David Mech is a biologist who has devoted his entire career to the study of wolves. He is a wonderful scientist who has contributed a lot during his life to help us understand wolves better.
The text here is a translation of his 1999 article "Alpha status, dominance, and division of labor in wolf packs". Translation of the article and publication here is coordinated with David Mech.
Alpha status, dominance and division of labor in wolf packs
David Mech
Summary. According to the prevailing understanding, wolves (Wolf dog) a group of individuals in a herd, in which there is a struggle for the dominant role, and which is controlled by an "alpha pair" - an alpha male and an alpha female. However, many studies of wolf pack dynamics are based on unnatural populations of captive wolves. In this paper, I describe wolf pack social relationships as they occur in the wild, analyze the concept of alpha, social dominance, and submission, and provide data on the exact relationships of free-ranging packs based on a review of the literature and 13 summers spent tracking wolves on Ellesmere Island in northwestern Canada. According to my conclusion, a typical wolf pack family is one in which adult parents manage the group's activities based on a division of labor system, in which the female primarily controls activities such as care and protection of the pups, and the male foraging and associated migration.
Introduction
of wolves (Wolf dog) herds have long been used to describe the behavioral relationships between members of social groups. The topic of social dominance and alpha status has received considerable attention (Schenkel 1947; Rabb et al. 1967; Fox 1971b; Zimen 1975, 1982) and the prevailing view is that a wolf pack is a group of individuals competing for dominance and controlled by an "alpha pair"—an alpha male and an alpha female (Murie 1944; Mech 1966, 1970; Haber 1977; Peterson 1977).
However, most studies of wolf pack dynamics are based on unnatural populations living in captivity. These captive packs usually consisted of wolves from different locations, brought together and allowed to breed freely (Schenkel 1947; Rabb et al. 1967; Zimen 1975, 1982). This approach reflected the notion that in the wild “pack formation begins at the onset of winter” (Schenkel 1947), referring to the annual aggregation of independent wolves. (Schenkel did consider the possibility that a herd is a family, as Murie (1944) had already suggested, but only as a footnote.)
In captive packs, wolves that did not know each other formed hierarchies consisting of alpha, beta, omega members, etc. In such communities, these definitions were probably relevant because most captive species tend to share roles in similar ways.
However, a wolf pack living in the wild is not such a community. Instead, it is a family (Murie 1944; Young and Goldman 1944; Mech 1970, 1988; Clark 1971; Haber 1977) consisting of a breeding pair and its offspring of the previous 1-3 years, or sometimes two or three such families (Murie 1944; Haber 1977). 1977; Mech et al. 1998).
Occasionally a wolf unrelated to the others joins the pack (Van Ballenberghe 1983; Lehman et al. Mech et al. 1998) or is a relative of one breeding individual (Mech and Nelson 1990) or the dead parent is replaced by a wolf outside the pack (Rothman and Mech 1979; Fritts and Mech 1981) and one of the offspring, which is of the opposite sex to the newcomer, replaces its parent and breeds with the foster parent (Fritts and Mech 1981; Mech and Hertel 1983).
However, these variations are exceptional, and even in these situations the flock consists of a breeding pair and its young offspring (Mech 1970; Rothman and Mech 1979; Fritts and Mech 1981; Mech and Hertel 1983; Peterson Peterson et al. 1984). The herd acts uniformly throughout the year (Mech 1970, 1988, 1995b).
When offspring begin to reach sexual maturity, they leave the herd at nine months of age at the earliest (Fritts and Mech 1981; Messier 1985; Mech 1987; Fuller 1989; Gese and Mech 1991). Most leave at 1-2 years of age, and few remain in the herd for more than three years (Mech et al. 1998). Thus, young members form a temporary part of most flocks, and the only long-term members are the breeding pair. In contrast, captive herds often contain members forced to live together for many years (Rabb et al. 1967; Zimen 1982; Fentress et al. 1987).
