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研究生:林泰榮
研究生(外文):Tai-jung Lin
論文名稱:壽山台灣獼猴之理毛行為
論文名稱(外文):Grooming Behavior of Formosan macaques (Macaca cyclopis) at Mt. Longevity, Taiwan
指導教授:徐芝敏徐芝敏引用關係
指導教授(外文):Hsu, Minna J.
學位類別:博士
校院名稱:國立中山大學
系所名稱:生物科學系研究所
學門:生命科學學門
學類:生物學類
論文種類:學術論文
論文出版年:2009
畢業學年度:97
語文別:中文
論文頁數:188
中文關鍵詞:優勢位階親屬選擇社會理毛台灣獼猴年齡層季節
外文關鍵詞:Macaca cyclopisSocial groomingKin selectionDominance rankAgeSeason
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本研究是探討高雄市壽山地區台灣獼猴(Macaca cyclopis)成年雌猴之社會理毛在親屬、位序、年齡與季節變化之關係,主要的研究對象是C群與Cd群成年獼猴。從2000年8月到2003年2月止,歷經3個交配季與2個非交配季,共188個工作天,約1248.8個小時。實際記錄C群660.6小時,Cd群為244.5小時。研究期間C群成年雄猴有8-13隻,成年雌猴有14-15隻;Cd群成年雄猴有1-3隻,成年雌猴2-4隻。研究成年雌猴區分為高低位階及老年(>13歲)和年輕(5-12歲)。另於理毛對中,分別判定二者相對高低位序以及親屬或非親屬的關係。而成年雄猴則區分為群內雄猴與周邊雄猴兩類型。
成年雌猴個體之異體理毛活動佔本身日間行為的37.62% ± 13.59%(n = 15)。在台灣獼猴社會理毛活動中,成年雌猴給予嬰、幼猴理毛多於接受者(p < 0.001)。成年雌猴間的社會理毛活動在非交配季比較頻繁,約是交配季的2.12倍。無論在交配季或非交配季,成年雌猴與親屬雌猴間的理毛頻度皆高於和非親屬關係者(p < 0.001)。另外在交配季,成年雌猴與相對低、高位序成年雌猴理毛頻度受親屬關係影響(p < 0.05),不受個體位階影響。但以高階成年雌猴給予相對低位序親屬雌猴理毛的頻度最高(0.38 ± 0.40次/ 100 scans, n = 7)。在非交配季,成年雌猴與相對低位序雌猴理毛頻度受親屬關係影響(p < 0.05),以老年成年雌猴與相對低位序親屬雌猴的理毛頻度最高(1.57 ± 1.74次/ 100 scans, n = 8)。相對的,年輕成年雌猴比老年者與相對高位序親屬雌猴的理毛頻度高(p < 0.05)。
季節、親屬關係對成年雌猴給予或接受成年雌猴理毛之頻度皆有顯著影響(兩者p < 0.01)。成年雌猴和相對低位序成年雌猴理毛頻度受親屬關係及季節之影響(p < 0.01),以非交配季成年雌猴和相對低位序親屬雌猴的理毛頻度最高,100次掃瞄中有1.25次(± 1.48, n = 11)。在沒有親屬關係下,低階成年雌猴比高階成年雌猴給相對高位序雌猴較多的理毛頻度(p < 0.05)。成年雌猴間的社會理毛次數有65%發生在親屬雌猴之間。40%成年雌猴給親屬雌猴理毛比例高於給非親屬者,20%給非親屬雌猴理毛比例高於給親屬者。而40%成年雌猴與親屬、非親屬雌猴理毛無顯著差異。而非親屬成年雌猴間的社會理毛有8.68%為互惠理毛。
成年雌猴給或接受親屬雌猴理毛伙伴比例皆高於非親屬關係者(兩者p < 0.05)。成年雌猴接受高階雌猴理毛的伙伴比例高於接受低階者(p < 0.01)。而成年雌猴給或接受老年、年輕雌猴理毛伙伴比例則皆相近。另外成年雌猴無論給或接受成年雄猴之社會理毛頻度,在交配季皆比非交配季多(p < 0.05)。而高階成年雌猴比低階者與成年雄猴有較頻繁的社會理毛。
C和Cd群的成猴性別比在交配季相近(AM:AF = 1:1.8)。兩群成年雌猴與群內及周邊雄猴的社會理毛在交配季多於在非交配季。雄猴類別(群內、周邊雄猴)、社群(C、Cd群)對成年雄猴之間的理毛頻度在交配季有顯著影響(p < 0.05),但是在非交配季則無顯著影響(p > 0.1)。且群內雄猴比周邊雄猴有較多的理毛伙伴,其主要的理毛伙伴為成年雌猴。周邊雄猴則比群內雄猴有較多的雄猴理毛伙伴。另外在交配季,低階雄猴與優勢雄猴的理毛對中常為被理毛者,但是在非交配季則為理毛者。雄猴之間的衝突行為較常發生在交配季,且周邊雄猴間的衝突頻度是群內雄猴間的1.8倍(p < 0.05)。
壽山台灣獼猴C和Cd群成員異體理毛與自體理毛的偏好部位有相關性(p < 0.001)。獼猴異體理毛偏好於背部,而陰肛最少。自體理毛偏好於下肢部位,而背部和臉部則都沒有記錄資料。不同性別成年個體對獼猴頭部、背部、臉部理毛之頻度有顯著差異(p < 0.05)。有親屬關係之成年雌猴間頭部理毛之頻度多於非親屬關係者(p < 0.05)。而位階、年齡層對成年雌猴間的理毛部位則皆無顯著影響(所有p > 0.1)。
