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Chapter 29

COMMUNICATIONIN THE WITH EMPHASIS ON SCENT MARKING AND CONTACT PATTERNS

Christen Wemmer and Kate Scow

Introduction Methods

It is ironic that students of social behavior Ourstudies have taken place at the Chicago and communication have not shown as muchin- Zoological Park (Brookfield Zoo). Unsystematic terest in as the general public has. Fewer observations have been made during the past than a third of the thirty-six living species of cats two years on the following species: African have been studied from an ethological stand- ( leo), Indian (P. tigris), (P. point. Because of the scarcity of information, pardus), (P. onca), (P. uncia), moststudies have been of a general nature, and ( nebulosa), few have been addressedto particular aspects of ( jubatus), (Puma concolor), communicative behavior. Critical, detailed inves- (Lynx lynx canadensis), sand (Felix margarita), tigation of auditory, visual, and tactile modes of (F. silvestris), Pallas’ cat (F. manul), interaction is to our knowledge greatly under- golden cat (Profelis temmincki), represented. At the time of writing there are only ( bengalensis), (P. viverinus), a handful of studies published on olfactory as- ( wiedu), (Her- pects of communication (Fiedler, 1957; Palen petailurus yagouaroundi). A focal method of and Goddard, 1966; Verberne, 1970). For re- observation (Altmann, 1974) has been employed views of cat behavior the reader is referred to to monitorsocial interaction in twolitters of Pal- Leyhausen’s pioneering studies (1956, 1960) las cat containing four (one male, three female) and the more recent works of Schaller (1967, and five (one male, four female) young, respec- 1972). This paper is not intended as such a re- tively. Daily observations of varying length (ten view, but reports on certain aspects of communi- to eighty minutes) were made on twolitters of cation involving contact directed to objects or leopard cat (two male, two female, respectively) companions. It is hoped that our statements and onelitter of sandcats (four male). In these concerning the ecological determinants and de- cases we continuously sampled patterns of con- sign features of felid communication will stim- tact in all . ulate more complete analysis than we can now A brief review of felid natural history is neces- offer. sary to understand the ecological setting to 749 750 Communication in Selected Groups which the felid mode of communication 1s Secondly, the feline estrous cycle and in- adapted. duced ovulation place several constraints on courtship. During proestrus the female becomes hyperactive and announces her emerging recep- The Felid Habitus tivity by rubbing and calling (Rabb, 1959; Mi- The felid body plan is progressive, but its chael, 1961). This attracts a number of males, uniformity between speciesis striking when com- and severe rivalry, which is manifested by a pared with most other families of Carnivores. seemingly disproportionate amount of display The small size and forest habitats of mostliving relative to combat, gives rise to a dominantani- cats are probably primitive adaptationsfor utiliz- mal (Ewer, 1973). The female’s repulsion of the ing the relatively diverse small vertebrate fauna male diminishes after further repeated intense inhabiting such regions (Kleiman and Eisenberg, and often strikingly dramatic transactions. In- 1973). The most economical schemefor exploit- duced ovulation requires that the female copu- ing such preyis a system of solitary land tenure. late repeatedly, often over a period of days The cat occupies a more or less exclusive hunt- before ova are produced and conception can oc- ing ground and probably encounters other com- cur (Ewer, 1973; Schaller, 1972). Often as the munity members infrequently. The mother sexual motivation of the female waxes, that of family, the most complex social unit, 1s but a the male wanes (Ewer, 1974). Sexual exhaustion brief, usually seasonal association. High-inten- of the first male mayresult in his retirement from sity vocalization and locus-specific marking with further involvement, and another male maythen scent are the two predominant methodsby which step in. The observation that males are more various species space themselves and avoid con- tolerant of one another than are females may in frontations (Muckenhirn and Eisenberg, 1973; part be explained by the adaptivenessof nonfatal Schaller, 1967, 1972). In more open habitats inter-male competition during these circum- movement may be_ regulated by vision stances (Leyhausen, 1965a; Berrie, 1973; Pro- (Leyhausen, 1965a). It is also likely that neigh- vost et al., 1973). In summary, the usual solitary bors recognize one another,andthatthe ‘“‘broth- existence of the mature but sexually inactive cat erhood”’ is characterized by a loose social breaks down when females comeinto estrus and hierarchy in whichtheterritory insures even the becomehighly attractive to males. The proximity lowest-ranking cat of priorities in resources and of rival males provokes intensive display and space (Leyhausen 1965a). fighting. Further agonistic behavior developsbe- Since nearly all cats can kill prey as large as tween the dominant male and female. In these themselves, hostilities between conspecifics are contexts and in occasionalterritorial disputes potentially lethal (Leyhausen, 1960, 1965b; motivational differences between animals are re- Schenkel, 1968; Schaller, 1972). This fact has solved throughintenseand highly modulated in- been instrumental in evolving a more stereo- teractions. typed repertory of distance signals on the one hand, and a highly graded repertory of proximal Some General Features of Felid signals on the other. Furthermore,thereis a par- Communication allel between the dependence on vision and au- dition in the localization and captureof prey, and In view of the paucity of information for a the linkage of visual (facial expression) and audi- variety of species, it is difficult to make general- tory signals in proximal agonistic interactions. izations. However, the available information Communication in the Felidae 751

