|
- mise à jour
du
- 23 mars
2014
- Anim
Cogn.
- 2026;29(1):60
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- Contagious
yawning is stronger in wolves
- than in
dog-admixed wolves
- Amici F, Liebal K, Oña L, Iacuzzi M,
Altdörfer T,
- Gretschzel A, Kamanga G, Delaunay M, Ciucci
P.
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- Tous
les articles sur la contagion du
bâillement
- All
articles about contagious
yawning
-
- Variation
in behavioural contagion patterns
-
- Behavioural contagion occurs when the
behaviour of one individual elicits the same
behaviour in another individual observing the
event, and it is considered one of the
fundamental mechanisms promoting group
coordination and cohesion in social species,
including humans.
-
- Empirical research on behavioural contagion
has focused primarily on yawning and has shown
evidence of it in several animal species. Here,
the authors studied behavioural contagion in 2
groups of captive wolves (N_=_20) and 3 groups
of captive dog-admixed wolves (N_=_13). They
conducted behavioural observations of naturally
occurring yawning events and monitored the
yawning behaviour of the other group members in
the two minutes following each event. Their
results showed that, in both admixed and
non-admixed wolves, individuals who had the
opportunity to observe the initial yawn were
significantly more likely to yawn within the
subsequent two minutes than those who had not
observed the event.
-
- Crucially, this effect was stronger in
non-admixed than in admixed wolves. Furthermore,
individuals that were more socially integrated
were more likely to elicit behavioural contagion
in both admixed and non-admixed wolves. In
contrast, neither individual rank nor dyadic
relationship quality between the yawner and the
observer mediated the probability of
contagion.
-
- By revealing variation in behavioural
contagion patterns between admixed and
non-admixed wolves, this study provides insights
into how introgressive hybridization with
domestic species may influence mechanisms
underlying social coordination and pack cohesion
in wolves.
-
- Variations
dans les schémas de contagion
comportementale
-
- La contagion comportementale se produit
lorsque le comportement d'un individu suscite le
même comportement chez un autre individu
qui observe la scène ; elle est
considérée comme l'un des
mécanismes fondamentaux favorisant la
coordination et la cohésion au sein des
groupes chez les espèces sociales, y
compris les humains.
-
- Les recherches empiriques sur la contagion
comportementale se sont principalement
concentrées sur le bâillement et
ont mis en évidence ce
phénomène chez plusieurs
espèces animales. Ici, les auteurs ont
étudié la contagion
comportementale au sein de deux groupes de loups
en captivité (N = 20) et de trois groupes
de loups métissés avec des chiens
en captivité (N = 13). Ils ont
mené des observations comportementales
d'épisodes de bâillement survenant
naturellement et ont suivi le comportement de
bâillement des autres membres du groupe au
cours des deux minutes suivant chaque
épisode. Leurs résultats ont
montré que, tant chez les loups hybrides
que chez les loups non hybrides, les individus
qui avaient eu l'occasion d'observer le
bâillement initial étaient
significativement plus susceptibles de
bâiller au cours des deux minutes
suivantes que ceux qui n'avaient pas
observé l'événement.
-
- Il est important de noter que cet effet
était plus marqué chez les loups
non métissés que chez les loups
métissés. De plus, les individus
les mieux intégrés socialement
étaient plus susceptibles de
déclencher une contagion comportementale,
tant chez les loups métissés que
chez les loups non métissés. En
revanche, ni le rang social de l'individu ni la
qualité de la relation dyadique entre
celui qui bâille et l'observateur n'ont
influencé la probabilité de
contagion.
-
- En mettant en évidence les variations
dans les schémas de contagion
comportementale entre les loups
métissés et non
métissés, cette étude
apporte un éclairage sur la
manière dont l'hybridation introgressive
avec des espèces domestiques peut
influencer les mécanismes sous-jacents
à la coordination sociale et à la
cohésion de la meute chez les loups.
-
- Introduction
- The ability to coordinate behaviour with
conspecifics can confer substantial fitness
benefits to group-living animals, for example by
strengthening social cohesion, facilitating
collective vigilance, improving the
effectiveness of anti-predator strategies and
increasing foraging efficiency through
cooperative hunting (Conradt and Roper 2000;
Duranton and Gaunet 2016; Gallup and Gallup
2007; Lakin et al. 2003). One mechanism that may
support such coordination is behavioural
contagion, which occurs when observing the
behaviour of one individual increases the
probability of performing the same behaviour
(Zentall 2003; Palagi et al. 2020). By promoting
temporal alignment of activities within dyads
and groups, behavioural contagion may contribute
to behavioural synchrony without requiring
explicit signalling or complex cognitive skills,
and it is therefore considered a powerful tool
to facilitate coordination and collective
behaviour (Duranton and Gaunet 2016; Massen and
Gallup 2017).
