摘 要
本研究針對壽山地區的台灣獼猴進行調查，野外觀察時間自1999年10月至2000年10月，共記錄了375小時，其中包含55小時的掃描取樣(scan sampling)時間及320小時的焦點追蹤(focal animal sampling)時間。壽山地區台灣獼猴的叫聲至少有25種不同的聲音類型 (vocal pattern)，其中包括32種不同的聲音 (vocal type)。在這25種聲音類型中，出現頻度最高的三種類型分別是coo calls (18.59%)、hack (17.78%) 及contact rattle (11.85%)。其餘的聲音類型還有squeak (11.34%)、noise and undulated scream (6.61%)、sneeze (4.54%)、greeting (3.77%)、squeal (3.41%)、vibrato growl (3.02%)、growl (2.86%)、cluck (2.54%)、female copulation call (2.45%)、squawk (2.14%)、tonal scream (2.03%)、alarm call (1.74%)、threat rattle (1.66%)、chuckle (0.92%)、rise (0.61%)、weeping (0.56%)、male copulation call (0.32%)、babble (0.30%)、whine (0.29%)、roar (0.27%)、bark (0.24%)及mounting grunt (0.15%)。
所有的聲音類型依其間有無中間型存在可分為二類：獨立訊號(Discrete signal)及層級訊號(Graded signal)。獨立訊號就是在各聲音類型間缺乏中間型聲音，包括coo call、greeting、contact rattle、mounting grunt、male and female copulation calls, babble、whine及weeping。而層級訊號則是在各聲音類型中有中間型態的聲音存在，包括了屬於敵對叫聲的threat rattle、growl、bark、roar、vibrato growl和alarm call, 還有chuckle及屬於屈服叫聲的noise and undulated scream、squeal、 tonal scream and squeak以及屬於難過不舒服(distress)的叫聲(包括squawk、hack、cluck和sneeze)。從發聲的情境來看，屬於接近聯繫的叫聲有coo calls, greeting and contact rattles，以減少或維持與其他成員之間的距離。Growls、threat rattles和vibrato growls 屬於敵對的叫聲，發生在群內成員間位序高者威脅位序低者或群對抗及威嚇其他物種時。屈服的叫聲包含noise and undulated scream, squeal, tonal scream及兩種squeak，其聲音結構有明顯變化。而交配叫聲及警戒聲則與特定的互動或特定的外在刺激有關。
特定社會因素、發聲者的動機及激動狀態可能會影響聲音結構。屈服叫聲的結構可能與敵對互動時，有否身體接觸有關。Noise and undulated scream 較常發生在有身體接觸的敵對互動；而squeal、tonal scream 及 tonal and compound squeak 較常出現在無身體接觸的敵對互動。
台灣獼猴之聲音行為在年齡間的差異比在性別間的差異更大。各年齡性別層的相對頻度及發聲率上，從嬰猴到成年猴遞降。在所有的聲音表現中各年齡性別層裡，嬰猴有最高的發聲率(每10小時有1142.2個聲音單位)及相對頻度(44.58%)。隨著年齡增加，視覺訊號不只補充了聲音訊號，也可能取代了聲音訊號。有四種聲音類型(coo call、greeting、squeak及hack)在各年齡性別層中的相對頻度是從嬰猴到成年猴遞降；而threat rattle及growl 這二種聲音類型的相對頻度則是從嬰猴到成年猴遞增。cluck、sneeze、whine、weeping及babble這五種聲音類型僅在幼猴及嬰猴出現。 而在squeak (64 %)、hack (63.6 %)、squawk (76%)、cluck (85.7 %)、sneeze (90 %)、weeping (88 %)及babble (100%)中，大多由嬰猴產生，而且其中有三種聲音類型主要是由年齡階層II的嬰猴發出。另外，只有嬰猴會發出babble；而roar及bark只由成年或亞成年猴產生。
在聲音類型的相對頻度上，成年猴間有較明顯的性別差異。除了roar外，大部分的聲音類型其在成年母猴的相對頻度及發聲率皆高於其在成年公猴的。另外，母猴在六種聲音類型中(vibrato growl、chuckle、contact rattle、squeal、squeak及hack)的相對頻度及發聲率皆高於同年齡層的公猴。此外，mounting grunt、male copulation call只出現在成年公猴；而female copulation call 則只出現在成年與亞成年母猴。
特定年齡性別層中發出或不發出某些聲音，可能與社會組織、生理形態特性及環境經驗有關。而台灣獼猴的聲音表現與其他同屬的獼猴，尤其是日本獼猴(M. fuscata)、恆河猴(M. mulatta)和綺帽獼猴(M. radiata)有高度相似性。這可能是由於親緣關係、棲地和社會組織相似的緣故。

