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Rochelle Yuan
Writing 39c
Prof. Haas
July 24, 2016
Review of the Scientific Literature: The Social Behaviors of Canines
In Empire of the Summer Moon, S. C. Gwynne writes of an 1860 army attack on a camp
in which the first canines had started defend their masters; this was the beginning of the humanwolf domesticated relationship. Barry Lopezs classic 1978 book on the subject, Of Wolves and
Men, details how the wolf exerts a powerful influence on the human imagination (Humans
versus Wolves). Now, scientific researchers believe that these canines have a human-like social
cognition, (Safina 223) as the studies I shall review here as examples of captive and freeranging canine social behavior, specifically the wolf and occasionally its relation to its
counterpart, the dog. Much of the violence toward wolves has been fueled by human
imagination, and often what humans imagine to be true about wolves is countered by close
observation (Humans versus Wolves). Therefore, I shall emphasize early accounts of the
human documentation of canines up to how they have been currently studying them in 2016. The
review begins with the effects of domestication on social behaviors and development as
documented in 1982 shown by a comparison between timber wolves and Alaskan Malamutes
raised in the same environment. It then addresses whether grey wolves show a hierarchy of
dominance or conflict management through agonistic interactions and observations of captive
wolves in 1982 in comparison to free-range wolves in 2014, respectively. I will then follow the
social play of captive, young wolves, slightly touching on adult ones as well, and how their

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social behavior is affected by relationship, aggression, or sex, or if they would display an
egalitarian-style behavior despite contrasting levels of dominance. The research reviewed here
focuses on the social behaviors of wolves to justify that human and wolf antics and behaviors are
unequivocally similar. Its even been said that no two species are more alike than wolves and
humans (Safina 236). Knowing this, how then do we proceed with what we have been doing,
such as in continuing to allow the mistreatment of such animals and diminishing what little land
they have left because of our own actions?
Adolescent humans and wolves behave in a way that is more juvenile because they are
still growing and learning, a common factor that applies to both species. Harry Frank and Martha
Gialdini Franks 1982 study On the effects of domestication on canine social development and
behavior published in Applied Animal Ethology, compares four Eastern timber wolves and four
Alaskan Malamutes, who were fostered by the same female wolf and reared under conditions
deemed natural only to domestic dogs, to discuss the well-known aggression in wolves and the
effects of domestication on the same type of behavior in domestic dogs (508). The Franks noted
how the wolf pups show agonistic behavior that may be innate, but that also involves their
maturation and interaction probably mediated by an innate teaching mechanism, considering
how they were not raised in the wild; on the other hand, the effects of domestication on
dominance behavior in dogs cause selection pressures against aggression to have relaxed, as dogs
probably elicit submissive responses more for attention rather than in response to domination
(517-519). It appears that as wolves transitioned to dogs in human households and became more
domesticated (less aggressive), humans have [also] undergone a reduction in environmental
awareness in parallel to domestic species (Safina 235) according to archaeologist Colin Groves,

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such that they relied on each other for protection (by wolves) or for food and shelter (by
humans).
Conflict is a part of every relationship, whether man or wolf. Besides the fact that wolves
and humans alike have domesticated themselves (Safina 234) and evolved with each other, a
hierarchy still stands in each species. Greg Moran, obtained a Ph.D. focused on the detailed
structure of wolf social interaction, leading to his study of Long-term patterns of agonistic
interactions in a captive group of wolves (Canis lupus) that was published in the well-known
journal, Animal Behaviour. Moran recorded agonistic interactions in a group of captive, grey
wolves for 21 months from a trailer so he could view them in a large open area (75-76). He
described various aspects of their interactions in as detailed a manner as possible and categorized
different types of approaches and described relationships between different wolves, leading to a
total of 2823 interactions of all kinds (77-81). He thus concludes that some aspects of the
distribution of agonistic interactions observed during the study are consistent with the predictions
of an underlying dominance hierarchy social organization. Nevertheless, he could not accurately
claim that there is this dominance hierarchy because the agonistic interactions alone cannot
determine social relationships, only partially, especially since his study was limited to a small
number of pairs (81-83). Moran was able to see a sort of hierarchical social organization within a
pack, similar to how humans live with families that usually contain a male and female head,
alphas, and subordinates. Often, the subordinates would listen to their alphas in both species or
end up fighting, trying to assert dominance.
Tying into this struggle for dominance, Candice Baan, from the University of Neuchtel
in Switzerland, observed free-living wolves in Yellowstone National Park in 2008-2009, and in

