Feeding-order in an urban feral domestic cat colony:

relationship to dominance rank, sex and age

ROBERTO BONANNI* , SIMONA CAFAZZO* , CLAUDIO FANTINI†, DOMINIQUE PONTIER‡ & EUGENIA NATOLI†

*Dipartimento di Biologia Evolutiva e Funzionale, Universita di Parma

yAzienda USL Roma D, Area Dipartimentale Sanita Pubblica Veterinaria

zUMR-CNRS 5558 ‘‘Biometrie and Biologie Evolutive’’, Universite 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 asymmetriesin resource-holding potential (RHP) between competitors of different ageesex classes with males usuallybeing dominant and feeding first, followed by females and then juveniles. In this study we investigatedhow variables such as sex and age affected dominance rank and feeding order in a social group of feral do-mestic cats, Felis silvestris catus, a sexually dimorphic species in which males are larger than females and donot take part in parental care. Intersexual dominance relationships varied depending on the competitivecontext: males occupied top rank positions away from food, whereas females increased in rank at the ex-pense of males in a feeding context. Around the age of 4e6 months, kittens were significantly more likelythan 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 smallercompetitor when asymmetries in both the value of winning and in the cost of winning inappropriatelymay compensate for the smaller competitor’s lower RHP. It is suggested that the results are not an artifactof 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 main-tain constantly a peak physical condition to defend a group of females and protect offspring frominfanticide.

� 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

ANIMAL BEHAVIOUR, 2007, 74, 1369e1379doi:10.1016/j.anbehav.2007.02.029

Social dominance has historically been defined as priorityof access to resources (Wilson 1975). Since food is consid-ered a major determinant of the reproductive success ofindividuals, several authors have tested the influence ofdominance rank on access to food and it has been verified

Correspondence and present address: R. Bonanni, Dipartimento diBiologia Evolutiva e Funzionale, Universita di Parma, Parco Area delleScienze 11/A, 43100 Parma, Italy (email: roberto.bonanni@inwind.it).C. Fantini is at the Azienda USL Roma D, Area Dipartimentale SanitaPubblica Veterinaria, Direzione, via Portuense 1397, 00050 Ponte Ga-leria (Roma), Italia. D. Pontier is at the UMR-CNRS 5558 ‘‘Biometrieand Biologie Evolutive’’, Universite de Lyon, Universite Lyon 1, 43 Bddu 11 nevembre 1918, 69622 Villeurbanne cedex, France. E. Natoliis at the Azienda USL Roma D, Area Dipartimentale Sanita PubblicaVeterinaria, Ospedale Veterinario, via della Magliana 856, 00148Roma, Italy.

10003e3472/07/$30.00/0 � 2007 The Association for the

that higher ranking members of a social group enjoy prior-ity of access to food over subordinates in an array of ani-mal species (e.g. red deer, Cervus elaphus, Appleby 1980;rhesus monkeys, Macaca mulatta, Deutsch & Lee 1991; ol-ive baboons, Papio anubis, Barton & Whiten 1993; chim-panzees, Pan troglodytes, Wittig & Boesch 2003; brownbears, Ursus arctos, Gende & Quinn 2004).

Patterns of dominance relationships and food accesscan be expected to follow predictions of classic game-theory models, with contest outcomes largely affected byasymmetries in both resource-holding potential (RHP),which is a measure of fighting ability, and relative resourcevalue between contestants (Parker & Rubenstein 1981;Hammerstein & Parker 1982; Enquist & Leimar 1987). Pre-cisely, competitors should be more likely to escalate andeventually win conflicts, the higher their own RHP andthe higher the value of the resource to them.

369Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

ANIMAL BEHAVIOUR, 74, 51370

In many species, body weight (and/or size) is a goodindicator of a competitor’s RHP (Parker 1974; Archer1988), and where asymmetries in body weight exist be-tween competitors belonging to different ageesex classesin a social group, these usually predict conflict outcomeover food resources: a well-known example is that of thelion, Panthera leo, in which males are the largest membersof the group and dominate both females and juveniles infeeding competition situations (Van Orsdol 1986; Packeret al. 2001). In species where females invest substantiallymore energy than males in parental care, food may beworth more to them than to males. In some mammalianspecies showing a lack of sexual size dimorphism, suchas the spotted hyaena, Crocuta crocuta, and several Lemur-iformes, 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 underwhich contest outcome may be predicted in favour ofthe smaller competitor. Hand (1986), following Parker &Rubenstein (1981), pointed out that, while the fitnesscosts in terms of injuries sustained increase at rates in-versely correlated with RHP during a contest, there couldbe an additional fitness cost arising from winning a con-flict when it is inappropriate from the perspective offitness to do so, that may, under some conditions, behigher for the larger competitor. This may occur in con-flicts between opponents which share some individualfitness interests, such as conflicts between mates or pa-renteoffspring conflicts. For instance, a male could reducedirectly his fitness by driving the mother of his offspringor the offspring themselves away from critical food re-sources because, in both cases, he would probably decreaseoffspring survival. Since both the female and the off-spring, which usually are smaller than the male, are likelyto have less fitness to lose from winning than does themale, they may have a ‘leverage’ advantage over themale. If such asymmetry in the cost of winning equalizesthe total costs for both opponents, conflict outcome willprobably be decided by asymmetries in resource value(Hand 1986). Since both the value of the resource andthe cost of winning (called ‘leverage cost’) may changeconsiderably depending on the competitive context, thedominance relationship in a given dyad of individualsmay also vary according to the context (Hand 1986; seeNoe et al. 1980 for an empirical example). This modelmay help to explain the cases of male food deference tofemales during periods corresponding to egg laying or oes-trus (e.g. Western Gulls, Larus spp., Hand 1986; chimpan-zees, Stopka et al. 2001), when the food is of particularlyhigh value to females, and also why adult individuals ofboth sexes allow juveniles’ feeding priority in a varietyof species (e.g. wild dogs, Lycaon pictus, Malcom & Marten1982; spotted hyenas, Frank 1986; several primates,Hrdy 1981).

In this paper we examine how variables such as sex andage affect dominance and access to food in a social groupof feral domestic cats, Felis silvestris catus, living in an ur-ban environment, and test whether the pattern found fol-low the predictions of evolutionary game-theory models.

In urban areas, domestic cats live at very high densities(up to 2000 cats/km2) and form multimaleemultifemalesocial groups subsisting on an abundant, patchily distrib-uted, source of food provided by ‘cat lovers’ at a few tradi-tional feeding sites (Izawa et al. 1982; Natoli 1985; Liberget al. 2000). Such groups provide an ideal situation tostudy social factors affecting feeding competition becauseseveral individuals of all ageesex classes usually gather atthe places where food is distributed by human beings. Ithas been suggested that domestication may have fosteredthe evolution of sociality in cats through an increase inbehavioural plasticity and social tolerance of conspecifics(Macdonald et al. 2000). However, the cat’s history as a do-mesticated animal is very short, about 4000 years, in evo-lutionary terms (Serpell 2000), and many observedbehavioural patterns may have evolved in their solitaryancestor, the African wild cat, Felis silvestris lybica, beforedomestication and urbanization.

Whether a linear dominance hierarchy exists amongdomestic cats is controversial probably because, as a resultof missing interactions between some individuals, it is notalways easy to reach a statistically significant level thatprove the occurrence of linearity (see for example Natoli &De Vito 1991). Recently, Natoli et al. (2001) found a signif-icantly linear dominance hierarchy in a rural cat colony,based on the outcome of agonistic encounters in the ab-sence of any sources of competition, in which a femaledominated some males. The last could be surprising be-cause, as the lion, the domestic cat is a sexually dimorphicspecies in which males are 15e40% heavier (Pontier et al.1995, 1998) and possess 19% longer canines (D. Pontier,unpublished data) than females. Possibly, in domesticcats female dominance is achieved by other mechanismsthan physical strength and fighting ability (Natoli et al.2001). To improve our comprehension of such mecha-nisms it seems useful to investigate dominance relation-ships among cats in feeding competition. Since food isthought to be a valuable resource for female cats, becausemale cats do not take part in parental care (Deag et al.2000), it can be expected that females would gain furtherrank positions in a feeding context, at least duringpregnancy and lactation, where the value of winningthe resource, together with the possible higher costs tomales of winning inappropriately, may offset asymmetriesin RHP.

Although male feeding deference to juveniles is com-moner in species in which males provide substantialparental care (Hand 1986), it could be expected that, ifmale domestic cats allow female dominance due to thehigh cost of winning a feeding conflict against the motherof their offspring, they would also display a certain degreeof feeding tolerance to kittens.

Here, we ascertained whether a dominance hierarchybased on outcome of agonistic encounters in the absenceof any sources of competition (food or receptive females)exists in an urban cat group. Then, we examined thedominance relations in the feeding context, and com-pared the rank found (feeding rank henceforth) with thatobtained in the absence of food. At the same time, werecorded which cats were the first to feed among all thosepresent at the feeding station (feeding order). Finally, we

BONANNI ET AL.: FEEDING ORDER IN DOMESTIC CATS 1371

sought to answer the following questions: is high socialrank associated with priority of access to food? Which sexis dominant? Are kittens tolerated by adults in competi-tive feeding situations?