suggests several parallels and contrasts with excitation and sound intensity seem to be com- other carnivore groups. mon concurrent features of such sequences. There are six calls that are commonto the Whengrowling is resumedit is usually after an repertories of most small cats that have been interruption in sound production. Extensive studied andthese can be groupedinto categories sonographic analysis is needed, however, to con- of discrete and gradedcall types. Some physical firm this observation, for each call type may be properties of these calls are listed in Table 1. All considerably modulated in frequency, intensity, of these vocalizations, with the exception of spit- and noise level. As Schaller (1972) noted, the ting, vary in intensity, duration, and emission repertory of cats is smaller than it seems because rate. Spitting and hissing share features of broad discrete signals are uncommon. nontonal energy distribution. Sonographed ex- These calls share certain features with the amples of the two calls can be arranged on a graded vocalizations of terrestrial primates dis- continuum from long, moderateintensity hisses cussed by Marler (1965, 1967). They are ad- to brief, loud, and explosive sounding spits. dressed to conspecifics at close range and are However, most examples fall at the extremes of accompanied by highly varying facial expres- the continuum,andtherefore the labeling of two sions. The potential of such visual-vocal signal basic calls seemsjustified. Whether the variants systems for communicating fine-grain motiva- are lumpedorsplit, the call(s) cannot be consid- tional changeshas been discussed by Marler. In ered graded because the variation between ex- cats these signals occur mainly during proximal tremes does not involve qualitative differences. agonistic interactions, territorial skirmishes, and Intermediates exhibit quantitative variation preambles to copulation. The sounds may within the same physical parameters. broadcast and attract neighboringcats, but this Graded calls among the small cats display seems to be a secondary or inadvertent side several characteristics. Call transitions often oc- effect. The discrete, high-intensity calls that cur without an interruption in the air column. serve to attract or space neighbors seem to be This producesa usually short intermediate seg- lacking, or are at most only poorly developed, mentof sound that shares certain characteristics among the small cats that have been studied. of the preceding and followingcalls. This kind of Tomsare certainly attracted by the discrete and noninterrupted, inter-call gradation has been repeated miau of the estrous domestic cat, but described for a number of small carnivores this call cannot be comparedonrelative grounds (Wemmer, in press) and primates, particularly with the roaring and sawing calls of the great open-habitat terrestrial forms (Marler, 1965, cats. 1967). The features of this kind of acoustical This leads us to considerthe sensory assort- gradingareto be distinguished from the tempo- ment of the feline signal repertory. Based on rally discrete (interrupted) but graded calls of Leyhausen’s (1960) studies of the domestic cat, other (e.g., red colobus monkey: Eisenberg (1973) tabulated twenty-five visual Marler, 1970). patterns (facial expressions, tail and body pos- A second feature of these graded vocaliza- tures), sixteen of which occur in combination. tions is that gradation often seems to be one- There were eight vocal patterns and three olfac- directional. For example, howling (an harmonic tory patterns, andto this list can be added about call) may arise from growling (a pulsedcall) and seven contact patterns (body rubbing, clasp, terminate with a scream (a noisy, high-frequency mount,bite, lick, pat, hind leg pump). A similar call) followed by a rapid volley of spitting. Rising profile emerges from Schaller’s (1972) lion 752 Communication in Selected Groups study; there are at least seventeen visual pat- ing thatit assists in orienting the direction of the terns, thirteen vocalizations (including graded spray (Fig. la). More often than this, however, series), seven contact patterns, andfive olfactory the terminal part of the tail undergoes a marked patterns. On the basis of this cursory examina- erratic twitching lasting several seconds. The tion, vision and audition seem to be the twotop- pattern occurs in snow leopard, lynx, leopard ranking signal modalities, a pattern that agrees cat, and domestic cat, but is apparently absent in with the general impression that cats are essen- the other Pantherinae. While Schaller states that tially sight- and sound-oriented animals. spraying is purely an olfactory pattern in , the conspicuous character of this movement in Scent Marking the above-mentioned cats implies it may also have a visual signal function. Alternately it may Urine, feces, and glandular exudates of the simply be an autonomic manifestation of urine skin are potential carriers of chemical informa- emission. Schaller (1972) states that in lions one tion in nearlyall terrestrial mammals.In the Fell- to twenty jets of urine are emitted during the dae feces seem to lack the widespread assumption of the characteristic stance andthat communicative significance that they havein the it travels three to four meters. canids, but urine 1s undoubtedly an important Defecation is similar between sexes and spe- information carrier. The five patterns of object- cies, but its association with other patterns of oriented contact that can be distinguished vary behavior differs between species. In many small in expression between species and possibly be- felids (particularly the ) feces are tween sexes; interspecific postural differences deposited in areas where they can be covered by and variation in sequencing also exist, but de- repeated scratching motions of a forefoot. tailed studies on these aspects are lacking. Largercats (Panthera) make no attemptto cover Urination occurs in a squatting or standing feces; defecation does not occur in conspicuous position. The latter is nearly a universal male areas and is not locus-specific, as it is in many felid trait, but it is also seen in the females of other carnivores. Lindemann (1955) reported somespecies (Table 2). It is assumed that retro- that both the European wildcat and the lynx bury mingent urination against upright objects (Harn- their feces at specific localities within their terri- Spritzen of Leyhausen, 1956; spraying or urine tories, but leave them uncovered on stones and spraying of Schaller, 1967, 1972) has evolved tree stumps in the spaces between territories specifically as a scent-markingpattern. The urine (Memandslandstreifen). In the lynx these “‘rendez- is spread overa larger area than if deposited on vous” sites are eagerly sought out during the the ground;it can be sniffed at head level; and mating period by all adult conspecifics (Lin- accordingto the diffusion model of Bossert and demann, 1955). Wilson (1963), a point source of scent above While anal scent glands occur in most small ground can produce an active space as much as cats, the buryingof feces suggests that anal scent twice as large as a scent source at groundlevel. secretion does not take place during defecation, Ewer and Wemmer(1974) have also pointed as it does in certain rodents. Adamson (in out that the height of a scent mark can be used Schaller, 1967) reported that anal scent was to judge the sex of the owner; however,this in- voided during urine spraying in her female lion formation might be redundantto sex identifiers Elsa, and Schaller (1967) observed that tiger in the scent itself. The upheld tail at times urine deposited by spraying has a strong musky touchespartof the object to be sprayed, suggest- odor compared with urine deposited in a squat-

