Alpha Status Dominance and Division of Labor in Wolf P - PDF document

on non-natural foraging and food-provisioning andthe travels associated with them.This resource is based on the following source (Northern Prairie Publication 1078):Mech, L. David. 1999. Alpha status, dominance, and division of labor in wolf packs. Canadian Journal of Zoology 77:1196-1203.This resource should be cited as:Mech, L. David. 1999. Alpha status, dominance, and division of labor in wolf packs. Canadian Journal of Zoology 77:1196-1203. Jamestown, ND: Northern Prairie Wildlife Research Center Home Page. http://www.npwrc.usgs.gov/resource/2000/alstat/alstat.htm (Version 16MAY2000).Table of Contents! Introduction! Methods! Results and Discussiono address: North Central Research Station, 1992 Folwell Avenue, St. Paul, MN55108, U.S.A.(e-mail: Mechx002@tc.umn.edu).IntroductionWolf (Canis lupus) packs have long been used as examples in descriptions of behavioral dominance andalpha status has gained considerable prominence (Schenkel 1947; Rabb et al. 1967; Fox1971b; Zimen 1975, 1982), and the prevailing view of a wolf pack is that of a group ofindividuals ever vying for dominance but held in check by the "alpha" pair, the alphamale and the alpha female (Murie 1944; Mech 1966, 1970; Haber 1977; Peterson 1977).Most research on the social dynamics of wolf packs, however, has been conducted onwolves in captivity. These captive packs were usually composed of appropriate, for the wolf pack is not such an assemblage. Rather, it is usually a andNelson 1990), or a dead parent is replaced by an outside wolf (Rothman and Mech 1979;Fritts and Mech 1981) and an offspring of opposite sex from the newcomer may thenreplace its parent and breed with the stepparent (Fritts and Mech 1981; Mech and Hertel1983).Nevertheless, these variations are exceptions, and the pack, even in of the alpha wolf as a "topdog" ruling a group of similar-aged compatriots (Schenkel 1947; Rabb et al. 1967; Fox1971a; Zimen 1975, 1982; Lockwood 1979; van Hooff et al. 1987) is particularlymisleading.Because wolves have been persecuted for so long (Young and Goldman 1944), they females under naturalconditions, and about the role of each in the pack and how dominance relates to these thesis, presented onlya quantified summary of the pack's hierarchical relationships, and Haber (1977) describedhis interpretation of a pack's social hierarchy but gave no supporting evidence. Thus, noone has yet quantified the hierarchical relationships in a wild wolf pack.Here I attempt to clarify the natural wolf-pack social order and to advance our knowledgeof wolf-pack social dynamics by discussing the alpha concept and social dominance and in free-livingpacks.This study was conducted during the summers of 1986-1998 on Ellesmere wolves prey a;National Geographic Society 1988).We noted each time a wolf submitted posturally to another wolf. Usually this deferencewas characterized by "licking up" to the of the other. I did notconsider "standing over" a dominance behavior (L.D. Mech, submitted for publication). 2The following is a summary of generalizations documented in the previous references,together with new quantified findings.Alpha status"Alpha" connotes top ranking in some kind of hierarchy, so an alpha wolf is by definitionthe top-ranking wolf. Because among wolves in captivity the hierarchies are gender-based, there are an alpha male and an alpha female (Schenkel 1947).The way in which alpha status has been viewed historically can be seen in studies inwhich cub, thepotential alpha animal (emphasis mine) of the (Fox1971a, p.307). This view implies that rank is innate or formed early, and that somewolves are destined to rule the pack, while others are not.Contrary to this view, I propose that all young wolves are potential breeders and thatwhen social rank (Fox 1971b; Fox and Andrews1973). Secondly, wolves in captivity breed readily, and I know of no mature captiveindividuals that failed to breed when paired apart from a group, as would be the case ifthere were inherently low-ranking, nonbreeders.Third, in the wild, most wolves disperse from their natal packs and attempt to pair withother maturing pack to its young offspring, so"alpha" adds no information. Why not status, which is trivial information, but its role as pack progenitor,which is critical information.The matriarch and can more appropriately be called the alphas. Evidence for such acontention would be an older breeder consistently dominating food disposition or thetravels of the pack.The point here is not so much the terminology but what the terminology falsely implies: arigid, force-based dominance hierarchy.The degree to which these arguments apply to (K Ina natural wolf pack, the dominance rules bear no resemblance to those of the peckingorder, that of a group of similar individuals competing for rank.The only consistent demonstration of rank in natural packs is the animals' postures duringsocial interaction. Dominant wolves assume the classic canid standing posture with In fact, submission itself may be as important as dominance interms of promoting friendly relations or reducing social distance.Schenkel (1967), who promoted the importance of submission, and I find active submission and food-begging indistinguishable. Thebegging or submissive wolf approaches another wolf excitedly, wagging the tail,lowering the ears, and "licking up" to the other wolf. The other wolf may or may not (Mech et al. 1999). In passive submission,the submissive wolf rolls over on its side or back, and the dominant wolf sniffs its groinor genitals (Schenkel 1967). Active submission was more common posturally to thebreeding male, both actively and passively (Schenkel 1967). The yearlings and 2-year-oldwolves and one old post-reproductive female submitted to both breeders. These Two-year-old female89Ñ421Two- year-old male400Ñ4Total25c9c2642cNote: Interactions do not include "standing over" or involve food, except for "food-begging."aThese are the yearlings in Table 2.bThe female parent dominated the 2-year-old female for 15 min at one of