ABSTRACT
Field observation and of sound recording of Formosan macaques (Macaca cyclopis) at Mt. Longevity were conducted from Oct. 1999 to Oct. 2000. Spectrographic analyses revealed at least 25 basic patterns with 32 different vocal types from a total of 375 hrs recordings (55 hrs for scan sampling and 320 hrs for focal animal sampling). The three most frequent calls in overall vocal repertoire were coo calls (18.59%), hack (17.78%) and contact rattle (11.85%). The other calls included squeak (11.34%), noise and undulated scream (6.61%), sneeze (4.54%), greeting (3.77%), squeal (3.41%), vibrato growl (3.02%), growl (2.86%), cluck (2.54%), female copulation call (2.45%), squawk (2.14%), tonal scream (2.03%), alarm call (1.74%), threat rattle (1.66%), chuckle (0.92%), rise (0.61%), weeping (0.56%), male copulation call (0.32%), babble (0.30%), whine (0.29%), roar (0.27%), bark (0.24%) and mounting grunt (0.15%).
Formosan macaques employ a complex vocal system composed of discrete signals as well as graded signals which vocal patterns connected by intermediate gradations. Discrete signal included coo call, greeting, contact rattle, mounting grunt, two copulation calls, babble, whine, weeping. Whereas the graded signals included the aggressive signals (threat rattle, growl, bark, roar, vibrato growl and alarm call), chuckle, the submissive signals (noise and undulated scream, squeal, tonal scream and squeak) and the distress signals (squawk, hack, cluck and sneeze).
According to the context of emission, some of vocalizations could be divided into: (1) Affiliative contact calls: The calls, as the coo calls, greeting and contact rattles, may be summarized as affinitive contact calls for reduction and/or maintenance of close proximity between group members. (2) Aggressive calls: The growls, threat rattles and vibrato growls were uttered by dominant animals menacing sub-dominate group mates, members of other groups or other species. (3) Submissive calls: these submissive calls (including the various form of scream, squeal and squeak) with obviously structure differences. Male and female copulation calls and alarm call of M. cyclopis were clearly related to particular interactions or specific external stimulus.
The acoustic structures might be influenced by specific social factors, motivation or the arousal state of the callers. The acoustic structure of submissive calls was associated with the aggression with or without physical contact. Noise and undulated scream were usually used in aggressive interactions with physical contact (68%), whereas squeal, tonal scream and two types of squeak were used without physical contact (75%).
Age specific difference of vocal behavior seemed to more pronounced than asymmetries between the sexes. Infants have the highest vocal rate (1142.2 vocalizations/10 h) and relative frequency of vocalization (44.58 %) than other age/sex classes and they decreased from infants to adults. With increasing age, visual signals could not only complement vocal signals but also replace the vocal cues. The relative frequency of four vocal patterns (coo call, greeting, squeak and hack) decreased from infants to adults, whereas that of threat rattle and growl increased from infant to adults. On the other hand, there were five vocal patterns (including cluck, sneeze, whine, weeping and babble) only used by juveniles and infants. In addition, seven vocal patterns included squeak (64 %), hack (63.6 %), squawk (76%), cluck (85.7 %), sneeze (90 %), weeping (88 %) and babble (100%) mainly were used by infants. Among them, three vocal patterns (squeak, hack and squawk) were produced mostly by infant II. Only infants used babble. Roar and bark were only present in adults and sub-adults.
Obvious sex differences of relative frequencies of vocal patterns occurred in adults, the vocal activity of females consistently higher compared to that of males. In six patterns (vibrato growl, chuckle contact rattle, squeal, squeak and hack), females uttered more often than male peers. Mounting grunts and male copulation calls were exclusively used by adult males whereas female copulation calls were only produced by adult/sub-adult females.
Vocalizations were produced/ceased by certain sex/age class that might associate with social organization, morphological feature and circumstances experience. The vocal repertoires of Formosan macaques revealed the high similarities in the species within genus Macaca, especially to M. fuscata, M. mulatta and M. radiata. It may be due to phylogeny, habitats and social organizations.

TABLE OF CONTENTS
誌謝.......................................i
中文摘要..................................ii
Abstract..................................iv
Introduction...... .........................1
Materials and Methods...... ................2
Study site...............................2
Study groups and methods.................3
Terminology and analysis.................4
Results.. ...................................6
Part I. Vocal categories of Formosan macaques.........................6
Part II. Comparative analysis of age- and sex- specific patterns.....27
Discussion........ .........................29
Vocal patterns....... ................29
Social contexts......................31
Acoustic strutures...................33
Age- and sex- specific patterns......35
In response to potential predators...38
Male loud calls...... ................39
Summary...............................40
References..................................41
Tables......................................48
Figures.....................................64

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