ANIMAL BEHAVIOUR, 2007, 74, 1369e1379
doi:10.1016/j.anbehav.2007.02.029
Feeding-order in an urban feral domestic cat colony:
relationship to dominance rank, sex and age
ROBERTO B ONANNI* , SIM O NA CA FA Z ZO* , CLA UDIO F ANTINI†, DOMI NI QUE PO NTI E R‡ & EUG EN IA N AT OLI†
*Dipartimento di Biologia Evolutiva e Funzionale, Università di Parma
yAzienda USL Roma D, Area Dipartimentale Sanità Pubblica Veterinaria
zUMR-CNRS 5558 ‘‘Biométrie and Biologie Evolutive’’, Université de Lyon
(Received 20 May 2006; initial acceptance 12 July 2006;
final acceptance 28 February 2007; published online 24 September 2007; MS. number: 8964R)
In social species, dominance relationships and access to food resources are often affected by asymmetries
in resource-holding potential (RHP) between competitors of different ageesex classes with males usually
being dominant and feeding first, followed by females and then juveniles. In this study we investigated
how variables such as sex and age affected dominance rank and feeding order in a social group of feral domestic cats, Felis silvestris catus, a sexually dimorphic species in which males are larger than females and do
not take part in parental care. Intersexual dominance relationships varied depending on the competitive
context: males occupied top rank positions away from food, whereas females increased in rank at the expense of males in a feeding context. Around the age of 4e6 months, kittens were significantly more likely
than adults of both sexes to be the first to feed, indicating that they received a certain level of tolerance.
These results provide support for game-theory models predicting conflict outcome in favour of the smaller
competitor when asymmetries in both the value of winning and in the cost of winning inappropriately
may compensate for the smaller competitor’s lower RHP. It is suggested that the results are not an artifact
of domestication: unlike male lions, Panthera leo, which usually dominate both females and cubs at kills,
male domestic cats may value the food less than females and juveniles, because they do not need to maintain constantly a peak physical condition to defend a group of females and protect offspring from
infanticide.
Ó 2007 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.
Keywords: feeding order; Felis silvestris catus; feral domestic cat; game theoretical approach; intersexual dominance;
leverage; social cats; tolerance to juveniles
Social dominance has historically been defined as priority
of access to resources (Wilson 1975). Since food is considered a major determinant of the reproductive success of
individuals, several authors have tested the influence of
dominance rank on access to food and it has been verified
Correspondence and present address: R. Bonanni, Dipartimento di
Biologia Evolutiva e Funzionale, Università di Parma, Parco Area delle
Scienze 11/A, 43100 Parma, Italy (email: [email protected]).
C. Fantini is at the Azienda USL Roma D, Area Dipartimentale Sanità
Pubblica Veterinaria, Direzione, via Portuense 1397, 00050 Ponte Galeria (Roma), Italia. D. Pontier is at the UMR-CNRS 5558 ‘‘Biométrie
and Biologie Evolutive’’, Université de Lyon, Université Lyon 1, 43 Bd
du 11 nevembre 1918, 69622 Villeurbanne cedex, France. E. Natoli
is at the Azienda USL Roma D, Area Dipartimentale Sanità Pubblica
Veterinaria, Ospedale Veterinario, via della Magliana 856, 00148
Roma, Italy.
0003e 3472/07/$30.00/0
that higher ranking members of a social group enjoy priority of access to food over subordinates in an array of animal species (e.g. red deer, Cervus elaphus, Appleby 1980;
rhesus monkeys, Macaca mulatta, Deutsch & Lee 1991; olive baboons, Papio anubis, Barton & Whiten 1993; chimpanzees, Pan troglodytes, Wittig & Boesch 2003; brown
bears, Ursus arctos, Gende & Quinn 2004).
Patterns of dominance relationships and food access
can be expected to follow predictions of classic gametheory models, with contest outcomes largely affected by
asymmetries in both resource-holding potential (RHP),
which is a measure of fighting ability, and relative resource
value between contestants (Parker & Rubenstein 1981;
Hammerstein & Parker 1982; Enquist & Leimar 1987). Precisely, competitors should be more likely to escalate and
eventually win conflicts, the higher their own RHP and
the higher the value of the resource to them.
1369
Ó 2007 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.
1370
ANIMAL BEHAVIOUR, 74, 5
In many species, body weight (and/or size) is a good
indicator of a competitor’s RHP (Parker 1974; Archer
1988), and where asymmetries in body weight exist between competitors belonging to different ageesex classes
in a social group, these usually predict conflict outcome
over food resources: a well-known example is that of the
lion, Panthera leo, in which males are the largest members
of the group and dominate both females and juveniles in
feeding competition situations (Van Orsdol 1986; Packer
et al. 2001). In species where females invest substantially
more energy than males in parental care, food may be
worth more to them than to males. In some mammalian
species showing a lack of sexual size dimorphism, such
as the spotted hyaena, Crocuta crocuta, and several Lemuriformes, females are dominant over males at feeding sites
(Roeder & Fornasieri 1995; Digby & Kahlenberg 2002; East
& Hofer 2002; Ostner et al. 2003; Pochron et al. 2003),
possibly due to higher relative food value.
Moreover, there are a number of conditions under
which contest outcome may be predicted in favour of
the smaller competitor. Hand (1986), following Parker &
Rubenstein (1981), pointed out that, while the fitness
costs in terms of injuries sustained increase at rates inversely correlated with RHP during a contest, there could
be an additional fitness cost arising from winning a conflict when it is inappropriate from the perspective of
fitness to do so, that may, under some conditions, be
higher for the larger competitor. This may occur in conflicts between opponents which share some individual
fitness interests, such as conflicts between mates or parenteoffspring conflicts. For instance, a male could reduce
directly his fitness by driving the mother of his offspring
or the offspring themselves away from critical food resources because, in both cases, he would probably decrease
offspring survival. Since both the female and the offspring, which usually are smaller than the male, are likely
to have less fitness to lose from winning than does the
male, they may have a ‘leverage’ advantage over the
male. If such asymmetry in the cost of winning equalizes
the total costs for both opponents, conflict outcome will
probably be decided by asymmetries in resource value
(Hand 1986). Since both the value of the resource and
the cost of winning (called ‘leverage cost’) may change
considerably depending on the competitive context, the
dominance relationship in a given dyad of individuals
may also vary according to the context (Hand 1986; see
Noë et al. 1980 for an empirical example). This model
may help to explain the cases of male food deference to
females during periods corresponding to egg laying or oestrus (e.g. Western Gulls, Larus spp., Hand 1986; chimpanzees, Stopka et al. 2001), when the food is of particularly
high value to females, and also why adult individuals of
both sexes allow juveniles’ feeding priority in a variety
of species (e.g. wild dogs, Lycaon pictus, Malcom & Marten
1982; spotted hyenas, Frank 1986; several primates,
Hrdy 1981).
