Ethical statement
This study was approved by the Ethics and Animal Welfare Committee of the University of Veterinary Medicine Vienna (ETK-023/02/2021). Written informed consent was obtained from all dog owners for their participation in the study and from the individuals in Supplementary Videos 1 and 2 for the publication of photographs and videos containing their images.
Subjects
44 pet dogs (22 males and 22 females) aged between 1 and 12 years old (M = 5.95, SD = 3.45) participated in the study (Table S1). They were recruited from a database of owners who volunteered to participate in behavioural studies at the Clever Dog Lab. Four dogs who were not motivated to participate in the study were excluded, resulting in a final sample of 40 dogs (20 males and 20 females, M = 5.95, SD = 3.40). No breeds were excluded.
Experimental design
To test the effect of age, we classified dogs into three age groups: 1-3 years as ‘young’ (early adulthood), 4-7 years as ‘adult’ (middle age and late adulthood), and 8-12 years as ‘senior’ (senior and geriatric). These age groups were based on previous papers on ageing in dogs: Wallis et al. (2020) categorised ‘early adulthood’ as >1-3 years and ‘middle age’ as >3-6 years; Chapagain et al. (2017) categorised ‘late adulthood’ as >6-8 years, ‘senior’ as >8-10 years, and ‘geriatric’ as ≥10 years. Our sample was fairly balanced in each age group, with 11 dogs in ‘young’, 15 dogs in ‘adult’, and 14 dogs in ‘senior’.
There were three conditions to test for reputation formation:
- Eavesdropping (n = 20): the subject observed two humans (henceforth ‘partners’) interact with a dog (henceforth ‘dog demonstrator’) to test for indirect reputation formation (i.e., eavesdropping). This condition consisted of two sessions, each with a single trial. Conducting more than one trial per session was avoided to prevent immediate influence from direct experience. However, we argue that the single trial in Session 2 was still based on observation rather than the brief interaction in Session 1, which took place on a previous day.
- Control (n = 20): the subject observed the humans perform the same actions as in the eavesdropping condition but without a dog demonstrator being present. This was conducted to determine whether the dogs' responses were due to the interactions between the humans and the dog demonstrator, or if the humans' actions alone were sufficient to allow a discrimination between them, should eavesdropping be observed in the eavesdropping condition. As with the eavesdropping condition, this condition also consisted of two sessions, each with a single trial.
- Direct experience (n = 40): to test for direct reputation formation, the humans directly interacted with the subjects. This condition consisted of a single session with 12 trials.
Dogs were assigned to the eavesdropping condition or the control condition and were matched regarding age and sex (Table S1). All dogs participated in the direct experience condition and were tested in a counterbalanced order (i.e., half of the dogs experienced the eavesdropping/control condition first and the other half experienced the direct experience condition first).
In total, 12 women, all unfamiliar to the dogs, volunteered to act as human partners in the study. Pairs of partners remained stable within each condition, but each dog encountered two different pairs of partners (e.g., partners A and B in the first condition and partners C and D in the second condition). During the experiment, one partner was dressed in white and the other in black. The roles and clothing colours were randomised and counterbalanced between subjects but remained fixed within subjects to avoid confusion and help the dogs to better distinguish between the individuals playing the different roles. For example, one subject might have had the generous partner in black and the selfish partner in white in both conditions, though different individuals acted as the partners in each condition. Each partner wore a hip bag containing 18 small pieces of sausage, but only the generous partner used them to feed the dog.
There were two dog demonstrators who interacted with the partners in the observation phase for the eavesdropping group: a neutered male dog (Jasper, 11 years, Labradoodle) served as the demonstrator for female subjects, and a spayed female dog (Emmi, 4 years, Labrador) served as the demonstrator for male subjects. Dogs of the opposite sex were chosen as demonstrators to reduce dominance behaviour and to ensure that subjects would pay more attention to the demonstrator.
Experimental setup
The experiment was conducted in the outdoor test enclosure of the Clever Dog Lab based at the University of Veterinary Medicine Vienna campus. A chair was placed in the centre of the enclosure, where the dog owner sat during the experiment, and there was a water bowl for the dog next to the chair. The partners stood behind a divider (175 × 180 cm) outside of the enclosure so that the subject could not see them and be distracted in the observation phase. A crate was placed outside the enclosure behind the divider, and the dog demonstrator was placed inside it after the observation phase to avoid disturbing the subject during the test phase.
