The social relationships of wolves are maintained through three types of communications: vocalization, postural signaling, and scent marking.
The exact emotions prompting a howl remain unknown, but scientists speculate that solo howls and group howling is brought on by restlessness and anxiety. Howling occurs frequently during the season of mating and courtship. The myth that wolves howl more often during a full moon may have arisen from the fact that on the cold and clear winter nights during courtship, the sound of a howl carried long and far across the landscape while a full moon lent an eerie aspect. Wolves also celebrate by howling, after a successful hunt or when a lone wolf returns from its wanderings.
Wolves also bark, growl and whine. The bark occurs very rarely and when it does, it is more like a soft "whoof" than a sharp dog-type bark. Barking also never occurs continuously. The bark is associated with surprise and warning. In the case of the latter, wolves, most commonly the alphas, will often emit a bark howl (two or three barks leading into a low, bellowing howl) that serves as a warning to outside threats and intruders.
Growling occurs during food challenges and is associated with threat behavior or assertion of rights. Pups will growl when they play. There also exists a type of growl that begins at a higher pitch moving toward a whine and it usually precedes a snapping lunge at another wolf. Whines are associated with greeting, feeding the pups, play, and other signs of anxiety, curiosity, and inquiring. These are probably the most intimate sounds.
Postural communication is composed of a variety of facial and tail expressions as well as piloerection (raising of the hackles). Lunging, chasing, body slamming, fighting, and more subtle gestures are all different modes of postural communication. A submissive animal may lower the body, lower the tail, turn slightly away from the dominating animal, lay the ears back, retract the corners of the mouth in a submissive grin, or even lick the dominating animal's muzzle. A dominant wolf might stand over or ride a submissive one by straddling the reclining wolf. If the subordinate does not submit, the dominant wolf might hip slam and push down the subordinate with its paws. Subordinates may respond to this by rolling over on the back and even urinating a few drops on itself. A stiff-legged approach from an alpha is a commonly seen movement of dominance.
With respects to ear posture, there are subtleties that can be observed. Ears that are pricked and turned to the side, for example, most often can be read as, "I'm not so sure I should be doing this." Ears that are pricked and turned forward are an open display of confidence among packmates, and an indication of a stimulated prey drive if the ears are also tilted slightly forward.
Mouthing another's muzzle is a friendly gesture. Even clamping down on a muzzle isn't a gesture of anger, though it might signal annoyance. Packmates, and especially siblings, often jaw spar with one another as a means of play. Throats may also be mouthed. Raised hackles and a raised tail can indicate annoyance. The fluffing of the fur is used to accentuate certain emotions and to make the wolf look more fierce. Tail twitching can indicate excitement and this motion is accentuated by the black tip at the end of the tail. Little is known about olfactory cues associated with scent markings and other glandular secretions.
Scent marks can be divided into four categories: raised-leg urination, squat urination, defecation, and ground scratching or dirt kicking. The first category is the most important. The alpha animals (primarily the males) make this scent mark. They are made against objects above ground level to ensure a large evaporative surface (this gives a stronger odor) and to keep the mark clear of rain or snow. Squat urination and defecation impart a personal scent (as anal glands can be triggered), though these mostly function to eliminate wastes. Dirt scratching probably serves as a visual display of dominance for the benefit of the other pack members.
However, if glands in the toes are stimulated, then another olfactory message is left behind. Like dogs, wolves also roll in putrid substances. This serves some form of communication, though what is communicated is unknown. Wolves will also roll on the ground as a way of marking their scent in a location. Wolves carry on a regular pattern of scent marking, visiting each section of their territory on the average of once every three weeks. Wolves also scent mark when in the pursuit of prey. Scent marks are used to warn off intruders but they also play a role in the maintenance of a sense of spatial organization for the resident pack. The markings may also help wolves find open territories and loners of the opposite sex, so that new packs can be formed. Male wolves will inspect the anal gland with dominant males readily presenting their anal parts for inspection while subordinate animals withdraw theirs or present them only reluctantly. Females rarely engage in anal inspection except during the breeding season.
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