摘要
　　臺灣獼猴（Macaca cyclopis）屬於臺灣特有種保育類生物，其野外社會行為的相關研究極少。在獼猴的社會行為中，分群行為（troop fission）屬於偶發且較罕見的行為。本實驗以高雄壽山地區發生分群的台灣獼猴社群B0群及C0群作為研究對象，自1999年11月及12月底分群發生開始，追蹤觀察至2000年5月31日，記錄分群後社群組成改變及互動狀況，並以此推測台灣獼猴分群發生的原因。記錄方式是以焦點取樣及掃描取樣方法，來收集四群猴群（主群B群、旁支群Ba群、主群C群及旁支群Ca群）成員間的個體行為、社會行為，及社群之棲地利用。B群及Ba群在分群發生六個月後，仍維持各自獨立的社群狀態，而Ca群成員則在分群發生後的兩個多月，即返回C群。在分群時，遷出原社群進入旁支群的個體，均為原社群內低位序或周圍個體。而這些個體在遷入旁支群後，位序較其在原社群中為高。分群推測是由成年雌猴發動，其餘成員藉機加入。與成年雄猴比較，成年雌猴為旁支群中較為固定的核心組成成員。在主群及旁支群的性別比結果中，僅Ca群雌猴比例較分群前低；而所有在旁支群中生育的雌猴，均在分群前即已受孕。由結果推斷：臺灣獼猴分群行為的發生，並非雌猴欲藉此增加交配對象或機會，而可能是由於雌猴在原群中的適存性喪失所致。分群後旁支群Ba群及Ca群的成員數目約為主群B群及C群的1/4~1/5。主群B群與旁支群Ba群相遇時的行為，顯示其較旁支群Ba群佔優勢。而在棲地利用的情形上，B群與Ba群對活動地區之利用範圍上有顯著差異（χ2= 49.195, d.f.= 14, p< 0.05），旁支群Ba群的活動範圍較主群B群及未分群前社群B0大。但在夜宿地的利用程度方面，旁支群Ba群與主群B群則無顯著差異（χ2= 11.6, d.f.= 13, p> 0.598）。

Abstract
Formosan macaque (Macaca cyclopis) is the endemic and threatening species of Taiwan, but information regarding their social behaviors is rare. Troop fission of macaque’s social behaviors was the rarely and accidental behavior. Two fission of Formosan macaques troops, from B0 and C0, at Mt. Longevity of Taiwan were the objects in this study. Field study was concentrated from November 1999 when fission occurred till May 2000. The records included the troop’s composition changes and interactions of troops after fission, and using these records to infer the reason of troop fission in Formosan macaques. The focal sampling method and scan sampling method were used to collect individuals’ behaviors、social behaviors and habitat utilization of four troops（main troop B, branch troop Ba, main troop C and branch troop Ca） after fission. Troop B and Ba remained separately six months after fission. But two months after fission, a fusion occurred between troop Ca and troop C. In the phenomenon of fission, the individuals who left the original troop and emigrated to the branch troop were all the subordinate or periphery individuals in the original troops. When they emigrated to the branch troops, their ranks were higher than that of the original troops. Adult females initiated the troop fission, and the other individuals joined later. Compared with the adult male, the composition of troop adult females were much stable and were the nucleus of branch troops. Only the adult sex ratios of troop Ca was lower than that of the original troop. Females delivered infants in branch troops were pregnant before troop fission. These two cases indicated that: fission occurred in Formosan macaque was not due to increasing mating partners or chances, but because of some adult females’ fitness lost in their original troop. After fission, the branch troops, Ba and Ca, were 1/4 ~ 1/5 troop size to the main troop. When main troop B met the branch troop Ba, the behaviors of main troop B indicated troop B was dominant to the branch troop Ba. The main troop B and branch troop Ba had a significant difference in home range using （χ2 = 49.195, d.f.= 14, p< 0.05）. The branch troop Ba’ home range was larger than that of the main troop B or of the original troop B0. The difference of night resting-sites between the main troop B and the branch troop Ba was not significant（χ2 = 11.6, d.f.= 13, p> 0.598）.