Fig. 1. Scent-marking patterns in a male snow leopard (Panthera uncia): a. Urine spraying (notice po- sition of tail). b. Scuffing with the hindfeet. c. Sniffing of leg prior to head and neck rubbing. d. Head and neck rubbing. The animal mustrise up on its hind legs to touch the elevated branch.

753 Table 1 Some physical properties of vocalizations common to members of the genera Felis, Prionailurus, and Lynx.

Harmonic Relative Frequency Vocalization structure Duration intensity modulation

Hiss _ Variable but usually brief Moderate — Spit — “Fixed” Moderately loud — — Variable, but usually long and repeated Soft — Growl — Variable, but usually long and repeated Moderate + Miau + Variable Moderately loud + Scream + Variable Loud +

Table 2 Distribution of scent-marking patterns in various Felidae.

Urine Scuffing Species spraying Feces (scraping) M F

African lion (Panthera leo) + — Scattered haphazardly, not buried + Tiger (P. tigris) + + Scattered and covered + Leopard (P. pardus) + — Occasionally deposited on a scrape + made by scuffing Jaguar (P. onca) + — Puma (Puma concolor) — — Deposited on scrapes + Snow leopard (Panthera uncia) + — Scattered, not buried + Clouded leopard (Neofelis nebulosa) + — Cheetah (Acinonyx jubatus) + — Occasionally deposited on scrapes + (Lynx lynx) + + Localized and covered and uncovered ? (Lynx rufus) + — Localized deposits ? Fishing cat (Prionailurus viverinna) | + ? In water ? — Leopardcat (P. bengalensis) + — Scattered, buried — Margay (Pardofelis wiedii) + — Not buried + Golden cat (Profelis temmincki) + ? ? ? Jaguarundi (F. yagouaroundi) + — ? + Pallas cat (F. manul) + — Feces covered — (F. margarita) + — Feces covered _ Domestic cat (F. catus) + — Covered Wildcat (F. silvestris) + — Locusspecific —

754

Communication in the Felidae 755 ted posture. He attributed this to a granular the entire length of its hind feet touching the white precipitate, but even in captivity where ground. Then each foot is alternately thrust spraying can be viewedatclose rangeit is not backward while the claws are usually extended. possible to witness the emission of the anal Upon completion of this scuffing motionthe foot scent, andit is possible that the white precipitate is almost always lifted as it is brought forward. seen was actually the glycerides excreted in the Occasionally one foot may repeat the scuffing urine by the cats. motion up to six times before the other foot Scuffing (scraping, treading) with the hind comesinto action (snow leopard). The emphasis feet is commonly associated with urination (Fig. of the contact along the length of the foot proba- Ib). The animal squats on its hindquarters with bly varies between species, as a numberofvaria-

Table 2 (continued) ee Head Recumbent head rubbing Claw raking rubbing Reference

ee, + + + Fiedler (1957), Schaller (1972) + + + Fiedler (1957), Schaller (1967) + + + Eisenberg (1970), Fiedler (1957), Muckenhirn and Eisenberg (1973), Schaller (1972) + + Fiedler (1957), Wemmer and Scow (pers. obs.) + + + Fiedler (1957), Hornocker (1969), Seidenstickeret al. |(1973) + + + Hemmer(1968, 1972) + + ? Hemmer(1968) + + + Eaton (1970) + + + Wemmerand Scow(pers. obs.), Lindemann (1955) — — ? Provostet al. (1973) + + + pers. obs. + + + pers. obs. + + + M.Peterson (pers. comm.) + + pers. obs. + + + Ewer(pers. comm.) + + + pers. obs. + + + Hemmer(1974), Wemmerand Scow(pers. obs.)

+ + + Lindemann (1955) eee 756 Communication in Selected Groups

tions have been described in the occursfor brief periods during or near the termi- (Wemmer, in press). The tempo of the move- nation of the scuffing bout or after the move- ment is moderately fast, but varies somewhat menthas stopped. Urination during scuffing may with the size of the species. be associated with a slight deceleration or even It can be seen in Fig. 2 that the variation a brief pause in leg motion. The churningeffect within an individualis slight and manifestsa typ!- of the feet mixes urine into the soil, and distrib- cal intensity in rate of delivery, but not in bout utes it along the length of the foot. The fur on length. The result of the movementis a charac- the sidesof the feet of captive snow may teristic scrape in which loosenedsoil is heapedat acquire a yellowish tinge from the activity, but the posterior end. The movement may also this may result from both unusually high levels of scrape bark loose or scratch the surface of hori- scuffing and its repeated occurrence at onesite zontal or diagonal logs (jaguarundi). Urination in captivity. In the African lion the urine often wets the hind legs (Schaller, 1972). Thus it may act as a solvent transferring pedal scent to the substrate. Seidensticker et. al. (1973), who studied the mountain lion in the Idaho primitive area, found that nearly all the 86 scrapes examined were madebyresident adult males, and that the fre- quency of scraping was highest in the overlap zone between the home ranges of two radio- one to six scrapes were BOUT tracked males. From