METHODS

Study Area

The study was carried out in Rome, in a popular quartercalled ‘Garbatella’. The cat colony lived in a large court-yard (about 6000 m2) owned by the Istituto AutonomoCase Popolari of Rome. It was bounded by a wall and com-pletely isolated from road traffic. The courtyard contained11 buildings, open areas, flower-beds cultivated by inhab-itants, trees and bushes. Cats had free access to every partof the courtyard but they established their ‘core area’ ina partially wire-fenced sector located in the south side ofthe courtyard. Here, a lot of spontaneous vegetation of-fered good shelter for animals, especially for lactating fe-males with kittens. The experimental feeding site wasplaced in a small field (about 150 m2), adjacent to thecore area, in which spontaneous vegetation was periodi-cally removed and where cats could be easily observedwhile feeding without any disturbance. This was themost important and regular food source for the colony.Cats were rarely seen preying on small rodents, passerinebirds, feral pigeons, Columba domestica, and insects. Waterwas available from a fountain just outside the entrance ofthe courtyard.

Cat Group

The cat group living in the study area had no con-straints placed on breeding or movements by humanowners and very few individuals were tame enough to behandled.

We visually recognized all individuals by their coatcolour pattern and hair length. The sex of adult cats couldbe determined on the basis of some morphological (pres-ence of testes in males) and behavioural characteristics.Mature males have a wider face than females; moreoverthey show posterior urine spraying which is a behaviourrarely displayed by females.

Age of the subjects was obtained from cat lovers orestimated according to Pascal & Castanet (1978) method.Cats were considered to be juveniles up to the age of 11months and to be adults afterwards.

Throughout the period of study the number of individ-uals ranged from a minimum of 10 to a maximum of 17cats. In September 2001, the cat colony consisted of eightintact adult males, three intact adult females, two adultfemales that had been neutered long before the studystarted, and four kittens (all of them were the offspring ofthe same female, LAV). Subsequently, two adult malesdispersed from the study area to settle in two differentneighbouring colonies, while five other individuals werekilled by cars.

Nine adult cats (five males and four females) werecaptured during the study period: one female was trapped

using a double-door trap; the remaining individuals wereshot with anesthetizing darts by mean of a blow-pipe,since they were too elusive to be trapped. Cats wereanaesthetized with an intramuscular injection of ketaminechlorhydrate (Inoketam 1000, 5 mg/kg, Virbac, Milan, Italy)and medetomidine (Domitor, 0.005 mg/kg, Pfizer, Rome,Italy). For all captured animals the following measureswere recorded: body weight, head volume, length of theright arm, and average length of canines. The same mea-surements, except head volume, were taken for a furtheradult female after she was found dead. For males, testicles’volume was also recorded, although it was not analysedhere because of the low number of captured males.

Experimental Procedure

We used a focal animal sampling technique (Altmann1974) to study the social behaviour of 13 adult cats. A totalof 555.63 h of observation were made between September2001 and June 2002. Each individual’s observations wereequally distributed over the time period, as well as overdaytime between 0900 and 1800 hours.

Agonistic behaviour (including aggressive and submis-sive behaviour) in absence of any sources of competition(food, receptive females and shelters) was recorded by ‘alloccurrences’ method (Altmann 1974). Aggressive behav-iour included the following: threats (striking with a paw,biting, assuming threatening postures, pointing, staringat, baring of the canines), chasing, ritualized vocal duelsand real duels. Submissive behaviour included: crouchingwith the ears flattened, avoiding, retreating, fleeing andhissing at. Kittens were not included in this data collec-tion because, in absence of food, they were expected to in-teract agonistically too rarely with adults. The individualscores of all behaviour patterns were corrected for animalobservation time because this varied between individuals.

In order to determine the dominance hierarchy, thedistributions of dyadic aggressive and submissive interac-tions were ranked in two different squared matrices withperformers on one axis and recipients on the other, so asto minimize the number of dominance reversals. Thedominant animal of each dyad was the one that performedmore aggressions than it received, or received more sub-missions than it performed. Hissing was excluded from thematrix based on submissive interactions because it wasconsidered an ambiguous behaviour, with componentsboth of aggressiveness and subordination. We tested thetransitivity of dominance relationships between the mem-bers of the social group, based either on aggressive or onsubmissive behavioural patterns separately, by applyingan improved test of linearity developed by de Vries (1995)that is based on Landau’s linearity index, but takes into ac-count unknown and tied relationships between groupmembers.

