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The semantics of Cebus olivaceus alarm calls : object designation and attribution

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The semantics of Cebus olivaceus alarm calls : object designation and attribution
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Norris, Jeffrey Copeland, 1950-
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Predators ( jstor )
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Signals ( jstor )
Snakes ( jstor )
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THE SEMANTICS OF CEBUS OLVCU ALARM CALLS:
OBJECT DESIGNATION AND ATTRIBUTION


















BY

JEFFREY COPELAND NORRIS



















A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT
OF THE REQUIREMENTS FOR THE DEGREE OF
DOCTOR OF PHILOSOPHY UNIVERSITY OF FLORIDA 1990















ACKNOWLEDGEMENT


This research could not have been completed without the help of many individuals. Drs. John Robinson, John Eisenberg and Jay Whitehead provided guidance and assistance throughout, for which I offer my deepest gratitude. Sr. Tomas Blohm kindly offered me his hospitality at the ranch. He has been steadfast in his support of this and many other research projects, and a good bit of what is known about neotropical ecology is in his debt. Dr. Bill Hardy generously allowed me to use the equipment in the Bioacoustics Lab of the Florida Museum of Natural History. Lastly, I would like to thank two wonderful women, my mother Mrs. J.D. Folsom and Kim Martin, for helping me through many hard times.
















TABLE OF CONTENTS


page

ACKNOWLEDGMENTS ...................................

TABLE OF CONTENTS .................................

LIST OF TABLES .................................... v

LIST OF FIGURES ................................... vi

ABSTRACT .......................................... vii

CHAPTERS

1 INTRODUCTION ................................ 1

Traditional vs. Modern Perspective
on Animal Communication ................... 1
Previous Studies of Cebus olivaceus
Communication ............................. 4
Linguistics and Animal Communication ........ 6 Research Goals and Rationale ................ 39

2 VOCAL RESPONSES TO PREDATORS BY
CEBUS OLIVACEUS ........................... 40

Methods and Materials ....................... 40
Responses of Cebus olivaceus to Predators ... 46 Further Investigations ...................... 56

3 VOCAL RESPONSES TO RELEASED SNAKES .......... 58

Methods and Materials ....................... 58
Results ..................................... 61
Discussion .................................. 75

4 RESPONSES TO ALARM CALL PLAYBACKS ........... 79

Methods and Materials ....................... 79
Results ..................................... 88
Discussion .................................. 93



iii









5 THE ACOUSTICS OF CEBUS OLIVACEUS
AL.ARM4 CALLS................................. 102

Methods and Materials........................ 102
Results..................................... 116
Discussion.................................... 142

6 SUMMARY AND CONCLUSIONS........................ 157


REFERENCES..................................... .... 160

APPENDICES

A NARJROWBAND SPECTROGRAMS OF GRRAHS TO A
0.68 m BOA.................................... 168

B NARROWBAND SPECTROGRAMS OF ALL GRRAH
VA.RIAN~TS...................................... 172

C DEFINITIONS OF ACOUSTIC VARIABLES............. 178

BIOGRAPHICAL SKETCH................................ 180
































iv
















LIST OF TABLES

pages

TABLES

1 ANIMALS THAT ALARM CALL......................... 14

2 PRIMATES EXHIBITING PHONETIC DIFFERENCES
IN THEIR COMMUNICATION SYSTEM................ 25

3 COMPOSITION OF MAIN GROUP....................... 43

4 SNAKES USED IN CONTROLLED RELEASES............. 61

5 CONTEXTS OF GRRAH VARIANTS...................... 68

6 GRRAH DIVERSITY BY INDIVIDUALS.................. 71

7 MATRIX OF GRRAHS PRECEDING OTHER GRRAHS ... 73

8 RESPONSES TO ALARM CALL PLAYBACKS............... 81

9 RESPONSES TO PLAYBACKS OF GRRAHS................ 88

10 RESPONSE RATES FOR CALLS AND INDIVIDUALS 89

11 FUNDAMENTAL FREQUENCIES OF WAAHS AND GRRAHS. 117

12 FUNDAMENTAL FREQUENCIES FOR INDIVIDUALS ... 117

13 DESCRIPTIVE STATISTICS OF ACOUSTICS
PARAMETERS OF CEBUS ALARM CALLS............. 123

14 CORRELATION MATRIX OF ACOUSTIC VARIABLES 126

15 DEFINING VARIABLES FOR GRRAHS.................. 136

16 AMPLITUDE CHANGES AT HARMONIC INTERVALS ... 139

17 FUNDAMENTAL FREQUENCY DIFFERENCES BY SEX 142

18 DISTINCTIVE FEATURES FOR PRIMATE
VOCALIZATIONS................................ 150



V
















LIST OF FIGURES

pa._es

FIGURES

1 STUDY AREA .................................. 41

2 NARROWBAND SPECTROGRAMS OF GRRAHS TO
VARIOUS THREATS .......................... 53

3 NARROWBAND SPECTROGRAMS OF WAAHS ............... 55

4 KEY TO GRRAH VARIANTS ............................ 66

5 DIGITAL OSCILLOGRAM AND ANALOG NARROWBAND
SPECTROGRAM OF JM102 ..................... 109

6 WATERFALL DISPLAY OF JM102, GRRAH VARIANT
1111. (FOURIER DERIVED) .................... 110

7 WATERFALL DISPLAY OF JM102, GRRAH VARIANT
1111. (LPC DERIVED) .........................

8 COMPARISON OF FOURIER AND LPC DERIVED
SPECTRA OF MIDDLE SEGEMENT OF JM102 ...... 112

9 OSCILLOGRAM OF 20 MSEC OF GRRAH
VARIANT 13121 ............................ 118

10 A DFT SPECTRA OF A GRRAH VARIANT 13121 ...... 119 11 A CEPSTRUM WAVEFORM FOR A GRRAH
VARIANT 13121 ............................ 120

12 PLOT OF THE FIRST TWO PRINCIPAL COMPONENTS.. 131

13 CLASSIFICATION OF 59 GRRAHS .................... 133

14 PLOT OF THE FIRST TWO DISCRIMINANT FUNCTIONS 135

15 CLADOGRAM OF ACOUSTIC DISTINCTIVE FEATURES.. 138

16 COMPARISON OF FOURIER AND LPC DERIVED SPECTRA OF GR69, A GRRAH VARIANT 3001.... 146


vi















Abstract of Dissertation Presented to the Graduate School of
the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Doctor of Philosophy

THE SEMANTICS OF CEBUS OLIVACEUS ALARM CALLS:
OBJECT DESIGNATION AND ATTRIBUTION By

Jeffrey Copeland Norris

December, 1990

Chairman: Dr. John F. Eisenberg Major Department: Forest Resources and Conservation (Wildlife and Range Sciences) Wedge-capped capuchin monkeys, Cebus olivaceus, give acoustically distinct alarm calls to different predators. The semantics of these alarm calls were studied in central Venezuela in three stages: 1) through recordings of Cebus olivaceus vocal response to various predators, 2) through release of boa constrictors of three sizes (small, medium, and large) and two quantities (one and two), and 3) through playback of resulting calls. Alarm calls to released snakes were categorized by acoustic features into 15 variants, 2 of which were used solely when a snake was on the ground, and a third when a snake was in a tree. The playback of the locational calls showed that upon hearing a call a capuchin looked in a particular direction, into trees or toward the ground, at the appropriate call. Capuchins use alarm calls



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to not only designate objects but also to attribute qualities such as location to those objects.















































viii















CHAPTER 1
INTRODUCTI ON


The wedge-capped capuchin (Cebidae: -Cebus olivaceus=

Cebus nigrivittatus), also known as the weeper capuchin, is a medium sized neotropical primate inhabiting much of South America north of the the Amazon river (Eisenberg, 1989). As its appellation "weeper" implies, it is a notably vocal animal. Among its many calls are a series of alarm calls given to a variety of predators. These calls are the subject of this dissertation--under what circumstances they are used, their variability, and their meaning. This search for meaning is a difficult one for which I am best warned to remember the story of Captain Cook, recounted in Cherry's seminal work, On Human Communication (1978). When the famous explorer first saw a strange hopping animal he inquired of a native as to what it was. The local gentleman, of course, did not understand English and replied "kangaroo," meaning "I don't know."


Traditional vs. Modern Perspective on Animal Communication


The traditional view of animal communication,

universally held until only recently, is that there is a


1









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fundamental dichotomy between language and animal communication. The difference in perspective involves both what is communicated and how it is done. Traditionally animals were believed to communicate only their motivational states: the signal indicates the sender's arousal or intent for action. According to this perspective, animal calls are affective. For example, an alarm call communicates fear. This is in contrast with what humans communicate, where symbols refer to, among other things, objects as well as emotions.

A referent is the designatum of the signal (Green and

Marler, 1979). Reference may be to either an internal state or to an external physical object. Communication about an object may take any of several forms: iconic, indexical, or symbolic. If variations in the object's physical form are transformed along some parallel acoustic dimension then the communication is iconic (Green and Marler, 1979). For example, an alarm call that increases in duration or amplitude in parallel with increased size of the predator is iconically representing the size of the predator. Indexical communication refers to an object through the simple expedient of pointing at it (Green and Marler, 1979). That is, the communicator refers to object by addressing it physically by pointing at it. In the context of an alarm call, a monkey would indexically communicate about a snake by pointing at it. Symbolic communication uses the signal









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to represent the object, where the signal itself refers to the object (Cherry, 1978). The alarm call itself refers to the predator because the call designates the object through symbolic representation. It is conventionally held that animals cannot symbolically communicate about referents, internal or external, and that if they want to communicate about an object they must do it either by indexing the object, or by using some iconic signal. Altmann (1967) notes the power of combined affective and indexical communication and how prevalent it is among primates. For example, Menzel and Halperin (1975) show that a chimpanzee's walking gate communicates information to other animals about the quality of food sites.

Animal communication has traditionally been seen as fixed, stereotyped, and simple whereas language was perceived as complex, variant, and open. This perspective began to change with the pioneering investigations of bee communication by Karl von Frisch (1967) where it was shown that a bee could symbolically communicate the distance and direction from the hive to a food source. Subsequent investigations in a wide variety of taxa have shown that the traditional view was inadequate. There is mounting evidence that a new perspective describes the behavior of certain species better than the earlier theory. According to this new view, primates use signals to communicate about objects in a manner very similar to how we use words to identify









4

objects. I argue below that primates may not only be able to designate objects but also attribute qualities to those objects. Now we examine the previous studies of the wedgecapped capuchin to see how it fits into this framework.


Previous Studies of Cebus olivaceus Communication


Oppenheimer and Oppenheimer (1973) describe, in a brief study, eleven different calls from C. olivaceus living at Hato Masaquaral, Venezuela, the site of this study. They characterize one call, the grrah as being an inter-specific agonistic call. Of the 60 grrahs they recorded, 48 were directed at humans, 4 to overflying birds, and 2 at howler monkeys (Allouatta seniculus). The remaining 6 were used in unknown circumstances. If Qrrahs were directed at animals other than humans, they were often repeated and at variable intervals. If the call was directed at birds, the authors describe the monkeys as giving the call once then "dropping to lower branches and moving deeper inside the tree" (Oppenheimer and Oppenheimer, 1973, pg 422). They also note that grrahs were associated with other qrrahs 92% of the time.

Robinson (1982) described how three calls--huhs, hehs, and arrawhs, mediate spacing within a group. Arrawhs were given in two contexts. Loud arrawhs were given by an animal if it became separated from the group. Group members responded with more huhs and often replied with arrawhs.









5

Quiet arrawhs were given by an animal if it began lagging behind the group. Robinson concluded that arrawhs acted to reduce spacing between group members.

In a subsequent publication, Robinson (1984) identified five basic vocalizations that were often syntactically combined--chirps, trills, squaws, screams, and whistles. The acoustic parameters that he used to differentiate these social vocalizations are compared to alarm call acoustics in a later section. Robinson first showed that these five calls were used individually and their use varied predictably with social circumstance expressing a continuum of internal states ranging from contact seeking to contact avoidance. The combined calls were relatively common, comprising 38% of the 868 calls recorded. He concluded that "the distribution of social circumstances in which compound calls are given was intermediate between the distributions of the constituent call types, which presumably indicates an intermediate internal state" (Robinson, 1984, pg 76). This study will be considered more fully below in a discussion of how the syntactic structures of primate vocalizations : parallel in important ways those found in human language.

Before continuing I would like to mention a dilemma facing anyone trying to explain what is occurring when primates vocalize. As we shall see, many primates combine vocalizations; yet in my experience in both South America and Africa, monkeys most often give single, often apparently










6

repetitive vocalizations. If the single call is referring to an object, why is it given repetitiously? Correspondingly, if calls can refer to objects, to what do combined calls refer? These questions pose methodological problems of vocal classification and procedures for determining meaning of calls.


Linguistics and Animal Communication


Linguistics is an alternative explanatory system to the traditional model of animal communication (see Snowdon, 1982). Linguistics deals with one form of communication, language, at several levels including semantics, phonology, and syntax. Because linguistic considerations are essential to this project, I will define them, discuss how they apply to animal communication, and give examples of pertinent investigations.

Semantics

Semantics is the study of meaning. In linguistics, meaning is acquired through denotations--how words are defined in a standard text. Words are also understood connotatively through the speakers conception of how a word is used in normal conversation. Words have referents. A word may symbolically refer to a physical object such as a car, whereas other words have internal, abstract referents, for example, anger. Green and Marler (1979) point out that the assessment of any referent is initially an internal










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procedure, with the signaller processing sensory and cognitive inputs prior to the generation of a signal. I note this here because in subsequent discussions I will use the terms external referent and internal referent. It is important to remember that all signals possess internal reference to some degree, where neural inputs are transformed into vocal outputs. Objects of external referents are not simply named without several layers of cognitive processing before the signal is generated. Nevertheless, the word car refers to an external physical object whereas anger refers to an internal state.

The concept of meaning is central to linguistics and

communication. The theme of word and meaning is the subject of a pivotal work by Quine (1960). He notes that "meaning, supposedly is what a sentence shares with its translation; and translation at the present stage turns solely on correlation with non-verbal stimulation" (Quine, 1960, pg 32). It is the problem of translation of a language where no translators or dictionaries provide denotative meaning that prompts Quine to coin the term radical translator. This is the problem of a linguist who meets a previously unknown people or a wildlife biologist who is trying to determine the meaning of an animal's vocalizations. Initial steps at translation are more like correlation, where the vocalization is first translated when paired with conspicuous events. First attempts at detecting meaning are









8

connotative. Quine uses the example of a linguist trying to understand what a native is saying when a rabbit scurries by and the native says gavagai. Is the native in fact signifying food, animal, rabbit, or something else? Meaning or at least a working hypothesis of meaning is made by correlating a reasonable number of stimulus events and the accompanying utterance. The stimulus meaning is the class of all stimulations that prompt that vocalization. If the linguist hears the native say gavagai many times when presented with a rabbit, he may assume that gavagai and rabbit have the same stimulus meaning. Yet the linguist cannot be sure that his translation is correct; there will always be an indeterminacy between stimulus meaning and the denoted meaning. A sort of working dictionary may be compiled but the correspondence of word to stimulus event will be imperfect because this is a mechanism for translating discourse, not single words. Quine goes so far as to state, in the tradition I believe of Turing and G6del, "that rival systems of analytical hypotheses can fit the totality of speech behavior to perfection, and can fit the totality of dispositions to speech behavior as well, and still specify mutually incompatible translations of countless sentences insusceptible of independent control" (Quine, 1960, pg 72). That is to say, given the same text









9

two translators could come up with internally consistent yet mutually incompatible translations.

I return now to the concept of reference, for it is

here that we confront the subject of what animals mean when they vocalize. Take the situation where a primate sees a predator, a snake for example, and utters an alarm call. To what exactly does this vocalization refer? Let us, for the sake of argument, assume that it does not refer simply to an emotion, but rather to the snake. Is the call referring to some concept of snake or to the snake itself? Does the signaller have a concept of snake or is it simply naming objects? What is the difference? The former would indicate that the monkey uses conceptual modes for organizing its world, while the latter indicates that the animal only names objects and is not conscious of the properties that define a snake no matter how precise and accurate its identifications (see Marler, 1982; Cheney and Seyfarth, 1985, for a discussion of how primates and other animals categorize objects in their world).

To describe the dichotomy between a word and its

underlying conceptual reality I will borrow from set theory and logic the notions of intension (not to be confused with intention) and extension (Quine, 1960; Harre, 1984). "The intension of a class is the concept under which any member must fall, while its extension is the set of members which satisfy that requirement. Intentions are concepts,









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extensions are sets of things "1 (Harre, 1984, pg 101). The extensions of a class therefore are the members comprising that class, while intensions are the unifying concepts that define the class. The exclusive use of extensions would therefore mean that the user has no knowledge about what characterizes and differentiates that object from others. Here clearly we are speaking of what the animal knows and intends in its communication.

Quine notes that attributes are monadic intensions, described by the notation "to be an object x such that... X,11 while relations are dyadic intensions, described "to be objects z and y such that . X . y . ."1 (Quine, 1960, pg 164). In other words, for an object to have attributes or be a part of a relationship requires a conceptual grasp--an intensional grasp--of the object. I am therefore distinguishing communication using extensions from communication using intensions. Alarm calls ma~y indicate that the primate is using extensional information, for example, that the predator is a member of a list of predators. Attributing qualities to that predator however proves that the monkey conceptually grasps the defining character of that predator. I would argue that current evidence shows only that primates are able to name an object, x, and there is as yet no evidence to suggest that they are able to talk about such objects. There is still no evidence that nonhuman primates are able to symbolize









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something about x such that x has a property. To paraphrase Bertrand Russell, no matter how adept a vervet may be at identifying its mother, it's not able to explain that its parents are poor but honest. If this view is correct, a monkey's lexicon is comprised of extensions of classes. The animal is not able to conceptually describe aspects of the object or to talk about the object, it only names objects. Correspondingly, if it uses intentions of the class of predators it should be able to go beyond simple object naming and attribute qualities, such as number, size, or location, to the object.

To show that an animal has a conceptual grasp requires that we find situations in which the signaller adds specificity about the object, for example, by using adjectives. Adjectives specify attributes to objects. Quine (1960) describes four levels of reference where objects of reference are further specified, from the general to the specific. In the first phase, objects are named through a process of reinforcement, extinction (learning), and ostension. Ostension is the direct experiential association with the object of reference, for example, that a certain object is called snake by other monkeys. Word usage at this phase is of extensions of a class. In the second phase, these general terms are paired with demonstrative singular terms, for example, this snake or that snake. The third phase compounds general terms with









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attributes, for example, big snake or ground snake. In the fourth phase, analogy is used to apply relative terms to singular or general terms to form other general terms, for example, larger than a small snake. All phases beyond the first reference phase require conceptual knowledge of the object--using these three further levels requires intensional knowledge.

Attribution therefore involves the addition of

information. This may be accomplished in a variety of ways. In language we achieve this through the use of additional words or modifications of existing words. In any event, attribution involves specifying additional meaning to a word. In linguistics this additional element of meaning may be a phone, morpheme, or clause. These terms, to be defined below, are called recurrent partials. To discover the use and meaning of a partial requires finding recurrent use with another signal to similarly modify it. If capuchins use a signal to indicate the location of a snake, for example, in a tree or on the ground, the necessary and sufficient proof of the meaning of the partial (and therefore whether the animals are attributing qualities to objects) is to find it being used in connection with another object that could also be found in a tree or on the ground. Do the monkeys, for example, indicate whether a tayra is in a tree or on the ground using the same signals that they may use when making the same distinction about snakes? The goal of the









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investigation described here is to determine the necessary condition, that is, if _Q. olivaceus use calls to indicate whether they ascribe location to snakes. Subsequent research will be necessary to establish the necessary and sufficient condition of recurrent use of the partial, and therefore whether Cebus olivaceus attribute qualities to objects.

Examples of semantic communication

Ethologists have noted for some time that many birds and mammals use alarm calls, which often are viewed as examples of semantic communication. The studies sited in table 1 are important because they bear on questions of kin selection, evolution, sociality, and animal communication. Two questions are pertinent here : 1) are these vocalizations affective or symbolic?, and 2) is there continuity throughout these taxa in what is referred to by the alarm call. Evidence consistent with the symbolic perspective is the requirement that these signals refer to objects in an arbitrary manner (Sebeok, 1975; Hockett, 1960) and that the signal production is disassociated from its physiological manifestation. A separate view, held largely by those studying various species of ground squirrels, suggests that alarm calls do not classify the predator but rather signify the different time constraints necessary for predator avoidance.









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Table 1. Animals that Alarm Call.


Species Reference

Birds
chukar partridge Stokes, 1961
red-legged partridge Goodwin, 1953
California quail Williams, 1969
turkey Hale et al, 1969
quinea fowl Maier, 1982
domestic chicken Collias and Joos, 1953
Konishi, 1963
Gyger et al, 1986

Rodents
black-tailed prairie dog Hoogland,1983
Richardson's ground squirrel Davis, 1984 thirteen-lined
ground squirrel Schwagmeyer, 1980
Belding's ground squirrel S.R. Robinson, 1980, 1981
California ground squirrel Owings and Virginia,1978 Leger and Owings, 1978 Owings and Leger, 980 Leger and Owings, 1978
arctic ground squirrel Melchior, 1971

Primates
saddle-back tamarins Bartecki and Heymann,1987
black spider monkey Eisenberg, 1976
black-handed spider monkey Chapman et al, 1990 wedge-capped capuchin Oppenheimer and
Oppenheimer, 1973
Robinson, pers.comm.
Norris, pers.obs.
vervet monkey Struhsaker, 1967
Seyfarth et al, 1980
Japanese macaques Green, 1975
rhesus macaques Chapais and Schulman,1980
chimpanzees Marler and Tenaza, 1977









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Alarm calls and other semantic vocalizations have been investigated at length in several primate species, in particular the vervet, Cercopithecus aethiops. Struhsaker (1967) first noted that vervets use a variety of alarm calls, given to at least three different predators. Seyfarth, Cheney, and Marler (1980) further demonstrated through playback experiments that a particular call, for example a martial eagle alarm call, prompted a predictable, adaptive anti-predator response pattern even in the absence of an eagle. Additional adaptive responses were elicited by leopard and python alarm calls, with the response best suited to protect against the differing attack strategies. Each of these three calls was acoustically distinctive. From the perspective of the signaler, particular predators had unique names, while from the receiver's perspective, a certain signal meant a particular predator which required a specific adaptive response.

These signals satisfy the basic requirements of

symbolic communication by being arbitrary, disassociated from physiological manifestation, and non-iconic, (see Altmann, 1967, and Owren, 1985 for discussions of Hockett's defining properties of symbolic communication). The visual image of an eagle, for example, stimulates the vervet and through a process of internal reference to emotive and other dispositions that image is transformed into a signal that prompts an equivalent mental image in a receiver. Cheney,









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Seyfarth and Marler (1980) demonstrated that these vocalizations were not simply affective or prescriptive of particular responses. For example, they demonstrated in the playback experiments that when an animal heard an alarm call, it immediately looked in the appropriate direction for the predator symbolized by the call (for example, into the sky for an eagle) and not to the presumptive signaller to indexically find out about what it was calling. As discussed below, I maintain that the receiver's immediate response to a call--whether looking to the signaller or to the presumed external referent--should be the sine aua non test of reference in field experiments.

In addition to alarm calls, several authors have argued that various primates use symbolic communication in food calls and agonistic recruitment calls (reviewed by Marler, 1985). Food calls are known to be used by chimpanzees (Pan troQlodytes) (Wrangham, 1977; Marler and Tenaza, 1977), toque macaques (Macaca sinica) (Dittus, 1984), black spider monkeys (Ateles fuscipes) (Eisenberg, 1976), and wedgecapped capuchin monkeys (C. olivaceus) (Oppenheimer and Oppenheimer, 1973; Robinson, pers. comm.; Norris, personal observation). These authors do not all maintain that food calls are used as symbols for food. More typically the earlier reports view these calls as affective; the animals are indicating their high motivation to eat when presented with particularly desirable or abundant food. Only Wrangham









17

(1977) and Dittus (1984) explicitly state that the food calls of their study animals are symbolic. In the chimpanzees and toque macaques, and perhaps in other primates, these calls are given when a troop member locates a superabundant food source. Typically, if the food was not superabundant, the call was not given. Calls were apparently nondirected, given to the troop as a whole. In the macaque, when a feeding bout was initiated by a food call, feeding duration was longer than those where no call was given. Dittus concludes these calls "convey information about the presence of a food source, its quantity and location" (Dittus, 1984, pg 476).

Dittus (1984) attempts to show that the macaque food calls are not affective; that they are not communicating only emotional state or probability of future action. He argues that since the stimulus eliciting the call is preci se, specific to the discovery of abundant food in 98% of the calls, the specificity is inherent in the call. I would argue that, on the contrary, since this call is given only when food is abundant, it would unambiguously signal to others that the stimulus, food, is present, regardless of whether the call is referring to the food itself or to the emotional state stimulated by abundant food. In other words, contextual specificity here is not sufficient proof that the signal is symbolic. Dittus has demonstrated that the message of the signal is the presence of abundant food.









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He has not demonstrated the meaning of the signal. A cue to the meaning of the signal is the receiver's immediate response upon hearing the signal. Does it look to the signaller or look for the food? The author states, in his description of response to the food call, that "upon hearing a food call, an individual would immediately stop its activity, glance alertly in the direction of the source of the call and run there to feed" (Dittus, 1984, pp 473). Later he states that "upon hearing a food call, animals rapidly and directly approach the call site and feed there" (Dittus, 1984, pp. 476). This seems to imply that signal receivers may be turning to the signaller to find out what stimulated the call rather than turning to the food itself, i.e. that the food's presence is signalled indexically. A more rigorous test for reference would be to determine the receiver's response when the signaller is remote from the abundant food.

The majority of primate vocalizations are, as Cheney and Seyfarth (1982) point out, chirps, trills, grunts, and screams typically found in social situations. A variety of studies have attempted to determine the reference and therefore the meaning for several of these calls. Gouzoules, Gouzoules, and Marler (1984) describe five screams given by rhesus macaques during social interactions. The authors assert that these screams were representational signals referring to the social rank of the screamer's









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opponent and the likelihood of physical contact in agonistic interactions. For example, where a young animal is signalling that it is interacting with a dominant animal the mother should react more strongly than when her offspring is signalling it is interacting with a lower ranking animal. Through analysis of the probability, duration, and latency of response the authors concluded that the mother responded differentially to four call classes and that the calls referred to the opponent. An alternative hypothesis is that the call refers to the level of fear the animal is feeling during the encounter. The mother's response to a particular call may be a result of, first, her recognizing that the call is from her offspring (through voice recognition) and, second, that levels of fear elicited by the potential opponent's likelihood of harming her infant are encoded in the four calls. For example, one call, one emotion, is stimulated by large dominant animals where the chances of physical contact are high while another call is stimulated by a less dominant animal where contact is unlikely. The stimulus for the call is the same, but the referent is to relative levels of fear, not to the object eliciting the fear.

In a similar study of a series of acoustically similar vervet monkey grunts, Cheney and Seyfarth (1982) demonstrate that vervets may differentiate several grunts that are undifferentiable to humans, and that these grunts may








20

designate objects and events in the external world. Through a paired comparison experiment in the field, the authors tested the hypothesis that grunts convey specific contextindependent information. Five test stimuli were used; three grunts given during intratroop interactions (when meeting a dominant male, a dominant female, a subordinate female), another grunt used when another vervet group was seen, and the last when the signaller saw a troop member initiate motion into the open. Results indicated a differential response pattern to the grunts. While these experiments may establish differential responses independent of context, they do not prove that the signal was interpreted by the receiver as anything other than the signaller was upset or emoting. In fact, the evidence seems to indicate that the receiver looks to the signaller, the speaker in the playbacks, to see what was causing the signaller to grunt. The appropriate experiment to indicate, for example, that one grunt refers to a dominant animal would be to play a call to an animal in the presence of a dominant animal--as if another vervet is grunting about that dominant animal-and determine if the receiver looks to the speaker or to the dominant animal. Another test would be to determine if an animal is greeted with different grunts before and after changing its social rank relative to the vocalizer.

I conclude that the results of the semantic studies of vervet alarm calls demonstrate that vervets use symbolic









21

communication. The authors of the studies on food calls and social vocalizations have not, I believe, sufficiently established their claims of symbolic communication. Alarm calls: models for semantic communication

There could be no better subject than alarm calls in a study of semantics. First, if an alarm call is semantic it will refer to an object, a predator, that is remote in space from the animal. This is essential because when a monkey is responding to, for example, a boa, the snake is probably distant from the monkey and not within the group. Testing alarm call semanticity becomes a geometric consideration. The physical remoteness of predator, caller, and receiver makes response quantification relatively easy because it is clear when the monkey is looking at the predator and when it is looking to other monkeys (or speakers). Other semantic studies of primate calls, for example those on food calls, have had the problem of differentiating when a monkey was looking at another monkey and when it was looking at the food the monkey was eating. (This ambiguity would also occur when testing for response to an alarm call referring to a snake close to the caller.) Secondly, by the nature of predation, response to a predator alarm call must be immediate and adaptive. When a predator is sighted there typically is an immediate need to give an alarm call. Likewise when a monkey hears an alarm call there is an immediate need for response. This need for immediate









22

response and external reference make the study of alarm calls ideal for semantic research. Phonology

Phonology is the study of speech sounds--the phonetics and phonemics of a language. It includes studies of how sounds are made as well as the psychoacoustics of their reception. Where phonetics is used to describe and classify speech sounds, in phonology these descriptions and classifications are used to describe communication systems and explain sound processes (Sloat, Taylor and Hoard, 1978). The basic unit of phonetics is the phone, which simply is any speech sound. The basic unit of phonemics is the phoneme which is the smallest speech unit that distinguishes one linguistic utterance from another.

Acoustic variability may describe phonetic or

phonological differences. The perceived association between the signal and phones is the phonetic quality of the signal. This is in contrast to phonemes which are the abstract class of minimal phonological units consisting of phones (the phonetic unit) that are functionally identical.

If we extend this definition of phonology to the study of all vocal sounds, including those of non-human animals, then we can view each species as potentially having its own phonetic system. Taxonomists of animal vocalizations, lacking (at least initially) a semantic framework to distinguish sounds, face the problem of whether to lump or









23

split similar sounding vocalizations (for a discussion of this taxonomic question see Marler, 1982). There have been several phonetic studies of primate vocalizations that point to a richly variable repertoire where the variability may be described as phonetic.

The diversity of alarm calls seen in this study prompts the central question of my research, namely is this variability meaningful to the monkeys. Green (1975) faced a similar problem when describing the variability of coo vocalizations by Japanese macaques. He adopted the terms continuous/ discrete to describe morphological variability; a continuous call was one in which the gradations of variability were functional and meaningfully significant, whereas a discrete signal had no functionally intermediate forms (Green and Marler, 1979). Owren (1985) provides the useful insight that it is not the presence of variability alone that determines whether a signal is discrete or continuous, rather it is the function of that variability. Determining function is, of course, a difficult problem and may lead to seemingly contradictory results. For example, Winter (1969 a&b) concluded that squirrel monkey (Saimiri sciureus) used a discrete repertoire whereas Schott (1975) concluded that the repertoire was continuous (=graded). Such difficulties are perhaps not surprising. All vocal signals contain some variability because the production mechanism is a biological organ producing a continuously








24

varying energy wave that is subject to fatigue, distortion and other perturbations.

Characterization of speech signals in articulatory or auditory terms is described as distinctive features analysis. Research on the distinctive features of human speech has shown that a relatively small set of parameters can define phonemes (for example, see Ladefoged, 1975). This is an important step in the description of variability because distinctive features provide a mechanism for describing how a continuously varying acoustic signal can be described in discrete phonological units. The mechanisms by which humans and other primates perceive their signals is discussed below, but it is clear that many primates distinguish signal variability in a manner similar to that of humans.

Examples of Rhonetic descriptions

Phonetic differences in vocal behavior have been

described in a variety of primates (Table 2). For example, Cleveland and Snowdon (1982) were able to describe eight chirp variants of a tamarin, Saguinus oedipus, each given in a definably different behavioral context. The authors concluded that each call variant represented a particular motivational state, i.e. that while they might have been affective vocalizations, each call was uniquely paired to a particular motivational state. A distinctive features analysis showed that four statistically significant









25

parameters (presence/absence of a frequency upsweep, difference between peak and end frequency, peak frequency, and duration of frequency downsweep) differentiated the eight calls.


Table 2.

Primates Exhibiting Phonetic Differences
in Their Communication System

Species Reference

Japanese macaque Green, 1975
Talpoin monkey Gautier, 1974
Pygmy marmoset Pola and Snowdon, 1975
Snowdon and Pola, 1978
Cotton-top tamarin Cleveland and Snowdon,1982
Vervet Cheney and Seyfarth, 1982
Seyfarth and Cheney, 1984 Owren and Bernacki, 1988 Owren, 1985
Gelada baboon Richman, 1976
Squirrel monkey Newman et al, 1978
Black spider monkey Eisenberg, 1976
Wedge-capped capuchin Robinson, 1984


In another distinctive features investigation, Owren and Bernacki (1988) were able to define a single acoustic variable, spectral tilt, that correctly classified two Cercopithecus aethiops alarm calls, eagle vs snake, 97% of the time. If the tilt was falling, that is, if the amplitude decreased each successively higher spectral peak, the call was an eagle call, otherwise it was a snake call. Perceptually this means that eagle calls were lower pitched than snake calls because the higher spectral peaks of the snake call were relatively louder. From another









26

perspective, the monkey stressed the higher frequencies of the snake calls and stressed the lower frequencies of the eagle call.

Spectral characteristics could also differentiate these calls. "If two or three spectral peaks occur but do not differ more than 3 dB in amplitude, or one stronger peak above 1750 Hz occurs, it is a snake call. If two or three spectral peaks occur, one peak below 1750 Hz is at least 3 dB stronger than any other peak, and an identical peak occurs above 1750 Hz, it is a eagle call" (Owren and Bernacki, 1988, pg 1933). These features correctly classified the two calls 89% of the time. These features, while including frequency qualities, still primarily depend on the amplitude characteristics best described using the tilt feature. Spectrograms of these two calls (fig. 1, Owren and Bernacki, 1988) show that the snake alarm call is considerably more wide band than the eagle call. That there appears to be no energy above 4 kHz in the eagle call is an artifact of spectrograms, which have a limited amplitude resolution. The energy above 4 kHz, as seen in the spectra, simply is louder in snake calls than in eagle calls resulting in a greater spectral tilt in the eagle calls.

Seyfarth and Cheney (1984) concluded that vervets may use three acoustic features to differentiate four grunts given in different social situations: frequency of the spectral peak associated with the fundamental (F.),









27

frequency of the second peak (F,), and increasing frequency in the second spectral peak. These authors did not measure the spectral tilt of these calls, but an examination of the spectra shows that these frequency features are probably the most parsimonious.

Richman (1976) gave perhaps the most detailed

description of a primate vocal distinctive features, describing the vocal features used by gelada baboons (Theropithecus gelada). He presented a number of spectrograms illustrating a variety of calls with complex acoustical morphology. He reported that geladas produce long strings of alternating ingressive and egressive phonations. Through spectrographic analysis he concluded that they were capable of producing both vowel-like and consonant-like calls. He also described place and manner of articulation for these calls.

Geladas appear to be able to vary the relative

frequency positions of the first two formants, either through divergence of the formants like human back vs front vowels or raising or lowering of the first formant--the low vs high vowel contrast. (I should note that there has been some disagreement regarding the use of the term formant when discussing non-human vocalizations. Please see chapter five, the acoustics section). Geladas can also simultaneously lower both formant frequencies--rounding, in acoustic phonetic terms. Lastly, geladas can selectively









28

dampen and enhance frequency components of the entire spectrum.