In this paper we examine how variables such as sex and
age affect dominance and access to food in a social group
of feral domestic cats, Felis silvestris catus, living in an urban environment, and test whether the pattern found follow the predictions of evolutionary game-theory models.
In urban areas, domestic cats live at very high densities
(up to 2000 cats/km2) and form multimaleemultifemale
social groups subsisting on an abundant, patchily distributed, source of food provided by ‘cat lovers’ at a few traditional feeding sites (Izawa et al. 1982; Natoli 1985; Liberg
et al. 2000). Such groups provide an ideal situation to
study social factors affecting feeding competition because
several individuals of all ageesex classes usually gather at
the places where food is distributed by human beings. It
has been suggested that domestication may have fostered
the evolution of sociality in cats through an increase in
behavioural plasticity and social tolerance of conspecifics
(Macdonald et al. 2000). However, the cat’s history as a domesticated animal is very short, about 4000 years, in evolutionary terms (Serpell 2000), and many observed
behavioural patterns may have evolved in their solitary
ancestor, the African wild cat, Felis silvestris lybica, before
domestication and urbanization.
Whether a linear dominance hierarchy exists among
domestic cats is controversial probably because, as a result
of missing interactions between some individuals, it is not
always easy to reach a statistically significant level that
prove the occurrence of linearity (see for example Natoli &
De Vito 1991). Recently, Natoli et al. (2001) found a significantly linear dominance hierarchy in a rural cat colony,
based on the outcome of agonistic encounters in the absence of any sources of competition, in which a female
dominated some males. The last could be surprising because, as the lion, the domestic cat is a sexually dimorphic
species in which males are 15e40% heavier (Pontier et al.
1995, 1998) and possess 19% longer canines (D. Pontier,
unpublished data) than females. Possibly, in domestic
cats female dominance is achieved by other mechanisms
than physical strength and fighting ability (Natoli et al.
2001). To improve our comprehension of such mechanisms it seems useful to investigate dominance relationships among cats in feeding competition. Since food is
thought to be a valuable resource for female cats, because
male cats do not take part in parental care (Deag et al.
2000), it can be expected that females would gain further
rank positions in a feeding context, at least during
pregnancy and lactation, where the value of winning
the resource, together with the possible higher costs to
males of winning inappropriately, may offset asymmetries
in RHP.
Although male feeding deference to juveniles is commoner in species in which males provide substantial
parental care (Hand 1986), it could be expected that, if
male domestic cats allow female dominance due to the
high cost of winning a feeding conflict against the mother
of their offspring, they would also display a certain degree
of feeding tolerance to kittens.
Here, we ascertained whether a dominance hierarchy
based on outcome of agonistic encounters in the absence
of any sources of competition (food or receptive females)
exists in an urban cat group. Then, we examined the
dominance relations in the feeding context, and compared the rank found (feeding rank henceforth) with that
obtained in the absence of food. At the same time, we
recorded which cats were the first to feed among all those
present at the feeding station (feeding order). Finally, we
BONANNI ET AL.: FEEDING ORDER IN DOMESTIC CATS
sought to answer the following questions: is high social
rank associated with priority of access to food? Which sex
is dominant? Are kittens tolerated by adults in competitive feeding situations?
METHODS
Study Area
The study was carried out in Rome, in a popular quarter
called ‘Garbatella’. The cat colony lived in a large courtyard (about 6000 m2) owned by the Istituto Autonomo
Case Popolari of Rome. It was bounded by a wall and completely isolated from road traffic. The courtyard contained
11 buildings, open areas, flower-beds cultivated by inhabitants, trees and bushes. Cats had free access to every part
of the courtyard but they established their ‘core area’ in
a partially wire-fenced sector located in the south side of
the courtyard. Here, a lot of spontaneous vegetation offered good shelter for animals, especially for lactating females with kittens. The experimental feeding site was
placed in a small field (about 150 m2), adjacent to the
core area, in which spontaneous vegetation was periodically removed and where cats could be easily observed
while feeding without any disturbance. This was the
most important and regular food source for the colony.
Cats were rarely seen preying on small rodents, passerine
birds, feral pigeons, Columba domestica, and insects. Water
was available from a fountain just outside the entrance of
the courtyard.
Cat Group
The cat group living in the study area had no constraints placed on breeding or movements by human
owners and very few individuals were tame enough to be
handled.
We visually recognized all individuals by their coat
colour pattern and hair length. The sex of adult cats could
be determined on the basis of some morphological (presence of testes in males) and behavioural characteristics.
Mature males have a wider face than females; moreover
they show posterior urine spraying which is a behaviour
rarely displayed by females.
Age of the subjects was obtained from cat lovers or
estimated according to Pascal & Castanet (1978) method.
Cats were considered to be juveniles up to the age of 11
months and to be adults afterwards.
Throughout the period of study the number of individuals ranged from a minimum of 10 to a maximum of 17
cats. In September 2001, the cat colony consisted of eight
intact adult males, three intact adult females, two adult
females that had been neutered long before the study
started, and four kittens (all of them were the offspring of
the same female, LAV). Subsequently, two adult males
dispersed from the study area to settle in two different
neighbouring colonies, while five other individuals were
killed by cars.
Nine adult cats (five males and four females) were
captured during the study period: one female was trapped
using a double-door trap; the remaining individuals were
shot with anesthetizing darts by mean of a blow-pipe,
since they were too elusive to be trapped. Cats were
anaesthetized with an intramuscular injection of ketamine
chlorhydrate (Inoketam 1000, 5 mg/kg, Virbac, Milan, Italy)
and medetomidine (Domitor, 0.005 mg/kg, Pfizer, Rome,
Italy). For all captured animals the following measures
were recorded: body weight, head volume, length of the
right arm, and average length of canines. The same measurements, except head volume, were taken for a further
adult female after she was found dead. For males, testicles’
volume was also recorded, although it was not analysed
here because of the low number of captured males.
Experimental Procedure
We used a focal animal sampling technique (Altmann
1974) to study the social behaviour of 13 adult cats. A total
of 555.63 h of observation were made between September
2001 and June 2002. Each individual’s observations were
equally distributed over the time period, as well as over
daytime between 0900 and 1800 hours.
Agonistic behaviour (including aggressive and submissive behaviour) in absence of any sources of competition
(food, receptive females and shelters) was recorded by ‘all
occurrences’ method (Altmann 1974). Aggressive behaviour included the following: threats (striking with a paw,
biting, assuming threatening postures, pointing, staring
at, baring of the canines), chasing, ritualized vocal duels
and real duels. Submissive behaviour included: crouching
with the ears flattened, avoiding, retreating, fleeing and
hissing at. Kittens were not included in this data collection because, in absence of food, they were expected to interact agonistically too rarely with adults. The individual
scores of all behaviour patterns were corrected for animal
observation time because this varied between individuals.
