1977-Wemmer.Pdf

Chapter 29 COMMUNICATIONIN THE FELIDAE 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 cats 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 lion have been studied from an ethological stand- (Panthera leo), Indian tiger (P. tigris), leopard (P. point. Because of the scarcity of information, pardus), jaguar (P. onca), snow leopard(P. uncia), moststudies have been of a general nature, and clouded leopard (Neofelis nebulosa), cheetah few have been addressedto particular aspects of (Acinonyx jubatus), puma (Puma concolor), lynx communicative behavior. Critical, detailed inves- (Lynx lynx canadensis), sand cat (Felix margarita), tigation of auditory, visual, and tactile modes of wildcat (F. silvestris), Pallas’ cat (F. manul), interaction is to our knowledge greatly under- golden cat (Profelis temmincki), leopard cat represented. At the time of writing there are only (Prionailurus bengalensis), fishing cat (P. viverinus), a handful of studies published on olfactory as- margay (Pardofelis wiedu), Jaguarundi (Her- pects of communication (Fiedler, 1957; Palen petailurus yagouaroundi). A focal animal 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 animals. 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 mammals (e.g., red colobus monkey: Eisenberg (1973) tabulated twenty-five visual Marler,

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