Experimental Feeding Sessions

We performed a total of 174 experimental feedingsessions from August 2001 to June 2002, correspondingto 178.75 h of observation. Food was delivered to cats at

ANIMAL BEHAVIOUR, 74, 51372

0930 or at 1430 hours on a rotational daily basis. Each ses-sion started just after the food container was placed andcontinued until the food was completely consumed. We as-sembled a plastic container on a baked-clay support andfilled it with 400 g of cat food. This structure had a circularhole (10 cm in diameter) on the upper surface that pre-vented 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 ob-served all individuals present within 4 m of the food, in-cluding adults and kittens. Since subgroup compositionchanged during a session, we recorded the sequence inwhich cats arrived at the feeding site and left it.

To determine the feeding order we estimated, for eachcat, the likelihood of being the first individual to eat of allthose that were present around the food at the beginningof each session, by calculating the percentage of sessionsin which it was the first to feed, in the total number of itspresences around the food, and verified whether suchpercentage varied throughout the study period.

Agonistic behaviour within 1 m of food was recorded byAll Occurrences method (Altmann 1974). This included theaggressive and submissive patterns described above, plus‘interruption of feeding after receiving an aggression’ thatwas a submissive behaviour. Individual scores were cor-rected for presence scores around the food (within 4 m).In order to determine the dominance hierarchy within1 m of food (feeding rank), we constructed two differentmatrices for aggressions and submissions, respectively,and applied the same methods described above. Wherethere were no data on outcomes of submissive interactionsover food by means of which to rank individuals, they havebeen ordered according to the outcomes of aggressive inter-actions over food; similarly, where there were no data onoutcomes of aggressive interactions over food by mean ofwhich to rank individuals, they have been ordered accord-ing to the outcomes of submissive interactions over food.One remaining ambiguity within a specific dyad was re-solved by ordering the cats according to agonistic inter-actions observed between 1 and 4 m of food.

We performed nonparametric statistical tests usingSTATISTICA 99 edition (StatSoft Inc., Tulsa, OK, U.S.A.)

and the Improved linearity tests using MATMAN (NoldusNoldus Information Technology bv, Wageningen, TheNetherlands). All statistical tests are two tailed.

Interobserver Reliability

Interobserver reliability between two of us was deter-mined considering the proportion of agreements on totalfrequencies of behaviour (Caro et al. 1979). Average con-cordance between observers, calculated over seven feedingsessions conducted across the study time period, was 0.93for submissions and 0.77 for aggressions (corrected bychance as indicated in Martin & Bateson 1993). Averageconcordance for autogrooming behaviour, on a series ofeight focal animals, was 0.97 (corrected by chance). Auto-grooming was considered because it was displayedfrequently by each subject. Spearman correlation coeffi-cient between observers, over the same series of focal ani-mals, was rS ¼ 1.

RESULTS

Dominance Hierarchy in Absence of AnySources 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), probablydue to missing values. However, a significantly lineardominance hierarchy based on direction of submissive be-haviour 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 threeadult males, whereas three females were at the bottom(Table 1).

This rank order based on submissions was positivelycorrelated 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 malesand females did not differ significantly in aggressive be-haviour (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 Totp

LEO 1 1ANT 1 1 2 1 5SON 3 1 1 5LAV 8 1 1 2 12PIC 2 6 1 1 1 11PAL 2 5 1 2 1 11PEL 1 1 2RIG 5 69 23 7 1 1 1 2 109GNO 3 2 1 5 4 2 4 1 22RED 42 4 1 1 6 2 2 58CLA 19 2 2 5 2 1 21 52FIA 1 6 10 1 6 2 11 4 41SIL 2 3 1 6 5 4 10 31Totr 15 158 35 14 37 14 1 20 17 35 9 4 1

Bold type: females; standard type: males.Totp ¼ total submissions performed, Totr ¼ total submissions received.

BONANNI ET AL.: FEEDING ORDER IN DOMESTIC CATS 1373

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 seemedto affect the dominance relationships. Body weight waspositively correlated with rank (rS ¼ 0.693, N ¼ 10, P <0.03; Fig. 1): on average, males were 50% heavier than fe-males and the three top-ranking males were the heaviestcats in the group; similarly, the highest-ranking femalewas also the heaviest female. Head volume was also posi-tively correlated with rank (rS ¼ 0.783, N ¼ 9, P ¼ 0.013),whereas for arm length and canines length the correla-tions with rank failed to reach a statistically significantlevel (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 oldestmales being the highest in rank (correlation ageerank:

12345678910

Rank

2

3

4

5

6

Bod

y w

eigh

t (k

g)

Figure 1. Correlation between dominance rank and body weight for

10 adult cats of the studied group. C ¼males; B ¼ females.

rS ¼ 0.726, N ¼ 8, P < 0.05), but no relation existed be-tween age and rank considering all adult cats as a whole(rS ¼ 0.168, N ¼ 13, P ¼ 0.583).