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2014, with fellow researchers in her field, published Conflict management in free-ranging
wolves, Canis lupus in the journal Animal Behaviour. Baan and her team acquired their
evidence by video, filming two free-ranging packs of grey wolves in Yellowstone National Park,
who were accustomed to humans observing them, and recording all possible social interactions
so they could analyze and document them, while not using any kind of radio-collaring (Baan
328). Because of wolves high interdependence on each other for survival, Baan discusses how
reconciliation, although mostly asymmetric (initiated by subordinates towards dominants), is
highly favored in free-ranging wolves in order to reduce social tension and restore social peace
(331). However, the results suggest that this occurs because of the benefits the subordinates
would derive from cooperative hunting, cooperative breeding, and cooperation in defense of the
territory and related resources from their stronger and more dominant counterpart (Baan 332).
More often than not, subordinates would be the ones who end up taking the first step in
reconciling with their higher-ups. This could happen in said packs, studied by Baan and her
team, or in families in the human world. Of course, the more dominant individual would have the
best chance at helping the family or pack survive, so its inevitable that the more submissive
individual would benefit the most out of a healthy relationship between them.
Breaking away from the idea of conflict within packs and families, I move onto more
jovial topics, such as in social play; research in this field adds to our growing knowledge of the
complex cognitive abilities of wolves. Giada Cordoni, Evolutionary Biologist from the
University of Pisa, wrote Social play in captive wolves (Canis lupus): not only an immature
affair in 2009, which has been cited by 14 other articles and can be found in the journal,
Behaviour. Because animal and human social play represent an important part in social
assessment during early ages, Cordoni studied how a captive group of grey wolves housed in the

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Pistoia Zoo (Italy) performed more social play compared to solitary play and how their play was
not affected by relationship quality, aggression levels, or gender (1363-1367). She defines a play
session, an aggressive encounter, and a mating period, and reports on recordings of solitary and
social playful behavioral patterns. With her recordings, she then discusses each of her five
predictions in accordance to her findings, such as how solitary and social play sessions were
most frequent during non-mating periods and that aggression was higher during mating periods
than non-mating periods (1368-1379). Children enjoy playing with their peers rather than
themselves just as much as wolves. Then, when people mature and find significant others to have
children, they are less likely to approach others for said play sessions, especially when raising
young. Regardless of the difference between human and wolf mating relationships, the concept is
analogous.
Cordonis research on social play leads to Jennifer Esslers co-writing of Play Behavior
in Wolves: Using the 50:50 Rule to Test for Egalitarian Play Styles in 2009 and 2012, which
was published 2016. Essler, who works with wolves and dogs at the Wolf Science Center, found
that wolves would maintain the same dominance relationship in the play context, leading to the
conclusion that wolves would not follow the 50:50 rule (Essler 3). The researchers investigated
two wolf packs of hand-raised wolf puppies aged three to five months in relation to the 50:50
rule, which is the hypothesis where wolves play equitably in which normally, the stronger partner
would take a disadvantageous position, and to their dominance relationship outside of play
(Essler 4-5). They defined play by different indicators, observed submissive and dominance
interactions (but not aggressive ones) and charted them into different tables, while also using
models to further test whether individuals engaged in behavior achieving a 50:50 ratio (Essler 69). Although different levels of dominance did not affect whether or not dyads engaged in play,

