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Cats’ (Felis catus) communicative behaviour towards humans was explored using a social referencing paradigm in the presence of a potentially frightening object. One group of cats observed their owner delivering a positive emotional message, whereas another group received a negative emotional message. The aim was to evaluate whether cats use the emotional information provided by their owners about a novel/unfamiliar object to guide their own behaviour towards it. We assessed the presence of social referencing, in terms of referential looking towards the owner (defined as looking to the owner immediately before or after looking at the object), the behavioural regulation based on the owner’s emotional (positive vs negative) message (vocal and facial), and the observational conditioning following the owner’s actions towards the object. Most cats (79 %) exhibited referential looking between the owner and the object, and also to some extent changed their behaviour in line with the emotional message given by the owner. Results are discussed in relation to social referencing in other species (dogs in particular) and cats’ social organization and domestication history.
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Social referencing and cat–human communication
I. Merola M. Lazzaroni S. Marshall-Pescini
E. Prato-Previde
Received: 4 August 2014 / Revised: 23 December 2014 / Accepted: 24 December 2014
ÓSpringer-Verlag Berlin Heidelberg 2015
Abstract Cats’ (Felis catus) communicative behaviour
towards humans was explored using a social referencing
paradigm in the presence of a potentially frightening
object. One group of cats observed their owner delivering a
positive emotional message, whereas another group
received a negative emotional message. The aim was to
evaluate whether cats use the emotional information pro-
vided by their owners about a novel/unfamiliar object to
guide their own behaviour towards it. We assessed the
presence of social referencing, in terms of referential
looking towards the owner (defined as looking to the owner
immediately before or after looking at the object), the
behavioural regulation based on the owner’s emotional
(positive vs negative) message (vocal and facial), and the
observational conditioning following the owner’s actions
towards the object. Most cats (79 %) exhibited referential
looking between the owner and the object, and also to some
extent changed their behaviour in line with the emotional
message given by the owner. Results are discussed in
relation to social referencing in other species (dogs in
particular) and cats’ social organization and domestication
Keywords Social referencing Cats Gaze alternation
Social learning Human–cat communication
Cats (Felis catus) are one of the most widespread and
beloved companion animals: they are ubiquitous, share
their life with people and are perceived as social partners
by their owner (Karsh and Turner 1988). Recent findings
suggest that their association with humans can be traced
back to approximately 8,000–10,000 years ago (Davis
1987; Vigne et al. 2004) and not just to 4,000 years ago, as
previously thought (Serpell 2000). In fact, recent genetic
and archaeological evidence indicates that the cat was
actually domesticated in the Near East (Driscoll et al.
2007) about 10,000 years ago (Vigne et al. 2004). Thus,
like dogs (Canis familiaris), cats began to live in associa-
tion with humans in ancient times, though dogs’ history of
domestication is thought to be substantially longer (Savo-
lainen et al. 2002; von Holdt et al. 2010; Hu et al. 2014).
One hypothesis relating to cat domestication is that the
driving force behind this process was a mutualistic rela-
tionship established between people and cats, with wild
cats learning to exploit the human environment, feeding on
rodents attracted by humans’ stocks of grains and cereals
(Clutton-Brock 1988) and on food scraps found in human
settlements. These factors coupled with humans’ tolerant
attitude towards them, potentially due to the recognition of
their utility in keeping rodents at bay, would have con-
tributed to the rapid domestication process of this species
(Todd 1978; Driscoll et al. 2009; Hu et al. 2014). Thus, it
I. Merola M. Lazzaroni S. Marshall-Pescini
E. Prato-Previde
Dipartimento di Fisiopatologia Medico-Chirurgica e dei
Trapianti, Sezione di Neuroscienze, Universita
`degli Studi di
Milano, Via F.lli Cervi 93, 20090 Segrate, MI, Italy
I. Merola (&)
School of Life Sciences, Joseph Bank Laboratories, University
of Lincoln, Green Lane, Lincoln LN6 7DL, Lincolnshire, UK
S. Marshall-Pescini
Comparative Cognition, Messerli Research Institute, University
of Veterinary Medicine, Vienna, Medical University of Vienna,
University of Vienna, Vienna, Austria
Anim Cogn
DOI 10.1007/s10071-014-0832-2
has been suggested that the domestic cat represents a
product of self-domestication and natural selection and that
compared to dogs or other domestic species, it has been
exposed to a less strict and conscious process of artificial
selection (Clutton-Brock 1988; Driscoll et al. 2009).
Despite a shorter history of domestication and a less
intense artificial selection process than dogs, the living
environment of cats and humans has overlapped consid-
erably (Bradshaw et al. 1999), with cats establishing
enduring relationships with humans, which often start at
early stages of their development and last all their lives.
In recent years, an increasing number of studies have
investigated social cognitive skills in domestic species such
as dogs (see Bensky et al. 2013; Kaminski and Nitzschner
2013; Prato-Previde and Marshall-Pescini 2014 for recent
reviews), horses (McKinley and Sambrook 2000; Maros
et al. 2008; Proops and McComb 2010; Kru
¨ger et al. 2011;
Proops et al. 2013), pigs (e.g. Albiach-Serrano et al. 2012)
and goats (e.g. Kaminski et al. 2005), testing the hypothesis
that domestication has favoured, at least in some species,
the emergence of a number of behavioural changes and
cognitive skills evolved to better exploit the human world
and effectively communicate with humans (Kaminski et al.
2005; Miklo
´si and Soproni 2006; Hare et al. 2010; Udell
and Wynne 2010).
Cats and dogs are the most common non-human animals
interacting with us daily: they have fully adapted to the
human social environment and are capable of establishing
long-term social relationships with humans (Miklo
´si et al.
2005). However, whereas the communicative abilities and
the ability to discriminate human emotional expressions of
domestic dogs have received growing attention (Nagasawa
et al. 2011; Merola et al. 2013b), there are only a few
studies that have investigated cat cognitive abilities (e.g.
Pisa and Agrillo 2009; Whitt et al. 2009; Pan et al. 2013),
and even fewer looking at cat social cognition and cat–
human communication and expression (e.g. Miklo
´si et al.
2005; Saito and Shinozuka 2013).