Dominance Hierarchy in the Feeding Context

Contrary to what we found in the absence of food, thematrix based on aggressive interactions recorded within1 m of the food (Table 2) showed a highly significant levelof linearity (improved linearity test: h0 ¼ 0.567, P ¼0.0005), probably due to a higher number of interactions(N ¼ 565). A female (LAV) was at the top of this hierarchyand, altogether, all females became dominant over somemales from whom they were dominated in absence offood (see Table 2). All kittens were at the bottom positionsof the rank order (Table 2). It should be noted that threeadult cats (two males and one neutered female) were notconsidered in these analyses because they visited toorarely the feeding site before their disappearance fromthe study area.

Females showed the highest total level of aggressivebehaviour within 1 m from the food (mean rank ¼ 11),followed by males (mean rank ¼ 7) and by kittens (meanrank ¼ 4.75); nevertheless, the difference between thethree ageesex classes was not significant (KruskaleWallistest: H2 ¼ 4.614, P ¼ 0.10). Females were still the most ag-gressive class even restricting the analysis to periods whenthey were neither pregnant nor lactating and, indeed, thistime the difference was significant (KruskaleWallis test:H2 ¼ 7.831, P ¼ 0.02; mean ranks: females 12.25, kittens5.75, males 5.5).

A significantly linear hierarchy based on 307 submissiveinteractions (except hissing) recorded within 1 m of thefood was also found (improved linearity test: h0 ¼ 0.464,P < 0.009; Table 3). This feeding rank based on submissionpatterns was very similar to the previous one, based on ag-gressions, 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 wasjust 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 Totp

LAV 4 1 1 15 3 42 13 4 6 15 14 71 110 299ANT 2 1 3 6LEO 2 2 7 1 1 2 1 3 6 25PIC 2 1 3CLA 4 4 11 16 3 4 1 9 7 59PEL 2 2 1 5FIA 4 1 3 3 3 7 13 31 20 85SON 2 1 1 4 19 17 44RIG 1 1 1 3RED 1 1 1 3CAL 5 4 4 13NER 1 1 2SPO 1 1 15 17RUG 1 1Totr 13 5 1 3 17 9 66 37 16 16 26 34 141 181

Bold type: females; standard type: males; italic type: juveniles.Totp ¼ total aggressions performed, Totr ¼ total aggressions received.

ANIMAL BEHAVIOUR, 74, 51374

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 Totp

ANT 1 1LAV 2 4 1 7LEO 1 1PIC 1 1CLA 2 8 10PEL 2 2FIA 34 2 3 12 2 1 54SON 2 1 5 1 9RIG 2 4 2 2 1 6 17RED 2 1 1 2 1 7CAL 13 1 1 2 17NER 9 1 2 4 16SPO 36 2 7 1 6 5 1 3 61RUG 61 6 8 14 5 1 2 7 104Totr 8 169 15 3 43 5 27 18 2 1 9 0 7 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 ¼0.588, N ¼ 10, P ¼ 0.074).

Feeding Order

The total number of cats that were present around thefood at the beginning of a single session ranged from 2to 10 (X� SD ¼ 4:72� 1:85). For adult cats the probabil-ity of being the first to feed tended to decrease since Oc-tober 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 kittensregularly fed at feeding site, they were more likely thanadults 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; meanranks: kittens 9.5, females 6, males 4) and was signifi-cant in February 2002 (KruskaleWallis test: H2 ¼ 6.281,P ¼ 0.04; mean ranks: kittens 9.25, females 5.1, males4), when kittens were 4 and 6 month olds, respectively.Figure 2 shows that the statistical significant level wasreached probably because of the great difference be-tween kittens and adults, whereas the distance betweenfemales and males was smaller. Kittens’ feeding prioritywas not a consequence of the protection of their domi-nant mother: they were still more likely than adults tobe 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).

