Chapter 5: the Signalling Repertoire of the Domestic Cat and Its

5 The signalling repertoire ot the domestic cat and its undomesticatedrelatives

JOHN BRADSHAW AND CHARLOTTE CAMERON-BEAUMONT III: Social life

Introduction benefit by cooperation, they need the ability to resolve conflicts without resorting to physical Previous accounts of communication between violence, particularly when both protagonists are as domestic cats (e.g. Bradshaw, 1992) have been largely well-armedasa cat. It is not yet certain when this based on a traditional ethological approach. The ability arose, since the social biology of F. libyca has signals and the context in which they occur have been been little studied, but in the second part of this described, and related to the kind of environment chapter we have attempted to examine the extent to signaller and receiver can expect to find themselvesin, which domestication has influenced the signalling and to the sensory capabilities of the receiver. For repertoire of the domestic cat, by comparing it with example, this approach explainstheuseofscentsig- that of other, undomesticated, Felidae. Inthe first part nals by domestic cats as products of both their acute we describe the signals performed by the domestic cat sense of smell, which may have evolved primarily in itself, and their presumed functions. relation to detection of food,andalso their origin as territorial animals which needed to communciate with neighbours that they might rarely encounter Communication between domestic cats face-to-face. However, modern biological signalling Olfactory theory is equally concerned with what informationis communication being transferred and howit is transmitted (Grafen & The ancestral species of the domesticcat, F. s. libyca,is Johnstone, 1993). More specifically, it examines how probably exclusively territorial (Smithers, 1983; signals can becomeevolutionarily stable, given that Happold, 1987; Macdonald, 1996), as are most of the the interests of emitter and recipient are often not smaller species in the Felidae. Since widely-spaced identical. animals rarely encounter one another face-to-face, Communication is said to occur when one animal they tend to communicate by scent-marks, which respondstothe signals sent out by another. Thisis a permit a delay of several hours or days between the more general definition than normally applies to com- deposition of the signal andits reception. For well- munication between people, when it is usually armed carnivores, there is also the advantage that assumed that information is being exchanged,and is potentially dangerous encounters with rivals can be reasonably accurate. Unfortunately there has been a avoided by the use of olfactory signals, both those tendency to carry this ‘conventional’ definition over deposited on the substratum and those that are carried to communication between animals, implying that directly from the body surface by air currents. The animals that are signalling to one another agree about potential disadvantage of relying on scentsignals is the message being transmitted (Zahavi, 1993). In lack of control, both of the direction the message is manyinstancesthere is no reason to believe thatthis is carried in, which is at the mercy of the wind, and of the case; signallers often attempt to manipulate the whoreceivesit, since a scent-mark cannot be switched behaviourofrecipients to their own advantage, while off at will; both lead to potential exploitation of the recipients attempt to ‘mind-read’ these deceptions information that the scent contains. Despite these (Krebs & Dawkins, 1984). This kind of theoretical problems, membersof the Carnivorarely extensively framework has hardly ever been applied to signalling upon scent for communication (Gorman & in the domestic cat; in this chapter we have attempted Trowbridge, 1989). to speculate as to the evolutionary origins of some Many domesticcatslive at a density several orders signals, such as the odour of tom-cat urine, purring, of magnitude higher than their wild counterparts and agonistic visual signals. (see Chapters 6 and 7), and it is therefore possible The influence of domestication on signalling adds that their scent communication has been modified a further dimensionto the explanation of whysignals during the course of domestication. Cats thatlive in take the form they do. In the case of the cat, the groups can potentially not only exchange information ancestral species Felis silvestris libyca is thought to be through scents, but also exchange the scents them- exclusively territorial, and so its signalling repertoire selves to produce colony- or group-specific odours must presumably have changedas it evolved to live (Gorman & Trowbridge, 1989). Comparisons with at high densities and to becomefacultatively sociable. other species therefore suggest that the domestic cat When individual animals live close together, and should have a complex and versatile repertoire of Signalling repertoire 69 and other disulphides scent signals, so it is perhaps surprising that com- 3-methylthio-1-butanol (III), odours(Joulain paratively little research has been conducted in this and trisulphides, have strong ‘tom-cat’ Entire males area. While several sources of odours have been & Laurent, 1989; Hendricks et al., 1995a). offelinine, up to 95 mg/day, documented, their functions in communication are can excrete large amounts 20 mg/day, generally still speculative. whereas females produceless, up to about which correlates with the lesser pungency of female have suggested Urine sprayed urine. Hendrickset al. (1995b) effect on Cats can adopt two distinctly different postures for that this excretion may have a significant of urination, indicating that at least one has someusein the sulphur-containing amino-acid requirements signalling. Kittens, juveniles and adult females usually an entire male, since felinine is biosynthesised from squatto urinate and then usually cover the urine with cysteine and possibly taurine.It is therefore possible soil or litter. Although this can be interpreted as an that the amount offelinine in the urine, and hence the attempt to hide the urine, and so presumably the strength of its odour, is an accurate reflection on the information that its odour contains, such deposits success of the male in obtaining high-quality food, and are sniffed by other cats if encountered. Moreover, is therefore an ‘honest’ signal (Zahavi & Zahavi, 1997) the duration of sniffing tends to increase with the advertising his fitness as a mate (to females) and com- unfamiliarity of the depositor, suggesting that the petitor (to other males). sniffer is responding to and gathering information The territorial function of urine-spraying,if any,is from the odour (Passanisi & Macdonald, 1990). This unclear. Spray-marks are rarely observed to actas a may only be a common occurrence where cats are deterrent in their own right, but this is the case for living at high densities; the attempted concealment most territorial scent-marks (Gosling, 1982), even maybeeffective in widely-spaced territories. those which markthe edgesof territories, which those Deliberate scent-marking with urine is performed of tom-cats do not (Feldman, 1994a). It has also been by spraying, in which the cat backs up to a vertical suggested that since the odourof scent-marks changes surface, and urinates backwards, usually while quiv- with age, they could be used to assist cats to space ering its tail. While mature males are the most themselves out while hunting, so that they could frequent sprayers, adult females do also spray. In avoid areas which had been disturbed recently closed or high-density colonies there may be some (Leyhausen, 1979). However, this is unlikely to be a suppression of spraying in females and younger stable strategy; cats that did not spray-urinate could males, resulting in most spray-marks being produced put themselves at an advantage because other cats by a small numberof ‘dominant’ males (Natoli, 1985; would waste time and effort hunting in places where Feldman, 1994a). Spraying by tom-cats is enhanced prey was still wary due to the recent proximity of a by the proximity of oestrous females, resulting in predator. an annual peak (in the UK) in February/March All cats, but particularly adult males, investigate (Feldman,1994a). spray-marks intently (Natoli, 1985; Matter, 1987; The odourof sprayed urine is pungent, prompting Passanisi & Macdonald, 1990), particularly if they are speculation that it carries other secretions, possibly produced by oestrous females (Verberne & de Boer, from the preputial or anal glands (Wolski, 1982). The 1976) which suggests that they do contain relevant anal gland secretion, which is voided by very fright- information.Initial inspection is usually by sniffing, ened cats, certainly has a distinctive odour, butthis is often followed by flehmen, in which the upperlip is not, to the humannose,similarto that of sprayedurine. raised and the mouth held partially open; this may The odour of sprayed urine increases after deposition persist for half a minute or more. During flehmen the (Joulain & Laurent, 1989), and is probably largely cat may makephysical contact with the source of the due to the microbial and oxidative degradation of the odour, and movesits tongue to and fro behindits two unusual amino-acids which it contains, felinine incisors, where the openings of the ducts that lead to (L-2-amino-7-hydroxy-5,5-dimethyl-4-thiaheptanoic the vomeronasal organs (VNO)lie. Both airborne and acid, I) and isovalthene (2-amino-5-carboxy-6- fluid-borne molecules of the odorant are