-
- Behavioural contagion is thought to arise
from simple perception&endash;action mechanisms
such as non-conscious mimicry, whereby observing
a behaviour automatically activates the
corresponding motor representation in the
observer, without requiring explicit
representation of others' internal states
(Chartrand and Bargh 1999; Lakin et al. 2003;
Yoon and Tennie 2010). Empirical research on
behavioural contagion has focused primarily on
yawning, and to a lesser extent on other
behaviours such as scratching, laughter or play
(Massen and Gallup 2017). The primary focus on
yawning likely stems from its being relatively
common, highly stereotyped and easily
identifiable (Provine 1986; Yoon and Tennie
2010), which makes it particularly suitable also
for observational settings. Spontaneous yawning
is widespread in vertebrates and has been
associated to a variety of physiological
functions, including arousal regulation,
thermoregulation and brain oxygenation
(Baenninger 1997; Gallup 2010; Massen et al.
2014; Smith 1999). In contrast, contagious
yawning is triggered by social stimuli and
appears to occur in fewer taxa than spontaneous
yawning (Massen and Gallup 2017). Supporting
this notion, experimental studies have failed to
show contagious yawning in some species that are
solitary or facultatively social, such as
red-footed tortoises (Wilkinson et al. 2011).
This absence suggests that contagious yawning is
unlikely to arise solely from a simple
perceptual-motor reflex and may instead depend
on the underlying social or cognitive
conditions.
-
- Evidence for contagious yawning has been
reported primarily in non-human primates,
including most great apes (e.g., Amici et al.
2013; Anderson et al. 2004; Campbell et al.
2009; Demuru and Palagi 2012; Palagi et al.
2014), several catarrhines (e.g., Gallo et al.
2021; Palagi et al. 2009; Paukner and Anderson
2006) and, more recently, both platyrrhines
(Valdivieso-Cortadella et al. 2023) and
strepsirrhines (Lemes et al. 2024; Valente et
al. 2023). Beyond primates, evidence for
contagious yawning has also been reported in
other taxa, including wolves (Canis lupus;
Romero et al. 2014), pigs (Sus scrofa; Norscia
et al. 2021), sheep (Ovis aries, Yonezawa et al.
2017), African elephants (Loxodonta africana;
Rossman et al. 2020) and parrots (Melopsittacus
undulatus; Gallup et al. 2015; Miller et al.
2012). In contrast, evidence in some species
remains equivocal. In domestic dogs (Canis lupus
familiaris), for example, findings are mixed:
while several studies report yawning contagion
in response to human yawns (Joly-Mascheroni et
al. 2008; Madsen and Persson 2013; Neilands et
al. 2020; Romero et al. 2013; Silva et al.
2012), others have found no such effect (Buttner
and Strasser 2014; Harr et al. 2009; O'Hara and
Reeve 2011). Moreover, there is currently no
evidence supporting contagious yawning in
dog&endash;dog dyads (Harr et al. 2009; O'Hara
and Reeve 2011), suggesting that behavioural
contagion in dogs might have been shaped over
the course of domestication for interspecific
interactions with humans rather than conspecific
partners (see Call et al. 2003, and Palagi and
Cordoni 2020).
-
- Some authors have also linked behavioural
contagion to empathy, arguing that
perception&endash;action mechanisms may allow
observers to match the emotional states of
others, giving rise to emotional contagion,
which is often considered a basic form of
empathy (Palagi et al. 2009, 2020; Platek et al.
2003; Preston and de Waal 2002). From this
perspective, behavioural contagion is expected
to vary across individuals and social contexts,
reflecting differences in empathic sensitivity.
For example, individuals sharing stronger social
bonds or higher familiarity are predicted to
show higher levels of behavioural contagion, as
empathy is assumed to be stronger in these dyads
(Palagi et al. 2009; Preston and de Waal 2002).
