HYPERWOLF -- Glossary

... Aggression is not a 'thing'. It is a concept. (p. 254)

Actions scored under the general heading of 'aggression' included: circling, raising hackles, biting, pinning, lunging, retracting the lips vertically, taking of food, snapping toward, supplanting pounces, raising tail, and forelimb wrestling. 'Recipient' behavior included: hunching-arching back, wheeling and facing, gaping, rolling on back, placing shoulder on ground, tucking hindquarters of tail, fleeing, or reciprocating aggression. (p. 255)

To this category belongs all behaviours of threat, attack, combat, imposition (subordination) and defense. (p. 341)

[Those activities on the part of one individual] directed towards another individual which could lead to physical injury to the latter and often results in settling status, precedence, or access to some object or space between the two. (p. 335)

A single agonistic episode was scored when one or more items within both classes of behavior ('aggression' plus 'recipient') were observed. The episode was considered finished when neither class of behavior continued to be expressed by the animals in question. (p. 255)

a surprise attack by persons in a concealed position

[with reference to aggressive behavior, generally without body contact] sudden threat attack (p. 285)

[with reference to actions used to intiate social play in 3 - 7 week old coyote, wolf, and beagle pups] The soliciting animal approaches its prospective playmate and then withdraws; withdrawal may involve stepping away slowly, or running away for a few meters and then approaching (and withdrawing) once again; it is not uncommon to observe the soliciting animal approach, stop, and then rock back-and-forth in one spot, making intention movements of running away. (p. 324)

Galloping and bouncing gaits are said to be asymmetrical because the footfalls of the two feet of a pair are unevenly spaced in time.(p.475)

[with reference to aggressive behavior, generally without body contact] sudden real attack from some distance (p. 285)

BARING TEETH is associated with threats in both attack and defense...Its characteristics are shortening the bridge of the nose, wrinkling the skin itself and lifting the lips, so that the foremost teeth (incisors, canines and premolars) are visible. ... When the teeth are bared, the MOUTH MAY BE OPENED TO A VARIABLE EXTENT. Enlargement of the opening-angle points to increased intensity of fight excitement. (p. 96)

... the relationship of a pup's trunk to the ground. ... The pitch is described as one of 6 levels:

• Level 5 - Standing, with all four legs extended.
• Level 4 - Sitting on the hind part of the body, with hind legs folded under and forelegs extended, or crouching on forelegs, hind legs extended.
• Level 3 - Lying with all four legs folded and the ventral side of the body touching the ground.
• Level 2 - Lying on either left or right flank.
• Level 1 - Lying on the back, all legs in the air.
• Level 0 - Unidentifiable body position, always a relatively fast transition between two levels. (p. 179)

body positions [wolf pups 15 - 85 days of age]:

• on back - lying on back.
  
• body low - standing with legs not fully extended.
  
• body high - standing erect with legs fully extended. (p. 107)
  

...an individual crouches on its forelimbs remains standing on its hindlegs, and may wag its tail and bark (BECKOFF, 1977). The bow is a stable posture from which the animal can move easily in many directions, allows the individual to stretch its muscles before and while engaging in play, and places the head of the bower below another animal in a non-threatening position. (p. 420)

...the nonrandom performance of bows during ongoing sequences of social play, supports the inference that bows might serve as a form of punctuation that clarifies the meaning of other actions that follow or precede them. In addition to sending the message "I want to play" when they are performed at the beginning of play, bows performed in a different context, namely during social play, might also carry the message "I want to play despite what I am going to do or just did. I still want to play"... (p. 426)

[with reference to actions used to initiate social play in 3 - 7 week old coyote, wolf, and beagle pups] The soliciting animal crouches on its forelegs and elevates its hind-end; ...from this position the animal is able to perform a wide range of other movements such as leaping, dodging, and springing back-and-forth (p. 324)

"mutually exclusive item dimensions" where item refers to "postural components and features of the interaction that can be simultaneously present" (p. 116).

The Canadian Oxford Dictionary, 1998

The act or an instance of pursuing

[with reference to aggressive behavior, generally without body contact] Full speed chase in order to catch opponent (p. 285).

In Circling, the interaction does not move significantly in space, but centers around a single area. (p. 212)

The interactants move slowly in a slightly off-anti-parallel orientation; the head of S [Supplanting animal] is closest to the hindquarters of D [Displaced animal], separated by at least half a wolf-length. ... Small fluctuations of one or more of the relational variables are interspersed with periods of actual maintenance of all three variables ... Both interactants shift front continuously and at the same time progress forward minimally. (pp. 1151 - 1152)

[with reference to seeing this behavior during play fighting in a 27-day-old coyote pup for the first time] Clasping a peer with the forelimbs around the waist or thorax ... (p. 244)

During eating, urinating, defecating and when rolling and rubbing in a strange odor, the ears were partially flattened and the eyes either narrowed or completely closed, or open and fixed in a 'middle distance' stare or 'glazed daydream' ... (p. 60)

• HEAD (H) = 1 + 2
• CHEST (C) = 3 + 4 + 7
• FRONT PAW (F) = 5 + 6
• HIP (P) = 8 + 9
• REAR PAW (R) = 10 + 11
• TAIL (T) = 12 + 13

...the point on each animal's body closest to its social partner. This may or may not comprise physical contact. (p. 264)

stoop low with the legs bent close to the body, esp. for concealment, or (of an animal) before pouncing; be in this position.

