Report

Domestication Does Not Explain the Presence of

Inequity Aversion in Dogs
Highlights Authors
d Pack-living dogs and wolves show both reward and quality Jennifer L. Essler,
inequity Sarah Marshall-Pescini,
Friederike Range
d More dominant individuals were more inequity averse for
both dogs and wolves Correspondence
d Inequity conditions influenced social behaviors in jennifer.essler@vetmeduni.ac.at
subsequent interactions
In Brief
d Inequity aversion is probably linked to the evolution of Essler et al. show that as pack-living dogs
cooperation in dogs and wolves and wolves react similarly to unequal
outcomes, this response was not a trait
that dogs evolved over the course of
domestication. Their results support the
hypothesis that inequity aversion is
closely linked to the evolution of
cooperation.

Essler et al., 2017, Current Biology 27, 1861–1865

June 19, 2017 ª 2017 Elsevier Ltd.
http://dx.doi.org/10.1016/j.cub.2017.05.061
Current Biology
Report
Domestication Does Not Explain
the Presence of Inequity Aversion in Dogs
Jennifer L. Essler,1,2,3,4,* Sarah Marshall-Pescini,1,2 and Friederike Range1,2
1Wolf Science Center, Messerli Research Institute, University of Veterinary Medicine, Vienna, Medical University of Vienna,
€rplatz 1, 1210 Vienna, Austria
and University of Vienna, Veterina
2Comparative Cognition, Messerli Research Institute, University of Veterinary Medicine, Vienna, Medical University of Vienna,
€rplatz 1, 1210 Vienna, Austria
and University of Vienna, Veterina
3Twitter: @jennyessler
4Lead Contact
*Correspondence: jennifer.essler@vetmeduni.ac.at
http://dx.doi.org/10.1016/j.cub.2017.05.061
SUMMARY
Sensitivity to inequity is thought to be an important
mechanism for recognizing undesirable cooperative
partners and thus crucial for the evolution of human
cooperation [1]. This link may not be unique to hu-
mans, as cooperative non-human primates also
react to unequal outcomes [2], whereas non-cooper-
ative species do not [3]. Although this hypothesis has
not been tested in non-primate species, studies re-
vealed that pet dogs show a limited form of inequity
aversion, responding to reward, but not quality ineq-
uity [4–6]. It has been proposed that this primitive
form of inequity aversion was selected for during
domestication and thus absent in their ancestors,
wolves. Alternatively, wolves, which hunt, raise
pups, and defend their territory cooperatively, are
similarly inequity averse as non-human primates, or
at least to the same degree as pet dogs. Testing simi-
larly raised and kept pack-living dogs and wolves, we
found both to be inequity averse when their partner
was being rewarded but they were not for performing
the same action. Additionally, both wolves and dogs
reacted to receiving a lower-quality reward than their
partner. These results suggest that the inequity
response found in pack-living dogs and wolves is
comparable to that observed in non-human pri-
mates; results from studies on pet dogs may be
confounded by the dogs’ relationship with humans.
Consequently, our results suggest that inequity aver-
sion was present already in the common—probably