Similarly, some researchers have suggested that
in some species females may show higher levels
of empathy and, consequently, more pronounced
behavioural contagion than males, reflecting sex
differences in caregiving and parental
investment (Norscia et al. 2016; see Gallup and
Massen 2016). However, whether such patterns
provide direct evidence for emotional contagion
remains debated, as they may instead reflect
attentional biases toward more familiar or
socially salient group members (e.g., Gallup
2021; Massen et al. 2012; Massen and Gallup
2017). From this perspective, contagion may be
more likely to occur when observing individuals
with prominent social roles, such as
high-ranking or socially well-integrated group
members. Supporting this view, in Tibetan
macaques (Macaca thibetana), the individuals
that are more central in the social network
elicit significantly higher levels of contagion
than more peripheral group members (Zhang et al.
2022).
-
- To date, several studies have investigated
whether behavioural contagion is modulated by
the relationship quality, which has been
operationalized in terms of familiarity, kinship
or strength of the dyadic bond (Silk et al.
2009). In chimpanzees, for instance, contagious
yawning is more frequent in ingroup or familiar
individuals, as compared to outgroup or
unfamiliar ones (Campbell and de Waal 2011,
2014). Similarly, contagious yawning is
modulated by relationship quality in bonobos
(Demuru and Palagi 2012; Palagi et al. 2014),
geladas (Theropithecus gelada; Palagi et al.
2009) and wolves (Romero et al. 2014). However,
other studies have found no effect of
relationship quality on contagious yawning
(chimpanzees: Massen et al. 2012; Madsen et al.
2013; spider monkeys: Valdivieso-Cortadella et
al. 2023; parrots: Gallup et al. 2015), or even
reported opposite patterns (in rats, Rattus
norvegicus: Moyaho et al. 2015). In domestic
dogs, some studies found that relationship
quality positively affected the likelihood of
yawning contagion in response to human stimuli
(Joly-Mascheroni et al. 2008; Romero et al.
2013; Silva et al. 2012), whereas others
reported no effect (Neilands et al. 2020; O'Hara
and Reeve 2011; Madsen and Persson 2013).
-
- With respect to sex differences, there is
currently no consistent support for the
prediction that females are generally more
likely than males to show contagious yawning
across mammals (Massen and Gallup 2017).
Although one study on wolves showed that females
had shorter reaction times than males to yawns
produced by close social partners (Romero et al.
2014), several other studies reported no sex
biases in contagious yawning (e.g., Campbell et
al. 2009; Valente et al. 2023). Given that these
patterns are inconsistent across species, some
researchers have suggested that they may align
more closely with an attention-bias account than
with the hypothesis that behavioural contagion
reflects emotional contagion or empathy (Massen
and Gallup 2017). In chimpanzees, for example,
yawns produced by males are more contagious than
those produced by females (Massen et al. 2012),
whereas the opposite pattern has been reported
in bonobos (Demuru and Palagi 2012), suggesting
attentional biases toward the dominant sex
(Massen and Gallup 2017). Similarly, female
geladas show higher levels of contagious yawning
than males, but only when the observed yawner is
female (Palagi et al. 2009). In dogs there is no
evidence for sex-based modulation of contagious
yawning (Neilands et al. 2020; Romero et al.
2013).
-
- In this study, we investigated contagious
yawning in wolves and wolves admixed with dogs
(hereafter, admixed wolves), the latter
comprising introgressed individuals of second-
or later generation backcrosses to wolves (see
below). From a theoretical perspective,
comparing admixed and non-admixed wolves can
shed light on the selective pressures shaping
behavioural contagion and, more broadly,
behavioural coordination. While evidence for
contagious yawning is relatively well
established in primates (e.g., Massen and Gallup
2017), patterns in canids are less clear. In
wolves, only a single study has documented
contagious yawning (Romero et al. 2014). In this
study, wolves were more likely to yawn after
observing a conspecific yawning, which the
authors interpreted as evidence that contagious
yawning may help highly social species such as
wolves synchronize behavioural and physiological
states within the group, thereby promoting
coordination and social cohesion within the
pack. In domestic dogs, however, there is no
clear evidence for dog&endash;dog contagion
(Harr et al. 2009; O'Hara and Reeve 2011), and
the role of familiarity and relationship quality
remains inconsistent. This raises the
possibility that domestication may have shaped
behavioural contagion (see Palagi and Cordoni
2020), favouring the emergence of traits that
enhance attention to humans (e.g.,
Miklósi et al. 2003) and sensitivity to
human social cues (e.g., Hare and Tomasello
2005). If these traits are inherited from dogs
through introgressive hybridization, admixed
wolves may show contagion patterns that are
intermediate between wolves and domestic dogs.
