Journal of Feline Medicine and Surgery (2004) 6, 19–28

www.elsevier.com/locate/jfms

Social organization in the cat: a modern

understanding
Sharon L. Crowell-Davis*, Terry M. Curtis, Rebecca J. Knowles
Department of Anatomy and Radiology, College of Veterinary Medicine, University of Georgia, Athens,
GA 30602, USA
Revised 13 August 2003; accepted 25 September 2003

Summary An increasing body of research work has made it clear that, while Felis
catus can survive in the solitary state, social groups with an internal structure, are
formed whenever there are sufficient food resources to support them. Most people
who have cats have two or more cats. Failure to understand what will promote either
friendly or aggressive behavior can lead to various behavior problems, including
aggression and conflict over resources, such as food, resting sites and litterboxes. An
understanding of the natural social organization, relationships and communication
between cats is therefore essential, and is the subject of this paper.
© 2004 ESFM and AAFP. Published by Elsevier Ltd. All rights reserved.

Introduction to or even partially on them. Also within the colony,

certain cats fail to form strong affiliative relation-
In the past two and a half decades, an increasing ships with certain other cats, thus producing a
body of research conducted by various scientists socially complex society in which alliances and
throughout the world has made it clear that, while antipathies can affect access to resources, fre-
the feral and free-living domestic cat, Felis catus, quency of friendly and agonistic behavior and other
can survive in the solitary state when food re- issues that we are just beginning to understand.
sources are so widely distributed as to be unable to Most people who have cats have two or more
support a group, social groups that have internal cats. Failure to understand what will promote
structure, and in which group members recognize friendly, amicable behavior and what will promote
each other and engage in a variety of social behav- aggressive behavior can lead to various behavior
iors, are formed whenever there are sufficient problems, including aggression and conflict over
food resources to support a group (e.g. Dards, 1978, resources, such as food, resting sites and litter-
1983; Kerby and Macdonald, 1988; Macdonald, boxes. Thus, it is critical that we understand
1983; Macdonald and Apps, 1978; Macdonald et al., the natural social organization, relationships and
1987, 2000; Mirmovitch, 1995; Natoli, 1985a,b; communication of the cat.
Natoli and De Vito, 1991; Natoli et al., 2001;
Panaman, 1981; Sung, 1998; Wolfe, 2001; Yamane
et al., 1996). In other words, they are a social
species. Within the group, commonly called a col- The colony
ony, cats form affiliative, or friendly, relationships,
with certain other cats, grooming them, rubbing At its core, the colony is matrilineal, and it is the
them, greeting them, and sleeping curled up next affiliative, co-operative relationships between
females that provide the social structure upon
* Corresponding author. Tel.: +1-706-542-8343; fax: +1-706- which the colony is based (e.g. Liberg and Sandell,
542-0051 1988; Macdonald et al., 2000). When the process of
E-mail address: scrowell@vet.uga.edu (S.L. Crowell-Davis). domestication first began in areas where humans

1084-2756/04/$ - see front matter © 2004 ESFM and AAFP. Published by Elsevier Ltd. All rights reserved.
doi:10.1016/j.jfms.2003.09.013
20 S.L. Crowell-Davis et al.

