Z. Tierpsychol., 38,304-314 (1975) @ 1975 Verlag Paul Parey, Berlin und Hamburg TSSN 0044-3573 / ASTM-Coden: ZETlAG

The Rockefeller University, New York, N. I’.

Dialects in Japanese Monkeys: Vocal Learning and Cultural Transmission

of Locale-specific Vocal Behavior?

By STEVEN GREEN

With one figure

Received: 2 . J. 1974

Abstract

Differences were detected by ear in vocalizations made during artificial feeding of Japanese monkey troops at three locations. Tape recording and sound spectrographic analysis confirmed a distinctive vocal pattern specific to each site and used only in the provisioning situation. The 3 different acoustic morphologies are variations on a shared tonal theme. Vocal learning by Macaca fuscata may have occurred separately at each site regulated by species-wide constraints on vocal production.

Methods

Three study areas where Japanese monkeys ( M . j . fuscata) are provisioned were visited repeatedly in the course of a 14-month study from July 196s through August 1969. These arc located a t Koshima, an islet on the southern coast of western Kyushu; Miyajima, an island near Hiroshima inhabited by a troop translocated from Shodoshinia; and Iwatayama, near Kyoto, where the Arashiyama monkeys dwell. The animals were all individually known.

Tape recordings were made on a Nagra 111 operated at 7.5 inches per second with a directional condenser microphone, the Sennheiser M K H 804. Sound spectrograms were pro- duced by the Kay Electric Co.’s “Sonagraph”, model 6061B, using a 3 0 0 H z filter to scan the SO-8,000 H z range.

More than 50 monkeys, a t least half of each troop, were sampled at each site, with over 6 X 104 vocalizations recorded in the 2X 10s observation hours. Of these, 104 were separately catalogued as given by an identified animal in known circumstances. Only sounds associated with provisioning are presented here.

Introduction

Unlike the many cases reported i n birds (e. g. MARLER 1952; MARLER and T A h l U K A 1962; see reviews by THIELCKE 1969, MARLEK and MUNDINGEK 1971 and NOTTEBOHM 1970 and 1972), there are few examples in mammals of geographic variation of sounds or local dialects. Vocal differences betwecii FSpUldtbllS of deer ( T i . M B K 3 C K 1965), langurs (VOGEL 1973), and squirrel

Dialects in Japanese Monkeys 305

monkeys (WINTER 1969) u-erc noted when comparing races or subspecies, and hence i t is likcly that hereditary influences rather than learning and tradition play the grater role in producing them. The subspecific affinities of pika populations exhibiting geographical vocal variation are ambiguous (SOMERS 1973). In the pulse-rate differences in populations of elephant seals reported by LEBOEUF and PETERSON (1969), it is not yet clear to what degree local factors such as density or level of aggression may be affecting the phenomenon. This report describes locale-specific vocalizations shared among Japanese mon- keys at three sites and given in similar circumstances a t each, and discusses the possible ontogeny of these sounds.

Results

Whenever a gamekeeper or researcher begins to distribute food to the monkeys, a chorus of vocalizations is initiated by one or a few animals in sight of the provisioning. The sounds reach a crescendo as vocal activity spreads contagiously among the troop, and then wane as nearby monkeys begin to eat and as the far-flung members run towards the provisioning area. The chorus may also be triggered by activities associated with provisioning. O n Koshima, for example, the cues which elicit a chorus include the sound of the outboard motor powering the small boat which ferries provisions, a person

A R A S H I Y A M A . *

M I Y A J l M A

I ' I * -.1

K O S H I M A

Fig . I : Locale-specific tonal vocalizations given by Japanese macaques during provisioning a t the 3 sites indicated. The last sounds on the Arashiyama spectrograms and the first on the Koshima ones illustrate the acoustic overlap resulting from simultaneous utterances occurring during a chorus. The last utterances on the Miyajima and Koshima spectrograms d o not show the locale-specific patterns. Sound spectrograms produced by Kay Electric Co.'s "Sonagraph"

using wideband (300 Hz) analysis

2. Tierpsychol. Bd. 38, Heft 3 20

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approaching the food storage h u t on the beach, or one of the familiar feeding personnel preparing to embark from the opposite shore. O n Miyajinia and a t the Arashiyama troops tourists are abundant, and sounds are also uttered in begging for food from them. All the sounds given i n these circumstances are basically tonal in structure, represented on sound spectrograms as an intense band with harmonic overtones. Other classes of vocalization, such as screams or girneys in which atonality or articulations pre-dominate, are not uttered a t these times.