IN found within a small area at each site; they were usually placed near but not on animaltrails, and

SCUFFS only 11 of the 86 sites were revisited. Feces or OF urine was detected at 17 of the sites, usually on the soil and needles heaped up by the scraping NUMBER action. Whetherscrapes are made withthe fore- or hind feet is not known. The authorsin this study concluded that scrapes demark the home ranges of adult males by indicating that the area is occupied. Schaffer (1940) described glands in the feet of the domestic cat, but the extent and type of glandular developmentin the feet of other spe- cies is not known. Scrapes made by are 5 10 15 20 25 30 35 40 45 also defecated upon if only in small amounts; DURATION OF BOUT (SEC) these scrapes are more evident and perhaps more commonly performed during the mon- Fig. 2. The relationship between the numberof soon, while the feces persist longer than the bout and the duration of scuffs composinga scuffing scrape during the dry season (Schaller, 1967). snow leopard (Panthera uncia). the bout in a male pos- Bouts varied from 1 to 47 sec and contained 2 to 60 Head rubbingis associated with several individual leg movements (scuffs). tural variants and is almost always preceded by | Communication in the Felidae 757 snifiing (Fig. Ic, d) or licking and biting of the rion, vomit, the feces of strange animals, and focal object, and by flehmen. Most commonly catmint. The latter pattern, which is part of the the face is rubbed as the animal stands beside a daily routine, characteristically occurs against branch or rockat head level. The contact phase objects that are also sprayed with urine. is often one-directional; it occurs as the body Depending on the position of the tree trunk, leans toward the object with the cheek or neck claw raking 1s performed in an upright or a hori- serving as the point of contact. Forward move- zontal position. Thecat generally grips the trunk mentof the bodyis achieved by a combination of with extended forelegs and depressed body, and neck extension, walking, and leaning forward. the claws are then drawn backward simulta- Upon breaking contact the cat may assume a neously or alternately in strokes of variable different positionin relation to the object or may length and speed. The motion often has a jerky repeat the movement from the samestarting quality that results from the intermittent snag- point. The head, cheeks, and neck may also be ging of the claws and raking. Theaction serves turned and rotated as the animal standsstill, to remove loose claw sheaths but also leaves a sometimessecuringthe branch with the claws of visual and possibly an olfactory trace having so- a forefoot. A similar style is employedby sitting cial significance to othercats. cat. If the branchis elevated the cat will grasp it The occurrence of these patterns in different with one or both forefeet while standing upright felid species is presented in Table 2, but there on the hind legs. Rubbing andsniffing very often have been few studies on the temporal organiza- lead shortly to salivation. A clear watery saliva tion of scent-marking behavior. Urine spraying appears on the closed lips and is wiped onto the occursbyitself and in association with otherpat- object and the cheeks and neck. A vigorous bout terns. Eisenberg (1970) reported that in the Cey- of rubbingvirtually soaks these areas. lon leopard scraping (scuffing) and urine The cheeks and neck are also rubbed against spraying occurred near the sloping trunks of novel or odoriferous substances on the ground. trees in which clawing with fore- and hind feet Herethe cat reclines with the object between the (scuffing?) and cheek rubbing took place. In the forefeet. After the object is sniffed and licked, snow leopard, cheek rubbing, scuffing, and the sides of the head and neck are pressed spraying are often associated acts (Fig. 3); against it and extended. The movementoften sniffing preceded cheek rubbing, scuffing, and alternates with sniffing, or licking and flehmen, spraying in a decreasing proportion of cases, and there is a tendency for the same side of the while urine spraying and cheek rubbing were the headto be used several times before switching to most common and second most commontermi- the opposite side. For example, the sequence of nal acts. In Pallas cats clawing and urine spraying actions in an adult male snow leopard in re- are coupled together, and Schaller (1972) re- sponse to an unknownscentin the soil was: sniff, ports that head rubbing mayprecedespraying by right, right, left, sniff, left, pause (22 sec) sniff, male Serengeti lions, particularly at spraying right, sniff and lick, right, right, left, lick, left, sites used by other males. stand. This recumbent rubbing, which is also Table 3 compares someattributes of the six characterized by salivation, may lead to rolling different scent-marking patterns. Urine spraying and writhing on the sides and back. There are is distinctive in being a one-directional marking two basic differences between recumbent head pattern, which by itself does nothing to modify rubbing and the other versions. The formeris the sender’s body odor. Clawing involves fore- clearly evoked by strong novel odors suchascar- limb contact, but it is doubtful that much of the 758 Communication in Selected Groups

Table 3 A comparison between characteristics of scent-marking and anointing patterns commonto the Felidae.

Material Substrate Anointing Habitual Possible information Pattern deposited disturbance material site usage potential

Urine spray Urine None None Yes Individual identity, sexual identity and condition.

Head and neck Saliva, skin None Saliva, glandular Yes Individual identity. rubbing exudates exudates

Scuffing Urine, feces, Considerable Urine Yes Individual identity, sexual skin exudates identity and condition. Clawing Skin exudates Considerable None Yes Individual identity.