因此,壽山台灣獼猴社群成年雌猴的社會理毛應該是以親屬選擇為主要基礎。無親屬關係成年雌猴的社會理毛則可能藉由互惠理毛達到結盟或提升位序之目的。而成年雄猴與成年雌猴的社會理毛在交配季多於在非交配季,此應該與雄猴的交配策略有關。而周邊雄猴比群內雄猴有較多的雄猴理毛伙伴,則可能是以結盟為目的。
I have investigated the social grooming in kinship, rank, age and seasonal change among adult female Formosan macaques (Macaca cyclopis) that inhabit Mt. Longevity, Kaohsiung. The major study groups were C and Cd groups. Field observations were conducted from August 2000 to February 2003 covering three mating seasons and two non-mating seasons. The observations covered a total of 188 work days including 1248.8 hours. I actually had recorded C group for 660.6 hours, and Cd group for 244.5 hours. During my study, C group consisted of 8-13 adult males and 14-15 adult females, while group Cd had 1-3 adult males and 2-4 adult females. In order to analyze grooming data, I divided 15 adult females into sub-groups such as dominant/ submissive groups, old (>13) /young age (5-12), relative higher/lower ranking and related/unrelated. I have also divided adult males into troop-males and periphery males.
Adult female allo-grooming activities accounted for 37.62% ± 13.59 (n = 15) of the behaviors in the daytime. I also found that adult females grooming infants and juveniles were greater than received from them (p < 0.001). The social grooming among adult females occurred mainly during non-mating seasons and its frequency was 2.12 times of mating seasons. Regardless of mating or non-mating seasons, the frequency of grooming among related females was significantly higher than among unrelated females (both p < 0.001). In addition, kinship affected the grooming frequency among female macaques with relative lower or higher ranking females during mating seasons (p < 0.05), while the dominant rank did not have the effect. However, the highest grooming frequency occurred in the high-ranking females who groomed relative lower ranking females within relatives (0.38 ± 0.40 bouts / 100 scans, n = 7). During non-mating seasons, kinship affected the grooming frequency among female macaques with relative lower ranking females (p < 0.05); the highest grooming frequency occurred in the old females who groomed relative lower ranking females within relatives (1.57 ± 1.74 bouts / 100 scans, n = 8).