were developing agriculture, there were concen-

trated food resources that could be effectively
defended by multiple cats. In this context, an ex-
tension of the mother–offspring relationship past
the weaning period would have been adaptive,
resulting in a queen and her adult offspring defend-
ing and monopolizing a valuable resource (Frank,
1998). The co-operative care of the kittens by a
queen and her female relatives, or other familiar
queens that exists today could readily have evolved
in this environment (Macdonald, 1983; Macdonald
and Carr, 1989). Today, food resources determine
colony size (Liberg et al., 2000). Large colonies
exist where food is abundant and small colonies
exist where food patches are still clumped, but less
abundant. Individual cats can survive in areas
where food is too widely dispersed to support a
colony, and it is in this context that we see the truly
solitary cat.
Figure 1 Two domestic cats greet each other with a
Relationships, social bonding and nose touch.
signaling within the colony
Cats recognize colony members vs. non-colony
members. Aggression is exhibited by most or all
colony members toward unfamiliar cats that are
not members of the colony. Thus, as is typical with
most social species, non-group members are not
allowed to casually approach and enter the group.
If non-colony members are persistent in attempts
to join the colony, they may eventually be inte-
grated into the group, but only by a gradual process
that involves many interactions (Macdonald et al.,
1987; Wolfe, 2001). Within the group, a number of
affiliative behaviors are exhibited, particularly be-
tween cats that are preferred associates. Preferred
associates are cats that can be found close together
(e.g. less than 1 m) more frequently than they are
found with other members of the colony. Preferred Figure 2 An allogrooming bout between three cats, a
associates can be found together in a variety of female and two of her adult offspring. Over the course of
contexts and locations: they do not simply go to several minutes, each cat groomed the other two cats.
preferred resources at the same time of day, but
come together because of the social bond that
exists between them (Wolfe, 2001).
In free-living and feral colonies of neutered or Nose-touch is a greeting behavior that is exhib-
intact cats, there is no effect of gender on which ited most commonly between preferred associates
cat approaches another, or which cats are pre- (Wolfe, 2001). There is no effect of gender: females
ferred associates if behavior during estrus of intact are equally likely to nose touch with females and
females is excluded (Sung, 1998; Wolfe, 2001). males, and males are equally likely to nose touch
Within a colony, some cats are close together less with females and males (Sung, 1998; Fig. 1).
often than is typical for the colony. In a colony of Allogrooming is a behavior in which one cat uses
intact cats, these pairs are disproportionately its tongue to groom another cat, usually on the
male–male pairs, while in a colony of neutered head and neck (Fig. 2). The recipient of the allo-
cats, there is no effect of gender (Wolfe, 2001). grooming is typically highly co-operative, tilting
This difference is probably due sexual competition and rotating its head to provide access to the
between certain males. groomer, and often purring. A cat may solicit
Social organization in the cat 21

Figure 3 The cat in the middle solicits allogrooming

from the cat on the right by lowering its head and flexing
its neck. The solicited cat responded by allogrooming the
head of the solicitor.

allogrooming by approaching another cat and flex-

ing its neck, exposing the dorsal surface and side
of the head to the cat being solicited (Fig. 3). Allo-
grooming is more frequent between preferred
associates than between non-preferred associates
(Wolfe, 2001). Allogrooming may or may not be
immediately reciprocated.
Colony members also allorub, a behavior in
which the cats rub up and down each other's sides
(Fig. 4a, b, c,). The head, sides and tail are all
involved in this behavior, which may go on for
several minutes. Like many cat behaviors, allor-
ubbing probably serves multiple functions. The
intense contact, particularly when the side of the
face is rubbed against the other cats face and body,
no doubt serves to facilitate exchange of scent.
There are probably tactile components to the be-
havior that are significant to the cats as well and
cats often purr during allorubbing. The existence of
allorubbing, combined with a high rate of sniffing
Figure 4 a, b, c. As part of an allogrooming sequence,
each other suggests that cats within a given colony two feral cats rub head to head, body to body and head to
develop a ‘colony odor’ that is maintained by the body.
exchanges of scent that occur during this behavior
(Bradshaw and Cameron-Beaumont, 2000).
Tail-up, in which the tail is held vertical to the Colony members of all ages will play with each
ground, signals friendly intentions upon approach. other, even in situations in which the cats are
Allorubbing is usually preceded by at least one cat chronically undernourished. The extended paw,
approaching with the tail-up, and is most likely to with claws retracted and no signaling of aggression
occur after mutual approach if both cats have the is a form of play-solicitation (Fig. 5). While play
tail-up (Cameron-Beaumont, 1997). Friendly cats continues into adulthood, it peaks at about 4 weeks
will also rub their tails against each other's bodies to 4 months of age, during which time social
and wrap their tails together so that the tails are relations among littermates are developed (West,
intertwined (Crowell-Davis, 2003). 1974).
22 S.L. Crowell-Davis et al.