to odours from othercats,it presumably gathers (and or whetherthere is considerable overlap. The secre- possiblystores) social information. tions of the glands on the head are rubbed on to prominent objects by a behaviour pattern known as Faeces bunting (Houpt & Wolski, 1982). The precise form of Manyspecies within the Carnivora use faeces, often this appears to depend upon the height of the object with glandularsecretions added, to convey informa- being rubbed, such that high objects are primarily tion (Gorman & Trowbridge, 1989), but the evidence marked with foreheadandears, objects at head height that domestic cats do this is only circumstantial. Near with a wipe ofthe head from the cornerof the mouth to the core of the home range, faeces are usually to the ear, and lower objects with the underside of the buried (Feldman 1994a), but they may beleft exposed chin and thenthe side of the throat (Verberne & de elsewhere (Macdonald et al., 1987). Cats usually sniff Boer, 1976). This plasticity suggests that similar the places where they have just buried faeces, but tend odoursare deposited from all parts of the head,either notto dosoafter leaving them exposed (Macdonald et because there is redundancy between the glandular al., 1987). This suggests that one of the functions of secretions themselves, or because they become tho- burying faeces is to minimise thelikelihoodthat the roughly mixed on the coat through grooming. olfactory information they contain will be detected Entire adult males tend to rub-mark morefre- by another cat, although hygiene may provide a more quently than do anoestrous females or juveniles parsimonious explanation. Attempts to demonstrate (Feldman,1994a) and occasionally spray urine on top that unburied faeces serve as territorial markers have of their own rub-marks (Dards, 1979; Panaman,1981 ) produced equivocal results (Dards, 1979; Macdonald or vice versa (Macdonald et al, 1987). Other rub- et al., 1987; Feldman, 1994a). marks, although performed on visually prominent objects, such as projecting twigs or corners of Scratching man-made structures, are not associated with any Although it undoubtedly has a role to play in the other visual or obviousolfactory cue and are thus not conditioningof the clawsofthe frontfeet, scratching obvious to the human observer. Cats, on the other must inevitably result in the deposition of scent hand,appearto be able to locate them easily, suggest- from the glands on the paws(interdigital glands) ing that they are quite pungent to the feline nose, (Ewer, 1973). The samescratchingsite is often used and frequently over-mark them with their own over and overagain,resultingin a clear visual marker cephalic secretions. The rub-marksof entire females which presumably drawsattention to the olfactory contain information aboutthe oestrus cycle, as indi- information, although there appear to be no pub- cated by the degree of interest shown by males lished studies which report the extent to which (Verberne & de Boer, 1976), but apart from this there scratched sites are sniffed. The scratching sites are is little published information on the function ofthis distributed along regularly-used routes, rather than behaviour. Some cats also rub-mark repeatedly in at the periphery of the territory or home-range the vicinity of humans, but this may possibly be a (Feldman,1994a). displaced version of cat-human rubbing (Moore & Stuttard, 1979). Skin glands Cat—cat rubbingis a visual and tactile display which Domesticcats haveseveral skin glands(Prescott, cited mustalso result in the exchange of odours between in Fox, 1974); in addition to the interdigital glands the pelages of the participating cats, although it is mentioned above, these include; the submandibular unclear whetherthis has any relevance, for example in gland beneath the chin, the perioral glands at the the establishment of ‘group odours’ shared by cats corners of the mouth, temporal glands on eachside that are friendly towards one another. When cats sniff of the forehead, a gland at the baseofthetail (which each other, they tend to concentrate on the head can over-secrete in entire males, giving rise to the region, rather than the flanks and tail where shared condition ‘stud-tail’), and caudal glands, which are odours would presumably accumulate, suggesting diffusely distributed along the tail (Wolski, 1982). The that even if group odoursdoexist, individual odours pinnae(externalears) also produce a waxysecretion. contain morevaluable information. It is unclear whether each of these glands produces a uniquesecretion, each with a well-defined function, ~Signalling repertoire 71

(Haskins, 1979). The call induced by cold is signifi- Auditory communication cantly higher pitched than the other two, although Cats’ vocalisations are largely restricted to four types this distinction disappears as the kitten becomes of interactions; agonistic, sexual, mother—young, and capable of thermoregulation at about four weeks of cat-human. Most of the aggressive and defensive age. Restraint inducesa call whichis similar in pitch to sounds(Table 5.1) are strained-intensity calls (Moelk, that caused byisolation, but is significantly longerin 1944), since under these circumstances the cat is duration, and theisolationcall is generally the loudest likely to be tensing its whole bodyin preparation for a (Haskins, 1979). It is therefore likely that mother fight. Tensionin the throat is presumably the reason cats can distinguish betweenthese calls, and respond whycats drool during fights, or have to break off from accordingly (Haskins, 1977) . vocalising to swallow repeatedly. The low pitch of the Purringis a ubiquitous vocalisation amongcats, but growl and the long duration of the yowl are presum- its function is not entirely understood and, until ably designed to conveythe size and strengthof the cat recently, its method of production was not entirely that is emitting them, and the abruptness and volume clear. It is produced during both inhalation andexhal- of the pain shriek may be designed to shock orstartle ation, except for a brief pause at the transition the attacker into loosening its grip. Both the female betweenthe phasesof the respiration cycle, and there- and male sexualcalls (Table 5.1) are also of high inten- fore soundsasif it is a continuous vocalisation. The sity, presumably advertising fitness to potential sexual sound is generated by a sudden build-up andrelease partnersandrivals of the same sex (see Chapter7). of pressure as the glottis is closed and then opened, The calls produced by kittens less than three weeks resulting in a sudden separation of the vocal folds, old are restricted to the defensivespit, purring, and a which generate the sound (Remmers & Gautier, distress call which has aural characteristics similar to 1972). The laryngeal muscles which movetheglottis the adult miaow (see Figure 5.1). The latter is given are driven by a free-running neural oscillator, gener- when the kitten becomesisolated, or cold, or trapped, ating a cycle of contraction and release every 30-40 for example,if its mother accidentally lies on top ofit milliseconds (Frazer-Sissom, Rice & Peters, 1991).

Table 5.1. Characteristics ofthe vocalsignals used by adult domestic cats, compiledfrom Moelk (1944), Brownetal. (1978) and Kiley-Worthington (1984), and the circumstances under which each is most commonly used.

Name Typical Fundamental Pitch change Circumstances duration(s) pitch (Hz) Sounds produced with the mouth closed Purr 2+ 25-30 ~ Contact Trill/chirrup (F)* 0.4-0.7 250-800 Rising Greeting, kitten contact Sounds produced while the mouthis open and gradually closed Miaow(B) 0.5-1.5 700-800 — Greeting Femalecall 0.5-1.5 ? Variable Sexual Mowl(malecall) ? ? Variable Sexual Howl (D) 0.8-1.5 700 - Aggressive Soundsproduced while the mouthis held open in oneposition Growl 0.54 100-225 — Aggressive Yowl (D) 3-10 200-600 Rising Aggressive Snarl 0.5-0.8 225-250 - Aggressive Hiss (E) 0.6-1.0 Atonal — Defensive Spit 0.02 Atonal — Defensive Pain shriek (C) 1-2.5 900 Slightrise Fear/pain

“Refers to Table 1 of Brownet al. (1978).