In order to determine the dominance hierarchy, the
distributions of dyadic aggressive and submissive interactions were ranked in two different squared matrices with
performers on one axis and recipients on the other, so as
to minimize the number of dominance reversals. The
dominant animal of each dyad was the one that performed
more aggressions than it received, or received more submissions than it performed. Hissing was excluded from the
matrix based on submissive interactions because it was
considered an ambiguous behaviour, with components
both of aggressiveness and subordination. We tested the
transitivity of dominance relationships between the members of the social group, based either on aggressive or on
submissive behavioural patterns separately, by applying
an improved test of linearity developed by de Vries (1995)
that is based on Landau’s linearity index, but takes into account unknown and tied relationships between group
members.
Experimental Feeding Sessions
We performed a total of 174 experimental feeding
sessions from August 2001 to June 2002, corresponding
to 178.75 h of observation. Food was delivered to cats at
1371
1372
ANIMAL BEHAVIOUR, 74, 5
and the Improved linearity tests using MATMAN (Noldus
Noldus Information Technology bv, Wageningen, The
Netherlands). All statistical tests are two tailed.
0930 or at 1430 hours on a rotational daily basis. Each session started just after the food container was placed and
continued until the food was completely consumed. We assembled a plastic container on a baked-clay support and
filled it with 400 g of cat food. This structure had a circular
hole (10 cm in diameter) on the upper surface that prevented more than one cat from feeding at the same time.
The focal subgroup sampling technique (Altmann 1974)
was applied to record cat behaviour while feeding: we observed all individuals present within 4 m of the food, including adults and kittens. Since subgroup composition
changed during a session, we recorded the sequence in
which cats arrived at the feeding site and left it.
To determine the feeding order we estimated, for each
cat, the likelihood of being the first individual to eat of all
those that were present around the food at the beginning
of each session, by calculating the percentage of sessions
in which it was the first to feed, in the total number of its
presences around the food, and verified whether such
percentage varied throughout the study period.
Agonistic behaviour within 1 m of food was recorded by
All Occurrences method (Altmann 1974). This included the
aggressive and submissive patterns described above, plus
‘interruption of feeding after receiving an aggression’ that
was a submissive behaviour. Individual scores were corrected for presence scores around the food (within 4 m).
In order to determine the dominance hierarchy within
1 m of food (feeding rank), we constructed two different
matrices for aggressions and submissions, respectively,
and applied the same methods described above. Where
there were no data on outcomes of submissive interactions
over food by means of which to rank individuals, they have
been ordered according to the outcomes of aggressive interactions over food; similarly, where there were no data on
outcomes of aggressive interactions over food by mean of
which to rank individuals, they have been ordered according to the outcomes of submissive interactions over food.
One remaining ambiguity within a specific dyad was resolved by ordering the cats according to agonistic interactions observed between 1 and 4 m of food.
We performed nonparametric statistical tests using
STATISTICA 99 edition (StatSoft Inc., Tulsa, OK, U.S.A.)
Interobserver Reliability
Interobserver reliability between two of us was determined considering the proportion of agreements on total
frequencies of behaviour (Caro et al. 1979). Average concordance between observers, calculated over seven feeding
sessions conducted across the study time period, was 0.93
for submissions and 0.77 for aggressions (corrected by
chance as indicated in Martin & Bateson 1993). Average
concordance for autogrooming behaviour, on a series of
eight focal animals, was 0.97 (corrected by chance). Autogrooming was considered because it was displayed
frequently by each subject. Spearman correlation coefficient between observers, over the same series of focal animals, was rS ¼ 1.
RESULTS
Dominance Hierarchy in Absence of Any
Sources of Competition
The matrix based on aggressive interactions (N ¼ 300)
recorded between adult cats showed a lack of transitivity
(improved linearity test: h0 ¼ 0.387, P ¼ 0.064), probably
due to missing values. However, a significantly linear
dominance hierarchy based on direction of submissive behaviour except hissing (N ¼ 360 interactions) was found
(Improved linearity test: h0 ¼ 0.448, P ¼ 0.026; Table 1).
The top positions of this hierarchy were occupied by three
adult males, whereas three females were at the bottom
(Table 1).
This rank order based on submissions was positively
correlated with aggressive behaviour (rS ¼ 0.709, N ¼ 13,
P ¼ 0.007). In other words, the higher cats were in rank,
more aggressive they were towards other cats. Adult males
and females did not differ significantly in aggressive behaviour (ManneWhitney U test: U ¼ 12, N1 ¼ 5, N2 ¼ 8,
Table 1. Submissive interactions (except hissing) in absence of any sources of competition
Cats
LEO
ANT
SON
LAV
PIC
PAL
PEL
RIG
GNO
RED
CLA
FIA
SIL
Totr
LEO
ANT
SON
LAV
PIC
1
2
1
PAL
PEL
RIG
GNO
RED
CLA
FIA
SIL
1
1
2
2
5
3
2
15
3
8
6
5
69
2
42
19
1
3
158
1
1
1
1
23
1
4
2
1
35
1
1
1
1
1
2
1
2
6
7
5
1
5
10
6
37
14
2
1
1
1
4
2
1
5
14
1
1
Bold type: females; standard type: males.
Totp ¼ total submissions performed, Totr ¼ total submissions received.
2
6
1
6
4
20
1
2
2
2
2
10
17
21
11
4
35
9
4
1
4
1
Totp
1
5
5
12
11
11
2
109
22
58
52
41
31
BONANNI ET AL.: FEEDING ORDER IN DOMESTIC CATS
rS ¼ 0.726, N ¼ 8, P < 0.05), but no relation existed between age and rank considering all adult cats as a whole
(rS ¼ 0.168, N ¼ 13, P ¼ 0.583).
6
Body weight (kg)
5
Dominance Hierarchy in the Feeding Context
4
3
2
10
9
8
7
6
5
4
3
2
1
Rank
Figure 1. Correlation between dominance rank and body weight for
10 adult cats of the studied group. C ¼ males; B ¼ females.
P ¼ 0.240), although males scored the highest values
(mean ranks: eight for males, 5.4 for females).
In this group individual differences in body size seemed
to affect the dominance relationships. Body weight was
positively correlated with rank (rS ¼ 0.693, N ¼ 10, P <
0.03; Fig. 1): on average, males were 50% heavier than females and the three top-ranking males were the heaviest
cats in the group; similarly, the highest-ranking female
was also the heaviest female. Head volume was also positively correlated with rank (rS ¼ 0.783, N ¼ 9, P ¼ 0.013),
whereas for arm length and canines length the correlations with rank failed to reach a statistically significant
level (rS ¼ 0.467, N ¼ 10, P ¼ 0.174; rS ¼ 0.358, N ¼ 10,
P ¼ 0.310; respectively).
