Lupine Behavior
(Part Two)

Lupine Ethology (Posturing)

eˇtholˇoˇgy n., the scientific study of animal behavior, especially as it occurs in a natural environment.

There are bunches of differing behavior postures that are recognized by Wolf experts. These behaviors, the study of which is called "ethology," have been classified into a list that is used by observers to standardize their notes. These ethological lists are sometimes called an "ethogram." The following is not a complete list, as there are probably hundreds of observed behaviors, but it should serve as a practical guide for those wanting to better understand lupine behavior. For a more comprehensive "ethogram," you might consider seeing Wolf Park's bookstore.

Ambivalence Display

An ambivalence display may be enacted when the Wolf is confused, afraid, or trying to warn off an intruder or submissive. The Wolf will bristles his or her pelt (raise hackles) in order to appear larger and more threatening, at the same time the eyes will take on an angry wild expression and the lips will curl back to expose the fangs and gums. It is felt that red is a threatening color in nature, thus baring the gums and tongue, which is pressed forward between the incisors during the display, makes for an especially effective threatening appearance. The purpose of this display is to look dangerous... it is a warning meant to avoid violence, not to incite it... and is not used during hunting or stalking, as prey animals are never warned, just attacked, killed, and eaten. Ambivalence display is often implemented during the defensive threat posture and during a dominance display in order to demand the respect of submissive Wolves.

Many experts consider the Ambivalence display as one of mixed emotions; where snarling and bearing of fangs represents aggression, and the extending of the tongue is a simultaneous act of submission. This, of course, is the very definition of ambivalence, but I have difficulty agreeing with this opinion, and think of this posture as more of threat display.


Defensive Threat

This is a bodily posture where the Wolf crouches and prepares for possible attack of an intruder or rival. It is a condition of readiness, more or less devised to warn off or threaten a possible combatant. The Wolf assumes an ambivalent facial expression while attempting to stare down an opponent. If pressed during this posture, wolves will generally resort to violence in order to defend themselves or take corrective action against subordinate pack members.


Dominance Display (See Submission, passive)


Dominant Parade

This is posture is generally implemented by an alpha male to show off his rank to subordinates. The Wolf parades around in proud fashion with head up high, ears forward, eyes secure and direct, and tail raised or level with the spine. An alpha Wolf who approaches a subordinate in this way expects respect and is often rewarded by an act of active submission to prove one's devotion.


Fighting, pin down

Wolves often spar, especially while they are pups and yearlings to establish place in the pack hierarchy. One of the most detrimental occurrences for a Wolf during a fight is to get brought down and pinned. During ritualistic combat this usually loses the match and proves the standing Wolf as dominant, but in deadly combat it can expose vital areas of the body, like the throat, leaving it prone to a killing bite.


Fighting, Snapping Attack

A snapping attack is an aggressive forward rush, where the Wolf is crouched with tail cocked, lips pulled back, fangs bared, ears forward, and eyes wild and threatening. The snapping attack rarely makes contact, however, and usually comes up short so that the teeth merely snap together with a loud clap. This posturing is more threat display/warning than actual attack, thus it is often referred to simply as "snapping" or "snap" behavior. This posture is often used during "dominance" where an alpha is attempting to regain control of subordinate Wolves.


Flagging

Flagging commonly occurs when a bitch enters estrus and is ready to mate. She will position her rump before the dog she wants to mate with and lift her tail up and to the side, offering her anus and vulva for him to sniff and lick while gazing back at him with a relaxed demeanor indicating her willingness to receive his attentions.

Another common situation, in which a male Wolf may flag, is when he presents himself so that another Wolf may sniff his anal scent glands during greetings or certain displays of dominance. Subordinates sometimes display their under-tail to dominant Wolves while standing, but most usually during acts of passive submission, while lying on their back.


Hunting and Stalking

When Wolves approach their prey they generally try to gain surprise by looking small or crouching below the level of the grass. Also, when making a snapping attack it benefits the attacker to maintain a low center of gravity so as not to loose balance and fall victim to a pin-down. To achieve this end, Wolves generally crouch and adopt a somewhat defensive posture as they approach their potential opponent. Note that the snarling Wolf in movies, looming around their victim is not only vilifying, but in general totally erroneous. Wolves only display ambivalence when they are attempting to avoid conflict and warn off a potential opponent. When Wolves hunt they seek every advantage of stealth, for their purpose is not to warn away, but to win a meal for themselves.


Mating Behavior

Mating behavior may be broken down into three distinct divisions:

Meeting/Bonding: A male and female meet. They stand shoulder to shoulder, not gazing directly into each other's eyes, but in a kind of mutual non-threatening dominance stance; both seeming hesitant to back off or advance. Within a few moments one or the other will break the ice and attempt a muzzle sniff, perhaps followed by tongue-flicking and/or nuzzling the other's cheek or lips. There are often somewhat exaggerated, stiff, reiterations of the non-threatening dominance stance. Eventually the Wolves relax; determining that no threat exists. Bonding then begins with running, playing, mutual nuzzling, licking of lips and noses, grooming, parallel walking, sniffing of each other's anus, genitals, and urine, and double marking; where the male urinates on the female's mark (urine and scat), or visa-versa.

Courtship: Intensified bonding behavior with the addition of one Wolf standing over the other, playful attempts at mounting from the front or side (sometimes called "riding up"), actual licking of the anus and genitals, parallel running which often incorporates nuzzling under the other's jaw or ear while in motion, pricking ears sideways while in each other's presence. The female generally enters proestrus during this time; her vulva swells and often bleeds. The male's arousal increases, becoming more apparent as his testicles grow more apparent in the scrotum.