72 IIT: Social life

(ii) Figure 5.1. Sonagraphsoftypical | 4kHz kitten and cat vocalisations. (1) Kitten isolation call. (ii) Maternal chirrup. (iii) Miaow (typical). (1v) Miaow(atypical). (v) Howl. (v1) Hiss. (vii) Pain shriek.(iii) and (iv) provided byJean-Luc Renck; others from Brownetal. (1978).

(iv) (v)

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Although it is traditional to interpret purring as response, based upon its effectiveness in getting indicating ‘pleasure’, it is produced in a widevariety humanattention.It is certainly very easy to train in of circumstances, most of which involve contact food-deprived cats; Farley et al. (1992) were able to between the cat and a person or anothercat. Kittens induce a rate of two miaowsper minute for a period are able to purr almost from birth, and do so primarily of two hours or more. There are also considerable whenthey are suckling, which may induce the mother variations in frequency, duration and form of the to continue to nurse them (Haskins, 1977). Adult cats miaow, both within and between individuals (Figure may purr when in contact with a familiar partner, and 5.1. 1, rv) (Moelk, 1944) which argue against the during tactile stimulation with inanimate objects, miaow havingan (intra)species-specific meaning.Itis such as whenrolling or rubbing (Kiley-Worthington, thereforelikely that each cat learns by simple associa- 1984). All of these circumstances can be conceived of tion that miaowing inducesfeeding,access to desired as potentially pleasurable to the cat, but there is one locations, and other resources provided by humans, serious exception to this: veterinarians commonly and that some cats can learn to produce different experiencecats that purr continuously whenthey are miaowsfor different purposes. chronicallyill or appearto be in severe pain (Beaver, 1992). Purring maytherefore function as a ‘manipula- tive’ contact- and care-soliciting signal, possibly Visual communication derived from its (presumed) functionin the neonate. Wild-type (striped tabby) domestic cats are cryptical- Apart from purring, the vocalisation that is com- ly marked, and have no obviousstructures that have monest in cat—humaninteractions is the miaow. This been specially adapted for signalling. Despite its is very rarely heard during cat—cat interactions relatively immobileflat face, compared with the wolf, (Brown, 1993) and may therefore be a learned the cat has quite a varied repertoire ofvisualsignals,

Signalling repertoire 73 a cat that wishes to mainlyused in regulating aggressive behaviour. There stand atits full height, whereas ouch on the ground, is no evidenceto suggest that any of the changesto the withdraw from a contest will cr raw its head into pelage introduced post-domestication (e.g. orange, flatten its ears (Figure 5.2), and withd t ready to launcha white spotting, long hair) have had any substantial its shoulders, indicatingthatit is no ve-aggressive effect upon ability to signal, in contrast to the pro- biting attack (Figure 5.3). The defensi resented found loss of visual signalling structures in some posture (bottom right of Figure 5.3) is p s attack breeds of dog (Goodwin, Bradshaw & Wickens, when the aggressor is aboutto press homeit 1997). (and also to potential predators such as dogs). Thisis Manyofthe postures adopted in agonistic encoun- usually adopted side-on to the opponent, doubtless to ters can be interpreted as attempts bythecatto alter maximiseits visual impact. Although more extreme,it its apparentsize, and thereby influence the outcome is similar in form to the ‘Side-step’ posture used by of the interaction. An aggressivecat will piloerect and kittens in play; since this posture tends to disrupt

EARS BACK AND FLAT EARS FORWARD AND ERECT

EARS BACK AND ERECT EARS FLAT

Figure 5.2. Ear postures associated with aggression and defence. From UK Cat Behaviour Working Group (1995). /4 IIT: Social life

Figure 5.3. Whole-body postures associated with aggression (increasing from left to right) and fear/submission (increasing from top to bottom). Redrawn from Leyhausen (1979).

bouts of social play (West, 1974),it is likely that one is or signals, the amountof time that the two cats looked the developmental antecedentof the other. at each other simultaneously (mutual gaze) was less Presumably all of these postures are interpreted than predicted from the total amountoftime that each by the cat’s opponent, and used in deciding how to spent looking at the other. In other words, each cat proceedin the encounter, butthereislittle direct evid- monitored the position of the other, but tended to ence as to how each posture influences its outcome. look away before being looked at: in these circum- Competitive encounters between animals of the same stances, mutual gaze may be being interpreted as a species tend to involve signals which are both unsub- threat signal. In encounters with no agonistic content, tle, and aimed at manipulating the behaviour of the the amountof mutual gaze was notdifferent from that recipient, which should attempt to combat this by predicted from the amountof time that each looked at ‘mind-reading’ (Krebs & Dawkins, 1984). The agon- the other, and so may notbe beingused a signal. istic displays of cats are certainly easy to see, but the Rolling is a component of female sexual (pro- extent to which each postureis a form of ‘bluffing’, estrus) behaviour, where it is usually accompanied and how effective each is at deceiving its recipient, by purring, stretching and rhythmic opening and remain to be investigated. closing of the claws, and is interspersed with bouts of In the preliminary stages of agonistic encounters, object-rubbing (Michael, 1961). Male-to-male rolling cats tend to avoid looking at one another. In a study appears to be a form of submissive or appeasement of staged 4-minute pairwise encounters between behaviour, since it is never directed by mature males neuteredcats from the same colony, D. Goodwin and towards immature males, and is often followed by the J. Bradshaw (unpublished data) recorded that each mature male ignoring or tolerating the immature cat looked at the other 1.8 times per minute on aver- male’s presence (Feldman, 1994b). age. In encounters that involved agonistic behaviour Thecat’s highly mobile tail, with its independently Signalling repertoire 7) movable tip, appears admirably suitable for use as a signalling organ as wellas assisting in balance. Thetail is tucked away between the hind legs in the submissive/defensive posture (bottom left of Figure 5.3), but this is unlikely to convey much informationthatis not already provided by the postureitself. Lashingof thetail from sideto side is a componentof aggressive behaviour (Kiley-Worthington, 1976), but its value as a signal is unknown. The vertically-held tail (tail-up, TU) is associated with affiliative behaviour (Brown, 1993; Bernstein & Strack, 1996), but its function as a signal has only recently been elucidated. In a colony of neutered feral cats, Cameron-Beaumont(1997) found that TU was Tail Up Othertail positions particularly associated with rubbing on and sniffing of another colony member (TU occurred in more

