Behaviour
Behaviour
Generally, canid conspecific behaviour involves a loose ranking system of dominant and subdominant individuals and a wide range of postures and behaviours in order to communicate information about rank or intent. C. lupus, C. latrans, and V. vulpes are all to some extent territorial, and extensively employ scent marking in the definition of boundaries.
The Canis species also howl to advertise their presence in and ownership of a territory. Latrans primarily howls individually, a high-pitched yelp with repeated intermittent barks throughout, and holds its head thrown far back, almost folded back into its shoulders, when it does so. Lupus howls individually to locate its pack or to advertise its readiness to mate, and group-howls to advertise the presence of the pack in a territory or to group the members together before (and sometimes after) a hunt. The head in lupus is held horizontal or slightly above, and there are few or no yips or barks in the howl. Vulpes does not howl, but will hiss and bark, and occasionally "screams" to scare off intruders.
The Canidae have the richest vocabulary of calls of all the carnivores. Besides the howl, the social species have a range of calls designed to increase social distance (snarling, growling, hissing): these are emitted rhythmically with an increased respiratory rate, and tend to be choked off at the end. Barking tends to be a call of warning in all three species (the mother wolf may have a distinct "warning bark" with which she warns pups away from danger), although modulations may be used as identifiers or greetings. Calls used to decrease social distance, such as whining, mewling or whimpering, are generally related to calls used by infants to elicit parental care and usually express submission during courtship, greeting, or aggressive encounters.
Scent plays an important role in communication, as expressed in the vast array of scent glands located on the head and tail regions (the most common places for sniffing) of canids. Mutual sniffing, first of the head and then of the anogenital regions, is characteristic of meetings between canids. Tail position may play a part in scent communication: a dominant animal raises its tail to facilitate scenting by the other animal, and a submissive animal will lower its tail to mask its scent. Bitches in heat carry their tails constantly raised and turned to one side, in effect broadcasting their condition. The violet gland on the surface of the tail is also checked during social sniffing.
As previously mentioned, all species scent-mark territorially, primarily
by spraying urine on the object of interest. Familiar
landmarks are routinely sprayed on a "patrol" around
the territory, but if something new or interesting appears it may be sprayed to
indicate ownership by the spraying animal. Dominant animals scent-mark most often and
most visibly; subordinate animals may squat like puppies to urinate and rarely mark objects as their own. Dominant animals usually
lift a leg and stand with tail raised during urination (dominant males will cock a leg high; dominant
females still squat, but often with one foot off the ground), and follow urination with a succession
of raking movements of the hind legs and general aggressive behaviour.
Scent-rolling is a peculiarity of canids which is thought to be a means of bringing information about interesting scents back to conspecifics which may not have accompanied the animal to the oderiferous site. All canids scent-roll to some degree: the animal first puts its nose in the smell and then slips down onto one shoulder, sliding first one flank through the smell and then the other, pausing in the middle to vigorously rub its back into the scent. Given the usual pattern of social sniffing, i.e., starting at the head and moving down the flank to the tail, a greeting animal would get a good sniff of everything the other animal had rolled in on its way from one end to the other.
Aggressive behaviour is shown almost exclusively to conspecifics: either as an attack against
an intruder on the individual's or pack's territory or as a move to dominate another animal. Ears are
held erect (or pushed to the side in "airplane ears"), lips puckered and closed or slightly open, eyes
wide open, hackles and tail erect, tail held stiffly. The animal carries its body rigidly, with
confidence and a stiff-legged walk, and stares at the object of aggression directly, keeping eye
contact. Two animals coming together in a ritual (rather than an attack) dominance interaction often
make a "T" formation, with the dominant animal pressing its chest into the other's flank. The
aggressor may press its chest into the other animal's shoulder or stand up with stiff forelegs braced
on the other's shoulder. Mounting is also occasionally seen as a dominance display.
Aggressive behaviour is often mistaken for behaviour exhibited during hunting. In truth, predators almost never display aggression behaviour to their prey. Hunting wolves trot or lope in a low stance with middle head carriage, a relaxed face, eyes wide and ears erect, tail extended flat rearward and waving gently, concentrating completely on their prey. Growling or snarling at the prey, as is popularly "reported" in low-budget documentaries, is rarely, if ever seen: prey is not regarded as a conspecific and communicative postures toward prey are uncommon.
Canids also employ more specialised strategies: hunting foxes may stand stock-still, listening in the direction of their prey, before leaping into the air and coming down on it. All three species have been credited with an unusual hunting tactic called the "fascination display", wherein the animal makes repeated strange and unusual movements such as leaping in the air, rolling on the ground, jumping, circling, or somersaulting, ostensibly in an effort to lure prey which cannot easily be run down within catching distance. The "play bow" (H in behaviour diagram) is a behaviour seen in many contexts, including play solicitation, but it also plays a role in hunting as a good "all- purpose" position from which the animal may easily break in any direction depending on the movements of the prey. Although this position is often mistaken for a play request or a suggestion that the hunting behaviour is less than a life-or-death ritual, the bowing animal is not soliciting play behaviour from its prey.
Defensive aggressive behaviour is displayed by animals
that would rather flee but are prepared to fight if necessary. The
animal is fearful but feels it may be able to hold its own if the
threat persists. Elements of threat and submission are combined
in their postures, or may visibly alternate as the animal "waffles"
between one option and the other. An animal may simultaneously
grimace in fear and snarl, or alternately growl and whine. The
hackles are erect, the back humped, but at the same time the
animal crouches submissively, keeps its tail and ears down, and
avoids staring directly at its opponent (more or less "J" in
behaviour diagram, previous page).
Submissive behaviour probably originated as soliciting
behaviour between offspring and parent; the movements often
parallel. Weaned pups food-beg from adults by approaching in a
submissive crouch -- back curled, head held low to the ground,
muzzle turned up -- and
licking the underside of the adult's muzzle; later in life, a crouch
and muzzle-lick from one adult animal to another is an active
submission gesture (G in the previous illustration). The
submissive animal may whine or paw at the dominant one and will
weave in and out in front of the animal's face, licking at its
muzzle. The tail is held between the legs or to one side and
wagging weakly; the eyes are narrowed and the head turned away,
the lips retracted in a submissive grin, or "fear-grimace". In
passive submission, the dominant animal initiates the submission
behaviour in the subdominant, often growling or staring to make
the subdominant animal lie down, flatten its ears, and roll over on
its back, in a purely indefensible position. The dominant animal
will often stand over the submissing one (C in behaviour diagram)
and growl at it or bite and mouth it to "remind" it of its position
in the social order.
Submission can be used obnoxiously as a means of "testing" both the patience and the rank of a more dominant animal. The subdominant animal will approach and repeatedly submit to the more dominant one in an attempt to "test" the other animal's rank ("G" in behaviour diagram).
"Play" in all three species is marked by the expression of any or all of these postures and gestures but without the sense of "formality" which accompanies non-play behaviour -- during play, subdominant animals may chase and harass more dominant ones, but when play is over, the original rank order returns. The essence of play is a relaxed face and repeated, apparently purposeless movements such as poking with the muzzle, "dancing" with the forepaws, bouncing or brief, invitational runs. Play is seen most frequently in pups of all species, but in the social animals (wolves) play behaviour may continue to a degree into adulthood. Adult wolves will invite play chases from one another with a "play face" -- lips drawn back horizontally, an open-mouthed grin, ears more or less erect, eyes partially closed and slanted -- or the play bow. Adult red foxes and coyotes exhibit playful behaviour primarily only during courtship.