Adaptive Strategies of wild wolves in the Bow Valley of Banff National Park

A review of wolf behaviour patterns in a human dominated environment

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Juvenile wolf and tourists
© Peter A. Dettling,

Günther Bloch & Dr. Mike Gibeau, 2010

There has been an ongoing debate among scientists, wildlife managers and the public about what is considered “wild”. However, “wild” is a human concept. In reality, individuals of many species can display a wide variety of behaviours. Some of these behaviours fall into the category of what we concider “wild” and some do not.

So what is "wild"?

In African national parks for instance an entire industry has been build upon close observations of all types of “wild animals”. All of these observations have included countless encounters with predators such as lions, leopards or African wild dogs. The phenomena of wolves adapting to a human dominated environment
and/or interacting with people, is also not new.

In fact, there are many documented examples of wolves coexisting with humans around the world. Here are some remarkable examples:

  1. Wolves in Africa: Over the last decade, wild Etheopean wolves were closely observed on a daily base by biologist D. Mcdonald & C. Sillero-Zubiri (2006). These unique wolves did not show significant signs of fear and adapted well to the regular presence of cross country vehicles. The wolves were hunting small prey species and even interacting with livestock guardian dogs in front as witnessed by field researchers.
  2. Wolves in Europe: In most European countries, wolves and other predators live in cultivated landscapes that are not considered “wilderness”. Zoologist E. Zimen (1986, 1990) studies in Italy showed wolves drinking well water in villages. Erik Zimen (2003) also observed “wild” wolves playing in the cornfields near the city of Leon, Spain, while tractors and farmers were present.
  3. Recently, there have been several TVreports from both Romania and Germany documenting some astonishing adaptive strategies of wild wolves. The members of a Romanian wolf pack regularly traveled through the suburbs of the city of Brassov as part of their normal routine. Likewise, it became a well known fact that several German wolf packs established their territories on military base properties. German wolves were not only traveling along coal slag heaps and villages, but also scent marking roads as part of their home ranges.
  4. Wolves in Canada/USA: Over three decades ago, field biologist E. Grace (1976) documented prolonged encounters with wolves near the Eureka weatherstation on Ellesmere Island. J. Halfpenny (2003), D. Smith & colleagues (2002-2008) as well as R. McIntire (2005) have chronicled a highly visible wolf population in the Yellowstone ecosystem. This has led to not only thousands of people watching wolves, but these wolves adaptation to humans as well. In Denali National Park, wolves also use human infrastructure (including roads), making them highly visual to people (Mech 1998).
  5. Wolves in Captivity: In the words of ethologist E. Klinghammer (2001), even wolves living in enclosures (that have been tamed by, and socialized with humans) should still be referred to as “wild”.
  6. Bow Valley Wolves: It is widely accepted among scientists that habitat dictates behaviour (Hummel 1991, Mech 1988, Hebblewhite 2000, Bekoff 2006, Mech & Boitani 2003). Consequently, there can be no doubt that the wolves of the Bow Valley show a high degree of behavioural flexibility. Over decades, their habitat specific behavior routines have been strongly influenced and shaped by a human-made infrastructure.

So the question, whether Bow Valley wolves should be considered “wild” or not can easily be answered: Yes, they were and still are wild predators.

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Summary of the history of wolves in the Bow Valley:

It is the understanding of the authors, that the Bow Valley in Banff National Park (BNP) should not be considered “wilderness”. Like other parts in the world, the Bow Valley is characterized by human infrastructure and a multitude of human activities. More than any other area in BNP, the Bow Valley is fragmented by the Highway (TCH), the CP Railway (CPR) as well as the Parkway (1A), many secondary off-roads, powerlines and campgrounds (Callaghan 2002). P. Paquet (2009) suggests, that wolves in the Bow Valley must contend with multiple threats to their livelihood. In essence, he maintains this impoverished environment is a “wilderness ghetto” for wolves.