Age affected dominance among males, the oldest
males being the highest in rank (correlation ageerank:
Contrary to what we found in the absence of food, the
matrix based on aggressive interactions recorded within
1 m of the food (Table 2) showed a highly significant level
of linearity (improved linearity test: h0 ¼ 0.567, P ¼
0.0005), probably due to a higher number of interactions
(N ¼ 565). A female (LAV) was at the top of this hierarchy
and, altogether, all females became dominant over some
males from whom they were dominated in absence of
food (see Table 2). All kittens were at the bottom positions
of the rank order (Table 2). It should be noted that three
adult cats (two males and one neutered female) were not
considered in these analyses because they visited too
rarely the feeding site before their disappearance from
the study area.
Females showed the highest total level of aggressive
behaviour within 1 m from the food (mean rank ¼ 11),
followed by males (mean rank ¼ 7) and by kittens (mean
rank ¼ 4.75); nevertheless, the difference between the
three ageesex classes was not significant (KruskaleWallis
test: H2 ¼ 4.614, P ¼ 0.10). Females were still the most aggressive class even restricting the analysis to periods when
they were neither pregnant nor lactating and, indeed, this
time the difference was significant (KruskaleWallis test:
H2 ¼ 7.831, P ¼ 0.02; mean ranks: females 12.25, kittens
5.75, males 5.5).
A significantly linear hierarchy based on 307 submissive
interactions (except hissing) recorded within 1 m of the
food was also found (improved linearity test: h0 ¼ 0.464,
P < 0.009; Table 3). This feeding rank based on submission
patterns was very similar to the previous one, based on aggressions, although a male occupied the first position
(here LAV was ranked as the second cat in the hierarchy;
see Table 3). This feeding rank based on submissions was
just moderately correlated with rank (based on submissions)
Table 2. Aggressive interactions within 1 m from the food
Cats
LAV
ANT
LEO
PIC
CLA
PEL
FIA
SON
RIG
RED
CAL
NER
SPO
RUG
Totr
LAV
ANT
LEO
PIC
CLA
PEL
FIA
SON
RIG
RED
CAL
NER
SPO
RUG
Totp
4
1
1
15
1
3
42
4
6
15
14
71
110
2
7
2
11
2
13
3
1
1
2
1
1
3
6
16
3
2
3
4
4
1
7
3
7
9
1
31
19
299
6
25
3
59
5
85
44
3
3
13
2
17
1
2
2
4
4
2
4
1
3
1
1
1
13
1
1
1
1
4
1
26
1
141
5
1
13
1
5
1
3
17
9
66
Bold type: females; standard type: males; italic type: juveniles.
Totp ¼ total aggressions performed, Totr ¼ total aggressions received.
37
16
16
20
17
1
1
4
15
34
181
1373
ANIMAL BEHAVIOUR, 74, 5
Table 3. Submissive interactions (except hissing) within 1 m from the food
Cats
ANT
LAV
LEO
PIC
CLA
PEL
FIA
SON
RIG
RED
CAL
NER
SPO
RUG
Totr
ANT
LAV
LEO
PIC
CLA
PEL
FIA
SON
RIG
RED
CAL
NER
SPO
RUG
1
2
4
1
2
2
2
8
8
34
2
4
1
13
9
36
61
169
2
1
2
12
5
2
2
1
3
1
1
1
2
6
15
7
8
43
3
2
2
1
1
1
6
1
1
5
2
2
6
14
27
5
5
18
4
3
2
9
1
1
2
1
7
7
0
Totp
1
7
1
1
10
2
54
9
17
7
17
16
61
104
1
1
0
Bold type: females; standard type: males; italic type: juveniles.
Totp ¼ total submissions performed, Totr ¼ total submissions received.
found in absence of any sources of competition (rS ¼
0.588, N ¼ 10, P ¼ 0.074).
Feeding Order
The total number of cats that were present around the
food at the beginning of a single session ranged from 2
to 10 (X SD ¼ 4:72 1:85). For adult cats the probability of being the first to feed tended to decrease since October 2001, when kittens appeared at the feeding site,
although not significantly (Wilcoxon signed-ranks test:
T ¼ 3.00, N ¼ 10, P ¼ 0.063; Fig. 2). In fact, when kittens
regularly fed at feeding site, they were more likely than
adults of both sexes to be the first individuals to feed;
the difference tended to be significant in December
2001 (KruskaleWallis test: H2 ¼ 5.930, P ¼ 0.05; mean
ranks: kittens 9.5, females 6, males 4) and was significant in February 2002 (KruskaleWallis test: H2 ¼ 6.281,
P ¼ 0.04; mean ranks: kittens 9.25, females 5.1, males
4), when kittens were 4 and 6 month olds, respectively.
Figure 2 shows that the statistical significant level was
reached probably because of the great difference between kittens and adults, whereas the distance between
females and males was smaller. Kittens’ feeding priority
was not a consequence of the protection of their dominant mother: they were still more likely than adults to
be the first to feed even in the absence of their mother,
the difference between classes being not significant
(KruskaleWallis test: H2 ¼ 1.735, P ¼ 0.420; mean ranks:
kittens 9, males 6.4, females 5.5).
80
Juveniles
Females
Males
70
Probability of being the first to feed
1374
60
50
*
40
30
*
20
10
0
Aug
Sep
Oct
Nov
Dec
Jan
Feb
Mar
Apr
May
Jun
Months
Figure 2. Mean percentage of sessions in which cats of each ageesex class were the first to feed in relation to the total number of their presences around the food and its variation throughout the study period. Asterisks indicate significant differences among the three ageesex classes.
BONANNI ET AL.: FEEDING ORDER IN DOMESTIC CATS
The feeding order (total probability of being the first to
feed) was not significantly correlated with feeding rank
when considering all group members as a whole (rS ¼
0.086, N ¼ 14, P ¼ 0.770 and rS ¼ 0.099, N ¼ 14, P ¼
0.735, for rank based on submissions and aggressions, respectively); however, for adult cats only, it was moderately, although not significantly, correlated with both
rank (rS ¼ 0.460, N ¼ 10, P ¼ 0.181) and feeding rank
(rS ¼ 0.460, N ¼ 10, P ¼ 0.181 and rS ¼ 0.497, N ¼ 10,
P ¼ 0.144, for rank based on submissions and aggressions,
respectively).
DISCUSSION
Intersexual Dominance and Social
Tolerance to Juveniles
In this study we have found that intersexual dominance
relationships in the cat group varied according to different
competitive contexts and to a minor extent to the
behaviour analysed: male cats were usually dominant
away from food, whereas females increased in rank at
the feeding site, especially when the feeding rank assessment was based on aggressive behaviour that is likely to
reflect motivation (Hurd 2006). Moreover, although juveniles were at the bottom of the feeding hierarchy, they
were significantly more likely than adults of both sexes
to be the first to feed around the age of 4e6 months, indicating that they received a certain level of tolerance (lack
of significance after the age of 6 months was probably due
to a decrease in the total number of cats present in the
study area). Social tolerance to kittens explains the total
lack of correlation between feeding rank and feeding order
that we have found considering the cat group as a whole,
since the level of the correlation became moderate once
the kittens were excluded from the analysis.