cooperative—ancestor of wolves and dogs and
thus support the hypothesis of a close link of cooper-
ation and inequity aversion.
RESULTS
Multiple hypotheses have been suggested to account for why
pet dogs (Canis familiaris) show a form of inequity aversion. First,
dogs may have acquired the capacity to respond to inequity due
Current Biology 27, 1861–1865, June 19, 2017 ª 2017 Elsevier Ltd. 1861
to an enhancement of their socio-cognitive abilities brought
about by domestication. More specifically, by evolving alongside
humans, dogs are thought to have gained unique abilities to suc-
cessfully cooperate with both humans and conspecifics [7, 8].
Based on this hypothesis, dogs’ closest living relatives, wolves
(Canis lupus), should not exhibit inequity aversion. Alternatively,
the ‘‘canine cooperation hypothesis’’ suggests that rather than
dogs gaining these abilities during the course of domestication,
they maintained such abilities from their ancestors, and hence
they should be shared with wolves [9]. This hypothesis is sup-
ported by the fact that the wolves’ social ecology remains highly
dependent on cooperation with conspecifics for activities such
as puppy raising, territory defense, and hunting [10–12], and in
primates at least, cooperation and inequity aversion appear to
be closely linked [3]. Compared to primates, however, pet
dogs show a reduced sensitivity to inequity in that they show
inequity aversion when receiving no reward next to a rewarded
partner [4, 6] but do not respond to differences in reward quali-
ties as some primates do [1]. This reduced sensitivity to inequity
of pet dogs in comparison to non-human primates may be due to
(1) canines showing in general a more primitive form of inequity
aversion, predicting that wolves also do not react to quality ineq-
uity; (2) the domestication process [13], which selected for un-
conditional cooperation with humans and perhaps a change in
intraspecific cooperation, predicting that although wolves react
to quality inequity, dogs do not, even if they have the same expe-
riences as wolves; or (3) socialization effects, which, due to the
close interaction of pet dogs with their human caregivers, lead
to ignoring unfavorable conditions to please the humans, pre-
dicting that dogs raised in packs with a lot less interactions
with humans would, like wolves, react to quality inequity.
We aimed to test these hypotheses using an inequity aversion
task based on earlier studies on pet dogs [4–6] by testing
similarly raised and kept pack-living wolves (n = 9) and dogs
(n = 10) at the Wolf Science Center, to determine whether they
show similar responses to inequity. Two animals were placed
in separate enclosures, each equipped with buzzer apparatuses
placed against the fencing (see Figure 1). The experimenter alter-
nately pointed to the two buzzers and asked the respective ani-
mal to press with a paw to receive a reward. Depending on the
condition, they were given a reward (or not) (see Table 1). In order
to keep the test as similar as possible between conditions, in
asocial conditions, the experimenter acted as if there were a

