Another time we watched Grand 2 with the big wolf Grand 4, her mate, and four other wolves as they picked their way along the far shore of the lake. We managed to stay parallel to them, walking through shallow snow. A strong wind bent the trees and covered our movement. Eventually they lay down in the sun on the snow-covered ice, and there they spent the afternoon, getting up, walking around a bit, lying down again, as “stationary” wolves always seem to do. It was a harmonious winter scene — snow, sun, forest, wolves.
Then in January 1991 something disturbed the pack. Maybe the death of the big male, trampled by a moose, destabilized them, but that had happened a year earlier. Maybe the recent dispersal of a young, radio-collared female was involved, but dispersal of young is common and not disruptive to wolf packs. The month before, the territory of the six wolves in the pack extended beyond both ends of Grand Lake. Then, Graham found Grand 2 with two other wolves outside the park in the Petawawa Military Reserve, ten kilometres northeast of her territory. They stayed there for two days, then moved another ten kilometres southeast. By mid-February, when Graham flew in search of them again, they were back in the eastern part of their territory and we thought the trio had only been off on an extra-territorial foray. After that, however, they never travelled beyond the midway point of Grand Lake, and Grand 2 never associated with her old den along the creek. And never that winter was she with more than the two other wolves. Meanwhile, the other three wolves of the original pack stayed put on the vacated west end of her old territory.
The pack had split in two, not because it had become too large, as happens occasionally — there had been only six — but because of some internal social dynamics known only to its members. Each pack maintained somewhat smaller territories than the original Grand Lake pack, although the following summer the eastern trio expanded its territory another few kilometres to the east into the range of an uncollared pack. In a new den, Grand 2 produced pups to become a pack of five. The western trio added one disperser from somewhere, then either lost adults or produced only one pup, because by winter they also were a pack of five. Despite probable genetic affiliations, and knowledge of each other, the two packs never associated again.
For a long time the Pretty Lake pack was an enigma. We knew it was there; several times each summer we heard its howls from our campsite among the pines. When we plotted the fixes of the Jack Pine and Mathews Lake packs to the west and east respectively, there was obviously a gap between them. But the missing pack eluded us until August 1991, when a yearling female was collared and began to fill this vacant place on the map with dots.
Pretty Lake, accessible from the Sand Lake road, turned out to be near the southern boundary of the pack’s territory, which extended north to, and sometimes beyond, the fast-flowing Petawawa River. The river posed a psychological more than real barrier in summer because it could be swum, and no barrier at all in winter. The collared yearling was in a pack of four. That December, as the deer migrated away, the pack made two extra-territorial forays south into lands occupied by packs in the Bonnechere Valley. On one of these trips the trespassers killed a deer.
In mid-March, Graham plotted aerial fixes after three consecutive flights that all fell close to the same little creek buried in a deeply incised, conifer-filled valley. A stationary pack such as that normally signals a moose carcass, so Mary, Michelle, and I loaded our snowmobile and drove to the “Pretty Lake trapping road.” The snow was hip-deep, and a strong wind left the sun cold. We cut down the two-metre-high snowbank at the edge of the road to get the snowmobile up onto the snowpack, then headed off with Michelle behind me on the seat and Mary in her accustomed place on the sled runners.
Six kilometres later, Pretty 1’s signal came in weakly. Our rule is to keep the snowmobile one or two kilometres back from any wolves, so we cut the noisy engine and donned our snowshoes to continue on foot. A trio of gray jays sailed in silently to look us over, concluded we were not good for a handout, and vanished. The paired tracks of a pine marten showed where, sometime ago, it had decided that red squirrel hunting was better in the pines on the opposite side of the road. In the spruce-fir low places, the tracks of snowshoe hares revealed where they had been dancing in the moonlight. The snow had enough body to hold them up; even the wolves sank only a few centimetres. Moose, though, were going in half a metre, deep enough to make them lift their hind legs high and run with knees splayed in an awkward gait. Wolves, under these conditions, can run circles around a moose without fear of its deadly hooves.
As we continued, the signal strengthened from the far side of the same gully Graham had seen from the air. Too steep and too densely covered with small, snow-bent pines, we took off our snowshoes and slithered to the bottom. Over another ridge we followed the wolf’s tracks to a knoll where she had been lying with a good view all around, as is typical. There, she had first heard us coming and bolted away with long strides, clearing a snow-laden log with a metre-high jump.
After searching unsuccessfully for the moose carcass, we floundered back to the snowmobile. In our absence, two wolves had come up the ravine, encountered our snowshoe tracks, and turned to walk beside them. They continued half a kilometre before crossing them and heading back into the ravine. Wolves have hesitated to cross our tracks on other occasions, but we have also seen instances of wolves walking right in them or crossing without hesitation.