Unlike their wild ancestors (Felis silvestris), domestic
cats are social animals (Crowell-Davis et al. 1997; Mac-
donald et al. 2000). They show intraspecific communica-
tion not present in other solitary felids (Bradshaw and
Cameron-Beaumont 2000) and entertain strict and complex
relationships with their owners, adapting flexibly to them
(Mertens 1991; Turner 1991; Leyhausen 1988; Rieger and
Turner 1999). There is evidence that cats react to unfa-
miliar and familiar humans differently (Collard 1967;
Casey and Bradshaw 2008) and recognize their owners’
voices, distinguishing them from other human voices (Saito
and Shinozuka 2013). Furthermore, it has been shown that
they become bonded to their owners and that the cat–owner
relationship appears to fulfil the behavioural criteria for an
attachment bond (Edwards et al. 2007).
However, to our knowledge, only one study has inves-
tigated cat–human communication by comparing the abil-
ity of both cats and dogs to communicate with humans by
either responding to a person’s gestural signals (i.e.
pointing) or by using attention-getting signals (i.e. gaze and
gaze alternation) to communicate in a feeding context
´si et al. 2005). This study showed that cats, like
dogs, were successful in using four different types of
human pointing cues (differing in visibility and duration of
the given cue and in the distance between the end of the
fingertip and signalled object) to locate hidden food in an
object-choice task. However, when facing a problem situ-
ation in which the food was hidden in an inaccessible place
and human intervention was needed, cats lacked certain
components of the attention-getting behaviour shown by
dogs. Indeed, dogs showed higher levels of gazing
behaviour, looked earlier and for longer at their owners,
and showed more gaze alternation between the hidden food
and the human compared to cats. Thus, these results sug-
gest the presence of similar abilities in ‘reading’ human
signals (i.e. pointing) in cats and dogs, but differences in
their tendency to communicate with humans in a problem
situation (i.e. the impossibility of reaching food).
It has recently been shown that dogs engage in human-
directed communication (looking behaviour and gaze
alternation) not only to request human intervention when
unable to obtain a desired goal, but, also in a context of
uncertainty, possibly as a way to monitor their human
partner’s behaviour and synchronize their own response to
an unfamiliar environment with him/her (see Prato-Previde
and Marshall-Pescini 2014 for a review). In particular,
Merola et al. (2012a,b,2013a) found that, like human
infants (Mumme et al. 1966; Vaish and Striano 2004;de
Rosnay et al. 2006) and human-raised chimpanzees (Rus-
sell et al. 1997), dogs look at humans when facing unfa-
miliar situations that are difficult to interpret, and act in
accordance with the informer’s positive or negative emo-
tional reactions, a process known as ‘social referencing’. In
these studies, when confronted with a new and potentially
frightening object (a fan with ribbons attached to it) in the
presence of their owner providing either a positive or a
negative emotional message towards it, dogs engaged in
visual communication with him/her (referential looking)
and also changed their behaviour towards the object in line
with the emotional message received (i.e. behavioural
The aim of the current study was to investigate cat–
human communication in a social referencing context.
Using a procedure similar to that previously used by Me-
rola et al. (2012a) with dogs, we (1) tested whether cats
would show referential looking (defined as looking to the
owner immediately before or after looking at the object),
when presented with a potentially frightening stimulus; (2)
Anim Cogn
assessed the effect on cats’ behaviour of the emotional
(facial and vocal) message (positive vs negative) expressed
by the owner towards the ambiguous object, and (3) eval-
uated whether cats would be influenced by their owner’s
overt approach versus avoidance behaviour towards the
object (observational conditioning).
To assess the potential presence of referential looking,
cats were initially confronted with an ambiguous stimulus
in the presence of their silent and neutral owner. Then, to
evaluate the behavioural regulation aspect of social refer-
encing, cats’ behaviour was recorded when the owner
delivered either a positive (happy) or a negative (fearful)
message about the ambiguous stimulus using only their
voice and facial expression. Finally, in the last stage of the
experiment, we evaluated whether cats would, through a
process of observational conditioning (Whiten et al. 2004;
Zentall 2006), be influenced by their owner’s approach
versus avoidance behaviour towards the stimulus.
We predicted that if cats used human-directed gazing
behaviour to obtain information about the new ambiguous
stimulus, they would look at it and rapidly look at the
owner (referential looking). Furthermore, if cats used
humans’ vocal and facial emotional expressions to guide
their behaviour, we would expect cats in the negative
group to show more interaction with the owner, gaze
alternating more often between the fan and the exit
screen, and showing more stress signals, whereas cats in
the positive group would vocalize, alternate gaze between
the fan and the owner more, and be more likely to enter
the fan zone.
Thirty-six cat–owner dyads participated in the study.
Owners were all female except one. Cats (20 neutered
males, 13 neutered females, and 3 tom cats), 8 of pure
and 28 of mixed breed, ranged in age from 2 to
13 years (mean =5.9 years; SD =3). All cats were pets
living at home with their owners. As inclusion criteria, we
required cats to be friendly towards strangers, not escaping
or showing aggressive behaviours towards them, but rather
seeking contact with them. Furthermore, they had to be
used to changes in their living environment (e.g. going on
holiday with their owners) and accustomed to travelling in
a carrier at least twice a month. To assess the above cri-
teria, we asked owners on the phone prior to inviting them
to the laboratory. No tests were carried out during the
preselection of the subjects. Following this initial selection,
cat–owner dyads were then randomly assigned to the
positive and negative message group.
Unfamiliar stimulus
The experimental stimulus was the same for all cats in both
conditions (positive and negative): a 50-cm-tall and 34-cm-
wide electric fan, with plastic green ribbons attached to it.
This stimulus was the same as used in previous studies on
social referencing in dogs (Merola et al. 2012a,b) and was
aimed at eliciting a mild fear reaction, i.e. neither very
positive (approaching directly and touching), nor very
negative (running in the opposite direction or strong stress
such as trembling, or hiding). It did not elicit predatory
The cats were individually tested in an unfamiliar
(3.5 94.5 m) room at the laboratory Canis Sapiens of the
University of Milan. The testing room was an empty space
with a black screen (h1m,w30 cm) at one end: the screen
hid a video camera and prevented the cats from going out
of sight (see Fig. 1). It had previously been used by testing
dogs, but was thoroughly cleaned before starting experi-
ments with cats. This screen was built in a way that cats
could hardly jump over it, but they could see the space
behind it. On arrival, the owner and the experimenter
entered the testing room and for 5 min the experimenter
explained the procedure to the owner, who was then asked
to repeat and also perform the entire procedure as if the cat
was being tested. During this time, the cat remained in
another room in its open carrying basket, free to stay inside
it or to move around. Then, the owner and experimenter
left the testing room, and the owner re-entered holding the
cat in his/her arms.
