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Facial Expressions and Vocalizations of Rhesus Macaques (Macaca Mulatta) Author(S): Sarah R

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Facial Expressions and Vocalizations of Rhesus Macaques (Macaca Mulatta) Author(S): Sarah R Single and Multichannel Signal Composition: Facial Expressions and Vocalizations of Rhesus Macaques (Macaca mulatta) Author(s): Sarah R. Partan Reviewed work(s): Source: Behaviour, Vol. 139, No. 8 (Aug., 2002), pp. 993-1027 Published by: BRILL Stable URL: http://www.jstor.org/stable/4535968 . Accessed: 28/07/2012 15:18 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp . JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. BRILL is collaborating with JSTOR to digitize, preserve and extend access to Behaviour. http://www.jstor.org SINGLE AND MULTICHANNEL SIGNAL COMPOSITION: FACIAL EXPRESSIONS AND VOCALIZATIONS OF RHESUS MACAQUES (MACACA MULATTA) by SARAH R. PARTAN12) (Animal BehaviorGroup, University of California,Davis) (Acc. 7-IV-2002) Summary A methodfor simultaneouslyexamining visual and vocal componentsof expressivebehavior is described, compiled from video recordings of social behavior of a free-ranginggroup of rhesus macaques on Cayo Santiago, Puerto Rico. I developed a catalog of expressive movements, and chronicled detailed informationon visual and vocal components of 1215 individual behaviors. Two thirds of the events recorded were silent, supportingthe idea 1)Department of Psychology, Universityof SouthFlorida St. Petersburg,1400 South Seventh Avenue, St. Petersburg,FL 33701, USA. e-mail: [email protected] 2) This work is based on my dissertationresearch in Peter Marler'slaboratory. I would like to give my heartfelt thanks to Dr. Marler for his encouragement,support, and advice, as well as for his comments on several drafts of the manuscript.I am gratefulto Marc Hauser for encouraging me to work on rhesus monkeys, and to William Mason for helping me to understandrhesus behavior during pilot observationsat the CaliforniaRegional Primate Research Center.This project was developed during discussions with Drs. Marler,Mason, Hauser,Christopher Evans and Joseph Macedonia.I would like to thank Charles Snowdon, CorreighGreene, Jill Soha, Katya Partan,and two anonymousreviewers for comments on the currentmanuscript; Dr. Mason, John Endler,and Arlene Alvaradofor commentson prior versions; and J.A.R.A.M. van Hooff for a suggestion on terminology.Neal Willits and Tim Allis provided key statistical advice; Renee Allen and Marc Fourriergave indispensable assistance in the field, and Jeannine Logan, Rebecca Wylie, and Virginia Price in the laboratory;Bill Patrickand Todd Hughes helped produce Figure 4. Finally, I would like to thank John Berard,Matt Kessler, the University of PuertoRico, and the CaribbeanPrimate ResearchCenter for providing access to the field site and the genealogical data, and Edgar Davila for introducingme to the monkeys.This study was fundedby grantsfrom the National Science Foundation,the L.S.B. Leakey Foundation,the Animal BehaviorSociety, and Sigma xi. ( KoninklijkeBrill NV, Leiden, 2002 Behaviour139, 993-1027 Also availableonline - 994 PARTAN that visual behaviors are primary for short distance communication in these macaques. Clustersof expressive components detected by PrincipalComponent Analysis and Multiple CorrespondenceAnalyses correspondedto threatening,submissive, and affiliativebehaviors describedpreviously, providing quantitative support both for these previous descriptionsand for the suggestion that these threepoles of behaviorare importantin daily social interaction. Silent expressions involved a greater variety of mouth positions than did vocalizations, which were produced with stereotyped mouth shapes. Other components of the face, not involved with articulation,were nonethelessassociated with particularvocalizations: specific associationswere found among barks,ears retracted,and head lowered on the one hand, and pant-threats,ears forward, and head raised on the other. Screams and squeaks were highly stereotyped,combined with prototypicalgrimace mouthpositions, crouchingand retreating. Girney vocalizations were accompanied by lipsmacking. Grunts were unaccompaniedby other expressivecomponents, evoking the suggestion thatthey may be predominantlyneutral in valence. Introduction Although communicationinvolves the use of multiple sensory channels, most researchersfocus on one sensory channel at a time, in isolation from the full repertoireof the animal. The combination of channels, however, can have importantramifications for signal meaning and efficacy (Marler, 1965; Partan& Marler, 1999; Rowe, 1999). Human facial expression and visual articulatorymovements, for