0

10

20

30

40

50

60

70

80

Aug Sep Oct Nov Dec Jan Feb Mar Apr May Jun

Months

Prob

abil

ity

of b

ein

g th

e fi

rst

to f

eed

JuvenilesFemalesMales

*

*

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 pres-

ences 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 1375

The feeding order (total probability of being the first tofeed) was not significantly correlated with feeding rankwhen considering all group members as a whole (rS ¼0.086, N ¼ 14, P ¼ 0.770 and rS ¼ 0.099, N ¼ 14, P ¼0.735, for rank based on submissions and aggressions, re-spectively); however, for adult cats only, it was moder-ately, although not significantly, correlated with bothrank (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 SocialTolerance to Juveniles

In this study we have found that intersexual dominancerelationships in the cat group varied according to differentcompetitive contexts and to a minor extent to thebehaviour analysed: male cats were usually dominantaway from food, whereas females increased in rank atthe feeding site, especially when the feeding rank assess-ment was based on aggressive behaviour that is likely toreflect motivation (Hurd 2006). Moreover, although juve-niles were at the bottom of the feeding hierarchy, theywere significantly more likely than adults of both sexesto be the first to feed around the age of 4e6 months, indi-cating that they received a certain level of tolerance (lackof significance after the age of 6 months was probably dueto a decrease in the total number of cats present in thestudy area). Social tolerance to kittens explains the totallack of correlation between feeding rank and feeding orderthat we have found considering the cat group as a whole,since the level of the correlation became moderate oncethe kittens were excluded from the analysis.

Yamane et al. (1997), who investigated the feeding orderin a cat colony living in a fishing village in Japan, founda general tendency for males to feed before females andkittens before adults of both sexes, although they didnot provide data on agonistic interactions, making it diffi-cult to compare their results with ours. On the other hand,Knowles et al. (2004) found results similar to those of thepresent study (females gaining rank position in feedingcompetition), although these authors studied a neuteredcat colony living in a closed environment.

The present work is, to our knowledge, one of the fewstudies documenting female dominance over considerablylarger 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 basedon direction of agonistic behaviour in dyadic conflicts,that is female dominance over larger males was notacquired through coalition formation, which results inan increase of RHP, as occurs in several primate species(e.g. bonobos, Pan paniscus, Furuichi 1989; patas mon-keys, Erythrocebus patas, rhesus monkeys, Kappeler 1993).The results shown here seem to provide support forgame-theory models predicting conflict outcome in favourof the smaller competitor, when asymmetries in resource

value and in the cost of winning inappropriately maycompensate for the smaller competitor’s RHP. We assumethat conflicts away from food are for general dominanceand the value of winning is roughly similar for both sexes.Here the outcome is, thus, mainly affected by asymmetriesin RHP with male cats being in advantage over females. Inaccordance with studies conducted on other species (e.g.hens, Gallus gallus domesticus, Cloutier & Newberry 2000;red deer, Veiberg et al. 2004; brown bears, Gende & Quinn2004; fallow deer, Dama dama, Jennings et al. 2006), dom-inance rank (or RHP) was positively correlated with bodyweight and size in our cat colony. Head size, in particular,may be used by cats as a cue to assess each other’s fightingability.

Conversely, it seems reasonable to assume that femalecats value the food more highly than males so as tocompensate for their lower RHP. In fact, reproduction canplace a particularly high level of stress on female domesticcats: it has been shown that their energy intake increasesup to 1.7 times maintenance levels during pregnancy andup to 3.5 times at peak lactation, when large litters arereared (Loveridge 1986), whereas a tom 1.5 times as heavyas a female should have an energy intake just 1.35 timesmaintenance levels of a female. However, in this study fe-male cats were more aggressive than males even duringnonreproductive periods. To explain this pattern, weshould consider that females are typically pregnant or lac-tating during the period JanuaryeOctober, that is for themajority of the year, and it is possible that, once estab-lished, dominance relationships may subsequently con-tinue to influence the decisions to escalate feedingconflicts.

Several authors have stressed the importance of leveragein affecting payoff asymmetries in animal contests (Hand1986; Smuts 1987; Noe et al. 1991; Kappeler 1993; Lewis2002). It has been argued that the smaller competitorcan become dominant over the larger one because of a le-verage advantage: although the female dominance ob-served among domestic cats in this study and in somemammalian species where sexes do not differ markedlyin 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 highlythan males, it is also possible that, in some of these spe-cies, males may suffer a leverage cost resulting from win-ning inappropriately a feeding conflict against females.In spotted hyenas, where females exercise a strong degreeof mate choice, male food deference may serve to affect fe-male preferences (East & Hofer 2002). In lemurs, malefood deference may enable females to meet unusuallyhigh energetic demands of reproduction and eventuallyto successfully wean infants (Kappeler 1993; Pochronet al. 2003). Similarly, for male domestic cats it might beconvenient to allow both female dominance and juve-niles’ feeding priority to occur. Even if male domesticcats play no direct part in rearing their kittens, they couldcontribute to increase kittens’ survival by tolerating bothfemales and kittens in feeding situation. A leverage cost