... in a supplanting interaction, D moved away from the direction of an approach by S. (p. 1147)

Having only the digits and distal ends of the metapodials in contact with the ground when standing or moving, as for cats and dogs. (p.623)

... the length of the interaction from beginning to end in seconds. The beginning was defined as when one wolf directed an approach or looked toward another wolf within one wolf length (excluding tail). However, if the wolves were already within one wolf length of each other (i.e. eating the same carcass prior to the interaction), the beginning of the interaction was defined as the point at which one wolf initiated some action (i.e., directed gaze at or movement towards) at the other wolf. The interaction was considered terminated in one of two ways depending on the outcome of the interaction. If the wolves separated, the end of the interaction was defined as the point at which the participants were one wolf length away and no longer interacting with each other. However, if the participants remained closer together than one wolf length, the end was defined as the time when all postures, movements, and actions which characterized the interaction ceased in all participants and there was no further interaction between them. If the interaction was already in progress when the video recording began or was not terminated when the video recording ended this was noted. (p. 15)

ear positions [ wolf pups 15 - 85 days of age]:

• ears flat - ears flattened against the back of the head and rotated backward (open to rear).
  
• ears to side - ears extended horizontally to sides and opened downward. (p. 108)

[with reference to actions used to initiate social play in 3-7 week old coyote, wolf, and beagle pups] The soliciting animal approaches its prospective play-mate in a 'loose, bouncy' gait, at a speed greater than that observed during normal walking; this has also been called a 'play rush' and 'gamboling' in the non-human primate literature; during the approach, the shoulders and head are frequently moved from side-to-side in an exaggerated fashion. (p. 324)

eye positions [wolf pups 15 - 85 days of age]:

• look toward - eyes open and directed toward partner.
  
• look away - eyes open and directed away from partner.
  

[with reference to actions used to initiate social play in 3 - 7 day old coyote, wolf, and beagle pups] This action involves extension of one of the forelimbs toward the face of the other animal ... it is not uncommon to observe rapid extension and flexion of the forelimb when the animals are at a few meters from one another (in this case the action is called a 'paw intention'). (p. 325 - 326)

[A] relational configuration that is only rarely observed during extended supplanting interactions is a face- to-face, anti-parallel, at near-contact or contact configuration. ....the two wolves often rear up on their hindquarters, pushing forward with all their weight, leaning on the partner's shoulders with forequarters, and biting extensively. S [Supplanting animal] eventually pushes over D [Displaced animal], who loses balance and falls on her back or side, as S commences to bite. (p. 1164)

[with reference to an action observed only by red foxes to initiate social play] ...the animal lies flat on the ground and moves its head from side-to-side. (p. 326)

During Following, because of the parallel orientation, the animals cover much more ground, [than circling], travelling over relatively long distances with frequent turns to avoid obstacles. (p. 212)

[with reference to aggressive behavior, generally without body contact] Walking or running behind opponent, often in imposing posture, without trying to catch up. (p. 285)

...during Following, opposition is between the head region of S [Supplanting animal] and the rear of D [Displaced animal] and the interactants are separated by at least half a wolf-length. A relative orientation near parallel is maintained for most of the sequence. ... Opposition is maintained with minimal or no variation. Both relative distance and orientation show greater variations. Following involves extensive movement, and obstacles are ultimately encountered and turns necessitated. Large deviations from parallel often indicate such turns by the two wolves... The leading wolf (D) turns; S replicates that turn at the same point in space; and finally, the pair resumes following in parallel. The faithful replication of the path of the leading animal by the second wolf is striking ... The smaller deviations in relative orientation ... are the product of commonly seen small shifts of front by the leading animal (D), as if to 'look back' at the following wolf. Relative distance is typically maintained steady for short periods of time, and then is shifted to a new maintained value, but these values are always within the range of half a wolf-length or greater. Relative orientation fluctuates in the range of 0 to 45 degrees; a range distinct from that seen in Circling. Maintained points of opposition are similar in Following and Circling. ...

Following and Circling frequently alternated during an interaction. Although no systematic records of sequence duration were kept, Following tended to be longer than Circling. A particular Following pattern lasted over 30 min, and sequences lasting a minute were common. (p. 1152-1153)

1) The CONTRACTION OF THE FOREHEAD towards the ridge of the head and base of the nose. Thereby, the forehead appears to be bulging, particularly above the eyes. This shape of the forehead belongs to the expression of threat, or readiness to fight.