than 80 per cent of these interactions). Almostall Frequency/dyad bouts of cat—cat rubbing were preceded bythe initiat- ing cat approachingwithits tail up, and the probability of the rubbing occurring was further enhancedif the recipient cat also raised its tail (Figure 5.4). She confirmedthe role of TU as signal, and not simply a correlate, of affiliative behaviour, by presenting pet cats with silhouettes identical apart from the position of the ‘tail’. The TU silhouette (Figure 5.5) was significantly more likely to induce TU whenit wasfirst ; sighted by the responding cat, and was also approached faster than the silhouette with its tail A down, which induced sometail-swishing or tail- tucked postures. The verticaltail therefore signals an Tail Up Othertail positions intention to interact amicably; presumablyit is neces- 5.4. Association between the Tail Up posture by the potentially dire consequences of Figure sary because of the initiator (upper graph) and recipient (lower graph), and rubbing being approached by a cat whose intentions are (Ml) and othertypes of interaction (A), comparedtoal] othertail unknown. postures (the rare Tail Half-Up posture is omitted). Only Tail Upapproachesbytheinitiatorare includedin the lower graph. Frequencies are averages per dyad ina free-ranging neutered Tactile communication colony (2 male, 3 female) during 34 hours of observation. From Cameron-Beaumont(1997). Although simple physical contact, as when two cats rest together, may have social significance, the two most obvious forms of tactile communication are cat-cat rubbing their heads, flanks or tails on one (a) from adult females to the male, (b) within adult another (allorubbing), and one cat licking another females, (c) from kittens to adult females (Figure 5.6). (allogrooming). Asymmetryin the flow of rubbing within dyads was Even though Macdonald et al. (1987) proposed also detected by Brown (1993) among neuteredferal that ‘cats in net receipt of rubbing would enjoy the cats. She also foundthat interactions involvingsitting benefits of dominance and, within their sex, greater together and allogrooming were unlikely to be pre- inclusivefitness’, little evidence has been forthcoming ceded (or followed) by rubbing, which supports the subsequently to confirm or refute this. In a breeding suggestion of Macdonald et al. (1987) that rubbing farm colony, they foundthat the flow of rubbing was tends to take place between cats of unequalsize or asymmetrical in the majority of dyads, being skewed status. Further research is needed to fully elucidate

/6 IIT: Social life

Figure 5.5. Cat-sized silhouette used to investigate the Figure 5.6. Frequencies of cat—-cat rubbingin a farm colony of signalling function of the TU posture. Thesilhouette used for five cats, comprisinga female, her two adult daughters, an adult comparisonhadits tail sloped down towardsthe ground, with male, and a male kitten. Widths of arrowsare proportional to its tip horizontal. the square rootof the numberof rubbinginteractions per- formedover an 8-monthperiod (6 monthsfor the kitten) by each age/sex-class towardsevery other, and within the female the social meaning of rubbing, including whetherthe sex class. Actual numbersof interactionsare indicated for the largest, smallest and one intermediate arrow. All transfer of scent that must inevitably take place has pairs of arrows indicate a significantly assymetric performance of rubbing, any significance. except that between the Female and Daughter 1. Data from While grooming of one member of a social group Macdonaldet al. (1987). by anotherhassignificance in manyspecies (Wilson, 1975) itis only recently that Ruud van den Bos (1998) has begunto elucidate its role in the domestic cat. In Functional organisation of signals between an indoor colonyconsisting of 14 neutered males and domestic cats 11 neutered females, the more aggressive individuals groomedtheless aggressive moreoften than the other Various techniques have been used to combine way around. In about one-third of the interactions, communicative patterns together into groups with groomers were also aggressive towards the cats they overlapping functions, including subjective methods were grooming, often immediately after the bout of (Kerby, 1987), differences between pairwise rela- grooming had finished. These results are consistent tionships (van den Bos & de Vries, 1996) and with the ideathat allogrooming in the domestic cat is a probability of performance by an individual cat form of redirected aggression or dominance behav- within a single interaction (Brown, 1993; Cameron- iour. He found no evidencefor any effect of kinship Beaumont, 1997). Direct comparisons between these on the choice of partners for allogrooming (related- studies are not straightforward, since different ness coefficients within the colony varied between ethograms have been used, and different social com- 0 and >0.6), which tends to argue against a role in positions observed (Kerby: free-ranging breeding maintaining bonds between kin. However,the possi- farm cats; van den Bosand de Vries: indoor colonies bility remains that allogrooming has other roles in of breeding females; Brown, Cameron-Beaumont: free-ranging breeding colonies. neutered, mixed-sex indoor and free-ranging colonies). Cameron-Beaumont, reanalysing data collected by Brown from three neutered colonies, Signalling repertoire 77 two free-ranging and one indoor, detected five main 1996), as well as the morphology ofskulls (Werdelin, groupings: contact including allogroom, rubbing, 1983). Three major lineages are thought to exist aggressive, defensive, and play (Figure 5.7); sexual and (Figure 5.8): the ocelot lineage, which includes the maternal behaviour were inevitably not included in small South American cats; the domestic catlineage, these groups. The vertically-raised tail (TU) was which includes the small Mediterranean cats; and the associated with both the contact and rubbing groups. pantherine lineage, made up of large and small cats In three colonies of entire females, groups of offen- from several continents. sive, defensive and contact (including allogrooming) with patterns were detected; allorubbing was grouped Spatial organisation in undomesticated Felidae sexual behaviour(rolling, lordosis) (van den Bos & de Vries, 1996) (Table 5.2). Both the function of a signal and the modality These groupingsarelikely to be affected by theage, employed are highly dependent on the distance sex and reproductivestatusof the individualcat. They between the emitter and the receiver. Communication mayalso be affected by genetics and early experience; is therefore intimately related to spatial organisation. the signalling patterns used by McCune (1995) in For any predator feeding on sparsely distributed measuring cats’ reactions to familiar and unfamiliar small prey, non-overlapping hunting areas are pre- people (see Chapter 4) show somedifferential effects dicted (Ewer, 1973; Kleiman & Eisenberg, 1973; of paternity (genetics) and early socialisation. Of the Milinski & Parker, 1991). Field studies have shown defensive vocalisations (directed towards a person), this to be the case for most wild undomesticated cats, growl wasinhibited by socialisation but unaffected by including Felis silvestris (F. s. silvestris: Corbett, 1979, paternity, whereas hiss showed stronger paternal Stahl, Artois & Aubert, 1988; F. s. libyca: Fuller, effects. The frequency of TU was highest in both Biknevicius & Kat, 1988). There are three notable friendly-fathered and socialised cats, but purring was exceptions: the lion Panthera leo (Schaller, 1972), the not affected by paternity, and only enhanced by cheetah Acinonyx jubatus (Eaton, 1970; Caro & socialisation in the presenceof a familiar person. Collins, 1987; Caro, 1989), and the domestic cat (see Chapter 7), all of which have been foundliving greg- ariously. The domestic cat is, however, by no means Communication in the undomesticated an obligate group-living species, and has been fre- felids: the effect of domestication on quently documented to be solitary when foodis at signalling behaviour low density and sparsely distributed (Chapter 7). Given the small numberof generations since domesti- Group-living is most often triggered by anartificial cation, it is reasonable to assume that the domestic clumping of food associated with humansettlements. cat’s repertoire of signals is largely unchanged from The change in niche caused by domestication that of its direct ancestor, the African wildcat F. s. maytherefore cause a decrease in the adaptive value of libyca. However, domestication has substantially solitary life, and a corresponding change in intra- increased the requirementfor social communication, specific communication. both intra- and interspecific. It should therefore be possible to investigate the effect of domestication on Communication in the undomesticated through a comparison of communication behaviour Felidae; differences between lineages signalling in the domestic cat with that of undomesticated felids. Even in solitary species or individuals, signalling 1s necessary for mating, parent—younginteractions, and maintenanceofterritorial boundaries. The wide range Phylogeny of the Felidae of signals exhibited by these largely solitary animalsis Current ideas on the phylogeny of the Felidae are demonstrated by the ethogramsin Tables 5.3 and 5.4; largely based upon molecular techniques, including mostspecies have been found to exhibit a rich reper- albumin immunological distance (Collier & O’Brien, toire of signals despite being predominantly solitary. 1985) and isozyme genetic distance (O’Brien et al, However, the frequently nocturnal and solitary 1987) (for review see Wayneet al., 1989) and mito- behaviour of these species hinders the study of com- chondrial gene sequence analysis (Masuda et al, munication, and as a result much of the published

