Wild Hunters 3: Toklat’s Shift To Hares
 
The Toklat wolf family has persisted for at least the 41 years of my research in Denali.  But now its history has taken a twist that represents a sad loss even as it showcases high intelligence.  Toklat’s spectacular, decades-old winter sheep-hunting skills and routines have virtually disappeared since the last experienced adults were trapped and shot in 2005.  The young survivors have focused instead on a much easier food source, snowshoe hares (Lepus americanus), during a coincidental peak in hare abundance.  Their adjustments to the now-declining hare peak are fascinating, illustrate high intelligence, and have enabled them to extend their reproductive success.  
 
Toklat’s previous sheep-hunting activities likewise illustrated high intelligence and other impressive skills, and, unlike the new reliance on hares, have proven to be of central importance for long-term success in this area.  Probably for good reason, no other wolf research has reported heavy “boom-bust” reliance on hares, nor have I observed it for any other Denali groups or for Toklat during earlier hare peaks.  Refer to the August 2007 report (pp. 3-9) and December 2007 paper (Reports2 page) and the January 19, 2008 and November 9, 2007 blog entries for background.  There are also related materials on the Reports page, especially in the December 2006 report/museum exhibit.
 
Past sheep-hunting skills
Although this is still an unfolding story, already it seems unlikely that Toklat’s sheep-hunting skills will reappear anytime soon with the remarkable tuning that was in place.  The experienced Toklat wolves seemed to know every significant sheep mountain in their 600-700 square mile (1,550-1,800 sq. km) territory and checked most of these regularly throughout the winter.  They could pick out sheep at almost unbelievable distances, especially when scanning from one high ridge or mountaintop to another.  Once I observed them intently study 20 (white) sheep sleeping on a slope against a fresh (white) snow cover from another slope eight miles (13 km) away, then excitedly head straight to the sheep (traveling with the wind) and catch one.  They developed impressive search and pursuit tactics that utilized the treacherous, rugged terrain to advantage, including various kinds of ambushes and especially surprising sheep from above, forcing them to flee downslope away from the best escape terrain.  One tactic seemed to involve deception.  Usually the wolves left soon after a sheep fled beyond reach onto a narrow ledge, spire, etc.  However, sometimes they waited quietly nearby, out of view, for up to a day or two.  Unable to eat, drink, or often even lay down, the sheep would finally come out to a vulnerable location where the wolves were afforded an opportunity to ambush it.  
 
Toklat wolves, led by the alpha female and alpha male, hunt sheep along a steep, narrow 4,800 ft (1,463m) ridge.  March 2004.
  
 
 
 
 
 
 
 
 
 
Toklat wolves contemplate several sheep staring up at them from a short distance below a 5,000 ft (1,524 m) ridge.  In this case the wolves decided not to pursue.  March 2004.  
 
 
 
 
 
 
 
 
 
 
 
Just learning to negotiate the high, icy knife ridges, peaks, and slopes usually required extensive guidance from experienced adults.  And alongside these skills, Toklat displayed impressive physical prowess, regularly climbing up and down the steep mountains like they weren’t even there.  Ascending a 50-60º snow-covered slope with an elevation change of 1,800 ft (550 m) in 18 minutes - no rest stops, then breaking into intense play at the top, is among the typical examples I recorded.  
 
Several times pilots and I watched in disbelief as Toklat wolves literally skied, stiff-legged, straight down icy chutes, adjusting their upraised tails back and forth for balance.  We watched three Toklat wolves do this while trying to intercept sheep that were fleeing below from several other Toklat wolves chasing from another direction.  The three bounded over the top of a high, narrow, rocky ridge at a full run and jumped immediately - apparently without looking first - into the top of an icy chute and skied straight down the 60-70º, 2,000-3,000-ft run (600-900 m); they zipped down this chute faster than any Olympic downhill ski-racer I’ve watched.  At the bottom they switched immediately to a full run, although in this case the sheep escaped to safety on a nearby ridge.  
 
The Savage family, Toklat’s former neighbor to the east, developed similar sheep-hunting skills over its history of at least 17 years.  The 12 Savage wolves disappeared in winter 1982-83, with indications of illegal aerial wolf hunting at the time.  Three groups - Headquarters, Sanctuary, Margaret - successively recolonized major portions of the Savage vacancy over subsequent years, but each terminated due largely to human causes.  None hunted sheep or developed sheep-hunting skills to the extent I observed for Savage - nothing even close.  There hasn’t been enough continuity to facilitate the necessary learning.
 