Richman further described the distinctions of vocal

onset and finds them similar to those employed by humans in making consonants: gradual, sudden, and fricative. These articulation features produce, in human speech, glides, liquids, stops, and fricatives. He concluded that it appears possible that geladas produce these consonant-like sounds by changes in three places of articulation: labial, a velarlike position, and an intermediate position. The author made no attempt to correlate these acoustic phonetic differences with changes in behavior.

These studies illustrate that calls with similar

acoustic structure often have variants recognizable to the vocalizing animals, though perhaps not to humans. Call variants are often correlated with different behavioral functions. Future taxonomists of animal calls should recognize that phonological variability in animal vocalizations is important yet often very difficult to discern.

Acousticpercerption in Primates

The above examples of how various primates use

linguistic-like modifications of their communication signals assume that the animals perceive these changes. Recently there has been a significant increase in research on animal psycho-acoustics investigating both the signal








29

characteristics attended to by animals and whether perceptual strategies are similar to those employed by humans in speech. Ideally investigators will identify perceptual sensitivities to particular signal characteristics that are of communicative importance.

Investigators have studied species-specific perceptual processing of communication signals in a limited set of old World primates. The results of various studies by Sinnott (1976, 1985) suggest that 1) primates do not discriminate pure tone changes, the vowel cue, nearly as well as humans, and 2) their discriminatory abilities used in differentiating consonants, where both frequency and intensity cues are crucial, are roughly similar to humans.

Petersen and others investigated the perception of coo vocalizations by japanese macaques (Macaca fuscata) (reviewed in Petersen, 1982) finding, for example, similarities in neural lateralization patterns between primates and humans. They also found that peak frequency location, a factor central to discriminating among the various macaque 'coo' vocalizations, shows a right ear advantage. This is in contrast to how macaques hear pitch changes, which shows a left or no ear advantage. These findings are comparable to similar results in humans. There is a right ear advantage in humans for stop consonants and either a right or no ear advantage for vowels. In regard to selective attention to changes of particular acoustic









30

parameters, it was found that, when pitch is varied, there is selective attention to peak frequency but when, in the reverse experiment, peak frequency is varied, there is poor attention to pitch. These results indicate that Japanese macaques take a discretely used acoustic cue, peak position, and use it to partition vocalizations where peak frequency position continuously varies. The perceptual processes involved, selective attention, perceptual compensation, and partitioning, closely match those used by humans in speech perception. These results indicate that both human and nonhuman primates apparently detect communicatively salient features in their signals in similar manners. Syntax

Syntax is the study of word order. Syntax describes the ways in which words are put together to form phrases, clauses, and sentences. Syntax describes another layer of meaning, one provided by word order. It is through syntax that we distinguish 'dog bites man' from 'man bites dog'.

The concept of syntax itself is layered, with the

distinctions centered around the use of meaning. At the linguistic level Chomsky defined syntax as a "system constituted by rules that interact to determine the form and intrinsic meaning of a potentially infinite number of sentences" (Chomsky, 1972, pg 69). At a functional level several students of animal communication (Robinson, 1979, 1984; Snowdon, 1982; Cleveland and Snowdon, 1982) have









31

adopted a more general notion of syntax. To them syntax is simply a system of rules that will generate and predict sequences of signals, thus they view syntax as a probabilistic phenomenon. Altmann (1965) noted for example that in many vocal sequences first-order Markov (predictable) processes describe transition probabilities and are equivalent to grammars.

Marler and Tenaza (1977) differentiated two forms of syntax: phonological (called phonetic by Snowdon, 1982) syntax and lexical syntax. Phonological syntax is a system for arranging communicative components into more than a single pattern, each with a distinctive function. In language it is analogous to word formation through phonemic rearrangement. In lexical syntax "compound signals derive their meaning from the multiplexing of the meanings of the components as used separately or in other combinations" (Marler and Tenaza, 1977, pg 25). Linguistically it is analogous to phrase formation through word combination so that the product is the sum of the meanings of the individual elements.

Several additional points about syntax are pertinent. First, in parallel to the semantic classes of agent, patient, instrument, and object there are syntactic classes such as noun phrase, verb phrase, and determiners. These syntactic classes are necessary for modeling human language. Premack (1985) makes the valuable point "that syntax cannot









32

be derived from semantics. No metamorphosis has been demonstrated for turning the semantic caterpillar into the syntactic butterfly: agent, recipient, and the like, no matter how abstractly construed, will not turn into noun phrase, verb phrase, etc." (pg 284). If primates can, as I suspect, name objects and are able to differentiate agent from patient, there still is no evidence to suggest that they are capable of the purely linguistic differentiation of noun phrase from verb phrase. Secondly, Chomsky's conception of syntax has progressed beyond the definition given above. He now views mental representations, traditionally within the purview of semantics, as being a form of syntax.

The study of the relation of syntactic structures
to models, "pictures," and the like, should be regarded as pure syntax, the study of mental representations, to
be supplemented by a theory of the relation these
mental objects bear to the world . Thus, the shift
-towards a computational theory of mind encompasses a
substantial part of what has been called "semantics" as
well. (Chomsky, 1986, pg 45).

Additionally, Chomsky views universal grammars as being endowments due to innate, biologically determined principles. This is our language faculty, our "language acquisition device, an innate component of the human mind that yields a particular language through interaction with presented experience, a device that converts experience into a system of knowledge attained: knowledge of one or another languagell (Chomsky, 1986, pg 3). In this new formulation I









33

perceive a weakening in his position that animals are categorically incapable of a language; instead, an animal 'language' will be the product of that species' own innate language faculty. All animals convert experience into a system of knowledge, some perhaps using vocal behavior to differentiate objects. Without begging the definition of language, if we accept that some species now use 'words' to identify objects in their environment, the separation between human language and animal communication becomes smaller, though still very real. The difference between our language and the communication systems used by animals may be a difference in degree, not kind. Alarm call syntax

If certain alarm calls are used to indicate

attributions about an object, for example, the location of a snake, they are used in some sense like adjectives. one would expect then that these calls might be associated with other calls. For example, if one vocalization means 'snake' and another means 'ground snake' there must be some additional segment to the latter signal carrying the additional information, either as another call or as an infix in the original signal. In English we modify the meaning of a word both ways, either by using another word, an adjective, or by internally modifying the original word with prefixes or suffixes, e.g. usual and unusual.









34

Investigating such matters requires knowing the phonetic structure of the communication system. All biological signals have variability but only some of it is communicatively meaningful. The alarm call classification scheme described below parses the various alarm calls into a number of variants. Without knowing the phonetic structures of the communication system, I am unable to determine if C. olivaceus alters its signals with internal modifications. I am, however, able to determine whether calls are combined. I will limit this study to the description of associations between grrahs alone because to accurately determine the association between particular alarm calls and other calls, huhs for example, requires knowing how the other calls vary. Examples of syntactic communication

A wide assortment of primates use syntactic

combinations of different calls--long calls of gray-cheeked mangabeys (Cercocebus albigena) (Waser, 1975), chimpanzees (Pan troglodytes) (Marler and Hobbet, 1975), various gibbon species (Tembrock, 1974; Tenaza, 1976; Marshall and Marshall, 1976); alarm calls of cotton-top tamarin (Saguinus oedipus) (Cleveland and Snowdon, 1982); intragroup calls in the pygmy marmoset (Cebuella Dvgmea) (Snowdon, 1982) and wedge-capped capuchin monkey (g. olivaceus) (Robinson, 1984) and "singing" in titi monkeys (Callicebus moloch) (Moynihan, 1966; Robinson, 1979). Some of these combinatory signals use lexical syntax to impart additional information to the









35

signals, while in others the combined signal elements have a separate meaning--phonological syntax.

Cleveland and Snowdon (1982) demonstrated lexical syntax in cotton-top tamarins (Sacruinus oedinus). They found that two frequently used calls, an alarm chirp and a low arousal alerting call, were combined such that animals responded to this combined call as if it was the sum of the constituent elements.

Robinson (1979) described the remarkably complex vocal behavior of titi monkeys (Callicebus moloch). He found an elaborate hierarchical system where calls are repeated to form phrases, which may then be variably combined to form sequences. He distinguished six types of sequences by their different structure, as disclosed by transition probabilities, their situational context, and the sex of the caller. For example he describes duetting as "an alternation of pant and bellow phrases follows the introductory moaning phrase. As the sequence continues the animals begin to add pumps, either after pants or bellows, and insert honks between pant phrases and after pumps.. Honking usually ends the sequence" (Robinson, 1979, pg 393).

The driving force behind sequencing, and the effect sequences have on meaning remains unknown; without such knowledge we cannot determine if these syntactic patterns are lexical or phonological. Robinson (1979) notes that the hierarchies found in titi monkey calls are similar to those









36

derived by phrase structure grammars. Further analysis of sequencing is severely hampered by our failure to understand what the vocalizations mean; we lack a semantic context to place the calls. Chomsky differentiated phrase structure grammars from more simple sequential models by the characteristics of nested dependencies: 'if. . then' structures. Syntactic structure may depend on the occurrence of particular words, with independent order for intervening words (see Robinson, 1979). Alternatively, responses to different sequences may be due to different proportions of phrases and not to their syntactic order. Robinson (1979) points out, however, that titi monkeys clearly distinguish phrase types and concludes that variation in phrase order is important and has communicatory significance.

Robinson (1984) has also investigated the syntactic

structures in vocalizations of Q. olivaceus. He found that these monkeys use lexical compounding rules to generate new sequences, where the new compound call is produced in contexts intermediate between those of the constituent vocalizations. Additionally, he found a second, more simple sequence type where two call types are blended to form a transitional third call. These new calls are without apparent syntactic structure. He concluded that these lexical syntactic rules are analogous to the rules we use to generate words from morphemes, they are not analogous to the grammatical rules of language.









37

The studies of syntax in both Callicebus and C. olivaceus by Robinson are important because they best illustrate that at least some primates use rudimentary ordering schemes in vocal production and these syntactic patterning rules may be similar to those used by humans.

Several problems emerge from the above description of linguistics and primate communication. I have posited that primates are capable of naming objects but to date there is no evidence that they speak about them. If this is the case, syntax poses a particular problem because syntax provides another layer of meaning upon the individual lexical elements. Is syntax a linguistic operator requiring one form for extensional terms and another for intensional? The linguistic elements syntactically combined may imply differences in conceptual understanding. Evidence for this would be how elements are syntactically combined. Syntactic combinations of extensional terms would involve combinations of simple names and an operator, whereas a combination of intensional terms would involve, for example, analogy or other mechanisms requiring a conceptual grasp of the combined words. I believe there is a conceptual difference between the syntax of 'man bites dog' vs 'dog bites man' and that occurring when water is lexically combined with bird to form waterbird (where waterbird is used in a context separate from water and other birds to refer to a duck). The former construction requires only an ability to name two









38

objects and an action, whereas the latter seems to require a conceptual grasp of the terms.

The above duality suggests a novel hypothesis about how animals communicate about emotions and objects using both affective and symbolic communication. This communication system would use, first, purely affective vocalizations to communicate about emotions and combine those signals to refer to intermediate emotions, for example, the lexically and phonologically combined vocalizations in the capuchin and titi monkeys, and, second, use symbolic terms to refer to objects, for example, alarm calls. According to this hypothesis primates would combine calls to form other calls referring to intermediate emotions using lexical syntax and restrict their use of symbolic terms for named objects. Sequences would be restricted to combinations of affective vocalizations, where combining the emotive signals acts to fluidly transpose the emotions into a third transitional emotion that would have its own subjective reality. Primates would give symbolic calls singly (or repetitions of the same 'word') because they refer to an identifiable object and they would not use them in a sentential form because sentences cannot refer simply to objects. Quine (1960) argued that only words have reference and that sentences are identified by their logical truth. If my hypothesis is correct, it may explain much of the primate vocal behavior described above. They have, first, words for









39

objects and, second, calls and sequences for emotions, each reflecting an underlying reality of objects and feelings. The remainder of this study will address representational communication by g. olivaceus to determine whether they have words for objects.


Research Goals and Rationale


The goal of this research is twofold: 1) to establish whether . olivaceus semantically uses alarm calls to refer to objects and, if so, 2) whether it is then able to attribute qualities to those objects.

In this first chapter I reviewed the literature and

described the problems currently prominent in the study of primate communication. In Chapter 2 1 describe the alarm calls of C. olivaceus. In Chapter 3 1 describe the first of two experiments on the semantics of a particular alarm call type, the snake alarm call. The first experiment involves release of different sizes and numbers of snakes. In chapter

4 1 describe the second experiment where I play recordings of the alarm calls given to the released snakes back to the monkeys. The purpose of the first experiment is to determine if C. olivaceus uses alarm calls as names for particular predators and in the second experiment whether it attributes qualities to them. In Chapter 5 1 describe the acoustic structure of C. olivaceus alarm calls. I finish with a general summary of findings.














CHAPTER 2

VOCAL RESPONSES TO PREDATORS BY CEBUS OLIVACEUS


Methods and Materials


Research Site

This project was conducted at Hato Masaguaral in three periods: preliminary observations from May-July, 1986; snake release experiments from March-August, 1988; and playback experiments from April-August, 1989. Hato Masaguaral is an active cattle ranch as well as wildlife refuge owned by Sr. Tomas Blohin and is located 146 miles south southwest of Caracas in central Venezuela (8*34'N, 670351W). The study site was a 4 kin2 gallery forest (figure 1). It was bordered to the east by a seasonal river, the Cafio Caracol and, respectively, to the north and south by two recently deforested ranches, Finca Torres and Hato Flores Morades. To the west the gallery forest gradually became a more open forest. The study site seasonally flooded from approximately May to October after which it dried and the deciduous canopy became increasingly open. Rainfall in the region averaged 1450 mm/ year. The mean temperature was approximately 28*C. See Troth (1979) for a more detailed description of the ranch.



40










41




















NATO MASAGUARAL




N






NATO MASAGUARAL

















NATO FLORES MORADAS

0So
.S te'si







Figure 1. Study area showing adjoining ranches, the
bordering streams, and the trail system. Marks on trails
are at 25 m intervals (Robinson, 1986).









42

Recordings and Data Analysis Earuipment

Two recording systems were used. Initially, from May to July, 1986, a Sony TC-D5M cassette recorder and a Sennheiser MKh-805 directional microphone were used ad libitum to record monkey calls. Recording distances were less than 15m. The system's frequency sensitivity was flat from approximately 50-15,000 Hz. These recordings were used to determine the monkeys lexicon.

A Sony 8mm CCD-V9 video camera with a Sennheiser MK-415 directional microphone was used during the snake releases and alarm call playbacks as both the auditory and visual recording device. The camera image sensor produces 380,000 pixels and the audio dynamic range is >80 dB over a frequency range of 30-15,000Hz. The camera has multiple shutter speeds which allowed for high speed filming. Video recordings were played back on a 19"1 Sony Trinitron TV. Study Animals

The C. olivaceus studied here were part of a population of approximately 300 animals that inhabit Hato Masaguaral. This population contains approximately 12 groups, 4 of which were studied--Main, White, Red, and Splinter. Most work was done with Main group, which contained 21-27 animals throughout the study: a single adult male, 5-8 adult females, and assorted juveniles and infants. Table 3 shows its composition at the beginning of the 1989 field season. These groups have been continuously studied by Robinson









43

(1981, 1982, 1984, 1986) from 1976 to the present time. All animals in the four groups were individually identifiable by name, with identification made possible by age/ sex, size, and individual markings. All animals, particularly Main group, were habituated to my presence. This made it possible to study the animals at extremely close range, usually 210m. Familial relationships within these four groups have been known for at least ten years.



Table 3.
Composition of Main Group Males Females
Name Age* Name Age*

White Beard >10 Mo 18
Jefe 8 Amelia 17
Griffin 8 Beaut 17
Stu 7 Whitey 12
Mani 5 Pointy Face 12
Babas 4 Alexandra 8
Winston 3 Onica 7
Mike 1 Margo 7
Hanna 5
Mali 5
Puffy 4
Amanda 3
Modem 3

age as of 1989



Predators of Cebus Olivaceus at Hato Masaguaral

As Cheney and Wrangham (1987) note, predation on

primates is rarely observed. There have been no confirmed instances of predation on C. olivaceus at the ranch. The Harpy eagle (Harvia harovia) and the boa constrictor









44

(Constrictor constrictor) are the only confirmed predators of _. olivaceus (Rettig, 1978; Chapman, 1986). Boas are common on the ranch, but this eagle species does not occur there. Aside from human predation through hunting which is negligible at the ranch, presumed predators are aerial, terrestrial, or arboreal.

The largest raptor found at Hato Masaguaral is the

ornate hawk-eagle (Spizaetus ornatus). This bird is almost certainly capable of taking gbus as prey. Other potential avian predators are the spectacled owl (Pulsatrix perspicillata), great horned owl (Bubo virginianus), savanna hawk (Heterospizias meridionalis), zone-tailed hawk (Buteo albonotatus), black collared hawk (Busarellus nigricollis), collared forest falcon (Micrastur semitorquatus), laughing falcon (Herpetotheres cachinnans), and the road-side hawk (Buteo magnirostris).

The only strictly terrestrial predator likely to take C. olivaceus at Hato Masaguaral is the dog, Canis familiaris. All other potential predators are capable of following g. olivaceus into the trees. There are four felids at the ranch; jaguar (Felis onca), cougar (Puma concolor), ocelot (E. pardalis), and jaguarundi (f. yagouaroundi). Tayra (Eira barbara) have been seen chasing capuchins (Robinson, pers.comm.). In my experience, the relationship between tayra and C. olivaceus is apparently context sensitive; among the same individual animals during









45

some interactions the monkeys chase the tayra, while during others the monkeys appear quite alarmed, and during still others they apparently ignore each other.

The boa constictor (Constrictor constrictor) is the only large snake living in the forest at Hato Masaguaral that is likely to take C. olivaceus. Other potentially dangerous snakes are anacondas (Eunectes murinus) and a rattlesnake (Crotalus durissus) which is rarely found inside the forest.

Estimated predation levels on C. olivaceus at Masaguaral are moderate to low. In seven years of observations on approximately 175 animals Robinson estimated that 15 animals were taken by unknown predators, an estimated annual predation rate of 3% (Robinson in: Cheney and Wrangham, 1987, pg. 232). Predation rates were highest on smaller animals; of those presumed taken 70% were infants and 20% juveniles with the remainder split between adult males and females. Reuter (1986) discusses the influence of predation on foraging behavior of C. olivaceus.

I should emphasize that intraspecific aggression

appears to be a major source of mortality among the study animals, and is probably greater than predation. For example, in 1988 a new male, WB, entered Main group and within 6 weeks the four newborn infants had all disappeared, with one confirmed case of infanticide by that new male (Valderrama et al, 1990).









46

olivaceus appear to be predators on bird eggs and hatchlings during the nesting season. Such behaviors typically elicit active nest guarding. I observed Main group attack, kill, and eat a fledgling yellow-knobbed curassow (Crax daubentoni) and attempt to attack a nestling giant potoo (Nyctibius grandis), which was actively defended by an adult. Pairs of road-side hawks called and swooped at C. olivaceus most often during the nesting season. Active nest defense may be the reason that capuchins gave alarm calls at these relatively small raptors. Responses to attacks by nest guarding hawks presumably do not markedly differ from predation events.


Responses of Cebus olivaceus to Predators Anti-Predator Behavior

Responses to predators, both vocal and otherwise, varied according to the predator and the context of the interaction. The description of crrah usage by Oppenheimer and Oppenheimer (1973) is different from my experience in several ways. First, I consider qrrahs to be specifically alarm calls, not simply agonistic calls. Secondly, the Oppenheimers differentiated only a single alarm call, apparently lumping all alarm calls as grrahs. Lastly, I rarely saw capuchins take more vigorous anti-predator actions than just giving alarm calls. Also, in many interactions between C. olivaceus and howler monkeys









47

(Allouatta seniculus), I never heard an alarm call. In the case of raptors, Oppenheimer and Oppenheimer (1973) write that when a bird swiftly flew over the animals one or more C. olivaceus would give a single Qrrah then drop to lower branches or move deeper into the trees. I differ with the Oppenheimers by distinguishing alarms calls to birds as being acoustically distinct from other alarm calls, with waahs given to avian threats and Qrrahs given to terrestrial and arboreal threats.

In the presence of avian threats the monkeys typically gave a single waah, looked up, and on rare occasions moved away from the perceived threat. A variety of birds elicited waahs from C. olivaceus: ornate hawk eagles, road-side hawks, zoned tailed hawks, forest falcons, laughing falcons and various species of vultures. I should note that with some birds the monkeys waahed only after they appeared to be surprised as the birds flushed in front of them or as they flew low overhead: chachalacas (Ortalis ruficauda), currosows (Crax daubentoni), egrets, ibis, muscovy ducks (Oxvura dominica), and macaws (Ara macao). I will recount, from field notes, two examples of interactions between C. olivaceus and hawks.

On one occasion a road-side hawk attacked a juvenile
female crossing a tree gap on a vine. As the hawk
approached, other animals waahed then the hawk hit the
monkey's dorsal side without apparently hurting it.

On another occasion an adult female carrying a newborn was moving on a branch when a laughing falcon (that had been displaced by a harrassing monkeys) flew 2-3 m over









48

it. Other animals waahed and she imediately swung
beneath the branch to interpose the branch between the falcon and herself. It was not clear whether the bird was attacking the female or simply flying away from the
harassing individuals.

This was the only observed active defense, other than vocalizations, against avian predators. When the physical threat was nonavian the situation was quite different. Mobbing behavior, rarely directed towards raptors, was regularly used against non-avian threats. Q. olivaceus have been observed mobbing jaguar, puma, ocelot, tayra, boa constrictors, and humans. Additionally, they arrahed at donkeys, cows, and deer, though this appeared to happen only when the animal ran through the forest, presumably frightening the monkeys.

In a typical interaction, multiple animals cluster

near the potential predator and give many cirrahs. At least in the case of boas, which often remain still and therefore cryptic around monkeys, the animal discovering the snake often remains near it through much of the interaction, giving the most arrahs while other troop members move by also emitting alarm calls. Under these conditions when. a predator is mobbed there may be hundreds of crrrahs given.

The emission of grrahs is a highly directed behavior. Typically the alarming animal faces the threat and gives numerous calls in a clearly agitated manner. These may or may not elicit calls from other monkeys. Grrahs appear to function most clearly as an alarm, warning other animals of









49

the presence of the threat. That they serve to warn other animals of the presence and even location of the predator is the subject of the experiments described below.

On one occasion I observed White group mob an ocelot (Felis pardalis). The ocelot was walking on the ground while the monkeys followed it in the trees, crrrahing as they went. The monkeys appeared to be highly agitated whereas the cat appeared quite indifferent. A similar incident was witnessed with Red group. In both cases the monkeys were already in trees while the cat was on the ground, many animals vocalized, and the ocelot showed little interest in the monkeys.

On a variety of occasions I observed tayra (Eira barbara) move among White, Red, and Main group. For example, just as I was preparing to conduct a playback experiment with Red group I heard a grrah and turned to see a young tayra approach the area on the ground. I was standing below an adult male, Finger, and several other monkeys, less than 10m distance. The tayra saw me and approached to investigate. The tayra appeared to ignore the monkeys who casually watched it as they continued to feed. At another time, for two weeks Main group inhabited a region that was also frequented by a tayra. Sometimes the two species would amicably occupy the same tree while on other occasions the monkeys would chase the tayra out. on still another occasion I saw an adult tayra move through the same









50

trees as capuchins without incident. Lastly, I observed Main group approach a mother and infant tayra in a palm tree. When the adult tayra saw me it tried to leave by coming to the ground and running away, however the infant wouldn't leave and when the mother appeared to abandon it the C. olivaceus threatened the young tayra. This prompted the mother to return, which caused the Cebus to back off. The monkeys gave few qrrahs but gave many other threatening calls during the interaction. J. Robinson (personal comm.) reported seeing an adult tayra chasing C. olivaceus through the trees. I conclude that tayra are a potential threat to C. olivaceus, but the context of the interaction was important as to whether the monkeys reacted.

The response of animals hearing an alarm call is

typically immediate and marked. For example, I was filming a subadult male, GR, as he sat near me on a branch. Main group was spread out around us. A monkey approximately 60m away saw a tayra and Qrrahed. Griffin immediately turned to look at the caller and began scanning that same distant area.

Van Schaik and van Noordwijk (1989) described the role of male Cebus apella and C. albifrons in predator avoidance. They measured differences in vigilance and related behaviours between sexes of both species and, finding a difference, interpreted it as a special male role in predator avoidance. The same anti-predator behaviors of










51

~olivaceus were not measured here, specifically vigilance for predators. It was my impression however that males were not more likely to give alarm calls or mob a predator. As discussed below, in vocal responses to released snakes, females gave many more alarm calls than males, which is different from what van Schaik found in the other Cebus species.

The differences in response by C. olivaceus to avian

and terrestrial predators, particularly the number of calls given, is explained well in a geometric model of alarm call behavior (Taylor et al.., 1990). The authors show that slow moving predators should elicit more calls than fast moving predators. This was found to be the case here where waahs were typically given once while many grrahs were often given.

Alarm Calls

Cebus olivaceus give alarm calls to a variety of

animals, as is clear from the above descriptions. While I divide these calls into two types, waahs and Qrrahs, there appears to be great variability within these call types.I will examine in greater detail how C. olivaceus respond to snakes and the variability within the snake alarm calls in subsequent chapters. Here I will provide examples of arrahs and waahs given to a variety of predators and apparent threats. These calls can be compared to other C. olivaceus calls presented in Robinson (1982, 1984).









52

Grrahs

Graahs, in general, can be distinguished from other C. olivaceus calls by two characterstics: 1) falling frequency and 2) a breathy aspiration sound at the end. Other C. olivaceus calls are downswept, chirps and hehs are good examples, but the terminal aperiodic breathy sound is distinctive to arrahs. The amount of frequency drop and number of energy bands is highly variable, as will be seen in later chapters.

Grrahs to snakes. Figure 2a illustrates narrow band spectrograms of two qrrahs given by an adult male, SH, in Red group to a boa constrictor (Constrictor constrictor). Note that they have 3-4 continuously falling energy bands with the fundamental frequency beginning at approximately 1 kHz. Note also that the calls are less periodic, more noisy at the end. Figure 2b is another spectrogram of a qrrah to a boa by Quay, an adult male in Splinter group. In this call the pitch remains relatively steady until the end where it first climbs then precipitously falls. Note again the broad band noise at the end, particularly in the second formant. The acoustics of Qrrahs to snakes is the subject of chapter 5 which provides examples of many other grrah variants. Notice that the calls just described are similar to some of those given by Main group.

Graahs to humans. Unhabituated and, on occasion,

habituated capuchins would emit grrahs at humans. Figure 2c









53






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54

is a grrah at me given by an adult female, Irma, in White group. Note that it is noisy at the end and downswept.

Graahs to other animals. Figure 2d illustrates two Qrrahs to an ocelot (Felis pardalis) from an unidentified animal in White group as it watched the cat walk under it. Both calls have two energy bands and drop discontinuously. The terminal aspiration is slight. The pure tone at approximately 1.5 kHz is an artifact.

On several occasions C. olivaceus would arrah at

seemingly anomalous objects. Figure 2e provides two qrrahs to a donkey given by unknown animals. These calls are quite distinctive with a long fundamental having multiple energy bands only in the middle of the call after which the frequency drops.

Waahs

Waahs are, in general, longer duration, slightly lower in frequency and less forceful than grrahs. Like cirrahs the pitch falls, however since waahs typically have longer duration the slope is less. They may also be quite breathy or noisy. While the sample size for waahs is much smaller than for grrahs it is quite apparent there is also significant variability among waahs. The semantic utility of this variability is unknown.

Waahs to Road-side hawks. Waahs are most often

given to the road-side hawk (Buteo magnirostris). Figure 3a illustrates two waahs by adult females Gertrude and Irma











55







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IV I







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340














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8 4









56

from White group. These calls typically have multiple energy bands, which like arrahs, fall in the later half of the call. Notice however that the higher energy bands in some calls may be stressed.

Waahs to vultures. Figure 3b presents two waahs by

the dominant male in Red group, Finger, to a vulture, probably a turkey vulture (Cathartes aura). Again these are relatively long duration calls that fall in pitch through the majority of the call.

Waahs to caracara. Figure 3c is a waah to a crested

caracara (Caracara cheriway) by the adult male Brow from White group. This call is shorter in duration, with the second formant stressed.


Further InvestiQations


The anti-predator behavior of C. olivaceus suggests that, like the vervet (Cercopithecus aethiops), the alarm calls described above are used in a semantic manner, with the call referring to an object, in this case a predator, and not simply to an emotion. Demonstrating that C. olivaceus use these calls in a semantic manner requires investigating how a monkey responds to an alarm call. I therefore undertook a series of experiments to verify that C. olivaceus use their alarm calls to refer to objects. I first obtained a series of recordings of known animals calling at a predator and then played those calls to other









57

known animals to determine where they looked upon hearing the signal. These experiments replicate and extend Seyfarth, Cheney, and Marler's (1980) experiments on vervets. I hypothesized that if capuchins named objects, e.g. snakes, they might also attribute qualities to those objects. For example, would they vocally distinguish one snake from two, small from large, or provide information on a snake's location? To test these hypotheses, I released different numbers and sizes of snakes then recorded the monkeys responses. Following that I played to them selected calls to determine if a listening animal responded appropriately.















CHAPTER 3
VOCAL RESPONSES TO RELEASED SNAKES Methods and Materials


Data Analysis Methods

The alarm calls recorded during snake releases were digitized at 20,000 samples/ second using a Digital Translation DT2821 analog/digital- digital/analog board in a Compaq Portable II computer using the RDA and LDA routines in the ILS spectrum analysis system. Where necessary these calls were digitally filtered using ILS. These signal were then played into and analyzed on a Kay Elemetrics model 7029A spectrograph. The spectrograms were narrow band (resolution= 45Hz), 80-8,000 Hz bandwidth, with FL1 shaping.

An essential element to the analysis of these calls was the identity of the caller. This was determined by

1) identifying an animal as it calls on video,

2) identifying the caller in the field and calling out
its name while recording,

3) determining who was in the area during the
interaction and attributing caller identification
by listening, a process of elimination,

4) where calls sounded the same to me, I visually
identified the group of animals that was present
and calling and then compared the fundamental
frequency for similar calls by those same animals
at other times. This attribution was made by
comparing fundamental frequency of calls.

58









59

Where none of the procedures worked, I discarded the

call. Most of the attribution were done with the first two methods. Using these procedures I identified the caller for 190 of 279 arrahs recorded during releases.

The following data were input into the Q & A data base management program to correlate the various calls with the contexts of the snake releases,

1) Caller distance to snake and nearest neighbornear, medium, far

2) Identity of nearest neighbors

3) Number of snakes

4) Snake location (tree or ground) and size.

Additional information included vocalization number, call class (variant type, see below) and caller. Release Methodology and Release Schedule

Recordings of arrahs to snakes were made, with a single exception, by releasing a snake or snakes in front of Main group. In the exceptional case, the monkeys discovered a boa in a tree, I video recorded the interaction, then captured that snake for a later controlled release. In a controlled release, I approached the monkeys, stayed with them for an appropriate time to make sure my presence was not alarming, then removed the snake or snakes from a cloth bag in front of the monkeys. In some situations where I wanted to obtain recordings from a particular animal, I made sure it was alone or had few companions before releasing a









60

snake in front of it. Usually the snake began moving away, often into trees. I pulled the snake down to the ground if it began to move beyond my reach. Typically, the monkeys immediately began calling and mobbing the snake when it was released. Sometimes many other animals approached while at other times only the animals that first saw the snake called. On a few occasions the monkeys did not apparently see the snake and left without vocalizing. On still other occasions they obviously saw the snake but did not immediately begin to call.

There were eight boa releases in the following order, listed by snake length: medium, large, medium, large and medium, small, large and medium, large and medium (to a lone monkey), medium and small. Additionally, a second snake species, Drvmarchon corais, was used in a single release. The number of cirrahs recorded for the nine snake releases ranged from 9-58.

Snake Descriptions

Two species of snakes were used: boas (Constrictor

constrictor) and Drvmarchon corais, a yellow phase indigo snake. The ]2. corais, a snake very similar to the indigo snake, was used to determine if C. olivaceus used alarm calls only at boas or whether they used a general snake call. I used four boas of three sizes (table 4) in combinations of small, medium, large, small and medium, medium and large, and medium and medium.









61


Table 4.
Snakes Used in Controlled Releases

Species Length (m) Weight (kg)
----------------------------------------Boa 1.95 4.65
Boa 1.60 2.00
Boa 1.58 1.94
Boa 0.68 0.20
D.corais 1.80 2.50



Results


Responses by Monkeys to Snakes

I will first describe the release of a 0.68 m boa and provide examples of the grrahs recorded in response to it. The transcription below of my commentary on the video recording shows that while the calls are diverse, several animals give quite similar calls, and that some calls are given in only certain contexts. The commentary also provides a background for understanding the resulting investigations. The vocalizations referred to in the text are presented as spectrograms in appendix A. There were 87 calls during the release, of which 29 were arrah variants. It is clear from the spectrograms that these qrrahs are highly diverse, with many configurations. Five animals called during the 24 minutes interaction.

Snake release commentary

I release the small, 0.68m boa in front of Jefe as
he is sitting alone 4m above me eating a butterfly
larva. For the first 1 minutes he continues eating
without calling even though he appears to see the snake. once he finishes eating he begins calling,









62

JEl0l is his first grrah. He then gives a long series of huhs interspersed with a single grrah after a minute.
By the second minute other animals begin arriving and while other animals in the distance continue giving huhs an arriving animal grrahs, JE 118. By then there are three agitated monkeys over the snake, giving few grrahs. At 2:02 minutes into the interaction, Jefe gives cirrah JEll9, the first ground snake grrah variant (for a discussion of arrah variants, see below). For the next two minutes a juvenile female, Hanna, sits over the snake watching it without giving alarm calls. Jefe then gives another grrah, JEll7, which appears to prompt Hanna to intently look around, otherwise there is little action. Hanna returns to lying on a branch over the snake content in watching it. Other animals continue to huh in the background. overall the troop is relatively placid. Eventually Hanna and another small female moved.
Four minutes into the interaction Jefe gives
another ground snake arrah, JEl2l. Then, after another lag in action, Hanna gives another type of ground snake call, HAlO2, this one a loud brief call like a bark which she repeats 45 seconds later, HA 105. By now the snake, which was released on the ground, movs into water and neither the monkeys nor I can find it. Hanna gives arrah HA122 as she moves lower in a bush, intently trying to locate it.
A minute and a half later the snake begins to swim away, prompting Hanna to give cirrah HA 107. Other monkeys move in and give several desultory arrahs (that are too poor for analysis). Hanna then gives three calls, a loud cirrah, HA 108, followed immediately by HA 109, then another call like the loud call, HA 111. Next she gives four ground snake calls, JE 123, HA112, and JE 124 (one is masked by bird calls so it isn't analyzed), like Jefe's earlier ground snake call, JE119. The interaction is now 10 minutes long. The middle call, HA 112, prompts another monkey to intently look down. The boa then begins to climb into a vine. Hanna gives grrah HA125 followed by two similar calls, JE126 and HAllS. Notice the wide diversity in call types used by Hanna in these last 14 calls.
Over the next 2 minutes there are few calls, yet
Babas, a juvenile male, agitatedly brakes a branch near the snake. They do not appear to know where the snake is. The juvenile monkeys left after two minutes without more calling.
Three minutes after the last grrah Mo approaches, stopping frequently to look down to the ground where the boa had been before it moved into the vines. She is quite evidently looking for something. For two









63

minutes she looks for the snake, at which time I pull
it out of the vine and put it on the ground in front of her. She watches the boa for a minute without calling.
It is now seventeen minutes into the interaction.
Finally she gives a quiet grrah (not analyzed) and
another animal gives a huh. Her next call is MM101, a call quite similar to RA122 by Hanna. A half a minute later she becomes agitated, jumps, and then gives arrah
MM102 while looking down at the boa. Another animal huhs and she responds with MM103 as the snake swims
below her. Fifteen seconds later she gives arrah MM106, an alarm call used for snakes in a tree. I
cannot confirm the snakes location at that time, though
1h minutes later the boa was I meter up a vine. The
last calls recorded from Mo are the two similar calls, MM107 and MM108. At this last call, 30 seconds after
the tree snake call, she does appear to be looking
obliquely at the snake instead of down, i.e. the snake may be in a vine. She then begins to DILh at Hanna who
then rapidly moves down a vine emitting hehs. Hanna appears to direct the calls at the snake rather than
Mo. White Beard, the adult male of the group then
passes below her unperturbed by either the snake or Hanna's aggressive hehs. (On other occasions other
monkeys emitted hehs at boas).
Two minutes after the last grrah, Hanna is lying
on a vine and Mo is self grooming, both monkeys can
plainly see the small boa as it tries to climb a palm.
The interaction ends with Mo moving away, five minutes
after the last grrah.

