Northern Prairie Wildlife Research Center
However, few opportunities have existed for studying leadership in wild Wolf packs because of the elusive nature of Wolves. Only Murie (1944), Clark (1971), Haber (1977), and Mech (1988, 1995a, 1999) have studied behavior in free-ranging Wolf packs, and no one has examined the leadership concept critically or quantitatively. This article attempts to do so, based primarily on my 13 summers of observation of a free-ranging Wolf pack.
Because Wolf packs are basically families (Murie 1944; Mech 1970), or at least almost always include a breeding pair (Mech and Nelson 1990; Mech et al. 1998), it is only natural that some member of this pair would be the pack leader (Mech 1970). This is because most members of the pack would be the offspring of the breeding pair and would tend to follow their parents' initiatives. Sometimes, a post-reproductive animal remains with the pack (Mech 1995a). Conceivably, such an individual, being older and more experienced, would lead in some activities. On the other hand, deposed breeders usually become subordinate, and tend not to take initiatives involving the group (Zimen 1976). A post-reproductive female on Ellesmere was subordinate to the breeding pair (Mech 1999).
Therefore, determining which member(s) leads the pack in a given activity would usually involve determining whether it is the breeding male or breeding female. Murie (1944) identified a male "lord and master" of a pack to which all four other adults submitted. Murie concluded that this animal was not mated to any of the females, although there was no way he could have known (Haber 1977). One of the other adult males tended to lead the chases of Caribou, Rangifer tarandus, during Murie's study.
On Isle Royale, Mech (1966) observed from the air that in a large pack one member stood out as leading the pack, taking the initiative during hunts, and making decisions, but Mech could not identify the individual well enough to know whether it was always the same Wolf. It did, however, seem to be a male, and during the breeding season, a female sometimes led, with a male close behind her.
Later, Jordan et al. (1967) recognized a particular male that led the Isle Royale pack during travels for several winters. On the other hand, Peterson (1977), observing the Wolves during the breeding season, believed that females tended to lead the packs, as concluded by Pulliainen (1965: 236) who cited anecdotal literature.
Haber (1977) considered a "beta male" to be the leader of the Savage Pack in Denali Park. That animal tended to break trail, set the direction and pace of travel and resting, and initiated and ended most of the hunts and rest periods. However, this pack was highly unusual in that both the alpha male and beta male remained with the pack until 8 or 9-years old. No one else has ever reported two adult males remaining concurrently with a pack for even 4 years, including during a study of 13 packs for 4-9 years in the same area (Mech et al. 1998). In two other packs, Haber believed that high-ranking males generally led pack travels. During summer, leadership was less clear but Haber believed it also tended to involve high-ranking males. None of these studies provided quantitative behavioral data on leadership.