One characteristic sound associated with provisioning a t each site is, to the ear, recognizably typical of the monkeys a t that site only. The Arashiyama monkeys produce a low and rich sound of smooth and continuous tonality with the peak in frequency occurring at or near the end (Fig. I ) . O n Miya- jima, after a similar initiation, a frequency modulation is imposed on the tonal structuring of a somewhat higher and longer sound during a slowly falling phase. O n Koshinia, the continuity of the tonal band is interrupted by an abrupt upward shift in frequency and a characteristic range of more than three-fold results between the maximum and minimum frequencies reached by the fundamental band. These three patterns belong to the same generally tonal portion of the repertoire; the locale-specific qualities derive from the different modifications a t each site of the shared tonal theme. The locale-specific sounds were recorded from both sexes, but not from all age-classes. During .provision- ing, the same animals may also give tonal sounds lacking the site-specific features. Old animals, both 8 6 and 99, utter vocalizations of the same gene- rally tonal pattern during provisioning choruses, sometimes individually idio- syncratic, bu t always lacking the locale-sFecific modifications. Infants less than six months old rarely vocalized during provisioning and were not re- corded giving these sounds.

Discussion and Conclusions

Among possible explanations of the origin, spread and maintenance of

A. There are inherited differences between troops in vocal propensities or motivational control of vocalizations.

B. Contrasts between troops in the motivational states are reflected by voca- lizations, perhaps a function of the different ecological circumstances at each site a t the time of original provisioning 1. Original vocal differences were maintained as a superstitious habit

even if the original motivational contrasts between troops were later blurred

2. Contrasts in motivational state, and hence vocal pattern, were them- selves maintained after provisioning was well established.

C. Behavioral founder-effects account for the vocal differences between troops 1. The effects spread within each troop by socially-facilitated mimicry,

a reflection of the monkeys’ abilities to modify their behavior and to learn new motor patterns by observation, particularly for the ex- ploitation of new food resources.

2. The effects were maintained by inadvertent conditioning contiiigent upon provisioned food as reinforcer.

these locale-specific Japanese monkey sounds are the following:

Dialects in Japanesc Monkeys 307

Inherited differences

Individual vocal idiosyncracies aside, the remainder of the repertoire, presumably under the same genetic influences, is lully shared among the troops at the three sites. Hence, although the requisite degree of breeding isolation undoubtedly exists, the possibility is discounted that a genetically determined founder effect (MAYK 1966) produces the regional variation. A hereditary influence on thresholds or motivations affecting vocal behavior is also pos- sible, but, if present, was not revealed by any systematic differences between sites in social use of the repertoire.

Differences in motivational state

A link between vocal output and motivational or emotional states of monkeys and apes has been noted by many workers (e. g. BOLWIG 1959; ROWELL 1962; ROWELL and HINDE 1962; ITANI 1963; JAY 1965; GREEN 1975); a similar view is reflected i n the widely-held opinion that monkey sounds communicate affect or mood. If nionkeys at each of the three sites share distinctly different moods from that of the monkeys a t the other two, the sounds would differ among the sites.

Macaque sounds of the tonal pattern have been named “clear calls” (RO- W E L L and HINDE 1962) and are represented ononiatopoetically as “koo” (ALT- M A N 1962), “coo” (CHANCE 1956; BERTRAND 1969), “woo” (ANDREW 1962), and “houi”, “oui”, etc. (ITANI 1963). In all reported cases they are used in affinitive circumstances. Many intergraded varieties of tonal sounds, without locale-specific modifications, are used by Japanese monkeys to promote affini- Live contact and to mark position. They may also indicate lack, yet desirability, of contact, thus serving as contact solicitations. These uses do not differ i n form or function among the three sites and they may all be termed contact calls both motivationally and functionally. The fine details of acoustic struc- ture niay be correlated with different social nuances of the contact circum- stances and hence probably to subtle differenccs in the vocalizing animal’s state (GREEN 1975).