I conclude from this interaction that 1) C. olivaceus

gave a variety of acoustically distinct alarm calls to

snakes, 2) the alarm call variability was apparently not

random, instead different animals gave apparently similar

calls, 3) the alarm calls Mo heard informed her of the

snake's presence, 4) Mo may have called even before she

herself saw the snake, and 5) the presence of a snake does

not automatically prompt a monkey to call.

By all appearances Mo was looking for the snake. I

should note that this snake release occurred during the










64

early rainy season, August 3, 1988, and there is standing water in many places. The monkeys rarely come to the ground at this time of year so it is highly unlikely that she was looking for food or other monkeys when she scanned the ground. This is noteworthy because the snake had probably not been visible for four minutes.

I interpret Mols behavior to indicate that she understood the various arrahs to refer to a snake. Alternatively she could have seen the snake earlier or seen the other monkeys looking down. Call Types

The fact that different animals made similar calls suggests that the variability represents meaningful differences to the animals. Testing this hypothesis requires a classification schema. I therefore devised a key based on seven arrah features:

1) Format frequency drops/ increases
2) Frequency drop is continuous/ discontinuous 3) Frequency drop is extensive/ not extensive
4) Number of formats
5) Which format was stressed
6) Stressing at end was above/ below the tail
7) Extensive frequency drop in formats other than F1
8) Call duration

These seven features use all three acoustic domainsfrequency, amplitude, and duration. I concentrated on the frequency domain because the calls appeared to vary the most in this domain, and, a Priori, appeared to be the most robust domain to differentiate the calls. Call duration was









65

clearly variable yet duration variability did not appear, in advance of the analysis, to discriminate the calls as well as frequency components. The only amplitude component examined was the relative location of stressing, that is, the position of maximum amplitude. I could not compare calls by their overall amplitude, because the calls were neither recorded at a standard distance nor did I use a calibration tone when recording.

The grrah variant classification schema was defined solely on the basis of the appearance of the spectrograms rather than how the calls sounded to me, however there is, of course, agreement between a spectrogram's appearance and its sound. I assume there are acoustically relevant features that my hearing may not attend to yet may be important to the animals. I further assume that these features will be evident on a spectrogram.

Using these features I parsed the 279 calls recorded during ten snake releases into 15 grrah variants. The key, including the number of grrahs in each variant, is presented in figure 4. Two calls were found only once each and were excluded from further analysis. Initially, as a control on individual variation of call variants, only those calls from known vocalizers were classified, then once a robust classification system was devised all suitable qrrahs were included. Spectrograms of representative samples of each grrah variant are presented in appendix B. While there is










66

l..0.0.0 Formants only drop

1.1..0.0 Formant level then drops

1.1.1..0 Freq. drop cont. and extensive

1.1.1.1 One formant or F1 stressed27

1.1.1.2 F2 stressed 7

1.1.2..0 Freq. drop discont.and extensive 32

1.2..0.0 Formants drop continuously

1.2.1..0 One formant 17

1.2.2..0 Multiple formats

1.2.2.1. F1 stressed 22

1.2.2.2. F2 stressed 8

1.3..0.0 Formant drop not extensive

1.3.1..0 F1 stressed

1.3.1.1. Tail stressed below 36

1.3.1.2. Tail stressed above 17

1.3.1.2.1 F2 extensive drop 34

1.3.1.2.2 Fother extensive drop 6 1.3.2..0 F2 stressed, tail stressed below 27 2..0.0.0 Formants rise and fall 13

2.1..0.0 Formants rise and fall discontinuously 3 3..0.0.0 Others 18

3.0.0.1 Short duration 10

Total 277


Figure 4. Key to grrah variants by spectrographic configuration with totals for each variant.









67

some variability within call types, this schema does appear to group calls into similar appearing classes. Context of the Call

Once the grrahs were classified the specificity of

grrah usage was examined in three contexts: snake location, snake number, and snake size. Table 5 lists the context where each call was given, the number of different releases in which the call was given, and the number of different animals giving the call. The following call variants appear related to certain contexts:

Call Snake Context: On the Ground
Type

1111 26/27 calls by at least 5 animals in 5/8
possible situations, i.e. a snake was on the ground in 8 releases, during 5 of which the
monkeys gave qrrah variant 1111. In the
single exception there were two snakes--one in a vine and the other on the ground. The
call was given by an animal off camera so it was not possible to determine to which snake
the call was directed, i.e. the snake may
have been on the ground.

3001 10/10 calls by at least three animals in 4/8
possible situations.

Overall, when a snake was on the ground 184 calls of 14 types were used by at least 13
different animals.

Snake Context: In Trees

2000 11/13 calls by at least 4 animals in 4/4
situations. In one exception, by Mo, the
snake may have been in a vine, it certainly
was soon afterwards.

13122 5/6 calls by one animal in 1/4 situations.

In total 83 calls of 12 types by at least 10
animals are given to arboreal snakes.












68




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69

Snake Context: Near the Vocalizer

1222 7/8 calls by 4 monkeys in 3/5 situations.

Overall, 107 calls of 13 types by at least lo
animals. 3/8 calls given when snake was on the ground and 5/8 when snake was in a tree.

Snake Context: One Snake

13122 6/6 calls by 2 animals in 2 situations.

Overall, 151 calls of 14 call types by at
least 12 animals.

Snake Context: Two Snakes

2100 3/3 calls by 1 animal in 1 situation.

Overall, 121 calls of 14 types by at least 9
animals.

Snake Context: Medium Sized Snake

13122 6/6 calls by 2 animals in 2 situations.

In total, 101 grrahs were given by at least
13 different animals.

The evidence for other calls being given in particular

contexts is less compelling.

1122 5/7 calls to a snake on the ground, all by
unknown vocalizers in a single release; 2/7
in tree, by whitey in one release and another
by an unknown animal in another release. It
was also given 5/7 when a snake was near.
Its specificity is suspect because of small
sample size and it was given by several
animals with snakes in both locations.

1130 28/32 calls to a snake on the ground by at
least 6 animals in 4/8 situations, however
3/32 calls by 2 different animals in 2
situations were given to snakes in a tree.

1210 14/17 calls to a snake on the ground by at
least 10 animals in 6/8 situations. 3/17 calls were to a snake in a tree, all by a
single animals in one situation.









70

Calls by Age! Sex Class

No call given by more than one caller was given by only a single age/ sex class. There was, however, a significant skew to overall age/ sex class of all callers: 175/ 190 calls were given by adult females, subadult males and juvenile females. The remaining 15/ 190 calls by identified callers were given by adult males (2 calls), juvenile males (9 calls), and infants (4 calls).

There was no significant correlation (Kendall r= -.40, p=0.33) between dominance rank of adult females (the only age class for which sufficient data were available) and number of crrrahs given to released snakes. Dominance was defined operationally by social grooming patterns (T. O'Brien, pers. comm.).

Grrah Diversity

There was no clear pattern to the diversity of grrah usage. Females used a more diverse grrah vocabulary than males, with the Shannon diversity H Ifemaes=l.l3, whereas H'Ixtes=*9l (table 6). Overall diversity was H'=l.12. Notice that females used all variant types whereas males did not use five variants, including one indicating snake location, variant 3001. Notice also that males gave many fewer alarm calls than females, 41 vs 149. Two males, Stu and Griffin, produced almost half of the male data set. These two males are subadults, both born outside the group and approximately second and fourth in dominance among










71

Table 6. Grrah diversity by individuals.


Females Males
C all-- - - - - - - - - -
Type AHa MdMl MoPf PuWh Ba GrJe Mn St WB cf 9
---------------------------------------------------------1111 6 1 2 1 3 3 10
1112 1 1
1120 3 4 3 2 2 2 12
1210 3 5 1 1 1 1 1 1 1 4 11
1221 8 3 1 1 1 1 1 14
1222 2 3 1 2 2 6
1311 5 2 2 7 4 2 1 3 20
1312 2 3 3 2 2 7 5
13121 3 2 9 3 2 5 2 9 17
13122 5 0 5
1321 7 2 4 2 3 2 2 3 1 8 18
2000 8 1 1 1 2 9
2100 3 0 3
3000 3 2 5 1 0 11
3001 3 3 1 0 7
----------------------------------------------------------Tot. 15 22 1 26 47 10 7 21 2 7 9 4 7 10 2 41 149

H'= .45 0 .94 .67 0 .66 .47 0 .91
.87 .83 .65 .91 .56 .45 .59 1.13









72

males. White Beard, the dominant adult male, gave only two alarm calls. This low production of alarms was typical of him and apparently of other adult males, both for alarm calls and other vocalizations. Grrah Syntax

A matrix of grrah variants indicating call pairs is provided in Table 7. The data set is from all identified callers, with call variants modified from that provided in figure 4: variant 3000 includes 3001 and both 1112 and 2110 are excluded due to small sample size. One grrah followed another 158 times, approximately 78% of the time, so the most likely association is between arrahs. The most common association was call duplication, where the same call variant was simply repeated. This happened significantly more often than by chance (X2= 52.37, p= 2.3 E-7, d.f.= 11). This same statistic was not possible with other pairings because of the numerous null cells in the matrix, but it is clear that there are no strong associations between any calls. I conclude that except for duplicating calls C. olivaceus do not combine grrah types in a syntactic manner.

If C. olivaceus do not syntactically combine different qrrahs, do they combine calls to form a new call with intermediate form? This intermediacy in form could indicate that 1) the meanings of the two calls were combined, using phonological syntax or 2) that the signals are graded. If this is phonological syntax, I am suggesting that the













73












T14
O 4 -4N -4N NIV Q Nr-4 N

0
0 W4., 4
0 1-4





N


N ~ L H N H

1-4



41 C

0 -r4. ~ Wr4 r
'-4 N Nm Hr4Cj c


N~. V-4-4H'-4 r

1-4



4i fn C N 14~




o -4

.-I N N V-4 4-q 4N



H 0 HN-4 NN -40
H1- N I r4H f4 r- I rdC% r-4 H r-4 H 1-4 r4 N









74

monkeys are combining elements of different calls to form a new signal, with a new meaning. This would be analogous to word formation in human language. For these calls to be graded (=continuous) requires that the new form follow a continuum where the meaning of the signal is intermediate between the meaning of the two calls from which it is formed. This raises questions that are essentially epistemological, for if these alarm calls are referring to objects, what could be the intermediate meaning, e.g. what is the intermediate meaning for the word snake? For this reason, I doubt whether the calls are graded. Notice however that the physical evidence for both positions, phonological syntax or graded signal, is the same--calls whose spectrographic configuration is intermediate between two other calls. Examining appendix B shows that several variants appear intermediate in form, for example, variant 1130 seems intermediate between variants 1111 and 1210. Even if they were intermediate in form, I have no data to indicate whether they were used in contexts that were in some way intermediate between the contexts in which the other calls were used, that is, that they had an intermediate meaning.

Grrahs to Other Snakes

In the single release in which another snake species was released, two monkeys gave three different call types, including one call, AD11, of qrrah variant 3001, the ground









75

snake call. It seems that g. olivaceus do not restrict alarm calls only to boas.

other Responses

There were other responses to released snakes besides

grrahs. For example, on four occasions monkeys hit a branch while they or another animal were calling and each time a different call was used: variants 2000, 1221, 1210, and 3001. At least twice monkeys knocked the snake out of a tree by hitting the branch it was on.


Discussion


I conclude from the above data that C. olivaceus emit certain arrah variants when snakes are in particular locations: grrah variants 1111 and 3001 when a snake is on the ground, crrrah variant 2000 when it is in a tree, and variant 1222 when the snake is near the caller.

There is, at present, no convincing evidence that C. olivaceus communicate about snake numbers or size. I found the associations between grrah variants and number of snakes unconvincing, primarily because the sample size for each call was small and the calls were given by few animals. Grrah variant 13122 was given 6 times, of which 5 were given by Whitey, while variant 2100 was given only 3 times, all by Mo. A larger sample from more animals is needed before I am convinced of their specific utility. I feel that the specificity apparent in the usage of cirrah variant 13122 is









76

due to its limited application and we can conclude nothing about its usage.

Perhaps one of the most surprising aspects to the

capuchin's reactions to snakes was their calling at small boas in the same manner as they did larger snakes. For some reason the monkeys apparently felt threatened by a .68m snake. Thirty grrahs of nine different variants were given at it, twice as many as were given to the largest snake, which was almost three times as long and over twenty times as heavy.

Grrah variant 1130 was the second most commonly used grrah and apparently used almost exclusively to snakes on the ground. I remain skeptical, however, of concluding its specific usage because it was also used by two animals in two releases when snakes were in trees. Likewise, variant 1210 was used mostly toward snakes on the ground, but was also given to snakes in a tree.

The production of many of the other grrah variants was separated almost equally between one and two snakes or snakes on the ground and snakes in a tree.

Examination of these results shows that the above call variants are not always given when a snake is in the prescribed location. For example, call variant 1111 was given in only 5/8 situations when a snake was on the ground. The snakes location is apparently not a necessary condition for the utterance of a particular call. On the other hand,









77

the emission of a call is sufficient for determining the location of a snake. That is, whenever a monkey gives a arrah, any other monkey hearing it would know where a snake was located. Stated another way, the combined probability of grrah variant 1111 being given when a snake is on the ground is 5 of 8 whereas the probability that a snake is on the ground when arrah variant 1111 is given is 26 of 27.

The three vocational calls were not common; never more than 14% of grrahs given in a particular context. of course, meaning should not depend on the frequency of the call; a call given once would be sufficient to indicate location.

There were fewer calls to snakes in trees than to

snakes on the ground but the average number of calls of any variant per situation was approximately the same. For example, grrah variant 1311 was given 27 times in 7 situations (3.9 calls/ situation) when the snake was on the ground and 9 times in 3 situations (3 calls/ situation) when the snake was in a tree.

Examination of the various grrah variants in appendix B shows that the classification schema is robust, yet there is some overlap; differences between some variants is slight. These similarities may be due to their being syntactic combinations of separate calls or they may be, in fact, distinct calls representing different objects or emotions.









78

The next step in this translation process is to find out how a monkey will respond to these vocational call variants. For example, if grrah variant 3001 is always given to a snake on the ground, and in fact means 'ground snake' then we would expect a monkey to look down to the ground when it hears this call. The experiments described below will serve to confirm the above results.















CHAPTER 4
RESPONSES TO ALARM CALL PLAYBACKS Methods and Materials


Playback, Data Recording. and Analysis EcTuir)ment

These experiments required both playback and recording devices. Calls were played to the monkeys on a Sony TC-D5M audio cassette recorder over two Sony SRS-30a self powered speakers at the end of a 15m cable. The speakers were typically hidden in a green bag placed under leaves in a tree. The recording equipment was the same as that used in the snake releases: Sony video camera and Sennheiser directional microphone. The responses of the monkeys were scored in the field by watching the video recording of the playback through the camera (as a camcorder it was capable of both playback and recording). Results were verified by watching the playbacks on a larger TV.

Immediately after each playback a map was drawn of the experimental scene, indicating the location and height of the monkey, position of other animals, and the distance, angle and height of both the speaker and camera relative to the monkey. This map made it possible to judge where the monkey looked upon hearing a call: down to the ground, up



79









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toward the sky, into foliage, toward the camera, toward the speaker, or toward other animals. Playback Methodology and Schedule Test stimuli

The schedule, call type, call, caller, and receiver for 21 alarm calls of four variants used in playbacks are presented in table S. Each call was selected for clarity from the set of calls recorded during snake releases. Test trials were designed to test the response of single animals to single alarm call playback. Control trials involved either playing 0.5 seconds of noise or setting up the equipment and running a trial with no signal as if a signal were played.

The playback tapes were produced by taking digitized examples of the chosen calls and recording them onto 10 second tape loops. Amplitude of each call was equilibrated by making the maximum amplitude of each oscillogram equal. A single call was presented in each playback.

The calls used in the playbacks were the two ground snake calls, grrah variants 1111 (5 replicates), 3001 (6 replicates), and the tree snake call, graah variant 2000 (7 replicates).

Playback methodology

The major goal in a playback experiment was to present as realistic a stimulus as possible. verisimilitude was essential.









81
Table 8. Responses to Alarm Call Playbacks.
a) Responses to call variants 2000 and 1111 Responses
----m-----------------------m---No
Date Call Sp.Rec. Up/Sky Down Fol. Spkr. Res. Other
-----------------------------------------------------------ARBOREAL CALLS-2000
4/9 HQ125 WH HA X
4/9 HQ125 WH MI X
4/9 HQ125 WH STU X
4/26 MO126 MO AM X
4/26 MO126 MO BA X
4/26 MO126 MO GR X
4/26 MO129 MO Am X
4/27 MO129 MO WH X
4/27 MO129 MO BU X
4/30 MO126 MO ON,Am X
4/30 MO126 MO JE,STU X X
5/22 MG113 WH AM X
5/22 MG113 WH AM X
5/22 MG113 WH AL X
5/31 AB16 MI BA X
5/31 AB16 MI BU X
6/8 MO128 MO WH X
6/8 MO128 MO STU X
6/8 MO128 MO STU X
6/21 MO128 MO STU X
6/21 MO128 MO STU X
6/29 AB67 MO HA X
6/29 AB67 MO WB X
6/29 AB67 MO MO X
----------------------------------------------------------Totals 0 2 7 4 11 1

Ground Calls-lill
4/16 JM102 HA MD X
4/19 BB122 PU GR X
4/19 BB122 PU ML X
4/19 BB122 PU ML X
5/6 AF40 MD WH X
5/20 JE124 HA STU,MN XX
5/25 JE119 JE PF X
5/25 JE119 JE PF X
5/25 JE119 JE STU X
6/10 JE124 HA PF,ML X X
6/10 JE124 HA STU X
6/15 JE124 HA AM X
6/15 JE124 HA WB X
6/15 JE124 HA ML X
6/15 JE124 HA AM,Am X X
Totals 0 8 0 3 5 2-----------------------------------------------------------Totals 0 8 0 3 5 2









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Table 8-continued.
b) Responses to call variant 3001.

Responses
----------------------------------------------No
Date Call Sp. Rec. Up/Sky Down Fol. Spkr. Res. Other
---------------------------------------------------------------Ground Calls-3001
4/22 HA105 HA ML X
4/22 HA105 HA ML X
4/22 HA105 HA MO X
5/4 BB117 PF STU X
5/4 BB117 PF STU X
5/17 BB117 PF BA X
5/17 BB117 PF ON X
5/29 AF27 WH ON X
5/29 AF27 WH AM X
5/29 AF27 WH PU X
6/13 HA105 HA MO X
6/13 HA105 HA WH X
6/13 HA105 HA WB X
6/13 HA105 HA STU X
6/13 HA105 HA WH X
6/17 BB118 PF AM X
6/17 BB118 PF AM X
6/17 BB118 PF BA X
6/17 BB118 PF ML X
6/23 HA105 HA MG X
6/23 HA105 HA BU X
6/26 HA105 HA STU X
6/26 HA105 HA STU X
6/26 HA105 HA WB X
7/6 AD12 HA STU X
7/13 AD12 HA WB X
7/13 AD12 HA AL X
7/15 AD12 HA PF X
7/15 AD12 HA ON X
7/15 AD12 HA ? X
7/15 AD12 HA AM,ON X
7/15 AD12 HA MO X
7/15 AD12 HA MO X
7/15 AD12 HA MO X
7/17 BB126 PF MG X
7/17 BB126 PF AL,ON X
7/17 BB126 PF AL X
Totals 1 6 1 0 22 7----------------------------------------------------------------Totals 1 6 1 0 22 7








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Table 8-continued.
c) Responses to waahs and control trials

Responses
-------------------------------e-No
Date Call Sp. Rec. Up/Sky Down Fol. Spkr. Res. Other

Waahs
4/24 WA104 ? STU X
5/2 WA101 BR GERT X
5/18 WA104 ? FI X
5/18 WA104 ? FI X
5/22 WA103 GERT JUV d X
5/23 WA103 GERT FIL X
5/30 WA101 BR FF,GERT XX
5/30 WA101 BR IR X
6/12 WA104 ? 9 X
6/16 WA101 BR JUV d X
6/18 WA101 BR JUV C X
6/18 WA101 BR JUV d X
7/14 WA104 ? Juv X
-------------------------------------------------------------Total 8 0 0 3 3 0

Control Trials
4/16 EQ. MAIN X
4/22 EQ. MAIN PU X
6/7 EQ. MAIN X
7/12 NOISE WH IR X
7/12 NOISE WH BR X
7/12 NOISE RED JUV d X
7/12 NOISE RED FI X
7/12 NOISE RED FI X
7/12 NOISE RED JUV X
7/12 NOISE RED JUV 0 X
7/13 NOISE MAIN Am X
7/14 NOISE RED JUV d X
7/14 NOISE RED 9 x
7/15 NOISE MAIN MO,MD XX
7/17 NOISE MAIN ON,AL XX
7/17 NOISE MAIN PF X
---------------------------------------------------------------Total 0 0 0 6 10 2
----------------------------------------------------------------Up/Sky= animal looks up or to open sky; Down= animal looks down; Fol.= animal looks to foliage; Spkr.= animal looks to speaker; No Res.= No response; Other= Other responses.








84

once the subject animal for a playback was chosen, I followed that animal until the conditions for a playback were appropriate. I tried to set up each trial before a monkey was near, though this was at times difficult and the trials could not always be run. Speakers were always set in trees, never on the ground, because this was the typical position of a calling monkey and signals broadcast better off the ground. I ran a trial when the experimental subject

1) was relatively near the speaker,
2) in clear view and roughly facing the camera,
3) the speaker was out of sight of the monkey,
4) the animal producing the signal (recorded the
previous year) was out of sight.

If these conditions were met I then started filming the animal and then played the tape loop and continued filming for at least 30 seconds after the call. This playback protocol was developed in 1988, with Red group. Data taken during that period were not used in this analysis.

Playbacks often were to only one or a few animals,

which meant that other animals would not hear the playback. Thus, in some cases it was possible to do several playback experiments in a single day. I never repeated the same trial with the same animal on the same day.

For procedural reasons dealing with interpretation

difficulties, grrah variant 1222, the call given when snakes were near, was not tested. When considering a playback experiment it is instructive to think in geometrical terms. Playback experiments form a triangle, the three points being









85

the monkey, the alarm source (speakers), and the recordist. Ideally where the monkey looks--the response variable--is isolated from either of*the other two triangle points. For example, in a playback of a ground snake alarm call the monkey looks down, which is a different direction than either to the recordist or sound source. Unfortunately, in a playback of a near snake call, this condition could not be met. When the monkey hears this call, it should look near the sound source and determining whether it was looking at the sound source or near it for a snake would be difficult. For this reason, I did not test this call.

Playbacks of grrahs were done to 20 of the 21 available animals in Main group, the lone exception being Winston, a juvenile male. Playbacks of waahs were done to eight different animals in Red, White, and Main groups. Response Variable

This set of experiments tested where a monkey looked upon hearing an alarm call. The measured response was the direction the animal looked upon presentation of the stimulus. Response criteria were:

1) Up or to the sky- The animal looks toward the open
sky, either directly above it or towards a break
in the canopy.

2) Down- The animal looks down- either directly below
it or towards the ground?

3) Foliage- The animal looks to adjacent foliage,
either by looking around itself or looking up into
foliage?

4) Speaker- The monkey looks toward the speaker.









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5) No Response- The animal did not react to the
signal. Did it change its behavior,particularly
where it was looking, immediately upon
presentation of the signal?

6) Other- Were there other responses, such as looking
towards other monkeys, at the camera (me), or at
other animals?

Experimental Limitiations

There were several limitations to this experimental

design. First, regarding experimental replication there is a requirement that each event be independent. To produce independent replicates of these tests I could repeat them with another group of monkeys. Playing the calls used in these experiments back to another group of monkeys however might produce the confounding factor of the animals recognizing the vocalizations as not being from a member of their own group and responding to the stranger's voice rather than the message of the signal. Simple repetitions of the same stimulus only increase sample size. Replicates within the monkey group were accomplished by using a number of different stimuli given to a number of different animals. This design therefore avoided pseudoreplication. The playback design used here most closely follows design 1D among those described by Kroodsrna (Table 1, pg 601, 1989).

Another limitation was the inability to quantify a control response: responses to natural alarm calls. The monkeys did not always respond to the stimulus. Did they respond at a different rate to alarm calls given by actual









87

monkeys than to playbacks, or were they habituating to the presentations? To answer this requires a video record of the monkeys response to actual alarm calls. I have only one such record, where Jefe immediately looked in the direction of a monkey that called at an approaching tayra. I have no records of animals ignoring natural alarm calls. Lacking such records, I must judge whether these experiments represented realistic situations. Did the animals respond as if another animal had seen a snake and then give a call in alarm? Based on the behavior of responding animals compared to responses to actual predators I feel these responses were real, that the animals understood the calls and responded as if another monkey was giving alarm calls at a snake.

When a monkey did not respond, was this an indication that it had not heard the call? Was it not concerned about the snake? Lack of concern could be due to habituation or as a result of only a single call being heard rather than several. Lack of response is a difficult problem, particularly when I have no measure of its frequency in natural situations. I will assume that the specimen is the authority and have used as my data set only those responses where the experimental subject was apparently looking for a snake. I have not used responses where monkeys looked toward the speaker, other responses, or no responses.










88

Results


The results of the 111 playbacks are listed in table 8 above. There were adaptive responses to waahs and various grrahs 33 times, after discounting the "no responses", "looks to speaker" and "other responses." Grrahs

Responses to playbacks of the three grrah variants

indicating snake location are given in table 9. A Fisher's exact probabilities test indicates there is a highly significant difference in response probabilities (p< 0.00214), indicating that the monkeys respond differentially to each alarm call.


Table 9.
Responses to Playbacks of Grrahs

Responses Calls
2000 1111 3001
------------------- ------------------------Look Down 2 8 6
Look to Foliage 7 0 2
------------------- ------------------------Totals 9 8 8



Resiponse-variations

The question arises as to whether response differences are due to factors related to signal configurations such as signal duration, and intensity, or to other factors such as the age, sex, familial or social relationship of the caller and respondent. I found no correlation between signal









89

duration and percentage of correct responses (T=0.1521, Kendall rank correlation). Since the signal amplitudes were equilibrated there is no influence of amplitude differences on responses.

As for response rates to the various replicates,

within-call variant mean response rates for playbacks (for calls used more than once) are provided in table 10.



Table 10 '
Response Rates for Calls and Individuals Calls Individuals
------------ -----------Mean Range Mean Range

Grrah 1111
39% 33-50% 40% 33-55%

Grrah 3001
24% 0-50% 26% 17-33%

Grrah 2000
38% 0-67% 39% 33-50%

Waahs
66% 40-100% 66% 40-100%



Variation by caller. I will first examine response variations relative to the context of the caller. There were insufficient data to statistically compare differential responses by sex, familial relationship, or social dominance of the caller.

There were only five playbacks of calls from males,

which is too small for further analysis, whereas there were









90

73 playbacks of calls by females. Within the sample of female callers, there were no response differences by caller age-class, with 11/25 correct responses to juvenile females and 11/25 correct responses to adult females. (As a matter of definition, correct is used here to mean that the animal appropriately responded to the alarm call, which typically was to look for the predator.) There do not appear to be differential responses to females according to social dominance, though the sample is small. Of the 73 playbacks using female callers, 68 were done with the calls of four monkeys, three adult females (Whitey, Mo, and Pointy Face), and a single juvenile (Hanna). Whitey and Hanna are dominant within their age class, while Mo is intermediate between Whitey and Pointy Face, the lowest ranking adult female. Animals correctly responded to calls of the two dominant animals in 30% of the playbacks, 35% to the intermediate animal and 27% to the lowest ranking monkey.

There was only a single playback with related animals. Mo did not respond to a call by her youngest son, Mike. of the four females, Mo had the most offspring in the troop, Margo, Malli, Mike, and Modem, whereas Whitey had only a single son, Winston present. Pointy Face's only surviving offspring was a newborn in 1989, while Hanna had not reproduced.









91

Variation by respondent. Next we examine the

character of variations among respondents, by sex, ageclass, and familial and social relationships.

There was a significant difference (p=0.08, two-tailed Mann-Whitney test) in correct response rates between sexes of respondents by age class (adult, subadult, and juvenile). That is, males in three age classes responded correctly significantly more often than did females. There was no such relationship by sex of individuals outside of age-class.

The question next arises as to whether certain animals with many relatives in the group responded correctly more often than animals with few relatives. For example, did the members of Mols family respond differently than the unrelated group of males? Mols family contained five members, none of which were fathered by a male currently in the group. When Mols families rate of response is compared to that of the four presumably unrelated males (White Beard, Jefe, Griffin, and Stu) we find that there is no significant difference (p=0.53, two tailed Mann-Whitney) in response rate. This result is not surprising given that there was only the single test of an animal responding to a related animal's call. This explanation also assumes that the monkeys recognize each others' voices. A better test would compare data from responses to calls from related animals to those from unrelated animals. However if animals do not recognize each others' voices this result is somewhat









92

surprising, since one would expect that related animals would respond more often to each others' calls. This result can, by extension, be construed to further indicate that animals do perhaps recognize each others' calls.

Lastly, if we examine the relationship between social

dominance and rate of correct response within the five adult females we see that there is no correlation (r=0.4122, Kendalls rank correlation).

Waahs

Another series of playback experiments was undertaken using waahs. The same overall playback protocol was followed. Fourteen playbacks were done using three calls given by animals in two different groups (Table 8). In eight of the fourteen playbacks the respondent reacted by looking up to the open sky, three times they looked towards the speakers, and three times there was no response. I conclude from these experiments that when the monkeys responded to these calls as alarms they reacted in every case, 8/8 times, by looking up for an avian predator. ResRonse variations

Response variations by sex of the respondent is

interesting. In the eight calls played back to males, they responded by looking to the sky on three occasions, whereas females responded each of the five times. I was unable to sex one respondent. When the caller was a male, monkeys correctly responded 4/7 times while for females it was 2/2




Full Text
64
early rainy season, August 3, 1988, and there is standing
water in many places. The monkeys rarely come to the ground
at this time of year so it is highly unlikely that she was
looking for food or other monkeys when she scanned the
ground. This is noteworthy because the snake had probably
not been visible for four minutes.
I interpret Mo's behavior to indicate that she
understood the various arrahs to refer to a snake.
Alternatively she could have seen the snake earlier or seen
the other monkeys looking down.
Call Types
The fact that different animals made similar calls
suggests that the variability represents meaningful
differences to the animals. Testing this hypothesis requires
a classification schema. I therefore devised a key based on
seven grrah features:
1) Formant frequency drops/ increases
2) Frequency drop is continuous/ discontinuous
3) Frequency drop is extensive/ not extensive
4) Number of formants
5) Which formant was stressed
6) Stressing at end was above/ below the tail
7) Extensive frequency drop in formants other than F1
8) Call duration
These seven features use all three acoustic domains
frequency, amplitude, and duration. I concentrated on the
frequency domain because the calls appeared to vary the most
in this domain, and, a priori. appeared to be the most
robust domain to differentiate the calls. Call duration was


94
The responses of Q. olivaceus to alarm calls for
raptors and snakes are, in some aspects, very similar to
those of Cercopithecus aethiops: both monkey species, when
in trees, look up upon hearing raptor alarm calls and look
down when they hear snake alarm calls (Seyfarth, et al,
1980). In other ways, however, the anti-predator behavior
of the two primates is quite different. Descriptions of
vervet responses to predators recount how the monkeys
actively react, both vocally and otherwise, to protect
themselves. For example, Seyfarth et al. (1980) describe
vervets responding to leopard calls by running into trees or
to eagle calls by looking up then running into cover. Cebus
olivaceus rarely were so active in their anti-predator
behavior. This is perhaps due to their not being on the
ground as much as vervets and the lack of dangerous raptors.
I never saw them on the ground when a cat was present, so I
do not know their response in that context. When in a tree
C. olivaceus called and mobbed the cat but did little else.
I suspect that they did not run into cover when raptor calls
were given because 1) there was little real danger from a
large raptor and 2) most waahs were given to small,
relatively less dangerous hawks. In fact it still remains a
mystery why C. olivaceus give alarm calls at road-side
hawks, which almost certainly do not pose a mortal threat to
any but the smallest of monkeys. The anti-predator
behavior of C. olivaceus at Hato Masaguaral may not be


167
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132
Call 3001- Like 1111, maximum frequency and its
increase were intermediate to low, but like
2000, the total frequency drop was relatively
large with a late location of frequency
maxima.
Call 2000- maximum frequency and increase tended to be
high, the total frequency drop was relatively
large, and the location of maximum frequency
tended to be late.
The exemplars for the other call variants had
properties like the following locational calls, as
determined by the first two principal components:
Call 2000- Variant 1321
Call 1111- Variant 1210, variant 1222
Call 3001- Variant 1311, variant 13122
Intermediate between call 1111 and 3001- Variant 13121,
and variant 1221.
Call variant 13121 had similar component 1 values but widely
disparate component 2 values and therefore was unlike any
other call.
Discriminative functions analysis, A discriminative
functions analysis was next used to describe the acoustic
cues that a capuchin may use in differentiating the various
qrrah variants. This analysis was used to determine how
well these variables discriminate the various call variants.
Figure 13 shows how 4 variables discriminate 59 calls. This


134
allocation was accomplished using a subset of the variables
used above in the principal components analysis: call
duration (DUR), maximum frequency (MAX), the bandwidth of
the dominant formant (BWFORM), and the amount of increase in
this formant (INC). Overall 76% (45/59) of the calls were
correctly classified. Among the locational calls, the
extremely short duration call 3001 was most accurately
discriminated, followed by 1111, while only half of the call
2000 were correctly classified. All the other calls were,
with one exception, correctly classified.
Classification errors were instructive. Call 1111 is
typified by a large frequency drop from a single dominant
formant after an initial period of constant frequency. This
was most similar to call 1130, where the frequency drop was
also large but discontinuous (a descriptor not used here)
and call 1210, where there was a large frequency drop with
no initial period of level frequency. The mistaken
identities in call 2000 were to calls where there was little
frequency drop, calls 13xx. These two calls were different
in where and how the frequencies increased.
Few calls were misidentified as a locational call,
indicating that, at least with these variables, the calls
were quite distinctive. This can be seen in figure 14, a
map of the locational calls as arrayed by the first two
discriminant factors. The location of the group centroids
( + ) show that these calls were all distinctively different.