partner in the enclosure, including giving commands to and
showing the reward briefly to the partner enclosure; however,
after the food rewards were presented, they were discretely
moved back into the bowl. The bowl contained a large amount
of both food rewards, such that over the course of the session,
as foods were removed, the bowl remained sufficiently full.
Rewards—a high-value (HVR) and a low-value (LVR) reward—
were determined prior to testing based on the preferences of
the animals. Each session lasted a maximum of 30 trials per
animal. If an individual stopped performing the task, the experi-
menter gave up to ten commands (‘‘press,’’ or ‘‘come’’ if the
animal had moved away, up to ten times each). If the individual
refused to press or return to the apparatus, the session was
ended. Moreover, as we were interested in whether the inequity
condition experienced by the subject directly affected subse-
quent interactions with its partner and the experimenter, we
implemented an ‘‘interaction test’’ previously used with pet
dogs [4]. After each test condition, both the subject and the part-
ner were moved to a neutral enclosure where they had the choice
to interact with each other and/or the experimenter.
Each session consisted of one condition, and only one session
per day was conducted for each subject. In each condition, we
changed exactly one factor in regard to the baseline equity con-
dition (quality inequity: food quality of the partner’s food; food
control: shown HVR but given LVR [to test for frustration effects];
reward inequity: subject unrewarded; assessment control: pres-
ence of the partner). To further control that a possible response
in the reward inequity condition was due to the mere absence of
food and not the social context, we ran a further condition for
comparison (the no reward condition) in which, like in the ineq-
uity, the subject did not receive food, but this time the partner
was absent (see Table 1 for a summary of the conditions).
Conditions were randomized, except that no-reward conditions
(i.e., RI and NR; see Table 1) were never consecutive, to prevent
animals from losing motivation. Since the dominance relation-
ship is known to affect responses to inequity in primates [14],
we considered both the dominance relationship (dominant-
submissive) and rank distance between partners. See STAR
Methods for more information on all methods.
For both dogs and wolves, there were fewer trials completed
in the reward inequity condition than in the equity condition
(F = 6.698, df = 59, p < 0.001; Figure 2; Data S1), showing
that both wolves and dogs performed fewer trials when they
were unrewarded in the presence of a rewarded partner. Impor-
tantly, wolves and dogs also completed fewer trials in the reward
inequity than in the asocial no reward condition (F = 0.478,
df = 18, p < 0.001; Data S1), indicating that the presence of the
1862 Current Biology 27, 1861–1865, June 19, 2017
Figure 1. Set-Up View from One Enclosure
Fencing between the two enclosures allowed for
each individual to fully see their partner’s behavior.
The same setup was used for the dogs (left) and the
wolves (right). See also Figures S1 and S2.
partner, not the lack of reward, is driving
their refusals in the reward inequity condi-
tion. In other words, dogs and wolves
respond to the inequity in the social
context and not solely to the reward distribution. In contrast to
previous studies on pet dogs, both wolves and dogs also
completed fewer trials in the quality inequity condition than in
the equity condition (F = 1.998, df = 59, p = 0.050; Figure 2;
Data S1). Though there was no significant effect of species
(F = 1.490, df = 14, p = 0.158), it appears that the wolves re-
sponded more strongly than the dogs (see Figure 2). We did
not find any difference between the equity condition and our
food control condition (Data S1).
Furthermore, we investigated how many commands, normal-
ized by number of trials, were given by the experimenter in each
condition as a measure of their resistance. Wolves and dogs
both were given more commands in the reward inequity condi-
tion compared to the equity condition (F = 2.176, df = 64,
p = 0.033; Data S1), but there was no difference between the
reward inequity and no reward conditions (F = 1.675, df = 17,
p = 0.112; Data S1). We found a species interaction for the quality
inequity condition (species 3 QI: F = 2.753, df = 64, p = 0.008;
Data S1). Although wolves were given more commands in the
quality inequity condition compared to the equity condition,
this was not the case for the dogs (wolves, QI versus ET:
F = 3.415, df = 30, p = 0.002; dogs, QI versus ET: F = 0.688,
df = 34, p = 0.496; Data S1). Thus, in order to continue working
in the quality inequity condition (compared to the equity condi-
tion), the experimenter had to prompt the wolves significantly
more times per trial than the dogs, suggesting that the willing-
ness to work in that condition was lower in the wolves than in
the dogs. This finding, paired with the appearance that wolves
responded more strongly to the quality inequity condition,
Table 1. Summary of Conditions
Condition Subject Partner
Social Conditions
Equity test (ET; baseline) LVR LVR
Quality inequity (QI) LVR HVR
Reward inequity (RI) no reward HVR
Food control (FC) HVR shown, HVR shown,
LVR given LVR given
Asocial Conditions
Assessment control (AC) LVR LVRa
No reward control (NR) no reward HVRa
LVR, low-value reward; HVR, high-value reward.
a
In the asocial conditions, the partner’s site was empty; however, the
normal procedure was carried out. The food was moved to the partner’s
side and was then re-placed into the food bowl.

