Wolf Pair Bonds
 
Wolf reproductive bonds easily rival or exceed typical human marital bonds in their strength, and the bond between primary, i.e., “alpha,” breeders is the most important relationship in a group.  The following scenes, from my 2008 observations of the Toklat (East Fork) family of Denali National Park during the annual sexual activities in late February and early-mid March, illustrate the close bond between the present alpha male and female.  They also provide an indication of the intensity of courtship behavior and some of its ritualism.  It is worth emphasizing that close emotional ties and physical contact are not unique to the sexual activities, however.  The present male and female maintain similar high levels of emotional attachment and physical closeness year-round, and so have most of the other alpha and lower-ranking pairs of my research.        
 
In earlier years I was able to observe Toklat’s sexual activities as they progressed for about a week and then culminated with several days of copulating.  Unusually windy flying weather halted my close observations this year as the copulations seemed about to begin or intensify on March 3-4.  Radio collar signals indicated the two were still at the March 3-4 location on March 6 and thus that their sexual activities probably continued at least that long.  There was no further sexual activity when I was able to resume close observations on March 10.
 
Below (4 photos) and top of page, February 27 and March 2:  The charcoal-gray Toklat alpha male and his tan-gray mate close together, as usual, during courtship.  Overall he is the dominant wolf of the family, but she is often more assertive - including toward him - and commonly leads.  Both are in their prime at five years of age.
 
 
 
    
 
 
 
 
 
 
 
 
 
 
 
 
 
 
Courting wolves commonly “snuggle” while walking and laying together.  This behavior probably has the same adaptive value as the snuggling, hand-holding, arms-around, and related contact of human courtship.
 
                                    
 
                        
 
 
The male and female largely keep to themselves during courtship but usually do not separate very far from the other family members.  In all of these photos, the others - mostly their 1-3-year-old offspring - are within a hundred yards (meters) or so.  Below, on February 28, just prior to the most intensive sexual activities, mother tolerates some face-licking from one of the younger wolves.  Contrast this to her response (4 photos further down) when a young wolf approaches three days later.
 
                         
 
      
                    
                    
 
 
 
Above, left, and below (3 photos), March 2:  The female is receptive, standing for the male, but apparently he isn’t ready at this time.  First he sits, then he lays down.  Sometimes, as in these photos, the wolves look away in separate directions, almost as if trying to convey nonchalance amidst what amounts to a high-key interaction that requires considerable synchrony.              
 
 
    
 
 
 
 
 
                                                               
 
                                                                                                                                                                                    
Below (4 photos), March 2:  Seven of the other Toklat wolves about a hundred yards (meters) away seem restless. One of the young wolves approaches in a submissive, seemingly innocent way. Bad decision.  The female, who is perhaps a little on edge after the above sequence, immediately jumps the approaching wolf and reprimands it for several minutes.  The young wolf’s intrusion doesn’t seem to matter much to the male, although the mild interest that he does express suggests it might be a female that the older female is treating as potential competition.  The young wolf undoubtedly suffers hurt feelings but not any obvious injuries.              
 
                    
 
                
 
                      
 
            
 
 
Below, March 3:  Although the windy weather prevented me from observing subsequent copulations, the way the two are standing in this scene comes close to showing what a tie looks like.  The male mounts, thrusts, then slides off locked, standing back-to-back and/or side-by-side with her.  Most ties last for 10-13 minutes, and during this time there is commonly much mutual face-licking and other affection.  Usually within days if not hours after the sexual activities, and sometimes even before, the pair and others go to an established natal den where they may work together for hours cleaning it out, despite any lingering deep snow and even though they won’t occupy the den for another two months.  Sometimes the wolves mate right at a den.  They seem able to associate their sexual activities with the production of young.                      
 
                            
 
 
Some of the details of behavior I observed when a wolf lost a mate, using examples from Toklat’s recent history, further illustrate how closely and almost irreversibly wolves can bond but also that this doesn’t necessarily preclude additional matings:
 
In February 1997, the second-ranking Toklat male died in a snare just outside the northeast park boundary.  A month later, his mate, the second-ranking female with whom he had produced at least one previous litter (simultaneous to the alpha pair’s litter), left Toklat and joined another group, then disappeared after dispersing again two months after that.  Had these two survived in the Toklat family they likely would have assumed a more important role, given the aging Toklat alpha female’s reproductive failure in May 1997 and death in April 1998.    
 