Yamane et al. (1997), who investigated the feeding order
in a cat colony living in a fishing village in Japan, found
a general tendency for males to feed before females and
kittens before adults of both sexes, although they did
not provide data on agonistic interactions, making it difficult to compare their results with ours. On the other hand,
Knowles et al. (2004) found results similar to those of the
present study (females gaining rank position in feeding
competition), although these authors studied a neutered
cat colony living in a closed environment.
The present work is, to our knowledge, one of the few
studies documenting female dominance over considerably
larger males in a feeding context, in a mammalian species,
not restricted to limited periods of the reproductive cycle
(see examples in the introduction). Dominance was based
on direction of agonistic behaviour in dyadic conflicts,
that is female dominance over larger males was not
acquired through coalition formation, which results in
an increase of RHP, as occurs in several primate species
(e.g. bonobos, Pan paniscus, Furuichi 1989; patas monkeys, Erythrocebus patas, rhesus monkeys, Kappeler 1993).
The results shown here seem to provide support for
game-theory models predicting conflict outcome in favour
of the smaller competitor, when asymmetries in resource
value and in the cost of winning inappropriately may
compensate for the smaller competitor’s RHP. We assume
that conflicts away from food are for general dominance
and the value of winning is roughly similar for both sexes.
Here the outcome is, thus, mainly affected by asymmetries
in RHP with male cats being in advantage over females. In
accordance with studies conducted on other species (e.g.
hens, Gallus gallus domesticus, Cloutier & Newberry 2000;
red deer, Veiberg et al. 2004; brown bears, Gende & Quinn
2004; fallow deer, Dama dama, Jennings et al. 2006), dominance rank (or RHP) was positively correlated with body
weight and size in our cat colony. Head size, in particular,
may be used by cats as a cue to assess each other’s fighting
ability.
Conversely, it seems reasonable to assume that female
cats value the food more highly than males so as to
compensate for their lower RHP. In fact, reproduction can
place a particularly high level of stress on female domestic
cats: it has been shown that their energy intake increases
up to 1.7 times maintenance levels during pregnancy and
up to 3.5 times at peak lactation, when large litters are
reared (Loveridge 1986), whereas a tom 1.5 times as heavy
as a female should have an energy intake just 1.35 times
maintenance levels of a female. However, in this study female cats were more aggressive than males even during
nonreproductive periods. To explain this pattern, we
should consider that females are typically pregnant or lactating during the period JanuaryeOctober, that is for the
majority of the year, and it is possible that, once established, dominance relationships may subsequently continue to influence the decisions to escalate feeding
conflicts.
Several authors have stressed the importance of leverage
in affecting payoff asymmetries in animal contests (Hand
1986; Smuts 1987; Noë et al. 1991; Kappeler 1993; Lewis
2002). It has been argued that the smaller competitor
can become dominant over the larger one because of a leverage advantage: although the female dominance observed among domestic cats in this study and in some
mammalian species where sexes do not differ markedly
in RHP, such as the spotted hyaena (East & Hofer 2002)
and Malagasy lemurs (e. g. Eulemur fulvus mayottensis,
Roeder & Fornasieri 1995; Eulemur macaco flavifrons, Digby
& Kahlenberg 2002; Eulemur fulvus rufus, Ostner et al.
2003; Propithecus diadema edwardsi, Pochron et al. 2003),
may partly be due to females valuing the food more highly
than males, it is also possible that, in some of these species, males may suffer a leverage cost resulting from winning inappropriately a feeding conflict against females.
In spotted hyenas, where females exercise a strong degree
of mate choice, male food deference may serve to affect female preferences (East & Hofer 2002). In lemurs, male
food deference may enable females to meet unusually
high energetic demands of reproduction and eventually
to successfully wean infants (Kappeler 1993; Pochron
et al. 2003). Similarly, for male domestic cats it might be
convenient to allow both female dominance and juveniles’ feeding priority to occur. Even if male domestic
cats play no direct part in rearing their kittens, they could
contribute to increase kittens’ survival by tolerating both
females and kittens in feeding situation. A leverage cost
1375
1376
ANIMAL BEHAVIOUR, 74, 5
must be necessarily involved in cases of food deference to
juveniles by adults since asymmetries in RHP between
adults and juveniles appear to be too extreme to be overcome by asymmetries in resource value. Since to deny
food to a given offspring should be more costly to males
that invest heavily in few offspring, a correlation could
be expected between the degree of paternal investment
and this type of male tolerance (Hand 1986). However,
a more general condition may be that tolerance occurs
whenever winning elevates costs for the larger competitor
beyond a certain critical value (Hand 1986). Male food
deference to females can be found in association with juveniles’ feeding priority in some primate species wherein
males invest considerably in parental care (e.g. common
marmosets, Callithrix jacchus; golden lion tamarins, Leontopithecus rosalia; titi monkeys, Callicebus moloch; indri,
Indri indri, all reviewed in Hrdy 1981), as well as in a cooperative breeder carnivore such as the African wild dog
(Malcom & Marten 1982) and in the spotted hyaena
(Frank 1986), suggesting that in these species, as in the domestic cat, both the types of tolerance may be part of a single male reproductive strategy. In domestic cats, such male
tolerance could have evolved in the original environment
of adaptation in which, since males are expected to monopolize the access to oestrous females living within the
boundaries of their own territory (Say et al. 1999; Liberg
et al. 2000), there are high probabilities that the females
encountered are carrying or nursing the males’ offspring.
On the other hand, male tolerance may be a consequence
of the high level of paternal uncertainty resulting from the
promiscuous mating system found among domestic cats
in the urban environment (Natoli & De Vito 1991; Say
et al. 1999; Natoli et al. 2005).
In a similar way, female tolerance of kittens may be
a consequence of females having a weak capacity to
discriminate their own offspring from those of other
females, given that for a female cat living solitarily in the
original environment of adaptation the evolution of a finely
tuned parenteoffspring recognition is not necessary.
Comparison with the Dominance System
of Lions
The behaviour of domestic cats and lions has been
compared by several authors (see e.g. Macdonald et al.
1987; Natoli 1990; Natoli & De Vito 1991; Liberg et al.
2000) because they are the only two felids that have
been proven to be able to live socially. Their social behaviour shows some striking similarities. First, both lions
(Schaller 1972; Packer et al. 1991) and domestic cats (Liberg et al. 2000; Macdonald et al. 2000) form social groups
composed of genetically related females, their dependent
offspring and attached adult males. Second, both lionesses
(Pusey & Packer 1994a) and female domestic cats (Macdonald et al. 1987) show alloparental care of juveniles
and cooperation in territorial defence.
However, the feeding competition pattern and the
dominance system observed in this study among domestic
cats were quite different from those found in lions. Unlike
male domestic cats of our social group, which were less
aggressive than females at feeding sites and allowed
kittens’ feeding priority, male lions dominate all other
ageesex classes at kill, routinely supplanting both females
and cubs (Packer et al. 2001). Even if male lions have been
shown to be effective in hunting large prey (Funston et al.
