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Wolf packs often work cooperatively, as in this bison hunt at Yellowstone National Park.
A pack of coyotes in Yellowstone National Park
A hunting pack of African wild dogs
Pack size distribution of the African wild dog.

Pack is a social group of conspecific canids. Not all species of canids form packs; for example, small canids like the red fox do not. Pack size and social behaviour within packs varies across species.

Contents

Pack behavior in specific speciesEdit

African wild dogs (Lycaon pictus) live and hunt in packs. Males assist in raising the pups, and remain with their pack for life, while the females leave their birth pack at about the age of two and a half years old to join a pack with no females. Males outnumber the females in a pack, and usually only one female breeds with all of the males. African wild dogs are not territorial, and they hunt cooperatively in their packs, running down large game and tearing it apart. They cooperate in caring for wounded and sick pack members as well as the young.[1]

Gray wolves (Canis lupus) usually live in packs which consist of the adult parents and their offspring of perhaps the last 2 or 3 years. The adult parents are usually unrelated and other unrelated wolves may sometimes join the pack.[2] Wolves usually hunt in packs, but they hunt alone in spring and summer months when there is plenty of prey available. They are found in both Eurasia and North America.

Black-backed jackals (Canis mesomelas) in Southern and Eastern Africa and coyotes (Canis latrans) which are only found in North America have a single long term mate, but they usually either hunt alone or in pairs.[3][4] Both parents care for the young, and the parents and their current offspring are the pack. They occasionally cooperate in larger packs to hunt large game.

The Ethiopian wolf (Canis simensis) has different social behavior from the gray wolf: pack members hunt alone for rodents, and come together mainly to defend their territory from other packs.[5]

The Social Dynamics and Hierarchy of a Gray Wolf PackEdit

The gray wolf is one of the Earth’s most iconic pack-living animals, known for their fierce loyalty to and reliance on the other members of their group[6]. These intense social bonds have led to their tremendous success as hunters, as well as allowed for better territorial defense and preservation of the individual. Despite hierarchy, all individuals benefit from being a member of the pack; the weak are supported by the efforts of stronger wolves[7], and higher ranking individuals enjoy better and larger kills than could be taken on their own[8]. Protection is granted by sheer number, and larger, more plentiful territory can be won and sustained[9]. Care and protection of young is shared, and knowledge can be passed down through generations, creating a unique culture within each group[7].  In wolf society, the pack is greater than the sum of its parts.

The pack often consists of 5-10 (though in areas of high prey abundance can be up to 20)[10] mostly related individuals, with an often unrelated breeding pair, known as the alphas[11], and their various offspring. Occasionally, there may also be a handful of other wolves which can be related or not[12]. This familiarity lends to their intense bonding. Members of the pack can spend most of their lives alongside one another, resulting in a highly efficient and formidable unit. The pack will fiercely defend their own against enemies and territorial invasion, and it is not uncommon to see wolves die in this effort[6]. Despite this strong allegiance, membership may be fluid and is subject to change[13]. Outside wolves may be shunned or, though rarely, accepted, depending on the specific circumstances. Genetic variability can become limited within such an interrelated group, and so conditions for gene flow must exist[11]. Outside wolves can provide these opportunities[7]. A pack may be accepting of another wolf into their group if it is a distant relative[12], if reproduction rates are low due to loss or infertility of an alpha, or if their numbers are significantly reduced[11].

The Lone WolfEdit

These singular outside wolves, often referred to as lone wolves, are vulnerable to food scarcity and territorial attacks, and generally make up less than 15% of a wolf population. Lone wolves may result from young offspring leaving their parental pack, harassed subordinates who chose to disperse, or as a result of lost dominance; lower ranking wolves often challenge the alpha for leadership, and if won, the former alpha will often leave the pack. In times of prey scarcity, low ranking wolves may choose to go off on their own if the pack cannot supply sufficient food. These lone wolves may then attempt to join into an existing wolf pack or, more commonly, find a mate and begin a new pack family as the alphas[11].

The Alphas (Breeding Pair)Edit

Within the wolf pack, the alphas, now more commonly referred to as the breeding pair, are the only wolves in the group which breed and produce offspring; they are the matriarch and patriarch of the family[10]. The alpha or parent role consists of one male and one female which are pair-bonded and usually unrelated, although they may be distantly related or, in some cases, fully related. This variable degree of relation and inbreeding can significantly impact the viability of the pack, and inbreeding is known to commonly occur in both captive and wild wolf populations. It is common for an aging or sick alpha to be replaced by one of their offspring, with this new alpha now becoming the mate to his or her own parent. This inbreeding depression can cause weak pups, lowered reproduction rates, blindness, and reduced longevity[11].  