We drove the snowmobile south for a couple of kilometres, set off again by snowshoe, and soon found the tracks of what looked like the same two wolves. They had crossed a beaver pond and scratched all the snow off the top of a lodge, undoubtedly smelling beaver through the top vent. We have yet to find a successful excavation; the frozen mud and sticks form an impenetrable roof. Some freshly cut birches on shore indicated recent beaver activity. To get a meal, wolves must come upon beaver away from their lodges, which they do often enough, judging by beaver hairs in winter wolf scats, to make checking out beaver ponds worthwhile.
The collared wolf’s signal was still to the north, so we waited in some fir trees where we had a view across the pond. By then, late afternoon, the cold sun had disappeared, and an icy wind whipped the pines causing them to convulse in disarray. Tracks showed where a moose had browsed balsam firs, then bedded down making an ice-lined depression with its body. We watched a black-backed woodpecker flick bark scales off a red pine in endless pursuit of bark beetles. But the day was in shut-down mode.
Soon a full moon veiled by thin cirrus strands sailed into view among the tossing pines. The collared wolf was closer, but by then all we could hope to see was its shadow out on the pond. We were not going to find its moose. I howled; the wind grabbed the sound and transported it elsewhere. The wolves heard it, though they didn’t respond immediately but waited until we had started back along our snowshoe trail. Then they broke out into a wild chorus from the beaver pond where we had been, accompanied by the stereophonics of wind gusting in the pines, rising on one side of us, falling, building up again somewhere else.
We wound our way back in the moonlight, flashlights off. Twice we stopped to listen to the voice of a single wolf. No breaks in pitch, just smooth up, then down, in long, beautiful solos.
Pretty Lake wolves trespassed on occasion in Grand Lake West’s territory and in turn got trespassed on by the Mathews Lake pack. Once, a Pretty Lake wolf joined a Jack Pine wolf next door. But other than occasional winter excursions in response to migrating deer, they remained reasonably faithful to their own lands.
Our winter experiences with the Travers wolves were limited. Because they lived on the west side of Lake Travers, and only the southern tip of their territory was accessible by road, we rarely could reach them. They provided a few moose carcasses for us, and made their share of extra-territorial forays. Once, in mid-December they split into two groups, one of them feeding on poached moose in the Foys Lake pack’s territory while the other group was also off territory, twelve kilometres away. They appear not to have shared the moose, a puzzling lack of pa
ck coordination.
Then, tragedy befell the pack, repeated tragedy that made the Travers lands vacant through much of our study.
When we look at a handful of computer-drawn maps showing the locations of these wolves, most fixes fall within the loose boundaries of what we came to expect for each pack. Strings of dots occur where Graham backtracked a pack from the air, most frequently along lakes; linear Grand Lake looks like a highway. In places, concentrations of dots occur along roads, reflecting where we searched most intensively rather than wolf behaviour.
Except for the Grand Lake highway, we can discern no tendency by these packs to use distinct travel routes repeatedly, although we did for the Foys Lake pack to the south. Pack movements within territories seem to be largely random. Minimum pattern represents maximum flexibility. That undoubtedly is adaptive for wolves under the circumstances of the largely patternless and unpredictable movements of moose and deer. If wolves were creatures of habit, prey would be able to predict their movements and avoid them better.
Boundaries differed somewhat from one year to another. This tendency was partly due to the outer locations we managed to detect, but also to differences in the way various wolves used their territories over the years. The packs were definitely not nomadic. But they did not seem preoccupied with defending boundaries, either — we recorded only a few instances of boundary patrolling. If boundary patrolling was common, many more fixes would have helped define the outer limits of their territories. Instead, the points seem to peter out away from an ill-defined core. Boundaries must have been marked, or remembered and recognized, however, otherwise pack overlap would have been much more common.
Against this matrix of territoriality were frequent instances of trespass and even some cases of prey poaching or sharing between packs. Part of what we saw could be attributed to dispersing individuals with their apparent free pass to travel on any pack’s land. We tracked young dispersers, such as a young, female Grand Lake wolf who travelled widely before joining a pack to the south, and a few adults, such as Grand Lake I who wandered much of one winter before settling down in the Jack Pine pack. Until we realized that these dispersing wolves held free passes, their trespassing escapades perplexed us.
Then we developed an entirely new perspective on our population from the genetics work of Paul Wilson and Sonya Grewal. The genealogy of the population showed that we were studying one big extended family! Not only the Jack Pine pack had scattered its genes in most of the surrounding packs, but so had many others. The Basin Depot pack in the Bonnechere Valley, for example, sent three dispersers north to the Travers pack, one in 1994 and two in 1995. As in rhesus monkeys and chimpanzees, one sibling dispersing to a new pack may open the door for later ones who would have known each other back in their natal pack.
We found that we had collared eight great-grandchildren of the oldest wolf in the study, Redpole 4, who died when fifteen years old (confirmed by tooth cementum analysis), and these great-grandchildren lived in four different packs. We identified other relationships forged by dispersal between the Mathews and Pretty packs, Grand and Mathews, Redpole and Travers.… Our conclusions about the difficulty of Jack Pine wolves going anywhere without meeting kin was true for most other packs too.