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irefemales (n = 10, 10, 9) Table 5.2. Groupings ofbehaviourpatterns performed in three confined colonies ofent ee C Colony A B 3 F2 Fl F3 F5 F4 Factor Fi F2 ¥F4 F3 F5 F2 Fil F5 F4 F Ten XK XX XX XxX Rolling 91 XX XX XX XX XX XX 90 Lordosis 84 87 Rubbing 83 62 83 Biting — oe eee 95 Grooming 74 83 46 69 Sniffing 86 62 42 83 Nosing 93 82 78 Sniff rear 66 46 90 64 = Treading - = - - = 78 53 56 97 88 Defensive 91 ——eeeSSo_Ser— Staring 76 48 44 76 XX XX XX XX XX Offensive 92 92 75 eee Figures are percentage factor loadings (values <40 omitted) from separate varimax-rotated factor analyses performed on the patterns exchanged within each pairwise combination of cats in each colony. XX,insufficient data for analysis; —, pattern not includedin the ethogram for this group. From van den Bos & de Vries (1996).

data, particularly on small cats, has been collected on ever, the communciative function of this behaviouris captiveindividuals. not known.

Scraping Olfactory communication Undomesticated cats have additionally been docu- Olfactory signals are long-lasting and would there- mented to scrape their hind feet without urination fore be expected to play an importantpart in commu- or defaecation (Hornocker, 1969; Schaller, 1972; nication between both social and solitary membersof Seidensticker et al., 1973; Wemmer & Scow, 1977; the Felidae. Smith, McDougal & Miquelle, 1989). The absence of urine, faeces or anal gland secretions implies that Urine scrapes are acting asvisual signals as well as olfactory Urine is emitted in the two ways described for the ones (Smith et al., 1989), although scraping may help domestic cat, spraying or squat urination. Spraying to pass secretions from the glandsin thefeet on to the occurs more frequently in males than in females substrate (Wemmer & Scow, 1977). Seidensticker et (Wemmer & Scow, 1977; Mellen, 1993). Sprayed al. (1973) found that scrapes by mountain lions urine has been suggested to contain anal gland secre- Puma concolor demark homeranges,visually and/or tions, whereas squat urinations appear unlikely to chemically. contain any extra components(Schaller, 1972). Squat Mellen (1993) compared the presence and absence urinationsdiffer also in that the urine is usually raked of scraping in 20 species of small cats. Scraping into the soil with the hind feet (known as scuffing/ occurred in most of the species that she observed scraping or raking). It has been suggested that this within the ocelot and Panthera lineages, but in only action may mix urineinto the soil and aid the transfer one species within the domestic cat lineage, Pallas’s of urine scent (Verberne & Leyhausen, 1976), and cat (Otocolobus manul). This species probably possibly also the scent from the glands on the feet diverged from the remainderof the domesticcatline- (Wemmer & Scow, 1977) to the environment; how- age at an early stage (see Figure 5.8), in which. case this

80 IIT; Social life

. Pleistocene Pliocene Miocene

30 25 20 15 10 5 M.Y. Lott lion (PLE) GROUP

Cneereccvenncscsccecs PPA, PON, UUN Panthera

tiger (PTI)

Ovecccneereccceseoces PMA Canadian lynx (LCA) -- LLY, LRU Sevecucvevscnscoccces HYA PANTHERINE cheetah (AJU) LINEAGE setonaeennsesrestenueecust CCA — serval(LSE)

? Oe acceeeccvacenccescs IPL, PVI, PBE clouded leopard (NNE)

PAU Asian golden cat (CTE) puma (PCO)

geveeereansanereceseeceeneussssssansanin FMA,FNI

et Cee ccesecceceneseccanenearseee . FLI domestic cat (FCA) DOMESTIC CAT LINEAGE

ocelot (LPA)

; oeeceneeencccercccnensccccoeee Oco, LWI, LTI OCELOT LINEAGE

Lescececcsscassosscosececcocse sccscocese OGU, OGE

hyena (CCR)

AJU Acinonyx jubatus LTI Leopardustigrina CCA Caracal caracal LWI Leoparduswiedii CCR Crocuta crocuta NNE Neofelis nebulosa CTE Catopuma temmincki OCO Oncifelis colocolo FCAFelis silvestris catus OGEOncifelis geoffroyi FCH Felis chaus OGU Oncifelis guigna FLI Felis silvestris lybica OMA Otocolobus manul FMA Felis margarita PAU Profelis aurata FNI Felis nigripes PBE Prionailurus bengalensis FSI Felis silvestris silvestris PCO Puma concolor HYA Herpailurus PLE Panthera leo yagouaroundi PMA Pardofelis marmorata Figure 5.8. Evolutionary tree of the IPL ictailurus planiceps PON Panthera onca LCA Lynx canadensis PPA Panthera pardus Felidae, from Wayneetal. (1989). The LLY Lynx lynx PTI Pantheratigris positionsof species in bold are based LPA Leoparduspardalis PVI Prionailurus viverrinus on average reciprocal microcomple- LRU Lynx rufus UUN Uncia uncia LSE Leptailurus serval mentfixation measurements (Sneath & Sokal, 1973, Collier & O’Brien, 1985). The positions ofspecies attached by dotted line are based on albumin immunological distance (Collier & O’Brien, 1985). Systematic names have been altered to match those usedin this chapter.

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S a l 7 P o u n c e Cat leaps atoron toanothercat. Cameron-Beaumont 1 99 eaumont199 7, Social Play Socialplay has been divided into contactsocial play, and non-contactsocial Petersen 1979, Cameron-B play by Caro (1995).The various behavioural elements involvedi n thisare Bennett & Mellen 1983, Caro 1995

described inthispaper. Contactsocialplay has alsobeen described asjostle play (described as:one catstruggleswith anothercat, rakingwithits hindlegs and pullingthe opponent towardsits body with itsforepaws). ameron-Beaumont SniffCat Onecat smellsthebody ofanothercat. Itmay be subdividedinto: Mellen 1988, 1993, C Eaton & SniffNose (two cats sniffeach other’s noses), 1997, Free man 1983, Petersen 1977, & Scow,1977 SniffRear (onecat smells theperi-analareaofanothercat), and Velander 1977, W e mm e r SniffBody (one catsmells the head,flank or tailofanothercat). t 199 7, Petersen 1979 Touch Nose Twocats touch each other’s noses;thisisprobably accompanied by SniffNose Cameron-Beaumon 1988, 1993, M o u n t Onecat attempts,butfails to achieve,intromission. The mountingcat Bennett & Mellen 1983, Mellen normally holds therecipientfirmly atthenapeo f theneckwhilstmounting Petersen 1977, 1979

