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. 2018 Jul;141:137–150. doi: 10.1016/j.anbehav.2018.04.017

Table A1.

Behaviours used to determine agonistic and affiliative relationships outside the play context

Behaviour Definition
Rank indicator behaviours
Dominant behaviours
Stand tall A straightens up to full height, with a rigid posture and tail; may include raised hackles, ears erect and tail perpendicular or above the back
Stand over A is standing over R's body, with all four paws on the ground, with the tail held high. R may have either the whole body or just the forepaws under A's belly/side
No-play paw on A places one or both forepaws on R's back; outside of the play context
Ride up A mounts R from behind or from the side, exhibiting a thrusting motion
Head on A approaches R's shoulder/back and puts its head on it. Formation usually looks like a capital ‘T’
No-play muzzle bite A grabs the muzzle of R either softly or with enough pressure to make the other whimper; outside of the play context
Submissive behaviours
Crouch A lowers the head, sometimes bending the legs, arching the back, lowering the tail between the hindlegs and avoiding eye contact
Passive submission A lies on its back showing its stomach and holding its tail between its legs. The ears are held back and close to the head and A raises a hindleg for inguinal presentation
Active submission A has its tail tucked between the hindlegs sometimes wagging it while in a crouched position (with hindquarters lowered) and may attempt to paw and lick the side of R's muzzle. The behaviour may include urination
Withdrawing A withdraws from R moving away slowly in the opposite direction, displaying a submissive posture. It occurs when A has been threatened or attacked by R, or a fight has taken place
Flee A runs away from R with tail tucked between the legs and body ducked. It occurs when A has been threatened or attacked by R, or after a fight
Avoidance In response to R reducing the distance to it, A moves away displaying a submissive posture. A may also look at the individual it is trying to avoid
Aggressive behaviours
Threat A orients towards R performing one or more of the following behaviours: staring at, curling of the lips, baring of the canines, raising the hackles, snarling, growling and barking, sometimes with the tail perpendicular or above the back
Attack A runs into or jumps onto R with tail, ears and sometimes hackles up, often with bites at the neck
Knock down A strikes R sharply with the chest or shoulder so that R falls to the ground
Pin A grabs R at the neck or muzzle, forcing it down to the ground and holding it there
Fight A and R engage in reciprocal biting and aggressive physical contact
Chase A runs after R, exhibiting threatening behaviours (see ‘threat’ above)
Jaw spar A and R ‘fencing’ with open jaws
Snapping A snaps its teeth into the air towards R
Affiliative behaviours
Grooming A nips, licks or scratches the fur or skin, occasionally the neck of R
Lie friendly A lies on its back, tail-wags, sometimes kicking with the foreleg against R, often with open mouth
Stand friendly A stands with tail perpendicular to or below the plane of the back, wagging it, ears pointed forwards, while R is approaching it or orienting/looking towards it
Social sniff A sniffs R's body part except its anogenital area
Body rubbing A rubs its body against any part of R's body
Body contact A stays (for at least 10 s) with at least part of its body in contact with R; both A and R have a relaxed position

Adapted from the Wolf Science Center Social Behavior Ethogram. A: wolf performing the behaviour; R: wolf receiving the behaviour.