Some notes on canine estrus: Estrus begins with a period of bleeding and frequent urination called proestrus. This lasts for 7 to 10 days during which the bitch's vulva will swell significantly. Following proestrus she will enter actual estrus (often called heat) and bleeding may be replaced by a clear or yellowish discharge having a very erotic effect on the dogs. Nuzzling or sniffing the croup of her tail may cause her to flag; that is, lift her tail up to the side. Estrus lasts 4 to 7 days, and assuming she has selected a mate she will not only be receptive, but insistent that he mate with her over and over as many times as possible. The final stage of estrus is called diestrus, and like proestrus lasts 7 to 10 days. During diestrus the bitch will be unreceptive to mating and may snap or sit down if a dog attempts to mount.

Copulation: Copulatory behavior begins when the female enters estrus. She will flag the male and assume a posture sometimes called lordosis, where the hind legs are stretched back, the tail is averted, and the back is arched to lower the vulva to an optimal angle of entry for the male's penis. The male will then mount from the rear and grapple the bitches hips in his forepaws, pulling her back as he thrusts and penetrates her vulva with his penis (called intromission). The male may alternately dance his right and left hind paws excitedly as he works his bulbus glandis into the bitch's vagina; there her vaginal sphincter muscles clasp onto his penis, the two Wolves tie, and insemination is achieved. The Wolves may remain in a copulatory tie for several minutes, during which they may stand tail to tail or lay together, licking and nuzzling each other.


Play Bow

Wolves love to play; it's one of their favorite group activities, right up there with hunting as a group. The play bow is an invitation to other Wolves to romp and play. The play bow is a deep forward bow, kind of like stretching, but there is no yawn, just an occasional woof-like vocalization with paws stretched forward, rump raised high, and tail straight out or wagging.


Running, fear

When a Wolf is afraid and runs to escape danger, the head is lowered, the tail is tucked, and the ears are laid back.


Running, play/hunting

When Wolves run while playing or chasing pray they tend to be relaxed and cheerful, with head raised, ears in a somewhat neutral posture, and tail out straight or free hanging. Wolves love games of chase where they race each other. The lead Wolf in such games often finds his or her tail grabbed by the "tailing" Wolf.


Scent Marking

Scent plays an important role in lupine communications. A wolf is capable of smelling the details of an odor with a dozen times more accurately than a domestic canine, and probably a hundred times more accurately than a human being. Wolves have good vision, but really don't trust their eyes when identifying each other; they trust their sense of smell though, and rely on it to identify their territory and each other.

Wolves scent mark by urinating on a tree stump, a bush, or over another Wolf's mark or scat. Both males and females will mark by lifting a hind leg while semi-squatting to releasing a squirt of urine containing their own individual musk. Special anal scent glands also excrete an individual's scent onto their scat as they defecate. Other Wolves smelling these marks will know where a particular Wolf has been and the extent of his or her territory. Wolves know these scent marks so well that they may even know where a Wolf has been and what a Wolf has been eating during the customary genital sniffing that occurs while greeting each other.

Scent marking is quite important during mating season, when the male will mark over the female's urine and scat, and visa-versa, to indicate their mutual scent to others as being mated. This procedure is often referred to as "double-marking," and is accomplished by following the female around and immediately lifting leg and urinating over where she has recently defecated or peed. Double-marking is a very important aspect of pair bonding, and it is thought that the pair comes to associate each other's union through the consistency of their mutual marks; this may well be a kind of lupine wedding band made of golden fluid, rather than golden metal.


Scent Rolling

Scent rolling is likely the way Wolves tell each other where they've been and what interesting smells they've discovered. Wolves often find things with unusual odors, and upon deciding something is pretty neat, will flop down and roll their whithers against the whatever-it-is to collect a sniff sample. Wolves not only collect carrion odors but also the scent of dung, flowers... almost anything the smell of might impress family and friends. It may be as simple as this: lupines enjoy wearing their style of perfume just as some humans do.


Stretching, yawning

Stretching is initiated by a low forward bow with paws straight ahead, toes spread, rump raised high, and tail raised or back straight. Then the position is reversed so that the fore legs and head are raised, hind paws are pushed back, paws are curled under, pads sometimes skyward, and the tail lowered or pressed to one side. The stretch is usually accompanied by a yawn.


Submission, active

Active submission often looks like a subordinate Wolf begging the dominant for forgiveness. The submissive assumes a crouching posture with curled down rump and tail tucked and/or wagging. Then the submissive nuzzles and licks at the dominant's chin, lips, nose, and muzzle as is often seen in ritual greeting. The dominant will usually gaze ahead, raising the muzzle while accepting this display of respect and/or affection, with a bristled, but cool, appearance. This posture likely represents bowing and flattering the king in order to stave off possible disfavor.


Submission, passive (Dominance)

Passive submission takes two modes, rolling over and standing. The most common passive submissive posture is indicated by the subordinate Wolf rolling over onto the back and presenting the belly to the dominant Wolf. The submissive usually folds the paws across the chest and lifts the hind quarters. Occasionally, if the dominant Wolf is being quite aggressive, the subordinate may urinate. The submissive Wolf's tail may or may not be tucked. The dominant Wolf then stands over and sniffs and/or licks the muzzle, throat, genitals, and anus of the prostate Wolf. A less common mode of passive submission is very much like active submission, minus the nuzzling and licking activity. The subordinate Wolf tucks rear and tail and crouches down with ears and muzzle lowered.



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