Recumbenthead Saliva, skin Minimal Carrion, feces, No Individual identity, sexual and neck rubbing exudates vomit, catmint identity and condition. Rolling Skin exudates Minimal Carrion,feces, No ? vomit, catmint cat’s personal odoris imparted to the substrate. radically changed, and anytrace of the animal’s Like scuffing, however, the behavior may consid- scentleft at the site must be slight compared with erably modify the substrate, and it is hkely that the intensity of the foreign scent source. the odors emanating from the lacerated bark and In passing, mention should be madeof moti- scraped soil modify and present a ‘“‘disturbance”’ vation and function. The difficulty in under- to the olfactory landscape. This substrate modifi- standing these aspects of felid scent marking is cation probably creates a detectable secondary that markingis often not characteristically linked cue that intensifies perceptbility of the signal’s to other motivationally distinctive (aggressive, location. sexual, or fearful) behavior. Recumbent head Cheek rubbing and scufhing,associated with rubbing and rolling is an exception. Todd (in urination, however, transfer body secretions Palen and Goddard, 1966) studied this pattern in (saliva and urine) to other body regions, namely, domestic cats as a response to catmint; he the head, cheeks, neck, shoulders, chest, and showedthatas a specific responseto this plantits hind feet. These parts of the body arealso tar- expression is controlled by a singleallele, and he gets of companion-oriented contact, a problem regarded it as a sexual display. to be considered in the following pages. Recum- Palen and Goddard revealed that as a reac- bent rubbing androlling, on the other hand, tion to refined catmint(trans-cis-nepetalactone) clearly differ from the above patterns in being rubbingand rolling were independentof sex and directed to decomposing animal matter. They gonadal state. The occurrence of the pattern in differ from each otherin the extent to which the the estrus female and as a reaction to this chem1- body is covered with the foreign scent, but it 1s cal led them to conclude that both conditions important to rememberthat these behaviors may increase skin sensitivity on the head. Thus, the occur in tandem.In any case, an overriding re- occurrence of catmint-evoked rubbing androll- sult of either act is that the cat’s body odoris ing is noncyclical in both sexes, while in the

Communication in the Felidae 759

pee

IASou aN se

7 ok M e 7 Ae 5ae tsLe ', hePParr cA Preter Bi) | hipaa am ee - (a ~ ee if “et ‘§er 7, 2 ole s ae 4 3 2 an pt tie Om ag: 3 oe 5 OP 4 N ES yeAB - fe” tw, tt : » 2 '

A Seek ‘

ia? > : 2 URINE\Y SPRAY

J CLAW>LICK/BITE

Fig. 3. Sequencing of scent-marking patterns in a tions from one pattern to another. For example the male snow leopard (Panthera uncia). The thickness of scuff-to-urine-spray arrow represents 66% of all tran- the arrowsis proportional to the percentageoftransi- sitions originating with scuff.

female spontaneous rubbing androlling cycles the proestrous female during this activity and with the estrousperiod.If the pattern evolved as attempts to mount her. The female’s orgiastic an olfactory signal the male would be expected postcoital rolling frequently provokes forelimb to sniff sites where the female has rubbed. sparring with the male. Therefore the display Palen and Goddard did not investigate the appears to be visual rather than olfactory; when pattern from a zoosemiotic standpoint; however, sniffing and licking occur in the sexual context Michael (1961) reported that the male watches they are directed to the female’s head and neck, 760 Communication in Selected Groups and the vulva. Possibly the orientation to the several points of contact exist between two or anterior bodyareas is reinforced by heightened more animals, the bodies are usually aligned in rubbing during proestrus. characteristic configurations. The configurations Less is known concerningtheother patterns. restrict the range of targets selected for contact Leyhausen (1965a) remarkedthat while territory (Wemmerand Fleming, 1974). In the following markingis often interpreted as having a warning pages the distributions of four contact patterns function in solitary mammals, there is no indica- (rubbing, sniffing, biting, and patting with the tion that a cat is intimidated uponsniffing a com- forepaw)to the bodytargets of siblings are com- panion’s scent mark. The receiver’s responseis pared amongthe leopard cat, sand cat, and Pal- neither overt nor immediate, and probablyde- las cat. pendson the identity of the ownerandits rela- Body rubbing betweensiblings and between tionship to the receiver. The owner of a fresh siblings and adults was infrequently seen in the mark might be aggressively sought by a domi- leopard cats (N=7) and sand cats (N=14). How- nant male receiver, while the mark left by an ever, 187 incidents of rubbing were recorded in estrous female might arouse sexual interest in the Pallas cats. In all three species the pattern the same animal. The assumption herethatre- resembles the body rubbing of domestic cats. mains to be tested is that cats can identify indi- The initiator presses the side of the head and viduals and sexual condition by olfactory traces neck or the torso against the companion’s body. in urine and skin secretion. Other hypotheses The position of the recipient’s body (standing, also require testing. When head rubbingfollows sitting, lying) to an extent determines the area urine spraying, is some additional information that is rubbed. Several juxtapositions are possi- incorporatedat the site? It 1s possible that differ- ble, but for simplicity we can considerfour basic ent sequence-linked patterns simply overlap in situations ranked in decreasing order of occur- message content and serveas cross-referencing rence: body to body (79%), body to head-neck or redundancy function (Birdwhistell, 1970)? (8.5%), head-neck to head-neck (6.4%), and head-neck to body (5.3%). The body-to-body category is clearly divisible into two groupings. Companion-Oriented Contact Kittens pressed their sides against the sitting or standing mother’s breast and lowerthroat re- Contact between animals takes many forms, gion on 81 occasions (43%), while the remaining and probably not all of them are communicative. body-to-body rubbing for the most part oc- At one extreme there are the fleeting, infre- curred between siblings and consisted of one quent, and seemingly inadvertent contacts that cat’s rubbingits side against the other’s. result from proximity due to a common concern Of the remainingthree patterns, snifing was or activity. An example is the crowding and exhibited least often (Fig. 4). Though it is diffi- bumpingof bodies that accompanythe flight of cult to assess the relative importance of compan- a family group into a burrow. Sender and re- ion-oriented sniffing in different species, it 1s our ceiver are difficult to delineate, and the points of impression that it is not as prevalent in these contact often vary without a predominatingpat- small cats as it is in other carnivores. For exam- tern. Most contact, however, is clearly inten- ple, in observed undersimilar condi- tional, of variable duration, and specifically tions the overall sniffing rate was 7.9/hr (data oriented to a part of the companion’s body. It from Wemmerand Fleming, 1974). In the leop- may be a single brief, one-directional act, but if ard cats the rate was 2.3/hr andin the Pallas cats Communication in the Felidae 761

