details pack numbers, park officials wrote "it is unfortunate to lose track of this long-tenured and well-followed pack," though they do note that the pack's lineage would continue in the members of other packs who have bred with the East Fork wolves.
Two other park packs, the Savage Pack and the Headquarters Pack, were previously destroyed by hunting and trapping, according to Wolf Song of Alaska, an organisation dedicated to preserving wolves.
These lands are your lands. They are not game farms managed for a slice of their diversity for the benefit of a few people who would call themselves hunters.
The more than 70 years of continuous study make the East Fork pack one of the longest-observed large mammal families, perhaps only rivaled by Jane Goodall's chimpanzees.
Observation of the pack began in 1939 when National Park Service biologist Adolph Murie began tracking the wolves, following them on foot for more than two years - through buggy summers and hair-freezing winters - as they traversed the park's 7800 square km. In 1944, he published a book, The Wolves of Mount McKinley, detailing his observations.
Things were better for the wolves then, it seems. Murie wrote "but as yet man's activities have probably not altered conditions sufficiently to seriously change the (wolves') natural relationships."
But there might be hope for the remaining Denali wolves.
Last week, the US Fish and Wildlife Service banned the hunting of predators in Alaska's 16 wildlife refuges unless needed "in response to a conservation concern." The change was a challenge to a continuous push by the Alaska Board of Game to loosen the regulation of predator hunting, which the board calls "intensive predator management".
Over the past few years, the board has approved a variety of controversial hunting methods, including targeting bears and wolves from planes and shooting wolves and their pups in their dens. In 2010, it eliminated a "buffer zone" that banned wolf hunting just outside of Denali's borders, near the East Fork's historic range. The zone was an effort to protect park wolves who wander outside of its boundaries. The last East Fork male was found dead in an area that would have been protected by the buffer.
"There comes a time when the US Fish & Wildlife Service must stand up for the authorities and principles that underpin our work and say 'no,' " the wildlife service's director, Dan Ashe, said about the new restrictions in a blog post published by the Huffington Post.
The state government "strongly opposes" the new rules, arguing that it is federal overreach into one of the state's most lucrative industries and shrinks the moose and caribou populations that Native American groups rely on for food, the Guardian reports. Guided hunting generated a total of $78 million in economic activity and more than 2,210 jobs in 2012, according to a study commissioned by the Alaska Professional Hunters Association.
"Ultimately, the new regulations would have significant impacts on Alaskans, particularly those living a subsistence way of life," Bruce Dale, director of the division of wildlife conservation, told the Guardian.
But dozens of scientists disputed Dale's claim that a larger wolf population would greatly affect the number of moose and caribou. The new rules are neither anti-hunter nor anti-native, Ashe said, but are meant to create sustainable hunting methods for future generations - hunters and park visitors alike.
"These lands are your lands," he wrote. "They are not game farms managed for a slice of their diversity for the benefit of a few people who would call themselves hunters."