(seeNapeBite).It isalso sometimes accompanied by treading movementso f the hindlegs. Nuzz le Onecatpushesits head againstthehead or,more uncommonly,the bodyo f Cameron-Beaumont 1 99 7 anothercat. This resembles the form ofabriefRub butthere isno rubbing action, only agentlepush. 993, W e mm e r & Scow Paw/pat Onecat pats another individualwith itsforepaw, keeping clawsretracted. Foster 1977, Mellen 1 1977, Cameron-Beaumont 1997 ow 1977, Bite Onecat snapsits teethator succeedsi n nipping another animal. Mellen 1993, W e mm e r & Sc Cameron-Beaumont 1 99 7

Napebite The hold used by the male catwhilstmounting thefemale.The female’s necki s Mellen 1993 held in a firm bite-hold. tt & Mellen 1983, Cuff Onecatstrikes another catwithits forepaw, usuallywith claws extended. Mellen 1988, 1993, Benne Cameron-Beaumont 1997 ett & Mellen 1983, Cameron-Beaumont SitWith A catsits next to, or very near to, anothercat. Benn eee 1997 one has beencollated forthe Signalling behaviour inundomesticatedfelids has been discussed by various authors, but withouta definitive ethogram, although stic cat.Descriptions arebased on domestic cat(UK Cat Behaviour Working Group, 1995). Manyo f thebehaviours described aresimilartothose ofthedome thereferences giveninthe thirdcolumn. © | $$

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“ F Signalling repertoire 85 on objects previously behaviour may imply an evolutionary loss/change many species of small felids rub & Scow,1977). Both of amongst an ancestral member of the domestic cat sprayed with urine (Wemmer Mellen’s (1993) data lineage. Wemmer & Scow (1977) similarly found that these theories were supported by picked up (e.g. this behaviour was absent in the genus Felis (used in in which scents were seen being both by a variety of the strict sense to meanonly cats in the samelineage as urine) and deposited (e.g, saliva) from many species the domesticcat). species. Thirdly, observations suggest that object-rubbingacts as a visual signaldur- Faeces deposition ing reproductive and oestrous behaviour (Acinonyx The method of faeces deposition varies according to jubatus: Foster, 1977; Leopardus wiedu: Petersen, F. s. species (reviewed in Wemmer & Scow, 1977). 1977; Puma concolor: Eaton & Velander, 1977; However,it is difficult to see if there is an evolution- silvestris: Ragni & Possenti, 1990; Oncifelis geoffroyt: ary pattern to these differences or whether they are Cameron-Beaumont, 1997) and in many species of dependent on local conditions. Lindemann (1955, in small cats (Mellen, 1993), as it does in the domestic cat Wemmer & Scow, 1977), found that the Canadian (Rosenblatt & Aronson, 1958; Michael, 1961). Taken lynx (Lynx canadensis) and the European wildcat (F- s. together, these observations suggestthatall three line- silvestris) used two methods, dependent on where the ages of undomesticated cats use object-rubbing simi- defaecation took place; faeces were localised and cov- larly, as a signal of both visual and olfactory nature. ered within territories, but left uncovered in promi- The function of scent-marking was investigated in nent positions at points between territories (which tigers (Panthera tigris) by Smith et al. (1989), who were used as mating rendezvoussites in the lynx). proposedthatit playsa role in establishing and main- This finding suggests that the method of defaecation taining territories. They found that scent-marking may depend on local conditions rather than on was concentrated at potential contact zones where phylogeny. major routes of travel approachedterritorial borders, which supported the hypothesis that the density and Skin glands age of scent-marks give invaders some information As for the domestic cat, tree-scratching functions to aboutthe probability of encountering another animal, removeloose claw sheaths (Wemmer & Scow,1977), and therefore also aboutits risk of injury by being in butit is also used as part of the scent-marking routine that area. This fits with the oft-cited observation that in mostcats, often occurring in the sameareas as other scent-marks rarely act as an immediate deterrent to methods of scent-marking (Mellen, 1993). It may also invaders (Leyhausen, 1965; Schaller 1972; Mellen, leave a visual signal (Wemmer & Scow, 1977). This 1993). Previous hypotheses on the function of scent- behaviour occurs in a diverse range of felids (Pallas’s marking have tended to involve the idea that this cat, sand cat, fishing cat, Temminck’s goldencat, behaviour provides temporal information about the jungle cat, rusty spotted cat, Indian desertcat, serval, whereaboutsof each individual cat (Leyhausen, 1965; caracal, African golden cat, Geoffroy’s cat, jaguarun- Schaller, 1967, 1972; Hornocker, 1969; de Boer, 1977), di, ocelot, Scottish wildcat, Siberian lynx, Canadian which mayalso be the case, although as noted above, lynx: Mellen, 1993; margay: Petersen, 1979; tiger: the benefit to the producerof the signal is unclear. The Schaller, 1967, Smith et al., 1989; lion: Schaller, second function of scent-marking found by Smith et 1972; Canadian lynx, Pallas’s cat, jaguar, fishing cat, al. was that it serves to signal the onset of oestrus in leopard cat: Wemmer & Scow, 1977; snow leopard: the female. This was supported by Mellen (1993), who Hornocker, 1969; Seidensticker et al., 1973; cheetah: found that a change in the markingrate of the female Eaton, 1970; leopard: Bothma & Leriche, 1995), and was a goodindicator of reproduction in a variety of appears to have changedlittle in character or function small cat species. during the courseoffelid evolution. Object-rubbing has been suggested to have three Acoustic communication waysofacting:first, it acts as a methodof scent-marking by depositing gland secretions such as saliva on Acoustic signals in felids carry a wide variety of mes- objects (Ewer, 1973; Wemmer & Scow,1977). Rieger sages (Peters & Wozencraft, 1989), and are used & Walzthony (1979; see also Rieger, 1979) addi- across long distances as well as during close contact tionally suggest that object-rubbing picks up scent, as and in group-livingfelids as well as solitary ones. For 56 IIT: Social life