As soon as the door was closed, the owner reached
Location 1 holding the cat in his/her arms and then put the
cat on the floor. Prior to starting the test, the cat was
allowed to move around and explore the room for 1 min in
the presence of the silent and relaxed owner standing in
Location 1. After this exploration time, the test started. The
test lasted 125 s and was divided into four phases. During
the first three phases, the fan, placed at the far end of the
room (see Fig. 1), was in motion; during Phase 4, it was
switched off. Each test phase was characterized by the
owner behaving in a different way. Since in Phase 3 and 4
owners were required to move to specific locations in the
room, coloured sticky tape was placed in the appropriate
spots (Fig. 1). Each cat was allocated to one group only
and thus exposed either to the positive or negative
1 Norwegian forest, 1 Siamese, 1 Bengal, 3 Devon Rex, 2 Maine
Anim Cogn
Test phases were as follows:
Phase 1: this phase was the same for both the positive and
negative groups. The owner, while standing at Location
1, called the cat by name as soon as the cat approached
he/she crouched over it with his/her body and turned so as
to have their backs to the door whilst petting the cat. This
position and action prevented the cat from seeing the
experimenter opening the door and positioning the fan in
front of it. The fan was then activated via remote control.
Immediately after the experimenter left the room, the
owner released the cat and stood at Location 1 facing the
fan, while the cat was free to move around the room. At
this moment, we activated a stopwatch. The owner
remained silent looking at the fan with a neutral facial
expression for 25 s.
Phase 2: in this phase, regardless of the cat’s behaviour,
the owner stood in Location 1 and talked throughout the
whole phase, using either a happy (positive group) or
fearful (negative group) voice and facial expression and
gaze alternating between the cat and the fan continu-
ously for 25 s.
Phase 3: in the positive group, the owner approached the
fan reaching Location 2, crouched down facing the fan
and started to touch it, whilst still talking in a happy
voice and expressing a positive emotion and gaze
alternating between the cat and the fan continuously;
in the negative group, the owner moved away from the
fan reaching Location 3, crouching down whilst talking
with a fearful voice, expressing a negative emotion and
gaze alternating between the cat and the fan. In both
groups, the phase lasted 25 s.
Phase 4: This phase lasted 50 s and started when the
experimenter turned the fan off from an adjacent room
using a remote control. In the positive group, whilst still
crouching in Location 2, talking in a positive manner
and gaze alternating between the cat and the fan
continuously, the owner touched the fan and the ribbons
for the entire phase. In the negative group, the owner
stayed crouched down in Location 3 whilst continuing to
talk with a negative tone of voice for the entire phase
and gaze alternating between the cat and the fan
In both groups, in Phases 2, 3 and 4, the owners were
instructed to continue talking throughout the entire phase
and to communicate with their cats as they would nor-
mally, using typical phrases such as ‘that’s nice’ or ‘that’s
scary’, accompanied by either a smiling happy face or a
scared worried expression. They were also explicitly told
not to use the cat’s name and potential directions (look,
come, touch, etc.). Finally, they were instructed to show,
through facial and vocal expressions, the feeling either that
the cat could safely and happily approach the object, or that
the object was to be avoided. After the test ended, the
researcher entered the room with a handful of treats and
asked to the owner to sit next to the fan, giving the cat
treats when it came into proximity of the fan. If the cat was
not eating the treat, the owner and the experimenter sat
next to the fan until the cat started to explore the room in a
relaxed manner. All cats, regardless of condition, received
this treatment so that they would not become sensitive to
Data collection and analysis
The test was recorded by two video cameras (Panasonic
NVGS330) and analysed using Solomon Coder (beta
081122, Copyright 2006–2008 by Andra
Five nonmutually exclusive categories of behaviour
were recorded: gaze, action, body posture, stress signals
(following van den Bos 1998) and vocalization. In addi-
tion, the location of the cats when in closer proximity to the
fan (i.e. within 50 cm), during each phase of the test, was
recorded (fan zone) (see Table 1).
As in previous studies on dogs (Merola et al. 2012a,b,
2013a), in line with Russell et al. (1997), referential
Fig. 1 Experimental room (3.4 m 93.9 m) with owner’s location
during the different phases of the test (Location 1 (L1), Location 2
(L2), Location 3 (L3). The black thicker line is representing the
screen, while the two thin lines show the owner and fan zone. The
distance between L1 and L2 was 1.6 m, while the distance between
L1 and L3 was 1.3 m
Anim Cogn
looking was defined as a gaze towards the owner that was
preceded by a look to the fan, and gaze alternation as a
consecutive sequence of three looking behaviours (fan–
owner–fan or owner–fan–owner). The percentage of cats
carrying out these behaviours is recorded for Phase 1 for
comparison with other species. A generalized linear model
analysis (binomial distribution) was carried out to assess
whether the likelihood of alternating the gaze between
owner and fan and fan and screen (hence two-way alter-
nations) would vary according to group.
The screen was a barrier placed at the far end of the
room behind the initial location of cat and owner and far-
thest away from the fan. As mentioned above, this barrier
hid the video camera placed on a tripod but was not tall
enough to reach the ceiling; hence, cats could see that there
was space behind it. The fact that the screen was the fur-
thest location from the fan and that cats seemed aware of
there being space behind it led us to analyse both gazing
and interaction behaviours directed at the screen as possi-
ble indication that cats were looking for a way out of the
room (keeping in mind that the door from which they had
entered was located directly behind the fan during testing).
A series of generalized linear models (Poisson distri-
bution) were used to evaluate the potential group differ-
ences on the frequencies of behaviours. Given their
relatively low frequencies, the three behaviours towards the
owner were considered as a sum (interact owner, rubbing
against the owner and tail up whilst interacting with the
owner). A generalized linear model (binomial distribution)
was also used to assess whether the likelihood of entering
the fan zone was affected by group and phase. Finally, we
analysed the potential group differences in the latency and
the duration of being static and moving around the room
(locomotion), using a Mann–Whitney test, since despite
transformation the residuals were not normally distributed.
The cats’ behaviour was coded by M. L., and a second
independent coder (E. P.) analysed 25 % of the data. The
interobserver reliability on the duration of the major
behavioural categories analysed was calculated using
Cronbach’s alpha (gaze fan a=0.97; gaze owner
a=0.79; gaze screen a=0.89; static a=0.98; interact
owner a=0.94; tail up interact owner a=0.85; vocali-
zation a=0.98). GLM analyses were conducted using the
lme4 package in R ( All other analyses
were conducted in SPSS v.19. All tests were two-tailed,
and the alpha level was set at 0.05.