目錄
誌謝 i
中文摘要 ii
英文摘要 iii
前言 1
材料與方法 4
一、實驗地 4
二、觀察對象 4
三、研究分析方法 5
結果 7
一、分群初期記錄 7
(一) 旁支群Ba群的分群初期狀況及組成改變 7
(二) 旁支群Ca群的分群初期狀況及組成改變 8
二、分群前後雌雄猴之位序變化 9
(一) 成年雄猴的位序變化 9
(二) 成年雌猴的位序變化 9
(三) 旁支群雌猴遷回主群後之位序變化 10
三、分群前後之社群組成變化 11
(一) 分群後社群成員數目與成年性別比之變化 11
(二) 雌雄猴的交配情形 11
(三) 雌雄猴間的互動 12
(四) 分群後旁支群雌猴懷孕之情形 13
四、社群間的互動 14
(一) 旁支群與主群的互動情形 14
(二) 旁支群與其他社群相遇時的表現 16
五、棲地利用 16
(一) 主群與旁支群對活動地區及夜宿地的利用情形 16
(二) 旁支群接近主群夜宿地時的特殊情形 18
討論 21
一、與”Dittus”五項假說之討論 21
二、”雌猴在群內的適存性降低”假說之討論 21
三、分群後旁支群與主群的互動關係 25
四、分群後旁支群與主群之棲地利用關係 25
結論 27
參考文獻 28
圖表 34
附錄 52

參考文獻
鹿野忠雄，1929，臺灣產哺乳類的分布及習性。動物學雜誌，41 (489): 332-340。
彭明聰、賴義隆，1968，臺灣獼猴之性週期與其他種猴之比較。師大生物學報，3: 10-13。
吳海音，1986，墾丁地區臺灣獼猴的行為與生態學研究。國立台灣大學動物學研究所碩士論文，107頁。
林進丁，王建平，1991，高雄市壽山地區臺灣獼猴之社群分佈與棲地利用調查。行政院農委會，50頁。
林金福，徐芝敏，1998，壽山臺灣獼猴活動範圍與變遷之因素探討。科學教育學術研討會短篇論文集，863頁~871頁。
儲瑞華，1998，柴山台灣獼猴成年雌猴的親緣、階級與社會行為。國立台灣大學動物學研究所碩士論文，60頁。
Altmann, J., 1974. Observational study of behavior: sampling method. Behavoiour, 49: 227-262.
Caldecott, J. D., 1986. Mating patterns, societies and the ecogeography of macaques. Anim. Behav., 34: 208-220.
Chagnon, N. A., 1975. Genealogy, solidarity, and relatedness: limits to local group size in an expanding population. Ybk phys. Anthropol., 19: 95-110.
Chalmers, N., 1979. Social behavioer in primates. Edward Arnold, London.
Chepko-sade, D.B & D. S. Sade, 1979. Patterns of group splitting within matrilineal kinship groups. A study of social group structure in Macaca mulatta (Cercopithecidae: Primates). Behav. Ecol. Sociobiol., 5: 67-86.
Cords, M. & Rowell, T. E., 1986. Group fission in blue monkeys of the Kakamega Forest, Kenya. Folia primatol., 46: 70-82.
Dien, Z. M., 1958. The Formosan rock-monkey. Quart. J. Taiwan Museum, 11: 345- 348.
Dittus, W. P. J., 1977. The social regulation of population density and age-sex distribution in the toque monkey. Behaviour, 63: 281-322.
Dittus, W. P. J., 1979. The evolution of behaviours regulating density and age-specific sex ratios in a primate population. Behaviour, 69: 265-302.
Dittus, W. P. J.,1982. Population regulation: the effects of severe environmental changes on the demography and behavior of wild toque macaques. Int. J. Primatol., 3: 276.
Dittus, W. P. J., 1985. The influence of cyclones on the dry evergreen forest of Sri Lanka. Biotropica, 17: 1-14.
Dittus, W. P. J., 1986. Sex differences in fitness following a group take-over among Toque macaques: testing models of social evolution. Behav. Ecol. Sociobiol., 19: 257-266.
Dittus, W. P. J., 1988. Group fission among wild toque macaques as a consequence of female resource competition and environmental stress. Anim. Behav., 36: 1626-1645.
Dunbar, R. I. M. & E. P. Dunbar, 1977. Dominance and reproductive success among female gelada baboons. Nature, Lond., 266: 351-352.
Dunbar, R. I. .M., 1980. Demographic and life history variables of a population of gelada baboons (Theropithecus gelada). J. Anim. Ecol., 49: 485-506.
Eisenberg, J. F., 1966. The social organization of mammals. Handbk Zool. 10: 1-92.
Eudey, A. A., 1980. Pleistocene glacial phenomena and the evolution of Asian macaques. In: Studies in ecology, behavior and evolution. Lindburg, D. G. (ed.) Van Nostrand Reinhold Company, New York, pp. 52-83.