By the late 1980’s BNP Wardens verified that two or more wolf packs had become established in or near the central Bow Valley (Mickle et al. 1996). In winter 1987, the warden service radio-collared 1 adult female wolf in the lower Bow Valley (Paquet 1993). Rick Kurnelius was the first to observe a wolf interacting with people on the Parkway. In 1987, he observed a young female wolf known as “Gaby” following the road maintenance crew in a curious but not aggressive manner. In the early 1990’s, as wolves were becoming more common, a section of the Bow Valley Parkway became known as “wolf alley” due to frequent sightings on the road.

From 1995 onward, the pack dynamics of both the “Castle Mountain Pack” and the “Spray Pack” changed, resulting in only one pack known as the “Bow Valley Pack”. All these wolves were observed using the railway regularly as a travel corridor (Callaghan 2002). Additionally, wolves traveling the Parkway were documented on a nearly daily basis (Bloch 2002, 2004). Pups were raised in the “traditional and cultural” environment of their parents and emulated established patterns of behaviour (Paquet 2009). All these observations included passing parked cars, as well as both walking and resting on the roadway. The wolves appeared not to discriminate among various vehicles, displaying these behaviours around the parked cars of visitors, CP, powerline and gas maintenance crews (Bloch & Bloch 2002).

In addition to the “Bow Valley Pack” another group of wolves emerged in 1999 and became known as the “Faireholme Pack”. As theses wolves adapted to human presence, they too started using human infrastructure including roads and rails as travel routes. The Faireholme wolves even were hunting and killing so called “town elk” very successful. By 2002 both packs had collapsed due to high mortality rates and a low prey base. In 2009 the last original Bow Valley wolves were displaced by a new group, later known as the “Pipestone Pack”. Interestingly, this pack has also adapted to the human dominated environment extremely quickly and is now using the Parkway, railway and other infrastrucure on a regular basis.

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Understanding behavioural dynamics of wild wolf packs:

D. Mech (1999) described the social structure of wild wolves as “parent-offspring-dominance-systems”. Life in social groups holds a variety of benefits such as protection from other predators, cooperative defence of food, and collective rearing of offspring as well as the opportunity for long-ranging social relationships (Fengler 2009). For the same reasons, Bow Valley wolves also tend to build family groups so that they can profit from them.

Use of technology such as radio-collars and even remote cameras have helped provide some ecological insights, but the important and intimate details that are the essence of behaviour are best revealed via direct observation (Paquet 2009). It is widely agreed among scientists that long-term ground observations are the basis to understand the social organization of wolf packs, to determine the sex, social status, colour, individual fitness and character of each adult and juvenile wolf (Murie 1985, Coscia 1991, Carbyn et al. 1993, Mech 1998, Bloch & Callaghan 2000, Peterson 2008, Smith et al. 2005). Therefore, we have been involved in direct behavior observations since summer 1992 under the direction of Paul Paquet, from 1995 under the direction of Carolyn Callaghan (Central Rockies Wolf Project) and since 2003 with the carnivore specialist Mike Gibeau.

Our most important finding is that the social organisation of Bow Valley wolves did not conform to the generally accepted and conventional view of rigid and hierarchical group relationsships. Infact, dominance interactions occurred frequently between females and males (Bloch & Dettling 2009, Paquet 2009). Even breeding females that were no longer fertile maintained their high ranking social status while allowing other females in the pack to produce pups. Individual wolves responded differently to roads, railways and wildlife crossing structures. These responses were developed through different social bonding with breeding wolves (parents), early experiences with roads, railways and through individual temperament. The movement patterns of the wolves in the Bow Valley that consisted of individuals with variable behavioural character types were not always cohesive (Bloch & Bloch 2002).

We have documented many changes in wolf population dynamics, improved our understanding of the adaptation and/or habituation process in wolves living in areas close to humans and their vehicles, and contributed to knowledge of the effetcs of human activities, infrastructure and cars on movement patterns, hunting patterns and mortality risks (Bloch 2002, Bloch & Dettling 2009).

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The difference between adaptation and habituation:

Even today there seems to be some confusion among park personal, wildlife managers, photographers and the general public about the terms “adaptation” versus “habituation”. However, German ethologist D. Feddersen-Petersen (2004) describes:

a.) adaptation as “a characteristic which increases individuals’ overall fitness through adjusting or conforming”
b.) habituation as “a learning process, in which individuals stop reacting to stimuli that have no consequences”.