1998), they can acquire the bulk of their food intake by
appropriating kills made by lionesses (Van Orsdol 1986).
In contrast to males, lionesses usually do not displace juveniles from a specific feeding site around a carcass (Packer
et al. 2001).
To explain such differences we should consider the
different reproductive tactics followed by these two feline
species. Male lions form cooperative coalitions which
compete against other coalitions for exclusive possession
of a female pride (Bygott et al. 1979; Packer et al. 1988;
Grinnell et al. 1995). As demonstrated by paternity analysis, a resident coalition is very effective in preventing extra-pride males from fathering cubs (Packer et al. 1991)
and, consequently, the possession of a pride of females
is the only chance for a male lion to gain mating opportunities. The male group tenure is retained for a variable period and typically ends when they are displaced from their
pride by a new group of males (Bygott et al. 1979; Packer
et al. 1988). Incoming males accelerate the females’ return
to oestrus by killing unweaned cubs and evicting subadults
(Pusey & Packer 1994b), and thus prolonged residence is essential for successful reproduction. Male reproductive success increases with coalition size, mostly because larger
coalitions maintain residence for longer periods, thus
fathering more cubs and postponing the next episode of
infanticide (Bygott et al. 1979; Packer et al. 1988). We hypothesize that selective pressures favouring male dominance and aggressiveness at feeding occasions would be
particularly strong in lions, because male lions need to
maintain constantly a peak physical condition so as
to ward off rivals, to prolong their tenure in a pride and
to protect their cubs from the risk of infanticide. Paradoxically, if male lions were more tolerant to cubs, they could
not be nourished enough to protect them from infanticidal rivals. Recent findings seem to support our hypothesis:
dark-maned lions, which have higher levels of testosterone and hence of aggressiveness, are better nourished
and enjoy both longer tenures and higher offspring survival (West & Packer 2002).
Furthermore, among lions, females would not probably
have a leverage advantage over males: lionesses themselves, and not just males, may incur a leverage cost by
winning a feeding conflict against a competitor of the
opposite sex, because they probably need male help in
protecting offspring from infanticide.
These arguments do not apply to domestic cats. Male
domestic cats are not known to form coalitions which
cooperate to take-over a group of females. Also, defence of
females is not the only mating tactic available to them:
even in rural populations, where the mating system is
polygyny, territorial males are not completely successful
in preventing transient males from copulating with
females (Say et al. 1999). In addition, infanticide has not
been documented in urban domestic cats and rarely documented in rural cats (Natoli 1990; Pontier & Natoli 1999).
This is not surprising because infanticide is thought to be
BONANNI ET AL.: FEEDING ORDER IN DOMESTIC CATS
advantageous as a male reproductive tactic in species that,
like lions, do not breed seasonally and show a very long
interbirth interval (Hausfater et al. 1982; Packer & Pusey
1984). Unlike lions, domestic cats are seasonal breeders
with a short interbirth interval (Schmidt et al. 1983):
a mated female domestic cat is unavailable to other males
for about 4 months from the beginning of oestrus to the
weaning of kittens; furthermore, in the present study, a female cat came back into oestrus and performed fertile matings before the weaning of her current litter. Thus, for
a male domestic cat infanticide is not crucial for speeding
up the reproductive cycle of females, whereas for a male
lion it is the only chance to bring a female back into oestrus in reasonable time (Natoli 1990). In conclusion, we
believe that male domestic cats do not behave aggressively
at feeding site in the same way as lions, or do not value the
food in the same way, because they do not need to maintain constantly a peak physical condition in order to retain exclusive control of a group of females and to
defend their offspring from infanticidal rivals.
Domestication
Domestication is thought to have led to a decrease in
intraspecific aggression and loss of social inhibition in
a variety of species, mostly because of artificial selection
and relaxation of natural selective pressures under human
influence which has provided animals with abundant
resources (Price 1984). It has been suggested that the
same applies to domestic cats because they apparently
show greater sociability and behavioural flexibility than
their wild solitary ancestor, the African wild cat (Macdonald et al. 2000). Thus, it may be thought that the social tolerance displayed, in this study, by male domestic cats to
both females and kittens is a by-product of domestication,
which would have caused a decrease in male aggressiveness, rather than the result of natural selection mechanisms. However, in our opinion, it seems unlikely that
domestication would have caused a differential decrease
in aggression in the two sexes and differentially in different competitive contexts. Given that our results seem to
be in accordance with predictions of game-theory models,
they are more likely to reflect selective processes possibly
operating in the original environment of adaptation before domestication and urbanization.
Moreover, evolution of sociality does not necessarily
imply a reduction in intraspecific aggression: studies conducted on social insects indicate that it involves evolution
of conditional aggressive behaviour rather than unconditional intraspecific tolerance (Cahan 2001).
Indeed, whether or not the evolution of sociality in cats
has been promoted by domestication is still a debatable
point: feral domestic cats live solitarily when depending
on widely dispersed natural prey, whereas they form social
groups in the presence of clumped, rich food resources
provided by human beings (Liberg et al. 2000); even if
fragmentary evidence indicates that African wild cats do
not form colonies where they are exposed to abundant
human refuse (Macdonald et al. 2000), it is intriguing
that, in captivity, female African wild cats have assisted
mothers in provisioning of kittens with food (Nowell &
Jackson 1996), a behaviour observed in feral domestic
cats colonies, but not in any other solitary cat species.
Domestic cats do not necessarily have a greater level of
behavioural flexibility than their wild ancestor, but they
may have simply been exposed to a far greater diversity
of ecological conditions (Leyhausen 1988).
Acknowledgments
Special thanks are due to Antonio Buogo for his availability to all our requests during the study, to all veterinarians
working at the Veterinary Hospital of Rome for their help
in handling cats and, in particular, to Giuseppe Cariola for
his unvaluable help in trapping cats, anaesthetizing them
and collecting blood samples. We wish to thank Masako
Izawa for procuring relevant literature, Alessandro Giuliani for valuable suggestions, Marion L. East for helping
with the figures and the referees for their valuable
comments that helped us to greatly improve the manuscript. Finally, a special thanks goes to Prof. Alberto
Fanfani from the Department of Animal and Human
Biology, University of Rome ‘La Sapienza’, who offered
support and facilities.
References
Altmann, J. 1974. Observational study of behavior: sampling
methods. Behaviour, 49, 227e267.
Archer, J. 1988. The Behavioural Biology of Aggression. Cambridge:
Cambridge University Press.
Appleby, M. C. 1980. Social rank and food access in red deer stags.
Behaviour, 74, 295e309.
Barton, A. & Whiten, A. 1993. Feeding competition among female
olive baboons, Papio anubis. Animal Behaviour, 46, 777e789.