example, calls can display territorial advertisement ever, no evidence of social rolling in undomesticated (Eisenberg & Lockhart, 1972), defensive and offen- felids being used in the submissive manner sive described threat(spit, hiss, growl, snarl: Wemmer & Scow, for the domestic cat. 1977; Peters, 1983; Cameron-Beaumont, 1997), close With the exception of Tail Under, which occurs in range affiliation (prusten, gurgle, puffing: Peters, conjunction with Crouchas a defensive posture, no 1984a, b), mating signals, both for sexual advertise- othertail position appears to act asa signal in undom- ment (male and female sexual calls: Kleiman, 1974; esticated cats (Cameron-Beaumont, 1997). The one Foster, 1977; Petersen, 1977; Seager & Demorest, exceptionis the lion, which has been reported to show 1978; Peters, 1980; Rieger & Peters, 1981; Ragni & a Tail Up position in conjunction with rubbing, Possenti, 1990) and during copulation (Peters, 1978; although it was not described as a signal (Schaller, Rieger & Peters, 1981), infant signals of contact (purr, 1972); this is discussed furtherin the section on effects miaow:Schaller, 1972) and distress (miaow); identifi- of domestication(p.87). cation messages (call sequence duration in lions: There has been no investigation into the use of Peters, 1978); and to encourage assembly of a group body andface signals in undomesticated felids, with (roaring of lions: Schaller, 1972). Table 5.4 lists the the exception of Schaller (1972), who describes the use most commonly cited calls, and those which have of these visualsignals in the lion. been described in somedetail. Unfortunately, however,it is impossible to create an exhaustive ethogram of felid calls because detailed Tactile communication information on many species is sparse. There are Tactile communication in free-rangingsolitary felids anecdotal mentions of other sounds (e.g. Schaller, generally occurs as a componentofeither mating or 1972; Foster, 1977; Petersen, 1979; Cameron- mother—young behaviour. However, in zoos (where Beaumont, 1997) butit is not usually possible to tell unrelated adult cats are often kept together), tactile whethertheseare distinct soundsorjust a grading of a communication between adult cats is regularly previously recorded call, or a slight call variation observed in a more general social context, although between species. For the well-detailed or well-known the rates of tactile contact vary. Many naturally calls listed in Table 5.4, however, most appearto be solitary cats have been observedto be sociablytactile relatively uniform acrossthe three lineages, although in captivity (Mellen, 1993; Cameron-Beaumont, the roar is found only in the Pantheralineage. Other 1997), which demonstratesthefelids’ ability to adapt ditferences include the close-rangefriendly affiliation their behaviour according to the prevalent conditions, call described by Peters (1984a, b), which differs in although othercaptive studies have foundlowratesof structure across the three lineages, there being three tactile contact (Tonkin & Kohler, 1981; Bennett & types (gurgle, prusten and puffing), all of which are Mellen, 1983). The rate appears to depend onindivid- thoughtto have the same functionin differentspecies. uals rather than on thespecies involved. Threat and infant sounds appear to be relatively In social cats, particularly the lion and the domestic uniform. The less commonlycited calls include the cat, tactile signals are frequently used as general social wah-wah andthe chatter, both described by Peters signals as well as morespecifically in a reproductive or (1983, 1987). It is not known how widespread these parental context. Interestingly, tactile signals appear two soundsare acrossthe lineages. to be used in a similar manner in these two social species, despite their different evolutionary lineages. Social rubbing amongst small felids Visual communication has not been documentedin the wild; in captivity, it may be derived As in domestic cats, social rolling in undomesticated from the mating ritual, due to its occurrence during felids is a component of sexual behaviour (Uncia reproductive behaviour (F. s. silvestris: Ragni & uncia: Freeman, 1983; A. jubatus: Foster, 1977; L. Possenti, 1990; P. leo: Schaller, 1972; O. geoffroyi: wiedit: Petersen, 1977; F. s. silvestris: Ragni & Cameron-Beaumont, 1997; several species of small cats: Possenti, 1990; several species, Mellen, 1993), Mellen, 1993). However, some publications on felid although in captive cats it does also occur in general reproductive behaviour do not mentionsocial rubbing, social situations (O. geoffroyi, F. chaus, Caracal despite mentioning object-rubbing(A. jubatus: Foster, caracal: Cameron-Beaumont, 1997). There is, how- 1977; P. concolor: Eaton & Velander, 1977). 87 Signalling repertoire wherebya signaldiversi- Social rubbing amongstlions has been reported in ritualisation can then occur, distinct signals, more detail, occurring as an affiliative behaviour fies, givingrise to several functionally between adults. Schaller (1972) found that rubbing via the followingstages: had occurredparticularly after membersof the group 1 Signal occurs in one contextonly. been separated, and also after agonistic interactions. 2 Signal appears in two contexts, assuming a He suggested that this behaviour indicates that the secondfunction, but remains structurally that intentions of the animal are peaceful. He found unchanged. males rarely rubbed on females or cubs, while females 3. The two signals becomestructurally distinct in rubbed on both males and females, and cubs rubbed the two contexts. mostly on females; this is compatible with the explanation that rubbingacts as a placatory gesture, Stage 2 is therefore an essentially transient phase producing morebenefit for a subordinate animalthan between Stages 1 and 3 (Otte, 1974). for a dominant. Interestingly, this system has also Domestication can provide an insight into the been proposed for the other group-living cat, the processof ritualisation of signals, becauseit is possi- domestic cat (Macdonald et al., 1987). The fact that ble to compare the domestic cat with relatives that both lions (Panthera lineage) and domestic cats behave verysimilarly to its ancestor; thusit is possible (domestic cat lineage) appear to use social rubbing to determine whether any diversification or ritualisa- as a placatory signal implies that rubbing may have a tion of signals has occurred during domestication. similar function in mating behaviour amongstsolitary Differences between signals used by the domesticcat cats, i.e. indicating that the intentionsof the animalare and undomesticated felids are therefore discussed peaceful, both before and after copulation. If this is below, these being differences which may have been the case, then it is understandable that this signal has caused by domestication, both by altering the circum- diversified to be used in othersocial contexts amongst stances in which intraspecific behaviour is expressed the two gregarious species, F. s. catus and P. leo, (e.g. high local population densities), and byintro- despite their different lineages. ducing a need for interspecific (i.e. cat—human) Social groomingin solitary cats occurs both as part communication. of mating behaviour (Schaller, 1972; Foster, 1977; Freeman, 1977; Petersen, 1977; Cameron-Beaumont, (1) The evolution of a new signal from a 1997) and in mother—younginteractions, in which it non-signal behaviour: Tail Up has a utilitarian function of maintaining the cubs’ cleanliness. In the gregarious lion it occurs in these The action of Tail Up,as an integrative part of urine- two situations, and additionally in a non-specific spraying, is thought to occurin all species of felids, social situation, frequently when two are resting domestic and undomesticated. Thetail is raised verti- together (Schaller, 1972). The function of this has not cally during spraying and then immediately lowered been elucidated. Normally solitary cats kept in cap- (Hornocker, 1969; Schaller, 1972; Wemmer & Scow, tivity also use social grooming in this non-specific 1977; Smith et al., 1989; Mellen, 1993; Bothma & manner (Mellen, 1993; Cameron-Beaumont, 1997). Leriche, 1995; Solokov, Naidenko & Serbenyuk 1995). However, in domestic cats, Tail Up has additionally been shown to act as an affiliative signal The effect of domestication on cat—cat (Cameron-Beaumont, 1997). The Tail Up affiliative behaviour signalling signal differs from the raised tail that occurs During domestication F. silvestris must have adapted during urine-spraying in both context (being linked to living at higher densities than previously, and then to affiliative behaviours, in particular social rubbing) subsequently adopted group-living. Sincethesignals and structure (occurring for prolonged periods of needed by solitary animals have different properties time, often remaining upright during locomotion) from those needed by group-living individuals, this (Cameron-Beaumont, 1997). move may haveled to an evolutionary change in the Cameron-Beaumont(1997) investigated the point signalling patterns used by this species. at which the Tail Up affiliative signal might have Signals must be derived originally from non-signal evolved by lookingforits presence in undomesticated movements, by ritualisation (Harper, 1991). Further felids, using representatives from all three evolution- 58 ITT: Social life