We used spray paint on the grass to mark where individuals would stand during the experiment. A cross was marked 3 m away from the owner’s seat and two crosses were marked 2 m perpendicular from the central cross. A circle with a radius of 50 cm was marked around the left (P1) and right (P2) crosses (Fig. 1). A camera (GoPro Hero 4 Black) was placed on a tripod in the enclosure and filmed the whole experiment.
Procedure
The study was conducted from May to August 2021 and generally followed the procedure outlined in Jim et al. (2022) (see Fig. 2 for an overview).
Eavesdropping/control condition
This condition comprised two sessions and there was a break of 3-38 days between sessions. The first session began with a habituation phase, where the owner and main experimenter took the subject and the dog demonstrator on a short walk around the university campus (5-10 minutes) to ensure that they tolerated each other and could then focus during the experiment (if the dogs had been uncomfortable with each other, the experiment would have been terminated, but this never happened). The subject and the owner then entered the enclosure, and the subject could explore it freely for five minutes while the main experimenter explained the procedure to the owner.
Before the main procedure, a baseline was conducted to determine whether dogs, as a group, exhibited a preference for one partner over the other before observing any third-party interactions. The owner sat with their dog between their legs, holding the dog by the collar, and remained blindfolded throughout the experiment to prevent them from influencing the dog’s behaviour. The partners entered the enclosure and stood on P1 and P2 (their positions were randomised and counterbalanced across subjects) without making eye contact with the dog. Each person held a piece of sausage in their hand, keeping their arms relaxed by their sides so it was not obvious to the dog that they were holding food. When the main experimenter said “ok”, she started the timer for one minute and the owner let go of the dog’s collar. The owner could give a short prompt if the dog did not move by themselves, such as a gentle nudge or saying “ok” to indicate to the dog that they were free to move, but was instructed not to gesture in a specific direction. If the subject walked towards a partner and at least one front paw was within or on the circle marked on the floor whilst looking at her, this was considered as a choice and the partner fed the dog. After the subject ate the food, the other partner who was not chosen called the subject to get their attention and fed the dog too to ensure they did not develop a preference for one partner. If the subject did not approach either partner after one minute, it was considered a ‘no choice’ (this never happened). Then, the main experimenter asked the owner to call the dog back. The owner remained seated, holding their dog by the collar, and the partners left the enclosure. The baseline ended after this single trial (Supplementary Video 1).
The main procedure comprised two phases:
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Observation phase
In the eavesdropping condition, the main experimenter entered the enclosure with the dog demonstrator on a short lead and stood on P1 or P2 (randomised and counterbalanced across subjects). Then, one partner entered the enclosure and walked to the opposite standing point, holding a piece of sausage in her hand. The main experimenter walked alongside the dog demonstrator toward the partner but then stayed behind, allowing the dog demonstrator to approach the partner alone. When the dog demonstrator reached the partner, she raised her arm to show the food. The subject witnessed one of the following scenarios, depending on which partner the dog demonstrator interacted with:
- Generous: The partner said, “here you go!” or “du kannst es haben!” in a friendly tone and fed the dog demonstrator (the partners spoke in their preferred language to ensure their behaviour was natural). After the dog demonstrator had eaten the food, the main experimenter walked the dog demonstrator back to the starting point, while the partner left the enclosure.
- Selfish: The partner said, “you can’t have it!” or “du kriegst es nicht!” in an unfriendly tone, crossed her arms, and turned away, keeping the food in her hand. After the interaction ended a few seconds later, the main experimenter walked the dog demonstrator back to the starting point, and the partner left the enclosure.
This phase was identical in the control condition, with one difference: the main experimenter and dog demonstrator were absent. Thus, the subject observed the partners ‘interact’ with an invisible dog in the same way as in the scenarios described above (Supplementary Video 1).
