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.