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研究生:林芷頡
研究生(外文):Chih-chieh Lin
論文名稱:奧氏後相手蟹降海釋幼之導向行為機制研究
論文名稱(外文):Orientation mechanisms of larval release migration by the sesarmid crab, Metasesarma aubryi
指導教授:劉烘昌劉烘昌引用關係黃興倬黃興倬引用關係
指導教授(外文):Hung-chang LiuHsin-drow Huang
學位類別:碩士
校院名稱:靜宜大學
系所名稱:生態學研究所
學門:生命科學學門
學類:生態學類
論文種類:學術論文
論文出版年:2010
畢業學年度:98
語文別:中文
論文頁數:58
中文關鍵詞:遷移定向化學性訊號物理性訊號抱卵陸蟹
外文關鍵詞:orientationphysical cueschemical cuesmigrationovigerous land crab
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許多動物善用存在環境中的訊號,作為導向遷移之依據。陸蟹在其生活史中,必須為了繁殖返回海洋釋放幼蟲。長距離遷移使其易暴露於危險環境,亦較容易受到人類活動的干擾,進而影響陸蟹的生存繁衍。本研究旨在探討陸蟹降海釋幼遷移可能的導向機制,以恆春半島目前族群數量最多的奧氏後相手蟹(Metasesarma aubryi)為材料,進行野外行為觀察與實驗室行為實驗。在實驗室中,螃蟹被放置在直徑 1 公尺的圓盤中進行物理訊號(光照)的導向反應實驗;在四選一的化學選擇裝置中,分別測試螃蟹對三種化學訊號(淡水、海水、和同類氣味)的導向反應。物理實驗數據以圓形統計法(circular statistics)測試導向的顯著性;化學實驗數據則以比例檢驗(proportion test)進行差異比較。

野外觀察發現奧氏後相手蟹的繁殖期取決於每年雨季開始和結束的時間,抱卵雌蟹通常於月亮盈虧的最後四分之一週期間降海,在黎明前集體於海中釋放幼蟲。行為實驗結果顯示:在沒有提供任何採集地點熟悉訊號的黑暗環境中,抱卵雌蟹仍然具有顯著朝海方向移動的反應,釋幼後雌蟹呈現隨機移動或靜止,且釋幼後可能並不立即折返;面對不同方向的聚光刺激,抱卵雌蟹呈現顯著的趨光反應。化學訊號實驗則顯示奧氏後相手蟹抱卵雌蟹明顯受到淡、海水的氣味所吸引,釋幼後雌蟹受海水氣味吸引的比率明顯降低,且不以海水氣味為訊息,判別背海移動方向。同類氣味對奧氏後相手蟹則不具有吸引力。

綜合各項研究結果顯示,奧氏後相手蟹抱卵雌蟹在降海遷移的過程中,其導向行為不僅受到釋幼的生理需求所驅策,同時也受陸生適應壓力所牽制,面對不同生理階段的優先需求差異,其導向行為也隨之做出不同的選擇判斷。奧氏後相手蟹抱卵雌蟹在降海遷移前期,傾向選擇基質潮濕或富含水氣的路徑進行遷移,以減低降海遷移途中的水分喪失;在降海遷移後期,利用地磁訊號作為基本的導航依據,同時整合天光和海水氣味作為識別海岸方向的導向訊號;而釋幼後雌蟹演化出較長的回遷時間並且發展出延後回遷的行為,來增加回遷個體的存活率。途中的人造設施,明顯會對遷移中的雌蟹造成定向的影響。
Animals employ environmental cues for orientation in migration. In their life history, land crabs have to migrate to the sea for larval release . Larval release migration renders them exposed in dangerous environments and they are likely to expose themselves to dangerous environment and tend to be disturbed by human activities during their migration and ultimately, which would impact their survival and reproduction.
The aim of this study is to investigate the orientation mechanisms of larval release of the land crabs Metasesarma aubryi, the most abundant land crab species in Hengchun Peninsula. It was hypothesized that M. aubryi use both physical and chemical environmental cues for orientation. In laboratory, crabs were tested in a circular arena for orientation mediated by different illumination conditions; a choice chamber apparatus was employed for testing chemical cues from freshwater, seawater, and conspecifics. The data were analyzed by circular statistics for orientation significance and compared by proportion test for differences.
In the field, we observed that the breeding season of Metasesarma aubryi is variable, depending on the beginning and ending of the rainy season, and ovigerous females release larvae during the last quarter of the lunar cycle. Migration of mass ovigerous M. aubryi commences after midnight from the highland forest; crabs waits until the time before dawn for larval release. In laboratory, ovigerous crabs oriented to the sea in the dark. After larval release, however, female crabs would rest or move randomly, and they may not move back to land immediately. Both in field and in laboratory, ovigerous crabs were attracted to spot lights from directions. The scents of freshwater and seawater showed obvious attraction to ovigerous crabs, but showed less influence after larval release. After larval release, females did not tell the direction away the sea by the scent of seawater. The scent of conspecifics did not play roles in the orientation, while the scent did not appeal to the scent of other congeners.
The orientation of Metasesarma aubryi is not only prompted by its physiological drives for larval release, but also restrained by the pressure of terrestrial adaptation, their physiological needs influenced by the environments would determine the priority in orientation. M. aubryi tends to choose damp substrate or humid route for migration to reduce water loss during the early stage of larval release migration. When close to the littoral zone, ovigerous crabs may use geomagnetic fields to navigate, and integrate signals of skylight before dawn and the scent of seawater for seaward orientation. To increase the survival of the individual, female crabs evolve a longer period to return and delay the time to move inland after larval release. Moreover, artificial illumination en route cause negative effect on migrating crabs.
摘要……IV
Abstract……VI
表圖目錄……XI
謝誌……… XII


