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Leadership Behavior in Relation to Dominance and
Reproductive Status in Gray Wolves, Canis lupus

Introduction


In social groups or packs of gray wolves, Canis lupus, dominant individuals are those that successfully control the behavior of others (Schenkel 1947) and pack leaders are those that control pack movements (Mech 1970, p. 73, 2000; Boinski 2000). How pack leadership relates to dominance and breeding is not well documented for wild wolves, particularly in packs containing more than two breeding individuals (6-8% of all packs; Mech 2000). Furthermore, there is disagreement in the literature about the relative roles of male and female breeding wolves in directing pack activities. Murie (1944) referred to the dominant, or alpha, male as the lord and master of the pack; Haber (1977) agreed, and Fox (1980, p. 128) described the alpha male as the leader of the pack (italics added). Yet Mech (1966) and Peterson (1977) reported that the dominant female usually leads a pack during travel, at least during the midwinter breeding season. Mech (1970) referred to the dominant male as the one who initiates and guides attacks on intruding wolves from outside the pack. A recent review emphasized that it is alpha male wolves that are the pack leaders (Holekamp et al. 2000).

While it is generally understood, especially from studies of wolves in captivity (e.g., Fox 1980; Zimen 1981), that in wolves, leadership has some relationship to dominance and breeding status, quantitative data on leadership behavior in the wild have been reported for only one pack in summer (Mech 2000). Mech (1999) asserted that the significance of dominance relationships within pack society has been overrated, and he argued that wolf packs are best understood as family groups in which a breeding pair "shares leadership in a division of labor system in which the breeding female initiates pup care and the breeding male leads in foraging and food provisioning". According to this view, breeding wolves provide leadership because offspring tend to follow their parents' initiative. Yet we know little about how leadership roles might change in packs with multiple breeders (Mech 1999), or how individual age or pack size might influence the process of leading.

The primary objective of this study was to document and compare the leadership roles of breeding and nonbreeding wolves, dominant and subordinate breeding females, and dominant males and females. The reintroduction of wolves to Yellowstone National Park in 1995, following their extirpation by 1926 (Bangs and Fritts 1996; Bangs et al. 1998), afforded an opportunity to observe the behavior of wolves of known sex and age in free-ranging wild packs. We studied leadership behavior in three packs, including one with a simple structure (a breeding pair and their offspring) and two with multiple breeding females. All three packs contained offspring from multiple years. During the study there were two turnovers in dominant breeding individuals, shedding further light on the relationship between social status and leadership role.


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