Bygott, J. D., Bertram, B. C. R. & Hanby, J. P. 1979. Male lions in
large coalitions gain reproductive advantages. Nature, 282,
839e841.
Cahan, S. H. 2001. Cooperation and conflict in ant foundress associations: insights from geographical variation. Animal Behaviour,
61, 819e825.
Caro, T. M., Roper, R., Young, M. & Dank, G. R. 1979. Interobserver reliability. Behaviour, 69, 303e315.
Cloutier, S. & Newberry, R. C. 2000. Recent social experience,
body weight and initial patterns of attack predict social status
attained by unfamiliar hens in a new group. Behaviour, 137,
705e726.
Deag, J. M., Manning, A. & Lawrence, C. E. 2000. Factors influencing the mother-kitten relationship. In: The Domestic Cat: the Biology of its Behaviour. 2nd edn (Ed. by D. C. Turner & P. Bateson),
pp. 23e45. Cambridge: Cambridge University Press.
Deutsch, J. C. & Lee, P. C. 1991. Dominance and feeding competition in captive rhesus monkeys. International Journal of Primatology,
12, 615e628.
Digby, L. J. & Kahlenberg, S. M. 2002. Female dominance in
blue-eyed black lemurs (Eulemur macaco flavifrons). Primates,
43, 191e199.
East, M. L. & Hofer, H. 2002. Conflict and cooperation in a female
dominated society: a reassessment of the ‘‘hyperaggressive’’
image of spotted hyenas. Advances in the Study of Behavior, 31,
1e30.
1377
1378
ANIMAL BEHAVIOUR, 74, 5
Enquist, M. & Leimar, O. 1987. Evolution of fighting behaviour: the
effect of variation in resource value. Journal of Theoretical Biology,
127, 187e205.
Frank, L. G. 1986. Social organization of the spotted hyaena Crocuta
crocuta. II. Dominance and reproduction. Animal Behaviour, 34,
1510e1527.
Funston, P. J., Mills, M. G. L., Biggs, H. C. & Richardson, P. R. K.
1998. Hunting by male lions: ecological influences and socioecological implications. Animal Behaviour, 56, 1333e1345.
Furuichi, T. 1989. Social interactions and the life history of female
Pan paniscus in Wamba, Zaire. International Journal of Primatology,
10, 173e197.
Gende, S. M. & Quinn, T. P. 2004. The relative importance of prey
density and social dominance in determining energy intake by
bears feeding on Pacific salmon. Canadian Journal of Zoology, 82,
75e85.
Grinnell, J., Packer, C. & Pusey, A. E. 1995. Cooperation in male
lions: kinship, reciprocity or mutualism? Animal Behaviour, 49,
95e105.
Hammerstein, P. & Parker, G. A. 1982. The asymmetric war of attrition. Journal of Theoretical Biology, 96, 647e682.
Hand, J. L. 1986. Resolution of social conflicts: dominance, egalitarianism, spheres of dominance, and game theory. Quarterly Review
of Biology, 61, 201e220.
Hausfater, G., Aref, S. & Cairns, S. J. 1982. Infanticide as an alternative male reproductive strategy in langurs: a mathematical
model. Journal of Theoretical Biology, 94, 391e412.
Hrdy, S. B. 1981. The Woman that Never Evolved. Cambridge, Massachusetts: Harvard University Press.
Hurd, P. L. 2006. Resource holding potential, subjective resource
value, and game theoretical models of aggressiveness signalling.
Journal of Theoretical Biology, 241, 639e648.
Izawa, M., Doi, T. & Ono, Y. 1982. Grouping patterns of feral cats
(Felis catus) living on a small island in Japan. Japanese Journal of
Ecology, 32, 373e382.
Jennings, D. J., Gammell, M. P., Carlin, C. M. & Hayden, T. J.
2006. Is difference in body weight, antler length, age or dominance rank related to the number of fights between fallow deer
(Dama dama)? Ethology, 112, 258e269.
Kappeler, P. M. 1993. Female dominance in primates and other
mammals. In: Behaviour and Evolution (Ed. by P. P. G. Bateson),
pp. 143e158. New York: Plenum Press.
Knowles, R. J., Curtis, T. M. & Crowell-Davis, S. L. 2004. Correlation of dominance as determined by agonistic interactions with
feeding order in cats. American Journal of Veterinary Research, 65,
1548e1556.
Lewis, R. J. 2002. Beyond dominance: the importance of leverage.
Quarterly Review of Biology, 77, 149e164.
Leyhausen, P. 1988. The tame and the wild e another just-so story? In:
The Domestic Cat: the Biology of its Behaviour (Ed. by D. C. Turner &
P. Bateson), pp. 57e66. Cambridge: Cambridge University Press.
Liberg, O., Sandell, M., Pontier, D. & Natoli, E. 2000. Density, spatial organisation and reproductive tactics in the domestic cat and
other felids. In: The Domestic Cat: the Biology of its Behaviour. 2nd
edn (Ed. by D. C. Turner & P. Bateson), pp. 119e147. Cambridge:
Cambridge University Press.
Loveridge, G. G. 1986. Bodyweight changes and energy intake of
cats during gestation and lactation. Animal Technology, 37, 7e15.
Macdonald, D. W., Apps, P. J., Carr, G. M. & Kerby, G. 1987. Social dynamics, nursing coalitions and infanticide among farm cats,
Felis catus. Advances in Ethology, 28, 1e64.
Macdonald, D. W., Yamaguchi, N. & Kerby, G. 2000. Group-living
in the domestic cat: sociobiology and epidemiology. In: The Domestic Cat: the Biology of its Behaviour. 2nd edn (Ed. by D. C.
Turner & P. Bateson), pp. 95e118. Cambridge: Cambridge University Press.
Malcom, J. R. & Marten, K. 1982. Natural selection and the communal rearing of pups in African wild dogs (Lycaon pictus). Behavioral Ecology and Sociobiology, 10, 1e13.
Martin, P. & Bateson, P. 1993. Measuring Behaviour. Cambridge:
Cambridge University Press.
Natoli, E. 1985. Spacing pattern in a colony of urban stray cats (Felis
catus L.) in the historic centre of Rome. Applied Animal Behaviour
Science, 14, 289e304.
Natoli, E. 1990. Mating strategies in cats: a comparison of the role
and importance of infanticide in domestic cats, Felis catus L.,
and lions, Panthera leo L. Animal Behaviour, 40, 183e186.
Natoli, E. & De Vito, E. 1991. Agonistic behaviour, dominance rank
and copulatory success in a large multi-male feral cat, Felis catus L.,
colony in central Rome. Animal Behaviour, 42, 227e241.
Natoli, E., Baggio, A. & Pontier, D. 2001. Male and female
agonistic and affiliative relationships in a social group of farm
cats (Felis catus L.). Behavioural Processes, 53, 137e143.
