以黃豹天蠶蛾屬為例 Evolutionary Plasticity and Functional Diversity of Eyespot Wing Pattern in Loepa Silkmoths

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以黃豹天蠶蛾屬為例 Evolutionary Plasticity and Functional Diversity of Eyespot Wing Pattern in Loepa Silkmoths 國立中山大學生物科學系 碩士論文 Department of Biological Sciences National Sun Yat-sen University Master Thesis 翅面眼紋之演化可塑性與功能多樣性- 以黃豹天蠶蛾屬為例 Evolutionary plasticity and functional diversity of eyespot wing pattern in Loepa silkmoths 研究生:蘇昱任 Yu-Jen Su 指導教授:顏聖紘 博士 Dr. Shen-Horn Yen 中華民國 106 年 7 月 July 2017 致 謝 非常感謝顏聖紘副教授,不辭辛勞地從我一開始入學對研究毫無頭緒,到制 定研究方向與題目、想法討論、方法的修改與問題指正,以及海報與論文等書面 修正等,一路指導教育我。 感謝小鳥們健康的活著陪我完成研究,沒有因為被我做的假餌嚇到而拒絕進 食或行為異常。 曾經在研究過程中提供給我幫助的朋友們:陳鍾瑋,引發我部份實驗設計的 構想。陳怡潔,提供我鳥類飼養與部分研究基礎認知。韋家軒,提出我實驗設計 上的缺失並點出我思考方向與研究中可能遭遇的問題與癥結。杜士豪、周育霆、 楊昕,協助我飼養鳥隻。非常謝謝你們的幫助。此外還有不少朋友們,在研究以 外的事務上提供我不少協助,使我能有更多心力專注於研究,感謝各位提供的幫 助,雖然無法將你們的名字在此一一列出,但你們的功勞我依然會銘記在心。 另外感謝一些商家與團體所提供無償的協助:中華民國生物奧林匹亞委員會, 借我使用攝影器材與印刷設備。社團法人高雄市野鳥協會,提供部分實驗用的鳥 隻。北站鳥園、五甲鳥藝坊、藍寶石鳥獸寵物大賣場,提供我不同鳥類飼養的技 巧與經驗分享。 最後我要在此鄭重感謝,高雄醫學大學生物醫學暨環境生物學系,謝寶森副 教授,以及國立中山大學生物科學系,黃淑萍助理教授。兩位老師願意抽空擔任 口試委員,並在短時間內閱讀與修改我的論文,提供我論文修訂上莫大的幫助。 iii 摘 要 眼紋(eye spot)是一種近圓形且具有同心圓環的斑紋。眼紋存在於許多動物體 表,目前最廣為人知的功能是防禦天敵。眼紋禦敵的機制包含威嚇天敵或誘導攻 擊失誤。黃豹天蠶蛾(Loepa)是天蠶蛾科(Saturniidae)下的一屬夜行性蛾類,其屬內 所有物種皆具有眼紋,但研究不如日行性的蛺蝶如此豐富透徹。黃豹天蠶蛾的翅 面斑紋變化不像蛺蝶那麼複雜,僅具有弧度或位置上些微差異,眼紋位置也相似, 推斷應有相同的禦敵機制,但卻在尺寸上有劇烈的變化,甚至有少數物種具有其 他斑紋使眼紋醒目程度下降或比眼紋更顯眼。這些變化與誠實訊號(honest signaling) 及威嚇理論有所牴觸,因此我們認為有必要釐清黃豹天蠶蛾眼紋的變化與其功能 性。首先我們分析了黃豹天蠶蛾所有物種的翅面紋路,歸納出翅紋的模式與少數 具其他斑紋的物種。然而因為黃豹天蠶蛾翅面具大面積黃色,鳥類可能會具有先 天色彩偏見,因此利用人工假餌檢測鳥類先天色彩偏見,之後才進行眼紋尺寸的 變化與其功能性差異的檢測。最後檢測部分具有其他斑紋的物種,這些斑紋是否 具有不同禦敵功效。結果顯示,黃豹天蠶蛾的眼紋效果確實與威嚇理論相符,但 是遭遇不同天敵時的效果有所差異。具有其他斑紋的少數物種在特定情況下反而 能呈現隱蔽作用,而比眼紋顯眼的斑紋則也有類似眼紋的威嚇作用。因此我們認 為在探討斑紋功能時,除了檢測現有功能假說外,還要考慮掠食者的認知多樣性 及共域物種的性狀,亦不可忽略其有功能多樣性的可能,如此才能更了解真實的 獵物-獵食者關係。 