LEOPARD CAT

CONTACT

TOTAL SAND CAT

OF N=519

PERCENTAGE

PALLAS CAT N=40 SNIFF BITE PAT

Fig. 4. The percentage distribution of sniffing, neck, b = forelimbs (including feet), c = torso, d = biting, and patting to general bodyareasofsiblings in hind limbs(including feet), e = ano-genital region, f the leopard cat, sand cat, and Pallas cat. a = head- = tail.

2.1/hr (data for sand cats are not in comparable the leopard cats and least sniffed in the sandcats, form). but the sample size for the latter species was In all three species the head-neck and tail small. In all species the ano-genital region was rank high amongthe general bodyareassniffed infrequently checked, which is in marked con- (Fig. 4); however, there are differences among trast to other species of carnivores, particularly species. In the leopard cats and sand cats the canids and viverrids (Ewer, 1973; Wemmer, in head-neck was the area most often sniffed, while press; Wemmer and Fleming, 1974). Michael this area and the torso received equal attention (1961) has observed in fact that the ano-genital in the Pallas cats. The torso washighly sniffed in region of anestrous domesticcats has a repelling 762 Communication in Selected Groups effect when sniffed by sexually mature males. Discussion Whenthehistogramsfor these three species are comparedstatistically a significant positive cor- Scent transfer between cat and environment relation is found for the leopard cats and Pallas is mediated through elimination patterns and cats (r, = .940, P < .05, Spearmanrankcorrela- body contact with inanimate objects. Notall tion coefhcient). Unfortunately, the small num- parts of the body are used in scent marking ber of sand cat observations are inadequate for throughcontact. For example, the head and neck Statistical treatment. are employed in two distinct patternsby all the In all three species the head-neck is the most species reviewed, while scuffing seemsto be re- frequent biting target, with the torso ranking stricted at least to the Pantherinae and cheetah, second and third in importance (Fig. 4). Pallas but is probably absent in the genus Felis (Table and leopard cats differ from the sand cat in that 2). The zone of contact between body and object the tail is the second most commonbitingtarget. has properties similar in some ways to a mechan- All other areas (limbs and ano-genital region) ical joint or an articular facet (I. Golani, pers. receive a relatively small portion of bites com- comm.). The relationship between the object pared with these three areas. The ranked distri- and other, more distal body parts can vary con- bution of biting in the Pallas and leopardcats are siderably, but movementat the joint is morere- identical, and a correlation of .829 (P = .05) was stricted. Contact between animals can be found between the sand cats and leopard cats, regarded in a similar way. The body has certain and sand cats andPallas cats. When specific areas focal points for contact, and indeed it can be of the torso are considered there is no general visualized as a field of valences; the shape or agreementin biting (Fig. 5). Only in the sand cat “relief” of the field may differ for different types did one body area receive more than 27% of all of contact(sniffing, biting, or licking). The distri- bites. In the leopard cat the rump wasthe fa- bution of valences (or the form of the relief) is vored target, and in the Pallas cat the side and probably determined on the one hand by topo- belly. In all three species chest and haunches graphical features such as the location of sense were always low-rankingtargets. In comparison organs and glandular areas, and on the other by with the general picture no specific target pre- characteristics of body movement and orienta- dominates the torso (Fig. 5). Between species tion to one another. tests of specific head-neck and bodytargets (Fig. Recumbent head and neck rubbinginthefel- 5) producerelatively high correlations, but only ids has analogs of varying similarity in certain Pallas cat and sand cat exhibit a statistically sig- membersofall but one family of the , nificant correlation at P = .Q1. including three species of (Viverridae: Fig. 4 showsthat patting in the three species Wemmer, in press; Ewer and Wemmer, 1974), is directed mainly to the forebody, infrequently the spotted (Kruuk, 1972), various to the hindlegs and ano-genital region, but more (, 1971), the polar , and the (a often to the tail. Ear, cheek, and neck stand out memberof the , pers. obs.). The spe- amongthe specific head-neck targets, while back cific use of the head and neck in these patterns and side predominatein the torso (Fig. 5). There suggests the hypothesis that the additional scent are no significant correlations among head re- may enhancethe attractiveness of those regions gions of the three species; however, values for for certain types of contact received from com- leopard cats and Pallas cats, and leopard cats and panions. In all three cat species the head-neck sand cats were significantly correlated (P > .O1, region was the predominant sniffing target, > .05, respectively). thoughthetail also received considerable atten- Communication in the Felidae 763