The alphas represent the most dominant rank in the wolf pack, and so are held in the highest esteem by the other members of the unit[14]. Along with sole breeding, the alphas are responsible for control and leadership of the pack[15], including establishing pack territory, leading hunts, and accepting or rejecting outside wolves[16]. The loss of an alpha can spell complete dissolution for the entire pack[12].  Because of this, the alphas always eat first, and are always awarded the biggest and best portions of the kill[15]. Lower-ranking individuals showing disrespect to the alpha will be punished, or sometimes banished from the pack. The societal rules existing within the wolf pack do so in order to preserve the whole and protect the alpha, and these rules must be imposed. Due to the alphas’ vital importance to the viability of the pack, their safety is a first priority, and so the majority of enforcement falls to the next-in-command: the betas[14].

The BetasEdit

Because of their role, the betas are often referred to as the “enforcers” of the pack[14]. These wolves rank directly below the alphas, and generally consisting of one or two individuals[15]. They are often the wolves which will first confront a newcomer or potential threat[14]. The betas serve both as enforcement and second-in-command, often taking over control while the alphas are away or preoccupied with breeding. Should anything happen to an alpha, a beta is next in line to satisfy the vacancy[17]. As such, these wolves are also highly respected within the pack, and receive the next choicest portion of the kill[14]. The betas are believed to be the most trusted and most bonded companions of the alpha pair, acting as “generals” in the pack hierarchy[15].

Mid-Ranking WolvesEdit

All other wolves in the group are considered subordinates, though they serve important roles in the pack dynamic[15]. The mid-ranking wolves fall between the betas and the lowest rank, the omega. Mid-ranking wolves have been organized and re-organized into various divisions in an attempt to understand their complex interactions, but in reality the middle ranks are constantly interchanging, with dominance awarded to the fittest individual for the particular endeavour at hand[18]. These wolves often serve as babysitters to the pups when the alphas and betas are away, and conditional to the pack’s size, certain individuals may fill the roles of watchers, scouts, teachers, hunters, or protectors, depending on what is needed at the time[15].

The OmegaEdit

Omegas are the lowest ranking members of the group, usually consisting of one individual of either sex. Omegas sometimes represent the weakest of the pack, or may have been demoted to the position as a consequence of bad behavior. Life as an omega can be difficult, eating last and taking the brunt of aggression from the rest of the pack[15]. These individuals may chose to leave the group in search of a higher ranking position or to breed[11], and can be replaced as ranks are shuffled overtime. Older omegas are often allowed to retire, rescinding their position and being looked after by the pack[19]. Alleviating aggression is the omegas principle role, and this is often done through initiating play as a way to burn off excess negative energy[15]. As a result, long-time omegas often bear the scars of their submissive position[14].

The Pups

Importance of the alpha is rivaled only by that of the pups[14]. The fundamental purpose of the pack is the successful production of offspring, and so raising of the litter is a collaborative venture – all members contribute to their development[7]. Despite this committed involvement, pup mortality is high, with researchers citing that only roughly 30% survive their first year of life[20]. Due to this fragility, priority is given to the alphas rather than the pups, as more litters are needed to compensate for this high rate of mortality[14]. Those who survive, however, grow up with the added advantage of being surrounded by numerous care-takers and teachers. There exists a culture within wolf packs, and this is passed on to the offspring by the elders of the group.  Pups learn something from each member of the pack, and attain the vital social skills required to create those powerful bonds upon which wolf societal structure relies[7].

Without a pack a wolf is constantly searching. Like humans, wolves instinctually seek out members of their own species for socialization.  Undoubtedly, this was a commonality early humans recognized, leading to the eventual domestication of the wolf into our modern day companion dogs[7]. Understanding pack dynamics, and the specific roles our canine companions are suited for, is crucial for a healthy and functioning interspecies relationship. Further research and interpretation into the structure and dynamics of wolf hierarchy is ongoing, and will provide a deeper understanding into the roles humans play for both wild and domesticated canines.

 
White huskies dog sledding

Dominance and the alpha wolfEdit

Animals which typically predominate over others are associated with the term alpha. Among pack-living wolves, alpha wolves are the genetic parents of most cubs in the pack. Such access to mating females creates strong selective pressure for intra-sex competition[21].

