Toklat’s High Value
The Toklat wolves of eastern Denali National Park live together as one of the world’s oldest known non-human social groups in the wild, a family lineage that has persisted for at least 42-70 years. As such, Toklat, also known as East Fork, is a biological treasure, a gold mine of research opportunities for understanding more about the characteristics of a successful society and cooperative behavior in general. Toklat is also important to the biological integrity of Denali as one (the oldest) of only about 20 predator functional units that interact year-round with the ungulate populations and subpopulations of this ecosystem. It is the world’s most viewed and probably most photographed group of wolves and provides a wealth of educational opportunities. The history Toklat embodies in this ecosystem gives it a special esthetic presence as well, something like the aura of a 150-year-old bowhead whale or 2,000-year-old redwood.
Below (2 photos): The Toklat family of 17 (one is ahead, outside the photo), in late January 2008, three weeks before losing seven wolves during a trip into the northeast park boundary trapping area. Second photo: Seven of the 17 (others are outside the photo) try to cross a river without getting wet on a snowy November 2007 day. The alpha female is in the midst of a leap, while a black female (to her left) who cooperatively nursed the pups with her at the spring natal den contemplates doing the same. Just behind the black female is the charcoal-gray alpha male, and behind him is a young adult black male, one of the largest wolves in the family. The black female and black male were among the seven wolves apparently snared on the February trip.
My emphasis in this essay is on the behavior that makes Toklat - and made its former neighbor to the east, Savage - so successful, interesting, and important to study.
[The Savage family was well established in 1966 but disappeared in winter 1982-83, probably due to illegal aerial hunting. None of Savage’s successors - Headquarters, Sanctuary, Margaret, Toklat East - has lasted long in the presence of continued trapping and hunting just outside the east and northeast park boundaries.]
When I began this research in 1966, Toklat was a well established family. Adolph Murie, the famous naturalist-scientist who was still studying bears and wolves when I arrived, told me this was the same family of wolves he began observing in 1939 and described in his classic 1944 monograph, The Wolves of Mount McKinley. From my own observations, I know Toklat is at least 42 years old, and from what Murie told me, it is probably at least 70 years old. Wolves of this family have been radio-collared continuously for the last 21 years.
This ranks the Toklat wolves and a 46-year-old or older community of chimpanzees in Gombe National Park, Tanzania as the two oldest-known, longest-studied large mammal social groups anywhere in the wild.
Under natural conditions, longevity and levels of sociality can be expected to vary widely among groups of wolves, in areawide patterns related directly and indirectly to prey resources. Toklat, Savage, and Savage’s successors have been the primary resident groups of eastern Denali, the only major area of the northside park/preserve where mountain sheep as well as moose and caribou are available to wolves. This richer, more diverse prey base has enabled Toklat and (formerly) Savage to avoid much of the direct competition and strife with other groups that results in higher natural rates of turnover and probably lower levels of sociality for wolves in surrounding areas to the north and west (Haber 2007a). Sheep provide a key additional winter hunting opportunity that reduces the need of the eastern groups to venture to the more dangerous caribou wintering areas. The eastern groups also largely escape competition from migrating wolves because the latter are much more attracted to the nearby caribou wintering areas, in part because caribou are easier to hunt than sheep.
Eleven Toklat wolves bunch together while chorus howling. February 1973.
Toklat wolves chase a fox that approached too closely. March 1973.
Characteristics of success
The Denali research suggests that being able to accommodate major prey changes, durable relationships among key individuals, and heavy reliance on altruism and other sophisticated cooperation are among the most important reasons why Toklat still persists and Savage lasted at least 17 years.
Although both groups benefited from good overall prey resources, there were differences in the way these resources were distributed between their territories and major seasonal and long-term prey changes. Both groups seemed to adapt primarily through social adjustments (Haber 2007a). Adapting efficiently to food constraints is obviously vital for any society’s long-term success and in this case is consistent with other evidence (Haber 1977; Walters et al 1981) that the eastern Denali wolves conserve their prey as “prudent predators.”
Durable relationships characterized both groups, especially their primary reproductive bonds. For example, over a span of 33 years there were only 5-6 Toklat alpha (primary breeding) females, one of which retained the position for 13-14 years until she died naturally. The behavior of three subsequent Toklat alpha and beta pairs in their prime and of the surviving mates after human-caused breakups (April 8 blog entry, last nine paragraphs) leaves little doubt that these and other relationships would have lasted much longer under natural conditions. In the Savage family, the same two males occupied the alpha and beta positions and maintained an efficient division of leadership for at least 7-8 years.