ary lineages. There was no evidenceofits presencein quantitative differences in the character of signals. any of the three species studied (O, geoffroyi (ocelot Thusit would prove particularly interesting if the Tail lineage), 2 = 14: C. caracal (Panthera lineage), n = 13; Upaffiliative signalis found to have evolved asa result F. chaus (domestic cat lineage), n = 12) during total of domestication. of 539 hours of observation. All three species carried out social and object rubbing without raising their (2) An established tail; this is in contrast to domestic cats, where rubbing signal diversifies to develop a secondary is almost exclusively carried out with thetail held ver- function (i.e. occurs in a new tical. Theraisedtail during spraying was, in contrast, context), but does not change in structure observed in all species. None of the publications Social rolling in undomesticatedfelids is a sexual sig- which discuss felid communication and behaviour nal, occurring as part of the reproductive repertoire. (with the exception of the lion: see below) mention In domestic cats it is still used in this reproductive Tail Up occurring in any context other than urine- manner (Rosenblatt & Aronson, 1958; Michael, spraying (Wemmer & Scow, 1977; Mellen, 1993; 1961), but is additionally used as a submissive gesture Table 5.3). in groups of domestic cats (Feldman, 1994b). Thereis This study appears to suggest that Tail Up may no evidence that undomesticated felids use social have evolved as an affiliative signal during domestica- rolling for this function, althoughit is possible thatits tion, perhaps consecutively with increased sociality, role in sexual behaviouris a submissive one, in which which may havecaused the necessity of an additional case it is only a small step to its general (non-sexual) visual signal. However,it cannot be ruled outthat the use as a submissive behaviour in groups of domestic Tail Up signal may have evolved at an earlier stage, cats. possibly amongst one of the undomesticated formsof Social rubbing and social grooming are also both F, silvestris. There are few behavioural studies on the sexualsignals in undomesticatedfelids. In the domes- undomesticated subspeciesof F.silvestris, particularly tic cat, however, they are additionally used in a the African subspecies, which may account for the general social greeting situation. However, this absence of any mention of Tail Up. change in context and thus in function cannot be The one exceptionto this is Schaller’s description of attributed to domestication, because adult undomes- social behaviourin lions (Panthera leo), in which he ticated cats in zoos exhibit the same changes,i.e. an states that social rubbing (in both mating and general increased use of head-rubbing and grooming in non- social situations) frequently occurs with the tail sexual situations (Cameron-Beaumont, 1997). Thus raised. He writes: ‘During head-rubbing and anal- the use of these behaviours in a wide variety of sniffing contacts the animals raise their tail so thatit affiliative contexts is probably a natural ability of all either arches overtheir back or tips towards the other felids rather than a product of domestication. animal.’ He gives no moredetail about the contextual nature of this behaviour, but the fact that it occurs Neotenised signals with the affiliative behaviours of rubbing and anal Miaow,knead and purrare all generally considered to sniffing implies thatit is being used in a different way be juvenile behaviours, with the possible exception of from the raised tail during spraying. Its function in purr, which also occurs in adult cats (Peters, 1981). lions may even be similarto that in domestic cats(i.e. However, in the cat—humanrelationship, adult cats as an affiliative signal). The occurrence of a Tail Up use all three of these signals habitually (e.g. Turner, affiliative signal only in F. s. catus and P. leo, from 1991; Bradshaw & Cook, 1996). Cameron-Beaumont different evolutionary lineages, but not in any other (1997), in a survey of zoo keepers, found that adult undomesticated species of felid, implies that this undomesticatedcats in captivity were very unlikely to signal may have evolved separately in the two species, perform any of these three vocalisations towards possibly as a result of similar selective pressures acting humans, suggesting that undomesticated cats cannot only on the two mostsocialspeciesofcats. naturally revert to performing kitten behaviours Various previousinvestigations have looked for the when adult. This discrepancy suggests that the emergence of a new behavioural pattern as a result of domestic cat has evolved (either culturally or geneti- domestication (reviewed in Kruska, 1988), but to date cally) the ability to use kitten behaviours towards no new behaviours have been found, despite many humanswhenadult (neoteny). Signalling repertoire 89

animal as a whole, although Cameron-Beaumont signal diversifies in both (3) An established (1997) has achieved it for the Tail Upposture. a different structure and function to become Thereis also a possibility that the domestic cat pro- signal duces subtle signals which have yet to be identified as difficult to There are no definite examples of this in the cat—cat such. Cooperative signals may be very relationship, but a change in signal structure does detect experimentally, since they should be produced and appear to have occurred in cat—humansignals. with the minimum amountof energy required, possible Cameron-Beaumont(1997) investigated the use of should keep the signaller as inconspicuousas & rubbing in the domestic cat in both cat-cat inter- to minimise detection by predators (Krebs by actions and in cat—-humaninteractions. In the human- Dawkins, 1984). For example, the grunts emitted the directed situation, rubbing occurs at a higher vervet monkeys, although indistinguishable to frequency and at a higher intensity than it does in humanear, are producedin atleast two distinct forms the cat—cat situation. This difference is likely to have with different meanings (Cheney & Seyfarth 1982). occurred partly becauseof the change in recetver psy- Since sociality in the domestic cat may be somewhat chology (Guilford & Dawkins, 1991), but also primitive, and may even have evolved as a conse- because of the change in the meaningofthe signal;it is quence of cats’ association with humans, we might likely that much of human-directed cat behaviour1s not expect such signals to have emerged as yet. exhibited as either a food- or attention-getting signal However, this may be something of a circular argu- (see also Mertens & Turner, 1988). This is in contrast ment, i.e. we may regard the cat’s social system as to the message given in the cat—cat situation (whereit primitive because we have notyetidentified all the acts as a subtle affiliative signal). A food-eliciting sig- signals by which relationships are established and nal would favour a ‘loud’ prominent signal, whereas maintained, and also do not yet fully understand an affiliative cooperative signal between membersof a those we haveidentified. colony would favour a subtle cue (Krebs & Dawkins, It is still unclear, for example, whether convention- 1984). Thus the difference in the type of message that al concepts of ‘dominance’, which are so useful in is being given by rubbing maycausea difference in the interpretingthe social behaviourof otherspecies, can frequency andintensity with whichthe signalis given. be usefully applied to the cat. In termsofsignals, the This ritualisation of an established cat—cat signal in the roles of allogrooming and rubbingin redirecting and cat-humansituation may have also occurred in other averting aggression warrant further investigation. commoncat—humansignals such as the miaow. Our understanding of the role of scent-signals in social behaviour has also lagged behind that of some other mammals, particularly since synthetic Concluding remarks analogues of the so-called ‘facial pheromones’ of the Despite a substantial literature on communication in cat are now becoming commercially available for the cat family, several important issues remain to be the control of indoor urination (White & Mills, 1997) resolved. Thefirst is whether everything that has been and aggressive behaviour (Pageat & Tessier, 1998). described as communicationreally involves transmis- All of the scent-marking performed by cats is in sion of information from onecat to another, and con- need of reappraisal in termsof the benefits accrued by versely, whether all the signals produced by the the depositor, as well as the recipient, as we have domestic cat have been identified. Most signals have attemptedto do for spray-urination by males. been defined on the basis that they are behaviour Finally, the cat offers considerable opportunities to patterns that are obvious (to humans) and which examine the effects of domestication on signalling. appearto elicit responses from other cats. However, This may have occurred in two non-exclusive ways: rigorousinterpretation of a behaviourpatternasa sig- either ‘hard-wired’ changes in the structure and/or nal requiresthat it should be tested independently of meaning of signals which are inherited genetically, the context in which it normally occurs. This is more or an enhanced ability to learn to communicate in easily achieved for vocal signals (playback exper- new ways, particularly whensignalling to humans. ments) and chemical signals (presentation of isolated We suggest that the appearance of the Tail Upsignal or synthetic odours) than for visual or tactile cues, is an exampleof the former, appearing as a methodfor where the signal is difficult to separate from the avoiding unnecessary conflict as cats adapted to 90 LIT: Social life

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