大鉤蛾亞科為大蛾類中一個規模較小，分布於亞洲古北區與東方區的分類群，包含了分別置於Cyclidia Guenée, 1857以及Mimozethes Warren, 1901屬的13個物種。本亞科之幼蟲為山茱萸科八角楓屬的專食者，並具有狀似警戒的體表斑紋與聚集性。大鉤蛾亞科在過去研究中的分類地位並不穩定，由時被處理為獨立科，有時則被降為鉤蛾科的一個亞科。雖然目前已有兩個包含鉤蛾總科的分子親緣關係假說並支持鉤蛾科的單系性，然而大鉤蛾亞科與鉤蛾科的形態共衍徵尚未以支序系統分析方法進行檢測。本研究的目的為探索大鉤蛾亞科內部之系統發生以及與其它亞科之親緣關係。共選取16個大鉤蛾亞科的種級分類群(共13種，其中包含4亞種)，以及鉤蛾亞科(Drepaninae)之Drepana pallida nigromaculata與Oreta insignis、波紋蛾亞科(Thyatirinae)之Thyatira batis formosicola與Hypsidia屬的Hypsidia robinsoni與Hypsidia niphosema代表性物種做為內群；另由金蛾科(Cimeliidae)之Axia margarita與Epicimelia theresiae屬及兜蛾科(Doidae)之Doa ampla作為外群。我總共選取72個成蟲形態特徵(32二態、40多態)，並以最簡約法分別建構「包含翅紋」與「不包含翅紋」等兩種特徵包含方案所得之最簡約公同樹。結果顯示無論是何種方案下，大鉤蛾的兩個屬、各鉤蛾亞科與整個鉤蛾科的單系性均被支持，而大鉤蛾之姐妹群為((波紋蛾亞科+Hypsidia)+鉤蛾亞科)。雖然包含翅紋並未顯著提高大鉤蛾科內部的親緣關係解析度，然而這對大鉤蛾亞科內部份物種的親緣關係位置具有顯著的效應。此外本研究結果亦顯示許多由過去研究所倡議的大鉤蛾衍徵多為祖徵。

The subfamily Cyclidiinae is a small macroheteroceran group, which is endemic to the Oriental region and comprising 13 species in 2 genera, e.g. Cyclidia Guenée, 1857 and Mimozethes Warren, 1901. The larvae of the subfamily are specialists on Alangium (Cornaceae) and exhibit conspicuous colouration and gregariousness that may possibly function as aposematism. Historically this subfamily was inconsistently treated as either a separate family from Drepanidae, or lumped as a subordination of Drepanidae. Although both the two recent molecular phylogenetic hypotheses support the monophyly of Drepanidae, the morphological characters supporting the monophyletic status of Cyclidiinae and Drepanidae, however, have never been examined using modern methods. In the present study I investigated the phylogenetic relationships of Cyclidiinae based on morphological characters obtained from all the described species (13 species, including 4 subspecies) and the representatives of Drepaninae, Thyatirinae and Hypsidia. Cimeliidae and Doidae were chosen to be the outgroup taxa. In total 72 adult characters (32 binary, 40 multistate) were obtained and analyzed using Maximum Parsimony based on two character inclusion schemes (including or excluding wing pattern characters). The results showed the monophyletic status of the two Cyclidiinae genera, each subfamily of Drepanidae and the entire Drepanidae were recovered no matter which character inclusion scheme was adopted. The sistergroup of Cyclidiinae is confirmed to be ((Thyatirinae + Hypsidia) + Drepaninae). Although including wing pattern characters in the analysis does not significantly increase phylogenetic resolution within the Cyclidiinae, this scheme has great influence on the internal relationship of the subfamily. Meanwhile the present study reveals that most synapomorphies of Cyclidiinae proposed by previous studies are in fact plesiomorphic.

Contents
論文審定書 i
誌謝 ii
摘要 iii
Abstractiv
Contents vi
Index to Figures vii
Index to Tables ix
1. Introduction 1
1-1. The Drepanoidea: historical review 1
1-2. The Cyclidiinae: review of its systematic history 2
1-3. The questions to be addressed and aims of present study 3
2. Material and Methods 5
2-1. Taxon sampling & specimen acquisition 5
2-2. Morphological measurement and terminology 6
2-3. Character analysis 6
2-4. Character coding 27
2-5. Phylogenetic analyses 28
3. Results 29
3-1. Relationships between ingroup and outgroup taxa based on whole dataset 30
3-2. Relationships between ingroup and outgroup taxa based on deducted dataset 33
3-3. Testing previous hypotheses involving Cyclidiinae 36
4. Discussion 36
4-1. Monophyly and morphological synapomorphies of Drepanidae 36
4-2. Inter-relationships between major lineages of Drepanidae 37
4-3. Phylogenetic pattern of Cyclidiinae 38
4-4. Future context 40
5. Referenes 43
Appendices 76

Index to Figures
Figure 1. Figure 1. Eggs and larvae of Drepanidae and Doidae 52
Figure 2. Larvae of Drepaninae and Cyclidiinae 53
Figure 3. Pupal morphology of Drepaninae (A-B) and Cyclidiinae (C) 54
Figure 4. Previous phylogenetic hypotheses of the major lineages of Drepanoidea 55
Figure 5. Outgroups and ingroup taxa 56
Figure 6. Characters of frontoclypeus and labial palpus 57
Figure 7. Characters of thoracic sclerites and appendages (A-D) and zonation (E) 58
Figure 8. Wing venation of outgroup and ingroup taxa 59
Figure 9. Selected characters coded for wing patterns 60
Figure 10. Male scent organs, abdominal sclerites and tympanal organs 61
Figure 11. Male genitalia of outgroup and ingroup taxa 62
Figure 12. Male genitalia of Cyclidia 63
Figure 13. Male genitalia of Cyclidia 64
Figure 14. Female genitalia of Cyclidia 65
Figure 15. Genitalia of Mimozethes 66
Figure 16. Stylized drawings of selected characters of male genitalia 67
Figure 17. Strict consensus of the 12 MPTs based on the whole dataset (colour pattern characters included) under EW 68
Figure 18. The MPT based on the whole dataset (colour pattern characters included) under SAW 69
Figure 19. Strict consensus of the 120 MPTs based on the deducted dataset (colour pattern characters excluded) under EW 70
Figure 20. The MPT based on the deducted dataset (colour pattern characters excluded) under SAW 71

Index to tables
Table 1. Checklist of the 3 outgroup and 21 ingroup taxa included in the phylogenetic analysis. The cyclidiine species are listed in alphabetical order 72
Table 2. The phylogenetic hypotheses involving Cyclidiinae based on molecular evidence 73
Table 3. The contents of the topological constraints generated in PAUP* for testing the previous phylogenetic involving Drepanidae 74
Table 4. Comparison of the tree description using different datasets and weighting schemes 74
Table 5. Comparison between unconstrained trees and the trees obtained from each constraint analysis by using Templeton (1983) and Kishino-Hasegawa (1989) tests 75

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