suggests that there may be a difference between dogs and
wolves in how they respond, or how strongly they respond, to
the quality inequity condition. We did not find any difference be-
tween the equity condition and the food control condition for
number of commands per trial (Data S1).
We found an effect of rank on the number of trials completed.
Regardless of species, the more dominant animals were to their
partner (i.e., the larger the rank distance between individuals in
the dyad), the fewer trials they completed in the reward inequity
condition compared to the equity condition and the fewer trials
they completed in the reward inequity condition compared to
the no reward condition (RI versus ET: F = 2.364, df = 59,
p = 0.021; RI versus NR: F = 2.491, df = 15, p = 0.025; Data
S1; Figure 3). This suggests that the more dominant an individual
is to their partner, the more sensitive they are toward the reward
inequity. Additionally, there was an interaction between species
and rank distance for the rate of stress behaviors (e.g., lip licking;
see Table S1) in the social test conditions (species 3 rank dis-
tance: F = 3.101, df = 13, p = 0.008; Data S1). Further analysis
showed that the more subordinate dogs had a tendency for a
higher rate of stress behaviors seen in the test conditions; this
was not the case in wolves (dogs, rank distance: F = 2.340,
df = 6, p = 0.058; wolves, rank distance: F = 0.765, df = 7,
p = 0.469; Data S1). However, it is not clear that the stress
seen in the subordinate dogs is due to working next to a more
dominant partner, because there was no effect of condition be-
tween the assessment control condition and the equity condition
(ET versus AC: F = 0.841, df = 60, p = 0.403; Data S1).
The test conditions also affected the subject’s subsequent in-
teractions with both the experimenter and the partner. Dogs took
longer to go to the experimenter (F = 2.184, df = 27.83, p = 0.038;
Data S2) and spent less time with the experimenter (F = 2.225,
df = 35.33, p = 0.033; Data S2) in the quality inequity condition
compared to the equity condition. Independent of condition,
there was an effect of rank distance, where the more subordinate
a dog was to their partner, the lower their latency to go to the
experimenter (F = 4.646, df = 6.080, p = 0.003; Data S2).
For wolves, we found that subjects went to the experimenter
faster in the quality inequity compared to the equity condition
(F = 2.261, df = 32, p = 0.031; Data S2), but there was no differ-
ence in time spent with the experimenter between the test con-
ditions (F = 1.398, df = 32, p = 0.172; Data S2).
In regards to the latency to be within 1 m of their partner,
there was an interaction between rank distance and condition
Figure 2. Number of Trials Completed by
Condition
Data are represented as mean ± SE.
(F = 2.516, df = 41.25, p = 0.016; Data
S2). Rank distance affected the latency
to be within 1 m of the partner in the
reward inequity condition (F = 6.943,
df = 1.15, p = 0.019), but not in the
equity condition (F = 0.156, df = 1.13,
p = 0.699): the higher ranking the subject
was, the faster they went to their partner
after the test in the reward inequity condi-
tion. Finally, both dogs and wolves spent more time within 1 m of
their partner after the equity condition compared to the reward
inequity condition (F = 2.834, df = 53.08, p = 0.006; Data S2).
DISCUSSION
The inequity aversion response shown here in wolves and dogs
is more similar to that of non-human primates than previously
thought, as dogs and wolves responded to both reward inequity
and quality inequity. Moreover, the more dominant individuals (of
both wolves and dogs) responded similarly to the more dominant
non-human primates [14] by refusing to participate earlier when
receiving nothing in the presence of a rewarded, lower-ranking
partner [15]. We found no dominance effect in the control condi-
tion; thus, it is the relationship between subject and partner, not
merely a lack of reward, that drives this response in these
subjects.
These results support hypotheses that these cognitive skills in
dogs are not an effect of domestication, but rather were main-
tained from their ancestors [9]. In fact, the results here and in pre-
vious studies on pet dogs suggest that rather than increasing
dogs’ response to unequal treatment, their relationship to hu-
mans may result in a higher tolerance for unequal treatment, at
least from humans. Life-long positive interactions and training
with their human caregivers might prevent dogs from refusing
to continue to participate in the experiment due to their willing-
ness to please the human experimenter. The dogs in the present
study, though highly socialized with humans in their first weeks of
life, do not have a pet-owner relationship with humans and so
may not have the same eagerness to please humans as a pet
dog would. Nevertheless, they were still more eager to please
the human experimenter than were the wolves, as the wolves
needed more prompts than dogs to comply in the quality inequity
condition compared to the equity condition.
If inequity aversion does function to maintain cooperative in-
teractions, we would expect that being treated unequally would
have consequences for subsequent interactions. And indeed,
we found that, like pet dogs, both wolves and pack-living dogs
spent less time with their partner after the reward inequity condi-
tion than after the equity condition. Moreover, rank also influ-
enced this behavior, since, if the dominant animal of a dyad
was tested beforehand, the latency to approach each other
was shorter in the reward inequity than the equity condition.
This again supports our results that dominant animals seem to
Current Biology 27, 1861–1865, June 19, 2017 1863