APPENDIX A
NARROWBAND SPECTROGRAMS OF GRRAHS TO A RELEASED 0.68 m BOA
VOCALIZATION NUMBER AND CALLER NAME IS GIVEN FOR EACH CALL
BANDWIDTH= 80-8000 HZ, RESOLUTION= 45 HZ.


CHAPTER 6
SUMMARY AND CONCLUSION
The major findings in this investigation are two: Cebus
olivaceus use alarm calls to designate predators and they
use at least some alarm calls to attribute location to these
predators. Additionally, C. olivaceus may use a limited set
of acoustic features to differentiate among the variants of
their grrahs to snakes.
These results raise the question not only of what is
being communicated by the monkeys but how. Cebus olivaceus
appear to designate objects and attribute qualities to those
objects. They apparently do this by using acoustic
variations of other alarm calls. They modify the meaning of
the call 'snake', for example, into ground snake by
modifying a basic 'snake' call, as opposed to adding
separate signals syntactically, through the use of
additional distinct calls.
Earlier I distinguished the use of general terms as
being extensions of a class of objects vs the use of
intensional, or conceptual uses of a term. Evidence to
prove whether an animal conceptually grasped the defining
characteristics of that class included the use of
attributions. This study of C. olivaceus alarm call
157


37
The studies of syntax in both Callicebus and C.
olivaceus by Robinson are important because they best
illustrate that at least some primates use rudimentary
ordering schemes in vocal production and these syntactic
patterning rules may be similar to those used by humans.
Several problems emerge from the above description of
linguistics and primate communication. I have posited that
primates are capable of naming objects but to date there is
no evidence that they speak about them. If this is the case,
syntax poses a particular problem because syntax provides
another layer of meaning upon the individual lexical
elements. Is syntax a linguistic operator requiring one
form for extensional terms and another for intensional? The
linguistic elements syntactically combined may imply
differences in conceptual understanding. Evidence for this
would be how elements are syntactically combined. Syntactic
combinations of extensional terms would involve combinations
of simple names and an operator, whereas a combination of
intensional terms would involve, for example, analogy or
other mechanisms requiring a conceptual grasp of the
combined words. I believe there is a conceptual difference
between the syntax of 'man bites dog' vs 'dog bites man' and
that occurring when water is lexically combined with bird to
form waterbird (where waterbird is used in a context
separate from water and other birds to refer to a duck).
The former construction requires only an ability to name two


7
procedure, with the signaller processing sensory and
cognitive inputs prior to the generation of a signal. I
note this here because in subsequent discussions I will use
the terms external referent and internal referent. It is
important to remember that all signals possess internal
reference to some degree, where neural inputs are
transformed into vocal outputs. Objects of external
referents are not simply named without several layers of
cognitive processing before the signal is generated.
Nevertheless, the word car refers to an external physical
object whereas anger refers to an internal state.
The concept of meaning is central to linguistics and
communication. The theme of word and meaning is the subject
of a pivotal work by Quine (1960). He notes that "meaning,
supposedly is what a sentence shares with its translation;
and translation at the present stage turns solely on
correlation with non-verbal stimulation" (Quine, 1960, pg
32). It is the problem of translation of a language where
no translators or dictionaries provide denotative meaning
that prompts Quine to coin the term radical translator.
This is the problem of a linguist who meets a previously
unknown people or a wildlife biologist who is trying to
determine the meaning of an animal's vocalizations. Initial
steps at translation are more like correlation, where the
vocalization is first translated when paired with
conspicuous events. First attempts at detecting meaning are


54
is a arrah at me given by an adult female, Irma, in White
group. Note that it is noisy at the end and downswept.
Graahs to other animals. Figure 2d illustrates two
arrahs to an ocelot (Fells pardalis) from an unidentified
animal in White group as it watched the cat walk under it.
Both calls have two energy bands and drop discontinuously.
The terminal aspiration is slight. The pure tone at
approximately 1.5 kHz is an artifact.
On several occasions C. olivaceus would grrah at
seemingly anomalous objects. Figure 2e provides two grrahs
to a donkey given by unknown animals. These calls are quite
distinctive with a long fundamental having multiple energy
bands only in the middle of the call after which the
frequency drops.
Waahs
Waahs are, in general, longer duration, slightly lower
in frequency and less forceful than grrahs. Like grrahs the
pitch falls, however since waahs typically have longer
duration the slope is less. They may also be quite breathy
or noisy. While the sample size for waahs is much smaller
than for grrahs it is quite apparent there is also
significant variability among waahs. The semantic utility
of this variability is unknown.
Waahs to Road-side hawks. Waahs are most often
given to the road-side hawk (Buteo magnirostris). Figure 3a
illustrates two waahs by adult females Gertrude and Irma


100
and Seyfarth (1981) could not give similar data for response
differences to snakes because they observed vervets
interacting with pythons on only two occasions. This can be
compared to data for C. olivaceus where 92% of calls to
released snakes were from three age/sex classes: adult
females, subadult males, and juvenile females. Adult and
juvenile males rarely gave alarm calls (6%) Therefore in
both primate species there is a skew in the frequency with
which certain age/sex classes emit alarm calls.
A similar comparison cannot be made between vervets and
Cebus with regard to age/ sex class of the alarmist in
playback experiments. This is due to the lack of alarm
calls from male C. olivaceus. I had only two. Among the
vervets there was no significant difference for eagle or
snake alarm calls, but there was for leopard calls. When
the age/sex class of the alarmist was examined regardless of
call type, no significant difference emerged (Seyfarth et
al., 1980).
Among vervets high ranking animals give more calls than
low ranking animals. This was not the case in C. olivaceus,
where there was no relationship between dominance among
adult females (the only group for which sufficient data were
available) and number of grrahs given to released snakes.
Other animals
Several species of ground squirrels give alarm calls to
a variety of predators. Initial research on the these calls


43
(1981, 1982, 1984, 1986) from 1976 to the present time. All
animals in the four groups were individually identifiable by
name, with identification made possible by age/ sex, size,
and individual markings. All animals, particularly Main
group, were habituated to my presence. This made it possible
to study the animals at extremely close range, usually 2-
10m. Familial relationships within these four groups have
been known for at least ten years.
Table
3 .
Composition of
Main Group
Males
Females
Name
Age*
Name
Age*
White Beard
>10
Mo
18
Jefe
8
Amelia
17
Griffin
8
Beaut
17
Stu
7
Whitey
12
Mani
5
Pointy Face
12
Babas
4
Alexandra
8
Winston
3
Onica
7
Mike
1
Margo
7
Hanna
5
Mali
5
Puffy
4
Amanda
3
Modem
3
* age as of
1989
Predators of Cebus Olivaceus at Hato Masaquaral
As Cheney and Wrangham (1987) note, predation on
primates is rarely observed. There have been no confirmed
instances of predation on C. olivaceus at the ranch. The
Harpy eagle (Harpia harpyia) and the boa constrictor


72
males. White Beard, the dominant adult male, gave only two
alarm calls. This low production of alarms was typical of
him and apparently of other adult males, both for alarm
calls and other vocalizations.
Grrah Syntax
A matrix of grrah variants indicating call pairs is
provided in Table 7. The data set is from all identified
callers, with call variants modified from that provided in
figure 4: variant 3000 includes 3001 and both 1112 and 2110
are excluded due to small sample size. One grrah followed
another 158 times, approximately 78% of the time, so the
most likely association is between grrahs. The most common
association was call duplication, where the same call
variant was simply repeated. This happened significantly
more often than by chance (X2 = 52.37 p= 2.3 E'7, d.f.= 11).
This same statistic was not possible with other pairings
because of the numerous null cells in the matrix, but it is
clear that there are no strong associations between any
calls. I conclude that except for duplicating calls C.
olivaceus do not combine grrah types in a syntactic manner.
If C. olivaceus do not syntactically combine different
grrahs. do they combine calls to form a new call with
intermediate form? This intermediacy in form could indicate
that 1) the meanings of the two calls were combined, using
phonological syntax or 2) that the signals are graded. If
this is phonological syntax, I am suggesting that the


88
Results
The results of the 111 playbacks are listed in table 8
above. There were adaptive responses to waahs and various
arrahs 33 times, after discounting the "no responses",
"looks to speaker" and "other responses."
Grrahs
Responses to playbacks of the three qrrah variants
indicating snake location are given in table 9. A Fisher's
exact probabilities test indicates there is a highly
significant difference in response probabilities
(p< 0.00214), indicating that the monkeys respond
differentially to each alarm call.
Table 9.
Responses to
Playbacks
of Grrahs
Responses
2000
Calls
1111
3001
Look Down
2
8
6
Look to Foliage
7
0
2
Totals
9
8
8
Response variations
The question arises as to whether response differences
are due to factors related to signal configurations such as
signal duration, and intensity, or to other factors such as
the age, sex, familial or social relationship of the caller
and respondent. I found no correlation between signal


127
There was a negative correlation between end frequency
(END) and location of no dominant formant (NODOM). This
correlation indicates that where calls ended at higher
frequencies the location of no dominating formant, that is,
where all formants were of the same amplitude, was likely to
be earlier. Likewise if the end frequency was low the
location of no dominance was likely to be closer to the end
of the call; these lower calls would sound more noisy.
NODOM had different interpretations based on its location in
a call. In some grrahs, particularly those with more
aperiodicity at the end, NODOM determined where the
aperiodicity occurred. In grrahs with many formants, if all
formants were equally stressed NODOM was near the beginning.
The amount of increase in the first formant (INC) was
positively correlated with the location of maximum frequency
of that formant (MAXLOC): calls with the greatest frequency
increase were more likely to have the increase near the end
than if the increase was slight. This was best seen in the
alarm call to avian predators, variant 2000. Unfortunately,
there was neither a correlation, nor partial correlation,
between either of these two variables and the position of
maximum amplitude (DBLOC). If so that would indicate that
when a large frequency increase was located near the call's
end the increase would have been emphasized. As it turns
out, the position of maximum amplitude was not correlated


45
some interactions the monkeys chase the tayra, while during
others the monkeys appear quite alarmed, and during still
others they apparently ignore each other.
The boa constictor (Constrictor constrictor) is the
only large snake living in the forest at Hato Masaguaral
that is likely to take C. olivaceus. Other potentially
dangerous snakes are anacondas (Eunectes murinus) and a
rattlesnake (Crotalus durissus) which is rarely found inside
the forest.
Estimated predation levels on C. olivaceus at
Masaguaral are moderate to low. In seven years of
observations on approximately 175 animals Robinson estimated
that 15 animals were taken by unknown predators, an
estimated annual predation rate of 3% (Robinson in: Cheney
and Wrangham, 1987, pg. 232). Predation rates were highest
on smaller animals; of those presumed taken 70% were infants
and 20% juveniles with the remainder split between adult
males and females. Reuter (1986) discusses the influence of
predation on foraging behavior of C. olivaceus.
I should emphasize that intraspecific aggression
appears to be a major source of mortality among the study
animals, and is probably greater than predation. For
example, in 1988 a new male, WB, entered Main group and
within 6 weeks the four newborn infants had all disappeared,
with one confirmed case of infanticide by that new male
(Valderrama et al, 1990).


158
behavior shows that this primate uses a concept of a class
of predatorssnaketo order some as being located in a
tree and others as being on the ground. This
differentiation may involve the same individual snake at
different times. This is the first evidence that a non
human primate has a conceptual grasp of categorical
distinctions.
These distinctions naturally raise the question of
whether C. olivaceus uses language. As stated above,
language is notoriously difficult to define. We are no
longer safe in the axiom that we may not know how to define
it but we know what it is when we hear it. This is a
homocentric subterfuge, because we may forever remain unable
to distinguish the different types of calls used in a
particular species communication system and therefore will
be unable to determine the meaning of the various signals.
There is no evidence to date, and none is provided here, to
indicate that any non-human primate is capable of the
linguistic facility shown daily by humans. This is not to
say that they are incapable of language. On the contrary, I
think that we may be unable to ever fully decipher the
meaning of the majority of a primates calls, their many
calls given in social situations for example, so we will be
unable to determine if they have the openness, sensu Hockett
and Chomsky, that is seen in our languages. It is my
position that primates, at least C. olivaceus. use a


22
response and external reference make the study of alarm
calls ideal for semantic research.
Phonology
Phonology is the study of speech soundsthe phonetics
and phonemics of a language. It includes studies of how
sounds are made as well as the psychoacoustics of their
reception. Where phonetics is used to describe and classify
speech sounds, in phonology these descriptions and
classifications are used to describe communication systems
and explain sound processes (Sloat, Taylor and Hoard, 1978).
The basic unit of phonetics is the phone, which simply is
any speech sound. The basic unit of phonemics is the
phoneme which is the smallest speech unit that distinguishes
one linguistic utterance from another.
Acoustic variability may describe phonetic or
phonological differences. The perceived association between
the signal and phones is the phonetic quality of the signal.
This is in contrast to phonemes which are the abstract class
of minimal phonological units consisting of phones (the
phonetic unit) that are functionally identical.
If we extend this definition of phonology to the study
of all vocal sounds, including those of non-human animals,
then we can view each species as potentially having its own
phonetic system. Taxonomists of animal vocalizations,
lacking (at least initially) a semantic framework to
distinguish sounds, face the problem of whether to lump or


106
window and the a's are regression coefficients to be
calculated. The error term an is the difference between the
actual sample on and the estimated sample n
a = o eq. 2
The regression coefficients, a's, are determined such that
they minimize the mean square of the error terms, an's. An
important assumption of this technique is, functionally
speaking, that the signal is periodic over a reasonably
short time window. Calculation of the regression
coefficient requires that the statistical distribution
characteristics of the signal are constant. This assumption
is met when the signal is periodic and fails when the signal
is aperiodic. This time window typically is 10 msec. The
duration of alarm calls range from 30-140 msec and therefore
values from many windows are used.
Linear prediction coding is employed here to calculate
the formant frequencies, amplitudes and bandwidths used in
principal components and multi-variate analyses.
Accurate representations of most human speech are
produced by LPC, in particular vowels and voiced consonants.
It produces less accurate representations of less periodic
sounds, those with higher frequency components such as
unvoiced consonants, fricatives, and some nasalized sounds.
Does LPC accurately represent non-human vocalizations,
particularly Cebus calls? This question is necessary
because, first LPC was designed to describe the lower


103
conversion board. Anti-alias filtering was done with an
Allison Labs model 2AB variable filter set at 9.6 kHz.
Digital signal processing and speech acoustics
Digital recording and analysis is fundamentally
different from analog recording and analysis. In digital
recording the waveform is recorded as a sequence of digital
samples taken at a particular sampling rate. Provided that
this sampling rate is fast enough any signal of limited
bandwidth can be accurately represented. A minimum of two
samples per wave will describe a waveform, therefore the
spectral bandwidth is \ the sampling frequency. The
sampling rate used here was 20,000 samples/ second,
resulting in a 10 kHz spectral bandwidth. Digital
processing systems therefore transform a continuous waveform
into discrete events.
Digital representations of a communication signal can
be classified into either waveform or parametric
representations. Both types are used here. The waveform
representation preserves the analog waveform through
sampling and quantization processes, from which frequency,
time, and amplitude information can be obtained from, for
example, digital oscillograms and Fourier transforms. The
Fourier transform is an algorithm that translates the time
domain waveforms into the frequency domain, producing
frequency spectra. Parametric representations, developed in
large part to model human speech, represent the acoustic


CHAPTER 2
VOCAL RESPONSES TO PREDATORS BY CEBUS OLIVACEUS
Methods and Materials
Research Site
This project was conducted at Hato Masaguaral in three
periods: preliminary observations from May-July, 1986; snake
release experiments from March-August, 1988; and playback
experiments from April-August, 1989. Hato Masaguaral is an
active cattle ranch as well as wildlife refuge owned by Sr.
Tomas Blohm and is located 146 miles south southwest of
Caracas in central Venezuela (834'N, 6735'W). The study
site was a 4 km2 gallery forest (figure 1). It was bordered
to the east by a seasonal river, the Cao Caracol and,
respectively, to the north and south by two recently
deforested ranches, Finca Torres and Hato Flores Morades.
To the west the gallery forest gradually became a more open
forest. The study site seasonally flooded from
approximately May to October after which it dried and the
deciduous canopy became increasingly open. Rainfall in the
region averaged 1450 mm/ year. The mean temperature was
approximately 28C. See Troth (1979) for a more detailed
description of the ranch.
40


to not only designate objects but also to attribute
qualities such as location to those objects.
viii


149
Overall, vervet and capuchin alarm calls were more
acoustically different than alike. These differences should
not mask the similarity in function: both serve to designate
objects. In fact, the apparent differences in acoustical
structure combined with the similarities in function point
to a convergence in communicatory purpose that may parallel
that between humans and these two primates.
Distinctive Features
Table 18 presents the vocal distinctive features for
cotton-top tamarin (Cleveland and Snowdon, 1982) chirps,
pygmy marmoset trills (Pola and Snowdon, 1975; Snowdon and
Pola, 1978), vervet grunts (Cheney and Seyfarth, 1982) and
alarm calls (Owren and Bernacki, 1988; Owren 1985), C.
olivaceus tonal calls (Robinson, 1984) and alarm calls.
Frequency cues
Bandwidth of the frequency change differentiated calls
in all but the vervets. Bandwidth most accurately defined
how much the calls changed in frequency; typically how much
frequency dropped. Bandwidth was an important determiner
when the call was tonal. Bandwidth was not, however, a
sensitive factor in differentiating vervet calls because
both alarm calls and grunts were very short duration and
therefore were equally wide band. (Typically very short
duration calls are wide band and noisy. This is due to the
difficulty in controlling vocal folds vibrations in brief
calls so that they vibrate at a single frequency.) Peak


59
Where none of the procedures worked, I discarded the
call. Most of the attibutions were done with the first two
methods. Using these procedures I identified the caller for
190 of 279 arrahs recorded during releases.
The following data were input into the Q & A data base
management program to correlate the various calls with the
contexts of the snake releases,
1) Caller distance to snake and nearest neighbor
near, medium, far
2) Identity of nearest neighbors
3) Number of snakes
4) Snake location (tree or ground) and size.
Additional information included vocalization number, call
class (variant type, see below) and caller.
Release Methodology and Release Schedule
Recordings of orrahs to snakes were made, with a single
exception, by releasing a snake or snakes in front of Main
group. In the exceptional case, the monkeys discovered a
boa in a tree, I video recorded the interaction, then
captured that snake for a later controlled release. In a
controlled release, I approached the monkeys, stayed with
them for an appropriate time to make sure my presence was
not alarming, then removed the snake or snakes from a cloth
bag in front of the monkeys. In some situations where I
wanted to obtain recordings from a particular animal, I made
sure it was alone or had few companions before releasing a


112

Figure 8. Comparison of Fourier and LPC derived spectra of
middle segment of JM102, a grrah variant 1111. The LPC
spectra is smooth and overlays the FFT spectra.


Table 5. Contexts for grrah variants with total calls given, number of situations possible, and
number of callers.
Call
Ground
Tree
Near
One
Two
Small
Medium
Large
Yellow
Type
Tot
. #
S.
A.
#
S.
A.
#
S.
A.
#
S.
A.
#
S.
A.
#
S.
A.
#
S.
A.
#
S.
A.
#
S.
A.
1111
27
26
6
5+
6
1
1 +
7
2
3
20
3
4 +
5
1
2
2
1
1
1112
7
5
1
1+
2
2
2
5
2
1+
4
2
2 +
1
1
1
4
2
1+
1120
32
28
4
6+
3
2
2
10
3
3 +
11
3
5
21
2
3 +
6
1
3
2
1
1
3
1
1
1210
17
14
6
10+
3
1
1
5
2
2
12
4
6
5
3
3 +
4
1
3
5
1
2
2
1
2
1
1
1
1221
22
13
5
5+
8
1
2
7
1
2
11
2
4 +
11
3
2 +
10
1
3 +
1
1
1
1222
8
3
2
2
5
3
3
7
3
4
4
2
3
4
1
2
4
2
3
1311
36
27
7
6+
9
3
4
18
5
4 +
15
4
5+
19
4
4 +
3
1
2
9
2
3 +
3
1
1
1312
17
9
2
3 +
8
3
3 +
4
1
1+
14
3
4 +
3
1
1
2
1
1
12
2
3 +
1321
27
13
6
7 + 13
2
5
11
2
3 +
21
4
9
6
4
3 +
3
1
2
15
2
6
3
1
2
13121
34
21
6
6
10
3
5
12
3
4 +
21
3
7 +
12
4
3 +
4
1
2
17
2
6+
13122
6
1
1
1
5
1
1
4
2
2
6
2
2
6
2
2
2000
13
2
2
2
11
3
4 +
5
4
3 +
9
3
3
4
1
3 +
1
1
1
8
2
3
2110
3
3
1
1
3000
18
10
4
3 +
6
3
2 +
11
4
3 +
10
3
4 +
7
3
1+
9
2
3+
1
1
1
3001
10
10
4
3 +
6
3
2
4
2
1+
2
1
1
3
1
1
1
1
1
Total
279
184
83
107
151
121
30
101
15
5
o\
oo


175
HQ178
Ml
Variant 1221
HS8
Pu
I Kb *
A. I ^ A* ^
-
HP91 MM104
Ml Ha
Variant 1311
HQ125 HA115
Ml Ha
Variant 1312


113
those calls that were periodic at the end, the fundamental
was also measured.
I used two other techniques to verify the CLA results:
a discrete Fourier transform (DFT) and analysis using the
CEPSTRUM algorithm. The discrete Fourier transform allows
the user to specify the bandwidth and therefore the
resolution of the resulting spectra. I use a 200-1400 Hz
bandwidth. (The more normally used fast Fourier transform
(FFT) gives a spectrum that is equal to the entire
bandwidth, in this case, 10kHz, with the resulting decrease
in frequency resolution). The CEPSTRUM technique can also be
used to estimate pitch as well as to determine if the signal
is voiced or unvoiced (Rabiner and Schafer, 1978). This
technique takes the inverse Fourier transform of the log of
the power spectrum. The pitch can then be estimated by
taking the period of CEPSTRUM waveform. I will use a 4 msec
window in the analysis.
Other computing software
Two other software programs, besides ILS, were used to
develop and analyze the results given below: SuperCalc IV, a
spreadsheet program, and Statgraphics, a statistical
package. The acoustical data on three formants (frequency,
bandwidth, and dB level) was transferred from ILS to
SuperCalc where a spreadsheet calculated values for the
twenty variables. Values for these variables were then
transferred to Statgraphics where they were analyzed.


57
known animals to determine where they looked upon hearing
the signal. These experiments replicate and extend
Seyfarth, Cheney, and Marler's (1980) experiments on
vervets. I hypothesized that if capuchins named objects,
e.g. snakes, they might also attribute qualities to those
objects. For example, would they vocally distinguish one
snake from two, small from large, or provide information on
a snake's location? To test these hypotheses, I released
different numbers and sizes of snakes then recorded the
monkeys responses. Following that I played to them selected
calls to determine if a listening animal responded
appropriately.


Figure 3. Narrowband spectrograms of waahs to a) road-side hawk, b) vultures, and c)
caracara. (Bandwidth 80-8000 Hz, resolution= 45 Hz). Marks at 1 kHz intervals.


93
times. Lastly I should note that in one playback using
WA104, a call from Red group, the respondent was Stu, a male
in Main group. He responded to the call by looking up to
the sky.
Discussion
I conclude from these experiments that C. olivaceus use
their alarm calls to, first, identify objectssnakes and
avian predators, and second, to attribute qualities to those
objects by identifying the snake's location.
This is not an exhaustive catalog of what C. olivaceus
do with their alarm calls; there may easily be other calls
that identify other objects, give other attributions, or use
the same attributions for other objects.
The results of the alarm call playback experiments
confirm, I believe, the conclusions of the snake release
experiments; C. olivaceus both give specific alarm calls to
different predators and to specify predator location.
Comparisons with Other Animals
Vervets
The following is a detailed comparison between C.
olivaceus and Cercopithecus aethiops alarm call behavior.
These two species are ecologically similar and the
semanticity of their alarm calls are the most fully
described.


78
The next step in this translation process is to find
out how a monkey will respond to these locational call
variants. For example, if qrrah variant 3001 is always
given to a snake on the ground, and in fact means 'ground
snake' then we would expect a monkey to look down to the
ground when it hears this call. The experiments described
below will serve to confirm the above results.


117
Table 11.
Fundamental Frequencies (Hz)
for Waahs and Grrahs
Mean
S.D.
Range
Grrahs
Beginning
1173
182
833-1428
Middle
969
381
588-2000
End
881
290
370-1538
Waahs
Beginning
825
190
606-952
Middle
749
381
526-870
These same values were also calculated for
vocalizations by identified individuals (table 12).
Table 12.
Fundamental
Frequencies
for
Individuals
(Hz)
Individual
HA
HA
MO
Call Type
1111
3001
2000
Beginning
1334
1271
973
Middle
880
965
1261
End
558
909
1138
These fundamental frequency values, on the order of
1200Hz, are much higher than for vervets, 230-250 Hz
(Seyfarth and Cheney, 1984). These values were verified
using DFT and CEPSTRUM techniques. Figure 9 is 20 msec of a
qrrah variant 13121. It is clearly periodic. Figure 10 is
the DFT of this portion. The fundamental frequency by this
measure is 1187 Hz. Figure 11 is the CEPSTRUM of the first


25
parameters (presence/absence of a frequency upsweep,
difference between peak and end frequency, peak frequency,
and duration of frequency downsweep) differentiated the
eight calls.
Table 2.
Primates Exhibiting
Phonetic Differences
in Their Communication System
Species
Reference
Japanese macaque
Green, 1975
Talpoin monkey
Gautier, 1974
Pygmy marmoset
Pola and Snowdon, 1975
Snowdon and Pola, 1978
Cotton-top tamarin
Cleveland and Snowdon,1982
Vervet
Cheney and Seyfarth, 1982
Seyfarth and Cheney, 1984
Owren and Bernacki, 1988
Owren, 1985
Gelada baboon
Richman, 1976
Squirrel monkey
Newman et al, 1978
Black spider monkey
Eisenberg, 1976
Wedge-capped capuchin
Robinson, 1984
In another distinctive features investigation, Owren
and Bernacki (1988) were able to define a single acoustic
variable, spectral tilt, that correctly classified two
Cercopithecus aethiops alarm calls, eagle vs snake, 97% of
the time. If the tilt was falling, that is, if the
amplitude decreased each successively higher spectral peak,
the call was an eagle call, otherwise it was a snake call.
Perceptually this means that eagle calls were lower pitched
than snake calls because the higher spectral peaks of the
snake call were relatively louder. From another


I
of9the signal011109^111 f 2 mseC f ^rrah variant 13121 illustrating the periodicity
118


174


3
to represent the object, where the signal itself refers to
the object (Cherry, 1978). The alarm call itself refers to
the predator because the call designates the object through
symbolic representation. It is conventionally held that
animals cannot symbolically communicate about referents,
internal or external, and that if they want to communicate
about an object they must do it either by indexing the
object, or by using some iconic signal. Altmann (1967)
notes the power of combined affective and indexical
communication and how prevalent it is among primates. For
example, Menzel and Halperin (1975) show that a chimpanzee's
walking gate communicates information to other animals about
the quality of food sites.
Animal communication has traditionally been seen as
fixed, stereotyped, and simple whereas language was
perceived as complex, variant, and open. This perspective
began to change with the pioneering investigations of bee
communication by Karl von Frisch (1967) where it was shown
that a bee could symbolically communicate the distance and
direction from the hive to a food source. Subsequent
investigations in a wide variety of taxa have shown that the
traditional view was inadequate. There is mounting evidence
that a new perspective describes the behavior of certain
species better than the earlier theory. According to this
new view, primates use signals to communicate about objects
in a manner very similar to how we use words to identify


BIOGRAPHICAL SKETCH
I was born November 29, 1950 and raised in Muskegon,
Michigan, where I graduated from high school in 1969. I
graduated, with a BA in philosophy, from Michigan State
University in 1973. My Master of Science is from San Diego
State University, granted in 1981. My thesis subject was
hearing in whales and dolphins. My research interests
include communication in primates, cetaceans, and other
mammals, as well as conservation in Africa.
180


63
minutes she looks for the snake, at which time I pull
it out of the vine and put it on the ground in front of
her. She watches the boa for a minute without calling.
It is now seventeen minutes into the interaction.
Finally she gives a quiet qrrah (not analyzed) and
another animal gives a huh. Her next call is MM101, a
call quite similar to HA122 by Hanna. A half a minute
later she becomes agitated, jumps, and then gives qrrah
MM102 while looking down at the boa. Another animal
huhs and she responds with MM103 as the snake swims
below her. Fifteen seconds later she gives qrrah
MM106, an alarm call used for snakes in a tree. I
cannot confirm the snakes location at that time, though
Ik minutes later the boa was 1 meter up a vine. The
last calls recorded from Mo are the two similar calls,
MM107 and MM108. At this last call, 30 seconds after
the tree snake call, she does appear to be looking
obliquely at the snake instead of down, i.e. the snake
may be in a vine. She then begins to huh at Hanna who
then rapidly moves down a vine emitting hehs. Hanna
appears to direct the calls at the snake rather than
Mo. White Beard, the adult male of the group then
passes below her unperturbed by either the snake or
Hanna's aggressive hehs. (On other occasions other
monkeys emitted hehs at boas).
Two minutes after the last qrrah. Hanna is lying
on a vine and Mo is self grooming, both monkeys can
plainly see the small boa as it tries to climb a palm.
The interaction ends with Mo moving away, five minutes
after the last qrrah.
I conclude from this interaction that 1) C. olivaceus
gave a variety of acoustically distinct alarm calls to
snakes, 2) the alarm call variability was apparently not
random, instead different animals gave apparently similar
calls, 3) the alarm calls Mo heard informed her of the
snake's presence, 4) Mo may have called even before she
herself saw the snake, and 5) the presence of a snake does
not automatically prompt a monkey to call.
By all appearances Mo was looking for the snake. I
should note that this snake release occurred during the


83
Table 8-continued.
c) Responses to waahs and control trials
Responses
No
Date
Call
Sp.
Rec.
Up/Sky
Down
Fol. Spkr.
Res.
Other
Waahs
4/24
WA104
7
STU
X
5/2
WA101
BR
GERT
X
5/18
WA104
7
FI
X
5/18
WA104
7
FI
X
5/22
WA103
GERT
juv a
X
5/23
WA103
GERT
FIL
X
5/30
WA101
BR
FF,GERT
XX
5/30
WA101
BR
IR
X
6/12
WA104
7
9
X
6/16
WA101
BR
juv a
X
6/18
WA101
BR
JUV cf
X
6/18
WA101
BR
JUV cf
X
7/14
WA104
7
Juv
X
Total
8
0
0 3
3
0
Control Trials
4/16
EQ.
MAIN
X
4/22
EQ.
MAIN
PU
X
6/7
EQ.
MAIN
X
7/12
NOISE
WH
IR
X
7/12
NOISE
WH
BR
X
7/12
NOISE
RED
JUV cf
X
7/12
NOISE
RED
FI
X
7/12
NOISE
RED
FI
X
7/12
NOISE
RED
JUV
X
7/12
NOISE
RED
JUV cf
X
7/13
NOISE
MAIN
Am
X
7/14
NOISE
RED
JUV cf
X
7/14
NOISE
RED
9
X
7/15
NOISE
MAIN
MO, MD
XX
7/17
NOISE
MAIN
ON, AL
XX
7/17
NOISE
MAIN
PF
X
Total
0
0
0 6
10
2
Up/Sky= animal looks up or to open sky; Down= animal looks down;
Fol.= animal looks to foliage; Spkr.= animal looks to speaker;
No Res.= No response; Other= Other responses.


123
Table 13. Descriptive statistics of acoustic
parameters of Cebus alarm calls.
95% SIGN.
VAR TYPE
X
a
CONF
. INT.
F-ratio
LEVEL
DUR 1110
. 095
. 005
.085
. 104
27.68
. 0000
2000
. 100
.063
. 089
. Ill
3001
.044
. 004
.031
.057
BEG 1110
1372
46
1209
1535
4.959
. 0122
2000
1607
147
1403
1813
3001
1136
108
910
1360
END 1110
539
61
313
765
13.609
. 0000
2000
1474
191
1189
1758
3001
966
180
654
1278
BWFREQ
1110
832
67
662
1002
17.730
.0000
2000
134
100
0
348
3001
169
157
0
403
MAX 1110
1400
54
1220
1579
9.479
. 0005
2000
1885
160
1659
2111
3001
1207
120
959
1454
MAXLOC
1110
13.5
3.2
4.2
22.8
9.170
. 0006
2000
45.2
6.6
33.5
56.9
3001
26.2
8.1
13.4
39.0
MIN 1110
464
49
314
615
23.195
. 0000
2000
1274
111
1085
1463
3001
713
130
506
920
MINLOC
1110
94.0
1.4
83
105
2.887
. 0681
2000
78.4
10.4
64.8
92.1
3001
74.4
8.1
59.4
89.4
BWFORM
1110
935
60
789
1081
7.750
. 0015
2000
611
123
428
795
3001
494
76
293
694
INC 1110
28
12
0
95
11.408
. 0001
2000
277
65
193
362
3001
71
47
0
164


166
Schwagmeyer, P.L. (1980). Alarm calling behavior of the
thirteen-lined ground squirrel, Spermoohilus
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Stokes, A.W. (1961). Voice and social behavior of the chukar
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Struhsaker, T.T. (1967). Auditory communication among
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148
vervet alarm calls. The grunts given by vervets in four
social situations are also quite brief, averaging 125.5 msec
(range 75-235 msec). It is unlikely however that these
durational contrasts are of primary communicative
importance.
An interesting phenomenon found in vervet calls and not
seen among the C. olivaceus alarm calls was the pattern of
iterations and phrases that make up calls. Owren (1985)
defined iterations as any temporally continuous signal,
phrases as groups of iterations, and finally that calls were
comprised of a series of phrases. Iterations were the
smallest visible spectrogram unit with iterations occurring
repetitively in phrases. Calls could be either single or
grouped phrases. This classification system is reminiscent
of that required by Robinson (1979) and others who work on
the syntax of primate calls, where various elements of a
vocal system are combined, often in novel and
communicatively important ways. I found no evidence of
phrases, as defined here. The C. olivaceus alarm calls were
discrete vocalizations and were neither made up of definable
segments nor combined with other calls to form new
vocalizations. I found no evidence of either lexical or
phonological syntax. It would be interesting to see if
there were combinatory rules for vervet calls, such as seen
in other C. olivaceus calls (Robinson, 1984) or among titi
monkey (Callicebus moloch) calls (Robinson, 1979) .