Attempts to transfer information gathered about the behavior of unrelated captive wolf groups to the family structure of wild packs has led to much confusion. This approach is analogous to trying to make generalizations about people's family dynamics by studying people living in a refugee camp. Most misleading is the perception of the alpha wolf as a "boss" who rules over a group of peers of a similar age (Schenkel 1947; Rabb et al. 1967; Fox 1971a; Zimen 1975, 1982; Lockwood 1979; from Hooff et al. 1987)
Because wolves have been persecuted for so long (Young and Goldman 1944), their study in the wild has been difficult (Mech 1974), and thus information on free-living pack members has been slow to gather. We don't know much about the relationship between breeding males and females in the wild, what their role is in the herd, and how dominance affects those relationships.
Some people have observed the social behavior of wild wolves in the vicinity of a den, but Murie (1944) provided only anecdotal descriptions, Clark (1971) is an unpublished thesis that provided only a quantified summary of pack hierarchical relationships, and Haber (1977) described his interpretation of pack social hierarchy, without providing supporting evidence. This means that no one has yet quantified the hierarchical relationships of a wild wolf pack.
In this article, I try to explain the social order of a wolf pack living in the wild and to add to our knowledge about the social dynamics of wolf packs by analyzing the concept of "alpha" and social dominance and presenting data on dominance relationships between members of free-living packs.
Methods
The research was carried out between 1986 and 1998. in the summers of Ellesmere Island in the northwestern part of Canada (80°N, 86°W). There wolves hunt arctic hares (Arctic hare), musk cattle (Ovibos moschatus) and Peary caribou (Rangifer tarandus pearyi) and live far enough from human exploitation and persecution that they have relatively little shyness towards humans (Mech 1988, 1995a). I trained the wolf pack there with my presence during 1986 and strengthened that relationship every summer. The pack visited the same area every summer and usually used the same burrow or nearby burrows. Presence practice allowed me and my assistant to spend time with wolves every day, learning to distinguish individuals and observe them even from as little as 1 meter away (Mech 1988, 1995a; National Geographic Society 1988).
We noted all times when a wolf surrendered to another wolf with its posture. Typically, this submission was characterized by licking the dominant animal's mouth from a position of "active submission" (Fig. 5, source Schenkel 1967) similar to what Darwin (1877) described in domesticated dogs. This behavior often occurred when the animal returned to the burrow from foraging, and sometimes the returning individual ejected food to the behaving wolf (Mech 1988; Mech et al. 1999). Other behaviors recorded included "pushing" or passive submission (Schenkel 1967), in which the dominant wolf threatened another who dropped on it, and "standing over," in which one wolf stands over another, often indifferently lying, individual and sometimes sniffs the genitalia of another individual. I did not count 'standing' as a dominant behavior (L.D. Mech, submitted for publication).
The following is a summary of the generalizations documented in the sources below, along with new quantified findings.
Results and discussion
Alpha status
"Alpha" refers to the first rank in some kind of hierarchy, so an alpha wolf is by definition the highest ranking wolf. Because hierarchies are gendered among captive wolves, there is an alpha male and an alpha female in the pack (Schenkel 1947).
How alpha status has been treated in the past can be read in research papers where attempts are made to distinguish future alphas among captive wolf pups. For example, one hypothesis was that "the emotional reactivity of a dominant pup, the potential alpha animal of a pack, may be markedly different from that of subordinates" and that "therefore it may be possible to distinguish between the temperamental characteristics or emotional reactivity of potential alphas, or leader wolves, and subordinates" (Fox 1971b, page 299). Furthermore, under 'normal field conditions', timid and low-ranking wolves seem unlikely to breed (Fox 1971a, p. 307). This understanding suggests that order is innate or established early, and that some wolves are destined to lead the pack, while others are not.