Figure 3. Number of Trials Completed by Rank Distance between
Partners for Reward Inequity, No Reward, and Equity Conditions
react stronger to being treated worse than their subordinate
partners. This might be partly due to the fact that usually it is
the subordinate individual that is at a disadvantage, e.g., has
later access to food [16, 17], leading to a violation of expectancy
in the dominant partner if these roles are reversed. Thus, not only
does unequal treatment appear to have consequences for
subsequent interactions between pack-living dogs and wolves,
but these consequences are possibly augmented further when
the unequal treatment is directed toward the more dominant of
the two individuals. This suggests that inequity between individ-
uals that are part of the same social hierarchy may vary, with
unequal treatment representing something quite different de-
pending on the relationship between those involved and equal
division of reward not necessarily being as simple as being
treated the same (e.g., receiving the same food reward) as
another individual.
Interestingly, prior experience of inequity influenced the ani-
mals’ subsequent behaviors not only toward each other, but
also toward the experimenter. Whereas the wolves had a lower
latency to go to the experimenter after the quality inequity condi-
tion than in the equity condition, the dogs had a higher latency to
go to the experimenter and also spent less time with her. We
have no good explanation for why wolves and dogs reacted in
opposite ways toward the experimenter, but pet dogs behaved
similarly to our pack dogs [4]. The lack of a differentiated
response to the experimenter after the reward inequity condition
in the present study may be due to the fact that the subjects
continued to work for longer in the quality inequity condition, re-
sulting in a more sustained unequitable situation in the quality
inequity condition. This may have elicited a stronger response
toward the experimenter. In general, subordinate dogs had a
lower latency to go to the experimenter, which might be tied to
subordinate dogs being more stressed during the buzzer test
and thus looking for social support from the experimenter
(i.e., humans as an attachment figure [18–20]).
In summary, the results presented here suggest that canines
behave similarly to non-human primates in regard to unequal
outcomes and that this sensitivity was already present in their
common ancestor. Furthermore, the study once again highlights
that it is important to test different populations of dogs and
1864 Current Biology 27, 1861–1865, June 19, 2017
possibly wolves to get a better insight into the effects of domes-
tication. These results are in line with the hypothesis that inequity
aversion evolved alongside cooperation [3], allowing individuals
to recognize disadvantageous pairings in cooperative situations.
Since wolves, and thus most likely the common ancestor of
wolves and dogs, are a highly cooperative species in terms of
hunting, breeding, and territory defense, this probably led early
wolves to develop and maintain responses to inequity in order
to regulate cooperation. However, the fact that wolves and
pack-living dogs respond to the quality inequity condition similar
to primates supports also the possibility of an old evolutionary
origin, rather than convergent evolution.
STAR+METHODS
Detailed methods are provided in the online version of this paper
and include the following:
d KEY RESOURCES TABLE
d CONTACT FOR RESOURCE SHARING
d SUBJECT DETAILS
d METHOD DETAILS
B Pre-Testing
B Training
B Experimental Setup
B Test Procedure
d QUANTIFICATION AND STATISTICAL ANALYSIS
B Behavioral Coding
B Social Relationships
B Statistical Analyses
d DATA AVAILABILITY
SUPPLEMENTAL INFORMATION
Supplemental Information includes two figures, two tables, and three data files
and can be found with this article online at http://dx.doi.org/10.1016/j.cub.
2017.05.061.
AUTHOR CONTRIBUTIONS
Conceptualization, J.L.E. and F.R.; Methodology, J.L.E., S.M., and F.R.; Inves-
tigation, J.L.E.; Formal Analysis, J.L.E.; Writing – Original Draft, J.L.E., S.M.,
and F.R.; Writing – Review & Editing, J.L.E., S.M., and F.R.; Funding Acquisi-
tion, F.R.
ACKNOWLEDGMENTS
The Wolf Science Center was established by Zsófia Virányi, Kurt Kotrschal,
and Friederike Range, and we thank all of our helpers who made this
possible and thus indirectly supported this research. We thank the many stu-
dents at the lab that helped with this study, in particular Julie Carpenter and
Judith Zembrot. We additionally would like to give huge thanks to Marleen
Hentrup and János Tardi for building the apparatuses, Jeremy Cain for
graphical support, Rooobert Bayer for photos, Marianne Heberlein and
Stephan Reber for statistical support, Jennifer Bentlage for administrative
support, and four anonymous reviewers for their comments. This study
was supported by funding from the European Research Council (ERC) under
the European Union’s Seventh Framework Programme (FP/2007-2013)/ERC
grant agreement no. 311870. The authors further thank many private spon-
sors including Royal Canin for financial support and the Game Park Ernst-
brunn for hosting the Wolf Science Center. The funders had no role in study
design, data collection and analysis, decision to publish, or preparation of
the manuscript.
Received: March 6, 2017
Revised: March 29, 2017
Accepted: May 18, 2017
Published: June 8, 2017
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Current Biology 27, 1861–1865, June 19, 2017 1865
STAR+METHODS