ANIMAL BEHAVIOUR, 74, 51376

must be necessarily involved in cases of food deference tojuveniles by adults since asymmetries in RHP betweenadults and juveniles appear to be too extreme to be over-come by asymmetries in resource value. Since to denyfood to a given offspring should be more costly to malesthat invest heavily in few offspring, a correlation couldbe expected between the degree of paternal investmentand this type of male tolerance (Hand 1986). However,a more general condition may be that tolerance occurswhenever winning elevates costs for the larger competitorbeyond a certain critical value (Hand 1986). Male fooddeference to females can be found in association with ju-veniles’ feeding priority in some primate species whereinmales invest considerably in parental care (e.g. commonmarmosets, Callithrix jacchus; golden lion tamarins, Leon-topithecus rosalia; titi monkeys, Callicebus moloch; indri,Indri indri, all reviewed in Hrdy 1981), as well as in a coop-erative 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 do-mestic cat, both the types of tolerance may be part of a sin-gle male reproductive strategy. In domestic cats, such maletolerance could have evolved in the original environmentof adaptation in which, since males are expected to mo-nopolize the access to oestrous females living within theboundaries of their own territory (Say et al. 1999; Liberget al. 2000), there are high probabilities that the femalesencountered are carrying or nursing the males’ offspring.On the other hand, male tolerance may be a consequenceof the high level of paternal uncertainty resulting from thepromiscuous mating system found among domestic catsin the urban environment (Natoli & De Vito 1991; Sayet al. 1999; Natoli et al. 2005).

In a similar way, female tolerance of kittens may bea consequence of females having a weak capacity todiscriminate their own offspring from those of otherfemales, given that for a female cat living solitarily in theoriginal environment of adaptation the evolution of a finelytuned parenteoffspring recognition is not necessary.

Comparison with the Dominance Systemof Lions

The behaviour of domestic cats and lions has beencompared 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 havebeen proven to be able to live socially. Their social behav-iour shows some striking similarities. First, both lions(Schaller 1972; Packer et al. 1991) and domestic cats (Lib-erg et al. 2000; Macdonald et al. 2000) form social groupscomposed of genetically related females, their dependentoffspring and attached adult males. Second, both lionesses(Pusey & Packer 1994a) and female domestic cats (Mac-donald et al. 1987) show alloparental care of juvenilesand cooperation in territorial defence.

However, the feeding competition pattern and thedominance system observed in this study among domesticcats were quite different from those found in lions. Unlikemale domestic cats of our social group, which were less

aggressive than females at feeding sites and allowedkittens’ feeding priority, male lions dominate all otherageesex classes at kill, routinely supplanting both femalesand cubs (Packer et al. 2001). Even if male lions have beenshown to be effective in hunting large prey (Funston et al.1998), they can acquire the bulk of their food intake byappropriating kills made by lionesses (Van Orsdol 1986).In contrast to males, lionesses usually do not displace ju-veniles from a specific feeding site around a carcass (Packeret al. 2001).

To explain such differences we should consider thedifferent reproductive tactics followed by these two felinespecies. Male lions form cooperative coalitions whichcompete against other coalitions for exclusive possessionof a female pride (Bygott et al. 1979; Packer et al. 1988;Grinnell et al. 1995). As demonstrated by paternity analy-sis, a resident coalition is very effective in preventing ex-tra-pride males from fathering cubs (Packer et al. 1991)and, consequently, the possession of a pride of femalesis the only chance for a male lion to gain mating opportu-nities. The male group tenure is retained for a variable pe-riod and typically ends when they are displaced from theirpride by a new group of males (Bygott et al. 1979; Packeret al. 1988). Incoming males accelerate the females’ returnto oestrus by killing unweaned cubs and evicting subadults(Pusey & Packer 1994b), and thus prolonged residence is es-sential for successful reproduction. Male reproductive suc-cess increases with coalition size, mostly because largercoalitions maintain residence for longer periods, thusfathering more cubs and postponing the next episode ofinfanticide (Bygott et al. 1979; Packer et al. 1988). We hy-pothesize that selective pressures favouring male domi-nance and aggressiveness at feeding occasions would beparticularly strong in lions, because male lions need tomaintain constantly a peak physical condition so asto ward off rivals, to prolong their tenure in a pride andto protect their cubs from the risk of infanticide. Paradox-ically, if male lions were more tolerant to cubs, they couldnot be nourished enough to protect them from infantici-dal rivals. Recent findings seem to support our hypothesis:dark-maned lions, which have higher levels of testoster-one and hence of aggressiveness, are better nourishedand enjoy both longer tenures and higher offspring sur-vival (West & Packer 2002).