As a counterpoint to this, I propose that all young wolves are potentially reproductive and, if they reproduce, automatically become alpha (Mech 1970). Even in captive packs, individuals gain or lose alpha status (Zimen 1976), so certain wolves have no innate social status, even though captive pups exhibit psychological and behavioral differences corresponding to the prevailing social rank (Fox 1971b; Fox and Andrews 1973). Second, wolves breed freely in captivity, and I don't know of any mature captives that failed to breed with a mate from outside the group, even if it was a low-ranking non-breeding animal.
Third, in the wild, most wolves leave their natal pack and attempt to mate with other wolves that have left, have pups, and establish their own pack (Rothman and Mech 1979; Fritts and Mech 1981; Messier 1985; Mech 1987; Gese and Mech 1991; Mech et al. 1998). I know of no permanently departed specimen that failed to breed if it survived long enough to do so.
There is great variability among wolves in age, distance, direction, etc. of departure (see previous references) and is expected to be related to the within-litter differences described above (Fox 1971b; Fox and Andrews 1973). However, unless a sexually mature herd member inherits a position that allows her to breed within herd with her foster parent (Fritts and Mech 1981; Mech and Hertel 1983), sooner or later she will leave the herd and attempt to breed elsewhere.
Designating a higher-ranking wolf as alpha emphasizes its position in the dominance hierarchy. In natural wolf packs, however, the alpha male and female are merely breeding animals, the parents of the pack, and rarely, if ever, compete with other wolves for dominance. In my 13 summers following the Ellesmere Island herd, I saw no such cases.
Thus, calling a wolf alpha is no more appropriate than referring to a human parent or a deer male as alpha. All parents dominate their offspring, so the "alpha" adds no information. Why not refer to the alpha female as the female parent, the breeding female, the matriarch, or simply the mother? Such a definition emphasizes not the animal's dominant status, which is superficial information, but its role as a herd ancestor, which is critical information.
One use of the term "alpha" that may remain in use is in relation to the relatively few large wolf packs that consist of multiple litters. Although the genetic relationships of the mothers of these packs are still unknown, the mothers likely include the original matriarch and at least one daughter, and the fathers are likely the patriarch and unrelated adopted wolves (Mech et al. 1998). In such cases, the older breeding animals are likely to have a dominant attitude towards the younger breeding animals, and thus it is more appropriate to call them alphas. Examples of such competition would be the continued dominance of food distribution and herd movement by the parent breeding animal.
The problem here is not so much the concepts, but what these concepts falsely refer to: a rigid, force-based hierarchy of dominance.
The extent to which these arguments apply to other species will undoubtedly vary considerably and is beyond the scope of this article. However, it is significant that similar arguments can be made for hyena dogs (Lycaon painted) that are ecologically similar to wolves (Mech 1975). Although some researchers have described this species as not exhibiting rank-based behavior (Kuhme 1965; Estes and Goddard 1967), others describe it as an “alpha animal” (Creel and Creel 1996).
Dominance and submission among herd members
The concept, nature and nature of dominance hierarchy or social hierarchy (Schjelderup-Ebbe 1922) has been questioned in many species (summarized in Wilson 1975). Similarly, dominance in the wild wolf pack does not manifest itself as a social hierarchy and appears to be much less important than has been suggested in studies of wolves in captivity (Schenkel 1947, 1967; Rabb et al. 1967; Zimen 1975, 1982; Lockwood 1979). In a natural wolf pack, the rules of dominance do not to some extent resemble those of a social hierarchy, in which a group of similar individuals competes to achieve a certain rank.
The only consistent demonstration of rank in natural herds is the body posture of animals in social encounters. Dominant wolves assume the classic wolf stance, raising their tails to at least horizontal, and subordinate or submissive individuals lower their bodies and “smash” (Darwin 1877). Whereas submission itself can be just as important as dominance, promoting friendly understanding and reducing social withdrawal.
Table 1. Dominance exposures, i.e. the number of times individual wolves dominated or submitted to others, among breeding individuals of the Ellesmere Island wolf pack during summer in the absence of other individuals.
