Known pups surviving to yearling age over those years totalled fourteen. Known losses include four dispersers, six yearlings or adults dead, and one vanished, for a total of eleven. During the winter of 1997-98, the pack consisted of four wolves: the alpha-male and three pups. His mate had died of unknown causes the previous August.
Genetics showed that long-lived Redpole 4, who lived two territories distant to the west, had made an important contribution to the Basin Depot pack. He was the grandfather of three Basin Depot wolves and the great-grandfather of three more. His descendants, however, are spread through five different packs.
Recovery of a wolf population is an uneven process. Even though the ban reduced wolf killing, wolves continued to die, just enough to keep disrupting continuity and erasing gains. The population struggled, with life expectancy remaining low, dispersal exceeding immigration in the Bonnechere Valley, and low numbers of pups making it to yearling age. In 1996-97, mortality was 36 per cent and the population of our entire study area declined again by 28 per cent. Human killing in townships outside the ban area accounted for 75 per cent of these losses. In 1997-98, the population remained stable and low.
Until the Basin Depot pack expanded its territory to the west to meet the eastern boundary of the Jocko Lake pack in the summer of 1995, a big, packless piece of land about ten kilometres in diameter remained between them. That was not the only persistent wolfless hole. To the north in the Petawawa drainage, the territory of the Travers pack took two years to refill. North of Grand Lake a fifteen-kilometre hole between the Pretty Lake pack and the newly formed Hardwood Lake pack took three years to fill, and still there is a gap.
Just when the landscape seemed mended and the population had patched most of its holes, new ones appeared, the consequence of the 1996-97 decline. After we lost two radio-collared Pretty Lake wolves, both dying north of the park, the Pretty Lake territory remained vacant throughout the summer of 1997. A crew worked there for six weeks and only found evidence of wandering singles. Then in January 1998 the adjacent Jack Pine pack claimed the western part of the land. Something happened, as well, to the land lying north of Grand Lake — it was packless again.
As they resettled the land, the struggling reoccupants carved out largely different territories than those held by their predecessors, making it obvious that topography does not define boundaries. Grand Lake may always separate packs, as may the Petawawa River in summer, although wolves can swim both. Other than these two features, territorial boundaries must exist in the minds of individuals. They may fall where wolves first turn around because they are too far from a den or rendezvous site or other pack members, or where they encounter the scent of unknown wolves, and those places are remembered, retraced, and remarked over time.
In an exploited population, boundaries may be forgotten. Survivors may have no memory of den sites, rendezvous sites, ambush sites, or easy travel routes along esker ridges and across beaver dams, leaving the population less fit. The relationship between animals and ecosystem is fractured. A pack — especially a small, reforming one — may need all the subtle advantages it can get to bring enough pups through. The inexperience of breeding animals may contribute to the low number of young that reach yearling age that typifies the Algonquin population.
Up to a point, a wolf population has the biological capacity to offset its human-caused losses. A numerical balance between births and deaths may be acceptable today as a conservation objective over much wolf-occupied land.
Someday, however, we may re-evaluate our place in nature and decide that it is no more acceptable to kill wolves than it is to kill lions or cheetahs or other big cats. And, even today, is the mere maintenance of such a balance between births and deaths good enough for special places we call parks — supposed holding places of biological treasures? Should wolf populations even there be continually chipped away at, their adaptations driven by human influence? Can’t we allow nature to run anything any more — just on principle, just out of humility?
Unless officials that control policy in the MNR come to see parks as places to provide this ecosystem-sensitive level of protection and complete the task of boundary protection all around Algonquin Park, the wolf population there, as in so many other places, will never heal. At best it will only persist.
SPACE GAMES AND WOLF-DEER RELATIONSHIPS
COLD CLOAKS the snowy Algonquin landscape, and a deep quiet settles into the hills. Tourists have departed. Pine-softened wind sometimes brings the distant call of ravens, or a jay. Up in the hardwoods a downy woodpecker hammers, or some siskins with their questioning calls dip by. Even these quiet sounds cease as shadows swallow the day. Winter Algonquin is a place of snow-filled silences.
Wolves can find “peopleless” sanctuary there, can concentrate on being predator, not prey. Across vast stretches of the park, no human tracks disturb the snow. No threat of a bullet in the open places, no risk of a snare along the trails or strychnine in a chunk of meat. Just wind, cloud, ice, snow, moose — the wild harmony of winter.
Yet many wolves choose to leave the park and go where they daily encounter snowmobiles, cars, trucks, people, dogs, houses, barns. There they wait in ditches for vehicles to pass — at least the cautious ones do. They circle open fields and avoid barking dogs. They feed on deer behind the store at Bonnechere or the public school in Round Lake Centre. They die in traps and snares and on roads — less than before the ban, but it still happens.
An evolutionary irony, wolves are only partially adapted to shun humans. They would fare much better if the first hint of human presence triggered greater fear, if they always associated snowmobiles and cars with danger. But they hedge their bets and often lose.