2) The"PULLING-APART/STRETCHING" of the forehead in the direction of: corner of the eyes-lower edge of the ears-sides of the neck. The forehead is smoothed by means of this "PULLING-APART/STRETCHING", the brows disappear, the eyes become slit-shaped. This occurrence becomes evident in all forms of insecurity or submissiveness ... (p. 97)

The foot of a pair that strikes the ground first in each couplet of footfalls is called the trailing foot; the other is the leading foot. If the lead is the same, fore and hind, the gait is a transverse gallop (horse); it is is different, fore and hind, the gait is the more maneuverable, but less stable rotary gallop (cheetah). Unless performed slowly, asymmetrical gaits increase length of stride by introducing periods of suspension when all feet area off the ground. The suspension may come when the legs are gathered under the animal (horse), stretched out fore and hind (deer), or both (cheetah, pronghorn). (p. 475)

The galloping canid has two unsupported periods, one when the body is flexed and another when it is extended. (p. 464)

The gallop, sometimes also called the canter or lope, is the fastest gait of most animals, with the body often unsupported following a push-off with the front legs (flexed suspension) and sometimes with the hind legs (extended suspension). It is usually asymmetrical, with right and left legs doing different movements in a stride. If the last hind leg to touch the ground is followed by the front leg on the same side, it is a rotatory, rotary or lateral gallop ... [typically used by canids] (p. 71)

The typical gait of the dog is the light lateral gallop, which changes to the heavy lateral gallop when the animal slows down. (p. 204)

Apart from a change in the order of limb movements the distinctive feature of a rotatory gallop appears to be that all four feet are off the ground during two out of the eight phases of the cycle. The most striking visible difference between the transverse gallop of a horse and the rotatory gallop of a lightly built dog is the extent to which the latter increases the effective length of its steps, and therefore the length of its stride, by flexing and extending its back (figure 10.19), the hind feet being brought down to the ground much further in advance of the point at which the fore limbs are lifted from the ground than is the case of a horse. (p. 278)

skip or frolic playfully

[with respect to actions used to initiate social play in 3 - 7 week old coyote, wolf, and beagle pups] These include movements of the head and eyes, such as head-tossing and eye-rolling, and also body movements such as shoulder swaying; stalking of the prospective playmate is also included--one animal slowly circles its partner, and then slowly, stealthily approaches; the approach after a stalk may be exaggerated, and may also involve rapid approach/withdrawals. (p. 324 - 325)

In the coyote, wolf and many breeds of domestic dog, the shoulder hackles are comprised of longer hairs with black tips; structural differences have therefore evolved to enhance the signal value of this area (p. 64).

It should be emphasized that the 'hackles' of the neck region or scruff extends laterally on each side behind the shoulders as a pale vertical band of hair in many breeds of domesticated dog and in wolf and coyote. (p. 249)

As peripheral visual structures of expression, the trunk, colouration and build of the back-coat have to be mentioned in particular. Due to the black colouring of the tips of the guard hair (in the large breed), the otherwise milky or tan base-colour of the back appears to be veiled blackish. The shading effect differs depending upon local hair length, and is strongly influenced by the activity of hair-raising muscles and body posture. The hair of the shoulder and the withers is longer than that of the middle of the back. When the BACK HAIRS STAND ON END during fight excitement or impressing...a striking contour of the back is formed. (p.98)

[especially for infant coyotes] Actions such as biting accompanied by rapid side-to-side shaking of the head are used in aggressive interactions and also during predation and could be misinterpreted when used in play. (p.421)

[with reference to aggressive behavior with uninhibited biting] While biting, vigorous lateral head shaking

[with reference to red and Arctic fox] ...side to side head movements which occur just prior to the animal running off in a zig- zag 'catch me if you can' pattern. (p.59)

The Hip Thrust is similar in relativistic structure to the Twist-and-Turn. The essential difference is that a more nearly parallel orientation is maintained and contact is more extensive on the dominant animal. There is also significant pressure exerted by the dominant animal at hindquarters and the animals move sideways together. (p. 212)

... the Hip-Thrust features actual physical contact between the interactants, a near-parallel relative orientation, and points of opposition on both animals slightly to the rear of those maintained during the Twist-and-Turn. Actual thrusting alternates with static configurations, which can be seen in [frames 296, 352, and 407 of Figure 29] S [Supplanting animal] is turned away from D [Displaced animal] and the points of opposition on the two interactants are similar to those seen during thrusting.

... Thrusts alternate with periods of no movement by either animal. ... A shift of front by D results in a change in relative orientation from off-parallel to parallel (the 'thrust' configuration itself). The return to off-parallel is, similarly, the result of a movement by D alone. The progression during the actual maintenance of the parallel configuration is away from S and toward D. In other words, S is doing the 'thrusting' ...

Hip-Thrusts are almost always accompanied by snarling and snapping by both animals near the head and neck regions. The wolves growled loudly, and S frequently displayed vigorous, large tail movements resulting in actual contact with the upper surface of the +back of D. Far more baring of the teeth by both interactants was seen during this than in other stable configurations. (pp. 1154 - 1155).