The next Toklat alpha pair began producing litters in 1998 and maintained one of the closest, most efficient bonds I have seen in 41 years of research.  In March 2001, the male died in his prime during radio-collaring (April 2002 report, Reports page), a week after mating with the female, who likewise was in her prime.  Two adult sibling males from 170 miles (270 km) away showed up at the natal den in early June shortly after the female had produced her dead mate’s new pups (July 2002 report, Reports page).  One of them soon became the new Toklat alpha male, and both helped raise the unrelated dead male’s pups without any obvious difference in effort or affection vis-a-vis the mother and her seven older offspring.  A 2-3-year-old daughter bonded with the newcomer alpha male and they became the primary mating pair during the next breeding season, in March 2002.  The male also copulated with her mother - at the same location, two hours later, with the daughter’s apparent enthusiasm and cooperation (via oral stimulation during the tie).  
 
A month later the mother separated from the others and for the most part remained alone, probably of her own volition, while continuing to range within the established Toklat territory.  An adult male from 30 miles (48 km) away was with her for a few days but then left.  She did not enjoy much hunting success on her own.  In May, she occupied the same natal complex that she and her dead mate used for their first litter in 1998.  She produced 1-2 pups but they died inside the den soon after birth; she was in advancing stages of starvation by that time and probably wasn’t lactating much if at all (these details determined from a later necropsy).  I watched her dying alone in early July, so weak and emaciated that she could no longer stand for more than a few seconds at a time.    
 
Meanwhile, her daughter, other offspring, and the new alpha male attended five pups at the natal den the family had used since 1999 (and in certain earlier years).  However, following a mid-summer move to a rendezvous site, they lost the entire litter, apparently to a marauding bear.  They produced surviving litters in each of the next two years, in 2004 with help from a young newcomer female who became the pups’ primary attendant during the homesite period.  
 
In late January 2005, the female was caught in a trap and a snare just outside the northeast park boundary.  Her GPS radio collar locations, necropsy results, and other information indicated that she probably struggled for two weeks while caught, until the trapper shot her and took her away on February 11 (see the bottom two photos and caption in the March 8 blog entry).  The male left the trapping area with the nine others on February 11, apparently having remained with or near her during most if not all of her ordeal.  They went 14 miles (22 km) straight to the natal den where he and the female had produced most of the others in 2003 and 2004.  There they cleaned out the burrows despite the 2-3 ft (1m) snow cover and even though the normal courtship-mating and denning periods were at least two weeks and two and a half months away, respectively.  After the alpha male of the neighboring Margaret family died in a snare in the same general area in February 2004, his mate did virtually the same thing, traveling to their established natal den 10 miles away.
 
The Toklat male retraced the 14 miles almost straight back to the trapping area the next day, with such a rapid pace that at times the others lagged a mile or two behind.  His demeanor was unmistakable and seemed almost obsessive.  He was clearly intent on finding his mate.  When we left, he was sitting alone atop a high plateau, howling over and over again in obvious distress toward the trapping site a few miles (5 km) away (second to last photo, March 8 entry).  He continued returning to the trapping area through mid March, and two more wolves of the group were caught there - a 2004 pup and the newcomer female attendant.  After essentially separating from six remaining young wolves back inside the established territory, he mated with another female (probably also one of his young) on March 8-12.  But his focus seemed to remain on the dead female.  Overnight on March 12, for example, he went 20 miles back to the trapping site, again with the same unmistakable, almost obsessive behavior, including by moving so rapidly that the female with whom he had just copulated could barely keep up.
 
He and the above female became separated a few miles east of the trapping area by March 17.  She ended up dispersing 70 miles (112 km) southwestward and eventually joined the Eagle pair, though with no indication that she produced any pups (p. 12 of the August 2007 report, Reports2 page).  He began a month-long series of travels primarily back and forth along the east park boundary, at least 15 miles from his surviving young and established territory.  He joined an unrelated female in the east boundary area in late March or early April.  Hunters shot this female while the two wolves were together just outside the southeast corner of the park on April 8 or 9; he was lucky to escape.  
 
On April 12, he returned to the Toklat territory for the first time in a month but remained for only 1-2 nights and apparently did not reunite with his six surviving young.  By April 14 he was back along the east park boundary, 15 miles outside the Toklat territory.  Three days later a hunter shot him outside the southeast corner of the park, near the area where hunters shot the female he was with nine days earlier.  Now all that remained of Toklat were the six yearlings and two-year-olds, whose story continues in the August 2007 report, earlier blog entries (esp. Jan 19 and Feb 3), and above.
 
 
Apr 8, 2008
The Toklat alpha male and female walk together affectionately during courtship activities in late February 2008.