<0B>
FREQUENCY (HZ)
Figure 6. Waterfall display of JM102, grrah variant 1111. This Fourier derived
display has frequency on x-axis, time on z-axis, and amplitude (dB) on Y-axis. The
individual spectra marked with the asterisk is shown, with an overlaid LPC-derieved
spectra, in figure 8.
110


42
Recording and Data Analysis Equipment
Two recording systems were used. Initially, from May to
July, 1986, a Sony TC-D5M cassette recorder and a Sennheiser
MKh-805 directional microphone were used ad libitum to
record monkey calls. Recording distances were less than
15m. The system's frequency sensitivity was flat from
approximately 50-15,000 Hz. These recordings were used to
determine the monkeys lexicon.
A Sony 8mm CCD-V9 video camera with a Sennheiser MK-415
directional microphone was used during the snake releases
and alarm call playbacks as both the auditory and visual
recording device. The camera image sensor produces 380,000
pixels and the audio dynamic range is >80 dB over a
frequency range of 30-15,000Hz. The camera has multiple
shutter speeds which allowed for high speed filming. Video
recordings were played back on a 19" Sony Trinitron TV.
Study Animals
The C. olivaceus studied here were part of a population
of approximately 300 animals that inhabit Hato Masaguaral.
This population contains approximately 12 groups, 4 of which
were studiedMain, White, Red, and Splinter. Most work was
done with Main group, which contained 21-27 animals
throughout the study: a single adult male, 5-8 adult
females, and assorted juveniles and infants. Table 3 shows
its composition at the beginning of the 1989 field season.
These groups have been continuously studied by Robinson


141
acoustic relationships compared to normal voicing. I should
not assume that other primates act otherwise. The question
then becomes how to determine what structural variability
has communicative importance. I will limit my statistical
analysis to differences between sexes because of a limited
sample. I will also point out overall differences between
individuals without a strict statistical analysis.
Sexual differences
We acoustically determine the sex of a human speaker by
their fundamental frequency: males typically have lower
voices than females, in large part, due to differences in
length of the vocal folds. I examined fundamental frequency
of similarly aged subadult C. olivaceus giving the same two
calls to determine if their was an analogous pattern.
The beginning fundamental frequency and caller identity
for qrrah variants 1312 and 13121 are given in table 17. A
Wilcoxon pairs analysis showed that for both calls there is
a weakly significant difference between fundamental
frequencies of males vs females (p= 0.069 for qrrah 1312 and
p=0.061 for qrrah 13121). Unfortunately these differences
are not uniform. The mean fundamental frequency for males
giving qrrah 1312 is higher than for females (x= 1351Hz,
o=344 vs x=1170Hz, 0=368) whereas it is lower for call 13121
(x=1255Hz, o=129 vs x=1328Hz, o=156). Given that the same
animals made these calls, these differences are difficult to
interpret except to say that the difference is unlike humans


17
(1977) and Dittus (1984) explicitly state that the food
calls of their study animals are symbolic. In the
chimpanzees and toque macaques, and perhaps in other
primates, these calls are given when a troop member locates
a superabundant food source. Typically, if the food was not
superabundant, the call was not given. Calls were
apparently nondirected, given to the troop as a whole. In
the macaque, when a feeding bout was initiated by a food
call, feeding duration was longer than those where no call
was given. Dittus concludes these calls convey information
about the presence of a food source, its quantity and
location" (Dittus, 1984, pg 476).
Dittus (1984) attempts to show that the macaque food
calls are not affective; that they are not communicating
only emotional state or probability of future action. He
argues that since the stimulus eliciting the call is
precise, specific to the discovery of abundant food in 98%
of the calls, the specificity is inherent in the call. I
would argue that, on the contrary, since this call is given
only when food is abundant, it would unambiguously signal to
others that the stimulus, food, is present, regardless of
whether the call is referring to the food itself or to the
emotional state stimulated by abundant food. In other
words, contextual specificity here is not sufficient proof
that the signal is symbolic. Dittus has demonstrated that
the message of the signal is the presence of abundant food.


122
The essential information then is 1) the fundamental
frequency for C. olivaceus alarm calls consistently begin
around 1000Hz and 2) with one exception, fundamental
frequencies remain relatively level at the beginning and
middle of most alarm calls, ranging around 1100Hz at the
beginning, 1000Hz at the middle, then dropping appreciably
at the end to around 800Hz. In the single exception, the
call to avian predators, variant 2000, the pitch goes up
from, on average, 973 Hz at the beginning to 1261 Hz in the
middle, then descends at the end to 1138 Hz.
Descriptive Statistics
Values for 20 variables describing 41 qrrahs from the
three locational call variants, including an analysis of
variance between the three groups, are given in table 13.
By examining the confidence intervals for overlapping
values we can see that certain calls are uniquely defined by
some of these variables. The following call characteristics
are evident:
Call 1111- least time increasing, the greatest
bandwidth, duration of frequency change is
longest, has the most inflections, and the
location of maximum dB and no dominance is
closest to the end of the call.
Call 3001- shortest duration and least dB difference.


8
connotative. Quine uses the example of a linguist trying to
understand what a native is saying when a rabbit scurries by
and the native says gavagai. Is the native in fact
signifying food, animal, rabbit, or something else? Meaning
or at least a working hypothesis of meaning is made by
correlating a reasonable number of stimulus events and the
accompanying utterance. The stimulus meaning is the class
of all stimulations that prompt that vocalization. If the
linguist hears the native say gavagai many times when
presented with a rabbit, he may assume that gavagai and
rabbit have the same stimulus meaning. Yet the linguist
cannot be sure that his translation is correct; there will
always be an indeterminacy between stimulus meaning and the
denoted meaning. A sort of working dictionary may be
compiled but the correspondence of word to stimulus event
will be imperfect because this is a mechanism for
translating discourse, not single words. Quine goes so far
as to state, in the tradition I believe of Turing and Godel,
"that rival systems of analytical hypotheses can fit the
totality of speech behavior to perfection, and can fit the
totality of dispositions to speech behavior as well, and
still specify mutually incompatible translations of
countless sentences insusceptible of independent control"
(Quine, 1960, pg 72). That is to say, given the same text


92
surprising, since one would expect that related animals
would respond more often to each others' calls. This result
can, by extension, be construed to further indicate that
animals do perhaps recognize each others' calls.
Lastly, if we examine the relationship between social
dominance and rate of correct response within the five adult
females we see that there is no correlation (r=0.4122,
Kendalls rank correlation).
Waahs
Another series of playback experiments was undertaken
using waahs. The same overall playback protocol was
followed. Fourteen playbacks were done using three calls
given by animals in two different groups (Table 8). In
eight of the fourteen playbacks the respondent reacted by
looking up to the open sky, three times they looked towards
the speakers, and three times there was no response. I
conclude from these experiments that when the monkeys
responded to these calls as alarms they reacted in every
case, 8/8 times, by looking up for an avian predator.
Response variations
Response variations by sex of the respondent is
interesting. In the eight calls played back to males, they
responded by looking to the sky on three occasions, whereas
females responded each of the five times. I was unable to
sex one respondent. When the caller was a male, monkeys
correctly responded 4/7 times while for females it was 2/2


Table 7. Matrix of graahs preceding other grrahs
1111
1120
1210
1221
1222
1311
1312
13121
13122
1321
2000
3001
1111
2
2
1
1120
3
1210 1221
1
1
2
2
1
3
2
1
2
Following Call
1222 1311 1312 13121
1
1
1
1
2
2
2
3
1
1
2
3
2
1
2
3
4
2
5
1
2
1
13122
1
2
1
1
1321
1
7
1
5
2000
1
1
2
1
1
2
3001
1
1
2
1
2
4
2
1
2
u>


I certify that I have read this study and that in my
opinion it conforms to acceptable standards of scholarly
presentation and is fully adequate, in scope and quality,
as a dissertation for the degree of Doctor of Philosophy.
This dissertation was submitted to the Graduate Faculty
of the School of Forest Resources and Conservation in the
College of Agriculture and to the Graduate School and was
accepted as partial fulfillment of the requirements for the
Dean, Graduate School


29
characteristics attended to by animals and whether
perceptual strategies are similar to those employed by
humans in speech. Ideally investigators will identify
perceptual sensitivities to particular signal
characteristics that are of communicative importance.
Investigators have studied species-specific perceptual
processing of communication signals in a limited set of old
World primates. The results of various studies by Sinnott
(1976, 1985) suggest that 1) primates do not discriminate
pure tone changes, the vowel cue, nearly as well as humans,
and 2) their discriminatory abilities used in
differentiating consonants, where both frequency and
intensity cues are crucial, are roughly similar to humans.
Petersen and others investigated the perception of coo
vocalizations by japanese macaques (Macaca fuscata)
(reviewed in Petersen, 1982) finding, for example,
similarities in neural lateralization patterns between
primates and humans. They also found that peak frequency
location, a factor central to discriminating among the
various macaque 'coo' vocalizations, shows a right ear
advantage. This is in contrast to how macaques hear pitch
changes, which shows a left or no ear advantage. These
findings are comparable to similar results in humans. There
is a right ear advantage in humans for stop consonants and
either a right or no ear advantage for vowels. In regard to
selective attention to changes of particular acoustic


177
AA77 AA79
MO MO
Variant 2100
MG106
WH
Variant 1112


87
monkeys than to playbacks, or were they habituating to the
presentations? To answer this requires a video record of
the monkeys response to actual alarm calls. I have only one
such record, where Jefe immediately looked in the direction
of a monkey that called at an approaching tayra. I have no
records of animals ignoring natural alarm calls. Lacking
such records, I must judge whether these experiments
represented realistic situations. Did the animals respond
as if another animal had seen a snake and then give a call
in alarm? Based on the behavior of responding animals
compared to responses to actual predators I feel these
responses were real, that the animals understood the calls
and responded as if another monkey was giving alarm calls at
a snake.
When a monkey did not respond, was this an indication
that it had not heard the call? Was it not concerned about
the snake? Lack of concern could be due to habituation or
as a result of only a single call being heard rather than
several. Lack of response is a difficult problem,
particularly when I have no measure of its frequency in
natural situations. I will assume that the specimen is the
authority and have used as my data set only those responses
where the experimental subject was apparently looking for a
snake. I have not used responses where monkeys looked
toward the speaker, other responses, or no responses.


38
objects and an action, whereas the latter seems to require a
conceptual grasp of the terms.
The above duality suggests a novel hypothesis about how
animals communicate about emotions and objects using both
affective and symbolic communication. This communication
system would use, first, purely affective vocalizations to
communicate about emotions and combine those signals to
refer to intermediate emotions, for example, the lexically
and phonologically combined vocalizations in the capuchin
and titi monkeys, and, second, use symbolic terms to refer
to objects, for example, alarm calls. According to this
hypothesis primates would combine calls to form other calls
referring to intermediate emotions using lexical syntax and
restrict their use of symbolic terms for named objects.
Sequences would be restricted to combinations of affective
vocalizations, where combining the emotive signals acts to
fluidly transpose the emotions into a third transitional
emotion that would have its own subjective reality.
Primates would give symbolic calls singly (or repetitions of
the same 'word') because they refer to an identifiable
object and they would not use them in a sentential form
because sentences cannot refer simply to objects. Quine
(1960) argued that only words have reference and that
sentences are identified by their logical truth. If my
hypothesis is correct, it may explain much of the primate
vocal behavior described above. They have, first, words for


155
trouble distinguishing primate calls. This is exactly the
opposite of what Green (1975) found with macaque coos and
Cheney and Seyfarth with both vervet grunts and alarm calls.
I suggest that primates use a variety of subtle acoustic
tactics to differentiate their calls and may involve species
specific sensory processing mechanismsboth at the level of
production and reception.
Distinctive Features and Language Capabilities
Lieberman (1969, 1984) has written extensively about
primate communication and the question of language
capability. After acoustic analysis of a limited set of
vocalizations and morphological analysis of various primate
vocal tracts he concluded that non-human primates were
incapable of producing human vowels due to limits in the
capability to modify the vocal tract. He concluded that
nonhuman primates are therefore incapable of language. I
differ with him at a fundamental level. A language requires
a diversity of perceptually distinct sounds. The more
sounds, the larger the potential lexicon. Humans achieve
this, in part, through vowel production. There are,
however, a wide diversity of human speech sounds that are
not vowels, including stop consonants, fricatives, and
glides. The fact that many if not all primates do not make
human-like vowels (but see Richman, 1976) does not mean that
they are therefore incapable of language. Instead primates
mav develop a language using entirely different distinctive


146
(DB)
Figure 16. Comparison of Fourier and LPC derived spectra of
GR69, a grrah variant 3001. The smooth LPC spectra overlays
the FFT spectra. Notice that both spectra are bimodal, at
approximately 1200 and 2400 Hz.


52
Grrahs
Graahs. in general, can be distinguished from other C.
olivaceus calls by two characterstics: 1) falling frequency
and 2) a breathy aspiration sound at the end. Other C.
olivaceus calls are downswept, chirps and hehs are good
examples, but the terminal aperiodic breathy sound is
distinctive to grrahs. The amount of frequency drop and
number of energy bands is highly variable, as will be seen
in later chapters.
Grrahs to snakes. Figure 2a illustrates narrow band
spectrograms of two grrahs given by an adult male, SH, in
Red group to a boa constrictor (Constrictor constrictor).
Note that they have 3-4 continuously falling energy bands
with the fundamental frequency beginning at approximately 1
kHz. Note also that the calls are less periodic, more noisy
at the end. Figure 2b is another spectrogram of a grrah to
a boa by Quay, an adult male in Splinter group. In this
call the pitch remains relatively steady until the end where
it first climbs then precipitously falls. Note again the
broad band noise at the end, particularly in the second
formant. The acoustics of grrahs to snakes is the subject
of chapter 5 which provides examples of many other grrah
variants. Notice that the calls just described are similar
to some of those given by Main group.
Graahs to humans. Unhabituated and, on occasion,
habituated capuchins would emit grrahs at humans. Figure 2c


ACKNOWLEDGEMENT
This research could not have been completed without the
help of many individuals. Drs. John Robinson, John
Eisenberg and Jay Whitehead provided guidance and assistance
throughout, for which I offer my deepest gratitude. Sr.
Tomas Blohm kindly offered me his hospitality at the ranch.
He has been steadfast in his support of this and many other
research projects, and a good bit of what is known about
neotropical ecology is in his debt. Dr. Bill Hardy
generously allowed me to use the equipment in the
Bioacoustics Lab of the Florida Museum of Natural History.
Lastly, I would like to thank two wonderful women, my mother
Mrs. J.D. Folsom and Kim Martin, for helping me through many
hard times.
ii


28
dampen and enhance frequency components of the entire
spectrum.
Richman further described the distinctions of vocal
onset and finds them similar to those employed by humans in
making consonants: gradual, sudden, and fricative. These
articulation features produce, in human speech, glides,
liquids, stops, and fricatives. He concluded that it appears
possible that geladas produce these consonant-like sounds by
changes in three places of articulation: labial, a velar
like position, and an intermediate position. The author made
no attempt to correlate these acoustic phonetic differences
with changes in behavior.
These studies illustrate that calls with similar
acoustic structure often have variants recognizable to the
vocalizing animals, though perhaps not to humans. Call
variants are often correlated with different behavioral
functions. Future taxonomists of animal calls should
recognize that phonological variability in animal
vocalizations is important yet often very difficult to
discern.
Acoustic perception in primates
The above examples of how various primates use
linguistic-like modifications of their communication signals
assume that the animals perceive these changes. Recently
there has been a significant increase in research on animal
psycho-acoustics investigating both the signal


131
-1.9 0.1 2.1 4.1 6.1
Figure 12. Plot of the first two principal components and
how they group 59 grrahs of all variants. The three
locational call variants are circled.


104
signal as the output of a model of speech production. These
speech production parameters include excitation parameters
related to the waveform source (typically vocal fold
vibration) and vocal tract response parameters (parameters
modelling the vocal tract resonance patterns).
Speech production, or in general vocal production by
most primates, involves activities of the larynx, pharynx,
and the oral cavity. The parametric representation of a
speech signal requires data on the fundamental frequency and
amplitude of the vibrating organ in the larynx with the
resulting harmonic frequencies and bandwidth. As a matter
of definition, the fundamental frequency of the signal is
the vibration frequency of the sound production organ. This
fundamental frequency may also have harmonics, which are
energy bands at integer multiples of that fundamental
frequency. For example, in one C. olivaceus alarm call the
fundamental frequency is 1250 Hz with harmonics at 2500 and
3750 Hz. Like harmonics, formants are acoustic energy bands
that may be present within a vocalization. They are
resonances of the fundamental frequency of the vibrating
organ and are produced higher in the vocal tract, in the
pharynx and oral cavity. Note that formants may simply be
harmonics, or, more complexly, as excursions from those
harmonic values caused by resonances of the vocal tract
produced by alterations in the vocal tract shape resulting
in impedance changes. In practice, formants are those


16
Seyfarth and Marler (1980) demonstrated that these
vocalizations were not simply affective or prescriptive of
particular responses. For example, they demonstrated in the
playback experiments that when an animal heard an alarm
call, it immediately looked in the appropriate direction for
the predator symbolized by the call (for example, into the
sky for an eagle) and not to the presumptive signaller to
indexically find out about what it was calling. As
discussed below, I maintain that the receiver's immediate
response to a callwhether looking to the signaller or to
the presumed external referentshould be the sine qua non
test of reference in field experiments.
In addition to alarm calls, several authors have argued
that various primates use symbolic communication in food
calls and agonistic recruitment calls (reviewed by Marler,
1985). Food calls are known to be used by chimpanzees (Pan
troglodytes) (Wrangham, 1977; Marler and Tenaza, 1977),
toque macaques (Macaca snica) (Dittus, 1984), black spider
monkeys (Ateles fuscipes) (Eisenberg, 1976), and wedge-
capped capuchin monkeys (C. olivaceus) (Oppenheimer and
Oppenheimer, 1973; Robinson, pers. comm.; Norris, personal
observation). These authors do not all maintain that food
calls are used as symbols for food. More typically the
earlier reports view these calls as affective; the animals
are indicating their high motivation to eat when presented
with particularly desirable or abundant food. Only Wrangham


76
due to its limited application and we can conclude nothing
about its usage.
Perhaps one of the most surprising aspects to the
capuchin's reactions to snakes was their calling at small
boas in the same manner as they did larger snakes. For some
reason the monkeys apparently felt threatened by a .68m
snake. Thirty arrahs of nine different variants were given
at it, twice as many as were given to the largest snake,
which was almost three times as long and over twenty times
as heavy.
Grrah variant 1130 was the second most commonly used
grrah and apparently used almost exclusively to snakes on
the ground. I remain skeptical, however, of concluding its
specific usage because it was also used by two animals in
two releases when snakes were in trees. Likewise, variant
1210 was used mostly toward snakes on the ground, but was
also given to snakes in a tree.
The production of many of the other grrah variants was
separated almost equally between one and two snakes or
snakes on the ground and snakes in a tree.
Examination of these results shows that the above call
variants are not always given when a snake is in the
prescribed location. For example, call variant 1111 was
given in only 5/8 situations when a snake was on the ground.
The snakes location is apparently not a necessary condition
for the utterance of a particular call. On the other hand,


121
4 msec of the signal. The period of the wave, measured from
the beginning to the peak of the waveform, is 0.83 msec,
which equals a 1200 Hz fundamental. Both of these values,
1187 HZ and 1200 Hz compare favorably to the estimate using
the CLA method in ILS of 1250 Hz ( 39 Hz). I did other DFT
measures from other examples with similar results. Based on
the similarities of the three techniques I conclude that
1) CLA is an accurate technique to measure these Figure 9.
fundamentals and 2) that £. olivaceus fundamental
frequencies are in fact around 1200 Hz. Additionally, based
on the clear periodicity of the real waveform and its
CEPSTRUM pattern this vocalization and, in all likelihood,
all the other alarm calls were, in part, voiced. This point
is important because it indicates that the source mechanism
is either the same or analogous to that of humans.
A last bit of confirming evidence of the accuracy of
these fundamental frequency values is the presence of
harmonic bands at multiples of these fundamentals in many
calls. If the actual fundamental frequency was not 1200 Hz
but rather, for example, 300 Hz and what we were seeing was
only the fourth harmonic (300 x 4= 1200) it is likely that
some of the higher energy bands would be multiples of 300 Hz
(e.g. 300 x 6= 1800) and not of 1200 Hz. From an
examination of the spectrograms of these calls it is clear
that energy bands higher than the fundamental are all
multiples of fundamentals beginning around 1000 Hz.


I certify that I have read this study and that in my
opinion it conforms to acceptable standards of scholarly
presentation and is fully adequate, in scope and quality,
as a dissertation for the degree of Doctor of Philosophy.
Jt\n\F. Eisenbergt^^Chafir
Katharine Ordway Professor of
Ecosystem Conservation
I certify that I have read this study and that in my
opinion it conforms to acceptable standards of scholarly
presentation and is fully adequate, in scope and quality,
as a dissertation for the degree of Doctor of Philosophy.
John Jb. kobinson
Associate Professor of Forest
/Resources and Conservation
I certify that I have read this study and that in my
opinion it conforms to acceptable standards of scholarly
presentation and is fully adequate, in scope and quality,
as a dissertation for the degree of Doctor of Philosophy.
Al
Michael Collopy
Professor of Foresl
Conservation
sources and
I certify that I have read this study and that in my
opinion it conforms to acceptable standards of scholarly
presentation and is fully adequate, in scope and quality,
as a dissertation for the degree of Doctor of Philosophy.
Will^m J. Hardy
Professor of Zoology
'£b&k.


9
two translators could come up with internally consistent yet
mutually incompatible translations.
I return now to the concept of reference, for it is
here that we confront the subject of what animals mean when
they vocalize. Take the situation where a primate sees a
predator, a snake for example, and utters an alarm call. To
what exactly does this vocalization refer? Let us, for the
sake of argument, assume that it does not refer simply to an
emotion, but rather to the snake. Is the call referring to
some concept of snake or to the snake itself? Does the
signaller have a concept of snake or is it simply naming
objects? What is the difference? The former would indicate
that the monkey uses conceptual modes for organizing its
world, while the latter indicates that the animal only names
objects and is not conscious of the properties that define a
snake no matter how precise and accurate its identifications
(see Marler, 1982; Cheney and Seyfarth, 1985, for a
discussion of how primates and other animals categorize
objects in their world).
To describe the dichotomy between a word and its
underlying conceptual reality I will borrow from set theory
and logic the notions of intension (not to be confused with
intention) and extension (Quine, 1960; Harre, 1984). "The
intension of a class is the concept under which any member
must fall, while its extension is the set of members which
satisfy that requirement. Intensions are concepts,


90
73 playbacks of calls by females. Within the sample of
female callers, there were no response differences by caller
age-class, with 11/25 correct responses to juvenile females
and 11/25 correct responses to adult females. (As a matter
of definition, correct is used here to mean that the animal
appropriately responded to the alarm call, which typically
was to look for the predator.) There do not appear to be
differential responses to females according to social
dominance, though the sample is small. Of the 73 playbacks
using female callers, 68 were done with the calls of four
monkeys, three adult females, (Whitey, Mo, and Pointy Face),
and a single juvenile (Hanna). Whitey and Hanna are
dominant within their age class, while Mo is intermediate
between Whitey and Pointy Face, the lowest ranking adult
female. Animals correctly responded to calls of the two
dominant animals in 30% of the playbacks, 35% to the
intermediate animal and 27% to the lowest ranking monkey.
There was only a single playback with related animals.
Mo did not respond to a call by her youngest son, Mike. Of
the four females, Mo had the most offspring in the troop,
Margo, Malli, Mike, and Modem, whereas Whitey had only a
single son, Winston present. Pointy Face's only surviving
offspring was a newborn in 1989, while Hanna had not
reproduced.


142
and may perhaps be due to voluntary differences in how these
individuals give these calls. Unfortunately these calls
offered the best sample of same aged individuals to test
differences between sexes.
Table 17.
Fundamental Frequency (Hz) Differences by Sex
Males Females
Call#
ID
Fund.
Call#
ID
Fund
Grrah
PU120
1312
Gr
1053
HA115
Ha
1177
PU121
Gr
1127
HQ175
Ml
702
MN105
Mn
1100
HQ9 2
Ml
952
MN107
Mn
1967
HQ179
Ml
763
HP30
HP27
Stu
Stu
1428
1428
Grrah
13121
MN106
Mn
1124
HA114
Ha
1190
MN101
Mn
1092
HA108
Ha
1538
AF3 5
Stu
1212
HA111
Ha
1538
AF34
Stu
1273
HQ151
Ml
1454
HP21
Stu
1250
HS4
Ml
1428
HP19
Stu
1428
HP22
Stu
1408
Discussion
Comparative Acoustic Structures
The acoustic features of two Cercopithecus aethiops
vocalizations have been described in detail: alarm calls
(Owren, 1985; Owren and Bernacki, 1988), and grunts
(Seyfarth and Cheney, 1984). The alarm calls described by
Owren (1985) and Owren and Bernacki (1988) consisted of a
sample of the various alarm calls recorded by Cheney and


5
Quiet arrawhs were given by an animal if it began lagging
behind the group. Robinson concluded that arrawhs acted to
reduce spacing between group members.
In a subsequent publication, Robinson (1984) identified
five basic vocalizations that were often syntactically
combinedchirps, trills, squaws, screams, and whistles.
The acoustic parameters that he used to differentiate these
social vocalizations are compared to alarm call acoustics in
a later section. Robinson first showed that these five
calls were used individually and their use varied
predictably with social circumstance expressing a continuum
of internal states ranging from contact seeking to contact
avoidance. The combined calls were relatively common,
comprising 38% of the 868 calls recorded. He concluded that
"the distribution of social circumstances in which compound
calls are given was intermediate between the distributions
of the constituent call types, which presumably indicates an
intermediate internal state" (Robinson, 1984, pg 76). This
study will be considered more fully below in a discussion of
how the syntactic structures of primate vocalizations
parallel in important ways those found in human language.
Before continuing I would like to mention a dilemma
facing anyone trying to explain what is occurring when
primates vocalize. As we shall see, many primates combine
vocalizations; yet in my experience in both South America
and Africa, monkeys most often give single, often apparently


24
varying energy wave that is subject to fatigue, distortion
and other perturbations.
Characterization of speech signals in articulatory or
auditory terms is described as distinctive features
analysis. Research on the distinctive features of human
speech has shown that a relatively small set of parameters
can define phonemes (for example, see Ladefoged, 1975).
This is an important step in the description of variability
because distinctive features provide a mechanism for
describing how a continously varying acoustic signal can be
described in discrete phonological units. The mechanisms by
which humans and other primates perceive their signals is
discussed below, but it is clear that many primates
distinguish signal variability in a manner similar to that
of humans.
Examples of phonetic descriptions
Phonetic differences in vocal behavior have been
described in a variety of primates (Table 2). For example,
Cleveland and Snowdon (1982) were able to describe eight
chirp variants of a tamarin, Saquinus oedipus. each given in
a definably different behavioral context. The authors
concluded that each call variant represented a particular
motivational state, i.e. that while they might have been
affective vocalizations, each call was uniquely paired to a
particular motivational state. A distinctive features
analysis showed that four statistically significant


99
response ranged from 21-59% of trials. While these values
may be high, particularly the 59% no response rate for grrah
variant 3001, these values are perhaps within the ranges
reported above for vervets responding to actual alarm calls
(not playbacks).
Independence of response. Seyfarth et al (1980)
report that vervets on average showed a particular response
only after first looking to another animal that had begun
that same response in 25% of trials. They concluded that
responses by individuals were independent of the behavior of
other monkeys. A comparable examination of results here
indicates that C. olivaceus were even less likely to look to
another animal; this happened during only 4 of the 94 alarm
call playbacks. This may be further evidence that both of
these primates use the call alone as indicator of the
presence of a predator. In 38% of the times a vervet gave
alarm calls, another animal also called, whereas for C.
olivaceus this was rare for waahs. but very common for
grrahs Lastly, during playback experiments, vervets only
once and Cebus twice responded to the playback by giving
alarm calls. In all cases these calls were from juveniles.
Aqe/sex class differences. Cheney and Seyfarth
(1981) found that, in general, no age/sex class was more
likely to give alarm calls than another. However, adult
males and females were significantly more likely to call at
leopards than to eagles or baboons. Unfortunately, Cheney


159
combination of purely affective vocalizations to communicate
about emotions and motivations and symbolic callswordsto
identify objects in their world (at least predators, but
also perhaps other objects such as food, mates, and social
relationships, for which there is only mixed evidence). A
determination of whether this is language requires first an
accepted definition of language and secondly a means to
fully translate the calls. The definition awaits the
linguists and philosophers, the latter research awaits
biologists investigating whether Cebus use certain calls to
refer to the many other objects that directly effect their
world: other predators, food, mates and the many other
things that they interact with daily.


TABLE OF CONTENTS
Page
ACKNOWLEDGMENTS
TABLE OF CONTENTS iii
LIST OF TABLES V
LIST OF FIGURES vi
ABSTRACT vii
CHAPTERS
1 INTRODUCTION 1
Traditional vs. Modern Perspective
on Animal Communication 1
Previous Studies of Cebus olivaceus
Communication 4
Linguistics and Animal Communication 6
Research Goals and Rationale 39
2 VOCAL RESPONSES TO PREDATORS BY
CEBUS OLIVACEUS 4 0
Methods and Materials 4 0
Responses of Cebus olivaceus to Predators... 46
Further Investigations 56
3 VOCAL RESPONSES TO RELEASED SNAKES 58
Methods and Materials 58
Results 61
Discussion 75
4 RESPONSES TO ALARM CALL PLAYBACKS 79
Methods and Materials 79
Results 88
Discussion 93
iii


65
clearly variable yet duration variability did not appear, in
advance of the analysis, to discriminate the calls as well
as frequency components. The only amplitude component
examined was the relative location of stressing, that is,
the position of maximum amplitude. I could not compare calls
by their overall amplitude, because the calls were neither
recorded at a standard distance nor did I use a calibration
tone when recording.
The arrah variant classification schema was defined
solely on the basis of the appearance of the spectrograms
rather than how the calls sounded to me, however there is,
of course, agreement between a spectrogram's appearance and
its sound. I assume there are acoustically relevent
features that my hearing may not attend to yet may be
important to the animals. I further assume that these
features will be evident on a spectrogram.
Using these features I parsed the 279 calls recorded
during ten snake releases into 15 arrah variants. The key,
including the number of qrrahs in each variant, is presented
in figure 4. Two calls were found only once each and were
excluded from further analysis. Initially, as a control on
individual variation of call variants, only those calls from
known vocalizers were classified, then once a robust
classification system was devised all suitable qrrahs were
included. Spectrograms of representative samples of each
qrrah variant are presented in appendix B. While there is


39
objects and, second, calls and sequences for emotions, each
reflecting an underlying reality of objects and feelings.
The remainder of this study will address representational
communication by C. olivaceus to determine whether they have
words for objects.
Research Goals and Rationale
The goal of this research is twofold: 1) to establish
whether Q. olivaceus semantically uses alarm calls to refer
to objects and, if so, 2) whether it is then able to
attribute qualities to those objects.
In this first chapter I reviewed the literature and
described the problems currently prominent in the study of
primate communication. In Chapter 2 I describe the alarm
calls of C. olivaceus. In Chapter 3 I describe the first of
two experiments on the semantics of a particular alarm call
type, the snake alarm call. The first experiment involves
release of different sizes and numbers of snakes. In chapter
4 I describe the second experiment where I play recordings
of the alarm calls given to the released snakes back to the
monkeys. The purpose of the first experiment is to determine
if C. olivaceus uses alarm calls as names for particular
predators and in the second experiment whether it attributes
qualities to them. In Chapter 5 I describe the acoustic
structure of C. olivaceus alarm calls. I finish with a
general summary of findings.


75
snake call. It seems that C. olivaceus do not restrict
alarm calls only to boas.
Other Responses
There were other responses to released snakes besides
qrrahs. For example, on four occasions monkeys hit a branch
while they or another animal were calling and each time a
different call was used: variants 2000, 1221, 1210, and
3001. At least twice monkeys knocked the snake out of a
tree by hitting the branch it was on.
Discussion
I conclude from the above data that C. olivaceus emit
certain qrrah variants when snakes are in particular
locations: qrrah variants 1111 and 3001 when a snake is on
the ground, qrrah variant 2000 when it is in a tree, and
variant 1222 when the snake is near the caller.
There is, at present, no convincing evidence that C.
olivaceus communicate about snake numbers or size. I found
the associations between qrrah variants and number of snakes
unconvincing, primarily because the sample size for each
call was small and the calls were given by few animals.
Grrah variant 13122 was given 6 times, of which 5 were given
by Whitey, while variant 2100 was given only 3 times, all by
Mo. A larger sample from more animals is needed before I am
convinced of their specific utility. I feel that the
specificity apparent in the usage of qrrah variant 13122 is


129
Principal components analysis. A principal
components analysis of the twenty acoustic variables found
that a linear combination of five variables defined four
components which described 95% of the variance within all
the arrahs:
Comp.1: .35 DUR +.60 Max +.37 BWFORM +.52 INC +.33 MAXLOC
Comp.2: -.23 DUR -.05 Max -.68 BWFORM +.26 INC +.67 MAXLOC
Comp.3: -.90 DUR +.35 Max +.21 BWFORM +.12 INC -.11 MAXLOC
Comp.4: -.01 DUR -.37 Max -.11 BWFORM +.80 INC -.45 MAXLOC
The first component, describing 40% of variance, most
heavily loads on the dominant formants' maximum frequency
and its increase. The other three variables are all equally
loaded, at 35% correlation. This component can therefore be
viewed as describing the behavior of the dominant formants
upper frequency. Notice that all the variables are
positively correlated.
In the second component, describing 27% of variance,
the amount of frequency change in the dominant formant
(BWFORM) is negatively correlated though equally loaded with
the location of maximum frequency (MAXLOC). This means that
where frequency change is great, the highest frequency is
likely to be early in the call. This component therefore
deals with place and amount of frequency drop.
The third component, describing 17% of variance, is
heavily loaded on call duration, which is negatively
correlated with three other variables, each of which has


*n r
. M0165 JE101
Mn Je
Variant 1320
-?Tr



* tr
~ |
*
^L

1?
-* *fc S"'
ir^l
- ib* -i
- V
i i§:
* ^
*> 7
Hai*_
sr ^
HS4 MN106
Ml Mn
Variant 13121
MG125 MG117
Wh Wh
Variant 13122


23
split similar sounding vocalizations (for a discussion of
this taxonomic question see Marler, 1982). There have been
several phonetic studies of primate vocalizations that point
to a richly variable repertoire where the variability may be
described as phonetic.
The diversity of alarm calls seen in this study prompts
the central question of my research, namely is this
variability meaningful to the monkeys. Green (1975) faced a
similar problem when describing the variability of coo
vocalizations by Japanese macaques. He adopted the terms
continuous/ discrete to describe morphological variability;
a continuous call was one in which the gradations of
variability were functional and meaningfully significant,
whereas a discrete signal had no functionally intermediate
forms (Green and Marler, 1979). Owren (1985) provides the
useful insight that it is not the presence of variability
alone that determines whether a signal is discrete or
continuous, rather it is the function of that variability.
Determining function is, of course, a difficult problem and
may lead to seemingly contradictory results. For example,
Winter (1969 a&b) concluded that squirrel monkey (Saimir
sciureus) used a discrete repertoire whereas Schott (1975)
concluded that the repertoire was continuous (=graded).
Such difficulties are perhaps not surprising. All vocal
signals contain some variability because the production
mechanism is a biological organ producing a continuously


162
Goodwin, D. (1953). Observations on voice and behaviour of
the red-legged partridge Alectoris rufa.
Ibis, 95, 581-614.
Gouzoules, S., Gouzoules, H., & Marler, P. (1984). Rhesus
monkey (Macaca mulatta) screams: representational
signalling in the recruitment of agonistic aid.
Anim. Behav., 32, 182-193.
Green S. (1975). Variations of vocal pattern with social
situation in the Japanese monkey (Macaca fuscata): a
field study. In Primate Behavior, vol 4, Developments
in Field and Laboratory Research, ed. L.A. Rosenblum,
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Green, S., & Marler, P. (1979). The analysis of animal
communcation. In Handbook of Behavioral Neurobiology,
vol 3, Social Behavior and Communication, ed. P. Marler
& J.G. Vandenbergh, New York, Plenum Press, 73-158.
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an avian alarm call system: the male domestic fowl?
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Hale, B., Schleidt, W.M., & Schein, M.W. (1969). The
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Baltimore, Maryland, Williams and Wilkins.
Harre, R. (1984). Vocabularies and theories. In The
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Hockett, C.F. (1960). Logical considerations in the study of
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Communication, ed. W.E. Lanyon and W.N. Tavolga,
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Hoogland, J.L. (1983). Nepotism and alarm calling in the
black-tailed prairie dog (Cvnomvs ludovicianus^.
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Konishi, M. (1963). The role of auditory feedback in the
vocal behaviour of the domestic fowl.
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Ladefoged, P. (1975). A Course in Phonetics. New York,
Harcourt Brace Jovanovich.