Of the 36 cats tested, nine (6 males and 3 females) were
excluded from the analyses: one subject (male) because of
procedural errors committed by the owner during testing,
and eight (five males and three females) because they
succeeded in jumping or hiding behind the screen during
the test.
Of the remaining 27 subjects, 3 (all males) approached
and touched the fan during Phase 1 exhibiting a confident
and positive attitude towards the stimulus. These cats were
excluded both in the analysis of referential looking/gaze
alternation behaviour (Phase 1), and of behavioural regu-
lation, since a precondition for the test was that cats had an
Table 1 Ethogram of the behaviours analysed during the four phases of the study
Gaze Gaze owner Cat’s head is oriented towards the owner
Gaze fan Cat’s head is oriented towards the fan
Gaze screen Cat’s head is oriented towards the screen at the end of the room
Action Static Cat is in a position which does involve movement, i.e. standing, sitting or lying
Locomotion Any behaviour involving moving around the room whether walking with head down/sniffing, or pacing
whilst looking at the owner/object
Interact owner Cat is in physical contact with the owner
Rubbing owner Cat is around the owner and is rubbing with one side of his/her body (face or lateral)
Interact fan Cat is in physical contact with the fan
Interact screen Cat is in physical contact with the screen at the end of the room
Tail up
Tail down
Tail up Interaction
Stress Stress signals Yawning, head shaking, lips licking
Vocalization Meowing
Zone Fan zone 1 m 93 m 50 cm around the fan
Anim Cogn
ambiguous (or mildly fearful) behaviour towards the
stimulus object (Feinman 1982; Gunnar and Stone 1984;
Rosen et al. 1992).
Referential looking and gaze alternation
To assess whether cats carried out referential looking when
confronted with the ambiguous stimulus, we analysed cats’
referential looking and gaze alternation behaviour in Phase
1 (regardless of group since this phase was the same for all
cats). Of the 24 cats considered (9 F and 15 M), 19 (79 %)
showed referential looking towards the owner at least once
(and a maximum of eight times). As regards gaze alter-
nation, 13 out of 24 cats (54 %) showed at least one and a
maximum of three gaze alternation sequences (i.e. fan–
owner–fan or owner–fan–owner).
Behavioural regulation
Given that cats showed referential looking towards the
owner in Phase 1, when confronted with an ambiguous
stimulus, we then evaluated whether they would be dif-
ferently affected by the owners’ positive versus negative
emotional expressions. Of the 24 cats that showed an
ambiguous approach towards the fan in Phase 1, 12 (6 FN
and 6 MN) were in the positive message group and 12 (3
FN, 8 MN and 1 MUn) in the negative message group (in
this group, there was the only male owner).
There was no interaction (glm z=0.75, P=0.46) and
no main effect for neither phase (glm z=0.35, P=0.73)
nor group (glm z=1.76, P=0.08) on the likelihood of
cats gaze alternating between the fan and the owner. There
was no effect of phase (glm z =1.7, P=0.09) and no
interaction (glm z=0.46, P=0.64) between phase and
group on the likelihood of cats showing gaze alternation
between the screen and the fan. However, overall, cats in
the negative group were significantly more likely to gaze
alternate between the screen and the fan than cats in the
positive group (glm z=3.9, P\0.001).
Results showed no interaction between group and phase
(glm Phase 2 z=1.69, P=0.09; Phase 3 z=1.22,
P=0.22; Phase 4 z=0.42, P=0.67), but a main effect
of both group (glm z=4.43, P\0.001) and phase with
cats alternating their gaze significantly less between the
screen and the fan in Phase 4 than Phase 1 (glm Phase 4
z=5.7 P\0.001; Phase 2 z=1.7 P=0.08; Phase 3
z=0.02, P=0.9), and cats in the positive group alter-
nating their gaze between the screen and the fan signifi-
cantly less than cats in the negative group.
An interaction emerged between phase and group in
gaze alternation between fan and owner in which the pat-
tern of results was significantly different for groups in
Phase 4 than in Phase 1 (glm group*Phase 4 z=0.36,
P\0.001; group*Phase 3, z=0.02, P=0.98; group*-
Phase 2 z=1.73, P=0.08). Since we were mainly
interested in group differences in each phase of the test, we
ran separate models comparing the behaviour of cats in the
two groups in each phase. No significant difference
emerged in Phase 1 and Phase 2 (glm Phase 1 z=0.7
P=0.5; Phase 2 z=0.4, P=0.7); however, cats in the
positive group alternated their gaze between fan and owner
significantly more than cats in the negative group both in
Phase 3 (glm z=54.8, P\0.001) and Phase 4 (glm
z=3.05, P\0.001).
No interaction emerged between group and phase in the
frequency of interacting with the owner (glm Phase 2,
z=0.08, P=0.93; Phase 3, z=0/9, P=0.37; Phase 4,
z=0.35, P=0.72). However, overall, cats in the negative
group interacted with their owner more frequently than cats
in the positive group (glm z=3.2 P\0.001), and cats
interacted more in Phase 3 and Phase 4 compared to Phase
1, but no differences emerged between Phase 1 and Phase 2
(glm Phase 2, z=0.53, P=0.14; Phase 3, z=4.36,
P\0.001; Phase 4, z=3.53, P\0.001).
An interaction between group and phase emerged on the
frequency of meowing (glm Phase 2, z=1.7, P=0.09;
Phase 3, z=2.55, P=0.01; Phase 4 z=2.44,
P=0.014). Because we were predominantly interested in
the potential differences between groups, we ran sub-
sequent models comparing cats in the positive versus
negative group in each phase. Cats in the positive group
meowed more frequently than cats in the negative group
but only in Phase 4 (glm Phase 1, z=0.74, P=0.46;
Phase 2, z=0.94, P=0.34; Phase 3, z=1.64, P=0.11;
Phase 4, z=1.97, P=0.048).
Stress signals were infrequent, with only eight cats
showing between 1 and 2 stress signals during the whole
test. Hence, we ran a generalized linear model (with
binomial distribution) including Phases 2 to 4 as a whole
and analysing whether the likelihood of a cat expressing a
stress signal was affected by which group they were in. No
such effect was found (glm z=0.86, P=0.4). Further-
more, the likelihood that cats would enter the ‘fan zone’
was not affected by the group they belonged to in none of
the test phases (glm Phase 2, z=0, P=1; Phase 3,
z=0.12, P=0.9; Phase 4, z=0.82, P=0.4).