關鍵字:眼紋(Eye spot)、黃豹天蠶蛾(Loepa)、威嚇作用(Intimidating effect)、色彩 偏見(Color bias)、翅紋功能(Wing pattern function) iv Abstract The so-called “eye spot” refers to a peculiar color pattern which has a concentric structure. This pattern is widely present on external structure of a wide range of animal taxa, and its anticipated function is anti-predation, of which two conflicting hypotheses are involved, intimidation and deflection hypotheses. The majority of studies on the anti-predator function of eyespot focuses on diurnal butterflies of which the size of eyespots tend to be consistent within a genus. However, after having an overall examination of size, structural complexity, coloration, shape and contrast with background color of eyespots of all the known species of Loepa, I found that eyespots of Loepa have great interspecific variation, which is unusual for butterflies. Meanwhile, the fact that the eyespots of few species are not circular and their conspicuousness is weakened by the interruptive coloration raises a question whether they are still honest signals for anti-predation. First, I analyzed the wing pattern elements and variations of all species of Loepa. Second, prior to assessing the anti-predator function of the eyespots, I evaluated whether the avian predator had color and size bias toward the moths. I then conducted a series of experiment to evaluate the function of eyespots with a variety both in size, structure and conspicuousness. The results show that the function of eyespots of Loepa is conform to intimidation. The patches that makes eye spot less conspicuous can increase crypsis of the moth when situated in specific surrounding background. Furthermore, although the apical spot on forewing of Loepa oberthuri seems to be much more conspicuous than the discoidal eyespots, that eyespot did not show intimidating effect. In conclusion, I suggest to consider the variability of predator’s reaction toward the same prey and the phenotypic similarity among sympatric species when inferring the function of eyespots. Key words: Eye spot, Loepa, Intimidating effect, Color bias, Wing pattern function v 目 錄 論文審定書………………………………………………………… i 論文公開授權書…………………………………………………… ii 