The howl is a continuous sound from .5 to 11 sec. long. It consists of a fundamental frequency between 150 and 780 Hz., and has up to 12 harmonically related overtones. The frequency is usually constant or varies smoothly, and may change direction up to four or five times. The intensity does not vary greatly throughout (Theberge and Falls,1967) (p.20)

(1)bend or arch into a hump, (2)thrust out or up to form a hump

... inguinal presentation in canids, namely in the grey fox, wolf, coyote and domesticated dog, is a social display associated with social interaction and investigation, submission and appeasement... (p. 88)

During social interaction between dogs, one individual initiates inguinal contact and genital investigation while the other remains passive, and may elevate the hind leg... The adult individual who makes inguinal contact is usually dominant over the one who stands still and allows inguinal contact and genital investigation. During active submission (greeting) in dog, wolf, and coyote, one side (inguinal) may be presented as an animal approaches (the 'C' posture); during passive submission in these species, the uppermost hind limb may be raised to expose the inguinal (genital) area and urination may occur as the animal rolls over on to its side. (p. 83)

...defined as biting which did not cause the recipient to vocalize (yelp) or did not cause a visible wound. (p. 17)

An interaction was defined as a period during which the movements of two animals were judged to be reciprocally influenced following an approach by one of the interactants. (p. 1147)

The term "interaction" is often restricted to the overt expression of activity patterns, whereas less obvious factors, such as the mere presence of one or more other animals, may also exert important influences upon the overt behavior patterns observed.(p. 250)

... will refer to postural components and features of the interaction that can be simultaneously present... (p. 116)

Jaw wrestling was well developed in the wolf cubs by 45 days of age, occurring frequently during agonistic play and during more occasional actual fights. As in the coyote, jaw wrestling during fighting or intense play was accompanied by whines and occasional growls, variable flattening of the ears, eye narrowing and simultaneous vertical and horizontal retraction of the lips. (p.57)

...is a behaviour pattern that appears chiefly in connection with active submission. Members of the pack gather around the senior animals, press against them, jump up at them, lick them in the face, and so on. (p. 176)

See also MUZZLE LICK

LOOKING BACK

a sudden movement forward.

The Lunge/Swivel is a transition from the Follow stable configuration...to a Twist-and-Turn, Stand-Across, or Hip-Thrust... [In three examples] relative distance decreases to actual contact, orientation shifts through almost 360 degrees and the point of opposition on D [Displaced animal] moves from the rear to the shoulder area.

[The movement contributions of the two interactants:] The initial Follow stable configuration is disturbed by a decrease in rate of progression and a small shift of front by D. As the distance between the two wolves approaches minimal values, D performs a large shift of front of near 360 degrees. Only then does S [Supplanting animal] make any active contribution to the change in configuration, by way of a rapid decrease and termination of forward progression. The ordering and type of movement displayed by each interactant are consistent across examples of the Lunge/Swivel, but duration and amount of movement may vary and shifts of front in both directions are observed.

The Swivel/Stand-Across and Lunge/Swivel are somewhat similar transitions. Both involve a change from stable configurations ... Both involve the same pattern of change in relative distance and opposition. They do differ, however, in consistent aspects, which prompted the separate classification used here. The Swivel/Stand-Across is initiated from a Circle and D exhibits a shift of front of about 180 degrees, whereas the Lunge/Swivel always begins from a Follow and involves a 360 degree shift of front by D. ... In addition, the Lunge/Swivel is a faster, more vigourous transition because the larger shift of front by D is performed in a shorter peiod of time than the smaller shift of front in the Swivel/Stand-Across. (pp. 1157 - 1158)

[with reference to to aggressive behavior, generally without body contact] One, but more often two or more, wolves encircling defensive opponent. (p. 285)

[with reference to sexual behavior] Almost exclusively males mounting females from behind. (p. 286)

...the angle between the horizontal projections of the pups' longitudinal axes (regardless of body pitch). The longitudinal axis of each pup was obtained by drawing an imaginary line from the hips to the shoulders, by visual approximation. Only when the spine was fully curved (such as when a dog tries to catch its tail) did we draw the longitudinal axis as a curve. This curve was considered as beginning at the tip of the nose and extending to the tail.

Mutual orientation was scored by drawing the longitudinal axes of the pups as arrows oriented toward each other with an angular resolution of 45 degrees (an orientation thus scored as 90 degrees means that the angle between the pups could be anywhere from 67.5 degrees to 112.5 degrees) In addition to the angular relation between the pups, the location of each pup's head relative to the other pup's body axis was maintained in the arrow drawings. For example, if pup X's snout was oriented toward pup Y's shoulders, both the angle between the longitudinal axes of the interactants, and the location of the snout vis-a-vis the shoulders were retained in the drawing. In cases where a pup [was in contact with the other pup] ..., the origin of the arrowhead, but not its angular orientation, was a direct function of this variable. (p. 180)

... tongue flicks directed toward the muzzle or corner of the mouth [in wolf pups 15 - 85 days of age]. (p. 42)

push gently or gradually.

touch or rub gently with the nose.

...any wolf in the frame "obviously looking" toward the interaction, but not directly involved in it. "Obviously looking" was determined by looking at the wolf's eyes, if possible, to determine the direction of its gaze. If this was not possible, the orientation of the head was used as an indication of whether or not the wolf was looking at the interaction. If the wolf's eyes or head were in line with the action and there was no other known activity occurring in the area that may be capturing the wolf's attention, then the wolf was considered an onlooker. (p. 16)

...(i.e. did the interacting wolves remain together or did they separate greater than one wolf length apart at the end of the interaction?) If the wolves remained together, it was noted whether or not they resumed the activity they were engaged in before the interaction began or changed the activity. This was noted for the initiator and the recipient. If the wolves separated, it was noted who left first - the initiator, recipient, or both. (p. 18)