6
repetitive vocalizations. If the single call is referring to
an object, why is it given repetitiously? Correspondingly,
if calls can refer to objects, to what do combined calls
refer? These questions pose methodological problems of
vocal classification and procedures for determining meaning
of calls.
Linguistics and Animal Communication
Linguistics is an alternative explanatory system to the
traditional model of animal communication (see Snowdon,
1982). Linguistics deals with one form of communication,
language, at several levels including semantics, phonology,
and syntax. Because linguistic considerations are essential
to this project, I will define them, discuss how they apply
to animal communication, and give examples of pertinent
investigations.
Semantics
Semantics is the study of meaning. In linguistics,
meaning is acquired through denotationshow words are
defined in a standard text. Words are also understood
connotatively through the speakers conception of how a word
is used in normal conversation. Words have referents. A
word may symbolically refer to a physical object such as a
car, whereas other words have internal, abstract referents,
for example, anger. Green and Marler (1979) point out that
the assessment of any referent is initially an internal


165
Rabiner, L.R., & Schafer, R.W. (1978). Digital Processing
of Speech Signals. Englewood Cliffs, New Jersey,
Prentice-Hall Inc.
Rettig, N. (1978). Breeding behavior of the harpy eagle
(Harpa harvia). Auk, 629-643.
Reuter, J.R. (1986). The influence of group size on predator
scanning and foraging behaviour of wedgecapped capuchin
monkeys (Cebus olivaceus). Behaviour, 98, 240-258.
Richman, B. (1976). Some vocal distinctive features used by
gelada baboons. J. Acoust. Soc. Am., 60, 718-724.
Robinson, J.G. (1979) An analysis of the organization of
vocal communication in the titi monkey Callicebus
moloch. Z. Tierpschol., 49, 381-405.
Robinson, J.G. (1981). Spatial structure in foraging groups
of wedge-capped capuchin monkeys Cebus neorivittatus.
Anim. Behav., 29, 1036-1056.
Robinson, J.G. (1982). Vocal systems regulating within-
group spacing. In Primate Communication, ed. C.T.
Snowdon, C.H. Brown, & M.R. Petersen, Cambridge,
Cambridge Univ. Press., 94-116.
Robinson, J.G. (1984). Syntactic structures in the
vocalizations of wedge-capped capuchin monkeys, Cebus
olivaceus. Behaviour, 90, 46-79.
Robinson, J.G. (1986). Seasonal variation in use of time
and space by the wedge-capped capuchin monkey Cebus
olivaceus: implications for foraging theory.
Smithson. Contrib. Zool. 431
Robinson, S.R. (1980). Antipredator behavior and predator
recognition in Belding's ground squirrel.
Anim. Behav., 28, 840-852.
Robinson, S.R. (1981). Alarm communication in Belding's
ground squirrels. Z. Tierpsychol., 56, 150-168.
Schleidt, W.M. (1973). Tonic communication: continual
effects of discrete signs in animal communication
systems. J. Theor. Biology, 42, 359-386.
Schott, D. (1975). Quantiatvie analysis of the voalcal
reperoire of squirrel monkeys (Saimir sciureus ).
Z. Tierpsychol., 38, 225-250.


5 THE ACOUSTICS OF CEBUS OLIVACEUS
ALARM CALLS 102
Methods and Materials 102
Results 116
Discussion 142
6 SUMMARY AND CONCLUSIONS 157
REFERENCES 160
APPENDICES
A NARROWBAND SPECTROGRAMS OF GRRAHS TO A
0.68 m BOA 168
B NARROWBAND SPECTROGRAMS OF ALL GRRAH
VARIANTS 172
C DEFINITIONS OF ACOUSTIC VARIABLES 178
BIOGRAPHICAL SKETCH 18 0
iv


LIST OF TABLES
pages
TABLES
1 ANIMALS THAT ALARM CALL 14
2 PRIMATES EXHIBITING PHONETIC DIFFERENCES
IN THEIR COMMUNICATION SYSTEM 2 5
3 COMPOSITION OF MAIN GROUP 4 3
4 SNAKES USED IN CONTROLLED RELEASES 61
5 CONTEXTS OF GRRAH VARIANTS 68
6 GRRAH DIVERSITY BY INDIVIDUALS 71
7 MATRIX OF GRRAHS PRECEDING OTHER GRRAHS 73
8 RESPONSES TO ALARM CALL PLAYBACKS 81
9 RESPONSES TO PLAYBACKS OF GRRAHS 88
10 RESPONSE RATES FOR CALLS AND INDIVIDUALS.... 89
11 FUNDAMENTAL FREQUENCIES OF WAAHS AND GRRAHS. 117
12 FUNDAMENTAL FREQUENCIES FOR INDIVIDUALS 117
13 DESCRIPTIVE STATISTICS OF ACOUSTICS
PARAMETERS OF CEBUS ALARM CALLS 123
14 CORRELATION MATRIX OF ACOUSTIC VARIABLES.... 126
15 DEFINING VARIABLES FOR GRRAHS 13 6
16 AMPLITUDE CHANGES AT HARMONIC INTERVALS 139
17 FUNDAMENTAL FREQUENCY DIFFERENCES BY SEX.... 142
18 DISTINCTIVE FEATURES FOR PRIMATE
VOCALIZATIONS 150
v


2
fundamental dichotomy between language and animal
communication. The difference in perspective involves both
what is communicated and how it is done. Traditionally
animals were believed to communicate only their motivational
states: the signal indicates the sender's arousal or intent
for action. According to this perspective, animal calls are
affective. For example, an alarm call communicates fear.
This is in contrast with what humans communicate, where
symbols refer to, among other things, objects as well as
emotions.
A referent is the designatum of the signal (Green and
Marler, 1979). Reference may be to either an internal state
or to an external physical object. Communication about an
object may take any of several forms: iconic, indexical, or
symbolic. If variations in the object's physical form are
transformed along some parallel acoustic dimension then the
communication is iconic (Green and Marler, 1979). For
example, an alarm call that increases in duration or
amplitude in parallel with increased size of the predator is
iconically representing the size of the predator. Indexical
communication refers to an object through the simple
expedient of pointing at it (Green and Marler, 1979). That
is, the communicator refers to object by addressing it
physically by pointing at it. In the context of an alarm
call, a monkey would indexically communicate about a snake
by pointing at it. Symbolic communication uses the signal


85
the monkey, the alarm source (speakers), and the recordist.
Ideally where the monkey looksthe response variableis
isolated from either of the other two triangle points. For
example, in a playback of a ground snake alarm call the
monkey looks down, which is a different direction than
either to the recordist or sound source. Unfortunately, in
a playback of a near snake call, this condition could not be
met. When the monkey hears this call, it should look near
the sound source and determining whether it was looking at
the sound source or near it for a snake would be difficult.
For this reason, I did not test this call.
Playbacks of arrahs were done to 20 of the 21 available
animals in Main group, the lone exception being Winston, a
juvenile male. Playbacks of waahs were done to eight
different animals in Red, White, and Main groups.
Response Variable
This set of experiments tested where a monkey looked
upon hearing an alarm call. The measured response was the
direction the animal looked upon presentation of the
stimulus. Response criteria were:
1) Up or to the sky- The animal looks toward the open
sky, either directly above it or towards a break
in the canopy.
2) Down- The animal looks down- either directly below
it or towards the ground?
3) Foliage- The animal looks to adjacent foliage,
either by looking around itself or looking up into
foliage?
4) Speaker- The monkey looks toward the speaker.


101
interpreted the message to the receiver as being determined
by the sender's arousal level and other contextual
information (Leger, et al, 1979, Owings and Virginia, 1978).
This interpretation is consistent with the traditional
perspective of animal communication, that the meaning
depends on contextual and motivational information conveyed
about the caller's likely behavior. A re-examination of the
California ground squirrel acoustic data, however, showed
that particular calls, whistles and chatters, occur in
particular contexts (Leger, et al, 1980; Owings, et al,
1980). Playbacks of alarm calls showed that the call alone
was insuffient to elicit differential responses appropriate
for different predators. However when contextual
information about the numbers of animals calling was
provided with the alarm call, the squirrels responded
adaptively. Therefore, while the alarm calls alone are not
representational they can provide information about an
external object when simultaneously combined with other
information.
This requirement for additional, supplementary
information with ground squirrel alarm calls is in sharp
contrast to the alarm calls of both C. olivaceus and
Cercopithecus aethiops. In both primate species the calls
themselves were the only necessary and sufficient
communicatory element required to elicit adaptive, anti
predator responses.


140
This configuration is not found in human speech and is
probably not produced by vocal fold vibration in any way
analogous to humans (Rothman, personal comm.)*
I conclude from the spectral tilt data and the unusual
spectrogram configurations that the vocal production
mechanism of C. olivaceus needs extensive research. It is
important to note that however Cebus produce their signals
it is clear that they are capable of creating richly
variable signals.
Alarm Call Variability
Calls variability is an important yet difficult
subject. Variability may be due to the animal using the
variability to encode different meaning, i.e. the calls are
different, or the variability is due to random changes
during, for example, production. The problem then becomes to
determine which is random. This begs the question as to the
message of the signalthe call will have meaning at a
variety of levels, including the meaning of the call itself,
as well information on the sex, age, and possibly emotional
status of the caller. Lastly, only examining structural
variability through analysis of the acoustic elements may
miss the underlying purpose of that variability. This is,
in part, due to our uncertainty as to how the meaning of a
vocalization is encoded. Consider that human language can be
equally well understood when spoken in a falsetto or
whisper. These voicing changes drastically alter some of the


105
energy bands that diverge from the harmonic values of the
fundamental.
Linear prediction coding. A variety of algorithms
have been devised to accurately produce parametric
representations of human speech. Much of the acoustic data
used here is obtained using an algorithm called linear
predictive coding (LPC). This is an auto-regressive system
known in statistics as the linear mean square regression
technique. It is used to calculate three parameters
formant frequency, bandwidth, and amplitude. With these
calculated values a frequency spectra of the speech sound is
produced. Typically LPC parameters are used to reconstruct
a vocalization so that it sounds like the original, not
necessarily looks like it. Therefore a LPC analysis will
often have the same fundamental frequency as displayed in a
Fourier analysis while higher formants may not match
exactly, though to the ear the end products will sound the
same.
Linear prediction coding is used to predict a best
estimate of the next sample of a signal by observing past
samples and then minimizing the error between the predicted
value and the actual value. The estimated value, is
computed as an expansion of the previous M samples taken
during a particular time window
n=Q!10n-1 + a2n-2 + + amn.m eq.
where the various o's are M observed samples in the time


CHAPTER 1
INTRODUCTION
The wedge-capped capuchin (Cebidae: Cebus olivaceus =
Cebus niqrivittatus), also known as the weeper capuchin, is
a medium sized neotropical primate inhabiting much of South
America north of the the Amazon river (Eisenberg, 1989) As
its appellation ''weeper" implies, it is a notably vocal
animal. Among its many calls are a series of alarm calls
given to a variety of predators. These calls are the
subject of this dissertationunder what circumstances they
are used, their variability, and their meaning. This
search for meaning is a difficult one for which I am best
warned to remember the story of Captain Cook, recounted in
Cherry's seminal work, On Human Communication (1978). When
the famous explorer first saw a strange hopping animal he
inquired of a native as to what it was. The local
gentleman, of course, did not understand English and replied
"kangaroo," meaning "I don't know."
Traditional vs. Modern Perspective
on Animal Communication
The traditional view of animal communication,
universally held until only recently, is that there is a
1


133
Predicted Croup Actual Croup
1110
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*9
90.00
Figure 13. Classification of 59 grrahs of all variants
using a discriminate functions analysis of acoustic
variables DUR, Max, BWFORM, AND INC.


108
the entire call). Figure 5 illustrates both the digital
oscillogram and its analog spectrogram. They show a grrah
with a fundamental starting at 1250 Hz then descending to
approximately 450 Hz. Figures 6 and 7 are, respectively, a
Fourier derived waterfall and a LPC derived waterfall. In
general, the two waterfalls are alike. Next, in figure 8,
we examine individual spectra 40 msec into the call, near
the middle comparing the Fourier and LPC spectra. (These
same spectra occur in the waterfall, marked by the
asterisk). In this comparison the LPC spectra is smoothed
and overlays the FFT. Notice that the major frequency peaks
align well while higher formant peaks are somewhat offset.
This is due to the LPC system representing multiple higher
frequency peaks as a combination of fewer frequency bands.
I should note that spectral elements that are more than 10-
15 dB less intense than the peak frequency will have little
audible effect on the sound. I conclude that the LPC
representations of the alarm calls are accurate and assume
that they correctly portray the actual call.
Fundamental frequency
The fundamental frequency of a vocalization was
calculated using an ILS subroutine called CLA. With this
procedure period peaks are marked and fundamental frequency
is calculated as the reciprocal of the duration of a single
period. Five to 7 periods were measured at the beginning
and middle of exemplar calls of each alarm call variant. In


APPENDIX B
NARROWBAND SPECTROGRAMS OF ALL GRRAH VARIANTS,
INCLUDING CALL NUMBER, AND CALLER NAME.
BANDWIDTH= 80-8000 HZ, RESOLUTION= 45 HZ.


91
Variation bv respondent. Next we examine the
character of variations among respondents, by sex, age-
class, and familial and social relationships.
There was a significant difference (p=0.08, two-tailed
Mann-Whitney test) in correct response rates between sexes
of respondents by age class (adult, subadult, and juvenile).
That is, males in three age classes responded correctly
significantly more often than did females. There was no such
relationship by sex of individuals outside of age-class.
The question next arises as to whether certain animals
with many relatives in the group responded correctly more
often than animals with few relatives. For example, did the
members of Mo's family respond differently than the
unrelated group of males? Mo's family contained five
members, none of which were fathered by a male currently in
the group. When Mo's families rate of response is compared
to that of the four presumably unrelated males (White
Beard, Jefe, Griffin, and Stu) we find that there is no
significant difference (p=0.53, two tailed Mann-Whitney) in
response rate. This result is not surprising given that
there was only the single test of an animal responding to a
related animal's call. This explanation also assumes that
the monkeys recognize each others' voices. A better test
would compare data from responses to calls from related
animals to those from unrelated animals. However f animals
do not recognize each others' voices this result is somewhat


35
signals, while in others the combined signal elements have a
separate meaningphonological syntax.
Cleveland and Snowdon (1982) demonstrated lexical
syntax in cotton-top tamarins (Saguinus oedipus). They
found that two frequently used calls, an alarm chirp and a
low arousal alerting call, were combined such that animals
responded to this combined call as if it was the sum of the
constituent elements.
Robinson (1979) described the remarkably complex vocal
behavior of titi monkeys (Cal1icebus moloch). He found an
elaborate hierarchical system where calls are repeated to
form phrases, which may then be variably combined to form
sequences. He distinguished six types of sequences by their
different structure, as disclosed by transition
probabilities, their situational context, and the sex of the
caller. For example he describes duetting as "an
alternation of pant and bellow phrases follows the
introductory moaning phrase. As the sequence continues the
animals begin to add pumps, either after pants or bellows,
and insert honks between pant phrases and after pumps.
Honking usually ends the sequence" (Robinson, 1979, pg 393).
The driving force behind sequencing, and the effect
sequences have on meaning remains unknown; without such
knowledge we cannot determine if these syntactic patterns
are lexical or phonological. Robinson (1979) notes that the
hierarchies found in titi monkey calls are similar to those


CHAPTER 5
THE ACOUSTICS OF CEBUS OLIVACEUS ALARM CALLS
Methods and Materials
Recordings
The recordings analyzed here were those recorded during
the snake release experiments described above. The
recording equipment was the Sony video camera described in
Chapter 2.
Analysis System
The acoustics of C. olivaceus alarm calls were
described using two systems: spectrograms, both digital and
analog and the Interactive Laboratory System (ILS), a
digital spectrum analysis software system.
The analog spectrograms were produced on a Kay
Elemetrics model 7029A with a 80-8000 Hz bandwidth using
narrow band (resolution= 45 Hz). Digital spectrograms were
produced on a Uniscan II real-time spectrograph.
The ILS digital spectrum analysis package is a set of
92 subroutines used to digitally record and analyze sound.
Analysis was done on a Compaq Portable II computer using a
Data Translation DT2821 analog/ digital and digital/ analog
102


115
sensory phenomena where just noticeable differences (JND)
are constant across differing magnitudes of change. For
example, to perceive a change in pitch of a low frequency
tone requires only a small frequency change, whereas a JND
for a higher pitched tone requires a correspondingly larger
change. The ratio of *F/F2 remains constant, where *F is
the frequency change between two succeeding frequency
values, F^Fj. I reasoned that the difference limens
represented a fractional perceptible change at a particular
frequency and that this value could be viewed as a Weber
fraction and therefore would remain constant at any
frequency. The mean DL reported for primates by Sinnott
(1985) at 1000 Hz was 18.6 Hz, in other words, at 1000 Hz a
frequency change had to be greater than 18.6Hz to be
perceived. This DL could then be translated into a Weber
fraction of 18.6/1000= 0.0186. Therefore a 19Hz change at
1000Hz or a 27 Hz change at 1428 Hz (1428 x 0.0186= 27) were
both perceptible. In the following analysis I defined any
frequency change as being perceptible if the frequency
change ratio was greater than 0.0186. The determination of
when a frequency change was perceptible was important in
determining 1) place and number of inflections and 2)
percent of time that a call was increasing, decreasing, or
remaining steady.
The other important criterion was the dominant formant.
Remember that dB levels from the first three formants were


96
was to look in the direction of the call, then run passed
the speaker towards other animals. I suspect that she was
looking for her son.
The second and most important result reported here for
C. olivaceus alarm calls, that they provide locational
information about snakes, is not comparable to anything
reported for either vervets or other primate species.
Responses to other objects. Like C. olivaceus.
vervets gave alarm calls not only to confirmed and potential
predators but also to many non-predatory species, including
warthogs, various raptors and vultures, spoonbills, pigeons,
falling leaves, tortoises and mice (Seyfarth, et al, 1980).
Response frequency. Because predation levels were
much higher at Amboseli, Seyfarth and Cheney observed more
examples of responses to predators. It is instructive to
examine the number of responses during those situations when
actual monkey calls (as opposed to playback) were given. I
will limit this discussion to those vervet responses that
occurred when the monkeys were in trees, because this is the
situation most comparable with that of C. olivaceus. where
all playbacks were to animals in trees.
The vervet population under observation contained three
groups with a total of approximately 65 animals. On the 24
occasions when at least one individual showed a particular
response (other than calling) to a leopard alarm, they twice
ran higher in the tree. For 33 eagle calls, there were 13


13
investigation described here is to determine the necessary
condition, that is, if C. olivaceus use calls to indicate
whether they ascribe location to snakes. Subsequent
research will be necessary to establish the necessary and
sufficient condition of recurrent use of the partial, and
therefore whether Cebus olivaceus attribute qualities to
objects.
Examples of semantic communication
Ethologists have noted for some time that many birds
and mammals use alarm calls, which often are viewed as
examples of semantic communication. The studies sited in
table 1 are important because they bear on questions of kin
selection, evolution, sociality, and animal communication.
Two questions are pertinent here : 1) are these
vocalizations affective or symbolic?, and 2) is there
continuity throughout these taxa in what is referred to by
the alarm call. Evidence consistent with the symbolic
perspective is the requirement that these signals refer to
objects in an arbitrary manner (Sebeok, 1975; Hockett, 1960)
and that the signal production is disassociated from its
physiological manifestation. A separate view, held largely
by those studying various species of ground squirrels,
suggests that alarm calls do not classify the predator but
rather signify the different time constraints necessary for
predator avoidance.


]
I
JM107
JE119
AF27
BB117
MG113
M0126
Ha
Je
Wh
PF
Wh
Mo
Variant 1111 Variant 3001 Variant 2000
173


JelOl
Jel 16
Jell8
Jell9
Jel20
Je
Je
Ba
Je
Je
-

j
A
*
=
k
j
i
1
V
V
V
%
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Jel21
Ha 101
Hal02
Hal04
Je
Je
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Ha
169


69
Snake Context: Near the Vocalizer
1222 7/8 calls by 4 monkeys in 3/5 situations.
Overall, 107 calls of 13 types by at least 10
animals. 3/8 calls given when snake was on
the ground and 5/8 when snake was in a tree.
Snake Context: One Snake
13122 6/6 calls by 2 animals in 2 situations.
Overall, 151 calls of 14 call types by at
least 12 animals.
Snake Context: Two Snakes
2100 3/3 calls by 1 animal in 1 situation.
Overall, 121 calls of 14 types by at least 9
animals.
Snake Context: Medium Sized Snake
13122 6/6 calls by 2 animals in 2 situations.
In total, 101 grrahs were given by at least
13 different animals.
The evidence for other calls being given in particular
contexts is less compelling.
1122 5/7 calls to a snake on the ground, all by
unknown vocalizers in a single release; 2/7
in tree, by Whitey in one release and another
by an unknown animal in another release. It
was also given 5/7 when a snake was near.
Its specificity is suspect because of small
sample size and it was given by several
animals with snakes in both locations.
1130 28/32 calls to a snake on the ground by at
least 6 animals in 4/8 situations, however
3/32 calls by 2 different animals in 2
situations were given to snakes in a tree.
1210 14/17 calls to a snake on the ground by at
least 10 animals in 6/8 situations. 3/17
calls were to a snake in a tree, all by a
single animals in one situation.


154
single factor that separated two calls. There was no clear
pattern among Q. olivaceus. I felt that the relative
location of maximum dB within a call would be an important
cue and, as it turns out, was the distinguishing feature for
a single arrah variant. Like duration, variance of this
value may reflect its use in communicating other aspects to
the call, for example, emphasis.
The feature Q (the ratio of peak frequency to the
spectral bandwidth at that peak frequency) was useful in
distinguishing vervets alarm calls, but was used only in
that study.
In summary, four distinctive factors were used in at
least three of the four species calls: frequency bandwidth,
peak frequency and its increase, and relative durations of
frequency change or constancy. No single factor was used in
all calls by all species.
Based on these findings one would expect that primates
should be particularly sensitive to frequency changes.
Sinnott (1976, 1985) found that several primates were not as
sensitive to frequency changes as were humans. This of
course does not mean that they can not use frequency changes
as distinctive factors. Instead, it suggests that the
amount of frequency change needed to distinguish primate
calls may be greater than used in human vocal communication
and therefore may appear more extreme to humans. If this
was the case, we would expect that humans would have little


20
designate objects and events in the external world. Through
a paired comparison experiment in the field, the authors
tested the hypothesis that grunts convey specific context-
independent information. Five test stimuli were used; three
grunts given during intratroop interactions (when meeting a
dominant male, a dominant female, a subordinant female),
another grunt used when another vervet group was seen, and
the last when the signaller saw a troop member initiate
motion into the open. Results indicated a differential
response pattern to the grunts. While these experiments may
establish differential responses independent of context,
they do not prove that the signal was interpreted by the
receiver as anything other than the signaller was upset or
emoting. In fact, the evidence seems to indicate that the
receiver looks to the signaller, the speaker in the
playbacks, to see what was causing the signaller to grunt.
The appropriate experiment to indicate, for example, that
one grunt refers to a dominant animal would be to play a
call to an animal in the presence of a dominant animalas
if another vervet is grunting about that dominant animal
and determine if the receiver looks to the speaker or to the
dominant animal. Another test would be to determine if an
animal is greeted with different grunts before and after
changing its social rank relative to the vocalizer.
I conclude that the results of the semantic studies of
vervet alarm calls demonstrate that vervets use symbolic


147
fundamental frequencies of vervet vocalizations, the
spectral peaks starting at approximately 1 kHz will be 3rd
and higher harmonics of that fundamental. Higher frequency
spectral peaks may reflect different resonating mechanisms.
Vervet grunts also contain two spectral peaks. The
first peak is centered near the fundamental frequency and
Seyfarth and Cheney (1984) conclude that it is associated
with the action of the vocal cords. The second spectral
peak is between 550 and 900 Hz. It appears that the least
periodic C. olivaceus alarm call is more periodic and less
noisy than either vervet alarm calls or grunts.
The many differences noted here, both in source and
filter functions indicate that there are many fundamental
acoustical differences between C. olivaceus and
Cercopithecus aethiops vocalizations.
Other acoustic differences
All of the semantic calls described for vervets, alarm
calls and grunts alike, are very short duration, broad band
calls. The average duration for eagle calls was 29.6 msec
and 37.1 msec for the snake call. (Owren and Bernacki,
1988). This compares with 44 msec for the shortest duration
C. olivaceus alarm call, variant 3001, a grrah that was
noticeably noisy. The other two snake location calls
averaged 95 msec and 100 msec for, respectively, variants
1111 and 2000. The hawk alarm call averaged 254 msec.
Clearly, most C. olivaceus alarm calls are much longer than


80
toward the sky, into foliage, toward the camera, toward the
speaker, or toward other animals.
Playback Methodology and Schedule
Test stimuli
The schedule, call type, call, caller, and receiver for
21 alarm calls of four variants used in playbacks are
presented in table 8. Each call was selected for clarity
from the set of calls recorded during snake releases. Test
trials were designed to test the response of single animals
to single alarm call playback. Control trials involved
either playing 0.5 seconds of noise or setting up the
equipment and running a trial with no signal as if a signal
were played.
The playback tapes were produced by taking digitized
examples of the chosen calls and recording them onto 10
second tape loops. Amplitude of each call was equilibrated
by making the maximum amplitude of each oscillogram equal.
A single call was presented in each playback.
The calls used in the playbacks were the two ground
snake calls, grrah variants 1111 (5 replicates), 3001 (6
replicates), and the tree snake call, graah variant 2000 (7
replicates).
Playback methodology
The major goal in a playback experiment was to present
as realistic a stimulus as possible. Verisimilitude was
essential.


95
representative of the species as a whole, particularly given
the lack of large raptors. There are other regions of South
America where harpy eagles are likely to be major predators
of Cebus and it would be interesting to compare their
responses under those conditions with C. olivaceus at
Masaguaral and vervets in Africa. In Manu, Peru, for
example, Terborgh (Cheney and Wrangham, 1986) reports that
the estimated predation rate on Cebus apella was 13%,
similar to the Amboseli vervets.
The above differences are perhaps not surprising given
that the vervet's habitat and the predator commmunity are
very different from that of C. olivaceus in central
Venezuela. In Venezuela the predator load is currently low.
While the monkeys at the ranch are highly vigilant, the
likelihood that they may be attacked by a predator is very
different from that of vervets in the open spaces of
Amboseli, Kenya. In Amboseli there is an estimated 59
predators of vervets compared to 15 £. olivaceus predators
at Hato Masaguaral. The overall estimated predation rate is
five times higher in Africa, 15% vs 3% (Cheney and Wrangham,
1986).
It is worth noting that in the few playbacks where C.
olivaceus were on the ground, the monkey's first response
was to jump into a tree then look around. In another
playback of a snake alarm call to Margo, whose infant son
was separated from her at the time, her immediate response


156
features. Using this series of distinctive sounds they may
then define a language, albeit smaller and less complex than
human language, but nevertheless a communication system that
is more like human language than it is different from it.
I would offer that we know so little about the meaning
of animal signals because there is no easy mechanism for
accurate translation. This is the problem of the radical
translator (Quine, 1960) revisited. It is not that animals
are incapable of language, rather it is that we are
incapable of understanding all but a limited segment of it.
I expect that we will be limited in determining what animals
actually mean except in those calls that are actually object
orientedalarm calls, food calls, and perhaps a few calls
used in agonistic situations. Analyses of deception also
offer a means to better understand how animals use their
calls in a semantic manner. This should not be taken to
mean that I maintain that animals use language, but rather
that I suspect they use elements of languageobject
designation and attribution.


67
some variability within call types, this schema does appear
to group calls into similar appearing classes.
Context of the Call
Once the arrahs were classified the specificity of
grrah usage was examined in three contexts: snake location,
snake number, and snake size. Table 5 lists the context
where each call was given, the number of different releases
in which the call was given, and the number of different
animals giving the call. The following call variants appear
related to certain contexts:
Call Snake Context: On the Ground
Type
1111 26/27 calls by at least 5 animals in 5/8
possible situations, i.e. a snake was on the
ground in 8 releases, during 5 of which the
monkeys gave grrah variant 1111. In the
single exception there were two snakesone
in a vine and the other on the ground. The
call was given by an animal off camera so it
was not possible to determine to which snake
the call was directed, i.e. the snake may
have been on the ground.
3001 10/10 calls by at least three animals in 4/8
possible situations.
Overall, when a snake was on the ground 184
calls of 14 types were used by at least 13
different animals.
Snake Context: In Trees
2000 11/13 calls by at least 4 animals in 4/4
situations. In one exception, by Mo, the
snake may have been in a vine, it certainly
was soon afterwards.
13122 5/6 calls by one animal in 1/4 situations.
In total 83 calls of 12 types by at least 10
animals are given to arboreal snakes.


49
the presence of the threat. That they serve to warn other
animals of the presence and even location of the predator is
the subject of the experiments described below.
On one occasion I observed White group mob an ocelot
(Felis pardalis). The ocelot was walking on the ground
while the monkeys followed it in the trees, qrrahing as they
went. The monkeys appeared to be highly agitated whereas
the cat appeared quite indifferent. A similar incident was
witnessed with Red group. In both cases the monkeys were
already in trees while the cat was on the ground, many
animals vocalized, and the ocelot showed little interest in
the monkeys.
On a variety of occasions I observed tayra (Eira
barbara) move among White, Red, and Main group. For
example, just as I was preparing to conduct a playback
experiment with Red group I heard a qrrah and turned to see
a young tayra approach the area on the ground. I was
standing below an adult male, Finger, and several other
monkeys, less than 10m distance. The tayra saw me and
approached to investigate. The tayra appeared to ignore the
monkeys who casually watched it as they continued to feed.
At another time, for two weeks Main group inhabited a region
that was also frequented by a tayra. Sometimes the two
species would amicably occupy the same tree while on other
occasions the monkeys would chase the tayra out. On still
another occasion I saw an adult tayra move through the same


97
responsestypically looking up. For 10 snake calls there
were 17 responsesevenly divided between looking down and
approaching the snake.
A rough index of response frequency can be calculated
by taking the total number of responses, N, and dividing it
by the approximate average numbers of animals in a
troop(22), A, times the number of times in which at least
one animal responded, R.
% Responding= (N/ (A*R))*100
For example, for responses to snake calls, this is 17/220=
7.7%, for leopard calls it is 2/528= 0.4% and for eagle
calls it is 13/726= 1.8%. Even if only half of the troop
was visible to the observers (in which case these values
would need to be doubled), these results indicate that a
uniformly small percentage of the troop responded to any
particular alarm call. Similar results for vervets on the
ground were snake response= 5%, leopard= 3%, and eagle= 4%.
These are all higher than for vervets in trees but still low
overall.
These results are difficult to compare to results from
C. olivaceus because I did not quantify non-vocal response
to predators. It was my impression that non-vocal response
rates varied according to alarm call and presumed level of
threat. If an animal gave a waah to a raptor most animals
looked up but, with one exception, never ran into cover. If
a snake was discovered, most animals eventually joined in


CHAPTER 4
RESPONSES TO ALARM CALL PLAYBACKS
Methods and Materials
Playback, Data Recording, and Analysis Equipment
These experiments required both playback and recording
devices. Calls were played to the monkeys on a Sony TC-D5M
audio cassette recorder over two Sony SRS-30a self powered
speakers at the end of a 15m cable. The speakers were
typically hidden in a green bag placed under leaves in a
tree. The recording equipment was the same as that used in
the snake releases: Sony video camera and Sennheiser
directional microphone. The responses of the monkeys were
scored in the field by watching the video recording of the
playback through the camera (as a camcorder it was capable
of both playback and recording). Results were verified by
watching the playbacks on a larger TV.
Immediately after each playback a map was drawn of the
experimental scene, indicating the location and height of
the monkey, position of other animals, and the distance,
angle and height of both the speaker and camera relative to
the monkey. This map made it possible to judge where the
monkey looked upon hearing a call: down to the ground, up
79


84
Once the subject animal for a playback was chosen, I
followed that animal until the conditions for a playback
were appropriate. I tried to set up each trial before a
monkey was near, though this was at times difficult and the
trials could not always be run. Speakers were always set in
trees, never on the ground, because this was the typical
position of a calling monkey and signals broadcast better
off the ground. I ran a trial when the experimental subject
1) was relatively near the speaker,
2) in clear view and roughly facing the camera,
3) the speaker was out of sight of the monkey,
4) the animal producing the signal (recorded the
previous year) was out of sight.
If these conditions were met I then started filming the
animal and then played the tape loop and continued filming
for at least 30 seconds after the call. This playback
protocol was developed in 1988, with Red group. Data taken
during that period were not used in this analysis.
Playbacks often were to only one or a few animals,
which meant that other animals would not hear the playback.
Thus, in some cases it was possible to do several playback
experiments in a single day. I never repeated the same
trial with the same animal on the same day.
For procedural reasons dealing with interpretation
difficulties, grrah variant 1222, the call given when snakes
were near, was not tested. When considering a playback
experiment it is instructive to think in geometrical terms.
Playback experiments form a triangle, the three points being


98
mobbing the snake and produced alarm calls. On the three
occasions I saw monkeys responding to an ocelot, nearly all
animals appeared to watch the predator, some called, but few
more actively protected themselves. Seyfarth, Cheney, and
Marler (1980) report no response to a leopard call except
running into a tree or into cover. That is, the vervets did
not first look for the predator, but instead immediately
moved to protect themselves. This is contrast to their
response to a raptor in which case they most often looked
up. This is most likely a product of the severity of the
threat. Overall, I conclude that both C. olivaceus and
Cercopithecus aethiops gave relatively few non-vocal
responses other than looking for a predator. This may be an
indication of their reliance on alarm calls for their major
protection against predators.
It would be valuable to compare capuchin and vervet
rates of no response to alarm call playbacks. Unfortunately
this is difficult due to procedural differences. In this
study, I focused on the responses of a single animal while
Seyfarth et al (1980) report results from many animals
combined over many trials. For example, in the 9 trials
where a snake alarm call was presented to vervets in trees
(the comparable context to trials in this study) there were
a total of 27 responses. Unfortunately, the authors do not
report how many animals could have possibly responded. In
the playback experiments reported here, the rate of no


82
Table 8-continued.
b) Responses to call variant 3001.
Responses
No
Date Call Sp. Rec. Up/Sky Down Fol. Spkr. Res. Other
Ground Calls-3001
4/22
HA105
HA
ML
X
4/22
HA105
HA
ML
X
4/22
HA105
HA
MO
X
5/4
BB117
PF
STU
X
5/4
BB117
PF
STU
X
5/17
BB117
PF
BA
X
5/17
BB117
PF
ON X
5/29
AF27
WH
ON
5/29
AF27
WH
AM
5/29
AF27
WH
PU
X
6/13
HA105
HA
MO
X
6/13
HA105
HA
WH
X
6/13
HA105
HA
WB
X
6/13
HA105
HA
STU
6/13
HA105
HA
WH
6/17
BB118
PF
AM
X
6/17
BB118
PF
AM
X
6/17
BB118
PF
BA
X
6/17
BB118
PF
ML
X
6/23
HA105
HA
MG
6/23
HA105
HA
BU
X
6/26
HA105
HA
STU
X
6/26
HA105
HA
STU
X
6/26
HA105
HA
WB
X
7/6
AD12
HA
STU
X
7/13
AD12
HA
WB
X
7/13
AD12
HA
AL
X
7/15
AD12
HA
PF
X
7/15
AD12
HA
ON
X
7/15
AD12
HA
7
X
7/15
AD12
HA
AM, ON
7/15
AD12
HA
MO
X
7/15
AD12
HA
MO
X
7/15
AD12
HA
MO
X
7/17
BB126
PF
MG
7/17
BB126
PF
AL, ON
X
7/17
BB126
PF
AL
X
Totals 1 6 1 0 22 7