Note: Exposures consisted primarily of active submission, but three cases of passive submission are also included (Schenkel 1967); they do not include 'standing' or food contact other than 'asking for food'.
Speaking about the importance of submission, Schenkel (1967) distinguished two main types - active and passive. He believed that active submission came from asking for food, and in my opinion, active submission and asking for food are indistinguishable. A wolf asking for food or submissive approaches another wolf excitedly, wagging its tail, pulling its ears to its head and licking the other wolf's muzzle. Depending on the situation, the other wolf may or may not eject food for him (Mech et al. 1999). In passive submission, the submissive wolf lies on its side or back and the dominant wolf sniffs its groin or genitalia (Schenkel 1967). Active submission was more common among the Ellesmere Island herd.
In this herd, all members, including the breeding female, submitted to the breeding male both actively and passively with body posture (Schenkel 1967). One-year-old and two-year-old wolves and an older female past the breeding age surrendered to both breeding animals. These rules applied regardless of pack members: breeding pair or breeding pair with pups (Table 1), breeding pair with yearlings (Table 2), breeding pair with yearlings and pups (Table 3), breeding pair with pups and 2-year-old outside wolves (Table 4) or a breeding pair with puppies and a female out of breeding age (Table 5).
On June 22, 1991, I observed a remarkable example of how submissive rules foster friendly relationships. Out of breeding, the female returned to the burrow with a very dry rabbit carcass, which was more interesting entertainment than food. Instead of taking the dried rabbit directly to the puppies, she took it in a specially submissive way to the breeding male, who immediately grabbed it from him. He refused the pleas of both this female and even the breeding female and chewed it himself for 20-30 minutes.
Other notable dominance rules I observed included scent marking and possession and transfer of food. Both the breeding female and male mark with their scent, but inferior animals do not unless they are seeking a dominant position (Packard 1989; Asa et al. 1990) and I did not see such exceptions. Possession and transfer of food - when there were pups or yearlings in the herd, the breeding male I observed either ejected food or dropped it in front of his mate, or let his mate take the food, or took the food directly to his offspring. With the exception of taking food to the pups, each wolf appeared to have a possession zone around its mouth (Mech 1970), and regardless of the order of the animal that invaded it, the owner tried to keep the food in its possession, as Lockwood (1979) also observed in captive wolves. Wolves of any order could try to steal food from another wolf of any order, but each wolf defended its own food (Table 6). Overall, dominant wolves appeared to be more successful at stealing food, but the sample size was too small to draw definitive conclusions.
Table 2. Dominance relationships, i.e. the number of times individual wolves dominated or were subjugated among breeding animals and yearlings of the Ellesmere Island wolf pack during the summer of 1993 (no pups and same parents as in Table 1).
Note: Exposures do not include 'standing' or food, except for 'asking for food'. aMale parent vs. female parent, h2 = 0,94, P = 0,33, df = 1.
Table 3. Dominance exposures, i.e. number of times individual wolves dominated or were subjugated among breeding animals and yearlings of the Ellesmere Island wolf pack in 1988 (pups present and parents same as 1990–1996).
Note: Exposures do not include 'standing' or food, except for 'asking for food'.
a* Includes one short sequence of five submissions
b* Includes one short sequence of four submissions
Table 4. Dominance exposures, i.e. number of times individual wolves dominated or submitted to others among breeding animals and 2-year-olds of the Ellesmere Island wolf pack in 1994 (pups present and parents same as Tables 1 and 2) .
Note: Exposures do not include 'standing' or food, except for 'asking for food'.
a* These are the annual animals presented in Table 2.
b* On one of these occasions, a 2-year-old female dominated the older female for 15 minutes. No order was included where it was not clear whether the female parent or the 2-year-old female was dominant.
c* Male parent vs. female older, χ2 = 3.99, P = 0.05.
Two other behaviors between pack members may have been related to dominance, but there is insufficient data to confirm this. They stood over each other and "hugged". In "standing up", one wolf (Schenkel 1947) stands over a lying wolf in such a way that its groin is over the sleeping wolf's nose. Sometimes a lying wolf sniffed the genitals or groin of a standing wolf.