In the pace, the two feet on the same side of the body swing more or less in unison, which avoids interference between fore and hind feet. The camel family and some large dogs pace naturally when moving moderately fast ... (p.475)

...any wolf directly involved in the interaction either initiating some action toward another wolf or receiving some action from a wolf in the coded interaction. (p. 16)

[with reference to actions used to initiate social play by 3 - 7 week old coyote, wolf, and beagle pups] ...rapid extension and flexion of the forelimb when the animals are at a few meters from one another. (p. 326)

• pinning down with head: lying opponent is pinned to the ground by dominant, keeping head or wide-open muzzle over head, neck, or throat. (p.285)
• pinning down with body: opponent is pressed to the ground by dominant lying on top of it. (p.285)

Having the sole of the foot in contact with the ground when standing or moving, as for man and bears. (p. 629)

... crouching with tail wagging [in wolf pups 15 - 85 days of age] (p. 44)

Both red and Arctic fox have a clearly developed 'play face' in which the mouth is opened 2-5 degrees and the lips are pulled back horizontally. (p. 59)

[A run] with exaggeratedly high raising of the forepaws [in wolf pups 15 to 85 days of age]. (p. 44)

This [exaggerated approach: the "loose, bouncy gait" of the soliciting animal when approaching a playmate at a speed greater than during normal walking] has also been called a "play rush" and "gamboling" in the non-human primate literature. (p. 324)

... the point on each animal that was closest to the social partner. (p.210)

... used to indicate the point on each animal's body to which the social partner was nearest. Opposition need not imply physical contact between the interactants. ... Changes in point of opposition can be produced by movements of either or both animals. (p. 1149)

One may assert without oversimplifying the circumstances that the PRESENTATION OF THE ANAL REGION -- olfactorially and visually -- as well as its CONTROL represent SOCIAL PRIVILEGES. (p. 92)

The opposite of withdrawal, the presentation of the anal region, is -- as mentioned earlier -- observed only in high-ranking wolves, and in females almost only during the pack and mating season. (p. 93)

In the wolf packs I observed, not only females, but also the males sniffed each other at the anal region rather rarely. This is also the case with male dogs which are kept together in packs. In permanent group life, this form of olfactory control appears much reduced compared to the norm we are used to seeing in encounters of separately kept domestic dogs. (p. 93)

Males -- and domestic dogs too -- are generally indifferent to being sniffed at the genitals; the social significance of control of the anus is much more prominent. The latter is actually a very effective expression of social initiative. Diffident individuals react immediately by withdrawing the anal region, the high-ranking wolf on the other hand is characterized by "unembarrassed" display of the anal regions, and control activity on its part. (p. 92)

...all social behavior that is not considered agonistic. (p. 20)

See also MOBBING.

RATED RESTRAINT

...assessed in terms of whether or not oral contact between two wolves resulted in injury or pain. Only when biting occurred was it possible to determine if restraint was involved (i.e. was the biting inhibited or not?) It was assumed that if a bite hurt or wounded the recipient, then the wolf would express this pain in the form of a vocalization such as a yelp or a wound would be visible. Therefore inhibited biting was defined as biting which did not cause the recipient to vocalize (yelp) or did not cause a visible wound. (p. 17)

Pilo-erection of the tail, back, shoulder and rump hackles occurs during agonistic encounters in all canids, except in the arctic fox ... . (p. 63)

[With regard to their scoring of a single agonistic episode] 'Recipient' behavior included: hunching-arching back, wheeling and facing, gaping, rolling on back, placing shoulder on ground, tucking hindquarters of tail, fleeing, or reciprocating aggression. (p. 255)

For example, if wolf A bites wolf B and wolf B responds by biting back at wolf A... (p. 16)

... the minimum distance between the two interactants estimated in "wolf lengths". (p. 210)

... the distance between the two interactants. "Wolf-length" (excluding tail) was used as the unit of measure because it vividly conveyed this aspect of the interaction and eliminated the problems associated with estimating a metric unit of distance from interactions filmed at various distances from the camera. (p. 1149)

... the orientation of each wolf described by polar coordinates zeroed by the longitudinal axis of the social partner. (p. 210)

The orientation of each animal was described relative to the simultaneous orientation of the social partner. This variable was measured using a set of compass-like coordinates zeroed by the longitudinal axis of the second wolf in the interaction. Changes in this variable could be the result of movements by either animal or both animals simultaneously. (p. 1149)

...defined as a reversal of the original roles in the interaction (i.e., biter becomes bittee; chaser becomes chasee; "top dog" becomes "bottom dog". Role reversals are not simply responses to some action. For example, if wolf A bites wolf B and wolf B responds by biting back at wolf A, this is not a role reversal, but instead is an example of reciprocating aggression. Role reversals are usually characterized by a short pause or another action occurring in between the reversal of roles. For example, A approaches and directs a playface toward B; B turns and gives A a playface; A bites at B's tail; B stops and looks at another wolf; and then turns and chases A; A runs away. The role reversal occurred when B pauses and then chases A. (p. 16)

In walking gaits, each foot is on the ground more than half the time; in running gaits, each foot is on the ground less than half the time. (p. 475)

Scent rubbing in wolves has been described as an "uncondtioned response to strong odors" in which the animal lowers a shoulder and rubs its chin, cheeks, temples, neck, shoulders, and (or) back in the odoriferous substance. (p. 573)

• presentation - females walking or standing in front of male holding tail sideways (p. 286)
  
• mounting - almost exclusively males mounting females from behind (p. 286)
  
• pelvic thrusts - while mounting (p. 286)

Zimen, 1982

[with reference to aggressive behavior generally with body contact] hip slam - full lateral body slam when running past opponent. (p. 285)

A certain defensive component is peculiar to it [snapping], sometimes more in the sense of protest or indignation, at other times more of self-defense. Therefore I call the gesture "DEFENSE SNAPPING".