They don’t need to leave the park. In the park’s northwest sector, despite a near-absence of deer year-round, the wolves have managed just fine. Their only adjustments are slightly larger territories and an increase in snowshoe hare in winter diets to about 10 per cent. Yet in the east, the wolves follow the deer out of the park even though moose and beaver are just as abundant as in the west. Apparently deer are so highly preferred that the wolves are willing to risk exposure to people to follow them. This deer storehouse may be just too attractive for their predatory genes to ignore.
Up to fifty-eight wolves, our highest one-time estimate, or about twenty to thirty on average, concentrate in one hundred to two hundred square kilometres of deer yard. This constitutes an almost unprecedented density, more than ten times greater than in the park. Normal wolf spacing is temporarily on hold, and so is aggression. Never have we recorded a wolf-killed wolf in the deer yard.
The events of one late February afternoon should have erupted into wolf violence. Instead, what we found was a dramatic example of tolerance. The big radio-collared Vireo male had travelled west of his usual location into high hills on the periphery of the deer yard. We took a bearing on his signal and crossed a snowy field to a stone fence where we picked up his tracks. Following them downhill into the trees we saw where he was joined by packmates eventually making four sets of footprints. Then, to our amazement, even more tracks appeared, exceeding the number of wolves in his pack. The tracks were strung out along a small creek. Soon we came to deer hair and a little farther found a vertebral column and skull. The wolves had pulled the deer down under spruce trees, where the snow was blood-stained and packed into ice. A typical, skinned-out deer pelt lay nearby, along with the pelvic girdle and two hind legs.
The Vireo wolf had moved off the carcass and was about one hundred metres to the south. While Mary unpacked our carcass kit, for no distinct reason I flipped the receiver to scan. At four-second intervals it switched from one frequency to the next and suddenly another signal came crackling in. Unbelievably, McDonald I was there too.
Just then a wolf howled not more than fifty metres behind us, deep, throaty howls that reverberated in the quiet woods, each emphasized by a few seconds of expectant silence. I pointed the antenna towards the wolf and scanned through the frequen
cies, but it was uncollared. So was a treble wolf who howled equally close to us from the creek.
Never had we found two packs at the same kill simultaneously, breaking all wolf rules. In summer, these packs were separated by one pack between. Yet there they were together.
We moved back along the creek, and still the Vireo and McDonald wolves shared the same bearing. Then we returned to the carcass and moved off in another direction, taking a third bearing with the same result. These two wolves and likely others were together, silent but near us, probably anxious to return. At the kill again, we hacked off the femur and mandibles, then in the fading daylight backtracked out of the wolf-filled woods.
Nobody has described wolves concentrating like this in response to the movements of deer. A wolf pack with a deer yard in part of its territory may centre its activities there, as François Potvin found in Quebec and Dave Mech in Minnesota. What we found was different: a migratory wolf population. Similar are cases of wolf concentrations around migratory caribou in Alaska reported by Warren Ballard, Bob Stephenson, and co-workers, and in the Northwest Territories by Gerry Parker. Wolves also have been found to concentrate around bison at Wood Buffalo National Park by Lu Carbyn and associates. In none of these instances, however, have the social and spacing dynamics been described.
We wanted to describe them here and in the process re-examine territorial behaviour. Were the wolves setting up and defending mini-territories in the deer yard, or were they roaming freely? If they set up territories, did packs return to the same ones each winter? Did packs avoid one another? Was wolf spacing a response to the distribution of deer, or to human disturbance, or to interactions among packs? Were wolves scavenging or killing deer, and were the deer they killed in poor shape, ready to die anyway? Why wasn’t the deer population increasing? After expanding moderately in the early years of the study, deer numbers had fluctuated erratically.
Among the possible explanations for the lack of aggression and resulting concentration of wolf packs is a superabundance of food. While not explaining why only the Algonquin wolves are migratory, the difference between our study and the ones with non-migratory wolves in Minnesota and Quebec may be only a matter of scale — ours features a complete or near-complete deer exodus over a very large area.
Alternatively, the high level of genetic relatedness among packs we found may turn off aggression and allow wolves to concentrate. Heavy human exploitation and consequent dispersal to vacant lands probably drove up this relatedness, although heavily exploited and highly dispersed wolf populations in both Minnesota and Quebec did not concentrate.
A third hypothesis is that on a range-edge for wolves, there are no defending wolf packs in the farmlands to repel migratory wolves. From the howls we heard in summer, we had an idea that only coyotes lived in the Round Lake area.
Whatever the cause, what we were seeing did not happen thirty-five years previously. Then, with many small winter deer yards scattered throughout the park, Doug Pimlott found that packs stayed on territory. Deer numbers were about ten times as great and snow was deeper. The system had changed, as systems always do.
Two master’s and two bachelor’s theses based upon five winters of data helped answer these questions. Each December after term exams we would head north, stopping to check for signals from the blue body-shop sign on the big Wilno hill. Looking over the flat extent of deer yard, we always wondered what new twists the wolf-deer system would take, as it invariably did. I arranged my lecture schedule to allow us at least one month in the field between early December and the end of March. We lived periodically for two winters in a small MNR staff house on the edge of the highway near Bonnechere Park, the first year dodging scaffolding as the house was being renovated. Then we rented various small cottages to fit our budget. One of them, the most comfortable, came with a wood stove and an outhouse.