The procedure was repeated three times per partner (e.g., six interactions with the main experimenter and dog demonstrator standing on P1 and the partners standing on P2). Then, the main experimenter and dog demonstrator swapped sides and the whole procedure was repeated (e.g., six interactions with the main experimenter and dog demonstrator now standing on P2 and the partners standing on P1). After these 12 interactions, the main experimenter and the dog demonstrator left the enclosure, and the dog demonstrator was placed in the crate, after which the test phase immediately followed.
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Test phase
This phase was identical in both the eavesdropping and control conditions. The partners entered the enclosure and stood on P1 and P2 (their positions were randomised and counterbalanced across subjects) without holding any food, standing still, and looking ahead without making eye contact with the dog. When the main experimenter said “ok”, she started the timer for 15 seconds, and the owner let go of the dog’s collar. As in the baseline, the owner could give a short prompt if the dog did not move by themselves. During this time, the dog could act freely while the partners remained still and did not react to the dog. At the end of this single trial, the main experimenter said “stop” to indicate to the owner to call the dog back. The owner remained seated, holding their dog by the collar, and the partners left the enclosure, which concluded the session.
Session 2 (conducted 3-38 days later) was identical to Session 1, with some minor changes. There was no habituation phase or baseline, so it started directly with the observation phase. The order of the partners and the side they stood on first was counterbalanced across sessions (e.g., if the selfish partner started and stood on P1 in Session 1, then the generous partner started and stood on P2 in Session 2). The positions of the generous and selfish partner were also counterbalanced across sessions in the test phase. Again, there was only one 15-second trial (identical to Session 1), and this single trial concluded the session.
Direct experience condition
Prior to testing, the subject was allowed to explore the enclosure freely for approximately five minutes while the main experimenter explained the procedure to the owner. This condition comprised a single session and no baseline was conducted. Hence, the procedure comprised two phases:
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Experience phase
This was identical to the observation phase in the eavesdropping/control condition, except that the owner replaced the main experimenter and the subject replaced the dog demonstrator. Thus, the owner entered the enclosure with the subject on a short lead and stood on P1 or P2 (randomised and counterbalanced across subjects). One partner then entered the enclosure, walked to the opposite standing point, and interacted with the subject in the same way they would with the dog demonstrator in the eavesdropping/control condition. This procedure was repeated three times per partner, resulting in six interactions with the owner and subject standing at P1 and the partners at P2. The owner and subject then swapped sides, and the procedure was repeated (i.e., six interactions with the owner and subject now standing at P2 and the partners at P1). After these 12 interactions, the test phase immediately followed.
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Test phase
Before the first trial, the owner exchanged the short lead for a 10-meter training lead and sat with their dog between their legs, holding the dog by the collar. The long lead allowed the dog to move freely around the enclosure during the trial but facilitated recall at the end of each trial.
Both partners entered the enclosure, with one standing on P1 and the other on P2, each holding a piece of sausage in their hand, with their arms relaxed by their sides. When the main experimenter said “ok”, indicating the owner to release their dog, she started the timer for 15 seconds. Simultaneously, the partners raised their hands to show the food to the dog, standing still and looking ahead without making eye contact with the dog. As in the test phase of the eavesdropping/control condition, the owner could give a short prompt if the dog did not move by themselves.
If the subject walked towards a partner and their front paws were within or on the circle marked on the floor, this was considered a choice. The partner then acted the same way as during the observation phase (i.e., the generous partner fed the subject, while the selfish partner did not). At the same time, the other partner moved her hands to her chest to prevent the dog from taking the food, and both partners ignored the dog for the remainder of the trial. If the dog did not approach either partner during the 15 seconds, it was considered a ‘no choice’.
At the end of the trial, the experimenter said “stop” to indicate to the owner to call the dog back and hold them by the collar. If the generous partner was chosen, she rebaited herself by taking another piece of sausage from her hip bag, and the selfish partner performed the same action simultaneously to control for stimulus enhancement. The partners’ positions were semi-randomised to ensure they did not stay in the same position more than twice consecutively. The main experimenter used hand signals to silently instruct the partners to either remain in their positions or swap places during each trial; this was done to minimise the likelihood of the blindfolded owner becoming aware of the partners’ positions, as having such knowledge could allow them to influence the dog’s behaviour. After these 12 trials, this concluded the session (Supplementary Video 2).