Furthermore, among lions, females would not probablyhave a leverage advantage over males: lionesses them-selves, and not just males, may incur a leverage cost bywinning a feeding conflict against a competitor of theopposite sex, because they probably need male help inprotecting offspring from infanticide.

These arguments do not apply to domestic cats. Maledomestic cats are not known to form coalitions whichcooperate to take-over a group of females. Also, defence offemales is not the only mating tactic available to them:even in rural populations, where the mating system ispolygyny, territorial males are not completely successfulin preventing transient males from copulating withfemales (Say et al. 1999). In addition, infanticide has notbeen documented in urban domestic cats and rarely docu-mented 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 1377

advantageous as a male reproductive tactic in species that,like lions, do not breed seasonally and show a very longinterbirth interval (Hausfater et al. 1982; Packer & Pusey1984). Unlike lions, domestic cats are seasonal breederswith a short interbirth interval (Schmidt et al. 1983):a mated female domestic cat is unavailable to other malesfor about 4 months from the beginning of oestrus to theweaning of kittens; furthermore, in the present study, a fe-male cat came back into oestrus and performed fertile mat-ings before the weaning of her current litter. Thus, fora male domestic cat infanticide is not crucial for speedingup the reproductive cycle of females, whereas for a malelion it is the only chance to bring a female back into oes-trus in reasonable time (Natoli 1990). In conclusion, webelieve that male domestic cats do not behave aggressivelyat feeding site in the same way as lions, or do not value thefood in the same way, because they do not need to main-tain constantly a peak physical condition in order to re-tain exclusive control of a group of females and todefend their offspring from infanticidal rivals.

Domestication

Domestication is thought to have led to a decrease inintraspecific aggression and loss of social inhibition ina variety of species, mostly because of artificial selectionand relaxation of natural selective pressures under humaninfluence which has provided animals with abundantresources (Price 1984). It has been suggested that thesame applies to domestic cats because they apparentlyshow greater sociability and behavioural flexibility thantheir wild solitary ancestor, the African wild cat (Macdon-ald et al. 2000). Thus, it may be thought that the social tol-erance displayed, in this study, by male domestic cats toboth females and kittens is a by-product of domestication,which would have caused a decrease in male aggressive-ness, rather than the result of natural selection mecha-nisms. However, in our opinion, it seems unlikely thatdomestication would have caused a differential decreasein aggression in the two sexes and differentially in differ-ent competitive contexts. Given that our results seem tobe in accordance with predictions of game-theory models,they are more likely to reflect selective processes possiblyoperating in the original environment of adaptation be-fore domestication and urbanization.

Moreover, evolution of sociality does not necessarilyimply a reduction in intraspecific aggression: studies con-ducted on social insects indicate that it involves evolutionof conditional aggressive behaviour rather than uncondi-tional intraspecific tolerance (Cahan 2001).

Indeed, whether or not the evolution of sociality in catshas been promoted by domestication is still a debatablepoint: feral domestic cats live solitarily when dependingon widely dispersed natural prey, whereas they form socialgroups in the presence of clumped, rich food resourcesprovided by human beings (Liberg et al. 2000); even iffragmentary evidence indicates that African wild cats donot form colonies where they are exposed to abundanthuman refuse (Macdonald et al. 2000), it is intriguingthat, in captivity, female African wild cats have assisted

mothers in provisioning of kittens with food (Nowell &Jackson 1996), a behaviour observed in feral domesticcats colonies, but not in any other solitary cat species.Domestic cats do not necessarily have a greater level ofbehavioural flexibility than their wild ancestor, but theymay have simply been exposed to a far greater diversityof ecological conditions (Leyhausen 1988).

Acknowledgments

Special thanks are due to Antonio Buogo for his availabil-ity to all our requests during the study, to all veterinariansworking at the Veterinary Hospital of Rome for their helpin handling cats and, in particular, to Giuseppe Cariola forhis unvaluable help in trapping cats, anaesthetizing themand collecting blood samples. We wish to thank MasakoIzawa for procuring relevant literature, Alessandro Giu-liani for valuable suggestions, Marion L. East for helpingwith the figures and the referees for their valuablecomments that helped us to greatly improve the manu-script. Finally, a special thanks goes to Prof. AlbertoFanfani from the Department of Animal and HumanBiology, University of Rome ‘La Sapienza’, who offeredsupport and facilities.

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