related, e.g. a mother–daughter pair, kin-selection

would support aiding in the care of related kittens
(Hamilton, 1963; Trivers, 1971). Even if the queens
are not related, the situation is ideal for the
phenomenon of reciprocal altruism to function
effectively. In reciprocal altruism, one animal aids
another with the expectation that the aid will be
returned in kind. Reciprocal altruism is most likely
to function when the ‘returned favor’ can be done
closely in time and be of similar value (Hamilton,
1963). When queens give birth only a few days
or weeks apart, the conditions for facilitating
reciprocal altruism can be readily met.
Queens have been observed to engage in ‘mid-
wifing’ behavior, in which one is present during
parturition by another. The nonparturient queen
Figure 5 Two feral cats engage in play behavior, one will clean the perineum of the queen giving birth
extending its paw to the other in solicitation of play.
and will also clean the kittens and consume
the amniotic membrane. Queens engaging in co-
operative rearing of the young will groom, nurse
and guard each other's kittens. Non-nursing queens
have been observed to bring food to nursing queens
(Macdonald et al., 1987). When kittens are moved
from one nest site to another, kittens of queens
engaged in co-operative rearing spend less time
alone than kittens of a queen attempting to rear
them alone. Since moving of the nest and being
alone can be a particularly hazardous time for
kittens, which are subject to predation, having
multiple caregivers can clearly be advantageous
(Feldman, 1993). Kittens from communal nests also
leave the nest about 10 days earlier than kittens
from nests with single mothers, suggesting
that care by multiple queens facilitates speedy
development (Feldman, 1993).
Figure 6 Two feral farm cats that are preferred associ-
ates rest together, one laying its head on top of the Male–male
other.
While adult, intact male cats may engage in intense
aggressive conflict, particularly when in the pres-
ence of an estrous female, they do not necessarily
In addition to the active social behaviors de- do so. Intact, adult male cats may be preferred
scribed above, cats engage in the affiliative social associates, allogroom and allorub (Sung, 1998;
interaction of simply lying together in physical con- Wolfe, 2001). They may also remain non-aggressive
tact. One cat may use another as a ‘pillow’, with in the presence of an estrus female, and simply
the ‘pillow’ readily allowing the position (Fig. 6). alternate copulating with the female, who is poly-
This behavior occurs even in conditions of extreme androus and will readily mate with multiple males
heat, indicating that it occurs as a consequence of (Fig. 7).
social bonding, rather than for thermoregulation.
Female–male and the mating system
Female–female
Affiliative and contact behavior between females
Most notable in female–female relationships among and males is not exclusive to the breeding situation.
feral cats is the co-operative behavior exhibited Intact and neutered females and males may be
during rearing of kittens. Females that aid each preferred associates, engaging in a variety of affili-
other may or may not be related. If they were ative behaviors (Wolfe, 2001). When a female and
Social organization in the cat 23