71
Table 6. Grrah diversity by individuals.
Females Males
uaix
Type
Am
Ha
Md
Ml
Mo
Pf
Pu
Wh
Ba
Gr
Je
Mi
Mn
St
WB
a
9
1111
6
1
2
1
3
3
10
1112
1
1
1120
3
4
3
2
2
2
12
1210
3
5
1
1
1
1
1
1
1
4
11
1221
8
3
1
1
1
1
1
14
1222
2
3
1
2
2
6
1311
5
2
2
7
4
2
1
3
20
1312
2
3
3
2
2
7
5
13121
3
2
9
3
2
5
2
9
17
13122
5
0
5
1321
7
2
4
2
3
2
2
3
1
8
18
2000
8
1
1
1
2
9
2100
3
0
3
3000
3
2
5
1
0
11
3001
3
3
1
0
7
Tot.
15
22
1
26
47
10
7
21
2
7
9
4
7
10
2
41
149
H' = .
45
0
.94
.67
0
. 66
.47
0
.91

87
.83
.65
.91
. 56
.45
. 59
1
. 13


143
Seyfarth in their investigation of the semantics of vervet
alarm calls. The description of grunts followed other
investigations of semantic communication by vervets and how
apparently identical grunts were used in different social
situations (Cheney and Seyfarth, 1982) .
Following the paradigm used to model human speech, I
will contrast the alarm call acoustics of vervets with those
of Q. olivaceus by comparing first, the source function of
the signal then the filter function, e.g. the source
waveform and how it is altered by the vocal tract. Next I
will compare the distinctive features possibly used by
several primates to distinguish their calls.
Source Function
The most striking difference between the calls of these
two primates is the fundamental frequency. These two
species are approximately the same size yet the fundamental
frequency of their calls are very different. For the two
vervet alarm calls, the fundamental frequencies were
statistically the same, averaging 331 Hz (range 225-368 Hz).
For the various C. olivaceus qrrahs this same value was 1173
Hz (range 833-1428 Hz). This difference is probably due to
a much shorter vibrating organ in the C. olivaceus vocal
production mechanism.
The lack of tonality to either vervet grunts or alarm
calls is striking. The spectrograms of their alarm calls
are almost uniform vertical lines as compared to the


15
Alarm calls and other semantic vocalizations have been
investigated at length in several primate species, in
particular the vervet, Cercopithecus aethiops. Struhsaker
(1967) first noted that vervets use a variety of alarm
calls, given to at least three different predators.
Seyfarth, Cheney, and Marler (1980) further demonstrated
through playback experiments that a particular call, for
example a martial eagle alarm call, prompted a predictable,
adaptive anti-predator response pattern even in the absence
of an eagle. Additional adaptive responses were elicited
by leopard and python alarm calls, with the response best
suited to protect against the differing attack strategies.
Each of these three calls was acoustically distinctive.
From the perspective of the signaler, particular predators
had unique names, while from the receiver's perspective, a
certain signal meant a particular predator which required a
specific adaptive response.
These signals satisfy the basic requirements of
symbolic communication by being arbitrary, disassociated
from physiological manifestation, and non-iconic, (see
Altmann, 1967, and Owren, 1985 for discussions of Hockett's
defining properties of symbolic communication). The visual
image of an eagle, for example, stimulates the vervet and
through a process of internal reference to emotive and other
dispositions that image is transformed into a signal that
prompts an equivalent mental image in a receiver. Cheney,


10
extensions are sets of things (Harre, 1984, pg 101). The
extensions of a class therefore are the members comprising
that class, while intensions are the unifying concepts that
define the class. The exclusive use of extensions would
therefore mean that the user has no knowledge about what
characterizes and differentiates that object from others.
Here clearly we are speaking of what the animal knows and
intends in its communication.
Quine notes that attributes are monadic intensions,
described by the notation "to be an object x such that . .
x," while relations are dyadic intensions, described "to be
objects x and y such that . x . y . ." (Quine,
1960, pg 164). In other words, for an object to have
attributes or be a part of a relationship requires a
conceptual graspan intensional graspof the object. I am
therefore distinguishing communication using extensions from
communication using intensions. Alarm calls may indicate
that the primate is using extensional information, for
example, that the predator is a member of a list of
predators. Attributing qualities to that predator however
proves that the monkey conceptually grasps the defining
character of that predator. I would argue that current
evidence shows only that primates are able to name an
object, x, and there is as yet no evidence to suggest that
they are able to talk about such objects. There is still no
evidence that nonhuman primates are able to symbolize


12
attributes, for example, big snake or ground snake. In the
fourth phase, analogy is used to apply relative terms to
singular or general terms to form other general terms, for
example, larger than a small snake. All phases beyond the
first reference phase require conceptual knowledge of the
objectusing these three further levels requires
intensional knowledge.
Attribution therefore involves the addition of
information. This may be accomplished in a variety of ways.
In language we achieve this through the use of additional
words or modifications of existing words. In any event,
attribution involves specifying additional meaning to a
word. In linguistics this additional element of meaning may
be a phone, morpheme, or clause. These terms, to be defined
below, are called recurrent partials. To discover the use
and meaning of a partial requires finding recurrent use with
another signal to similarly modify it. If capuchins use a
signal to indicate the location of a snake, for example, in
a tree or on the ground, the necessary and sufficient proof
of the meaning of the partial (and therefore whether the
animals are attributing qualities to objects) is to find it
being used in connection with another object that could also
be found in a tree or on the ground. Do the monkeys, for
example, indicate whether a tayra is in a tree or on the
ground using the same signals that they may use when making
the same distinction about snakes? The goal of the


47
(Allouatta seniculus), I never heard an alarm call. In the
case of raptors, Oppenheimer and Oppenheimer (1973) write
that when a bird swiftly flew over the animals one or more
C. olivaceus would give a single qrrah then drop to lower
branches or move deeper into the trees. I differ with the
Oppenheimers by distinguishing alarms calls to birds as
being acoustically distinct from other alarm calls, with
waahs given to avian threats and arrahs given to terrestrial
and arboreal threats.
In the presence of avian threats the monkeys typically
gave a single waah. looked up, and on rare occasions moved
away from the perceived threat. A variety of birds elicited
waahs from C. olivaceus: ornate hawk eagles, road-side
hawks, zoned tailed hawks, forest falcons, laughing falcons
and various species of vultures. I should note that with
some birds the monkeys waahed only after they appeared to be
surprised as the birds flushed in front of them or as they
flew low overhead: chachalacas (Ortalis ruficauda),
currosows (Crax daubentoni), egrets, ibis, muscovy ducks
(Oxvura dominica), and macaws (Ara macap). I will recount,
from field notes, two examples of interactions between C.
olivaceus and hawks.
On one occasion a road-side hawk attacked a juvenile
female crossing a tree gap on a vine. As the hawk
approached, other animals waahed then the hawk hit the
monkey's dorsal side without apparently hurting it.
On another occasion an adult female carrying a newborn
was moving on a branch when a laughing falcon (that had
been displaced by a harrassing monkeys) flew 2-3 m over


31
adopted a more general notion of syntax. To them syntax is
simply a system of rules that will generate and predict
sequences of signals, thus they view syntax as a
probabilistic phenomenon. Altmann (1965) noted for example
that in many vocal sequences first-order Markov
(predictable) processes describe transition probabilities
and are equivalent to grammars.
Marler and Tenaza (1977) differentiated two forms of
syntax: phonological (called phonetic by Snowdon, 1982)
syntax and lexical syntax. Phonological syntax is a system
for arranging communicative components into more than a
single pattern, each with a distinctive function. In
language it is analogous to word formation through phonemic
rearrangement. In lexical syntax "compound signals derive
their meaning from the multiplexing of the meanings of the
components as used separately or in other combinations"
(Marler and Tenaza, 1977, pg 25). Linguistically it is
analogous to phrase formation through word combination so
that the product is the sum of the meanings of the
individual elements.
Several additional points about syntax are pertinent.
First, in parallel to the semantic classes of agent,
patient, instrument, and object there are syntactic classes
such as noun phrase, verb phrase, and determiners. These
syntactic classes are necessary for modeling human language.
Premack (1985) makes the valuable point "that syntax cannot


30
parameters, it was found that, when pitch is varied, there
is selective attention to peak frequency but when, in the
reverse experiment, peak frequency is varied, there is poor
attention to pitch. These results indicate that Japanese
macaques take a discretely used acoustic cue, peak position,
and use it to partition vocalizations where peak frequency
position continuously varies. The perceptual processes
involved, selective attention, perceptual compensation, and
partitioning, closely match those used by humans in speech
perception. These results indicate that both human and
nonhuman primates apparently detect communicatively salient
features in their signals in similar manners.
Syntax
Syntax is the study of word order. Syntax describes
the ways in which words are put together to form phrases,
clauses, and sentences. Syntax describes another layer of
meaning, one provided by word order. It is through syntax
that we distinguish 'dog bites man' from 'man bites dog'.
The concept of syntax itself is layered, with the
distinctions centered around the use of meaning. At the
linguistic level Chomsky defined syntax as a "system
constituted by rules that interact to determine the form and
intrinsic meaning of a potentially infinite number of
sentences" (Chomsky, 1972, pg 69). At a functional level
several students of animal communication (Robinson, 1979,
1984; Snowdon, 1982; Cleveland and Snowdon, 1982) have


171
Mml03
Mo
Mml04
Mo
Hall5
Ha
*
MmlOl
Mo
Mml02
Mo
Mml06
Mo
Mml07
Mo
MmlO
Mo


32
be derived from semantics. No metamorphosis has been
demonstrated for turning the semantic caterpillar into the
syntactic butterfly: agent, recipient, and the like, no
matter how abstractly construed, will not turn into noun
phrase, verb phrase, etc." (pg 284). If primates can, as I
suspect, name objects and are able to differentiate agent
from patient, there still is no evidence to suggest that
they are capable of the purely linguistic differentiation of
noun phrase from verb phrase. Secondly, Chomsky's
conception of syntax has progressed beyond the definition
given above. He now views mental representations,
traditionally within the purview of semantics, as being a
form of syntax.
The study of the relation of syntactic structures
to models, "pictures," and the like, should be regarded
as pure syntax, the study of mental representations, to
be supplemented by a theory of the relation these
mental objects bear to the world . Thus, the shift
towards a computational theory of mind encompasses a
substantial part of what has been called "semantics" as
well. (Chomsky, 1986, pg 45).
Additionally, Chomsky views universal grammars as being
endowments due to innate, biologically determined
principles. This is our language faculty, our "language
acquisition device, an innate component of the human mind
that yields a particular language through interaction with
presented experience, a device that converts experience into
a system of knowledge attained: knowledge of one or another
language" (Chomsky, 1986, pg 3). In this new formulation I


34
Investigating such matters requires knowing the
phonetic structure of the communication system. All
biological signals have variability but only some of it is
communicatively meaningful. The alarm call classification
scheme described below parses the various alarm calls into a
number of variants. Without knowing the phonetic structures
of the communication system, I am unable to determine if C.
olivaceus alters its signals with internal modifications. I
am, however, able to determine whether calls are combined. I
will limit this study to the description of associations
between qrrahs alone because to accurately determine the
association between particular alarm calls and other calls,
huhs for example, requires knowing how the other calls vary.
Examples of syntactic communication
A wide assortment of primates use syntactic
combinations of different callslong calls of gray-cheeked
mangabeys (Cercocebus albiqena) (Waser, 1975), chimpanzees
(Pan troglodytes) (Marler and Hobbet, 1975), various gibbon
species (Tembrock, 1974; Tenaza, 1976; Marshall and
Marshall, 1976); alarm calls of cotton-top tamarin (Saquinus
oedipus) (Cleveland and Snowdon, 1982) ; intragroup calls in
the pygmy marmoset (Cebuella pygmea) (Snowdon, 1982) and
wedge-capped capuchin monkey (C. olivaceus) (Robinson, 1984)
and "singing" in titi monkeys (Callicebus moloch) (Moynihan,
1966; Robinson, 1979). Some of these combinatory signals
use lexical syntax to impart additional information to the


130
much less loading. This component clearly deals with call
duration.
The last component, describing only 11% of variance,
loads most heavily on frequency increase, but unlike the
first component, increase is oppositely correlated with
maximum frequency and its location. This component should be
viewed as describing primarily frequency increase.
Overall these components appear to describe C. olivaceus
arrahs using variables loading primarily on frequency and
not duration. The first component deals predominately with
the maximum frequency and its increase, while the second
component deals inversely with the place and amount of
frequency decrease. Next we shall see how these components
describe the individual calls.
Figure 12 is a plot of the first two principal
components and how they array 59 qrrahsall the locational
calls and 2 exemplars from each of the other call variants.
Values for component 1 are equally low for the two ground
calls, 1111 and 3001, while for the arboreal call, 2000,
they are typically higher. Values for component 2 are low
in call 1111 and equally high in calls 3001 and 2000. From
these component values the following can be inferred about
the calls:
Call 1111- While maximum frequency and its increase
were intermediate to low, a large frequency
drop from an early maximum was important.


120
Figure 11. A CEPSTRUM waveform for a grrah variant 13121.
The peak of the waveform is at 0.83 msec, indicating a
fundamental frequency of 1200 Hz.


11
something about x such that x has a property. To paraphrase
Bertrand Russell, no matter how adept a vervet may be at
identifying its mother, it's not able to explain that its
parents are poor but honest. If this view is correct, a
monkey's lexicon is comprised of extensions of classes.
The animal is not able to conceptually describe aspects of
the object or to talk about the object, it only names
objects. Correspondingly, if it uses intensions of the
class of predators it should be able to go beyond simple
object naming and attribute qualities, such as number, size,
or location, to the object.
To show that an animal has a conceptual grasp requires
that we find situations in which the signaller adds
specificity about the object, for example, by using
adjectives. Adjectives specify attributes to objects.
Quine (1960) describes four levels of reference where
objects of reference are further specified, from the general
to the specific. In the first phase, objects are named
through a process of reinforcement, extinction (learning),
and ostensin. Ostensin is the direct experiential
association with the object of reference, for example, that
a certain object is called snake by other monkeys. Word
usage at this phase is of extensions of a class. In the
second phase, these general terms are paired with
demonstrative singular terms, for example, this snake or
that snake. The third phase compounds general terms with


77
the emission of a call is sufficient for determining the
location of a snake. That is, whenever a monkey gives a
orrah. any other monkey hearing it would know where a snake
was located. Stated another way, the combined probability
of qrrah variant 1111 being given when a snake is on the
ground is 5 of 8 whereas the probability that a snake is on
the ground when qrrah variant 1111 is given is 26 of 27.
The three locational calls were not common; never more
than 14% of grrahs given in a particular context. Of
course, meaning should not depend on the frequency of the
call; a call given once would be sufficient to indicate
location.
There were fewer calls to snakes in trees than to
snakes on the ground but the average number of calls of any
variant per situation was approximately the same. For
example, qrrah variant 1311 was given 27 times in 7
situations (3.9 calls/ situation) when the snake was on the
ground and 9 times in 3 situations (3 calls/ situation) when
the snake was in a tree.
Examination of the various qrrah variants in appendix B
shows that the classification schema is robust, yet there is
some overlap; differences between some variants is slight.
These similarities may be due to their being syntactic
combinations of separate calls or they may be, in fact,
distinct calls representing different objects or emotions.


51
C. olivaceus were not measured here, specifically vigilance
for predators. It was my impression however that males were
not more likely to give alarm calls or mob a predator. As
discussed below, in vocal responses to released snakes,
females gave many more alarm calls than males, which is
different from what van Schaik found in the other Cebus
species.
The differences in response by C. olivaceus to avian
and terrestrial predators, particularly the number of calls
given, is explained well in a geometric model of alarm call
behavior (Taylor et al., 1990). The authors show that slow
moving predators should elicit more calls than fast moving
predators. This was found to be the case here where waahs
were typically given once while many grrahs were often
given.
Alarm Calls
Cebus olivaceus give alarm calls to a variety of
animals, as is clear from the above descriptions. While I
divide these calls into two types, waahs and grrahs. there
appears to be great variability within these call types. I
will examine in greater detail how C. olivaceus respond to
snakes and the variability within the snake alarm calls in
subsequent chapters. Here I will provide examples of grrahs
and waahs given to a variety of predators and apparent
threats. These calls can be compared to other C. olivaceus
calls presented in Robinson (1982, 1984).


26
perspective, the monkey stressed the higher frequencies of
the snake calls and stressed the lower frequencies of the
eagle call.
Spectral characteristics could also differentiate these
calls. "If two or three spectral peaks occur but do not
differ more than 3 dB in amplitude, or one stronger peak
above 1750 Hz occurs, it is a snake call. If two or three
spectral peaks occur, one peak below 1750 Hz is at least 3
dB stronger than any other peak, and an identical peak
occurs above 1750 Hz, it is a eagle call" (Owren and
Bernacki, 1988, pg 1933). These features correctly
classified the two calls 89% of the time. These features,
while including frequency qualities, still primarily depend
on the amplitude characteristics best described using the
tilt feature. Spectrograms of these two calls (fig. 1,
Owren and Bernacki, 1988) show that the snake alarm call is
considerably more wide band than the eagle call. That there
appears to be no energy above 4 kHz in the eagle call is an
artifact of spectrograms, which have a limited amplitude
resolution. The energy above 4 kHz, as seen in the spectra,
simply is louder in snake calls than in eagle calls
resulting in a greater spectral tilt in the eagle calls.
Seyfarth and Cheney (1984) concluded that vervets may
use three acoustic features to differentiate four grunts
given in different social situations: frequency of the
spectral peak associated with the fundamental (F0) ,


36
derived by phrase structure grammars. Further analysis of
sequencing is severely hampered by our failure to understand
what the vocalizations mean; we lack a semantic context to
place the calls. Chomsky differentiated phrase structure
grammars from more simple sequential models by the
characteristics of nested dependencies: 'if. . then'
structures. Syntactic structure may depend on the occurrence
of particular words, with independent order for intervening
words (see Robinson, 1979). Alternatively, responses to
different sequences may be due to different proportions of
phrases and not to their syntactic order. Robinson (1979)
points out, however, that titi monkeys clearly distinguish
phrase types and concludes that variation in phrase order is
important and has communicatory significance.
Robinson (1984) has also investigated the syntactic
structures in vocalizations of £. olivaceus. He found that
these monkeys use lexical compounding rules to generate new
sequences, where the new compound call is produced in
contexts intermediate between those of the constituent
vocalizations. Additionally, he found a second, more simple
sequence type where two call types are blended to form a
transitional third call. These new calls are without
apparent syntactic structure. He concluded that these
lexical syntactic rules are analogous to the rules we use to
generate words from morphemes, they are not analogous to the
grammatical rules of language.


137
Another perspective on these relationships is provided
by figure 15. Here the relationships between defining
variables and acoustically related calls are seen. This
figure shows that while a variety of calls are similar they
all can be acoustically differentiated by a limited set of
variables. Remember also from chapter 2 that some call
variants formed families by similarity of appearance in
spectrograms. These call families are related by similar
defining variables. Notice that in a call family, where
there were more than two variants, the defining variables
were related associatively. For example, call variant 1210
is conditionally defined by variables STIME and MAXLOC,
while variant 1221 is defined by MAXLOC, MAX, NODOM, and
variant 1222 is defined by NODOM and BWFORM.
Vocal Production Mechanisms
Several pertinent aspects of vocal production can be
mentioned here though it is clear that this subject requires
extensive research in a laboratory. First, through
examination of slow motion pictures of C. olivaceus
producing alarm calls, they keep their mouths open when
vocalizing. Sometimes the lips are pursed, thereby
lengthening the vocal tract. Pursed lips were not however
required for grrah production. Open mouth vocalizing is not
always used in other calls, for example, C. olivaceus often
keep their mouths closed when emitting huhs.


107
frequencies of human vocalizations and second, LPC is an
estimation process whereas the Fourier system actually
transforms the waveform into its frequency components and is
assumed to depict an accurate representation. To answer
this question requires information about the respective
signals.
Adult human speech typically has fundamental
frequencies around 110-140 Hz, with spectral energy going to
15 kHz. Comparative data for £. olivaceus are fundamentals
of 750-1300 Hz with energy to 15 kHz. In both signals most
energy is below 10 kHz and contains both periodic and
aperiodic sounds. The major problem arises when signals
become less periodic, as happens at the end of many C.
olivaceus alarm calls. The following analysis is used to
confirm the accuracy of the LPC analyses.
The LPC parameters used in the following analyses are:
sampling rate= 20 kHz; context= 40; analysis window, n=256;
filter order, m= 23; preemphasis, Pr=35%; number of peaks,
NP=3.
Confirmation of LPC analysis. Confirmation of the
accuracy of LPC analysis will be concordance in the results
of analyses of the same call using LPC and non-LPC
techniques, particularly in the frequency and amplitude of
frequency peaks. We will examine vocalization JM 102 using
oscillograms, analog spectrograms, and both Fourier and LPC
derived spectra, and waterfalls (repeated spectra covering


128
with any other variable except another measure of maximum
amplitude location (LOdBMX).
Another striking negative correlation was between the
position within the first formant of the minimum frequency
(MINLOC) and the amount of time that formant increased
(INTIME). This correlation indicates that where the minimum
frequency was early there was more time spent increasing.
The contrapositive is less surprising; where the minimum was
late there was little time increasing. This situation was
again best seen in call type 2000 where the minimum
frequency was sometimes at the beginning of the call.
The last relationship I will point out is the negative
correlation between dominant formant (MXFReither 1, 2, or
3) and the amount of time the first formant decreases
(DETIME). This correlation suggests that calls that had
lower dominant formants spent more time decreasing than
calls with higher dominant formants and vice versa. In
other words, if the first formant was stressed, it was more
likely there would be more time spent falling than if higher
formants were stressed.
A partial correlation analysis showed that three sets
of variables were all highly correlated: END and MIN, BEG
and MAX, and DETIME, INTIME, and STTIME. For that reason,
in subsequent principal component analyses END, BEG, DETIME,
and STTIME were dropped. Therefore, MIN, MAX, and INTIME
were used in subsequent analyzes.


41
Figure 1. Study area showing adjoining ranches, the
bordering streams, and the trail system. Marks on trails
are at 25 m intervals (Robinson, 1986).


124
Table 13-continued. Descriptive statistics of
acoustic parameters of Cebus alarm calls.
VAR TYPE
X
a
95%
CONF.INT.
F-ratio
DBDIFF
1110
20
1.4
17
23
6.310
2000
17
2.2
14
22
3001
12
1.5
8
16
DBLOC
1110
57.9
4.3
48.8
67.1
5.466
2000
41.7
6.8
30.2
53.2
3001
34.1
5.0
21.5
46.8
INFLECT
1110
13
.76
11
15
5.115
2000
10
1.3
8
12
3001
9
1.3
7
11
INFIST
1110
27.4
4.3
18.1
36.7
1.166
2000
26.0
8.0
14.3
37.7
3001
15.8
2.33
3.0
28.7
INTIME
1110
3.0
.75
.5
5.5
7.298
2000
8.1
1.4
5.0
11.3
3001
10.6
2.8
7.1
14.1
DETIME
1110
30.0
106
26.1
33.5
23.276
2000
13.5
2.1
8.8
18.2
3001
35.2
3.3
30.1
40.4
STTIME
1110
42.1
6.4
31.9
52.3
11.040
2000
4.7
2.0
0
17.5
3001
20.7
6.5
6.7
34.8
NODOM
1110
42.1
6.4
31.8
52.3
11.040
2000
4.7
2.0
0
17.5
3001
20.7
6.5
6.7
34.8
LOdBMX
1110
55.3
4.2
46.9
63.7
6.141
2000
38.3
5.6
27.7
48.8
3001
32.9
5.7
21.3
44.5
MXFR 1110
1.00
0.0
.84
1.2
14.670
2000
1.67
.14
1.5
1.9
3001
1.2
. 13
.99
1.4
SIGN.
LEVEL
. 0043
.0082
. 0108
. 3225
. 0021
. 0000
.0002
. 0002
. 0049
. 0000


CHAPTER 3
VOCAL RESPONSES TO RELEASED SNAKES
Methods and Materials
Data Analysis Methods
The alarm calls recorded during snake releases were
digitized at 20,000 samples/ second using a Digital
Translation DT2821 analog/digital- digital/analog board in a
Compaq Portable II computer using the RDA and LDA routines
in the ILS spectrum analysis system. Where necessary these
calls were digitally filtered using ILS. These signal were
then played into and analyzed on a Kay Elemetrics model
7029A spectrograph. The spectrograms were narrow band
(resolution= 45Hz), 80-8,000 Hz bandwidth, with FL1 shaping.
An essential element to the analysis of these calls was
the identity of the caller. This was determined by
1) identifying an animal as it calls on video,
2) identifying the caller in the field and calling out
its name while recording,
3) determining who was in the area during the
interaction and attributing caller identification
by listening, a process of elimination,
4) where calls sounded the same to me, I visually
identified the group of animals that was present
and calling and then compared the fundamental
frequency for similar calls by those same animals
at other times. This attribution was made by
comparing fundamental frequency of calls.
58


81
Table 8. Responses to Alarm Call Playbacks.
a) Responses to call variants 2000 and 1111
Responses
No
Date
Call
Sp,
, Rec.
Up/Sky Down
Fol.
Spkr.
Res.
Other
ARBOREAL CALLS-
-2000
4/9
HQ125
WH
HA
X
4/9
HQ125
WH
MI
X
4/9
HQ125
WH
STU
X
4/26
M0126
MO
AM
X
4/26
M0126
MO
BA
X
4/26
M0126
MO
GR
X
4/26
M0129
MO
Am
X
4/27
M0129
MO
WH
X
4/27
M0129
MO
BU
X
4/30
M0126
MO
ON, Am
X
4/30
M0126
MO
JE,STU
X
X
5/22
MG113
WH
AM
X
5/22
MG113
WH
AM
X
5/22
MG113
WH
AL
X
5/31
AB16
MI
BA
X
5/31
AB16
MI
BU
X
6/8
M0128
MO
WH
X
6/8
M012 8
MO
STU
X
6/8
M0128
MO
STU
X
6/21
M0128
MO
STU
X
6/21
M0128
MO
STU
X
6/29
AB67
MO
HA
X
6/29
AB67
MO
WB
X
6/29
AB67
MO
MO
X
Totals
0 2
7
4
11
1
Ground Calls-1111
4/16
JM102
HA
MD
X
4/19
BB122
PU
GR
X
4/19
BB122
PU
ML
X
4/19
BB122
PU
ML
X
5/6
AF40
MD
WH
X
5/20
JE124
HA
STU,MN
XX
5/25
JE119
JE
PF
X
5/25
JE119
JE
PF
X
5/25
JE119
JE
STU
X
6/10
JE124
HA
PF, ML
X
X
6/10
JE124
HA
STU
X
6/15
JE124
HA
AM
X
6/15
JE124
HA
WB
X
6/15
JE124
HA
ML
X
6/15
JE124
HA
AM,Am
X
X
Totals 0 8 0 3 5 2


Figure 2. Narrowband spectrograms of grrahs to various threats: a & b) boa,
c) human, d) ocelot, and e) donkey. (Bandwidth 80-8000 Hz, resolution=45 Hz).
Marks at 1 Hz intervals.


139
The human vocal production mechanism is a set of vocal
folds. It is normally assumed that this is also the case for
primates. In humans the spectrum of the glottal waveform
drops (tilts) by approximately 12 dB per octave (Fant,
1973). This 12 dB tilt is used in ILS as an indicator of
voicing. I examined the dB difference between the first
three spectral peaks at the beginning of two qrrahs that had
multiple harmonics, qrrahs 13121 and 2000 (table 16). I
found that the tilt of these calls was highly variable,
with some calls increasing while others diminished
dramatically. These results on source spectral tilt make
any conclusion regarding source mechanisms equivocal.
Table 16.
Amplitude Changes at
Harmonic
Intervals (dB)
F2-F1
F3
-F2
Mean
Range
Mean
Range
Grrah 2000
5.7
-6 18
-9.3
-17 -1
Grrah 13121
-3.0
-10 8
-10.0
-18 -1
Lastly, an examination of the spectrograms of several qrrah
variants, for example, 13121 and 13122 in appendix B, shows
that there are certain spectrogram configurations not found
in human speech. In both of these calls the first energy
band is level, e.g. the pitch of these bands do not change,
but at higher bands there is a dramatic drop in frequency.


50
trees as capuchins without incident. Lastly, I observed
Main group approach a mother and infant tayra in a palm
tree. When the adult tayra saw me it tried to leave by
coming to the ground and running away, however the infant
wouldn't leave and when the mother appeared to abandon it
the C. olivaceus threatened the young tayra. This prompted
the mother to return, which caused the Cebus to back off.
The monkeys gave few arrahs but gave many other threatening
calls during the interaction. J. Robinson (personal comm.)
reported seeing an adult tayra chasing C. olivaceus through
the trees. I conclude that tayra are a potential threat to
C. olivaceus, but the context of the interaction was
important as to whether the monkeys reacted.
The response of animals hearing an alarm call is
typically immediate and marked. For example, I was filming
a subadult male, GR, as he sat near me on a branch. Main
group was spread out around us. A monkey approximately 60m
away saw a tayra and qrrahed. Griffin immediately turned to
look at the caller and began scanning that same distant
area.
Van Schaik and van Noordwijk (1989) described the role
of male Cebus apella and C. albifrons in predator
avoidance. They measured differences in vigilance and
related behaviours between sexes of both species and,
finding a difference, interpreted it as a special male role
in predator avoidance. The same anti-predator behaviors of


161
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86
5) No Response- The animal did not react to the
signal. Did it change its behavior,particularly
where it was looking, immediately upon
presentation of the signal?
6) Other- Were there other responses, such as looking
towards other monkeys, at the camera (me), or at
other animals?
Experimental Limitiations
There were several limitations to this experimental
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a requirement that each event be independent. To produce
independent replicates of these tests I could repeat them
with another group of monkeys. Playing the calls used in
these experiments back to another group of monkeys however
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their own group and responding to the stranger's voice
rather than the message of the signal. Simple repetitions
of the same stimulus only increase sample size. Replicates
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respond at a different rate to alarm calls given by actual


135
3.1
2.1
t*
1.1
0.1
-0.9
-1.9
-2.9
-2.5 -0.5 '1.5 3.5 5.5
Figure 14. Plot of the first two discriminant functions and
how they array the three locational grrah variants. Group
centroids are indicated by +. (1= grrah variant 1111, 2=
grrah variant 2000, and 3= grrah variant 3001).


163
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Figure 5. Digital oscillogram and analog narrowband spectrogram (inset) of JM102, a
grran variant 1111. (spectrogram bandwidth 80-8000 Hz, resolution=45 Hz).
109


27
frequency of the second peak (F,), and increasing frequency
in the second spectral peak. These authors did not measure
the spectral tilt of these calls, but an examination of the
spectra shows that these frequency features are probably the
most parsimonious.
Richman (1976) gave perhaps the most detailed
description of a primate vocal distinctive features,
describing the vocal features used by gelada baboons
(Theropithecus gelada). He presented a number of
spectrograms illustrating a variety of calls with complex
acoustical morphology. He reported that geladas produce
long strings of alternating ingressive and egressive
phonations. Through spectrographic analysis he concluded
that they were capable of producing both vowel-like and
consonant-like calls. He also described place and manner of
articulation for these calls.
Geladas appear to be able to vary the relative
frequency positions of the first two formants, either
through divergence of the formants like human back vs front
vowels or raising or lowering of the first formantthe low
vs high vowel contrast. (I should note that there has been
some disagreement regarding the use of the term formant when
discussing non-human vocalizations. Please see chapter
five, the acoustics section). Geladas can also
simultaneously lower both formant frequenciesrounding, in
acoustic phonetic terms. Lastly, geladas can selectively


THE SEMANTICS OF CEBUS OLIVACEUS ALARM CALLS:
OBJECT DESIGNATION AND ATTRIBUTION
BY
JEFFREY COPELAND NORRIS
A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL
OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT
OF THE REQUIREMENTS FOR THE DEGREE OF
DOCTOR OF PHILOSOPHY
UNIVERSITY OF FLORIDA
1990


116
measured by ILS. A dominant formant was defined in two
ways. First, a formant was dominant if it was more than 6dB
louder than any of the others at the beginning of the call
or, secondly, lacking amplitude differences at the
beginning, the dominant formant was the one with the maximum
dB level at any point in the call. Once the dominant
formant was determined its frequency at the beginning and
end, as well as maximum and minimum were measured.
Additionally, the maximum dB level locations were determined
following the above two techniques. If a formant was >6dB
louder than any other formant, the maximum dB location
(MAXLOC) was the place within that formant of maximum dB, if
no formant was dominant at the beginning it was simply the
maximum dB of any formant (LOdBMX).
Results
Alarm Call Acoustics
Fundamental frequency
Mean fundamental frequencies from 32 alarm calls of
both types are given in table 11. Values from the beginning
and middle segments of each calls are provided. The end
segments of some qrrahs and all waahs were not measured
because they were aperiodic.


138
ACOUSTIC FEATURES
BW NO MAX dB ST
FORM MAX DOM INC LOC LOC DUR TIME
3001
Figure 15. Cladogram of acoustic distinctive features
illustrating relationships between 12 grrah variants.


19
opponent and the likelihood of physical contact in agonistic
interactions. For example, where a young animal is
signalling that it is interacting with a dominant animal the
mother should react more strongly than when her offspring is
signalling it is interacting with a lower ranking animal.
Through analysis of the probability, duration, and latency
of response the authors concluded that the mother responded
differentially to four call classes and that the calls
referred to the opponent. An alternative hypothesis is that
the call refers to the level of fear the animal is feeling
during the encounter. The mother's response to a particular
call may be a result of, first, her recognizing that the
call is from her offspring (through voice recognition) and,
second, that levels of fear elicited by the potential
opponent's likelihood of harming her infant are encoded in
the four calls. For example, one call, one emotion, is
stimulated by large dominant animals where the chances of
physical contact are high while another call is stimulated
by a less dominant animal where contact is unlikely. The
stimulus for the call is the same, but the referent is to
relative levels of fear, not to the object eliciting the
fear.
In a similar study of a series of acoustically similar
vervet monkey grunts, Cheney and Seyfarth (1982) demonstrate
that vervets may differentiate several grunts that are
undifferentiable to humans, and that these grunts may


125
Call 2000- the highest maximum frequency in the
dominant formant, the highest dominant
formant, and least amount of time frequency
decreasing.
The ANOVA showed that all variables except BEG, MINLOC,
INFLECT, and IN1ST were significantly different at p<0.01.
Correlations. An analysis was done to find out
which acoustic variables correlated, in particular, whether
certain frequency characteristics of the calls were
correlated with formant structure and stressing. This
analysis was motivated, in part, by a desire to find out
whether these acoustic variables explained the acoustic
relationships that I heard, for example, whether the
differences I heard between calls 1111 and 2000 were found
in correlations between particular variables.
Results of this analysis are in table 14. There were
96 significant (p<0.01) correlations (including
duplications), with only a single variable, INFIST (location
of first inflection), having no correlations. Minimum
frequency had the most, correlating with the 12 other
variables.
There was no correlation between call duration and any
frequency aspect of the call other than amount of time
decreasing and remaining steady. There was a significant
(p<0.01) positive correlation between duration and amount of
amplitude difference (DBDIFF).