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after analyzing the data, they concluded that this rule was not supported in the wolf puppies
during dyadic play, and that adult wolves did not demonstrate self-handicapping behaviors but
that dominant puppies did not take advantage of their higher position (Essler 10-15). Just like
between siblings, a stronger sibling does not often create an evenly matched playing field for a
weaker sibling, but is more likely to be lenient on this matter compared to adults, who are more
unlikely to exhibit self-handicapping behavior due to their desire for personal gain, possibly for
selfish reasons.
In conclusion, the social behavior of free and captive, young and old wolves detailed in
countless studies by prominent researchers sheds light on the topic of human and wolf similarity,
which forces us to acknowledge the impact our actions have on canine life. They are so similar to
us, yet we treat them so differently. There was once a time when they thrived in the territory they
lived; but, once Europeans arrived, by 1930, humans had wiped wolves off 95 percent of their
holdings in the lower forty-eight states (Safina 169). Over several recent years, the federal
government weakened wolf protections, saying how 30 breeding pairs and 300 wolves in the
entire northern Rocky Mountain region constituted a recovered population (Safina 169-170).
This caused hunting to begin occurring again and no limit to it, something that had originally
wiped out so much of the population before. People live in a world where they feel entitled to
being better than all other non-human animals; they dont consider the fact that animals such as
wolves, depend on each other and the land to survive, much as people need companionship and
space to survive. Essentially, we cast out wolves because they are not like us, rather, they
become a metaphor for the feral and pre-civilized, the gang, for the people living outside the
bounds of convention and conformity (Safina 170). Keeping in mind the knowledge of how
humans have come to take-over the land wolves once roamed, Douglas W. Smith, who received

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his Bachelor of Science in Wildlife Biology from the University of Idaho in 1985 and is the
current project leader for the Yellowstone Gray Wolf Restoration Project in Yellowstone National
Park, studying wolves for 23 years, speaks in a Q & A about wolves in Should people be afraid
of wolves? since they generally are in regards to the unknown:
https://www.nps.gov/yell/learn/photosmultimedia/qa-wolves.htm. In this video, Smith
emphasizes three points about why humans are afraid of wolves and that during the entire 20th
century in North America, there were about 20 attacks on humans. This is such a low number
when compared to a whole century, but yet, people continue to believe that wolves are so
dangerous, and that they will automatically attack a human. After learning about how wolf and
human social behaviors are not so different after all, the way we treat them should now be taken
into account.

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Works Cited
Baan, Candice, Ralph Bergmller, Douglas W. Smith, and Barbara Molnar. "Conflict
Management in Free-ranging Wolves, Canis Lupus." Animal Behaviour 90
(2014): 327- 34. doi:10.1016/j.anbehav.2014.01.033
Cordoni, Giada. "Social Play in Captive Wolves (Canis Lupus): Not Only an Immature Affair."
Behaviour 146.10 (2009): 1363-385. doi:10.1163/156853909X427722
Essler JL, Cafazzo S, Marshall-Pescini S, Virnyi Z, Kotrschal K, Range F. Play Behavior in
Wolves: Using the 50:50 Rule to Test for Egalitarian Play Styles. PLoS ONE 11.5
(2016): e0154150. doi:10.1371/journal.pone.0154150
Frank, Harry, and Martha Gialdini Frank. "On the Effects of Domestication on Canine Social
Development and Behavior." Applied Animal Ethology 8.6 (1982): 507-25.
doi:10.1016/0304-3762(82)90215-2
Moran, Greg. "Long-term Patterns of Agonistic Interactions in a Captive Group of Wolves
(Canis Lupus)." Animal Behaviour 30.1 (1982): 75-83. doi:10.1016/S0003
3472(82)80239-X
Safina, Carl. Beyond Words: What Animals Think and Feel. New York, NY: Henry Holt,
2015.
"Humans versus Wolves." WildEarth Guardians. N.p., n.d.
Should People Be Afraid of Wolves? Perf. Douglas Smith. Q & A: Wolves, Doug Smith. National
Park Service, n.d.