致謝………………………………………………………………… iii 中文摘要…………………………………………………………… vi 英文摘要…………………………………………………………… v 第一章 黃豹天蠶蛾屬(Loepa)之翅紋多樣性 1.1 摘要…………………………………………………………… 1 1.2 前言…………………………………………………………… 2 1.3 材料與方法…………………………………………………… 3 1.4 結果…………………………………………………………… 4 1.5 討論…………………………………………………………… 5 第二章 黃豹天蠶蛾屬的各種眼紋型式是否皆有威嚇效果? 2.1 摘要…………………………………………………………… 7 2.2 前言…………………………………………………………… 8 2.3 材料與方法………………………………………………… 11 2.4 結果…………………………………………………………… 13 2.5 討論…………………………………………………………… 14 第三章 暗帶黃豹天蠶蛾(Loepa obscuromaginata)的眼紋與 橙斑黃豹天蠶蛾(Loepa oberthuri)翅頂黑斑 具有威嚇功能嗎? 3.1 摘要…………………………………………………………… 16 3.2 前言…………………………………………………………… 17 3.3 材料與方法…………………………………………………… 19 3.4 結果…………………………………………………………… 20 3.5 討論…………………………………………………………… 21 參考文獻……………………………………………………………… 42 圖 次 圖 1-1 katinka-group 成蟲之翅紋…………………………………… 23 圖 1-2 miranda-group 成蟲之翅紋………………………………… 24 圖 1-3 oberthuri-group 成蟲之翅紋………………………………… 25 圖 1-4 眼紋測量位置圖…………………………………………… 25 圖 1-5 黃豹天蠶蛾眼紋各結構比對眼睛結構名稱……………… 26 圖 1-6 黃豹天蠶蛾屬翅面紋路模式圖…………………………… 26 圖 1-7 黃豹天蠶蛾翅面紋路……………………………………… 27 圖 1-8 翅基曲線斑紋變化………………………………………… 27 圖 1-9 miranda-group 親緣樹與眼紋寬度比、分布地區對應圖 28 圖 1-10 翅面具大面積非黃色斑塊之物種翅紋…………………… 29 圖 2-1 實驗一麵包蟲假餌調色示意圖…………………………… 30 圖 2-2 實驗一假餌翅面設計……………………………………… 30 圖 2-3 實驗二假餌設計與作用方式……………………………… 30 圖 2-4 實驗三假餌翅面設計……………………………………… 31 圖 2-5 實驗三假餌展示裝置……………………………………… 31 圖 2-6 實驗二攻擊前準備時間四分位數圖……………………… 32 圖 2-7 實驗三白腰鵲鴝對不同假餌之捕食頻率………………… 33 圖 2-8 實驗三白尾八哥對不同假餌之捕食頻率………………… 33 圖 2-9 實驗三白腰鵲鴝對不同假餌之攻擊前準備時間………… 34 圖 2-10 實驗三白尾八哥對不同假餌之攻擊前準備時間………… 34 圖 3-1 實驗一假餌裝置…………………………………………… 35 圖 3-2 實驗二假餌裝置…………………………………………… 36 圖 3-3 實驗一訓練與測驗結果…………………………………… 37 圖 3-4 實驗二裝置被攻擊率與鳥隻攻擊前準備時間…………… 37 表 次 表 1-1 黃豹天蠶蛾翅面紋路分析表……………………………… 38 表 1-2 黃豹天蠶蛾眼紋結構分析表……………………………… 40 第一章 黃豹天蠶蛾屬(Loepa)之翅紋多樣性 1.1 摘要 根據日行性蝶類的研究顯示相對醒目的眼紋通常被認為具有嚇阻天敵的作用,而 相對隱匿的眼紋則有助於降低被天敵偵測到的風險。眼紋在蛺蝶科被認為是一個 經歷過多次退化的祖徵,而眼紋在同一種蛺蝶上的變化則與季節有關。然而不具 備季節多態性的天蠶蛾的眼紋功能與演化趨勢卻很少被檢驗。大多數天蠶蛾的翅 盤(discal cell)斑紋在發育上是同源的,這個斑紋在部分物種小且不醒目,但在另一 些物種則是大而顯眼並形成眼紋與月眉紋。黃豹天蠶蛾(Loepa)位於翅盤的眼紋具 有明顯的尺寸種間差異,相較之下更適合做為檢測眼紋變異與禦敵功能差異的材 料。