Interaction with the fan, which occurred only in Phase 4,
was shown by only two cats in the positive group and one
in the negative group.
Duration and latency (static and locomotion)
In Phase 2 (when the owner was in location 1 expressing
either a positive or negative emotional message), there was
a tendency between groups in the latency to show loco-
motion (z=1.91, df =23, P=0.055), with cats in the
Anim Cogn
negative group showing this behaviour earlier than cats in
the positive group. No difference emerged in the duration
of neither static nor locomotion (static z=0.96, df =23,
P=0.33, locomotion z=1.5, df =23, P=0.13). How-
ever, in Phase 3 (when the owner either approached or
moved away from the fan), cats in the two groups differed
in the amount of time spent in static behaviour (z=2.36,
df =23, P=0.017), with cats in the positive group being
more static than those in the negative one. No difference
emerged in the latency to perform these behaviours in this
phase (static z=0.37, df =23, P=0.7, locomotion
z=1.43, df =23, P=0.14). Finally, in Phase 4, no dif-
ferences emerged in duration (static z=0.37, df =23,
P=0.7, locomotion z=0.34, df =23, P=0.72) and
latency (static z=0.76, df =23, P=0.44, locomotion
z=0.82, df =23, P=0.41) of neither static nor
The aim of this study was to investigate cat–human com-
munication by evaluating the presence of a social refer-
encing process in cat–human dyads. As no other studies
have been carried out in cats on this topic before, we aimed
at assessing the presence of referential looking and
behavioural regulation, based on the owners’ emotional
expression (vocal and facial). Furthermore, we evaluated
the occurrence of observational conditioning.
Results showed that 79 % of cats looked referentially
towards their owner and the fan and 54 % showed gaze
alternation (three consecutive looking behaviours) when
the owner was still and silent in the room. This percentage
was similar to the one found in previous studies on dogs,
where 76 % showed referential looking and 62 % showed
gaze alternation (Merola et al. 2012a,b); this suggests that
cats, like dogs, will look to their owner when faced with an
ambiguous object. Our results contrast with those found in
a previous study on gazing behaviour in cats (Miklo
´si et al.
2005), in which cats, when facing a situation in which food
was in an inaccessible place, showed lower rates of gazing
behaviour, looking later and for shorter periods of time as
well as showing less gaze alternation between the human
and food compared with dogs. One possible explanation for
the difference between these two studies could be the dif-
ferent motivation behind the two experimental situations:
in our study, cats were placed in a situation of uncertainty
where they could choose to use their owner as a guide to
action. In this situation, their initial response at least
appears to be similar to dogs in which both look at the
owner. It is not, however, clear what motivates this looking
behaviour, and one possibility is that cats sought comfort
from their owner. This interpretation, however, is not
supported by the cats’ behaviour: in fact, just 2 cats of the
19 looking referentially sought contact with the owner after
having looked at him/her. Another possible explanation is
the different environments in which the tests were carried
out: our study took place in a laboratory room, whilst
´si et al. (2005), tested cats in a room of the owner’s
flat. Finally, as put forward to explain dogs’ referential
looking in social referencing paradigms (Merola et al.
2012a,b), cats may have looked to the owner to coordinate/
synchronize their actions with theirs.
To best understand the potential significance of looking
behaviour in the second part of the study, we assessed
whether cats would take into account their owners’ reaction
to the fan and modulate their own actions accordingly.
Although there is some evidence that cats react differently
to unfamiliar versus familiar humans (Collard 1967; Casey
and Bradshaw 2008) and that they recognize their owners’
voices (Saito and Shinozuka 2013), no study has evaluated
cat’s behavioural reaction to human emotions so far.
Results provide some indication that cats could discrimi-
nate between their owner’s different reactions. In fact, both
the likelihood and frequency of gaze alternating between
the screen and the fan were higher in the cats exposed to
the owner showing a negative reaction to the object. As
described above, the screen was the only possible way out
and thus looking at the screen and then at the fan poten-
tially suggests the cats’ were worried about the fan and
wanted to get away from it. A further indication comes
from the fact that cats in the negative group showed a
tendency to start moving earlier than cats in the positive
group in Phase 2, potentially showing that they started
looking for an escape route sooner than cats in the group
where the owner was expressing a positive emotion.
These results show an influence of the owner’s emo-
tional expression on the cat’s behaviour, but they differ
from results of previous studies with dogs and infants
(Merola et al. 2012a,b; Hornik et al. 1987). In fact, both
infants and dogs when seeing their caregiver/owner
expressing a negative emotion spent more time being sta-
tic, whereas cats showed the opposite type of reaction, i.e. a
tendency to move earlier than cats whose owner had
expressed a positive emotion. This opposite reaction could
be explained by the different species-typical behaviour
where cats, being both a predator (Bradshaw 1992), like
dogs, but also a prey species, may be more inclined to use a
flight response when in a fearful situation.
Overall, cats in the negative group also showed a higher
frequency in their interaction with the owner than cats in
the positive group, potentially suggesting they were look-
ing for security from their owner. However, results as
regards the gaze alternation between screen and fan, and
the higher frequency of interacting with the owner shown
by cats in negative group, appeared across all phases, hence
Anim Cogn
including Phase 1, in which no emotional cues were
delivered as the owner had to remain silent with a neutral
expression. This was unexpected since we assigned the cats
randomly to the two groups. One possibility is that owners
in the negative group inadvertently carried out ‘negative’
behaviours already in Phase 1, and cats ‘picked up’ on
these subtle cues. Hence, in future studies, it may be
important to give instructions to owners only once the
initial ‘baseline’ phase is over or, as was carried out in a
previous study (Merola et al. 2012b), ask owners to
immediately deliver their emotional message, which in
dogs enhanced their response to the owner’s emotional
Finally, we wanted to assess whether cats would syn-
chronize their behaviour towards the object with that of
their owners. Results showed that when the owner started
to act towards the fan (and when they continued to do so
but the fan was turned off in Phase 4), cats in the positive
group were static for longer and alternated their gaze from
the fan to the owner more often than cats in the negative
group; they also tended to vocalize more often when the
fan was turned off. However, only three cats in these
phases approached the fan, two being in the positive and
one in the negative group. Hence, taken together, results
suggest that cats discriminated between the owner’s reac-
tions to the fan, but they did not adjust their distance from
the fan in accordance with their owner’s emotional
expression or behaviour.