The shared locale-specific patterns in tonal vocalizations were not heard outside circumstances usually associated with provisioning. They are linked exclusively to whatever motivational state uniquely accompanies these circum- stances. A t all three sites the locale-specific sounds derive from the tonal por- tion of the repertoire indicating a certain amount of non-arbitrariness in selec- tion of the vocal signals. This shared tonal theme is probably a reflection of the same phylogenetic constraints which yield parallel usage of tonal sounds for affinitive contact in other macaque species. The differences in pattern among the sites might indicate either some relaxation of the motivational control or differences in the motivational substrates accompanying the origin of the three vocal patterns. This latter possibility will be considered first.

A motivationally-linked vocal mechanism could yield locale-specific voca- lizations if there exist locale-specific internal states or motivational substrates. In this sense, the monkeys may “mean” something different at each site and so indicate i t with sounds that differ. This possibility invokes neither learning nor inherited differences yet could account for the pattern variation among sites.

If, for example, supplied food were of different importance to the mon- keys in different locales, then an indication that food is to be proffered could result in different levels of arousal characterizing the internal states of the

308 STEVEN GREEN

monkeys at each site and hence yield different sounds. Provisions may have been much more important to the Koshima troop with its limited range (ca. 75 ha) than to the Arashiyama monkeys who, before provisioning, ranged over 750 ha (HAZAMA 1962) where naturally-occurring food is abundant (HAZAMA and MURATA 1968). The initial years of provisioning a t Koshima could readily have provoked greater arousal there upon sight of a formerly limited resource which became super-abundant. Greater arousal would in turn yield a charac- teristic pattern of vocalization, and the structure of the sounds given by cho- rusing Koshima monkeys does indicate greater excitation of the phonatory apparatus than do the vocal patterns associated with provisioning a t the other sites (cf. GREEN 1975; LIEBERMAN 1967; and LIEBERMAN, HARRIS, WOLFF and RUSSELL 1971 ; GAUTIER in press).

Vocal differences superstitiously maintained

Once established, reinforcement by eating provisioned food would be sufficient to maintain the vocal patterns each individual utters preceding feed- ing as superstitious habit (SKINNER 1948). Such differences in vocal habit could long outlast the inter-troop differences in motivational states which originally determined the vocal patterns. Unlike the high degree of individual stereotypy characterizing superstitious behavior, however, individual monkeys are ob- served to give both locale-specific variants and also tonal sounds lacking features distinctive to the site. Thus the argument for superstitious propagation is not persuasive.

Vocal differences maintained by continuing differences in motivational substrate

Marked regional vocal variation could be maintained if the motivational states which the sounds reflect differed systematically among locales not only a t the advent of provisioning but also if these differences continued after pro- visioning became well-established. The locale-specific vocal patterns are shared a t each site yet show no overlap among sites. The occurrence of widely shared tonal modifications requires, by this explanation, an extremely similar moti- vational state among many individuals a t one site while the nonoverlap re- quires that the same state does not occur a t the others. The monkeys a t the three sites did not exhibit any differences in response to provisioning during the study aside from the vocal pattern. A t each location they had been depen- dent on provisions for many years. This explanation of vocal differences being maintained by contrasting motivational substrates long after provisionization was regularized must be considered unlikely.

