Raising Pups Cooperatively
Once again, the wolves of the decades-old Toklat family of Denali National Park are cooperating to raise their new offspring with a level of sophistication that is unusual even in human societies. Toklat, also known as East Fork, and other well established extended family groups of wolves display impressive cooperation during hunting and other activities, but it is as “cooperative breeders” (the proper technical term) that their sociality is most highly developed. For background about this and related behavior particularly in the Toklat family, read the June 17, 2008 and then other summer-fall 2008 entries in the Blog section and the May 2009 report on the Reports2 page.  Most if not all of the 11 Toklat wolves shown in the March 18 and May 1, 2009 blog entries are still present and working together to raise the new (May 2009) pups.
The extreme cooperation that one sees among family members and occasional newcomers in raising the young pups makes it largely meaningless to distinguish between parents and non-parents at this time of the year; they all act like parents. This cooperation includes nursing the pups, which continues until the pups are about a month and a half old. Mothers nurse each others’ pups interchangeably. Non-mothers also lactate, sometimes even one-year-olds despite the fact that wolves do not normally become sexually mature until they are 22 months old.
The Toklat wolves produced 9-10 pups this spring. At least two and probably three females nursed them - the dominant female and two likely daughters: a large, submissive young adult and a 1-2-year-old. The large, submissive female likely produced a second small litter in 2008 (see p.8 of the May 2009 report and photos of her in the September 22, 2008, March 18, 2009, and May 1, 2009 blog entries) and may have produced some of the 2009 pups, although the dominant female could have produced all 9-10. I saw nipples on the third female (directly and in photos) and was fairly certain that a female I observed nursing alongside the dominant female on one occasion was not the second (large) female.
I also observed some interesting related behavior again this year - displays of extra affection by dominant and other family members for the lactating females and by the lactating females for each other. Ranking individuals who would normally assert their dominance seem more inclined to act affectionately toward these females at this time of the year, including with kissing and unmistakeable body language (photo below). There seems to be an understanding of their importance in the early care and provisioning of the pups.  I’ve commented in previous blog entries about the similar way all of the wolves dote over the new pups prior to weaning, behavior that subsequently wanes (and is accompanied by disciplinary action) largely because of the rambunctious food-begging of the rapidly growing pups.                
More than a single lactating female likely ensures more reliable early nourishment for the pups, especially when there are many of them. This could translate into early growth advantages and help compensate for periods of potentially leaner provisioning over the next 2-3 months, after the pups are weaned, when most of what they eat depends directly on the day-to-day foraging success of the older wolves. The presence of more than one lactating female probably also allows the dominant female and others more flexibility to forage away from the den during the first month and a half and thus stay in better condition.
Following are some scenes from my June-early July 2009 ground and aerial observations of the Toklat wolves at their natal den. As usual, the pups were born in early-mid May inside a burrow complex, emerged for the first time about three weeks later, and no longer used the burrows after another week or two. On June 20-23, the adults moved the pups to an area of the den about 400 yards (365 m) away, where heavier forest cover meant I could no longer see them other than by circling in a small airplane, though I could still hear their howling.  [In the Denali region, a “den” is not just a hole in the ground. Well established dens, such as this one, consist of 2-3+ elaborate burrow complexes, beds, trails, and other activity areas spread over 20-30 acres (10+ ha) or more. Some of the Denali sites are easily centuries and probably millennia old and have also been used by early human hunters, as base camps].
Below: Two lactating females and at least eight pups (2 blacks, 6+ browns) can be seen near the end of a typical cooperative nursing session. On the left is the dominant female, likely the mother of most if not all of the pups. A brown pup is exploring atop her back. The other female, a large, submissive young adult, is likely her daughter. Two pups (black and brown, the brown still nursing) are with this female. The other pups have finished nursing from both females and are exploring nearby and at the right side of the photo. June 7, 2009.
Below (2 photos): Later, the two females walk off a short distance, with some of the pups following. The dominant (alpha) female is on the right. Other adults and yearlings are sleeping in the adjacent forest. The pups are 3-4 weeks old in these three photos, 7-8 weeks old in the others. June 7, 2009.
Below: A dominant black yearling male (his coat largely shed) approaches a young lactating female. She shows the usual strong submission, but instead of asserting his dominance he shows affection with tongue-kissing and related behavior and body language. Higher-ranking individuals and others seem to recognize the importance of the lactating females and interact with them accordingly. July 4, 2009.
Below: The yearling male and young female with seven of the pups. Other pups are outside the photo. July 4, 2009.
Above, left, and below: Three of the pups climb, chew, and tug on the young female as a fourth pup approaches to join the fun. She not only tolerates but seems to enjoy their actions. July 4, 2009.
Below: The pups are not shy about approaching their year-old brother, but he is less tolerant of their vigorous play and so they stop short of the liberties they take with the young female. July 4, 2009.
Left: The pups have already seen ravens, eagles, hawks, and other large birds. These two pups are especially curious about the much larger bird circling them now. July 4, 2009.
Below (2 photos): The pups at rest. July 4, 2009.
Jul 23, 2009
Six of the 7-8-week-old Toklat pups rush to a 13-14-month-old brother as he gets up from rest nearby. Somewhat before and continuing after weaning, the pups are alert for any possibility that an older wolf might regurgitate for them. They “attack” and food beg whenever an older wolf arrives back from a hunt or approaches from within the homesite (den or rendezvous site). Successful hunters sometimes regurgitate more than once, even hours after returning from a hunt. Refer to various blog entries dating back to June 2008 to see this male and another black male from the 2008 litter growing from small pups to adult size.  July 4, 2009.