The lack of synchrony between the cats’ behaviour and
that of their owners’ contrasts with results found with dogs
and could depend both on the procedural differences
between studies and the evolutionary history of the two
species. The fact that the owner did not start expressing the
assigned emotional expression from the start may have
negatively affected the likelihood of cats being influenced
by their owner’s behaviour, as we observed in dogs (Me-
rola et al. 2012a,b).
A second potential reason is that the level of fear
induced by the unfamiliar stimulus may have been signif-
icantly different in the two species. Indeed, in the current
study, only three subjects (11 %) touched the fan in the first
phase, whereas in the dog studies (Merola et al. 2012a,b,
2013a) 30 % of dogs touched it in this phase. Hence, the
level of fear may have inhibited the cat’s tendency to
mirror the owner’s reaction to the fan. Consistent with this,
the positive group had a tendency to exhibit more meowing
than cats in the negative group in the fourth phase (i.e.
when the fan was switched off) and showed more static
behaviour and gaze alternation between the fan and the
owner, as if waiting to choose how to act and potentially
communicating with the owner to obtain more information.
The ‘meow’ seems to be specifically associated with vocal
communication directed at people (Nicastro 2004); it is
among the most common cat-to-human vocalizations
(Bradshaw and Cameron-Beaumont 2000), and its increase
in intensity and frequency is usually related with a growing
level of tension in the subject (McCune 1994).
There may be also reasons for the lack of synchrony
with owners in cats compared with dogs, based on the
evolutionary history of the two species, which may affect
the likelihood of displaying social referencing. Whereas
cats’ ancestors were solitary, and today this species is
dubbed ‘optionally social’ (i.e. likely to choose whether to
establish strong relationships or simply tolerate social sit-
uations, Bradshaw 2013), dogs’ ancestor were pack-living
animals and dogs today have been shown to organize in
multi-male, multi-female packs when food is prevalent
(Cafazzo et al. 2010). It would hence seem that for indi-
viduals of social species, being able to take into account a
conspecific reaction to external objects and mirror their
behaviour may be more relevant than for individuals of a
more solitary one. Furthermore, during the course of
domestication, dogs have been selected for work with
humans, which potentially enhanced the likelihood of their
being willing to coordinating actions with people (Soproni
et al. 2001; Hare et al. 2002; Miklo
´si et al. 2003), whereas
cats have not undergone this selection and their utility for
humans has mostly been linked with independent hunting
of small rodents (Clutton-Brock 1988). Hence, although
more studies are needed to assess cat’s social referencing
tendency, the dog–cat species differences observed may be
related to the social structure and domestication history
which characterize them.
Acknowledgments This research was supported by a PhD and
postdoctoral grant from Milan University to Isabella Merola and
Sarah Marshall-Pescini. A special thanks to Emanuela Ponzone for
her invaluable help in data collection and scoring, and Paola Val-
secchi for insightful comments on the manuscript. Finally, we would
like to thank all the owners and cats that participated as volunteers.
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Anim Cogn
... Cats display the ability to discriminate human emotional expressions (Galvan & Vonk 2016) and human attentional states (Ito et al. 2016;. They also exhibit referential looking toward their owner in the presence of a potentially frightening object (Merola et al. 2015 Across all species, communication implies emitters and receivers. During a dyadic interaction, an emitter sends signals through one or more channels to a receiver (Beauchaud 2020). ...
... Despite their worldwide popularity, characteristics of cats' behaviour are poorly studied, and people's abilities to read this apparently 'inscrutable' species have attracted negligible research . Indeed, most research investigating human -cat communication has focused on cats' socio-cognitive abilities (Galvan & Vonk 2016, Ito et al. 2016, Merola et al. 2015, Miklósi et al. 2005 In a pilot study, Schötz pointed the limited ability of human participants to categorize cats' ...
... De 9,7 millions d'individus en 2001, ce nombre est passé à 15,1 millions en 2021 -soit plus de deux fois les 7,5 millions de chiens recensés(FACCO 2021). De façon paradoxale, jusqu'à présent, bien plus de recherches ont été menées sur le chien que sur le chat(Merola et al. 2015) et on recense, comparativement, encore peu d'études scientifiques explorant les comportements de ce félin qui vit à nos côtés. Une des causes de cette disparité tient certainement à la difficulté d'observer ses comportements en situation standardisée, le chat exprimant une majorité de comportements liés au stress lorsqu'il sort de son environnement quotidien(Nibblett et al. 2015). ...
Dans une société où les animaux compagnons sont intégrés au cercle familial, beaucoup d’humains les considèrent comme des membres de la famille à part entière. La recherche doit suivre cette tendance et s’attacher à appréhender les mécanismes de relations qui se construisent entre différentes espèces amenées à cohabiter. L’objectif de cette thèse est d’enrichir et d’approfondir les connaissances scientifiques sur l’éthologie du chat compagnon (Felis catus), afin de mieux appréhender ses besoins et réponses comportementales, au sein d’un environnement souvent imposé par l’humain. Les travaux restitués sont principalement centrés sur la communication interspécifique entre l’humain et le chat. Soucieux d’explorer aussi bien la perspective de l’humain que celle du chat, nous avons étudié la façon dont chacun s’exprime et décode les messages de l’autre. Ainsi, nous nous sommes intéressés à la communication vocale et visuelle entre ces deux espèces différentes qui partagent un même milieu - et doivent apprendre à communiquer efficacement pour cohabiter sereinement. Nos études ont mis en évidence que les humains utilisaient un discours spécifique pour s’adresser à leur compagnons félins, caractérisé par l’utilisation d’une voix plus aiguë. Nous avons également rapporté que les chats étaient plus attentifs à ce type de discours, mais seulement lorsqu’il était prononcé par leur compagnon humain et non par un étranger. Dans une troisième étude, nous avons observé que les chats venaient plus volontiers au contact d’un humain peu familier si celui-ci proposait un contact bimodal ou visuel, plutôt que vocal. Enfin, nous avons vu que les humains comprenaient mieux les chats dans leurs expressions bimodales et visuelles que vocales. Ainsi, bien que communément utilisée par chaque émetteur de cette communication interspécifique, la modalité vocale ne semble pas être suffisante pour la transmission et la réception d’un signal clair. Ces résultats sont discutés à la lumière des notions d’attachement, d’anthropomorphisme et de bien-être animal.