LIST OF FIGURES
pages
FIGURES
1 STUDY AREA 41
2 NARROWBAND SPECTROGRAMS OF GRRAHS TO
VARIOUS THREATS 53
3 NARROWBAND SPECTROGRAMS OF WAAHS 55
4 KEY TO GRRAH VARIANTS 66
5 DIGITAL OSCILLOGRAM AND ANALOG NARROWBAND
SPECTROGRAM OF JM102 109
6 WATERFALL DISPLAY OF JM102, GRRAH VARIANT
1111. (FOURIER DERIVED) 110
7 WATERFALL DISPLAY OF JM102, GRRAH VARIANT
1111. (LPC DERIVED) Ill
8 COMPARISON OF FOURIER AND LPC DERIVED
SPECTRA OF MIDDLE SEGEMENT OF JM102 112
9 OSCILLOGRAM OF 20 MSEC OF GRRAH
VARIANT 13121 118
10 A DFT SPECTRA OF A GRRAH VARIANT 13121 119
11 A CEPSTRUM WAVEFORM FOR A GRRAH
VARIANT 13121 120
12 PLOT OF THE FIRST TWO PRINCIPAL COMPONENTS.. 131
13 CLASSIFICATION OF 59 GRRAHS 133
14 PLOT OF THE FIRST TWO DISCRIMINANT FUNCTIONS 135
15 CLADOGRAM OF ACOUSTIC DISTINCTIVE FEATURES.. 138
16 COMPARISON OF FOURIER AND LPC DERIVED
SPECTRA OF GR69, A GRRAH VARIANT 3001.... 146
vi


136
What variables describe the twelve call variants?
Table 15 illustrates how 8 variables completely define 59
calls in 12 variants. In every case no more than 3
variables are needed to define a call.
Table 15
Defining
Variables
for
Grrahs
BW
NO
MAX
dB
ST
FORM
MAX DOM
INC
LOC
DUR
LOC
TIME
Location
Calls
1111
+
+
2000
+
+
3001
+
Other
Calls
1130
+
+ +
1210
+
+
1221
+ +
+
1222
+
+
1311
+ +
+
1312
+
+
1321
+
+
13121
+
+
13122
+ +
;
Variable
Limits
750Hz
< BWFORM NODOM
<
5%
MAXLOC <
10%
MAX <
1600Hz 50Hz
<
INC
DUR
< .062
sec.
60% <
dBLOC STTIME <
60%
Among the locational calls:
Call 1111- bandwidth of the dominant formant > 750Hz
and maximum frequency < 1600Hz
Call 3001- duration < .062 seconds,
Call 2000- increase in the dominant formant > 50Hz and
location of no dominant formant at < 5% of
the call.


APPENDIX C
DEFINITIONS OF ACOUSTIC VARIABLES
Variable
Definition and (units)
DUR
Duration of the call, (sec)
BEG
Frequency at the beginning of the dominant
formant. (Hz)
END
Frequency at the end of the dominant formant. (Hz)
BWFREQ
The difference between the frequency at the
beginning and end of the call: END-BEG. (Hz)
MAX
The maximum frequency of the dominant formant.(Hz)
MAXLOC
The location of maximum frequency in the dominant
formant, as a percentage of total duration.
MIN
The minimum frequency in the dominant formant.(Hz)
MINLOC
The location of minimum frequency in the dominant
formant, as a percentage of the total duration.
BWFORM
The difference between the minimum and maximum
frequency of the dominant formant:MAX-MIN. (Hz)
INC
The amount of frequency increase from the
beginning of the call to the maximum
frequency:MAX-BEG. (Hz)
DBDIFF
The difference between dB at the beginning and
maximum dB in the dominant formant. (dB)
DBLOC
The location of the maximum dB in the dominant
formant, as a percentage of the total duration.
INFLECT
The number of times that the type of frequency
change changes,i.e. increasing, decreasing,
steady.
INTIME
The amount of time, as percentage of total, where
frequency increases in first formant.
178


18
He has not demonstrated the meaning of the signal. A cue to
the meaning of the signal is the receiver's immediate
response upon hearing the signal. Does it look to the
signaller or look for the food? The author states, in his
description of response to the food call, that "upon hearing
a food call, an individual would immediately stop its
activity, glance alertly in the direction of the source of
the call and run there to feed" (Dittus, 1984, pp 473).
Later he states that "upon hearing a food call, animals
rapidly and directly approach the call site and feed there"
(Dittus, 1984, pp. 476). This seems to imply that signal
receivers may be turning to the signaller to find out what
stimulated the call rather than turning to the food itself,
i.e. that the food's presence is signalled indexically. A
more rigorous test for reference would be to determine the
receiver's response when the signaller is remote from the
abundant food.
The majority of primate vocalizations are, as Cheney
and Seyfarth (1982) point out, chirps, trills, grunts, and
screams typically found in social situations. A variety of
studies have attempted to determine the reference and
therefore the meaning for several of these calls.
Gouzoules, Gouzoules, and Marler (1984) describe five
screams given by rhesus macaques during social interactions.
The authors assert that these screams were representational
signals referring to the social rank of the screamer's


114
Descriptive Variables
Twenty variables were used to describe the acoustics of
the alarm calls. They are defined in appendix C. These
variables were used to describe the multiple factors that
might determine how a call is perceived. I attempted to
model signal changes with how the animal may perceive them.
For ease of comparison, several variables duplicated as
closely as possible variables used by other animal
communication investigators.
Two important criteria were used in determining values
for these variables: perceptible frequency change and
formant dominance. As pitch changes during a call it is
important to know if the change is perceptible. For example,
if a pure tone centering at 1000 Hz drops by 15Hz or 19Hz
will the change be perceptible? Frequency cues play a
central role, for example, in differentiating vowels and are
discriminated in the same manner by not only humans but a
variety of other animals. While there have been few studies
of Cebus hearing (D'Amato and Salmon, 1982; 1984)
fortunately the various studies on primate psychoacoustics
(see above) may help determine if a frequency change is
perceptible.
I adapted the difference limens (DL) value reported by
Sinnott et al. (1985) and translated it into a Weber
fraction in order to judge whether frequency changes in the
Cebus calls were perceptible. Weber fractions represent a


62
JE101 is his first arrah. He then gives a long series
of huhs interspersed with a single qrrah after a
minute.
By the second minute other animals begin arriving
and while other animals in the distance continue giving
huhs an arriving animal grrahs. JE 118. By then there
are three agitated monkeys over the snake, giving few
grrahs. At 2:02 minutes into the interaction, Jefe
gives qrrah JE119, the first ground snake qrrah variant
(for a discussion of qrrah variants, see below). For
the next two minutes a juvenile female, Hanna, sits
over the snake watching it without giving alarm calls.
Jefe then gives another qrrah. JE117, which appears to
prompt Hanna to intently look around, otherwise there
is little action. Hanna returns to lying on a branch
over the snake content in watching it. Other animals
continue to huh in the background. Overall the troop is
relatively placid. Eventually Hanna and another small
female moved.
Four minutes into the interaction Jefe gives
another ground snake qrrah. JE121. Then, after another
lag in action, Hanna gives another type of ground snake
call, HA102, this one a loud brief call like a bark
which she repeats 45 seconds later, HA 105. By now the
snake, which was released on the ground, movs into
water and neither the monkeys nor I can find it. Hanna
gives qrrah HA122 as she moves lower in a bush,
intently trying to locate it.
A minute and a half later the snake begins to swim
away, prompting Hanna to give qrrah HA 107. Other
monkeys move in and give several desultory grrahs (that
are too poor for analysis). Hanna then gives three
calls, a loud qrrah. HA 108, followed immediately by HA
109, then another call like the loud call, HA 111.
Next she gives four ground snake calls, JE 123, HA112,
and JE 124 (one is masked by bird calls so it isn't
analyzed), like Jefe's earlier ground snake call,
JE119. The interaction is now 10 minutes long. The
middle call, HA 112, prompts another monkey to intently
look down. The boa then begins to climb into a vine.
Hanna gives qrrah HA125 followed by two similar calls,
JE126 and HA115. Notice the wide diversity in call
types used by Hanna in these last 14 calls.
Over the next 2 minutes there are few calls, yet
Babas, a juvenile male, agitatedly brakes a branch near
the snake. They do not appear to know where the snake
is. The juvenile monkeys left after two minutes
without more calling.
Three minutes after the last qrrah Mo approaches,
stopping frequently to look down to the ground where
the boa had been before it moved into the vines. She
is quite evidently looking for something. For two


48
it. Other animals waahed and she immediately swung
beneath the branch to interpose the branch between the
falcon and herself. It was not clear whether the bird
was attacking the female or simply flying away from the
harassing individuals.
This was the only observed active defense, other than
vocalizations, against avian predators. When the physical
threat was nonavian the situation was quite different.
Mobbing behavior, rarely directed towards raptors, was
regularly used against non-avian threats. C. olivaceus have
been observed mobbing jaguar, puma, ocelot, tayra, boa
constrictors, and humans. Additionally, they qrrahed at
donkeys, cows, and deer, though this appeared to happen only
when the animal ran through the forest, presumably
frightening the monkeys.
In a typical interaction, multiple animals cluster
near the potential predator and give many grrahs. At least
in the case of boas, which often remain still and therefore
cryptic around monkeys, the animal discovering the snake
often remains near it through much of the interaction,
giving the most grrahs while other troop members move by
also emitting alarm calls. Under these conditions when a
predator is mobbed there may be hundreds of grrahs given.
The emission of grrahs is a highly directed behavior.
Typically the alarming animal faces the threat and gives
numerous calls in a clearly agitated manner. These may or
may not elicit calls from other monkeys. Grrahs appear to
function most clearly as an alarm, warning other animals of


152
factor that defined a variety of japanese macaque coos, e.g.
smooth early high vs smooth late high calls (Green, 1975).
Rate of frequency change was only examined in £.
olivaceus tonal calls (Robinson, 1984).
The relative location where all formants had the same
amplitude (psychoacoustically affecting the pitch of the
call) was critical in differentiating six C. olivaceus
arrahs. Likewise, the total number of formants affects call
pitch and was important in discriminating C. olivaceus
chirps from trills and whistles.
Overall, the important frequency cues are determined by
the characteristics of the call. Noisy, atonal calls
(typically short duration) characteristically use peak
frequency and its change as critical factors. More tonal
calls also use peak frequency and its change as well as
bandwidth and the relative location of the peak frequency.
Duration cues
Duration was an infrequently used distinctive feature,
only used to distinguish a single qrrah and two trills. For
any cue to be a factor its variance must be low within a set
of calls. In this case, for duration to be a sensitive cue
between vocal types the range within a call would have to be
slight. Clearly this limits the duration over which a call
may range. This limitation may restrict the variability
that an animal wants to use in order to communicate other
aspects of the message of the call, for example, emphasis or


153
other measures of vocal affect. Therefore, it is not
surprising that duration by itelf is not an important cue.
The relative duration of a particular frequency change
or frequency constancy, however, was a cue in both C.
olivaceus call types and the similar Saauinus chirps. Among
C. olivaceus. the duration of constant frequency, as a
percent of total duration, differentiated some qrrahs while
the duration of high frequency was important in
differentiating certain £. olivaceus trills from whistles.
Lastly, the downsweep duration was critical in
differentiating Saauinus chirp variants.
The factor Interrupt was used to distinguish a single
discontinous marmoset trill. Interrupt could easily be
stated as a duration variable, but since only a single call
was discontinous, the discrete variable interrupt vs not
interrupted suited the authors needs.
Overall, relative duration of certain freqeuncy changes
provided several monkeys with means to possibly discriminate
their calls.
Amplitude cues
There is no clear pattern of amplitude factors being
distinctive features. Absolute dB is difficult to measure
in the field and dB values relative to signal background
amplitude were not measured in most studies. Spectral tilt
was measured in vervet alarm calls and estimated in C.
olivaceus qrrahs. Among vervet alarm calls tilt was the


60
snake in front of it. Usually the snake began moving away,
often into trees. I pulled the snake down to the ground if
it began to move beyond my reach. Typically, the monkeys
immediately began calling and mobbing the snake when it was
released. Sometimes many other animals approached while at
other times only the animals that first saw the snake
called. On a few occasions the monkeys did not apparently
see the snake and left without vocalizing. On still other
occasions they obviously saw the snake but did not
immediately begin to call.
There were eight boa releases in the following order,
listed by snake length: medium, large, medium, large and
medium, small, large and medium, large and medium (to a lone
monkey), medium and small. Additionally, a second snake
species, Drvmarchon coris. was used in a single release.
The number of qrrahs recorded for the nine snake releases
ranged from 9-58.
Snake Descriptions
Two species of snakes were used: boas (Constrictor
constrictor) and Drvmarchon coris. a yellow phase indigo
snake. The D. coris, a snake very similar to the indigo
snake, was used to determine if C. olivaceus used alarm
calls only at boas or whether they used a general snake
call. I used four boas of three sizes (table 4) in
combinations of small, medium, large, small and medium,
medium and large, and medium and medium.


61
Table 4.
Snakes Used
in Controlled
Releases
Species
Length (m)
Weight (kg)
Boa
1.95
4.65
Boa
1.60
2.00
Boa
1.58
1.94
Boa
0.68
0.20
D.coris
1.80
2.50
Results
Responses by Monkeys to Snakes
I will first describe the release of a 0.68 in boa and
provide examples of the qrrahs recorded in response to it.
The transcription below of my commentary on the video
recording shows that while the calls are diverse, several
animals give quite similar calls, and that some calls are
given in only certain contexts. The commentary also provides
a background for understanding the resulting investigations.
The vocalizations referred to in the text are presented as
spectrograms in appendix A. There were 87 calls during the
release, of which 29 were qrrah variants. It is clear from
the spectrograms that these qrrahs are highly diverse, with
many configurations. Five animals called during the 24
minutes interaction.
Snake release commentary
I release the small, 0.68m boa in front of Jefe as
he is sitting alone 4m above me eating a butterfly
larva. For the first 1^ minutes he continues eating
without calling even though he appears to see the
snake. Once he finishes eating he begins calling,


74
monkeys are combining elements of different calls to form a
new signal, with a new meaning. This would be analogous to
word formation in human language. For these calls to be
graded ^continuous) requires that the new form follow a
continuum where the meaning of the signal is intermediate
between the meaning of the two calls from which it is
formed. This raises questions that are essentially
epistimological, for if these alarm calls are referring to
objects, what could be the intermediate meaning, e.g. what
is the intermediate meaning for the word snake? For this
reason, I doubt whether the calls are graded. Notice
however that the physical evidence for both positions,
phonological syntax or graded signal, is the samecalls
whose spectrographic configuration is intermediate between
two other calls. Examining appendix B shows that several
variants appear intermediate in form, for example, variant
1130 seems intermediate between variants 1111 and 1210.
Even if they were intermediate in form, I have no data to
indicate whether they were used in contexts that were in
some way intermediate between the contexts in which the
other calls were used, that is, that they had an
intermediate meaning.
Grrahs to Other Snakes
In the single release in which another snake species
was released, two monkeys gave three different call types,
including one call, AD11, of grrah variant 3001, the ground



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66
1..0.0.0 Formants only drop
1.1..0.0 Formant level then drops
1.1.1..0 Freq. drop cont. and extensive
1.1.1.1 One formant or F1 stressed 27
1.1.1.2 F2 stressed 7
1.1.2..0 Freq. drop discont.and extensive 32
1.2..0.0 Formants drop continuously
1.2.1..0 One formant 17
1.2.2..0 Multiple formants
1.2.2.1. F, stressed 22
1.2.2.2. F2 stressed 8
1.3..0.0 Formant drop not extensive
1.3.1..0 F, stressed
1.3.1.1. Tail stressed below 36
1.3.1.2. Tail stressed above 17
1.3.1.2.1 F2 extensive drop 34
1.3.1.2.2 Fother extensive drop 6
1.3.2..0 F2 stressed, tail stressed below 27
2..0.0.0 Formants rise and fall 13
2.1..0.0 Formants rise and fall discontinuously 3
3..0.0.0 Others 18
3.0.0.1 Short duration 10
Total 277
Figure 4. Key to grrah variants by spectrographic
configuration with totals for each variant.


Table 14. Correlation matrix of acoustic variables
D
B
E
B
M
M
M
M
B
I
D
D
I
I
I
D
S
N
L
M
U
E
N
W
A
A
I
I
W
N
B
B
N
N
N
E
T
0
0
X
R
G
D
F
X
X
N
N
F
C
D
L
F
F
T
T
T
D
d
F
R
L
L
0
I
0
L
1
I
I
I
0
B
R
E
O
O
R
F
C
E
S
M
M
M
M
M
Q
C
C
M
F
C
T
E
E
E
X
T
DUR
\
#
#
- *
*
BEG
\
*
*
*

END
*
\
-*
*
*

#
#
-*
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*
BWFREQ
-*
\
-#
-*
*
*
-#
-*
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*
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*
*
\
#
*
*
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MAXLOC
*
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#
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*
#
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*
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*
*
-*
*

\
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#
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#
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*
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*
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BWFORM
*
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\

INC
#
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*
#
#
\
*
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#
\
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\
*
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#
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\
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#
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*
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*
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NO DOM
-#

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\
*= pcO.OOl #= p<0.01 -= negative correlation
126


89
duration and percentage of correct responses (r=0.1521,
Kendall rank correlation). Since the signal amplitudes were
equilibrated there is no influence of amplitude differences
on responses.
As for response rates to the various replicates,
within-call variant mean response rates for playbacks (for
calls used more than once) are provided in table 10.
Table 10.
Response Rates for Calls and Individuals
Calls
Individuals
Mean Range
Mean
Range
Grrah 1111
39% 33-50%
40%
33-55%
Grrah 3001
24% 0-50%
26%
17-33%
Grrah 2000
38% 0-67%
39%
33-50%
Waahs
66% 40-100%
66%
40-100%
Variation by caller. I will first examine response
variations relative to the context of the caller. There
were insufficient data to statistically compare differential
responses by sex, familial relationship, or social dominance
of the caller.
There were only five playbacks of calls from males,
which is too small for further analysis, whereas there were


145
are produced (this awaits laboratory investigation) it is
clear that capuchin alarm calls are far more tonal and all
involve a periodically vibrating mechanism. There are two
caveats; most arrahs contained an end segment that was
aperiodic and some calls contained spectral configurations
indicating a source function other than vibrating vocal
folds (or their analogous structure).
Filter Function
The spectral configurations of most of the calls of C.
olivaceus and Cercopithecus aethiops are very different,
though some of the calls have some qualities in common. A 5
kHz bandwidth spectra and LPC-derived smoothed spectra of a
vervet eagle call can be seen in Owren, 1985, (fig.2, pg
59). Figure 16 is a similar illustration of a C. olivaceus
qrrah. a 10 kHz bandwidth and LPC-derived smoothed spectra
of qrrah variant 3001, GR69 by Pointy Face. The differences
in bandwidth reflect differences in digital sampling rates.
Based on the LPC spectral envelop, both calls contain two
spectral peaks, both centered around 1 kHz. The second peak
though is rather different, with the C. olivaceus call
having a peak at the second harmonic. This points out an
important distinction between the two primates calls,
probably reflecting morphological differences in the
resonance chamber in the two species. When the C. olivaceus
calls have multiple energy bands they typically are
harmonics of the fundamental. Given the much lower


Abstract of Dissertation Presented to the Graduate School of
the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Doctor of Philosophy
THE SEMANTICS OF CEBUS OLIVACEUS ALARM CALLS:
OBJECT DESIGNATION AND ATTRIBUTION
By
Jeffrey Copeland Norris
December, 1990
Chairman: Dr. John F. Eisenberg
Major Department: Forest Resources and Conservation
(Wildlife and Range Sciences)
Wedge-capped capuchin monkeys, Cebus olivaceus. give
acoustically distinct alarm calls to different predators.
The semantics of these alarm calls were studied in central
Venezuela in three stages: 1) through recordings of Cebus
olivaceus vocal response to various predators, 2) through
release of boa constrictors of three sizes (small, medium,
and large) and two quantities (one and two), and 3) through
playback of resulting calls. Alarm calls to released snakes
were categorized by acoustic features into 15 variants, 2 of
which were used solely when a snake was on the ground, and a
third when a snake was in a tree. The playback of the
locational calls showed that upon hearing a call a capuchin
looked in a particular direction, into trees or toward the
ground, at the appropriate call. Capuchins use alarm calls
vii


150
Table 18. Distinctive features for primate vocalizations.
Cebus
Vervets
Marm.
Tam.
Alarm
Tonal
Alarm Grunt
Trill
Chirp
Cues
Freo.
BW
X
X
X
X
Peak
X
X X
X
Inc
X
X
X
PkLoc
X
X
Rate
Nodom
X
X
NBND
X
Dur.
Dur.
X
X
aF
X
X
X
Interrupt
X
i
dB
X
dBloc
Tilt
X
X
Other
Q
X
Definitions: BW=bandwidth; Peak=peak frequency;
Inc=increasing frequency; Rate=BW/Dur; PkLoc= location
of Peak; Nodom=location where all formant dB values
equal; NBND= number of formant bands; Dur= Duration;
*F=dur of a frequency change; interrupt= interruptions
in spectrograms; dB=dB at Peak; dBloc= location of
maximum dB; Tilt= slope of adjacent spectrum peaks,
dB/octave; Q=for a spectral peak, peak to bandwidth
ratio;


151
frequency was an important distinctive feature among all
species except Cebuella trills. This factor largely
determines the pitch of the call and may be the most
important frequency characteristic in a very brief call,
where changes in frequency are necessarily brief and masked
by the general noisy quality of the call. Peak frequency is
probably not an important factor in defining the Cebuella
trills because the frequency modulation of the call results
in a wide spectral peak. Therefore, bandwidth should be the
sensitive determining factor, as was the case. Likewise,
increasing frequency differentiated calls in all but
Cebuella trills. One C. olivaceus qrrah (variant 2000) was
very similar to a single Saquinus chirp (chirp variant B,
Snowdon, 1982, pg 219) when spectrograms are compared.
Increasing frequency would be a sensitive cue to
differentiate these calls from the other chirps and grrahs.
In fact, many £. olivaceus grrahs were similar to the
tamarin chirps, which may explain why these two calls shared
the most distinctive features.
Peak frequency location, where in the call the peak
frequency occurred, was the only variable used solely by the
two sets of C. olivaceus calls. One would not expect this
to be an important factor for the noisy, broadband vervet
call or the FM trills. It is surprising however that it is
not a factor in tamarin calls. Peak location is the single


Hal09
Ha
Hal05 Hal22
Ha Ha
Halil
Ha
t
Hal23
Ha
Hall2
Ha
Hal24
Ha


Figure 7. Waterfall display of JM102, grrah variant 1111. This is a LPC derived
display, unlike figure 6, which was Fourier derived. The individual spectra marked
by an asterisk can be seen overlaying an FFT spectra in figure 8.
Ill


179
DETIME
The amount of time, as percentage of total, where
frequency decreases in first formant.
STTIME
The amount of time, as percentage of total, where
there is no frequency change.
NODOM
The position, as a percentage of total duration,
where maximum dB in the dominant formant is within
6dB of maximum amplitude in any other formant.
LOdBMX
If there is no dominant formant, the position, as
a percentage of total duration, where any formant
has a maximum dB larger than any other formant's
maximum.
MXFR
The formant number (1, 2, or 3) of the dominant
formant.


14
Table 1. Animals that Alarm Call.
Species
Reference
Birds
chukar partridge
red-legged partridge
California quail
turkey
quinea fowl
domestic chicken
Rodents
black-tailed prairie dog
Richardson's ground squirrel
thirteen-lined
ground squirrel
Belding's ground squirrel
California ground squirrel
Owings and Leger, 980
arctic ground squirrel
Primates
saddle-back tamarins
black spider monkey
black-handed spider monkey
wedge-capped capuchin
vervet monkey
Japanese macaques
rhesus macaques
chimpanzees
Stokes, 1961
Goodwin, 1953
Williams, 1969
Hale et al, 1969
Maier, 1982
Collias and Joos, 1953
Konishi, 1963
Gyger et al, 1986
Hoogland,1983
Davis, 1984
Schwagmeyer, 1980
S.R. Robinson, 1980, 1981
Owings and Virginia,1978
Leger and Owings, 1978
Melchior, 1971
Bartecki and Heymann,1987
Eisenberg, 1976
Chapman et al, 1990
Oppenheimer and
Oppenheimer, 1973
Robinson, pers.comm.
Norris, pers.obs.
Struhsaker, 1967
Seyfarth et al, 1980
Green, 1975
Chapais and Schulman,1980
Marler and Tenaza, 1977


119
Figure 10. A discrete Fourier transform (DFT) spectra of a
grrah variant 13121. The main frequency peak is at 1187 Hz.


144
prominent horizontal energy bands of Q. olivaceus calls.
The oscillograms of each vervet alarm call also appear to be
considerably less periodic than those for C. olivaceus
(Owren, 1985, figure 4, pg 67). Owren and Bernacki (1988)
describe subtle yet consistent differences in source
periodicity between eagle and snake alarms. Analysis of
these two alarm calls showed that 18% of eagle calls
contained periodic elements compared to only 7% for snake
alarms. All of the C. olivaceus alarm calls contained at
least some periodicity.
A good indicator of source function is the spectral
tilt of the vocalization. As stated above, in many human
vowels, our most periodic vocalization, the spectral tilt is
12 dB (Fant, 1973). Owren (1988) found that the
Cercopithecus aethiops eagle alarm call had a 4 dB/octave
spectral tilt, while the snake alarm call was flat. The
results for C. olivaceus was mixed, with no clear pattern of
spectral tilt.
Based on the results of analyses of vervet alarm calls,
Owren (1985) concluded that the source for eagle calls was a
semi-periodic vibration mechanism which is different from
that used to produce snake alarm calls. Given that vervet
grunts were also brief and noisy, though somewhat periodic
(like eagle alarm calls) (Seyfarth and Cheney, 1984) it is
likely that they were produced in a similar fashion. While
I cannot state with assurance how C. olivaceus alarm calls


46
C. olivaceus appear to be predators on bird eggs and
hatchlings during the nesting season. Such behaviors
typically elicit active nest guarding. I observed Main
group attack, kill, and eat a fledgling yellow-knobbed
curassow (Crax daubentoni) and attempt to attack a nestling
giant potoo (Nvctibius grandis), which was actively defended
by an adult. Pairs of road-side hawks called and swooped at
C. olivaceus most often during the nesting season. Active
nest defense may be the reason that capuchins gave alarm
calls at these relatively small raptors. Responses to
attacks by nest guarding hawks presumably do not markedly
differ from predation events.
Responses of Cebus olivaceus to Predators
Anti-Predator Behavior
Responses to predators, both vocal and otherwise,
varied according to the predator and the context of the
interaction. The description of qrrah usage by Oppenheimer
and Oppenheimer (1973) is different from my experience in
several ways. First, I consider grrahs to be specifically
alarm calls, not simply agonistic calls. Secondly, the
Oppenheimers differentiated only a single alarm call,
apparently lumping all alarm calls as grrahs. Lastly, I
rarely saw capuchins take more vigorous anti-predator
actions than just giving alarm calls. Also, in many
interactions between C. olivaceus and howler monkeys


33
perceive a weakening in his position that animals are
categorically incapable of a language; instead, an animal
'language' will be the product of that species' own innate
language faculty. All animals convert experience into a
system of knowledge, some perhaps using vocal behavior to
differentiate objects. Without begging the definition of
language, if we accept that some species now use 'words' to
identify objects in their environment, the separation
between human language and animal communication becomes
smaller, though still very real. The difference between our
language and the communication systems used by animals may
be a difference in degree, not kind.
Alarm call syntax
If certain alarm calls are used to indicate
attributions about an object, for example, the location of a
snake, they are used in some sense like adjectives. One
would expect then that these calls might be associated with
other calls. For example, if one vocalization means 'snake'
and another means 'ground snake' there must be some
additional segment to the latter signal carrying the
additional information, either as another call or as an
infix in the original signal. In English we modify the
meaning of a word both ways, either by using another word,
an adjective, or by internally modifying the original word
with prefixes or suffixes, e.g. usual and unusual.


REFERENCES
Altxnann, S.A. (1965). Sociobiology of rhesus monkeys.
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ed. S.A. Altmann, Chicago, Univ. Chicago Press.
Bartecki, U., & Heymann, E.W. (1987). Field observation of
snake-mobbing in a group of saddle-back tamarins,
Saouinus fuscicollis niorifrons.
Folia Primatologica, 48, 199-202.
Chapis, B., & Schulman, S.R. (1980). Alarm responses to
raptors by rhesus monkeys at Cayo Santiago.
J. Mamm., 61(4),739-741.
Chapman, C.A. (1986). Boa constrictor predation and group
response in white-faced cebus monkeys.
Biotropica 18, 171-172.
Chapman, C.A., Chapman, L.J., & Lefebvre, L. (1990). Spider
monkey alarm calls: honest advertising or warning kin?
Anim. Behav., 197-198.
Cheney, D.L., & Seyfarth, R.L. (1981). Selective forces
affecting the predator alarm calls of vervet monkeys.
Behaviour 76, 25-61.
Cheney, D.L., & Seyfarth, R.L. (1982). How vervet monkeys
perceive their grunts: field playback experiments.
Anim. Behav. 30, 739-751.
Cheney, D.L., & Seyfarth, R.L. (1985). Social and non
social knowledge in vervet monkeys.
Phil. Trans. R. Soc. Lond. B 308, 187-201.
Cheney, D.L., & Wrangham, R.W. (1987). Predation. In
Primate Societies, ed. B.B. Smuts, D.L. Cheney, R.M.
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Cherry, C. (1978). On Human Communication, 3rd Ed.,
Cambridge, Mass., MIT Press.
160


70
Calls bv Age/ Sex Class
No call given by more than one caller was given by only
a single age/ sex class. There was, however, a significant
skew to overall age/ sex class of all callers: 175/ 190
calls were given by adult females, subadult males and
juvenile females. The remaining 15/ 190 calls by identified
callers were given by adult males (2 calls), juvenile males
(9 calls), and infants (4 calls).
There was no significant correlation (Kendall r= -.40,
p=0.33) between dominance rank of adult females (the only
age class for which sufficient data were available) and
number of grrahs given to released snakes. Dominance was
defined operationally by social grooming patterns (T.
O'Brien, pers. comm.).
Grrah Diversity
There was no clear pattern to the diversity of grrah
usage. Females used a more diverse grrah vocabulary than
males, with the Shannon diversity H' females=l. 13 whereas
H1 maies= 91 (table 6). Overall diversity was H' = 1.12. Notice
that females used all variant types whereas males did not
use five variants, including one indicating snake location,
variant 3001. Notice also that males gave many fewer alarm
calls than females, 41 vs 149. Two males, Stu and Griffin,
produced almost half of the male data set. These two males
are subadults, both born outside the group and approximately
second and fourth in dominance among


4
objects. I argue below that primates may not only be able to
designate objects but also attribute qualities to those
objects. Now we examine the previous studies of the wedge-
capped capuchin to see how it fits into this framework.
Previous Studies of Cebus olivaceus Communication
Oppenheimer and Oppenheimer (1973) describe, in a brief
study, eleven different calls from C. olivaceus living at
Hato Masaquaral, Venezuela, the site of this study. They
characterize one call, the arrah as being an inter-specific
agonistic call. Of the 60 grrahs they recorded, 48 were
directed at humans, 4 to overflying birds, and 2 at howler
monkeys (Allouatta seniculus). The remaining 6 were used in
unknown circumstances. If grrahs were directed at animals
other than humans, they were often repeated and at variable
intervals. If the call was directed at birds, the authors
describe the monkeys as giving the call once then "dropping
to lower branches and moving deeper inside the tree"
(Oppenheimer and Oppenheimer, 1973, pg 422). They also note
that grrahs were associated with other grrahs 92% of the
time.
Robinson (1982) described how three callshuhs. hehs.
and arrawhs, mediate spacing within a group. Arrawhs were
given in two contexts. Loud arrawhs were given by an animal
if it became separated from the group. Group members
responded with more huhs and often replied with arrawhs.


56
from White group. These calls typically have multiple
energy bands, which like arrahs. fall in the later half of
the call. Notice however that the higher energy bands in
some calls may be stressed.
Waahs to vultures. Figure 3b presents two waahs by
the dominant male in Red group, Finger, to a vulture,
probably a turkey vulture fCathartes aura). Again these are
relatively long duration calls that fall in pitch through
the majority of the call.
Waahs to caracara. Figure 3c is a waah to a crested
caracara (Caracara cheriway) by the adult male Brow from
White group. This call is shorter in duration, with the
second formant stressed.
Further Investigations
The anti-predator behavior of C. olivaceus suggests
that, like the vervet (Cercopithecus aethiops), the alarm
calls described above are used in a semantic manner, with
the call referring to an object, in this case a predator,
and not simply to an emotion. Demonstrating that C.
olivaceus use these calls in a semantic manner requires
investigating how a monkey responds to an alarm call. I
therefore undertook a series of experiments to verify that
C. olivaceus use their alarm calls to refer to objects. I
first obtained a series of recordings of known animals
calling at a predator and then played those calls to other


21
communication. The authors of the studies on food calls and
social vocalizations have not, I believe, sufficiently
established their claims of symbolic communication.
Alarm calls: models for semantic communication
There could be no better subject than alarm calls in a
study of semantics. First, if an alarm call is semantic it
will refer to an object, a predator, that is remote in space
from the animal. This is essential because when a monkey is
responding to, for example, a boa, the snake is probably
distant from the monkey and not within the group. Testing
alarm call semanticity becomes a geometric consideration.
The physical remoteness of predator, caller, and receiver
makes response quantification relatively easy because it is
clear when the monkey is looking at the predator and when it
is looking to other monkeys (or speakers). Other semantic
studies of primate calls, for example those on food calls,
have had the problem of differentiating when a monkey was
looking at another monkey and when it was looking at the
food the monkey was eating. (This ambiguity would also
occur when testing for response to an alarm call referring
to a snake close to the caller.) Secondly, by the nature of
predation, response to a predator alarm call must be
immediate and adaptive. When a predator is sighted there
typically is an immediate need to give an alarm call.
Likewise when a monkey hears an alarm call there is an
immediate need for response. This need for immediate


164
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vervet monkeys (Cercopithecus aethiops): perspectives
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M.I.T. Press.


44
(Constrictor constrictor) are the only confirmed predators
of £. olivaceus (Rettig, 1978; Chapman, 1986). Boas are
common on the ranch, but this eagle species does not occur
there. Aside from human predation through hunting which is
negligible at the ranch, presumed predators are aerial,
terrestrial, or arboreal.
The largest raptor found at Hato Masaguaral is the
ornate hawk-eagle (Spizaetus ornatus). This bird is almost
certainly capable of taking Cebus as prey. Other potential
avian predators are the spectacled owl (Pulsatrix
perspicillata), great horned owl (Bubo virginianus), savanna
hawk (Heterospizias meridionalis), zone-tailed hawk (Buteo
albonotatus), black collared hawk (Busarellus niqricollis),
collared forest falcon (Micrastur semitorouatus), laughing
falcon (Herpetotheres cachinnans), and the road-side hawk
(Buteo magnirostris) .
The only strictly terrestrial predator likely to take
C. olivaceus at Hato Masaguaral is the dog, Canis
familiaris. All other potential predators are capable of
following C. olivaceus into the trees. There are four
felids at the ranch; jaguar (Felis onca). cougar (Puma
concolor), ocelot (F. pardalis), and jaguarundi (F.
yaqouaroundi). Tayra (Eira barbara) have been seen chasing
capuchins (Robinson, pers.comm.). In my experience, the
relationship between tayra and C. olivaceus is apparently
context sensitive; among the same individual animals during