Behavioral Founder-Effect

A more attractive explanation for the origin of vocal differences com- bines the notion of a behavioral founder-effect (FRISCH 196s; BURTON and BICK 1972) with socially-facilitated mimicry, enhanced by cultural propaga- tion with feeding as a reinforcer. According to an earlier study by ITANI (1963)) sounds of the tonal class were used by the monkeys of Koshima not only during provisioning but also upon seeing a socially dominant animal. H e notes for his sound “A-12’’ (a tonal sound on his tape recording) that, “It is an interesting phenomenon that the same sounds are uttered either when they welcome their superior or when they know that they are going to be given food.” ITANI also indicates that these sounds became differentiated from the mass of other “A-Group’’ calls (those in his classification given by animals in

Dialects in Japanese Monkeys 309

peaceful states of emotion) only after provisioning began. I did not observe any such “welcoming” usage of the locale-specific sounds. Founding a vocal tradition would require a single monkey who vocalized with a tonal sound in the presence of more dominant monkeys and who then transferred this usage to human beings. People are often treated by these monkeys as dominant animals. If the monkeys were then fed by man, this individual would be con- ditioned so as to be more likely to utter a similar vocal pattern again in man’s presence.

As to the selection of signal pattern, the use of the tonal theme a t the three sites indicates that the broad acoustic structure is circumscribed by the genetic constraints linking tonal sounds to affinitive circumstances. Thus the vocal pattern of the individual founders was not completely arbitrary or accidental. The fine structure of the tonai sounds exhibited as site-specific modifications of tonality may, however, be under somewhat less rigorous control. It cannot now be determined, however, whether the vocal traditions were founded initially with improvised variations on the tonal theme, with idiosyncratic vocal patterns, with sounds reflecting precise internal states of the founder, or whether a variant structure produced within the normal range of acoustic variation became codified in conjunction with provisioning.

Social mimicry and vocal learning

Once well-established in a single individual, the vocal habit of greeting a provisioner with a tonal sound might well be copied by other animals since it would appear to “produce” food. Observational learning is particularly pronounced in monkeys (DARBY and RIOPELLE 1959; KAWAMUKA 1959; HALL 1963; HALL and GOSWELL 1964; STEPHENSON 1967) and Japanese monkeys are renowned for their abilities to copy each other and so transmit habits which allow exploitation of new food resources, e. g. sweet-potato washing (MIYADI 1959, 1965; KAWAI 1965), caratiiel-unwrapping (ITANI 1958), wheat- eating (YAMADA 1957), and placer-mining of grain (KAWAI 1965). Rhesus macaques can in the laboratory be conditioned to alter the fine structure of tonal sounds with food as a reward (LARSON, SUTTON, TAYLOR, and LINDEMAN 1973). The interaction of observational learning with conditioning contingent uFon food reward is perhaps sufficient to account for the locale-specific vocal behavior just as it accounts for other transmitted and shared motor patterns.

Old Japanese monkeys learn novel food habits last if a t all (FRISCH 1968; MIYADI 1959; KAWAr 1965). I n both man and birds vocal learning is more readily accomplished in young individuals (LENNEBERG 1967; NOTTEBOHM 1970; MARLER and MUNDINGER 1971). When the old Japanese monkeys in this study were in their youth, regular provisioning had not yet started. Their failure to show the locale-specific sounds hints strongly that some form of vocal learning may play a role in linking these sounds with provisioning.

Maintenance by conditioning

Conditioning may be inadvertently promoted by human investigators, giving rise to the propagation and maintenance of behaviors in a format usually ascribed to “cultural” transmission. I observed a t Koshima the woman who has been provisioning the animals regularly from the earliest days after ITANI and TOKUDA. She gave sweet potatoes differentially to those monkeys who washed them. Although this was done in the context of demonstrating the existence of Sweet Potato Washing (SPW) behavior to researchers and

310 STEVEN GREEN

tourists, during the course of the study this was the only time sweet potatoes were proffered. Direct reinforcement by human intervention may therefore affect the maintenance of this behavior. Furthermore, because matrilineal clustering characterizes the troop’s spatial organization, the intra-lineage pro- pagation of SPW behavior could have been guided more by the greater likeli- hood of related monkeys being close to each other and hence close to the human agent. Inadvertent conditioning of vocal behavior by man may well have occurred in a similar fashion by regular provisioning and is undoubtedly the case with those monkeys which vocalize in successfully begging food from tourists. The term cultural transmission could apply equally well to the spread of shared vocal signals in these circumstances as it does to the propagation of novel fecding motor patterns.