... Thus, despite domestic cats' relatively more recent domestication, we expect to discover similar proficiency in using human communicative cues given proper socialization. Recent research examining social cognition in cats provides some support, with cats displaying equivalent abilities to follow human pointing gestures as dogs (Miklosi, Pongracz, Lakatos, Topal, & Csanyi, 2005), as well as comparable levels of social referencing to owners in an uncertain situation (Merola, Lazzaroni, Marshall-Pescini, & Prato-Previde, 2015;Merola, Prato-Previde, & Marshall-Pescini, 2012a;Merola, Marshall-Pescini, & Prato-Previde, 2012b). Cats have also been successful at following dynamic human gaze both with ostensive and nonostensive cues (Pongrácz, Szapu, & Faragó, 2018). ...
... Although nearly all of the thirty-seven felid species are solitary as adults, tending to form strong territorial connections as opposed to bonds with conspecifics, freeranging domestic cats have been known to form colonies within which they display preferences toward particular associates (Bradshaw;Izawa & Doi, 1993;Macdonald, Yamaguchi, & Kerby, 2000). Further, just as domesticated dogs have shown a sensitivity to human communicative cues (Udell et al., 2010), domesticated cats have demonstrated similar abilities, in regards to following human pointing gestures (Miklosi et al., 2005), distinguishing between human voices (Saito & Shinozuka, 2013), engaging in social referencing in uncertain situations (Merola et al., 2015;Merola et al., 2012a;Merola et al., 2012b), and distinguishing between emotional states in their owners (Galvan & Vonk, 2016). Most relevant to the current study, domestic cats have been shown to learn via observation with observer cats learning more quickly to avoid an aversive stimulus than cats trained using shaping procedures (John, Chesler, Bartlett & Victor, 1968). ...
Full-text available
Although felids in general tend to be relatively asocial, domestic cats live closely with humans and other domestic species and thus, might be expected to respond to cues indicating, for example, the reputation of others. Furthermore, recent research with other less social species suggests that social learning is not unique to group-housed animals. Therefore, here we tested seven cat dyads with one cat interacting directly with unfamiliar humans, and another indirectly observing the interactions, to determine whether they would learn the ‘friendly’ and ‘aggressive’ reputations of the unfamiliar humans. Cats did not show a tendency to interact less, or more cautiously, with aggressive experimenters based on contact duration and latency to approach. Cats that observed the interactions indirectly spent more time near both experimenters and approached more quickly on test trials compared to cats that directly interacted with the experimenters, but this may have been due to spending more time crated between trials. We hesitate to conclude that cats are incapable of inferring reputation based on this small sample. It is possible that cats would behave more discriminately if tested in familiar environments.
... Cats and dogs are the most popular companion animals. They have fully adapted to the human social environment and are capable of establishing long-term social relationships with people [37]. The influence of the emotional connection on the hormonal interactions between humans and animals is the subject of much research. ...
... The social skills of domestic cats in the context of human-animal interactions have not been studied as thoroughly as for dogs [37]. Perhaps it is related to their shorter period of domestication and living with humans, as well as their higher sense of independence [10]. ...
Full-text available
Human–animal interactions and the emotional relationship of the owner with the pet are the subjects of many scientific studies and the constant interest of not only scientists but also pet owners. The aim of this study was to determine and compare the hair cortisol levels of dogs, cats, and their owners living in the same household. The owners were asked to complete a questionnaire concerning the frequency of their interactions with pets and emotional relationship with each of their cats and each of their dogs. The study involved 25 women who owned at least one dog and at least one cat. In total, 45 dogs and 55 cats from 25 households participated in the study. The average level of hair cortisol of the owners was 4.62 ng/mL, of the dogs 0.26 ng/mL, and in the hair of cats 0.45 ng/mL. There was no significant correlation between the hair cortisol level of the owner and dog or the owner and the cat and between dogs and cats living together. A significant positive correlation was observed between the hair cortisol level in the owner and the pet, for dogs in which the owner performs grooming treatments once a week and for cats which are never kissed. Although our study did not find many significant correlations, studies using other stress markers might have yielded different results.
... Consequently, cats' sociocognitive skills depend on both ontogenetic and evolutionary mechanisms. As pointed out by Jardat and Lansade (2022), these skills include: recognition of individual human features relying upon vocal cues (Saito and Shinozuka 2013), cross-modal and multimodal mental representations of owner (Takagi et al. 2019(Takagi et al. , 2021, perception of human emotions (Galvan and Vonk 2016;Quaranta et al. 2020), interpretation of humans' attentional state (Vitale and Udell 2019), interspecific communication (Miklósi et al. 2005;Miklósi and Soproni 2006;Humphrey et al. 2020), social referencing (Merola et al. 2015), and sensitivity to ostensive cues (Pongrácz et al. 2019;Pongrácz and Onofer 2020). Ostensive cues are signals given specifically to attract an auditor's attention and initiate an interaction (Jardat and Lansade 2022). ...
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In contemporary western cultures, most humans talk to their pet companions. Speech register addressed to companion animals shares common features with speech addressed to young children, which are distinct from the typical adult-directed speech (ADS). The way dogs respond to dog-directed speech (DDS) has raised scientists’ interest. In contrast, much less is known about how cats perceive and respond to cat-directed speech (CDS). The primary aim of this study was to evaluate whether cats are more responsive to CDS than ADS. Secondarily, we seek to examine if the cats’ responses to human vocal stimuli would differ when it was elicited by their owner or by a stranger. We performed playback experiments and tested a cohort of 16 companion cats in a habituation–dishabituation paradigm, which allows for the measurement of subjects’ reactions without extensive training. Here, we report new findings that cats can discriminate speech specifically addressed to them from speech addressed to adult humans, when sentences are uttered by their owners. When hearing sentences uttered by strangers, cats did not appear to discriminate between ADS and CDS. These findings bring a new dimension to the consideration of human–cat relationship, as they imply the development of a particular communication into human–cat dyads, that relies upon experience. We discuss these new findings in the light of recent literature investigating cats’ sociocognitive abilities and human–cat attachment. Our results highlight the importance of one-to-one relationships for cats, reinforcing recent literature regarding the ability for cats and humans to form strong bonds.
... For example, cats can use human pointing gestures to locate hidden food (Miklósi et al. 2005) and 78 follow the human gaze for referential information (Pongrácz et al. 2019). Cats also exhibit referential 79 looking toward their owner in the presence of a potentially frightening object (Merola et al. 2015 emitter sends signals through one or more channels to a receiver (Beauchaud 2020). The receiver 92 decodes these signals and is able to react accordingly, potentially turning into an emitter himself. ...
