水螟亞科是一個由700多種組成且大多數種類幼生期水生的分類群，廣布於全球除南極以外的各動物地理區，並在新熱帶區與東方區-澳洲區達到最高的多樣性。即使如此，該亞科仍被發現具陸棲幼生期的種類，而此陸生的起源可能說明了兩個截然不同的演化歷程：若此特徵是祖徵表示水螟是由陸地往水中發展，若為衍徵則相反。在具有陸棲幼生期的水螟中，分佈於東南亞地區、由8個種群約39種組成的河苔水螟屬是一個很獨特的例子。在過去，該屬幼蟲被認為全部生長於水域環境，然而近期研究卻發現幼生期陸生的種類。在鱗翅目甚至是所有完全變態類昆蟲中，除了河苔水螟，沒有其他案例在屬級尺度同時具有水陸兩棲的幼生期，而這樣的例子可能起因於非單系群。因此本研究以莎螟亞科的荸薺莎螟做為外群，利用成蟲、幼生期形態特徵與翅紋重建水螟亞科與河苔水螟的親緣關係，藉以探索「陸生事件在水螟亞科裡的演化過程」以及「駢系群或多系群是否為河苔水螟成為特例的原因」。內群從全球水螟亞科代表性屬中取樣屬的模式種或模式種之近似種共29屬38種，並選取河苔水螟8種群中的代表性種類26種。結果顯示陸棲水螟在水螟亞科中至少發生兩次，且都為演化後期出現的類群。河苔水螟屬為一單系群，鉤突基部側方的剛毛側方觀多列且背方觀重疊是重要的獨有衍徵之一；在僅使用成蟲特徵之加權樹中，其屬內關係分為水棲與陸棲兩大分支，抱器瓣遠體端末端的特化剛毛的形式、數量與產卵器長短等特徵的不同狀態各自支持這兩個分支的單系性。而產卵器長度與棲地、產卵策略之間的關聯性則是未來值得進一步探討的問題。研究結果顯示河苔水螟確實是完全變態類昆蟲中水陸兩棲的特例，惟以上關係皆未受到拔靴值支持且各分支的特徵大多為同塑特徵。

The Acentropinae represents a species-rich crambid group with more than 700 species distributed in all the zoogeographic regions except Antarctica, and the highest diversity occurs in the Oriental, Indo-Autralian and Neotropical regions. Most caterpillars of Acentropinae are widely adapted to feed on vegetation in different types of aquatic environments, such as ponds, streams and rivers. However, some of them are known to utilize mosses on land. Therefore the evolutionary relationships between the terrestrial and aquatic lineage become intriguing. The genus Paracymoriza Warren, 1890, is a moderate-sized acentropine genus comprising 39 species ranging throughout southeast and east Asia. Historically the genus was often confused with many unrelated genera due to the superficial resemblance in wing maculation, which is termed the “nymphuliform wing pattern”. The genus is currently divided into 8 species groups, with 5 of them known having aquatic immature stages, while the others having terrestrial immatures. The fact that Paracymoriza includes both truly aquatic and terrestrial species represents a unique case for the Lepidoptera, thus making the monophyly of the genus doubtful. In the present study, a phylogenetic hypothesis Paracymoriza was reconstructed by using morphological characters including wing pattern and immature stage characters obtained from 63 species representing 25 acentropine genera. The results suggest that Paracymoriza is monophyletic. The genus is composed of two major clades: one aquatic, and the other terrestrial. It suggests that Paracymoriza represents a unique example with amphibious immature stage in the Lepidoptera although the tree support values of this phylogenetic hypothesis remains relatively weak due to large amount of homoplasious characters.