If in a display the wolf growls and its hair is standing on edge, the defense-snapping has a protest character; if the former is not the case, then the wolf's snapping is purely defensive.

The PROTEST-SNAPPING is accompanied by very sharp barking-- FIGHT-BARKING. The more tense the preceding threat to bite appears and the closer the snapping motion is performed to the body of the opponent, the more confident is the wolf with regard to its rival.

The SELF-DEFENSE SNAPPING by the much inferior wolf takes place without barking; instead, THE JAWS BEAT TOGETHER WITH LOUD CHATTERING [Figure 39c]. The gesture is performed already at a great distance (up to approx. 10m) and in rapid succession. In comparison to protest-snapping, this gesture lacks precise direction. (p. 104)

While snuffling the sides of the neck, the tip of the nose is pushed through the coat down to the skin, so that it completely vanishes in the hair. The snuffler usually moves the head slightly whereby the tip of the nose shifts in the coat of the partner in a searching manner. (p. 91)

Social play in canids (and other mammals) may be characterized as follows: (i) actions from various contexts are incorporated into labile (unpredictable) temporal sequences; (ii) the "play bout" is typically preceded by a metacommunicative signal which indicates "what follows is play" (Bateson, 1955); these signals are also observed during the bout; (iii) certain actions may be repeated and performed in an exaggerated manner; (iv) the activity appears "pleasureable" to the participants (Bertrand, 1969; Csikszentmihalyi and Bennett, 1971; Csikszentmihalyi, personal communication; Bekoff, 1974; Reyonlds, unpublished). There is usually an initial decrease in distance between the interactants (Bekoff, 1972a), however, this distance may decrease and then increase, such as during chase and running play. (pp. 323 - 324)

The squeaking vocalization is composed of one or more squeaking phrases, each of which is comprised of one or more individual squeaks. Individual squeaks are brief (usually less than 300 msec), low-amplitude sounds with energy typically between 2 and 4 kHz. (p. 64)

... elements that fulfill the criteria of a squeak: A fundamental frequency in excess of 2 kHz, short duration, often with visible first and second harmonics, and emission in trains (Coscia et al., 1991; Fentress et al., 1978; and Field, 1978). (p. 171)

[with reference actions used to initiate social play in to 3 - 7 week old coyote, wolf, and beagle pups; included in a list of 'general movements'] ... stalking of the prospective playmate...one animal slowly circles its partner, and then slowly, stealthily approaches; the approach after a stalk may be exaggerated, and may also involve rapid approach/withdrawals. (p. 325)

The configuration observed during the Twist-and-Turn was frequently maintained without movement by either interactant. The static maintenance was labelled Stand-Across and is illustrated in [Figure 47]. In these cases, D was frequently seated. (p. 1153)

[in social play] ... one individual places its forepaws on the shoulder or back of another animal and incompletely or fully extends the forelegs. (p. 424)

[with reference to neutral social contacts] Standing transversely over lying partner. (p. 284)

[with reference to aggressive behavior generally with body contact] Standing stiff-legged over, or with one or two feet on lying opponent (p. 285)

[with respect to coyote, and presumably, wolf pups] Standing over a conspecific with the forepaws extended and placed on its shoulders or back emerged at this time [27 days of age for coyote pups]. (p. 244)

thrust out one's limbs and tighten one's muscles after being relaxed.

The wolf making the initial approach was designated the Supplanter (S). The second interactant was referred to as the Displaced Animal (D). In all supplanting interactions, D moved away from the direction of an approach by S. This class was further subdivided into those interactions that ended with the withdrawal of D (simple supplantings) and those interactions that featured a period of interaction following the initial approach and withdrawal. This latter group, called 'extended supplantings', was suited particularly well for detailed movement analysis. Extended supplantings featured the greatest amount of movement. These interactions were shown only by animals that had well-established relationships and resemble interactions described by Shenkel (1967) as "ritualized fighting".

Supplanting interactions comprise an important component of wolf social behavior. They accounted for fully 75% (2102 of 2823) of all interactions observed. Two pairs of animals ... accounted for the majority (66%) of supplantings. (pp. 1147 - 1148)

In all extended supplantings S and D exhibited consistent and distinctive positions of various body parts. These distinctions are illustrated in [Figure 47]. S exhibits erect ears turned toward the front of the head; whereas D holds the ears flat against the head and turned toward the rear. The tail of S is held at back level or above in contrast to that of D which lies between the rear legs and against the animal's underbelly. S exhibits a digitigrade stance with hind legs fully extended, while D holds the hind legs relatively flexed and walks and stands in a somewhat plantigrade fashion. These postural distinctions were observed in all extended and simple supplantings. Some individuals exhibited the characteristics of D in one relationship and those of S in another. (p. 1150)

...involves a change from a Circling stable configuration ... to a Twist-and-Turn, Stand- Across, or Hip-Thrust. [See Figure 37] Relative distance decreases from more than half a wolf-length to values near zero or actual physical contact; relative orientation changes from anti-parallel to near- parallel; and opposition on D shifts from the extreme rear to the shoulder region. ... The transition is relatively slow. ...