As noted in the April 8 entry, strong pair bonds do not necessarily preclude polygyny (“monogamy” can be a risky term). I directly observed one instance of cooperative polygyny in Toklat, in 2002 between the new alpha male, his primary mate, and her mother. This seemed to have little if any affect on his relationship with the younger female, which remained exceptionally close until she was trapped (and he was eventually shot) three years later. For almost two months the male continued returning almost obsessively to the site where she was trapped, even right after mating with another female. I suspected other polygynous matings with little effect on the primary bond, especially in the Toklat family with its relatively high frequency of multiple simultaneous litters (in 11-13 of the 38 years I could determine the number of litters); in 1990 there were at least three and probably four Toklat litters. Savage produced multiple litters (2) in only one of the 12 years I could determine the number of litters. In that case, I observed a young adult male making intense sexual advances on a young adult female during the mating period in March, after the older (alpha) primary pair had completed its courtship and mating activities.
Cooperative polygyny. March 2002.
The Toklat alpha male mates with a young adult Toklat female, his primary mate ...
... then after resting for a couple of hours mates with her mother, as the daughter stimulates them. The mother’s established mate, the former alpha male, died during radio-collaring a year earlier.
Non-breeders, especially young adult females, routinely assist - i.e., “help” - breeders and in the process often enhance their own potential future reproductive success (e.g., by practicing for motherhood or delaying dispersal until there are better opportunities elsewhere). However, much of the helping I observed in Toklat and Savage also appeared to be altruistic. Helpers included high-ranking, experienced adult males and females who deferred breeding even though they almost certainly could have dispersed from the group, found mates, and succeeded elsewhere. It was commonplace even for breeders to assist other breeders of the group, including by nursing and otherwise provisioning and protecting the young interchangeably. I observed cooperative nursing by non-breeding, pseudo-pregnant females as well.
The black Savage alpha male (center) plays with his mate near the peak of her estrus, shortly before mating with her. The beta male (bottom) is keenly interested but stays his distance as the alpha male warns him with a menacing stare and probably some growling. The beta male shows submission by turning his head to avoid the stare and by raising his right front paw. The wolves behind are younger offspring. Note from the raised left front paw of the young black who is sitting, watching, that the alpha male’s assertiveness leaves him a little uncertain as well. The beta male led many of Savage’s routine activities and had an otherwise friendly, cooperative relationship with the alpha male, but did not breed. March 1972.
Young Toklat and Savage wolves remained dependent beyond the summer homesite period, for as long as 2-3 winters while developing full hunting proficiency. Not surprisingly, much of the helping and related behavior that I observed, such as the division of leadership between the Savage alpha and beta males, was prominently on display year-around.
Some form of reciprocity apparently also operated. Two adult males from 170 miles (270 km) away took over the Toklat family two months after the established Toklat alpha male died and helped raise the dead male’s four unrelated newborn pups with what appeared to be the same dedicated behavior, including displays of affection, as their mother and older siblings. Yearling helpers play an important role in caring for new pups, thus there was a potential genetic payoff for one of the new males (the new alpha) the next breeding season when he produced his own litter.
Four Toklat adults on the hunt in July 2003. The alpha female is leading, followed by a younger sister, the beta male - likely her brother, and her mate, the (black) alpha male. The beta male returned in excellent condition in July 2003 after an absence of about eight months, and helped raise the alpha pair’s 2003 pups. The black male joined Toklat in 2001 and helped raise the unrelated dead alpha male’s pups.
Below (2 photos): Wolves of the Savage family on the hunt (February 1972) and playing after a fresh snowfall (March 1974).
There were also high levels of cooperation during hunting activities. Among the Toklat and Savage cooperative hunting tactics I have observed:
• coordinated setup of attacks on caribou, sheep, and moose from up to several miles away and several directions, including by driving them into difficult escape terrain,
• storming and then closely circling a moose to test it (pictured below),
• chasing sheep, caribou, and moose to one or more wolves waiting in ambush,
• ambushing moose or caribou at a mineral lick,
• decoying caribou while other wolves stalk from behind or on a flank,
• killing a moose by holding its nose and anchoring the dangerous front hoofs while other wolves attack from the rear and elsewhere (pictured below).