目錄
一、前言 1
(一)為何對生物的「水生」或「陸生」適應與演化感興趣？ 1
(二)水生昆蟲對水域環境的適應 1
(三)水生鳞翅目的多樣性與分類分布 2
(四)水螟的多樣性、分布、特徵與重要性 2
(五)多樣化的造巢模式 3
(六)水螟亞科分類與演化歷史的混亂與發展 4
(七)陸生水螟的發現與其演化意涵 5
(八)水陸兩棲的河苔水螟 6
(九)本研究要探索的問題 7
二、研究材料與策略 8
(一)內群與外群的選擇 8
(二)樣本來源與備置 8
(三)形態學術語與特徵分析 9
(四)特徵編碼與矩陣編輯 25
(六)幼生期特徵與翅紋特徵對親緣關係重建效應之測試 27
三、結果 28
(一)利用所有特徵所重建之親緣關係 28
(二)成蟲特徵加幼生期特徵所重建之親緣關係 29
(三)成蟲特徵加翅紋所重建之親緣關係 29
(四)翅紋加幼生期特徵所重建之親緣關係 30
(五)成蟲特徵所重建之親緣關係 30
(六)僅利用翅紋特徵所重建之親緣關係 32
(七)僅利用幼生期特徵所重建之親緣關係 32
四、討論 33
(一)幼生期特徵作為水螟亞科的獨有衍徵仍有待評估 33
(二)幼生期陸棲在水螟亞科的演化格局上是後衍特徵 34
(三)完全變態類的昆蟲裡的特例─水陸兩棲的河苔水螟 34
(四)產卵器長短與產卵策略 36
(五)翅紋對水螟亞科親緣關係重建之影響 36
五、未來展望 38
(一)完整幼生期資訊以評估水螟亞科的單系性 38
(二)利用產卵器特徵推測其可能棲息地 38
(三)水域環境指標生物的發展潛力 38
六、參考文獻 39

圖次
圖1、過去學者所提出的水螟亞科(Acentropinae)與其它草螟亞科群之親緣關係假說。 49
圖2、水螟亞科屬級的兩個親緣關係假說。 50
圖3、四種不同形態的幼蟲。 51
圖4、不同形態的幼蟲之棲地及造巢模式。 52
圖5、臺灣產不同種類的水螟。 53
圖6、水螟亞科頭部側面構造圖。 54
圖7、水螟亞科翅脈圖。 55
圖8、Yoshiyasu(1985)所提出之翅紋模式示意圖與本研究中選用的河苔水螟屬成員。 56
圖9、本研究中選用的水螟亞科各屬成員。 57
圖10、本研究中選用的水螟亞科各屬成員。 58
圖11、本研究中選用的水螟亞科成員與後翅眼紋等翅紋特徵狀態。 59
圖12、水螟亞科雄性生殖器側面示意圖。 60
圖13、水螟亞科雄性生殖器構造與特徵狀態示意圖， 61
圖14、水螟亞科抱器瓣特徵狀態示意圖。 62
圖15、水螟亞科抱器瓣特徵狀態示意圖。 63
圖16、水螟亞科雌性生殖器構造與特徵狀態示意圖。 64
圖17、利用所有特徵並以全部分類群進行親緣關係重建後所得之未加權嚴格公同樹與加權樹。 65
圖18、利用成蟲加幼生期特徵並以全部分類群進行親緣關係重建後所得之未加權嚴格公同樹與加權樹。 66
圖19、利用成蟲特徵加翅紋並以全部分類群進行親緣關係重建後所得之未加權嚴格公同樹與加權樹。 67
圖20、利用翅紋加幼生期特徵並以全部分類群進行親緣關係重建後所得之未加權嚴格公同樹與加權樹。 68
圖21、僅利用成蟲特徵並以全部分類群進行親緣關係重建後所得之未加權嚴格公同樹。 69
圖22、僅利用成蟲特徵並以全部分類群進行親緣關係重建後所得之加權樹。 70
圖23、僅利用翅紋並以全部分類群進行親緣關係重建後所得之未加權嚴格公同樹與加權樹。 71
圖24、在使用不同特徵組的條件下，以僅具完整資訊的分類群進行親緣關係重建後所得之未加權嚴格公同樹。 72

表次與附錄
表1、本研究中所取樣之物種。 73
表2、利用不同特徵組與分類群組所重建之未加權親緣關係樹。 76
附錄1、重建水螟亞科內部親緣關係的全矩陣。 77
附錄2、利用成蟲特徵所重建並經加權處理後親緣關係樹上各節點之特徵變化。 81

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