Schenkel (1947) saw "standing" only in "peacetime" and did not find it related to dominance. For cuddling, my sample size (5) was insufficient to determine if it was related to dominance.
The rules of dominance described above, which involve an age-based order with a breeding pair at the top and offspring and non-breeding animals subordinate to them, are so automatic that they are rarely tested. In this regard, social interactions between members of natural wolf packs are much more peaceful than those described by Schenkel (1947) and Zimen (1982) among captive wolves, as Clark (1971) also noted. Puppies also submit to adults and their older siblings in the same automatic and peaceful manner. It is debatable whether or when a hierarchy develops between pups (cf. Zimen 1975 and Fox and Andrews 1973; Haber 1977) and I cannot comment on that. Even among yearlings and 2-year-old animals, there were very few stage manifestations (Tables 2–5).
Table 5. Dominance encounters, i.e. the number of times individual wolves dominated or submitted to others between breeding animals and a non-breeding female in the Ellesmere Island wolf pack during the summers of 1990 and 1991 (pups were present and the male parent was the same in all years , except in 1998).
Note: Exposures do not include 'standing' or food, except for 'asking for food'.
a* Yearling female in 1988 (Table 1) and female parent in 1990–1996.
b* Female parent in 1988 and 1989 (Table 1).
c* The male surrendered when he approached the female in the burrow and the young pups.
χ2 = 12,64, P < 0,001, df = 1.
Table 6. Observed attempts to protect food from pack members in the Ellesmere Island wolf pack.
a* Does not include the taking of food from the breeding male by the breeding female.
b* A one-year-old female brought food to the pups and bit the air menacingly at the breeding female when she stole the food.
c* The one-year-old female that brought the rabbit kept vigil near the cub.
d* The breeding female could not stop the breeding male.
It is conceivable that social tensions increase during the breeding season (Schenkel 1947), but the fact that most herds have only one breeding pair makes it possible to prevent these tensions. Wolves living in the wild start breeding at the youngest age of 22 months (Ibid et al. 1979) and some individuals do not reach sexual maturity until at least 4 years of age (Haber 1977; Mech and Seal 1987). Since most wolves leave the pack before the age of 2 years, and almost all leave before the age of 3 years (Mech 1987; Gese and Mech 1991; Mech et al. 1998), in most herds there is no reason for breeding competition.
Thus, intense rivalries can only develop in those relatively few herds with multiple breeding pairs, as Haber (1977) described the breeding season in his unusual herd. On the other hand, it is possible that at least part of the described difference in "hostility" is caused by the different perspectives of the followers themselves. From time to time in the summer of 1944, I saw a mother forcefully push down her 2-year-old female offspring in what some might consider "hostile" behavior. To me, this behavior appeared to simply be a type of contact I observed between a mother and her out-of-control offspring. In any case, these kinds of exposures were not common at the time of my research.
When animals of a higher order control individuals of a lower order, the nature of this contact is extremely contingent. For example, larger prey such as adult moose (Alces alces) herd members of all ranks (ages) congregate around the carcass and feed at the same time without showing rank privilege (Mech 1966; Haber 1977). However, when the prey is smaller, such as a musk ox calf, dominant animals (breeders) may eat first and control when inferior individuals eat (Mech 1988; National Geographic 1988).
Similarly, puppies are inferior to their mother and father as well as their older siblings, while parents and even siblings give them food in priority order (Mech et al. 1999). When food is scarce, the breeding pair dominates their parent offspring and limits their food intake by feeding the pups instead. Thus, the most practical output of social dominance is the dominant individual's ability to control who has access to food.
The only other rank privilege I am aware of in the wild are situations where higher-ranking pups assert themselves more vigorously when adults bring food, and sometimes accompany younger-aged adults for foraging compared to lower-ranking pups (Haber 1977).
