We noted each time a wolf submitted posturally to another wolf. Usually this deference was characterized by "licking up" to the mouth of the dominant animal in the "active submission" posture (Fig. 5 in Schenkel 1967), similar to that described by Darwin (1877) for domestic dogs. Often this behavior took place as an animal returned to the den area after foraging, and sometimes the returning individual disgorged food to the soliciting wolf (Mech 1988; Mech et al. 1999). Other behavior noted included "pinning," or passive submission (Schenkel 1967), in which the dominant wolf threatened another, which then groveled, and "standing over," in which one wolf stands over another, which often lies nonchalantly but in a few cases sniffs the genitals of the other. I did not consider "standing over" a dominance behavior (L.D. Mech, submitted for publication). 2
The following is a summary of generalizations documented in the previous references, together with new quantified findings.
2 L.D. Mech. "Standing over" and "hugging" in wild wolves. Submitted for publication.