11 wolves of the Savage family test a moose with the “storm and circle” tactic that I’ve only seen this group use. The wolves storm the moose (2 wolves are lagging), apparently trying to rile it, then circle closely while evaluating whether to try for a kill. In this case, as in about 95% of wolf-moose encounters, the wolves judged the moose to be too dangerous and left within a few minutes. Read the left column of the Home (Alaska Wolves) page for related details. January 1971.
13 Toklat wolves attack an old moose. An experienced adult (black, tail up) grabs the nose, making it difficult to strike with the front hoofs, a moose’s main weapon. Others then attack from the rear and elsewhere. It still took the wolves two attempts and almost two hours to bring down this old cow.
March 1973.
Three Toklat wolves coordinate an attack on a grizzly bear to drive it from a nearby moose kill.
August 2000.
Perhaps the ultimate cooperation is the close inbreeding that Toklat, Savage, and other wolves seem able to do successfully. For example (Haber 2007b), six Toklat wolves, all almost certainly from the 2003-2004 litters, produced eight pups in 2005 and six in 2006 with high survival rates in both cases. The dominant pair of the (2003-2004) six also produced at least 5-6 pups in 2007 and 6-7 this spring (May 2008). There was high survival of the 2007 pups through at least 8-9 months of age, but the snaring losses in February (e.g., April 20 entry) confound a determination of natural survival thereafter.
The Savage alpha male, whose mate was shot in 1968, almost certainly mated with two daughters from the 1968 litter in 1970-1973 (with one in 1970-1971 until she disappeared, then with the other, who was subordinate to the first). These four primary matings produced 27 pups, including a litter of nine in 1970. I observed intense sexual activity between two of the 1970 pups in 1972. Two months later a young female produced a litter of at least four pups that was raised in the group together with the primary litter of five.
Inbreeding has negative consequences for many species. However, highly social species may avoid or offset these problems in unanticipated ways. Well established wolf social systems are relatively closed to newcomers, and, more importantly, newcomer-breeding; this may facilitate purging of genetic loads (deleterious recessive genes). Advantages accruing from sophisticated cooperation among close kin could also offset disadvantages from any losses of phenotypic variation or the effects of inbreeding depression. Recent research on a small Scandinavian wolf population indicates that inbreeding, at least to the level of sibling matings, does not necessarily lead to a major loss of genetic variation in the first place, least of all among the breeders (Bensch et al 2006).
Understanding the roots of cooperative behavior, including our own behavior, is among the most important endeavors in all of scientific inquiry (Kennedy and Norman 2005). To be able to observe the foregoing and other details of behavior for the same groups of wolves in the wild for decades presents one of the rarest and most direct opportunities for new insights about these roots. Given the likelihood that wolf social organization provides an excellent model for early human societies (better than non-human primates), continuing to allow the Toklat and other Denali wolves to be trapped and shot is especially foolhardy.
Below: Ten of the 11 Toklat wolves still together in late January 2005; one wolf is just outside the photo. The black alpha male is sitting near the top center of the photo, getting attention from a young female that the group adopted seven months earlier and who became the primary attendant of the six pups at the summer homesites. Standing just below the male, looking downslope, is his mate, the alpha female. The black male near the lower left (howling), one of the four pups this pair produced in 2003, is the current alpha male. Soon after I took this picture, the alpha female, the young female attendant of the pups, and a pup were snared and trapped just outside the northeast park boundary. Another young female dispersed after becoming separated from the alpha male in the trapping area. He was shot a month afterward in another area.
The six surviving Toklat wolves, following the Feburary-April 2005 losses described in the above photo caption. Three of the six were yearlings, and three were two-year-olds. The large black two-year-old, third in the line (howling, lower left in the above photo), became the new alpha male. He and one or two of the females have thus far produced at least 25-27 pups, 2005-2008. April 2005.
Toklat wolves at the natal den in May 2006. Several young wolves are playing with the three-week-old pups just inside one of the burrow entrances, lower right. All of the wolves relish the opportunity to play with and dote over the new pups each spring. The pups’ mother, the alpha female, is laying down on the far left. The black female on the far right cooperatively nursed the pups with the alpha female in at least 2005 and 2006. She was one of the seven Toklat wolves that disappeared on Toklat’s trip to the northeast park boundary trapping area February 21-23, 2008.