Adult–kitten/juvenile
The critical role of the queen in teaching her kit-
tens hunting techniques has long been recognized.
Among free-living cats, the mother starts bringing
her kittens prey when they are about 4 weeks of age
(Baerends-van Roon and Baerends, 1979). At first
she brings them dead prey, and later they are
brought live prey. The mother will release the live
prey at the nest, providing the kittens with an
opportunity to develop their hunting and killing
techniques. In the early stages of this learning
opportunity, the queen will often demonstrate
hunting techniques to the kittens. Both kittens and
adult cats are excellent observational learners.
Figure 7 One male copulates with an estrous queen They can learn arbitrary tasks that are not species
while another waits nearby. There was no aggression typical behaviors simply by observing another cat
between these two males, or an additional two males engaging in the behavior (e.g. Chesler, 1969). This
that waited nearby. ability has likely been selected for because rapid
learning of critical hunting skills is essential to
survival. While the cat is socially gregarious, hunt-
ing is conducted in a solitary fashion as a conse-
quence of the typical prey of the cat. The majority
of the hunted diet of free-living cats is small
rodents, and it requires several small rodents a day
to sustain a single cat. Sharing the kill, such as
happens with species that hunt large game, is
impractical.
As the first cat with which the kitten experiences
affiliative social interactions, the queen is critical
to the learning of social behavior. This learning can
extend well beyond kittenhood. For example, in
group living cats, the highest rate of allogrooming is
observed among cats whose mother is present in
the group (Curtis et al., 2003). Kittens appear to
look to their mother for information about how to
interact with the world. They socialize to humans
Figure 8 A male grooms the ear of an estrous queen most readily if their mother is present during
between copulations. Photo courtesy of Prince Royal socialization and is calm in the presence of humans.
Bengals. Kittens do not socialize to humans as readily if their
mother is absent (Rodel, 1986).
Toms are commonly attributed with having no
male are familiar with each other, mating may involvement in rearing of kittens. However, a
involve substantial courtship behavior, including number of observations have contradicted this
allogrooming between the queen and the tom, lying idea. Intact toms have been observed to join
side by side, and rubbing of each other occurring queens in defending kittens from invading toms
between copulations (Fig. 8). Mating is polygamous. (Macdonald et al., 1987; Feldman, 1993), to groom
Females mate with multiple males and males mate kittens (Feldman, 1993), to share food with juve-
with multiple females. Yamane et al. (1996) found niles and to rest curled up around kittens that have
that, while the largest males had the greatest been abandoned at a colony site (Crowell-Davis et
mating success overall, males that were members al., 1997). Males have also been observed to disrupt
of a colony had the greatest mating success within intense wrestling play of juveniles, using a fore-
that colony, even if they were small. Thus, social limb to pull them apart but not engaging in any
attachments between males and females affect overt aggression against either (Curtis, personal
mating success of males. observation).
24 S.L. Crowell-Davis et al.

Importance of relatedness and animals against a dominant for the specific re-
familiarity source or, in the case of mating, female choice, can
contradict this expectation. Nevertheless, under-
As discussed above, females form the core of cat standing of dominance relationships among cats
society. Each female's extended family includes and how those relationships affect access to re-
children and grandchildren, all of whom have grown sources such as food, water, toys, resting sites and
up in close relationship with each other. These litterboxes is critical if we are to appropriately
family members typically exhibit friendly relation- manage multi-cat households.
ships with each other more frequently than with In an established group of cats, subordinate
other cats. Cats living with both relatives and non- status is acknowledged and dominance status is
relatives are more likely to be close to and allo- maintained primarily by a set of ritualized signals,
groom with a relative than a non-relative. Among rather than by overt fighting (e.g. Natoli and
non-relatives, they are more likely to be close to De Vito, 1991; Knowles, 2003). Upon encountering
and allogroom with a cat with whom they are more cats that are dominant to them, subordinate cats
familiar than a cat with which they are less familiar will exhibit such subtle behaviors as looking away,
(Curtis et al., 2001). In contrast, cats that have lowering the ears slightly, turning the head away
lived together longer, i.e. are more familiar and leaning back. In more intense encounters, the
with each other, are less likely to exhibit overt subordinate will flatten the ears against the head,
aggression (Barry and Crowell-Davis, 1999). lower and curl the tail lateral to the thigh, turn the
head to the side and crouch. In the most extreme
Dominance cases, the subordinate will roll over (Konecny,
If one individual consistently submits or gives way 1983; Feldman, 1994b; Bradshaw and Cameron-
to another individual as a consequence of prior Beaumont, 2000). Often, close encounters with
experience with that individual, the animal that dominant cats are simply avoided by giving way
submits is considered to be subordinate, while the spatially (Knowles, 2003). If the dominant cat is
animal submitted to is considered to be dominant walking down the same path as the subordinate or
in that dyadic relationship (e.g. Bernstein, 1981; toward the chair the subordinate is lying on, the
Immelman and Beer, 1989). The submission need subordinate will simply deviate off the path to
not always happen to consider the relationship to allow the dominant to pass, or jump off the chair
be asymmetrical, with one animal being dominant and move away. Dominant cats signal their status
and the other subordinate. The subordinate animal by another set of signals. They will approach a
must simply show submission to the dominant ani- subordinate, stare, stiffen the limbs, stiffen the
mal more frequently during agonistic interactions ears erectly upright while rotating them so the
than would be expected by chance. When a group aperture opens laterally, and elevate the base of
of animals live together in the same social group, a the tail while allowing the remainder of the tail to
set of dominant-subordinate relationships are es- droop (e.g. Overall, 1997; Fig. 9). Sometimes the
tablished such that we can construct a hierarchy, dominant cat will mount the subordinate, but this is
e.g. A is dominant to B and C, while B is dominant to not a common display. It is uncommon for a domi-
C. Truly linear hierarchies are rare in the animal nant cat to give a complete display. Instead, a mild,
kingdom, especially among groups of animals larger partial display is usually sufficient to induce a sub-
than four or five. In most mammalian groups, there ordinate cat to defer, e.g. stare at while stiffening
are ties and reversals within the group, making the ears.
the hierarchy nonlinear (Lehner, 1996). Cats are Socially dominant cats have priority of access to
no exception. While small groups of three or four food over subordinate cats (Knowles, 2003; Fig. 10)
cats often have a simple, linear hierarchy, larger However, as with other species, they do not always
groups are likely to have one or more ties and invoke this priority, and occasionally defer to a
reversals. subordinate, presumably because they are not par-
The major function of dominance is presumably ticularly hungry. Among males, dominance may or
to allow priority of access to preferred resources, may not result in priority of access to estrous
such as food, water, resting sites and mates. How- females (Natoli and De Vito, 1991). The variables
ever, it is not always the case that the dominant affecting this phenomenon are poorly understood,
animal in a group always has first and greatest and both overt and cryptic female choice may be an
access to these resources (e.g. Natoli and De Vito, important factor that overrides male–male compe-
1991). Other variables such as motivation to obtain tition in determining which male fathers a given
the resource, coalitions by multiple subordinate females kittens (e.g. Eberhard, 1996).
Social organization in the cat 25