As stated previously, dogs were tested in the eavesdropping/control condition and direct experience condition in a counterbalanced order. If dogs completed the direct experience condition first, there was a break of 2-26 days between this condition and Session 1 of the eavesdropping/control condition. However, if dogs completed the eavesdropping/control condition first, we allowed them to participate in the direct experience condition on the same day after a break of 15-30 minutes due to unavoidable logistical constraints. This resulted in a break of 0-27 days between Session 2 of the eavesdropping/control condition and the direct experience condition. Importantly, the dogs interacted with two different sets of partners in the two conditions they experienced.
Behavioural analysis
The videos were coded on Loopy (http://loopb.io, Loopbio Gmbh, Vienna, Austria). We coded the dog’s choice in the baseline and test trials. The trial started when the experimenter said “ok” and stopped when the experimenter said “stop” after 15 seconds. We live coded the dog’s choice as a binomial variable (generous or selfish). If the dog did not make a choice within the trial, then it was a ‘no choice’ response and coded as NA and removed from the analysis. Additionally, the partner must have reacted to the subject for it to be coded as a choice in the direct experience condition (i.e., the generous partner must have fed the dog, and the selfish partner must have turned away). Interobserver reliability was not analysed for choice because it was clear which partner the dog approached first and we could review the video footage if necessary.
For the test trials, we also coded how much time the subject spent in proximity to each partner within the 15-second trial (see Table 1 for definitions). 20% of the videos were randomly selected for interobserver reliability, which were coded by KB and EMB and analysed with R (v4.4.0; R Core Team 2024) using the Intraclass Correlation Coefficient from the R package “irr” (v0.84.1; Gamer et al. 2019). Inter-rater agreement was excellent (ICC (two-way, agreement) = 0.993, F = 308, p < .001). KB and EBM then coded half of the remaining videos each.
Table 1 Definitions of coded behaviours
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Behaviour
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Definition
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Choice (generous / selfish / NA)
(Event)
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At least one of the subject’s front paws was within or on the circle marked on the floor whilst looking at the partner.
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Proximity to partner (generous and selfish were coded separately)
(Duration in seconds)
|
- At least one of the subject’s paws was within or on the circle marked on the floor.
- The subject moved their paw outside of the circle but stayed stationary near the partner until their whole body moved away.
- The subject stepped out of the circle but continued to look at the partner until they looked away.
- The subject was jumping on the partner and their paws left the circle.
- If the dog walked past the partner and entered the circle but did not stop or look at her, this was not coded.
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Statistical analyses
All analyses were performed using R in RStudio (v2024.04.0+735; Posit team 2024) and graphs were created using the R packages “ggplot2” (v3.5.1; Wickham 2016) and “ggeffects” (v1.7.0; Lüdecke 2018).
- Baseline
First, we analysed whether dogs, as a group, preferred one partner over the other before observing any third-party interactions. To do this, we conducted an exact binomial test to compare the proportion of dogs that chose one partner with the proportion expected by chance (0.5). However, even if the whole sample of dogs did not have a significant preference for one partner over the other, it is possible that a subsample in the different age groups or conditions (eavesdropping or control only) could have had a bias. Therefore, we conducted a General Linear Model (GLM) with a binomial error distribution using the function ‘glm’ to test whether age and condition (predictors) influenced dogs’ preference for one partner over the other. In these analyses, the individual who had been assigned the generous role for the main procedure was coded as a ‘success’ in the response variable. This model consisted of the single trial in the baseline from the 40 dogs.
- Test phase
2.1. Eavesdropping vs control condition
2.1.1. Partner choice. To examine whether the proportions of choices for the generous partner depended on age group (young, adult, senior) and/or condition (eavesdropping vs control), we conducted a Generalized Linear Mixed-Effects Model (GLMM) with a binomial error structure and logit link function (McCullagh and Nelder 1989), using the function ‘glmer’ from the R package “lme4” (v1.1.33; Bates et al. 2015). Age group and condition, along with their two-way interaction, were included as predictors. Trial number (1 or 2) and condition order (whether dogs experienced the control/eavesdropping condition before or after the direct experience condition) were added as control variables; trial number was treated as a factor with two levels, while condition order was treated as a continuous fixed effect. To account for repeated measures, we included subject ID as a random effect. After discarding six trials where dogs did not make any choice, we analysed data from 74 trials involving 39 dogs (one dog, Mozart, did not choose a partner in either of the two trials across both conditions).