Determining whether introgressed dog alleles
influence behavioural contagion, therefore, may
help clarify whether and how hybridization
affects group coordination and social cohesion
in admixed wolves, with important implications
for their ecology and social behaviour.
-
- Here, we made the following hypotheses and
predictions. Given that wolves form cohesive
packs characterised by strong affiliative bonds
and high levels of cooperation (Packard 2003,
2019), and that contagious yawning has been
documented in wolves (Romero et al. 2014) but
not in dog&endash;dog dyads (Harr et al. 2009;
O'Hara and Reeve 2011), we expected that both
admixed and non-admixed wolves would exhibit
behavioural contagion (Prediction 1a), but that
this effect would be stronger in wolves
(Prediction 1b). Furthermore, if the
distribution of behavioural contagion within
groups is shaped by attention- and/or
empathy-based biases, we predicted that
contagious yawning would be more likely between
individuals having stronger social bonds
(Prediction 2a); when the initial yawner was
socially more prominent, by having a higher rank
(Prediction 2b); or when the initial yawner
occupied a more central position in the social
network (Prediction 2c). However, such biases
may be more pronounced in cohesive groups, where
individuals form stronger affiliative
relationships, dominance hierarchies are more
stable, and individuals may differentially
allocate attention to socially salient partners
(see Amici et al. 2024, for a preliminary
comparison of social cohesion in admixed and
non-admixed wolves). Therefore, we also
predicted that the modulating effects of social
bond strength, rank and centrality would be
stronger in non-admixed than in admixed wolves
(Prediction 2d).
-
- Discussion
- In this study, we investigated behavioural
contagion in captive wolves and wolves admixed
(i.e., introgressed) with dogs. Our results
showed that, in both admixed and non-admixed
wolves, individuals who had the opportunity to
observe a yawn were significantly more likely to
yawn within the subsequent two minutes than
those who had not observed the event. Crucially,
this effect was stronger in non-admixed than
admixed wolves. Furthermore, individuals that
were more socially integrated in the group were
more likely to elicit behavioural contagion in
both admixed and non-admixed wolves. In
contrast, neither individual rank nor dyadic
relationship quality between the yawner and the
observer significantly influenced the
probability of yawning contagion.
-
- The presence of behavioural contagion in
both admixed and non-admixed wolves supports
Prediction 1a and aligns with previous findings
in wolves (Romero et al. 2014). More broadly,
our findings are consistent with evidence of
contagious yawning across a range of social
mammals and birds (e.g., Gallup et al. 2015;
Miller et al. 2012; Norscia et al. 2021; Rossman
et al. 2020), supporting the view that
behavioural contagion represents a widespread
mechanism facilitating behavioural synchrony
(Duranton and Gaunet 2016; Massen and Gallup
2017). Our study extends previous work by
showing that also admixed wolves, like
non-admixed ones, show behavioural contagion
when observing conspecifics yawning, suggesting
that the perception&endash;action mechanisms
underlying this phenomenon (Chartrand and Bargh
1999; Preston and de Waal 2002; Yoon and Tennie
2010) might be preserved in admixed wolves
despite introgression from domestic dogs, for
which evidence of behavioural contagion in
response to conspecifics is still lacking.
-
- At the same time, we found that behavioural
contagion was significantly stronger in
non-admixed than in admixed wolves, supporting
Prediction 1b. This difference is in line with
the hypothesis that dog domestication may have
favoured the emergence of social responsiveness
and attentional biases toward interspecific
rather than intraspecific partners, (Call et al.
2003; Miklósi et al. 2003; Hare and
Tomasello 2005), possibly decreasing the
likelihood of intraspecific behavioural
contagion in dogs and, through the introgression
of dog genes, in admixed wolves. Indeed, there
is no conclusive evidence yet for dog&endash;dog
contagious yawning (Harr et al. 2009; O'Hara and
Reeve 2011), although dogs have repeatedly been
shown to yawn contagiously in response to human
stimuli (Joly-Mascheroni et al. 2008; Romero et
al. 2013; Madsen and Persson 2013; Silva et al.