Wolves show deference to the alpha pair in their pack by allowing them to be the first to eat and, usually, the only pair to reproduce. Wolves use eye contact as an indicator of dominance or submission, but in order to establish a dominant position they often also show physical superiority through playing or fighting. The smaller and more nuclear a pack is, the status of alpha is less likely to be obtained through fighting, and young wolves instead leave the pack to find a mate and produce offspring of their own.[22] Larger or less-nuclear packs may operate differently and possess more complex and flexible social structures.[23][better source needed]

In the case of other wild canids, the alpha male may not have exclusive access to the alpha female;[24] moreover, other pack members may guard the maternity den used by the alpha female; as with the African wild dog, Lycaon pictus.[25]

As dominant roles may be deemed normal among social species with extended parenting, it has been suggested that the additional term alpha is not required merely to describe dominance due to its ubiquity, but should be reserved for where they are the predominant pack progenitor. For instance, wolf biologist L. David Mech stated

"calling a wolf an alpha is usually no more appropriate than referring to a human parent or a doe deer as an alpha. Any parent is dominant to its young offspring, so alpha adds no information. Why not refer to an alpha female as the female parent, the breeding female, the matriarch, or simply the mother? Such a designation emphasizes not the animal's dominant status, which is trivial information, but its role as pack progenitor, which is critical information. The one use we may still want to reserve for alpha is in the relatively few large wolf packs comprised of multiple litters. ... In such cases the older breeders are probably dominant to the younger breeders and perhaps can more appropriately be called the alphas. ... The point here is not so much the terminology but what the terminology falsely implies: a rigid, force-based dominance hierarchy."[26]

Use in dog trainingEdit

One of the most persistent theories in dog training literature is the idea of the alpha wolf, an individual gray wolf who uses body language and, when needed, physical force to maintain its dominance within the wolf pack. The idea was first reported in early wolf research.[2] It was subsequently adopted by dog trainers.[27] The term alpha was popularized as early as 1976 in the dog training book How to Be Your Dog's Best Friend (Monks of New Skete), which introduced the idea of the alpha roll, a technique for punishing unwanted dog behaviours. Psychologist and dog trainer Stanley Coren in the 2001 book How to Speak Dog says "you are the alpha dog...You must communicate that you are the pack leader and dominant".[28]

Training techniques assumed to be wolf pack related such as scruff shaking, the alpha roll and recommendations to be alpha to the dog continue to be used and recommended by some dog training instructors.[29] It has been suggested that the use of such techniques may have more to do with human psychology than with dog behaviour; "dominance hierarchies and dominance disputes and testing are a fundamental characteristic of all social groups... But perhaps only we humans learn to use punishment primarily to gain for ourselves the reward of being dominant.[30] Most leading veterinary and animal behavior associations, and most contemporary trainers would agree, advocating the use of rewards to teach commands and encourage good communication between owners and their pets. Modern best practices dictate an abandonment of outdated "pack" methods.[31] Some canine behaviourists suggest that kind, efficient training uses games to teach commands which can be utilised to benefit the owner's everyday life.[32]