KEY RESOURCES TABLE

REAGENT or RESOURCE SOURCE IDENTIFIER

Experimental Models: Organisms/Strains
Gray Wolf, Canis lupus Wolf Science Center http://wolfscience.at/en/
Domestic dog, Canis lupus familiaris Wolf Science Center http://wolfscience.at/en/
Software and Algorithms
R: A Language and Environment for [21] https://www.r-project.org/
Statistical Computing
‘R’ Package, ‘‘MASS’’ [22] http://www.stats.ox.ac.uk/pub/MASS4
‘R’ Package, ‘‘lme4’’ [23] https://cran.r-project.org/web/packages/lme4/lme4.pdf
‘R’ Package, ‘‘ggplot2’’ [24] http://ggplot2.org
‘R’ Package, ‘‘car’’ [25] http://socserv.socsci.mcmaster.ca/jfox/Books/Companion
‘R’ Package, ‘‘lmerTest’’ [26] http://cran.r-project.org/package=lmerTest
Observer XT 10.5 Noldus Information http://www.noldus.com/knowledge-base/observer-xt-10
Technologies
Other
Fuß- Grobhandtaster (‘Buzzer’) Conrad Electronic Eaton FAK-S/KC11/I IP67
GmbH & Co KG [AT]

CONTACT FOR RESOURCE SHARING

Further information and requests for resources should be directed to and will be fulfilled by the Lead Contact, Jennifer L. Essler
(jennifer.essler@vetmeduni.ac.at).

SUBJECT DETAILS

Ten dogs and nine timber wolves were tested at the Wolf Science Center (WSC) in Ernstbrunn, Austria. All animals except those dogs
in 2014 were brought to the center at ten days of age and hand-raised by professional experimenters, spending all of their time in the
presence of a human from the age of ten days to four months. After four months, individuals were introduced into previously estab-
lished packs of adult wolves or dogs, respectively. Dogs born in 2014 were born at the center, and spent at least four hours per day
with a trainer and other puppies (without the mother) before being left in their pack full-time at the age of four months (see [27] for
further information on hand-raising methods). Dogs and wolves are kept in a similar way and have the same life experiences in order
to compare the two species without major differences in life experience. This study was discussed and approved by the institutional
ethics and animal welfare committee at the University of Veterinary Medicine Vienna in accordance with Good Scientific Practice
guidelines and national legislation (Unser Zeichen #10/09/97/2013).