2012; Neilands et al. 2020). In the future, it
will be important to confirm these findings by
systematically testing dog&endash;dog social
responsiveness, attentional biases and
behavioural contagion in individuals living
under different conditions (e.g., dogs raised by
humans, dogs in kennels, free-ranging mongrel
dogs), to better disentangle evolutionary and
ontogenetic factors that might explain variation
in these behaviours. Moreover, our findings
align with previous work suggesting that social
networks might be more cohesive in non-admixed
than in admixed wolves (Amici et al. 2024). This
indicates that the introgression of dog genes
might potentially influence fine-scale
mechanisms of behavioural coordination, which,
in wolves, may facilitate synchrony and group
cohesion (Duranton and Gaunet 2016; Conradt and
Roper 2000). In the future, it will be essential
to confirm the differences in behavioural
coordination and group cohesion that we
revealed, by including more wolf and admixed
groups, possibly including recent hybrids (sensu
Stronen et al. 2025) and ideally in natural
settings.
-
- Individuals' integration in the social
network mediated behavioural contagion but, in
contrast to Prediction 2d, in a similar way for
admixed and non-admixed wolves. In all groups,
individuals that were more central in the social
network were more likely to elicit yawning
contagion, when the yawning event had been
observed. This suggests that socially
well-integrated individuals might exert greater
influence on the behavioural responses of group
members. Although the effect size appears
modest, it mirrors findings in primates, where
more central individuals elicited higher levels
of contagion (Zhang et al. 2022). Social
centrality may therefore represent a robust
proxy for social prominence, capturing an
individual's embeddedness within the group and,
consequently, its social salience (see Farine
2017). From an attention-based perspective
(Massen et al. 2012; Massen and Gallup 2017),
individuals that are more integrated may attract
greater visual monitoring from others, thereby
increasing opportunities for behavioural
contagion.
-
- In contrast, neither dominance rank nor
dyadic relationship quality (CSI) significantly
mediated behavioural contagion, neither in
admixed nor in non-admixed wolves. The absence
of a CSI effect is particularly noteworthy, as
previous studies have reported higher levels of
contagious yawning between closely bonded
individuals in chimpanzees and bonobos (Campbell
and de Waal 2011, 2014; Demuru and Palagi 2012;
Palagi et al. 2014), geladas (Palagi et al.
2009) and wolves (Romero et al. 2014). Our
findings therefore do not support Prediction 2a
and contrast with studies linking contagion to
relationship quality or familiarity. One
possible explanation is methodological. Our CSI
was calculated exclusively on spatial proximity
(Silk et al. 2009), which may not fully capture
the multidimensional nature of affiliative
bonds. Wolves, in particular, are known to form
complex and highly differentiated social
relationships within their packs (Packard 2003,
2012). These relationships are expressed not
only through spatial association but also
through coordinated and cooperative activities
such as hunting (MacNulty et al. 2009, 2012),
breeding (Mech 1999; Packard et al. 1992) and
territorial defence (Harrington and Mech 1979;
Packard 2003). By relying solely on proximity as
an indicator of bond strength, we may therefore
have overlooked important behavioural components
of social relationships, potentially limiting
our ability to detect bond-related modulation of
contagion. However, it is also possible that
social bond strength is simply not a relevant
factor in contagious yawning, in line with other
literature on domestic dogs (Neilands et al.
2020; O'Hara and Reeve 2011; Madsen and Persson
2013), primates (e.g., Massen et al. 2012;
Madsen et al. 2013; Pedruzzi et al. 2025;
Valdivieso-Cortadella et al. 2023) and parrots
(Gallup et al. 2015). Similarly, the absence of
rank effects contrasts with our Prediction 2b
and suggests that dominance status alone does
not significantly influence behavioural
contagion in admixed and non-admixed
wolves.
-
- Several limitations of our study should be
acknowledged. First, the number of study groups
and individuals was necessarily limited, as
facilities housing groups of wolves admixed with
dogs are extremely rare. Therefore, variation in
enclosure size, group size and composition, as
well as early life experiences, could not be
fully controlled for. In particular, most
admixed wolves spent few weeks in the wild with
their mother and were subsequently raised in
captivity, whereas the zoo-born wolves were born
and raised entirely in captivity, with regular
exposure to keepers and visitors. These
differences were mostly graded rather than
categorical, making it impossible to fully
capture them with simple descriptors. It
therefore cannot be excluded that such
developmental differences influenced social
responsiveness or attentional patterns.