See alsoEdit

ReferencesEdit

  1. ^ Animal Diversity Web. "Lycaon pictus: Information". University of Michigan. Retrieved 21 April 2008.
  2. ^ a b L. David Mech. "Schenkel's Classic Wolf Behavior Study Available in English". Retrieved 21 April 2008. ...Schenkel’s 1947 “Expressions Studies on Wolves”, the study that gave rise to the now outmoded notion of alpha wolves. That concept was based on the old idea that wolves fight within a pack to gain dominance and that the winner is the “alpha” wolf. Today we understand that most wolf packs consist of a pair of adults called “parents” or “breeders,” and their offspring.
  3. ^ Animal Diversity Web. "Canis mesomelas: Information". University of Michigan. Retrieved 21 April 2008.
  4. ^ "Coyotes 101 – CoyoteSmart". www.coyotesmarts.org. Retrieved 14 April 2018.
  5. ^ "Ethiopian Wolf". Animal Info. 2005-03-07. Retrieved 2015-02-26.
  6. ^ a b 1.      Nowak, K. (2015, June 15). 10 Things We Can Learn From Wolves. Retrieved February 20, 2019, from http://www.thecontentwolf.com/life/10-things-we-can-learn-from-wolves/
  7. ^ a b c d e f 1.      Living With Wolves. The Social Wolf. Retrieved February 20, 2019, from https://www.livingwithwolves.org/about-wolves/social-wolf/
  8. ^ 1.      Schmidt, P., & Mech, D. (1997). Wolf Pack Size And Food Acquisition. Am Nat, 150(4), 513-517. doi:10.1086/286079
  9. ^ 1.      Pack Power. (2015, March 30). The Economist. Retrieved March 2, 2019, from https://www.economist.com/science-and-technology/2015/05/30/pack-power
  10. ^ a b 1.      Western Wildlife Outreach. Wolf Ecology and Behavior. Retrieved February 24, 2019, from http://westernwildlife.org/gray-wolf-outreach-project/biology-behavior-4/
  11. ^ a b c d e f 1.      Tamonga Nature Park. (n.d.). Wolf Pack Society. Retrieved February 24, 2019, from http://www.tamonganaturepark.com/wolf-pack-society.html
  12. ^ a b c 1.      Mech, D. (1999). Alpha Status, Dominance, and Division of Labor in Wolf Packs. Canadian Journal of Zoology, 77, 1196-1203. Retrieved March 17, 2019, from https://www.wolf.org/wp-content/uploads/2013/09/267alphastatus_english.pdf.
  13. ^ 1.      International Wolf Center. (2014, January). Wolf FAQ's. Retrieved March 2, 2019, from https://www.wolf.org/wolf-info/basic-wolf-info/wolf-faqs/#g
  14. ^ a b c d e f g h 1.      Ellis, S., & Junor, P. (2009). The Man Who Lives With Wolves (1st ed.,). New York, NY: Harmony Books.
  15. ^ a b c d e f g h 1.      Ford, S. (2018, September). Wolf Pack Dynamics. Retrieved March 17, 2019, from https://wolvestuff.com/blogs/wolvestuff-blog/wolf-pack-dynamics
  16. ^ 1.      Foden, S. (n.d.). WOLF PACKS & THEIR PECKING ORDER. Retrieved March 23, 2019, from https://animals.mom.me/wolf-packs-pecking-order-3463.html
  17. ^ Dutcher, J. (n.d.). Beta Wolf. Retrieved March 20, 2019, from https://www.livingwithwolves.org/wp-content/uploads/2014/08/beta_cs4.pdf
  18. ^ 1.      Dutcher, J. (n.d.). Mid-Ranking Wolves. Retrieved March 20, 2019, from https://www.livingwithwolves.org/portfolio/mid-ranking-wolves/
  19. ^ 1.      Dutcher, J. (n.d.). Omega. Retrieved March 20, 2019, from https://www.livingwithwolves.org/portfolio/the-omega/
  20. ^ 1.      Smith, D. (2015, March 5). Wolf pup survival a fragile thing. StarTribune. Retrieved March 27, 2019, from http://www.startribune.com/wolf-pup-survival-a-fragile-thing/295226071/
  21. ^ Sillero-Zubiri, Gottelli, MacDonald (1996). "Male Philopatry, Extra-Pack Copulations and Inbreeding Avoidance in Ethiopian Wolves (Canis simensis)". Behavioral Ecology and Sociobiology. 38: 331–340 – via JSTOR.CS1 maint: Multiple names: authors list (link)
  22. ^ Mech, L. David. 1999.
  23. ^ Dutcher, Jim and Jamie. Wolves At Our Door, Simon and Schuster, 2002
  24. ^ Gary Greenberg and Maury M. Haraway. 1998
  25. ^ C. Michael Hogan. 2009
  26. ^ Mech, L. David (1999). "Alpha Status, Dominance, and Division of Labor in Wolf Packs". Jamestown, ND: Northern Prairie Wildlife Research Center Online. Canadian Journal of Zoology 77:1196-1203. Archived from the original on 2004-12-21. Retrieved 21 April 2008. The point here is not so much the terminology but what the terminology falsely implies: a rigid, force-based dominance hierarchy.
  27. ^ "ClickerSolutions Training Articles - The History and Misconceptions of Dominance Theory". Clickersolutions.com. Retrieved 2015-02-26.
  28. ^ Coren, Stanley (2001). "20". How to Speak Dog (First Fireside ed.). Simon & Schuster. p. 250. ISBN 0-684-86534-3.
  29. ^ "BC Boards". Bordercollie.org. Retrieved 2015-02-26.
  30. ^ Pryor, Karen (August 1999). "4". Don't Shoot the Dog! (Bantam trade paperback ed.). Bantam Books. p. 108. ISBN 0-553-38039-7.
  31. ^ "Position Statement on the Use of Dominance Theory in Behavior Modification of Animals" (PDF). American Veterinary Society of Animal Behavior. Retrieved 22 July 2017.
  32. ^ "Dog behaviour pack leaders debunked". Affinity Dog Training. Retrieved 2013-08-16.