In this context, the Bow Valley wolves, using human infrastruture, are often wrongly classified as being “too habituated”. In fact, what we are seeing is adaptation to their specific environment. According to wolf biologist D. Smith (2003) central to wolf management is an understanding of wolf behaviour, especially what constitutes normal versus abnormal. P. Paquet (2009) suggests that “individual wolves are unique in their behaviour and habits, the actions and traditions of wolf families reflect the collective interactions of these unique individuals”.

According to D. Smith (2003), “Wolves living in fragmented habitats, or areas with partial to frequent human presence, generally show a high degree of tolerance towards human activity and infrastructure, but still exhibit avoidance and fear in direct encounters. In such areas, wolves traveling near human developments, using roads as travel corridors, and showing a certain level of curiosity towards human activity, should all be considered normal wolf behaviour, because these behaviours reflect wolves’ natural ability to adapt to their surroundings”.

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Why are the wolves of the Bow Valley behave as they do?

Based on nearly 20 years work and over 9000 direct oberservations of behaviour patterns of Bow Valley wolves, we conclude that their behaviour has been shaped by the infrastructure of the Bow Valley. Between 39-59% of all direct behavior observations and sightings were of wolves in proximity to human infrastructure such as the Parkway and railway (Bloch & Dettling 2009). We can draw a number of general conclusions from these observations:

  1. Wolves often are described as “intelligent” animals (Bekoff 1977, Brandenburg 1990, Fox 1978, Mech 1977, Feddersen-Petersen 2008). However, “intelligence” is more complex than most people believe. Based on K. Kotrschal’s (2003) findings, there is not only social intelligence, but also “collective, emotional and habitat intelligence”. Adult wolves in the Bow Valley pass this “habitat intelligence” (including their adaptive behaviour strategies in using human infrastructure) on to young wolves over multiple generations as a form of “landscape mapping”
  2. K. Lorenz (1961, 1970) originally developed the conxept of social inprinting. A. Hess (1973) and E. Klinghammer (1994) later expanded the concept to include “food and habitat inprinting”. In the case of the Bow Valley wolves all, of their den sites are close to human infrastructure. This exposes wolf pups to the sights, smells and acoustics of humans at a very early age. Coppinger et al. (2000) found that by the age of 18 weeks, 80% of all adult characteristics were already “hard wired” in the brain. It is no wonder that Bow Valley wolves are accustomed to the presence of people, cars, trains and roadways.
  3. Wolves are forced to consume a lot of energy while traveling through deep snow (Pulliainen 1982). As part of adaptation to their environment, Bow Valley wolves take advantage of a number of energy saving opportunities. In the winter any smart wolf would not travel in deep or wet snow when they can save engery by walking on the road. Furthermore, in the late spring and early summer, wolves have been observed using roads and rails during the high water of spring runoff (Bloch & Dettling 2009). Lastly, because of the predominating influence of a low density prey base, wolves especially in the western part of the Bow Valley, must travel long distances. The older wolves get the more they maxamize use of human infrastructure to minimiye their energy use.
  4. Ethologists and behavioural biologist separate animal behaviour into two general personality types: the A type and B type (Bibikow 1988, Wilson 1998, Bloch 2004, Ganslosser 1998 + 2007, Bloch 2007). The A type is characterized as “bold, extrovert and daring”. The B type is characterized as “shy, introvert and reserved”. This explains why some wolf individuals are nonchaland around roads and people, while other individuals panic when contronted with new situations. In the same manner, some indivuals in the Bow Valley will walk right past parked car (A type), while others would detour around them (B type).
  5. It is recognized that wolf packs are not led exclusively by the so-called “alpha male” (Mech 2000, Bloch 2001, Smith 2002, Ganslosser 2007). To that end, a strict linear social rank order was not evident among the Bow Valley wolves. Social status was not related to sex, with females functioning as group leaders and acting as the primary decision makers within packs (Paquet 2009, Bloch & Dettling 2009). Given this, all combinations of age, sex and social rank could be perceived as “leading” the group in proximity to human infrastructure.
  6. All canids use scent marking as a primary means of communication. Bow Valley wolves often use human infrastructure to scent mark and delineate their central territory. These chemical messages are crucial to establish the Parkway as part of the territory for other wolves, coyotes, foxes and domestic dogs. They also more typically scent mark natural objects such as rocks, bushes, trees and deverging pathways.