Provisioning and the function of vocal dialects

Systematic regional variations in song or calls of birds are termed “dia- lects” by NOTTEBOHM (1969) if the populations sharing different vocal patterns neighbor on each other or potentially interbreed. In some examples, locale- specific patterns appear to be culturally transmitted (e. g. MARLER and TA- M U R A 1964, LEMON and SCOTT 1966), although in others the possibility of a genetic influence has not been eliminated (LEMON 1966; NOTTEBOHM and SELANDER 1972; HARRIS and LEMON 1972; KING 1972). Avian dialects usually involve those vocalizations used in mate selection or recognition of mates (THIELCKE 1969). NOTTEBOHM (1972) suggests that vocal learning of dialects in birds may, in combination with the role of sounds in breeding, be part of a mechanism circumscribing the genetic pool and therefore allowing a faster increase of genetically-linked adaptations to local conditions than panmixia would permit. This argument is strongest if the local conditioiis are trans- generational in duration and selective pressures have time to yield an in- creased incidence of the locally adaptive phenotype.

Just as it is advantageous to respond genetically to long term environ- nicntal changes, it is also advantageous to respond behaviorally to local con- ditions of shorter duration. Exploitation of novel foodstuffs with new motor patterns is one obvious example. A selective advantage would accrue to those individuals who are most successful in assimilating and responding to short- term, yet locally important, information, and, in Japanese monkeys or other animals with kin-group societies, to their kin who are spatially closer and thus more likely than distantly-related animals to imitate responses and so utilize the information. The spread of novel food habits in Japanese macaques is evidence of an inherited propensity to initiate and imitate new behaviors.

In addition to plasticity of manipulative motor patterns and a predispo- sition to observational learning, a role is played in the adaptability of these social animals by their communicative behavior. Just as changes in available foodstuffs will yield greater fitness to lineages which can best learn and socially transmit the appropriate new food behaviors, a similar advantage would be experienced by lineages able to communicate changes in resource availability. Animals whose vocalizations best reflect the nuances of excitement provoked by different degrees of concentration or abundance of food, for example, may permit their nearby kin cluster to forage with greater effectiveness. Natural selection is likely to favor those lineages in which communication of local and temporary events is most successful in permitting the kin group to exploit resources more efficiently. One direction such selection can take is towards a vocal process in which the control of fine details of acoustic structure is highly

Dialects i n Japanese Monkeys 311

developed and towards a comniunication system which is to some degree open, allowing for the production and imitation in a single generation of signals linked to local conditions which though they may be important, may also be novel and transient.

An inherited but latent ability to improvise motor and vocal behaviors would be revealed in responses to changes in major environinental influences. For the Japanese monkeys, provisioned food is the most important of the environmental variables affecting both behavior and reproductive rates. T h i s is implied by the booming Fopulations whenever food has been supplied them (FURUYA 1960; TOKUDA 1961-2; CARPENTER and NISHIMURA 1969) and by the shrinking over the years of the areas in which troops spend most of their time, finally narrowly focused on a feeding place (HAZAMA 1964). I suggest that in addition to the new food habits another response to these conditions is the vocal chorus containing unique tonal sounds which accompany provisioning. These locale-specific sounds reflect, I believe, a degree of openness in the Japanese monkeys’ vocal communication system within a pre-determined framework linking the tonal acoustic shapes to an affinitive state.

Although the communication system may be considered closed in its broad outline by the linkage of vocal patterns to internal states, plasticity in the fine gradations of acoustic structure still permits the animals to inform each other unambiguously of this most important event. One particular pat- tern of tonal modification a t each troop has been codified, after founding and mimicry, and is now firmly associated with the provisioning circumstance by conditioning. MARLER ( 1969, 1974) has proposed that the dissociation of im- mutable linkages between internal state and structure of vocalizations niay be one of the preconditions for language development, and it is tempting to speculate that the phenomenon reported here may be an early step in that direction.