Fooden, J., 1976. Provisional classification and key to living species of macaques. Folia Primatol, 25: 225-236.
Fujii, H., 1974. Structure of a natural group of Japanese monkeys at Katsuyama, based on individual relationships. Machikaneyama Ronso, 7: 1-18. (in Japanese)
Furuya, Y., 1960. An example of fission of a natural troop of Japanese monkey at Gagyuzan. Primates, 2: 149-180.
Furuya, Y., 1962. On the ecological survey of the wild crab-eating monkeys in Malaya. Primates, 3: 75-76.
Furuya, Y., 1968. On the fission of troops of Japanese monkeys. I. Five fission and social changes between 1955 and 1966 in Gagyuzan troop. Primates, 9: 323-350.
Furuya, Y., 1969. On the fission of troops of Japanese monkeys Ⅱ. Primate, 10: 47- 69.
Hsu, M. J. & G. Agoramoorthy, 1997. Wildlife conservation in Taiwan. Conserv Biol, 11: 834-836.
Hsu, M. J., Lin, J. F & G. Agoramoorthy, 2000. Occurrence of twins in wild Formosan Macaques, Macaca cyclopis, at Mt. Longevity, Taiwan. Folia Primatol, 71: 154-156.
Kleiman, D., 1977. Monogamy in mammals. Q. Rev. Biol., 52: 39-69.
Koyama, N., 1970. Changes in dominance rank and division of a wild Japanese monkey troop in Arashiyama. Primates, 11: 335-390.
Kudo, H., 1984. Dynamics of social relationships in the process of troop formation of Japanese monkey troop in Koshima-“working upon” and “impressing behavior”. J. Anthropol. Soc. Nippon., 92: 253-272. (in Japanese)
Malik, I., Seth, P. K. & C. H. Southwick, 1985. Group fission in free-ranging rhesus monkeys of Tughlaqabad , Northern India. Int. J. Primatol., 6: 411-422.
Martin, P. & P. Batesson, 1993. Measuring behaviour an introductory guide. Cambridge University press. 222pp.
Maruhashi, T., 1982. An ecological study of troop fissions of Japanese monkeys (Macaca fuscata yakui) on Yakushima Island, Japan. Primates, 23: 317-337.
McCullough, D. R., 1974. Status of larger mammals in Taiwan. Tourism Bureau, Taipei, Taiwan, R. O. C.
Mellen, J. D., Littlewood, A. P., Barrow, B. C. & V. J. Stevens, 1981. Individual and social behavior in a captive troop of mandrills (Mandrillus sphinx). Primates, 22: 206-220.
Melnick, D. J. & K. K. Kidd, 1983. The genetic consequences of social group fission in a wild population of rhesus monkeys (Macaca mulatta). Behav. Ecol. Sociobiol., 12: 229-236.
Moore, J., 1984. Female transfer in primates. Int. J. Primatol., 5: 537-589.
Murie, J., 1872. Observations on the macaques. Ⅲ. The Formosan or round-faced monkey. Proc. Zool. Soc. London, 1872: 771-774.
Nash, L. T., 1976. Troop fission in free-ranging baboons in the Gombe Stream National Park, Tanzania. Am. J. Phys. Anthropol., 44: 63-78.
Nishida, T., 1966. A sociological study of solitary male monkeys. Primates, 7: 141-204.
Nishimura, A., 1973. The third fission of a Japanese monkey group at Takasakiyama. In: Behavioral Regulators of Behavior in Primates. C. R. Carpenter (ed.), Bucknell Univ. Press, Lewisburg. pp. 115-123.
Nishimura, A., 1977. Group fissions of Japanese monkeys at Takasakiyama, from third to sixth fission. In: Population Dynamics of Japanese monkeys at Takasakiyama, Report of the Investigation 1971-1979, Oita City, pp. 37-91. (in Japanese)
Oi, T., 1988. Sociological study on the troop fissio of wild Japanese Monkeys (Macaca fuscata yakui) on Yakushima Island. Primates, 29: 1-19.
Peng, M. T., Lai, Y. L., Yang, C. S. & H. S. Chiang, 1973a. Formosa monkey (Macaca cyclopis): Present situation in Taiwan and its reproductive biology. Exp. Anim. 22(suppl.): 447-451.
Peng, M.T., Lai, Y. L., Yang, C. S. Chiang. H. S, New. A. E. & C. P. Chang, 1973b. Reproductive parameters of the Taiwan monkey (Macaca cyclopis). Primate, 14: 201-214.