1. 前言……1
1.1 遷移的定義與生態意義……1
1.2 定向與遷移的訊號利用……2
1.2.1 物理性環境訊號……3
1.2.1.1 視覺訊號……3
1.2.1.2 地磁……4
1.2.1.3 坡度與環境基質觸感……5
1.2.1.4 海流與潮汐……5
1.2.1.5 振動或聲音……5
1.2.2 化學性環境訊號……6
1.2.3 多重訊號的整合與利用……6
1.3 陸蟹……7
1.4 陸蟹的繁殖遷移……9
1.5 奧氏後相手蟹分類與生態簡……10
1.6 研究目的……12

2. 材料與方法……13
2.1 實驗前的田野觀察……13
2.2 實驗動物採集與操作……13
2.2.1 採集樣區……13
2.2.2 動物採集與實驗前準備……14
2.2.3 實驗時間的選定……15
2.3 物理性環境因子導向行為實驗……15
2.3.1 實驗假說……15
2.3.2 研究方法……15
2.3.3 資料分析……16
2.4 化學性環境因子導向行為實驗……16
2.4.1 實驗假說……16
2.4.2 研究方法……17
2.4.3 資料分析……17

3. 結果……19
3.1 野外行為觀察……19
3.1.1 公路抱卵雌蟹……19
3.1.2 海岸抱卵雌蟹……20
3.1.3 海岸釋幼後雌蟹……20
3.1.4 雄蟹……21
3.2 物理性環境因子導向行為實驗……21
3.2.1 公路抱卵雌蟹……21
3.2.2 海岸抱卵雌蟹……22
3.2.3 海岸釋幼後雌蟹……22
3.2.4 雄蟹……23
3.3 化學性環境因子導向行為實驗……23
3.3.1 公路抱卵雌蟹……24
3.3.2 海岸抱卵雌蟹……25
3.3.3 海岸釋幼後雌蟹……27

4. 討論……31
4.1 物理性環境因子導向行為實驗……31
4.1.1 公路抱卵雌蟹的遷移行為與導向……31
4.1.2 海岸抱卵雌蟹的遷移行為與導向-光照的影響……32
4.1.3 人工照明對抱卵雌蟹的影響……33
4.1.4 雌蟹的掠食者迴避反應……33
4.1.5 繁殖季雄蟹的行為……34
4.2 化學環境因子導向行為實驗……34
4.2.1 雌蟹在化學選擇實驗裝置中的行為……35
4.2.2 淡水氣味對遷移雌蟹的影響……35
4.2.3 海水氣味對釋幼雌蟹的影響……36
4.2.4 同類氣味的效應……37
4.3 釋幼後雌蟹回遷機制的推測……37
4.4 結語……39
引用文獻……40
表……45
圖……49
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