The transition begins from a Circling configuration. This stable configuration is disturbed by an increase in rate of shift of front by D and the final (Stand-Across) configuration is achieved by a termination of all movement by both interactants or by the resumption of compensatory movements in cases where the transition terminates with a Twist-and-Turn. In either case the transition is accomplished by active movement by D. (pp. 1156 - 1157)

If the footfalls of a pair of legs, fore or hind, are evenly spaced in time, as in pacing, walking or trotting, the gait is said to be symmetrical. (p. 492)

...one animal oriented perpendicularly to the other and making contact along the longitudinal dimension of its social partner. (pp. 268 - 269)

[based on observations of three wolf pups between three weeks and five months of age] Of all classes of mutual orientation one occurred at a very high rate during the interactions.

The "T" configuration is an orientation of 90 degress ... between the longitudinal axes of two pups, expect when they are at a snout to snout contact. (p. 184)

1. ... lifted at the base and hanging limply thereafter (See Figure 10f, adapted from Schenkel, 1947)
2. ... hanging at the base and lifted to an upward concave curve in the later part only (See Figure 10d, adapted from Schenkel, 1947)
3. ... jagged
4. ... sideways swinging and circling
5. ... extensive sideways swinging of the tail (wagging) with a soft course of motion
6. ... swings with a slightly tugged in tail, often with the entire hindquarters
7. ... very short and rapid sideways wagging with a straight tail or with the end of a lowered tail
8. ... the vertically positioned tail is sometimes in trembling motion. This 'tension-trembling' gives the impression of being charged with energy
9. ... sideward tail-beating (whipping)in the direction of the defendant

1. BEATING: Has a stacatto beat in contrast to the smoothness of a "wagging" tail.
   
   Schenkel, 1947

   A very short and rapid sideways wagging with a straight tail, or with the end of a lowered tail, occurs occasionally in a fighting mood; it is reminiscent , especially in the latter form, of tail movements of fox and cat. (p. 99)

   
   
2. LOOPING: The tail moves in circles.
   
   
3. WAGGING:
   
   McLeod, 1996

   [with respect to wolf pups between 8 and 59 days of age] lateral rhythmic movements of the tail (at any elevation) (p. 107)

   Schenkel, 1947

   Extensive sideways swinging of the tail (wagging) with a soft course of motion is indicative of friendly intentions in every height of rank or tail, respectively. (p. 99)

   
   
4. VERTICAL FLICKING: Rapid movement of the tail in the vertical plane.
   
   Schenkel, 1947

   During encounters of top ranking wolves, the vertically positioned tail is sometimes in trembling motion. This "tension-trembling" gives the impression of being charged with energe and it actually is the expression of highest social power. (p. 100)

tail positions [wolf pups 15 - 85 days of age]:

• tail tuck - tail tip between the plane formed between the posterior ends of the hind legs.
  
• tail high - tail raised at the base and held between 45 degrees below and 22.5 degrees above horiontal.
  
• tail wag - lateral rhythmic movements of the tail (at any elevation). (p. 107)

[with respect to wolf pups between 15 and 85 days of age] tail tip between the plane formed between the posterior ends of the hind legs (p. 107)

... the area on the wolf's body where contact was directed. (p. 17)

In the trot, which like the pace is usually performed at moderate speed by mammals, a fore and hind foot on opposite sides of the body swing approximately in unison. (p. 475)

An example ...[shown in Figure 40]. As shown in [the first frame], the pair begins the transition from a Twist-and-Turn configuration. [The next two frames] represent momentary points in the transition to the Follow configuration.

This transition commonly begins from either a Twist-and-Turn or a Stand-Across configuration. The Turn-to-Rear/Turn initially features an increase in relative distance and orientation and a shift of opposition to the front of D. Relative distance then remains near one wolf-length and the opposition shifts to the extreme rear of D. ... relative orientation can remain near anti- parallel or change to off-parallel values. The latter orientation indicates that the transition ends in a Follow, the former in a Circle configuration.

Like the Swivel/Stand-Across, and the Lunge/Swivel, however, the Turn-to-Rear/Turn moves through the rear rather than the front of the interaction space.

The Turn-to-Rear/Turn features greater variability of mechanism than any other transition type. (pp. 1159 - 1160)

... the animals are in contact or separated by a minimal distance and the point of opposition is between the head or shoulder area of the subordinate animal and the shoulder area of the dominant. The animals maintain an orientation that varies from slightly off-parallel to 45 degrees. This configuration is sometimes maintained because neither animal moves. Just as frequently, however, the animals maintain the configuration by simultaneous compensatory movement. That is, the dominant animal walks forward while turning and at the same time the subordinate pivots at its hindquarters. (p. 212)

The pair maintains a relative orientation between near 45 degrees and near perpendicular, with contact, or less than half a wolf-length, between the shoulder area of S [Supplanting animal] and the forepart of D [Displaced animal]. Maintained points of opposition range only from the shoulder to just back of the shoulder on S and are opposed only to the front and shoulder area of D. Although maintained values of relative orientation are typically near 45 degrees off-parallel, values ranging from near-parallel to near perpendicular sometimes occurred briefly during the Twist-and-Turn.

