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This picture depicts agonistic behavior. Photo courtesy
of Monty Sloan. For permission to use or for more information |
Social Organization
Gray wolves are very social animals, mainly living in packs of 5 to 10 individuals (Smith, 2002), although pack size is highly variable depending on environmental factors including ungulate availability, neighboring wolfpack density, and temperature. The pack is led by the alpha pair, one mated male and female. The rest of the pack typically consists of their offspring and any unrelated immigrant wolves that have been accepted and incorporated into the pack.
The dominance hierarchies are held in tact by the alphas. The alpha male is usually the dominant member of the entire pack; and there are two hierarchies that are established--one for males and for females (Smith, 2002). In case there is an injury or illness that claims the alpha male, the beta male is able to step up and fill in for the alpha male (Smith, 2002). However, unrelated immigrant wolves might be able to join the pack at this time and assume the leadership role to prevent the alpha female from forming an incestuous mated relationship with one of her kin (Smith, 1997). This social order is maintained through agonistic behavior.
Agonistic Behavior
Dominant individuals will demonstrate their authority over subordinates through postural cues and facial expressions (Smtih, 2002). Common threatening displays include showing the teeth, raised hair on the back, pointed ears, while one of the most common submissive movements is to show the belly to the opponent. Agonistic behavior actually works to reduce the amount of aggression within the pack. If there were less clarity in the dominance hierarchies, more fights would break out between members in an attempt to gain right to the privileges that dominant individuals enjoy. These rights include coordinating the group hunts and then eating first at the kill site. The young must be taught at an early age which behaviors are not tolerated by the older members. Until the young learn these social cues, they will not be able to effectively participate in group hunts or ritualized fights that will determine dominance amongst their kin.
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This picture depicts agonistic behavior. Photo courtesy
of Monty Sloan. For permission to use or for more information |
Communication
Howling is used as a territorial defense mechanism and for avoiding certain dangerous situations that a wolf might encounter (Harrington and Mech, 1979). When there were two packs that approached a common area, like a buffer zone, and when a pack returned back to an area that had not been used for a while, there were high howl reply rates. This is seen because scent marks have lost their maximal effect by this time and/or in these areas, so the wolves must use other forms of marking to assert their territory. The difference in these two types of territory marking is that scent marking is long-term and site specific, while howling is more immediate and long-range.
When wolves are "lone" or separated from their pack they generally are not as likely to respond as those in the pack (Harrington and Mech, 1979). If a wolf is separated from the pack, the howling reply rate depended on the social role of the wolf. Alpha males were the only adults to ever reply when alone. The alpha male was also the only adult that would approach the howler, every other adult usually retreated upon hearing the howl. Lone wolves, the authors suggest, try to remain low profile because they are surplus animals in a territorial population.
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Photos courtesy
of Monty Sloan. For permission to use or for more information
about wolf photographs please write wolfparkphoto@aol.com
It has also been found that howling reply rate varies throughout the year (Harrington and Mech, 1979). There were more responses in the mid-winter season during the breeding season. Through the whole year, there were high howling reply rates when the wolves were attending kill sites. The more food there was at a site, the higher the reply rates. This provides strong evidence that howling is used as a sign of territory maintenance.
Other functions of howling include locating and/or greeting one another, or an alpha calling the group together either for a hunt or to meet at the rendezvous or den site (Smith, 2002).
Another type of communication that establishes boundaries is scenting (Zub et al., 2003). This study investigated different types of territory markings, specifically urine and ground scratching in wolves from Poland. It was determined that there was a seasonal variation in urine and ground scratching behavior, with the highest amounts concentrated in the winter months (October through March). The peak came at the height of the mating season, in January and February. The densities of the the markings were the highest around the periphery of the territory and also near the very center of the territory. The alphas were likely to mark the boundary edges when patrolling the territory. The high number of markings at the territory center are likely due to the closeness of the den. This area is extremely valuable to the owners, so they will actively defend this area the most heatedly.