Natoli, E., Say, L., Cafazzo, S., Bonanni, R., Schmid, M. & Pontier,
D. 2005. Bold attitude makes male urban feral domestic cats more
vulnerable to Feline Immunodeficiency Virus. Neuroscience and
Biobehavioral Reviews, 29, 151e157.
Noë, R., de Waal, F. B. M. & van Hooff, J. A. R. A. M. 1980. Types
of dominance in a chimpanzee colony. Folia Primatologica, 34,
90e110.
Noë, R., van Schaik, C. P. & van Hooff, J. A. R. A. M. 1991. The
market effect: an explanation for pay-off asymmetries among
collaborating animals. Ethology, 87, 97e118.
Nowell, K. & Jackson, P. 1996. Wild cats. Status Survey and
Conservation Action Plan. Gland, Switzerland: IUCN.
Ostner, J., Heistermann, M. & Kappeler, P. M. 2003. Intersexual dominance, masculinized genitals and prenatal steroids: comparative
data from lemurid primates. Naturwissenschaften, 90, 141e144.
Packer, C. & Pusey, A. E. 1984. Infanticide in carnivores. In: Infanticide (Ed. by G. Hausfater & S. B. Hrdy), pp. 31e42. Chicago:
Aldine.
Packer, C., Pusey, A. E. & Eberly, L. E. 2001. Egalitarianism in female African lions. Science, 293, 690e693.
Packer, C., Herbst, L., Pusey, A. E., Bygott, J. D., Hanby, J. P.,
Cairns, S. J. & Borgerhoff-Mulder, M. 1988. Reproductive success of lions. In: Reproductive Success (Ed. by T. H. Clutton-Brock),
pp. 363e383. Chicago: University of Chicago Press.
Packer, C., Gilbert, D. A., Pusey, A. E. & O’Brien, S. J. 1991. A molecular genetic analysis of kinship and cooperation in African lions.
Nature, 351, 562e565.
Parker, G. A. 1974. Assessment strategy and the evolution of fighting behaviour. Journal of Theoretical Biology, 47, 223e243.
Parker, G. A. & Rubenstein, D. I. 1981. Role assessment, reserve
strategy, and acquisition of information in asymmetric animal conflicts. Animal Behaviour, 29, 221e240.
Pascal, M. & Castanet, J. 1978. Méthode de détermination de l’âge
chez le chat haret des ı̂les Kergulen. La terre et la Vie, 4, 529e555.
Pochron, S. T., Fitzgerald, J., Gilbert, C. C., Lawrence, D., Grgas,
M., Rakotonirina, G., Ratsimbazafy, R., Rakotosoa, R. &
Wright, P. C. 2003. Patterns of female dominance in Propithecus
diadema edwardsi of Ranomafana National Park, Madagascar.
American Journal of Primatology, 61, 173e185.
Pontier, D. & Natoli, E. 1999. Infanticide in rural male cats (Felis catus L.)
as a reproductive mating tactic. Aggressive Behavior, 25, 445e449.
Pontier, D., Fromont, E., Courchamp, F., Artois, M. & Yoccoz,
N. G. 1998. Retroviruses and sexual size dimorphism in domestic
cats (Felis catus L.). Proceedings of the Royal Society of London, Series B,
265, 167e173.
BONANNI ET AL.: FEEDING ORDER IN DOMESTIC CATS
Pontier, D., Rioux, N. & Heizmann, A. 1995. Evidence of selection
on the orange allele in the domestic cat Felis catus: the role of social structure. Oikos, 73, 299e308.
Price, E. O. 1984. Behavioral aspects of animal domestication. Quarterly Review of Biology, 59, 1e32.
Pusey, A. E. & Packer, C. 1994a. Non-offspring nursing in social carnivores: minimizing the costs. Behavioral Ecology, 5, 362e374.
Pusey, A. E. & Packer, C. 1994b. Infanticide in lions: consequences
and counterstrategies. In: Infanticide and Parental Care (Ed. by S.
Parmigiani & F. vom Saal), pp. 277e299. Chur, Switzerland: Harwood Academic Press.
Roeder, J. J. & Fornasieri, I. 1995. Does agonistic dominance imply
feeding priority in lemurs? A study in Eulemur fulvus mayottensis.
International Journal of Primatology, 16, 629e642.
Say, L., Pontier, D. & Natoli, E. 1999. High variation in multiple paternity of domestic cats (Felis catus L.) in relation to environmental
conditions. Proceedings of the Royal Society of London, Series B, 266,
2071e2074.
Schaller, G. B. 1972. The Serengeti Lion: a Study of PredatorePrey
Relations. Chicago: University of Chicago Press.
Schmidt, P. M., Chakraborty, P. K. & Wildt, D. E. 1983. Ovarian
activity, circulating hormones and sexual behaviour in the cat. II.
Relationships during pregnancy, parturition, lactation and postpartum oestrus. Biology of Reproduction, 28, 657e671.
Serpell, J. A. 2000. Domestication and history of the cat. In: The Domestic Cat: the Biology of its Behaviour. 2nd edn (Ed. by D. C.
Turner & P. Bateson), pp. 179e192. Cambridge: Cambridge University Press.
Smuts, B. B. 1987. Gender, aggression and influence. In: Primate Societies (Ed. by B. B. Smuts, D. L. Cheney, R. M. Seyfarth, R. M.
Wrangham & T. T. Struhsaker), pp. 400e412. Chicago: University
of Chicago Press.
Stopka, P., Johnson, D. D. & Barret, L. 2001. Friendship’’ for fitness
or ‘‘friendship for friendship’s’’ sake? Animal Behaviour, 61,
F19eF21.
Van Orsdol, K. G. 1986. Feeding behavior and food intake of lions in
Rwenzori National Park, Uganda. In: Cats of the World: Biology,
Conservation, and Management (Ed. by S. D. Miller & D. D. Everett), pp. 377e388. Washington D.C.: National Wildlife Federation.
Veiberg, V., Loe, L. E., Mysterud, A., Langvatn, R. & Stenseth,
N. C. 2004. Social rank, feeding and winter weight loss in red
deer: any evidence of interference competition? Oecologia, 138,
135e142.
de Vries, H. 1995. An improved test of linearity in dominance hierarchies containing unknown or tied relationships. Animal Behaviour, 50, 1375e1389.
West, P. M. & Packer, C. 2002. Sexual selection, temperature, and
the lion’s mane. Science, 297, 1339e1343.
Wilson, E. O. 1975. Sociobiology: the New Synthesis. Cambridge,
Massachusetts: Harvard University Press.
Wittig, R. M. & Boesch, C. 2003. Food competition and linear dominance hierarchy among female chimpanzees of the Ta€ı National
Park. International Journal of Primatology, 24, 847e867.
Yamane, A., Emoto, J. & Ota, N. 1997. Factors affecting feeding
order and social tolerance to kittens in the group-living feral cat
(Felis catus). Applied Animal Behaviour Science, 52, 119e127.
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