Family lineages and genetic dilution
A family’s history consists of social as well as genetic continuity. Observations of the Toklat wolves since 1966 indicate both kinds of continuity and confirm they are a single, 42-year-old or older family lineage rather than two or more consecutively recolonizing families.
The close radio-telemetry contact that has been maintained with these wolves since 1987 should not leave any doubt as to continuity from both standpoints over the last 21 years. The overlapping “timelines” of at least a dozen wolves that I (and others) could identify during the previous 21 years, including two wolves that were subsequently radio collared in 1987 and 1991 as the primary male and female breeders, provide one indication that a single family lineage spans both periods. Overlapping spatial and behavioral traditions provide another. For example, not only did wolves continue to use the same homesites during both periods but they also varied their within-summer occupancy among certain sites in the same ways. In the most striking example, adults repeatedly moved pups between two major denning complexes during both periods, using the same 14 mile (23 km) route each time.
In The Wolves of Denali (1998), Mech and his associates questioned my reference to Toklat as a continuing family lineage because, they argued, the original genes would be so “diluted” after such a long period of time.
First, this overlooks the importance of social continuity. Second, for what social group, of any species, do the original genes not become diluted? There may be much less dilution in groups like Toklat and Savage than in most human families, given that the wolves seem able to inbreed and newcomers become breeders less often.
Generations of the McDonald family have maintained the same Nebraska farmstead for more than 150 years. Some offspring have dispersed to distant areas, and some have started new farms nearby. But together with marital and adopted newcomers, other offspring remain on the farm and define a core family lineage. They have progressively diluted old McDonald’s (and Mrs. McDonald’s) genes, but there is also social continuity. The original (renovated) house and barns that old McDonald built are still home. They celebrate the harvest with remembrances of the old man and the farm’s trials and tribulations. Their children still gather mushrooms in the woodlot next to the south forty. Most of the original farmstead remains in production with much the same schedule of crop rotations, though some inferior acreage has been sold on one side and productive neighboring acreage annexed on another.
Distinct short-lived and persistent farm family lineages can be identified in mosaics across the landscape, the latter especially where soil and water conditions are the best. And for the oldest of these we recognize special cultural, esthetic, biological, and other values, just as we should recognize the enormous values of an old non-human family lineage such as the Toklat wolves of Denali National Park.
References cited
Bensch, S., H. Andren, B. Hansson, HC Pedersen, H Sand, et al. 2006. Selection for heterozygosity gives hope to a wild population of inbred wolves. PLoS One 1(1): e72. doi:10.1371/journal.pone.0000072
Haber, G.C. 1977. Go to link at bottom left of Home (Alaska Wolves) page.
Haber, G.C. 2007a. Go to December 2007 link on Reports2 page.
Haber, G.C. 2007b. Go to August 2007 link on Reports2 page.
Kennedy, D. and C. Norman (eds). 2005. What don’t we know? Special Section, Science 309: 75-102.
Mech, L.D., L.G. Adams, T.J. Meier, J.W. Burch, and B.W. Dale. 1998. The wolves of Denali. Univ. of Minnesota Press, Minneapolis. 227 pp.
Murie, A. 1944. The wolves of Mount McKinley. National Park Service Fauna Series 5. 238 pp.
Walters, C.J., M. Stocker, and G.C. Haber. 1981. Simulation and optimization models for a wolf-ungulate system. Ch. 16 (pp. 317-337) in C.W. Fowler and T.D. Smith (eds), Dynamics of large mammal populations. Wiley, New York. 477 pp. Republished 2004, Blackburn Press, Caldwell, New Jersey.
Jun 17, 2008
Eight wolves of the Toklat family depart from a rest stop; two are ahead, outside the photo. The alpha female is second in the line of six. The alpha male is at the rear, about to pass two of his grown young. Only six wolves, including the alpha pair, could be confirmed in the group as of early May 2008, just prior to the birth of at least 6-7 pups. I am still seeing only 5-6 adults and yearlings at the 2008 natal den. There were 17 wolves in the group as of early February 2008 but only 10 when they returned from a February 21-23 trip into the northeast park boundary trapping area. Two more disappeared after that. April 2008.