Figure 9 Two cats stare at each other briefly before the

cat on the right defers to the one the left by breaking eye
contact and moving away.

Cats are unique individuals and each group of

cats is a unique product of the combination and
interaction of the unique personalities within the
group. As with other species, some cats may be
high-ranking, but not make an issue of it unless
another cat confronts them over a resource they
happen to want at the moment. A household in
which the highest ranking cat or cats are like this is
likely to be relatively peaceful, as subordinate cats Figure 10 A subordinate cat waits while a higher-
ranking cat eats.
are able to obtain resources so long as they are
alert to the need to defer if they should encounter
a higher-ranking cat. Other high-ranking cats may
routinely move through the group, threatening Sounds made with the mouth closed include the
multiple individuals, especially those close to them purr and the trill. The purr is a friendly greeting and
in rank, and confiscate resources that they do not care-soliciting call that typically occurs during ami-
even appear to desire at the moment. Having such cable social interactions and when ill or injured.
‘bully’ cats in the household is likely to lead to The trill is a greeting call.
problems of serious intercat aggression and second- Sounds made with the mouth open and gradually
ary behavior problems that are a consequence of closing include a large variety of miaows. Miaows
subordinates being kept away from important re- are amicable greeting calls, uttered in a variety of
sources, such as litterboxes. The variables that situations of interaction with other cats, dogs and
produce bullies are not well understood, but poor humans.
early socialization may be an important factor in Sounds made with the mouth held open in a
some cases. relatively constant position are usually related to
aggression. These include the growl, yowl, snarl,
Auditory and olfactory communication hiss, spit and shriek.
Olfactory communication occurs via a variety
Cats are one of the most vocal carnivore species. of sebaceous glands located throughout the body,
The exact number of different vocalizations they particularly on the head, in the perianal area, and
have is subject to interpretation, depending on between the digits. Urine and feces are also used in
how much a given classifier wishes to subdivide olfactory communication.
broad categories. There are three major categories The temporal glands located in the temporal
of vocalization (Moelk, 1944; Kiley-Worthington, region, the submental gland under the chin and the
1984). circumoral glands around the lips are all rubbed
26 S.L. Crowell-Davis et al.