為了瞭解不同黃豹天蠶蛾物種之間,翅紋與眼紋的差異,同時便於後續眼紋 禦敵功能檢測之實驗設計,因此我們先針對黃豹天蠶蛾的眼紋尺寸、結構複雜, 以及翅面整體的紋路進行分析。紀錄黃豹天蠶蛾共有的斑紋樣式,並參照 Nymphalid Ground Plan(NGP)的建議繪製黃豹天蠶蛾屬的翅面紋路圖。隨後紀錄各 物種斑紋、色澤以及眼紋的差異,最後將具有較大差異的幾個物種單獨提出分析。 我們發現眼紋尺寸、複雜程度與翅基部曲線斑紋以及黃色色調的底色有一定的關 聯性。此外,有四個物種具有大面積非黃色斑塊出現在翅面,且其兩物種眼紋變 形,我們因此初步臆測其禦敵方式可能並非原先推測之威嚇禦敵模式。 關鍵字:黃豹天蠶蛾、眼紋結構、翅紋分析、禦敵機制、翅盤斑 1 1.2 前言 許多生物藉由展現具警戒性的視覺訊息降低受天敵捕食的風險。目前所知具有警 戒性的視覺訊息包含眼紋與警戒色,其作用方式雖然具有差異,但皆是藉由鮮明 顯眼的顏色使掠食者迴避捕食。眼紋存在於許多鱗翅目的翅面上,過去許多學者 針對眼紋所具有的功能提出不同假說,包含誘導天敵、融入背景以及種內競爭等 功能假說(Hingston 1933; Poulton 1890; Thayer 1909)。目前研究最多證據指出眼紋 確實有禦敵作用,不同眼紋的禦敵機制有所差異,如使天敵受到驚嚇以阻止獵物 被攻擊,或是誘導天敵攻擊翅緣藉以增加獵物脫逃機會(Blest 1957; Olofsson et al. 2010; Stevens et al. 2007; Stevens and Ruxton 2014)。 眼紋(eye spots)為蛺蝶科(Nymphlidae)的祖徵,且歷經多次的演化造就蛺蝶科眼紋 在尺寸與數量上的多樣化(Kodandaramaiah 2009; Monteior 2008)。此外蛺蝶科部分 物種的眼紋還具有季節性的個體發育差異,例如 Bicyclus 屬的潮濕季節羽化的成蟲 翅面邊緣會有許多眼紋,藉以誘導掠食者攻擊該位置,使其有更多的脫逃機會。 但在較乾燥的季節時,無眼紋的個體因環境改變而擁有更高的隱蔽性而能夠躲避 天敵攻擊(Brakefield and Larsen 1984; Lyytinen et al. 2004)。此外 Kodandaramaiah et al.(2013)在測量 Junonia 屬中各物種眼紋的尺寸與位置後,發現其變化的兩極端分 別符合威嚇天敵與偏離攻擊的眼紋模式。 天蠶蛾科(Saturniidae)翅面上位於翅盤的位置皆具有在發育上同源的斑紋,許多物 種的翅盤具有近似圓形且包含多個同心圓環結構的斑紋,此種形式的斑紋符合 Monteiro (2008)對於眼紋的定義:「近似圓形,且包含至少兩個以上同心圓的斑紋, 或是包含類似瞳孔斑點的單色圓形斑紋」。而另一些物種的翅盤斑紋則形成月眉紋 或其他推測有助於增加隱蔽性的斑紋。然而這些斑紋不會因季節變化而有個體發 育差異,相較於日行性的蛺蝶科眼紋而言穩定許多,但各屬之間除了位置與數量 2 之外,包含尺寸、顏色、複雜程度等都存在相當大的差異。 黃豹天蠶蛾(Loepa)是屬於蠶蛾總科(Bombycoidea)、天蠶蛾科(Saturniidae)下的一個 屬,目前已知種達 51 種,分布於亞洲,東亞至南亞各地諸如日本、台灣、中國、 菲律賓、爪哇以及蘇門答臘等地皆有黃豹天蠶蛾。黃豹天蠶蛾屬內目前可區分為 三 個 種 群 , 分 別 為 katinka-group 、 miranda-group 與 oberthuri-group ,其中 katinka-group 廣泛分布於中國、中南半島、印尼以及東亞島弧,包含日本、台灣與 菲律賓等地區;miranda-group 集中於亞洲大陸的山區,以及台灣也有一種; oberthuri-group 則主要分布於中國、越南與寮國等地區(Yen 2000)。 Loepa 屬內物種皆具有眼紋且位置皆位於翅盤,而眼紋尺寸卻有明顯種間差異,適 合用於做為眼紋功能研究材料,同時夜行性物種的眼紋研究並不如日行性物種如 蛺蝶來得如此豐富。然而我們發現 Loepa 屬內物種除了眼紋尺寸差異外,在眼紋 以外的翅紋部分仍具有些微差異,為了便於後續實驗設計,因此需要先分析 Loepa 屬內各物種的翅紋。 1.3 材料與方法 我首先以標本照片分析 Loepa 屬內已知的 51 個物種皆具有的翅紋,標本照片為 BOLD Systems v3 所提供的照片(圖 1-1 至 1-3),並參考過去學者提出過的 Nymphalid Ground Plan(NGP)繪製成一張翅面基礎圖(Monteiro 2008; Nijhout 2001)。 