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Across all species, a dyadic interaction entails that an emitter sends signals to a receiver through one or more channels. The receiver decodes the signals and reacts accordingly, potentially turning into an emitter himself. Like dogs and horses, cats can integrate visual and auditory signals sent by humans and modulate their behaviour according to the valence of the emotion perceived. However, the specific patterns and channels governing cat-to-human communication are poorly understood. This study addresses whether, in an extraspecific interaction, cats adapt their communication channel to those used by their human interlocutor. We examined three types of interactions: vocal, visual and bimodal (visual plus vocal), through coding video clips of 12 cats living in cat cafés. There was a significant effect of the modality of communication on the latency for cats to approach the human experimenter. Cats interacted significantly faster in the visual and bimodal compared to the “no communication” pattern as well as to the vocal condition. There was a significant effect of communication modality on the tail wagging behaviour. Cats significantly displayed more tail wagging when the experimenter engaged no communication compared to the visual and bimodal conditions. Cats also displayed more tail wagging in the vocal compared to the bimodal condition. Taken together, these results suggest that cats display a marked preference for both visual and bimodal cues addressed by non-familiar humans, over vocal cues only. Our data bring further evidence for the emergence of human-compatible socio-cognitive skills in cats, that favour their adaptation to a human driven niche.
... New Caledonian crows could learn to perform complementary behaviors in a collaborative task but behaved similarly regardless of whether a partner was present or absent or whether rewards were equal, causing the researchers to question their understanding of the task and their ability to transfer causal knowledge from the physical to the social domain (Jelbert et al., 2015). However, organisms can benefit from eavesdropping the interactions of others (Abdai & Miklósi, 2016), can acknowledge when they need to teach skills to others (as in the case of mothers teaching their young to hunt), and can reference others for information about how to proceed in ambiguous situations through social referencing (even in asocial species like cats; Merola et al., 2015). ...
Cats are unique amongst domestic species in that they have evolved from a solitary ancestral species to become one of the most beloved household pets today. Interestingly the cat's physical appearance and sensory systems remain almost identical to their wild counterparts. Recognition of the perceptual parameters allows us to better understand how the domestic cat responds to environment and communicates with social partners. Sociality is unequivocally the aspect of feline life most affected by the domestication process. Cats can display a wide range of social behaviors, and evidence indicates that early exposure to a variety of social and environmental stimuli is the most important postnatal factor for a well‐adjusted life in a domestic setting and resiliency to basic stressors. By gaining an understanding of feline natural behavior, communication, learning, and cognition, shelter staff can provide cats with an ideal environment, change unwanted behaviors, and improve the welfare of our cats.
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Humans communicate with each other through language, which enables us talk about things beyond time and space. Do non-human animals learn to associate human speech with specific objects in everyday life? We examined whether cats matched familiar cats’ names and faces (Exp.1) and human family members’ names and faces (Exp.2). Cats were presented with a photo of the familiar cat’s face on a laptop monitor after hearing the same cat’s name or another cat’s name called by the subject cat’s owner (Exp.1) or an experimenter (Exp.2). Half of the trials were in a congruent condition where the name and face matched, and half were in an incongruent (mismatch) condition. Results of Exp.1 showed that household cats paid attention to the monitor for longer in the incongruent condition, suggesting an expectancy violation effect; however, café cats did not. In Exp.2, cats living in larger human families were found to look at the monitor for increasingly longer durations in the incongruent condition. Furthermore, this tendency was stronger among cats that had lived with their human family for a longer time, although we could not rule out an effect of age. This study provides evidence that cats link a companion's name and corresponding face without explicit training.
Research into cat behavior has gained more attention in recent years. As one of the world’s most popular companion animals, work in this field has potential to have wide-reaching benefits. Cats living in shelters are posed with distinct welfare concerns. Shelter cat welfare can be increased through use of environmental enrichment to promote natural behaviors. This review focuses on relevant literature published to date on shelter cat enrichment. Several key areas of research were identified. These included sensory enrichment, feeding enrichment, physical enrichment, social enrichment, and assessments to determine cat preference for enrichment stimuli. Existing studies have examined the efficacy of enrichment to promote species-specific behaviors and to reduce stress in shelter cats. Studies have also explored housing conditions for shelter cats such as cage size, communal housing, or the general quality of the environment. Applications of this information are discussed in order to promote natural cat behavior and find ways to increase the welfare of shelter cats. A review of the literature highlights the importance of supplying novel items in shelter environments, providing a rotation of individually preferred items, the use of human social interaction as a way to increase interactive behaviors in shelter cats, and the importance of considering potentially aversive impacts of enrichment under certain situations.
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The natural habitats of most species are far from static, forcing animals to adapt to continuously changing conditions. Perhaps the most efficient strategy addressing this challenge consists of obtaining and acting upon pertinent information from others through social learning. We discuss how animals transfer information via social channels and what are the benefits of such exchanges, playing out on different levels, from theperception of socially delivered information to emotional sharing, manifesting themselves across different taxa of increasing biological complexity. We also discuss how social learning is influenced by different factors including pertinence of information for survival, the complexity of the environment, sex, genetic relatedness, and most notably, the relationship between interacting partners. The results appear to form a consistent picture once we shift our focus from emotional contagion as a prerequisite for empathy onto the role of shared emotions in providing vital information about the environment. From this point of view, we can propose approaches that are the most promising for further investigation of complex social phenomena, including learning from others.
Ever since the discovery of fossil remains of extinct animals associated with flint implements, bones and other animal remains have been providing invaluable information to the archaeologist. In the last 20 years many archaeologists and zoologists have taken to studying such "archaeofaunal" remains, and the science of "zoo-archaeology" has come into being. What was the nature of the environment in which our ancestors lived? In which season were sites occupied? When did our earliest ancestors start to hunt big game, and how efficient were they as hunters? Were early humans responsible for the extinction of so many species of large mammals 10-20,000 years ago? When, where and why were certain animals first domesticated? When did milking and horse-riding begin? Did the Romans influence our eating habits? What were sanitary conditions like in medieval England? And could the terrible pestilence which afflicted the English in the seventh century AD have been plague? These are some of the questions dealt with in this book. The book also describes the nature and development of bones and teeth, and some of the methods used in zoo-archaeology.
With reference to Felis catus, discusses the organisation of behaviour and how this is orientated towards particular goals. Tinbergen's model of the hierarchy of instincts is useful in examining cat behaviour, and comparisons are made between domestic cats and wild felids. F. catus is probably a self-domesticated animal. -S.J.Yates