Subject Partner Subject Sex Partner Sex Pack Birth Year LVR HVR
Wolves
Kaspar Aragorn M M 1 2008 Kibble Meat
Aragorn Kaspar M M 1 2008 Kibble Meat
Chitto Shima M F 1 2012 Kibble Meat
Shima Chitto F M 1 2008 Kibble Meat
Amarok Kenai M M 2 2012 Kibble Meat
Kenai Amarok M M 2 2011 Kibble Meat
Nanuk Una M F 3 2009 Kibble Meat
Geronimo Yukon M F 4 2009 Kibble Meat
Wamblee Yukon M F 4 2012 Kibble Meat
(Continued on next page)

e1 Current Biology 27, 1861–1865.e1–e3, June 19, 2017

Continued
Subject Partner Subject Sex Partner Sex Pack Birth Year LVR HVR
Dogs
Bansai Bora M F 4 2014 Kibble Sausage
Bora Bansai F M 4 2011 Kibble Sausage
Sahibu Gombo M M 5 2014 Kibble Sausage
Nia Sahibu F M 5 2011 Kibble Sausage
Zuri Layla F F 6 2011 Kibble Sausage
Layla Zuri F F 6 2011 Kibble Sausage
Nuru Layla M F 6 2011 Kibble Sausage
Maisha Binti M F 7 2009 Kibble Meat
Binti Maisha F M 7 2010 Kibble Meat
Meru Hiari M M 8 2009 Kibble Meat

METHOD DETAILS

Pre-Testing
We carried out a food preference test to establish a low- and high-value (LVR and HVR, respectively) reward to be used during testing.
A reward was considered to be of high value if they chose the HVR over the LVR at least 85% of the time, in two consecutive sessions
on different days. To be used in the test, the LVR had to be consumed by the animals at least ten times in a row.

Training
Prior to the test, all animals were trained to place their paw on a buzzer placed on a wooden apparatus (Figure S1) to obtain a reward
in response to an experimenter pointing at the apparatus/buzzer and giving the command ‘press.’ The buzzer was then moved to the
full-apparatus setup, and animals were considered ready for testing when they pressed the buzzer with their paw, in response to an
experimenter pointing and giving the command ‘press,’ ten times in a row. Reward used in the training were not used in the testing.

Experimental Setup
Individuals were placed in two familiar adjacent enclosures (7.3 m2 each). These enclosures were made with large wire mesh. On the
outside of each enclosure, one larger hole (18 cm x 18 cm) was cut into the fence for the buzzer (Eaton FAK-S/KC11/I) to be moved in
and out. Two buzzers were positioned each on a wooden table, so that the action of pressing the buzzer required the animals to lift a
paw upward in order to complete the task. Behind each table, but further obscured from the vision of the animals by opaque panels,
was one helper person per table who inserted and removed the buzzer on each trial. Between the two tables, and fully visible to the
animals, was the experimenter. Food reward for the test were placed in a bowl in front of the experimenter, where half the bowl held
the HVR and the other half held the LVR. Both food reward were present and visible in every condition, and the HVR was always closer
to the subjects than the LVR (see Figure S2). Outside of each enclosure was a camera to film every condition for later coding.
Each subject participated in six conditions across separate sessions on separate days (see Table 1). To test whether subjects
respond to receiving a different reward from their partner, we compared the equity baseline condition (ET), where both individuals
received the LVR, to the quality inequity (QI) condition (subject: LVR, partner: HVR) and the reward inequity (RI) condition (subject:
nothing, partner: HVR). To ensure that any differences in the previous tests were not due to a frustration effect of simply seeing
but not receiving the HVR, we further compared the ET baseline to a food control (FC) test, where both individuals were shown
the HVR but then given the LVR for completing the task. Further, to determine if any differences found between the ET and RI con-
dition were indeed due to not receiving a reward in the presence of a rewarded partner, and not simply due to not receiving any
reward (regardless of the partner), we compared the RI condition to a no reward (NR) solo condition, where the subject was tested
alone and received no reward, while the HVR was shown to the partner’s empty enclosure (controlling for the movement of the food).
In practice, the NR condition was the same as the RI condition, except that there was no animal in the partner’s enclosure.