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Aggression or not?

Our interpretation of “aggressive behaviour” in wolves has been confused or misunderstood with “predatory behaviour” in the past. There is a body of literature describing “functional systems” in canids (Immelmann 1979, Trumler 1987, Feddersen-Petersen 2004, Kappeler 2006, Zimen 2003). “Aggressive behaviour” is generaly related to the functional system of defending offspring, food and selfdefence. These three classic forms of aggression can only be intensified by the phenomena best decribed by behaviour biologists as priming (Ganslosser 2007). Furthermore, these forms of aggression may be both “interspecific” (aggression against non-group members) as well as “intraspecific” (aggression among group members). By contrast, predatory behaviour is related to food acquisition and has nothing do with aggression (Ganslosser 1998, Zimen 2003).

In addition, there are functional systems related to territoriality. Wolves tend to aggressively defend their core home ranges (preferred resting and sleeping spots, rendezvous areas, territory border lines) against competitors such as bears, cougars. coyotes and foxes. However, all of these functional systems come with a clear distinguishable set of body language (Ohl 1999, Feddersen-Petersen 2008). This is the reason why people need to understand and recognize differences in body language in order to correctly interpret wolf behaviour. For example:

  1. Wolves chasing prey are not aggressive! Any wolf approaching a prey species will show an “automatic” series of predatory sequences (eye-stalk-chase). It is important to note that predatory behaviour always comes along with a typical “streamlined” body postures (hackles and tail down, no stiff body posture, no growling or snarling).
  2. When a wolf approaches people it does not necessarily mean it is being “aggressive”. To the contrary, most of the time, it is often a young wolf who behave this way out of curiosity. Its body language is neutral and “relaxed” (hackles and tail down, no stiff body posture, no growling or snarling). However, it may let out a short “wuff” to express alarm in an unexpected situation.
  3. If a person were to approach either a carcass or a den site area, a wolf could display an antagonistic body language of aggression (hackles and tail up, stiff body posture, growling or snarling).
  4. Wolves that become “food conditioned” to human food or garbage are a serious problem that can result in aggression towards people. There is a progression of behaviour that starts with begging and may lead to demanding food in an aggressive way.

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Mortality - the down side to adaptation:

Many wolves have been killed over the years in the Bow Valley because of interacting with human infrastructure (TCH, CPR, Parkway). The situation is complicated by wolves accessing the TCH through holes in the existing fence. Hunting or chasing small prey (snowshoe hare, squirrels etc.) is one of the reasons why wolves continue to find holes in the TCH fence (Bloch & Dettling 2009). However, Bow Valley wolves’ familiarity with using human infrastructure elevates their probability of being struck by a vehicle or train. Even though these wolves are completely familiar with roads and vehicles, they do not pay attention to traffic while hunting (Bloch 2002). This single minded focus on hunting is hard wired, once the sequence is initiated (Ganslosser 2007). This is exacerbated by low prey densities as wolves in the Bow Valley have few opportunities to hunt large prey.

In the past, hazing and aversive conditioning have complicated both traveling and hunting as little or no attention was given to individual behaviour character types. Hazing “A type” wolves can be successful in driving them away from the Parkway, but it can also increase their mortality exposure to the railway and TCH. It is typical for “A type” wolves to become totally confused when confronted with a situation (in this case hazing) they can not control.

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Final remarks:

Adaptive behaviour is wide spread in many species around the globe. We have accepted this adaptability even in the Bow Valley with some species such as coyotes (Gibeau 1993). So why is there so much misunderstanding when it comes to adaptive behaviour strategies in wolves? One possible explanation is that wolves have become a symbol of our human notion of “wilderness”. The literature is full of references of wolves being an indicator of wilderness (Zimen 2003). Of course, wolves can live anywhere, as long as they are given the opportunity to adapt to their surroundings.

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