Dominance between breeding female and male
The relationship between the breeding female and male is complex and deserves further investigation. In captive herds, conflicting claims have been made regarding the dominance of "alpha males" and "alpha females" over each other and over subordinates. This question is closely related to the concept of leadership, but not necessarily exactly the same (L.D. Mech, submitted for publication).
It has been debated whether each gender has its own dominance hierarchy. As pointed out by Hooff et al. (1987, p. 248), Schenkel (1947) and Zimen (1982), each sex has a separate hierarchy in captive wolves. However, observations made in the wild do not agree with this. According to Clark's (1971) data, the breeding male dominated all other wolves and the breeding female dominated all but the breeding male. Haber (1977, p. 203) stated that the wolves he studied in the wild were dominated by males “with very few exceptions”. My records are consistent with breeding males dominating their body posture, but I have seen only one case where a breeding male also dominated the breeding female with posture (Tables 1-5).
Disagreements regarding the relationship between breeding females and males are likely due to the large differences between members and backgrounds of captive and wild herds, discussed earlier. Thus, it is useful to describe typical interactions between a breeding female and male in natural herds, as this has not been done before.
When the breeding female and male are separated, recognize each other and then meet, the breeding female approaches the male with a typical submissive posture – tail down or between legs, body hunched or close to the ground, ears back and nose up – and licks the male's mouth (Schenkel 1947 ). The male stands indifferently and sometimes raises his tail to a horizontal position.
In the summers when the herd I observed had pups or yearlings, such encounters often took place near them when the male returned from foraging. The male's response to the female's greeting was to drop the food in his mouth and/or expel it (Mech et al. 1999). The female then either ate the food or gave it to her offspring. I was unable to distinguish in which cases such a greeting was followed by the expulsion of food and in which cases it was not.
In 1998, when a breeding pair had no offspring, the four encounters between breeding pairs that I observed occurred immediately after the female had temporarily foraged alone or hid food from prey alone. Each time the female returned to her mate, she assumed an active submission posture when meeting the male, and on one of these occasions she submitted very expressively for about 90 seconds. Even on one occasion when a breeding female was chasing another wolf and her mate ran past her (June 17, 1991), she momentarily surrendered as the male passed. Based on these observational data, it is reasonable to conclude that the breeding female surrendered to her mate.
However, the practical purpose of these postures of submission is not obvious. The behavior did not always appear to be related to asking for food. For example, during one encounter in 1998, the female submitted to the male as described above when the female had a large bone from which she had just eaten a lot of meat. The male, who hadn't eaten for at least a few hours, tried to take the bone. However, the female defensively bit the air towards the male and successfully kept the bone despite the male repeatedly trying to take it away over the course of an hour.
Even if the purpose of the breeding female's active submission to her mate was to ask for food and not to express her inferiority, it can still be argued that sometimes the breeding female also passively submits to the male (Schenkel 1967). I observed it three times on Ellesmere Island (Table 1), but never saw a breeding male passively surrendering to a female. Because passive submission appears to be unrelated to food begging, these observational data clearly point to a lower order.
When attacking prey, the breeding female and male appear to be equally bonded in both adult musk oxen and calves, and they feed side by side, even if they sometimes keep yearlings at bay. A breeding pair also hunts hares together, but yearlings are also included in the hunt, and the breeding male appears to be more consistent than the female (Mech 1995b).
Both the breeding female and male mark with their scent, and either can initiate dual marking (Haber 1977; Rothman and Mech 1979), depending on what is waiting ahead at the particular moment of migration. For example, during a 4 km migration on 16 July 1993, a breeding pair on Ellesmere Island double scent-marked three times, two of which were initiated by the male. Both the female and the male raise their hind leg when marking with urine, but the male raises his leg higher (possibly for anatomical reasons) and both sometimes scratch the ground associated with marking.
During the early stage of pup care, a breeding pair exhibits a clear division of labor, with the female caring for the burrow territory and nursing the pups (Packard et al. 1992) and the male hunts away from the burrow and brings food to the female and pups (Mech et al. 1999).