The production by Japanese monkeys of a locale-specific sound may recur in other circumstances in which communication could aid the exploita- tion of a local condition. This hypothesis may be tested by examining the form and function of any vocal changes which occur in the Arashiyama troop which was recently translocated to Texas or by closely following the forma- tion of any vocal traditions which may develop as troops in Japan are newly provisioned.

Summary

1. Sound spectrograms of Japanese monkeys from three locations con- firm the field inipressi;.n that there are vocdizations characteristic of and spe- cific to each site. They occur, with other sounds, only during provisioning circumstances. Old animals whose youth antedates the advent of provisioning do not utter the locale-specific sounds.

2. The vocalizations have arisen independently a t each site yet they reflect modifications of a shared tonal theme. Tonal sounds are used by Japa- nese monkeys and other macaques in affinitive circumstances. This variation on the tonal acoustic theme indicates that the provisioning of food, a social attractant, is not communicated with an arbitrarily-selected vocal pattern, but rather with one determined in part by the genetic heritage of the animals.

3 . A number of possibilities are discussed concerning the origin and maintenance of vocal tradition. The locale-specificity of the sounds is ascribed to a behavioral founder-effect. The sharing a t each site is attributed to socially- facilitated mimicry, and the maintenance to vocal conditioning. The function

312 STEVEN GREEN

of the sounds reflects a degree of openness in the vocal communication system which allows the spread of a new signal pattern within a single generation, in a similar time span to the occurrence of the new phenomenon with which it is associated, namely provisioning.

Zusammenfassung

1. Ton-Spektrogramme japanischer Affen aus drei Gebieten beststigen den im Freien gewonnenen Eindruck, dai3 es fur jeden Bezirk charakteristische Lautgebungen gibt. Sie treten nur wahrend der Futterbeschaffilng auf. Altere Tiere, deren Jugend vor dem Beginn der kunstlichen Futterung liegt, i d e r n solche ortsgebundenen Laute nicht.

2 . Die Laute entwickeln sich in jedem Bezjrk unabhangig, sind jedoch Variationen eines Themas. Solche Laute werden auch von anderen Makaken unter entsprechenden Umstanden benutzt. Nahrungsbeschaffung als soziales Ereignis wird also nicht mit einem willkurlich gewahlten Laut verknupft, son- dern mit einem wohl genetisch dafur vorbestimmten.

3. Mogliche Ursprunge und Traditionen der Laute werden besprochen. Das Lokalkolorit der Laute wird einem Verhaltens-Grundereffekt zugeschrie- ben, die Einheitlichkeit in jedem Bezirk sozialer Imitation und die Beibe- haltung stimmlicher Konditionierung. Die Laute zeigen eine gewisse Freiheit in der stimmlichen Verstandigung, welche die Ausbreitung neuer Ausdrucks- weisen innerhalb einer Generation erlaubt, also ahnlich rasch, wie sich Futter- annehmen, mit dem es verknupft ist, ausgebreitet hat.

Acknowledgments

This research was supported by a U.S.P.H.S. behavioral sciences training grant (GM 1789), the New York Zoological Society, and the Rockefeller University’s graduate fellowship program. Partial support during manuscript preparation was provided by a N.S.F. grant (GO 16606) to P. MARLER. The field study was conducted under the auspices of the Japan-U.S. Cooperative Science Program; the many individuals in Japan whose help is gratefully acknow- ledged are listed in the major report of the research results (GREEN 1975). For their many helpful comments and discussion of this manuscript, I thank P. MARLER, K. MINKOWSKI, and F. NOTTEBOHM. I thank J. ITANI for supplying me with a copy of the tape “Vocal Sounds of Wild Japanese Monkeys - Series One”. The Sounds were recorded at Takasakiyama in 1960 by ITANI and M. NAKAJIMA and arranged by K. KAWANAKA in 1966 to illustrate ITANI’S vocal categorization scheme.

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Author’s address: Steven GREEN, The Rockefeller University, Field Research Center, Millbrozk, N. Y. 12545, U.S.A.