Moreover, wolf groups in our study were slightly
larger than admixed groups, which raises the
possibility that group size may have contributed
to the observed difference. In principle, larger
groups might promote behavioural contagion
because individuals are exposed to a greater
number of potential demonstrators and thus to
more opportunities for contagion. In the present
study, however, we accounted for this by
modelling behavioural contagion as the odds that
an individual yawned as a function of whether it
had observed a trigger event or not.
-
- This approach reduced the likelihood that
higher rates in larger groups simply reflect the
presence of more potential yawners.
Nevertheless, it is also possible that
behavioural contagion plays a stronger
functional role in larger groups, where
mechanisms promoting behavioural synchrony may
be especially important to maintain group
cohesion and coordinated activity. If so,
variation in behavioural contagion may reflect
the specific socio-ecological conditions
experienced by a group, rather than selective
pressures working at the evolutionary level, an
avenue that warrants further investigation.
Second, as in most previous studies of
contagious yawning (e.g., Romero et al. 2014;
Massen and Gallup 2017), our data were collected
in captive settings. Although our modelling
approach allowed us to control for differences
in the baseline occurrence of trigger events,
which might also be linked to specific living
conditions, caution is still warranted when
extrapolating these findings to wild
populations. Third, wolf-dog hybridization
represents a continuum, and even if our findings
were confirmed with larger samples, caution
would remain necessary when generalising to more
recent hybrids (sensu Stronen et al. 2025) or
introgressed individuals with different degrees
of admixture, as deviations from species-typical
wolf behaviour may be expected to scale with the
proportion of dog ancestry. Fourth, although we
conducted genetic analyses to confirm
introgression with dogs in our admixed groups,
we did not investigate the genomic architecture
potentially underlying behavioural variation.
Yet recent genomic research suggests that even
low levels of dog introgression may affect
specific brain function and behaviour in wolves
(Pilot et al. 2021), including variants involved
in neurotransmission and neurodevelopment. Given
the strong genetic basis of behavioural traits
in dogs (Morrill et al. 2022; Salomons et al.
2021), and the extensive impact of artificial
selection on the dog genome (Bergström et
al., 2020; Freedman et al., 2016), it is
plausible that introgressed alleles may
contribute to specific behavioural variation in
wild wolves subject to various extents of
introgression from dogs (Leonard et al. 2013).
-
- If the observed differences between admixed
and non-admixed wolves are replicated, future
research integrating behavioural data with
finer-scale genomic analyses will be essential
to determine whether specific introgressed gene
variants are associated with variation in social
responsiveness and coordination. Finally, for
logistic constraints, the timing of data
collection varied across the five groups
studied, introducing additional potential
confounding factors that we could not fully
account for. Temperature, for instance, is known
to affect yawning behaviour (Campos and Fedigan
2009; Eldakar et al. 2015; Gallup et al. 2011;
Massen et al. 2014), and although our models
partially controlled for baseline yawning
propensity (e.g., by also modelling individual's
likelihood to yawn when not observing the
trigger event), future studies should ideally
incorporate environmental temperature more
explicitly as a predictor in the models.
Similarly, we did not measure circulating
hormone levels or account for variation in
reproductive or gonadal status across
individuals and groups, although this would
ideally have been controlled for, given that
behavioural sampling occurred both within and
outside the breeding season. This may be
relevant, as hormonal state might be linked to
empathy-related processes and contagious yawning
(e.g., Kis et al. 2020). Future work should
therefore consider including hormonal status or
proxies thereof to better control for its
potential effects on behavioural contagion.
-
- In conclusion, our findings support
the view that dog introgression may modulate
behavioural contagion in admixed wolves,
possibly by modifying individual attentional
biases. Given that hybridization with dogs is an
emerging threat for several European wolf
populations (Ciucci et al. 2026), understanding
how introgression might affect behavioural
processes linked to pack cohesion is of growing
importance. While our findings should not be
interpreted as evidence of broad behavioural
disruption, they indicate that genetically
detectable levels of admixture may indeed
influence fine-scale social dynamics.
Behavioural modifications linked to admixture
could, in turn, have cascading ecological
consequences: changes in attentional biases and
social responsiveness may affect pack size and
cohesion, social interactions within and between
packs, reproductive success through alloparental
care and dispersal dynamics (Newsome et al.
2017; Sparkman et al. 2012). Integrating
behavioural analyses with ecological,
demographic and genomic data will therefore be
essential to determine whether such differences
translate into measurable consequences for group
coordination, cohesion and eventually the
ecological role of wolves.
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