We had funds for sixty to seventy hours of aerial radio-tracking each winter, which worked out to one three-hour flight every few days. On flight days we followed the same routine: a graduate student made early airborne reconnaissance, then Mary and I tracked to whatever kills were reasonably close. Often we arrived back at our lodgings long after dark. Occasionally we would howl the deer yard, but infrequently to avoid raising the ire of people and dogs. Saturday evenings during “Hockey Night in Canada” attracted the least attention.
As data accumulated, what we saw was almost patternless. While each winter we came to expect certain packs in certain areas, they rarely stayed for long. All packs shifted, some more frequently than others. Some stayed in the deer yard and never returned to the park all winter; others made up to eight return trips. Packs whose park territories lay closest to the deer yard travelled back and forth most often. When packs returned to the yard, sometimes they would go to the same area they had left, but other times they would centre their activities elsewhere. Each winter, a few packs did not come to the deer yard at all, or they went to the smaller Black Bay deer yard near the Ottawa River.
Some packs occupied only a few square kilometres in the deer yard; others travelled more widely. Joy Cook calculated that an average area covered by a pack over the period of one month was less than forty square kilometres. Some wolf movements were random, others clumped, others scattered. Single wolves without packs wandered everywhere.
We were seeing a maximum variety of responses to packs presented with the same environmental situation and stimuli, resulting in maximum unpredictability. That must have been confusing to the deer who could do nothing to reduce their exposure to wolves. From the wolves’ standpoint, it seemed almost planned, although a deliberate strategy for an entire population is impossible. Members of one pack can communicate frequently enough to act strategically. A whole population cannot.
We should have expected the wolves to display the unpredictable movements considering their circumstances of having no land to defend, hunting deer that were shifting in response to snow conditions and food availability, and trying to avoid other packs and human activity. Adding to the unpredictability were individual differences in experience, memory, social position, age, and pack size.
Yet, except for two instances when packs temporarily amalgamated, they were never in the same place at once. Close encounters were common, such as two or three packs only a few hundred metres apart in the same county forest, for example, or two packs heading towards one another but one turning aside. Occasionally, one pack was on a kill with other packs nearby.
Examining the data, Joy found that packs were deliberately avoiding each other. Close range, spatial-temporal avoidance was a principal conclusion in her master’s thesis. Ryan Norris, applying Joy’s analytical approaches to the previous two winters of data, found the same thing.
Still, it seemed improbable that avoidance could be so perfectly executed for such long periods each winter. At times avoidance did fail, as shown by the two-pack incidents at deer kills. As well, there was the singles bar phenomenon. Late winter is the breeding season and some wolves undoubtedly were looking for new alliances and opportunities. We know some packs went back to the park with extra members, or some single wolves associated with packs in the deer yard for a while.
Jack Pine 7, for example, joined a pack in a county forest in February 1997. Alone after losing her mate in the park, we found her bedded down in a young cut-over forest with four other wolves. Because she had been there for a few days, we expected to find a deer carcass. The snow was deep enough for snowshoes, but packed from a thaw. With us was Bob Chambers, a biologist from New York State interested in the ability of canids with different-sized feet to handle deep snow. He speculates that coyotes, with their small feet, can live in the Adirondacks, where snow is even deeper than in Algonquin, because periodic thaws condense the snow. By his reasoning, long periods of deep snow may help keep coyotes out of Algonquin Park.
Jack Pine 7 and her new associates had bedded in the open on snowy knolls, typically good vantage points. After counting the sites, we made a large c
ircle, noting every wolf track going in or out. The next day we returned and followed the pack a few kilometres as it skirted fields and travelled through forest patches. This was a temporary alliance; by spring Jack Pine 7 was alone again.
The other three instances of wolf concentration, in the Northwest Territories and Alaska, took place at caribou or bison aggregations. Similarly, grizzly bears leave their territories to come together at salmon runs in British Columbia, Yukon, and Alaska. Both grizzlies and black bears are well known to converge at garbage dumps. Spotted hyenas and silver-backed jackals take advantage of migratory herbivores on the Serengeti Plains. Coyotes come together to scavenge winter-killed elk in Wyoming and were reported foraging at a cattle-feed yard in Arizona. Once, Mary and I studied radio-collared red foxes that showed the same territorial breakdown we were seeing here by gathering each winter at a dump along the Alaska Highway. Local abundance of food induces many predators to leave their territories and congregate without displaying obvious aggression to one another. With abundant prey, there is simply no reason to pick a fight.
Was prey abundant for wolves in the Round Lake deer yard? While deer were plentiful, they may have been difficult to catch. Some biologists, most notably Mike Nelson and Dave Mech in Minnesota, concluded that deer yarding evolved particularly as a defence against predators. Encumbered by deep snow, lone deer are vulnerable, but in groups, an individual may benefit as it does in all herd species, by earlier detection of predators and lowered odds that it will be the one singled out for pursuit. If deer yarding reduces predation, then the apparent abundance may be only an illusion.
Wolf Country Page 25