LEOPARD CAT

N=155

ab c d e f

SAND CAT N=32

abcde f

PALLAS CAT

an N=15 ~N

CONTACT

TOTAL

OF

LEOPARD CAT

PERCENTAGE an

SAND CAT

i j ki PALLAS CAT N=39 SNIFF BITE Fig. 5. The percentage distribution of sniffing, three lines). a = chest, b = shoulder, c = back, d biting, and patting to specific regions of the torso (up- side, e = belly, f = haunch, g = rump, h ear, per three 1 lines) and to specific head regions (lower nose, j} = cheek, k = throat, 1 = neck. tion. The head-neck zonealso received most of was frequently observed only in the Pallas cats, = the bites, as well as a substantial proportion of involved the head-neck zone but minimally pats from the forelimbs. The data do not make (about 20%). In other words, support the hy- a compelling defense for the hypothesis. Fur- pothesis gains from the sniffing data is weakened thermore, anointing is only occasionally seen by the biting and patting data. even in captive situations, and rubbing, which Other species show similar trends whether 764 Communication in Selected Groups head and neck anointing1s in their repertory or the vulva of proestrousand estrous females. On not. The frequently and vigorously the whole, however, cats are not particularly ori- anoints the cheeks and neck with a variety of ented to this part of the body. substances, includingits food; up to 80% of all Small felids dispatch their prey with a fatal snaps andbites are directed to this contrastingly bite to the nape of the neck; the orientation to marked region, which is also presented to the the neck constriction is innate, but the precise biting animal. However, the same area receives orientation to the nape of the neck and the nec- less than 40% of all sniffing (Ewer and Wemmer, essary pressure for a lethal bite must be learned 1974; Wemmer,in press). In the , a viverrid through experience with live prey (Leyhausen, carnivore in which anointingis weakly developed 1960, 1965b). The youngcats of all three species and neck presentation is absent, these areas re- directed their bites to the neck more often than ceive about 70% of the bites and about 25% of to other parts, a situation that implies that this the sniffing (Wemmer,in press). Head and neck innate orientation 1s also operantin sibling in- rubbingis absent in the (a social-living teractions. However, the Pallas cats directed ), but these areas receive about 40% mostbites to the throat, while the other two spe- and 35% of all bites and sniffs (Wemmer and cies oriented more strongly to the nape (Fig. 5). Fleming, 1974). In most of the examples the Differences in neck structure undoubtedly con- head is the most commonsniffing target and the tribute to this behavioral contrast. The Pallas cat head and neck the prevailing biting target. The is a relatively short-necked species, and the con- anterior location of the sense organs undoubt- dition is particularly pronounced in the kittens. edly determines the predominance of thesere- The throat is probably more vulnerable than the gions during contact, but skin secretions and nape, but postural differences during interaction possibly strange scents may provide olfactory in- probably also play role. formation. This is supported by the observation There are other similarities and contrasts that the ear is the most commonlysniffed part of amongspecies and patterns for which explana- the head in the Pallas cats and sand cats (Fig. 5). tions are not apparent. The contact mapping Schaffer (in Kleiman and Eisenberg, 1973) re- methodprovidesa refined description for a cate- ports that the auditory meatusin cats is glandu- gory of behavior that often receiveslittle atten- lar, but his observations are based on the tion, but the general picture of contact we have domestic cat, and interspecific differences are presentedis the sum of occurrences from various likely to exist. contexts and postural configurations during The anal glands of the domestic cat and the early development. Contact is also integrated sand cat are voided during traumatic experi- with other concurrent communicative activities. ences, and the scent has a pungent, unpleasant No doubta better understanding of targetselec- odor. There is no indication that the glands are tion will be gained by relating modeand target used in scent marking,either alone or in combi- of contact to these variables. nation with urine spraying. The anal regionis glandular in many mammals, and in somespe- cies it is highly attractive to conspecifics (Fox, References 1971). There are twosituations in whichcats are Altmann, J., 1974. Observational study of behavior attentive to the ano-genital region. Motherssniff sampling methods. Behaviour, 197:227-67. andlick this region in theirinfants and ingestthe Berrie, P. M., 1973. Ecology andstatus of lynx (Lynx milk feces, and males respondin a similar way to canadensis) in interior Alaska. In: The World's Cats, Communication in the Felidae 765