...the animals maintain actual contact or near contact at the shoulder region of D. Relative orientation shows greater variation than either of the other variables, but remains principally near 45 degrees.

The configuration observed during the Twist-and-Turn was frequently maintained without movement by either interactant. The static maintenance was labelled Stand- Across and is illustrated in [Figure 47]. In these cases, D was frequently seated.(p. 1153)

• SQUAT URINATION: the wolf lowers the posterior half of the body by spreading the hind legs apart before urinating in this position.
  
• RAISED LEG URINATION: the wolf raises one hind leg before urinating with the leg raised.

Wolves have an extensive vocal repertoire but researchers disagree on how to categorize the sounds. Schassburger (1993) described the wolf vocal repertoire as a system divided into two sound groups: (1) harmonic sounds including whines, whimpers (squeaks), yelps and howls and (2) noisy sounds including growls, barks, snarls, whine-moans, moans, and growl-moans. Others, such as Theberge and Falls (1967), Harrington and Mech (1978), and Coscia, Phillips, and Fentress (1991), suggested that wolves produce 4-6 different sounds (growl, bark, yelp, whimper [squeak], howl, and whine) and considered the others as subclasses. (p. 3)

In walking gaits, each foot is on the ground more than half the time. ... At the usual walk, the four footfalls are independent; a forefoot strikes the ground next after the hind foot on the same side of the body (lateral sequence) in most tetrapods except primates, because interference between forefeet and hind feet is then avoided. The same gait can be done at the run but is unusual except for certain show horses. Gaits having independent footfalls provide the most stability (p. 475)

The walk is a slow, regular, symmetrical gait in which the left legs perform the same movements as the right legs but half a stride later... (p. 63)

An example of the Walk-Away/Walk transition is shown in [Figure 43]. In [the first frame] the pair maintains the Twist-and-Turn configuration. ... The transition also often begins from the Stand-Across or the Hip-Thrust configuations. In the next frames a transition to ... the Follow configuration is achieved via relatively small and direct changes of configuration. The Walk-Away/Walk sometimes flows directly into another transition (Lunge/Swivel) rather than a stable configuration.

The Walk-Away/Walk was the least frequently observed of the transitions. In all three [examples] a characteristic pattern of changes in relational variables is apparent during the Walk-Away/Walk transition itself: an increase in relative distance, the assumption or maintenance of a parallel relative orientation, and a shift of oppostion to the rear of D.

The disturbances of the Twist-and-Turn, Stand-Across, or Hip-Thrust are the result of movement by D alone. D first walks away from S, progressing slightly sideways, and makes a shift of front only after the distance between the two interactants has increased. At this point S initiates a forward progression and the Follow configuration begins. The order of these movements and the type of movement shown by each interactant are fixed. (p. 1161)

The transition begins from a Follow configuration as shown in [the first frame]. Between the first and second frame of the illustration, relative distance decreases to near zero. From the second to the third frame, opposition shifts to the shoulder region of both animals, and by the final frame, relative orientation changes to near 45 degrees and the animals assume the Twist-and-Turn stable configuration.

The Walk-Up/Stop contrasts sharply with the Lunge/Swivel and Swivel/Stand-Across. The latter transitions feature large changes in all relational variables, whereas the Walk-Up/Stop leaves the impression of a slow, almost leisurely pattern with relatively small changes in relational variables. ... Unlike the Swivel/Stand-Across and Lunge/Swivel, the Walk-Up/Stop is never followed by a Hip-Thrust.

Two equivalent forms were observed. In the form illustrated in [Figure 44] a Follow is disturbed by a decrease and cessation of forward progression by D. The continued progression by S [Supplanting animal] reduces relative distance and brings points of opposition into the shoulder region and the range of the Twist-and-Turn or Stand-Across configurations. The transition is completed by a small shift of front by S. In the second form of the transition, ... an increase in rate of forward progression by S precedes the cessation of forward progression by D [Displaced animal] described above. The two forms are equivalent in that they produce the same relational changes; both in terms of end points and form in the interaction space. S shows no systematic tendency to stand on one side or the other of D. (pp. 1158 - 1159)

...takes place by means of curving the back downwards, bending the hind-legs and "tugging-in" the tail ..., under some circumstances additionally by turning the hind-part of the body away sideways. (p. 92)

'Wolf-length' (excluding tail) was used as the unit of measure because it vivdly conveyed this aspect of the interaction and eliminated the problems associated with estimating a metric unit of distance from interactions filmed at various distances from the camera. (p. 1149)

Wolves, like dogs, yawn to demonstrate boredom or embarrassment. Yawning was originally a mechanism whereby muscles were stretched and lymphatic vessels compressed to ensure a flow of lymph after a period of inactivity. (p. 141)