Test Procedure
For each session for the buzzer task, individuals were placed in the two adjacent enclosures, with the sides randomly assigned. The
experimenter kneeled in-between the tables, with the bowl of food reward situated in front of her. Two helpers, one behind each table,
were responsible for moving the buzzers in and out of the enclosures for each trial. To start a session, the partner’s buzzer was moved
into his/her enclosure, while simultaneously the experimenter pointed to the buzzer and said ‘‘press.’’ Once the individual pressed the
buzzer, the experimenter picked up the food reward for that condition and held it between the two enclosures so both animals had the
opportunity to see the reward. The experimenter then gave the food to the partner animal. This was then repeated for the subject
animal, then continued back and forth until the session was over, with either 60 trials being reached (30 per animal) or one animal
refusing to continue.

Current Biology 27, 1861–1865.e1–e3, June 19, 2017 e2

 Animals could refuse to continue, thus ending the session, in different ways. First, if an animal did not press the buzzer, the exper-
imenter repeated the command ‘‘press’’ five times, followed by the name of the subject once, then again ‘‘press’’ five times. If the
animal did not press the buzzer after this, the session was ended. Second, if an animal left the immediate vicinity of the buzzer table
(1 m), the experimenter called the animal’s name five times, then called ‘‘come’’ once, then the animal’s name five times again. If the
animal did not return to the buzzer table after this, the session was ended. Finally, the session was ended if the subject destroyed the
apparatus by removing the buzzer from the table, as this was considered a frustration level at least as high as refusing to participate
further.
After each condition, a social interaction test (hereafter, Interaction Test) was conducted, where the experimenter walked around to
the opposite side of the larger, adjacent test enclosure and sat down on the outside of the fence. Both subject and partner (where
applicable) were then let into this enclosure and were free to interact with each other, or the experimenter, for five minutes. Each
animal was followed with a camera from outside the enclosure for later coding.

QUANTIFICATION AND STATISTICAL ANALYSIS

Behavioral Coding
Each condition, both during the Buzzer Test and afterward in the Interaction Test, was recoded with video cameras (Sony HDR-
CX220E Camcorder) and later coded in Observer XT 10.5 (Noldus Information Technologies). Behaviors coded included buzzer
presses, number of commands given and stress behaviors in the Buzzer Test, and time spent near partner/experimenter in the Inter-
action Test (see Table S1). 20% of the videos were coded by an independent outside individual (all behaviors above k = 0.85).

Social Relationships
Behavioral observations from the WSC were used to assess the social relationships for the analyses. These daily observations are
done by long-term students at the WSC who have previously shown interrater-reliability with established observers at the lab.
Observations are done when the animals are in their home enclosures with their full pack, where no individual is missing. Each obser-
vation is a ten minute focal on one individual where their social behaviors with their pack members are recorded by instantaneous
sampling (see Table S2). Data from these focals were used to calculate a David Score [28] for each individual and then subtracted
individual A’s score from individual B’s in order to obtain a value of rank distance for each pair.

Statistical Analyses
For all models in this study, we used a p value reduction method where we removed the least-significant variable from each model,
fully reducing it until only significant factors were left. Variables are listed in both the order they were removed (first to last) and with the
values they had in the model when they were removed. ‘‘FINAL’’ under ‘Order Removed’ denotes that the following variables were
present in the final model for that particular model. In all models, ‘‘:’’ between two variables denotes an interaction in the model. See
Data S1, Data S2.
Generalized linear mixed models (GLMMs) were run for the Buzzer Task using the total number of trials completed as the response
variable, and condition, rank distance, and species as predictor variables, with species by condition and by rank distance, run as
possible interactions. Additionally, GLMMs were run for both the number of commands (controlling for test duration) and the rate
of stress behaviors (together, whine, yawn, and lips licking). For the Interaction Test behaviors, we ran Linear Mixed Models
(LMMs) for time spent with experimenter/latency to experimenter and time spent with partner/latency to partner (where applicable)
as response variables and with the same predictor variables as in the Buzzer Task models. Analyses were run in R version 3.2.2 [21]
with the packages MASS [22], lme4 [23], ggplot2 [24], car [25], lmerTest [26].

DATA AVAILABILITY

See Data S3 for a description.

e3 Current Biology 27, 1861–1865.e1–e3, June 19, 2017