The dispersal we found differs only by degree from that described by other researchers. The greater amount of dispersal in the Algonquin population served to increase interrelatedness among packs and may help explain the tolerance and lack of aggression that characterizes our population. But Dave Mech and colleagues in northern Minnesota have found more killing and inter-pack rivalries than we have, despite a great deal of dispersion in the population they study.
Perhaps the difference in our results reflects differences in prey density or ease of capture, that is, some sort of cost:benefit analysis of defending a territory depending upon resource availability. Widely accepted theory explains that when prey is sparse and scattered and takes a long time to find, the energetic cost of maintaining a territory over a large area may be greater than the benefit of keeping all the prey to oneself. Under such circumstances in other species, nomadism may occur. But, scaling upward in prey availability, a situation may arise at moderate prey density where territories can be smaller and so defended at lower cost. In that situation, the predator benefits by not having to share prey and risk prey depletion by others. Then defended territories make sense. With even more prey, however, in the face of plenty of food, there may be little to gain by defending a territory.
The Algonquin wolf system, with some evidence of boundary patrolling but little active defence, may lie somewhere between the latter two situations. Subtle differences between years or even between seasons in prey availability or snow conditions may influence the level of aggression and territorial behaviour. The population may be balanced on the edge of an indifferent or avoidance-based home-range system and a competitive territorial system, depending on immediate resource availability.
A good analogy found in ecology books for what we were seeing is a “compressible rubber ball.” The more you squeeze it, the harder it is to squeeze until you cannot compress it any more. The behaviour of a landowner can be like that. The pack may be willing to make room for others for a while, but the more land it gives up, the less it is willing to concede.
As well, wolves with different temperaments, tolerances, thresholds of cooperation and competition may come and go in a population. Individual wolves have both behavioural plasticity and personality programmed in their genes.
The early Algonquin Indians faced many of the same survival problems as wolves. Both needed to strike a favourable ratio of energy expended to energy obtained where prey was sparse, discontinuous, scattered, its location unpredictable, and its chances of escape good. Both were co-predators on moose.
One winter, Mary pieced together a picture of the lives of these native people from archaeological and historical writings. She relied especially on the seventy-three-volume Jesuit Relations written by the first priests to make contact with them. Each winter the native people left their summer fishing camps along the Ottawa River and broke up into small family hunting units that spaced themselves out across the Algonquin dome, each with its own territory. Many family groups were related through a grown son, or a brother or sister. In times of need, one family could hunt on another’s land. It was much like the system used by the northern Cree even today.
With this flexible system of land allocation, the early Algonquin people did not operate in the stereotypic ways described for wolves. Their spacing mechanism was largely a kin-based, cooperative land division with undefended home ranges.
Direct comparisons are complicated by differences in mobility, efficiency in both detecting and dispatching prey, and ability to hunt at night; wolves have the edge in most of these traits. But, despite the evidence we found for competition and territoriality among wolf packs, we have just as much evidence for a home-range system similar to that used by the early Algonquin people. Admittedly, in our study we could not identify the importance of passive forms of defence such as howling or scent-marking to keep trespassing wolves away. And accommodating the needs of one another out of compassion is more highly developed in humans than in other species.
Nonetheless, the Pretty Lake pack appears to have behaved cooperatively by leaving room for a re-establishing Jack Pine pack. Similarly, there were many instances of trespass permitted by various packs on each other’s lands. As a survival strategy, some blend of competition, indifference, and even cooperation, some ability to show one or another depending upon circumstance, may be a fundamental part of the wolf’s success. At least in the Algonquin wolf, that is the way it is.
Harmonious relationships between the MNR, the landlord at Achray, and Algonquin College, the lessee, did not last. One winter we had the place much to ourselves, then it was declared surplus. The water pump was turned off, pipes drained, generator disassembled. One winter’s day a year later, we drove into the familiar parking lot
and found the big building gone. As if it had never been there, a smooth blanket of snow extended from the pines out back where the bird feeder used to be across to the parking lot in front. We sat for a long time in the sun-warmed truck cab reliving memories. By then the focus of our winter work, and most of the wolves, had shifted to the southeast, outside the park in a much more dramatic example of wolf territorial flexibility. Everything changed, for the wolves and for us. The system of land tenure we had been observing, it turned out, was only part of the social order, and the territories more transient than expected.
THE RED QUEEN AND WOLF-MOOSE RELATIONSHIPS
Predator and Prey
Predators can kill members of a prey population, yet not depress it — an apparent contradiction. A sharp-shinned hawk pinioning a flicker, a mink eating a shiner, a wolf killing a deer all result in one less prey animal. Predators get a bad rap, however, because the dramatic act of killing does not always lower the prey population in the expected way.
Several adjustments operate in predator-prey systems to accommodate such losses. Most significant are the compensations, first explained by Wisconsin ecologist Paul Errington. For example, prey populations drawn down by predation may increase their birth rate in response to better nutrition for the remaining animals.
Wolf Country Page 11