2.1.2. Time spent in proximity to the generous partner. We also tested whether dogs spent more time with the generous partner using a GLMM with a beta distribution, implemented with the R package “glmmTMB” (v1.1.7; Brooks et al. 2017). The predictors, their two-way interaction, control variables, and random effect were identical to those used in the binomial GLMM described above. The response variable was the proportion of time spent in proximity to the generous partner during each trial within each condition, calculated as the time in proximity to generous partner / (time in proximity to generous + selfish partner). After discarding eight trials where dogs did not spend any time in proximity to either partner, we analysed data from 72 trials involving 39 dogs (one dog, Mozart, did not approach a partner in either of the two trials across both conditions) to calculate proportions by trial. Lastly, as beta GLMMs cannot handle 0s and 1s, we transformed the response variable following the formula suggested by Smithson and Verkuilen (2006).
2.2. Direct experience condition
2.2.1. Partner choice. We analysed whether dogs preferred the generous partner after the experience phase at both an individual level and group level, using the proportion of choices for the generous partner as the response variable. This was calculated as the number of choices for the generous partner / (number of choices for the generous + selfish partner). At the individual level, we conducted exact binomial tests to compare this proportion against the value expected by chance (0.5). At the group level, we conducted a GLMM with a binomial error structure and logit link function to examine whether the proportion of choices for the generous partner varied by age group (young, adult, senior) and/or trial number (1-12, as more trials equated to more experience and opportunities to learn). Age group, trial number, and their two-way interaction were included as predictors, with condition order included as a control variable and treated as a continuous fixed effect. To account for repeated measures, we included subject ID as a random effect. After discarding 38 trials where dogs did not make any choice, we analysed data from 442 trials involving the 40 dogs.
2.2.2. Time spent in proximity to the generous partner. We tested whether dogs spent more time with the generous partner using a GLMM with a beta distribution. The predictors, their two-way interaction, control variable and random effect were identical to those used in the binomial GLMM described above. We calculated the proportion of time spent in proximity to the generous partner in each trial as the response variable. After discarding 100 trials where dogs did not spend any time in proximity to either partner, we analysed data from 380 trials involving the 40 dogs to calculate proportions by trial. Lastly, as beta GLMMs cannot handle 0s and 1s, we transformed the response variable following the formula suggested by Smithson and Verkuilen (2006).
We followed the same procedure to fit the GLMMs: we z-transformed condition order (a continuous fixed effect) to a mean of 0 and standard deviation of 1. To avoid inflated type I error caused by multiple testing (Forstmeier and Schielzeth 2011), we used a full-null model approach, comparing the significance of full models by means of a likelihood ratio test (Dobson and Barnett 2018) using the R function ‘anova’ with the argument test set to “Chisq”, with null models lacking the predictors but otherwise identical to their respective full model. p values for individual effects were based on likelihood ratio tests comparing the full model with reduced models lacking each term. If the model included an interaction that was not significant, we removed the interaction from the model and fitted a reduced model including only the single terms to ease interpretation of the single term’s estimations. We calculated profile likelihood confidence intervals based on the log-likelihood function.
We evaluated the quality of the models following a series of steps. We checked for model stability for the baseline GLM by means of dfbeta-values, and in the case of the GLMMs by excluding subjects one at a time from the data and comparing model estimates derived for these subsets with the main model, using a R function provided by Mundry (2023). All models were fairly stable. To check for collinearity problems, we inspected variance inflation factors (VIF) (Field 2005) from a linear model with the same terms as the full model excluding the interaction and random effects using the function ‘vif’ in the R package “car” (v3.1.2; Fox and Weisberg 2019). We did not detect any collinearity issues (all VIF close to 1). We checked whether there was a problem with overdispersion in the two beta GLMMs and found the dispersion parameter of both to be acceptable (eavesdropping vs control condition model = 1.155; direct experience condition model = 1.065). We inspected the histograms of the Best Linear Unbiased Predictors (BLUPs) to confirm that there was no deviation from a normal distribution.