Moreover, seasons (mating or non-mating) and kinship relationship had significant effects on grooming frequencies among female macaques (both giving and receiving p < 0.01), as well as on the grooming frequency of females groomed with relative lower ranking females (p < 0.01). The highest grooming frequency occurred on females groomed with relative lower ranking females within relatives during non-mating seasons (1.25 ± 1.48 bouts / 100 scans, n = 11).
Without kinship relationship, low-ranking females groomed relative higher ranking females more frequent than high-ranking females did (p < 0.05). Within adult females, 65% of social grooming was among relatives. However, 40% of adult females groomed equally with related and unrelated females, while 20% disproportionately groomed more with unrelated females than with related females. The grooming was kin-biased for 40%. About 8.68% of social grooming among unrelated females was being reciprocated.
On the other hand, the ratio of related grooming female partners to the total number of available related females was higher than that with unrelated females (p < 0.05). It also indicated that the ratio of each female received grooming from high-ranking grooming partners was higher than that from low-ranking females (p < 0.01).On the other hand, the ratio of the number of old or young grooming partners of adult females had similar values. The grooming frequency of adult females gave or received from adult males during mating seasons was higher than non-mating seasons (both p < 0.05) while high-rank females groomed adult males more than low-rank females did.
The socionomic sex ratios of these two social groups were similar during mating seasons (AM:AF = 1:1.8). Both troop and periphery males had significantly higher frequency of social grooming with adult females in mating seasons than in non-mating seasons. The types of males and social groups had significant effects on the allogrooming frequency among adult males in mating seasons (p < 0.05) but not in non-mating seasons (p > 0.1). Moreover, troop males had higher grooming partners than periphery males. Major grooming partners of troop males were adult females regardless of the seasons. Subordinate males were mostly the receivers in the grooming dyads with dominant males in the mating seasons, but the relationships changed during non-mating seasons. Agonistic interactions occurred mainly during mating seasons and its frequency among periphery males was 1.8 times of troop males (p < 0.05).
The preference grooming sites between allo-grooming and auto-grooming of C and Cd groups had varied significantly (p < 0.001). The back region was the preferred grooming site in allo-grooming of C and Cd group and the ano-genital region was the least groomed site. In auto-grooming, monkeys paid much attention to the legs ignoring the back and face. The result indicated that when the adult individuals groomed the head, back and face which showed significant difference in the frequency among AM and AF (p < 0.05). The related female adults groomed head more frequently than unrelated female adults (p < 0.05), but dominance rank and age-class did not apparently affect the corresponding values for the frequency of grooming sites among adult females (both p > 0.1).
The results indicated that social grooming among adult females took place more often during non-mating seasons, and more often in kin-related females than unrelated females. Moreover, the dominant females were likely to groom related females. Therefore social grooming among kin-related females may reinforce relationships while reciprocal grooming of unrelated females may serve to form alliance or ranking promotion in the social group. On the other hand, social grooming between adult male and female macaques more frequent in mating seasons than in non-mating seasons. This showed that adult male Formosan macaques employed complex strategies to achieve reproductive success. Nonetheless, the periphery males had more male grooming partners than troop males did which seemingly to enhance male coalitions.
目錄
誌謝......................................................................................i
摘要.....................................................................................iii
Abstract...............................................................................vi
目錄......................................................................................x
前言……………....................…………………….............1
材料與方法……………………………………….............6
一、研究地點…...…………..…………………….……...6
二、研究對象.....................................................................6
三、觀察記錄…....……………………………….……....8
四、行為定義………...………………………….….……9
五、資料分析………………………………………..…..12
結果….……………………………………..………….…21
一、社群成員變化........................................................21
二、社群成員與理毛行為............................................22
三、成年雌猴之社會理毛............................................23
四、成年雄猴之社會行為............................................34
五、獼猴理毛部位分佈................................................38
討論....................................................................................42
一、社群成員活動性之比較........................................42
二、成年雌猴之社會理毛............................................43
三、成年雄猴的社會行為............................................52
四、獼猴理毛部位之分佈............................................61
五、結論........................................................................63
參考文獻….……………..……………………..…….…..66
表........................................................................................80
圖…...……………………………………….……..........108
附錄..................................................................................135
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