同時藉由文獻描述來區別種群(Naumann and Löffler 2012; Naumann et al. 2012; Yen 2000)。再將不同的斑紋樣式提出檢視,包含斑紋顏色差異、底色色澤差異等。 眼紋的位置因為皆位於翅盤故不針對眼紋中心位置與翅緣或翅基距離進行測量, 僅測量眼紋尺寸以及結構複雜程度。由於我們發現,黃豹天蠶蛾中眼紋的最小寬 3 度等同中脈橫脈寬度,而較大的眼紋會擴散至其他翅室,因此測量中脈兩側延伸 至眼紋邊緣為做為眼紋寬度,再將眼紋寬度除以自身中脈橫脈寬度作為標準化, 得到眼紋寬度比例(圖 1-4)。同時我們整合了 Naumann and Löffler (2012)與 Naumann et al. (2012)所提出黃豹天蠶蛾屬 miranda-group 的親緣關係假說後,將前後翅眼紋 尺寸比例分別對應至各物種,檢視眼紋尺寸比例是否在親緣關係上具有演化趨勢, 或者具有高度的可塑性。而眼紋結構則是計算不同色之同心圓環或弧線結構數量 (圖 1-5)。 1.4 結果 黃豹天蠶蛾翅面紋路變化較不如蛺蝶豐富,容易歸納出一組模式翅紋(圖 1-6)。所 有物種底色皆為黃色,其中 katinka-group 為金黃色,而 miranda-group 的黃色稍微 較淡。前翅前緣呈現深褐色。由翅基至外緣一共四條波曲線斑紋,依序為翅基曲 線斑紋(basal undulate line)、中央曲線斑紋(medial undulate line),以及後中線雙曲 線斑紋(postmedial double undulate lines)(圖 1-7)。多數物種前翅基曲線斑紋呈現紅 棕色且向翅基漸層,部分為紅色或黑色線條(圖 1-8)。後翅翅基曲線斑紋距離翅基 較前翅遠,且僅有下緣會有紅色與黑色之差異(圖 1-4)。亞外緣具有白色弦月型斑 紋周圍略呈深黃色。翅頂部具有一紅色斑塊,外緣較深紅,中間有一條白色曲線。 下方則有一黑色斑塊,該斑塊外緣具有白色弦月斑紋,廣義上可稱為眼紋,最外 緣的黑色曲線會連接至此眼紋,全屬物種此處斑塊變異小,唯獨 L. oberthuri 因後 中線雙曲線斑紋於此處較粗黑,而使翅頂黑色斑塊更為顯眼。 黃豹天蠶蛾的翅盤眼紋位於中脈橫脈上,寬度最小者與橫脈等寬,較大者會橫跨 兩側中脈,眼紋寬與中脈橫脈寬比由 1 至 2.58 不等(表 1-1)。當我們將眼紋尺寸比 例對應至整合後的親緣關係樹之後,發現眼紋並沒有明顯由小至大或由大至小這 類的演化趨勢,反而具有高度的可塑性(圖 1-9)。 4 結構最複雜者包含 6 個環狀或弦月斑紋,最簡單者為 4 個斑紋組成(表 1-2)。將斑 紋比對眼睛由外而內可分別稱為眼眶(edge)、鞏膜(sclera)、鞏膜光澤(scleral luster)、 白色弦月反光(reflective)、虹膜(iris)以及瞳孔(pupil)。眼眶、鞏膜與虹膜為環狀結 構,鞏膜近翅基處有鞏膜光澤與反光等兩道拱型或弦月型斑紋(圖 1-5)。結構簡單 者可能是因為缺乏眼眶(E)、鞏膜光澤(L)、虹膜(I)或瞳孔(P)等結構(表 1-2)。 其中有 4 物種翅面出現大面積非黃色斑紋(圖 1-10)。L. anthera 與 L. oberthuri 的前 翅下緣呈現紅棕色,但只有 L. oberthuri 後翅下緣亦有大面積紅棕色斑塊,且此兩 物種的眼紋並非圓形,而是外緣凹陷的腎形(圖 1-10A、B)。而 L. obscuromarginata 與 L. sinjaevi 則是前翅的黑色曲線向翅基部靠近,且第二曲線以外的外緣皆呈現深 褐色,後翅則是第三區線以外的外緣呈現深褐色,此兩物種的眼紋雖圓,但結構 屬於最簡單的形式(圖 1-10C、D)。 1.5 討論 黃豹天蠶蛾的眼紋寬度最小者與中脈橫脈等寬,因此我們認為黃豹天蠶蛾的眼紋 尺寸變異會受到體型的影響,因而決定以中脈橫脈寬度對眼紋進行標準化。且獵 物的體型也會影響潛在天敵的體型範圍,譬如說體型小的鳥類較不會去追捕體型 碩大的蛾類。因此在進行比較時,將眼紋寬度標準化後不僅能排除眼紋受體型的 影響,更進一步能控制潛在天敵的體型的範圍差異。 黃豹天蠶蛾翅面斑紋變較不如蛺蝶如此複雜,主要差異在眼紋尺寸與黑色曲線的 曲折程度(圖 1-1、1-2)。其中 katinka-group 是眼紋尺寸比例大的種群,眼紋結構複 雜程度也較高,且同時容易於翅基外緣曲線斑紋處呈現紅色且漸層的斑紋(圖1-1)。 這樣的趨勢與眼紋威嚇假說相符,越複雜且越大的眼紋會具有更好的威嚇效果 (Stevens et al. 2008c)。而 miranda-group 是翅面黃色色澤較淡的種群,其眼紋結構
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