these times. Another time, when it wasunclear whether the female parent or 2-year-old female dominated, is not included.cFor male parent versus female parent, = 3.99, P = 0.05.Table 5. Dominance interactions, i.e., the number of times individual wolves dominated others or were submitted to,among breeders and a post-reproductive female in the Ellesmere Island wolf pack in the summers of 1990 and 1991(pups were present and the male parent was the same as in all other years in the study except 1998).!MaleparentFemaleparentaPost-reproductivefemalebTotalMale parentÑ1c00Female parenta35Ñ136Post-reproductive femaleb2617Ñ43Total6118180dNote: Interactions do not include "standing over" or involve food, except for "food-begging."aYearling female in 1988 (Table 1) and female parent in 1990-1996.bFemale parent in 1988 and 1989 (Table 1).cMale deferred when approaching a female and young pups in a den.d= 12.64, PThat these submission male, which instantly snatched it fromher. He refused entreaties by both that female and even the breeding female and to be an ownership zone (Mech 1970)around the mouth of each wolf, and regardless of the rank of a challenger, the owner triedto retain the food Failed1991-06-22Post-reproductive femaleBreeding maleSucceeded1993-07-11Yearling femaleYearling femaleFailed1994-07-16Pups and yearling maleYearling femaleFailed1996-07-15Pups/breeding femaleBreeding maledSucceeded1998-07-07Breeding femaleBreeding maleFailedaDoes not include the breeding female taking food from the breeding male.bYearling female had brought food to the pups and snapped at the breeding female when she stole it.cYearling female, who had brought a hare, stood guard near the pup.dBreeding female failed to stop the breeding male.Two other behaviors among pack members could have been dominance-related, althoughdata were insufficient to be certain. They were "standing over" and "hugging" (L.D.Mech, see footnote). In "standing over," one wolf would stand over (Schenkel 1947) alying wolf, positioning its groin above the nose of the lying wolf. Sometimes the lyingwolf sniffed at the groin or genitals of the standing wolf.Schenkel (1947) saw "standing over" only during "peaceful" times and did not seem toconsider it dom at the top and offspring or non-breeders subordinate, are so automatic and Zimen (1982)described for captive wolves, as Clark (1971) also noted. Similarly, pups defer to adultsand older siblings in the same automatic, peaceful way. When or whether a rank orderdevelops among pups is in dispute (cf. Zimen 1975 and Fox and Andrews 1973; Haber1977), and I cannot shed any light on that issue. Even among yearlings and "hostility" might bedue to different viewpoints of the observers. control when subordinates feed (Mech 1988; National Geographic 1988).Similarly, pups are subordinate to both parents and to older siblings, and sometimes accompanyadults on foraging trips at an earlier age than do subordinates (Haber 1977).Dominance between the breeding male and femaleThe relationship between the (L.D. Mech, submitted for publication).1Whether each gender has its own dominance tend to disagree. Clark's (1971) data indicated male and breeding female are separated, recognize each other, andthen meet, the breeding female approaches the male in a typical subordinate posture: withthe tail down or between the legs, body crouched or on the ground, ears back, and nosepointed the male as described above while shepossessed a long on 16 July 1993, during 4 km of travel, theEllesmere Island breeding pair double-scent-marked three times; the male initiated two ofthem. Both male and female raise a hind leg during urine-marking, although the maleraises his higher, possibly in food back to the female and the pups directions, I threw the male an adult hare carcass weighing about 5 kg. Themale to keep or regain the hare. I then gave themale a second hare of the same size. He ate the head and then took the rest of the carcass0.5 km to the female and gave it to her. She cached it. Similar tests with smaller piecesyielded similar results.Nevertheless, in keeping other pack members away from young pups, the breedingfemale seems to reign supreme, especially tending young his back end and tail like a interactions among wolf-pack members are not very different in intensity or quality from those among members ofany other group of related individuals. Even the much-touted wolf dominance hierarchyis primarily a natural reflection of the age, sex, and reproductive structure of the group,with the breeding male dominating all others posturally and the breeding femalegarnering food from the male while she is tending young pups.The typical wolf pack, then, should be viewed as a family with the adult parents guidingthe activities of the group and sharing group leadership in a division-of-labor system inwhich and food-provisioning and the travels associated a food-begging Association of J.Packard, R. Peterson, R. Ream, L. Shaffer, R. Sternal, and U. Swain. I also thank R.O.Peterson for critiquing an earlier draft of the manuscript and suggesting improvements.This is PCSP Paper 003298.ReferencesAsa, C. S., Mech, L. D., Seal, U. S., and Plotka, E. D. 1990. The influence of social and endocrine factors on urine-marking by captive wolves (Canis lupus). Horm. Behav. 24:497-509.Clark, K. R. F. 1971. Food habits and behavior of the tundra wolf on central Baffin island. Ph.D. thesis, University of Toronto. Toronto, Ont. [Available from the National Library of Canada, Ottawa, Ont.]Creel, S., and Creel, N. M. 1996. Rank and reproduction in cooperatively breeding African wild dogs: behavioral and endocrine correlates. Behav. Ecol. 8:298-306.Darwin, C. 1877. The expressions of the emotions in man and animals. Translated by J. V. Carus. 3rd ed. Stuttgart, Germany.Estes, R. D., and Goddard, J. 1967. Prey selection and hunting behavior of the African wild dog. J. Wildl. Manage. 31:52-70.Fentress, J. C., Ryon, J., McLeod, P. J., and Havkin Schjelderup-Ebbe, T. 1922. Beitrage zur Sozialpsychologie des Haushuhns. Z. Psychol. 88:225-252.Seal, U. S., Plotka