    
 
 
 
 
 
 
 
 
 
 
Three wolves of the Savage family chase two sheep downslope from a high plateau.  Both sheep escaped.  February 1973.
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
Wolves of the Savage family crowd together while eating a sheep just below the center of the photo, after a chase down a steep, icy mountain slope.  November 1972.  
 
 
 
 
 
 
 
 
 
 
 
 
Intelligent hare-hunting adjustments
During the past two winters of high hare densities, the Toklat wolves’ usual hunting tactic was to spread out as they entered a willow patch or spruce-willow area and flush hares.  Enough hares were soon jumping around in close proximity that the wolves could catch them fairly easily (see the August 2007 report for details).  The wolves spent most of their time in the eastern third of the traditional Toklat territory, often splitting temporarily into small subgroups and separating individually.
 
This winter, hares are much less abundant and continue to decline.  They seem to be changing their defensive tactics as they decline, and the Toklat wolves seem to be adjusting.  Hares are now relying more heavily on camouflage, sitting motionless in willow thickets with their white fur blending against the snow, typically not fleeing until a wolf has approached much more closely than last winter, sometimes within 2-3 feet - or not fleeing at all (and promptly getting eaten).  Driving and flushing do not work as well for this behavior.  Now the Toklat wolves examine the willow thickets more intently without as much driving, obviously recognizing the effectiveness of the hares’ camouflage and the potential for finding them this way.    
 
They are also ranging much more widely across the traditional territory and outside in northward and eastward directions, and are remaining together more as a group.  The winter observations suggest that so far most of this searching has been for better hare-hunting opportunities.  Ultimately this would be an exercise in futility because of the synchronous way hare numbers rise and fall across large regions.  However, twice during my observations over the last three months they also killed larger prey, in each case a single sheep they caught in relatively low terrain. This is a far cry from the past sheep-hunting skills and routines.  Nonetheless, it is reason for optimism.          
 
The following photos, from my October 2007-January 2008 observations, illustrate Toklat’s latest hare-hunting behavior.  
 
 
 
 
On a snowy November 2007 day, 15 Toklat wolves head across an open tundra divide into a willow-lined creek bottom where they expect to find some good hunting for snowshoe hares.  During the previous two winters, while hares were still at their latest peak in abundance, the wolves would have quickly flushed dozens of hares in a willow area such as this.  But now hare abundance is on the downside of the peak, and the wolves are relying more on their keen eyesight to spot camouflaged hares sitting motionless at more scattered locations in the thickets.  Upon arriving (below), at least a half dozen of the leaders begin looking for hares.      
 
 
The above willows do not produce many opportunities.  When we return the next day, the wolves (plus 2 others that rejoined them) have found better luck in another willow area five miles away.  During the hour that we circle, we see them find 15 hares as they search a 300-400-yard (meter) length of willow patches along a river.  Seven or eight of the 17 wolves catch and eat at least 10 of these hares:
 
 
A hare flees a willow thicket being examined by some of the wolves but zigs where it should have zagged, right into the three wolves on the left. The black wolf nails the hare and promptly eats it, without any challenges from other wolves. Others watch with keen interest but then return to searching the thickets.      
 
 
 
 
At their present lower densities, hares seem to sit motionless in the thickets more often when wolves approach, their white fur camouflaging them against the snow. The wolves have figured out this tactic and seem to be countering it by studying the thickets more intently, alone and in groups, apparently without prior cues (movements, etc.).  I saw them find hares in the third and sixth of these photos (alongside and below).        
 
 
 
 
 
 
 
 
 
 
 
Below, the bottom wolf has just caught a hare, and the wolf at top right is chasing another hare.
 
 
The next four photos, from an October 2007 hunt, illustrate playful stalking, short chases, and close contact to pick up scraps.  However I have not observed any serious attempts to steal hares, not even by wolves of higher rank.  
 
 
 
 
 
 
Ravens typically follow wolves to pick whatever they can from the remains of their kills and winter kills (scavenged carcasses), even when hares are the only prey.  Ravens also delight in teasing wolves.  In the first photo, below, the Toklat alpha male tries to ignore a raven that repeatedly flies low overhead (the wolf’s hefty appearance attests to how well he is doing on a diet of hares).  In the next two photos, a raven teases a younger wolf, who takes the bait.  November 2007.
 
 
        
 
Below:  The 17 Toklat wolves traveling in the typical single file manner.  The alpha female and alpha male are the fourth and fifth wolves in the line.  January 2008.
 
 
 
 
Feb 3, 2008
A Toklat wolf with a freshly caught snowshoe hare, carrying it a short distance from others of the family just outside the photo, October 2007.  The young Toklat survivors have thrived on hares since the experienced adults were trapped and shot in 2005, but the hare population is now on the downside of its latest 8-11-year peak in abundance.