The male shows considerable urge to surrender food to the breeding female. For example, on July 8, 1992, when the Ellesmere male and female were equidistant from me in different directions, I threw the carcass of an adult rabbit weighing about 5 kg to the male. The male grabbed it between his jaws, but the female immediately rushed to him, grabbed the carcass from his mouth and took it to the burrow. The male made no attempt to keep or retrieve the carcass. I then gave the male another rabbit of the same size. He ate the head, but took the rest of the carcass 0.5 km to the female and handed it over to her. The female grabbed it. Experiments with smaller pieces gave similar results.
However, the breeding female seems to have the upper hand in keeping other members of the pack away from the pups, especially when the pups are less than 3 weeks old. In the Ellesmere Island pack, a breeding female would often run to her young pups when approached by a breeding male or another wolf.
In addition, a breeding male adopted a submissive posture when approaching a breeding female caring for young pups. On June 26, 1990, I observed a breeding male approach a burrowing female "flagging his tail and body excitedly." I observed a similar situation on May 18, 1990 in Denali Park, Alaska, where radio-tagged male 251 of the Headquarters herd (Mech et al. 1998) approached a breeding female in a burrow with pups 307 by “wriggling” by flapping its rear and tail similar to how a submissive animal approaches a dominant one. The female emerged from the burrow and the male ejected food for her. These were the only times I saw a breeding male submit to any other wolf, and this seems to indicate that the breeding female temporarily dominates even the breeding male before the pups emerge from the den.
The breeding female takes care of the puppies and protects them more than any other member of the herd. For example, mothers were the only pack members I saw picking up and carrying puppies. In addition, I observed in one case how a breeding female of the Ellesmere Island herd behaved most aggressively with a muskox (L.D. Mech) standing at the entrance of the burrow. This coincides with the observational data of Joslin (1966) and Clark (1971). However, Murie (1944) stated that the breeding male was the most aggressive in scaring grizzlies away from the den containing cubs (Ursus arctos).
Conclusions
The aforementioned observational data show that, at least in summer, the social interaction of wolf pack members does not differ in intensity and quality from the relationships of members of any other group of related individuals. Even widespread dominance hierarchies primarily reflect the natural sex, age and reproductive ability structure of group members, in which the breeding male dominates all others with body care, and the breeding female receives food from the male when she cares for young pups.
Thus, a typical wolf pack should be viewed as a family in which adult parents manage the group's activities based on a division of labor system, in which the female primarily controls activities such as care and protection of pups, and the male foraging and related movement (L.D. Mech).
Dominance expression is not common, except when fighting for food. In this case, herd members let the parents capture the food and distribute it to the youngest offspring. Active submission seems to occur primarily for the purpose of begging for food or as a motivator for food gathering (Mech 1970). Further investigation into the role of active and passive submission between a breeding female and male when offspring are not present is needed.
Acknowledgments
The project was supported by the National Geographic Society, the United States Fish and Wildlife Service, the United States National Biological Service (now the United States Geological Survey's Biological Resources Division), and the United States Department of Agriculture's North and Central Forest Experiment Station. Logistical support was provided by the Polar Continental Shelf Project (PCSP), Natural Resources Canada; Atmospheric Environment Services, Environment Canada; and High Arctic International. Permits were issued by the Northwest Territories Department of Renewable Natural Resources and the Grise Fiord Hunting Association. I thank the following field assistants for logistical assistance: L. Adams, L. Boitani, D. Boyd, N. Gedgaudes, C. Johnson, J. Hutchinson, N. Gibson, T. Lebovsky, M. Maule, M. Ortiz, J. Packard, R. Peterson, R. Ream, L. Shaffer, R. Sternal, and U. Swain. In addition, I thank R.O. Peterson for reviewing and suggesting improvements to an earlier draft of the manuscript. This is PCSP document 003298.
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