example,play a role in speech perception (McGurk & MacDonald, 1976; Massaro, 1998). In birds, odor cues from prey can interact with visual stimuli of particularcolors to produce food aversionsthat do not occur without the odor (Rowe & Guilford, 1996), and odor plays an importantrole in combinationwith other cues during sexual behaviorin baboons (Goldfoot, 1982). Signal componentsin multichannel displays can be redundant(e.g. Conner,1987), or each componentmay play an independentrole (e.g. Fusani et al., 1997). It is difficult, however, to determinethe functionalrole of each componentof multicomponentsignals, as Green (1975, p. 87) mentioned for Japanese macaques: "...their vocal behavior is inextricably tied to simultaneous olfactory, tactile, and visual signals, hence considerationsolely of evoked responses cannot disentangle the roles of the concurrentsignals available by different sensory moda- lities .. Before parsingthe role of each componentin communication,one needs to determinewhich particularcomponents are combinedsimultaneously into multimodalsignals duringsignal production.Although researchers studying SIGNAL COMPOSITION 995 the receptiveside of communicationhave developed sophisticatedmethods for measuringthe perceptionof multichannelsignals (e.g. visual influences on human speech perception,Massaro, 1998), there are to date few estab- lished methodsfor quantifyingnatural multichannel signal production. This study describes how naturalvisual signals of facial expression and body postureare associatedwith vocalizationsof rhesus macaques(Macaca mulatta),bridging the separateanalyses of visual and vocal signals of this species (e.g. primarilyvisual analysesby Hinde & Rowell, 1962; van Hooff, 1962; Maxim, 1982, 1985; Zeller, 1986, 1996; Maestripieri& Wallen, 1997; and primarily vocal analyses by Rowell, 1962; Gouzoules et al., 1984; Hauser, 1991, 1996; Hauser& Marler, 1993). Although some authorshave described visual and vocal signals together (e.g. Altmann, 1962; Rowell & Hinde, 1962; Lindberg, 1971; Mason, 1985; Kalin et al., 1992), none have reportedthe frequencieswith which particularvisual componentsare associated with particularvocal components.This is an importantfirst step towards an understandingof the multimodalnature of communication.A similar approachhas been taken by Adams & Schoel (1982) in a study of stumptailmacaque (Macaca arctoides) communication:vocalizations were analyzed separatelybut in parallelwith the simultaneousfacial and postural componentsof behavior.My work also builds on the work of Hauseret al. (1993), who suggestedthat each class of rhesusvocalization is accompanied by an unique articulatorygesture. Here I not only examine mouth position, but also include facial expressionsand body posturesuninvolved with actual phonationthat nonethelesspredictably accompany each vocalization. One goal is to explorethe variabilityin facial expressivecomponents used during communication.I constructeda catalog of expressive movements of the face and head, grouping behaviors into 'morphological'categories (classifiedby body part:eye, ear,head, etc.; cf. Reynolds, 1976). Withineach category,I definedlogical, mutuallyexclusive states for each body part(e.g. the ear categoryincludes earsforward, retracted,orflapped backandforth). This categorizationscheme is more detailed than previous rhesus catalogs with regard to facial movements (see Zeller, 1986, 1996, for a detailed look at the face of other Macaca species) but may not be as detailed with regardto gross body movements.Reynolds (1976) thoroughlydocumented and cross-referencedterms used by early rhesus monkey observers, and I attempthere to use many of the terms of my predecessorsin the hope of achieving marginalconsistency and common understanding.My terms for 996 PARTAN the vocalizationsfollow most closely those used by Rowell & Hinde (1962) and Hauseret al. (1993). The behavior of the animals was recordedaction by action in sequence. An 'action' was identifiedas a discrete unit of behavior,which may include multiple simultaneous components. This designation was based on my impressionof a behavioral'unit.' Behavior,although it occurs in continuous streams, can be described as being made up of sequences of more or less discrete actions. The identificationof these action units can, however, be difficult. I separatedthe behavioral stream into discrete actions based partiallyon physical changes in the body (e.g. the head moving from raised to lowered), and partially on timing considerations (if two components of behavior occurred simultaneously, such as head lowered and bark, I consideredthem to be componentsof the same action, whereas if there was a time lag between them,
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