Poirier, F. E. & D. M. Davidson, 1979. A preliminary study of the Taiwan macaque. Quart. J. Taiwan Museum., 32: 123-191.
Pusey, A. E. & C. Packer, 1987. Dispersal and philopatry. In: Primate Societies, B. B. Smuts, D. L. Cheney, R. M. Seyfarth, R. W. Wrangham, & T. T. Struhasaker (eds.), The Univ. of Chicago Press, Chicago, pp. 250-266.
Richards, S.M., 1974. The concept of dominance and methods of assessment. Anim. Behav., 22: 914-930.
Sackett, G., Holm, R. & S. Landesman-Droyer, 1975. Vulnerability for abnormal Develoent: pregnancy outcomes and sex differences in macaque monkeys. In: Aberrant Develoent in Infancy. Human and Animal Studies. N. R. Ellis (ed.), John Wiley. N. Y., pp.59-76.
Sade, D., 1967. Determinants of dominance in a group of free-ranging rhesus monkeys. In: Social Communication in Primates. S. Altmann (ed.), The Univ. of Chicago Press, Chicago, pp. 99-114.
Silk, J. B., Clark-Wheatley, C. B., Rodman, P. S. & A. Samuel, 1981. Differential reproductive success and facultative adjustment of sex ratios among captive female bonnet macaques (Macaca radiata). Anim. Behav., 29: 1106-1120.
Small, M. F, 1983. Females without infants: mating strategies in two species of captive macaques. Folia Primatol, 40: 125-133.
Southwick, C. H. Beg, M. A. & M. R. Siddiqi, 1965. Rhesus monkeys in north India. In: Primate Behavior. I. DeVore (ed.), Holt, Rinehart & Winston Press, New York, pp. 111-159.
Sugiyama, Y., 1960. On the division of a natural troop of Japanese monkey at Takasakiyama. Primate, 2: 109-148.
Sugiyama, Y., & H. Ohsawa, 1974. Population dynamics of Japanese monkeys with special reference to the effect of artifitial feeding. Folia Primatol., 39: 238-263.
Sussman, R. W. & I. Tattersall, 1981. Behavior and ecology of Macaca fascicularis in Mauritius: a preliminary study. Primates, 22: 192-205.
Takizawa, H, 1983. Subgrouping in Japanese monkeys-A case study on Kamuri-A troop. Unpublished master’s thesis, Kanazawa Univ. (in Janpanese)
Tamaki, M., 1982. An ecological study of troop fission of Japanese monkeys (Macaca fuscata yakui) on Yakushima Island, Japan. Primates, 23: 317-337.
Teas, J., Richie, T., Taylor, H. & C. Southwick, 1980. Population patterns and behavioral ecology of rhesus monkeys (Macaca mulatta) in Nepal. In: Studies in ecology, behavior and evolution. Lindburg, D. G. (ed.), Van Nostrand Reinhold Company, New York, pp. 148-181.
Tokita, E., 1980. The fission of Shiga-A troop of Japanese monkeys. Nat. & Anim., 10: 26-30.
Tokuda, K, 1962. A study on the sexual behavior in the Janpanese monkey troop. Primates, 3:1-41.
Vogt, J. L., 1978. The social behavior of a marmoset （Saguinus fuscicollis）group Ⅱ: behavior patterns and social interaction.
Wheatley, B. P., 1980. Feeding and ranging of east Bornean Macaca Fascicilaris. In: Studies in ecology, behavior and evolution. Lindburg, D. G. (ed.), Van Nostrand Reinhold Company, New York, pp. 148-181.
Wrangham, R. W., 1979. On the evolution of ape social systems. Social Science Information, 18: 335-368.
Wrangham, R. W., 1980. An ecological model of female-bonded primate groups. Behaviour, 26: 105-120.
Wrangham, R. W., 1981. Drinking competition among vervet monkeys. Anim. Behav., 29: 904-910.
Wu, H. Y., 1986. The behavioral and ecological study of the Taiwan macaque (Macaca cyclopis) in Kenting area. Master thesis, National Taiwan Univ. (in Chinese)
Wu, H. Y., and, Y. S. Lin, 1992. Life history variables of wild troops of Formosan Macaques (Macaca cyclopis) in Kenting, Taiwan. Primates, 33: 85-97.
Yamagiwa, J., 1985. Socio-Sexual Factors of troop fission in wild Japanese monkeys (Macaca fuscata yakui) on Yakushima Island, Japan. Primates, 26: 105-120.