vol. 1, R. Eaton, ed. LagunaHills, Cal.: Lion Coun- solitary mammals. Symp. Zool. Soc. London, 14:249- try Safari, pp.4—41. 63. Birdwhistell, R. L., 1970. Kinesics and Context. New Leyhausen, P., 1965b. On the function of the relative York: Ballantine Books. hierarchy of moods (as exemplified by the phylo- Bossert, W. H., and Wilson, E. O., 1963. The analysis genetic and ontogenetic development of prey- of olfactory communication among animals. J. Theo- catching in carnivores). In: Motivation of Human and ret. Biol., 5:443-69. Animal Behavior: An Ethological View, K. Lorenz and P. Eaton, R. L., 1970. Group interactions, spacing and Leyhausen, trans. B. A. Tonkin. New York: Van territoriality in . Z. Tierpsychol., 27(4):481- Nostrand Reinhold, 1973, pp.120-43. 91. Lindemann, W., 1955. Uber die Jugendentwicklung Eisenberg, J. F., 1970. A splendid predator doesits beim Luchs (Lynx L. lynx. Kerr.) und bei der Wild- own thing untroubled by man. Smithson. 1:48-53. katze (Felis s. silvestris Schreb.). Behaviour, 8:1-45. Eisenberg, J. F., 1973. Mammalian social systems: are Marler, P., 1965. Communication in monkeys and primate social systems unique? Symp. Fourth Int. apes. In: Primate Behavior: Field Studies ofMonkeys and Congr. Primat., 1:232-49. Apes, 1. DeVore, ed. New York: Holt, Rinehart, and Ewer, R. F., 1973. The Carnivores. Ithaca: Cornell Uni- Winston, pp.544-84. versity Press. Marler, P., 1967. Animal communication signals. Ewer, R. F., 1974. Panel discussion. In: The World’s Science, 157:769-74. Cats, vol. 2, R. Eaton, ed. Seattle: Woodland Park Marler, P., 1970. Vocalizations of East African mon- Zoo. keys. I. Red. colobus. Folia Primat., 13:81-91. Ewer, R. F., and Wemmer,C. 1974. The behaviourin Michael, R. P., 1961. Observations upon the sexual captivity of the African civet, Civettictis civetta behaviour of the domestic cat (Felis catus L.) under (Schreber). Z. Tierpsychol., 34:359-94. laboratory conditions. Behaviour, 18:1-24. Fiedler, E., 1957. Beobachtungen zum Markierungs- Muckenhirn,N. A., and Eisenberg,J. F., 1973. Home verhalten einiger Sdugetiere. Z. Sdugetierk, 22:57- rangesand predation of the Ceylon leopard. In: The 76. World's Cats, vol. 1, R. Eaton, ed. LagunaHills, Cal.: Fox, M. W., 1971. Behaviour of Dogs, and Related Lion Country Safari, pp.142-75. Canids. London: Jonathan Cape. Palen, G. F., and Goddard, G. F., 1966. Catnip and Hemmer, H., 1968. Untersuchungen zur Stammes- estrous behaviorin the cat. Anim. Behav., 14:372-77. geschichte der Pantherkatzen (Pantherinae). TeilII, Provost, E. E.; Nelson, C. A.; and Marshall, A. Studien zur Ethologie D.; des Nebelparders Neofelis 1973. Population dynamics and behaviorin the bob- nebulosa (Griffith 1821) und des Irbis Uncia uncia cat. In: The World’s Cats, vol. 1, R. Eaton, ed. Laguna (Schreber 1775). Verdffentlichungen der Zoologischen Hills, Cal.: Lion Country Safari, pp.42-67. Staatssammlang Miinchen, 12:155-242. Rabb,G. B., 1959. Reproductive and vocal behaviorin Hemmer, H., 1972. Uncia uncia. Mammalian Species, No. captive pumas. 20:1-5. /. ., 40:616-17. Schaffer, J., 1940. Die Hautdriisenorgane der Sduge- Hemmer, H., 1974. Studien zur Systematik und Biolo- tiere. Berlin: Urban and Schwarzenberg. gie der Sandkatze (Felis margarita Loche, 1858). Z. des Kolner Zoo., 17(1):11-20. Schaller, G. B., 1967. The Deer and the Tiger. Chicago: University of Chicago Hornocker, M. G., 1969. Winterterritoriality in moun- Press. tain lions. J. Wildl. Mgt., 33:457-64. Schaller, G. B., 1972. The Serengeti Lion. Chicago: Uni- Kleiman,D. G., and Eisenberg,J. F., 1973. Compari- versity of Chicago Press. sons of canid andfelid social systems from an evolu- Schenkel, R., 1968. Toten Léwen ihre Artgenossen? tionary perspective. Anim. Behav., 21(4):637-59. Umschau in Wissenschaft und Technik, 6:172-74. Kruuk, H., 1972. The . Chicago: Univer- Seidensticker, J. C.; Hornocker, M. G.; Wiles, W. V.; sity of Chicago Press. Messick, J. P.; 1973. Mountain lion social organiza- Leyhausen, P., 1956. Das Verhalten der Katzen. Hand- tion in the Idaho primitive area. Wildlife Mono- buch der Zoologie, 8:1-34; Z. Tierpsychol., 16:666-70. graphs No. 35:1-60. Leyhausen, P., 1960. Verhaltensstudien an Katzen. Z. Verberne, G., 1970. Beobachtungen und Versuche Tierpsychol. Beiheft 2:1-120. liber das Flehmen katzenartiger Raubtiere. Z. Tier- Leyhausen, P., 1965a. The communal organization of psychol., 27:802-27. 766 Communication in Selected Groups

Wemmer, C., in press. Comparative ethology of the Verberne, Gerda, 1976. Chemocommunication large spotted genet, Genetta tigrina, and related viv- among domestic cats, mediatedby the olfactory and errid genera. Smithsonian Contributions to vomeronasal senses. II. The relation between the Zoology. function of Jacobson’s organ (vomeronasal organ) Wemmer,C., and Fleming, M. J., 1974. Ontogeny of and flehmen behavior. Z. Tierpsychol., 42:113-28. playful contact in a social mongoose, the meerkat, Verberne, Gerda, and Leyhausen, P., 1976. Marking Suricata suricatta. Amer. Zool., 14:415-26. behaviour of some Viverridae and Felidae time- interval analysis of the marking pattern. Behaviour, 58(3-4):192-253. Since the writing of this article, three importantarti- Verberne, Gerda, and de Boer, Jaap, 1976. Chemo- cles have appeared on thetopic of felid olfactory com- communication among domestic cats, mediated by munication that should be consulted by interested the olfactory and vomeronasal senses. I. Chemo- readers. communication. Z. Tierpsychol., 42:86-109.