against objects in the environment and against Implications for management and care
fellow colony members. This behavior is often ac- of cats
companied by purring. While there is no scientific
documentation of the specific molecules that are Failure of intraspecies socialization
deposited during these behaviors, or their exact
function, interpolating from the cats' behavior, the Social species are born with the capacity to learn
secretions appear to be related to (1) depositing a species-specific social skills, but they are not born
scent within a core area that labels the area and with the specific skills. In cats, as with humans,
(2) depositing a scent on a familiar conspecific dogs, horses and other social species, appropriate
with which the rubber has an amicable social experience with their own species is critical to the
relationship. development of appropriate species-typical social
While a lot is written about urine marking, behavior. Cats that are adopted as kittens and
primarily because people find it unacceptable subsequently kept in a one-cat household for sev-
when it occurs in the house, actual data from field eral months or years miss important learning and
situations is often conflicting regarding the appar- social bonding experiences that happen during late
ent function of the behavior (Feldman, 1994a; kittenhood and the juvenile period. While the
Gorman and Trowbridge, 1989; Natoli, 1985b; species as a whole is not asocial, such individual
Passanisi and Macdonald, 1990; Verberne and de cats may be, exhibiting a dysfunctional lack of
Boer, 1976). For example, while urine marking is knowledge of how to interact appropriately with
commonly credited with being a form of territorial their own species. If later attempts are made to
marking, Feldman (1994a) recorded deposition of introduce another cat, such asocial individuals are
urine and feces and found no evidence that either likely to exhibit uninhibited aggression or excessive
was used as a territorial marker. Indeed, while fear of the newcomer, fail to recognize species-
cats are often referred to as territorial, there is specific signals of greeting, dominance or submis-
no good evidence that they actually are terri- sion and fail to respond in a species-appropriate
torial, i.e. defend a piece of property. Urine manner.
marking occurs in a variety of contexts and doubt-
less has a variety of functions that probably Rejection of strangers and selection of cats
include:
for multi-cat households
• Communicating specific identifying infor-
mation, e.g. ‘intact female’ As discussed above, cat societies are fairly cohe-
• Communicating location information, e.g. ‘I sive. Friendly behavior toward familiar and related
entered this field X hours ago’ colony members coupled with agonistic behavior
• Communicating emotional information, e.g. ‘I toward unfamiliar, non-colony members is a gen-
am here and I am highly aroused’ eral rule that applies to most group-living, social
species, e.g. horses, wolves. If someone owns an
Attempting to attribute a single function or mes- established group of cats and abruptly brings in a
sage to urine marking is somewhat like attempting stranger, the situation is analogous to a govern-
to attribute a single message to all human speech. ment representative suddenly showing up at the
Substantial further study is required to truly door with a total stranger and expecting a human
understand this behavior. family to readily accept the stranger, i.e. share
their bed, food, bathroom and living space with
Feral cats have been observed to leave feces them. Given this similarity, it should not surprise us
unburied in the peripheral areas of their home that cats do not welcome the abrupt addition of a
range, but bury the feces in the core areas stranger any more than we would. This is an import-
of their home range (Macdonald et al., 1987; ant consideration for the pet-owner who wishes to
Feldman, 1994a). While it is possible that surface have multiple cats or who wishes to bring a strange
feces serve as a signal to strange cats entering the cat into an established group.
home range, there are no reports of strange cats First, instead of periodically adopting one cat,
departing from an area immediately upon encoun- pet-owners might have households with a greater
ter with feces of a resident cat. It may be the case rate of friendly behavior if they adopt two or three
that feces are typically buried in the core area for related cats at broad intervals of time, e.g. a
control of odors and parasites, but that the effort mother and two of her kittens.
of this behavior is not necessary in peripheral Second, when introducing a strange cat to an
areas. existing group, some degree of familiarity must be
Social organization in the cat 27

established before the cats are allowed to directly emphasis on the development of prey-catching. Verhaelin-
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