臺灣博碩士論文加值系統

木蝨為一群吸食植物汁液之昆蟲且具有高度寄主專一性，以往研究除了分類學及防治，也常著重於寄主植物共同種化之議題上，木蝨聲音訊號對於雌雄間求偶行為扮演著重要的角色，雌雄蟲藉由聲學訊號進行物種辨識進而交尾，但過去木蝨聲學研究不多，因此本研究首先目標為建立木蝨聲學資料庫，共錄製並描述木蝨總科8科22屬46種的聲學訊號及行為，雌雄蟲會在求偶時建立對唱行為，通常由雄蟲發出呼喚聲，再由雌蟲發出回應聲，雄蟲憑藉著雌蟲的回應聲搜尋雌蟲所在位置並進一步交尾，這樣的對唱在木蝨求偶行為中非常重要，由獨立性檢定顯示對唱及交尾具顯著關聯性，木蝨訊號的種間變異大且種內雌雄蟲訊號具差異。
就木蝨聲學訊號是否具有種間差異並可進一步作為物種辨識的依據進行探討，以兩種Cacopsylla及兩種Trioza做為測試體系，結果顯示這些近緣種確實會以聲音訊號作為生殖隔離的依據，而同種雌雄蟲在求偶時經由觀察量化所得的配對分數顯著高於異種雌雄蟲，顯示同種雌雄蟲相對於異種雌雄蟲較易進入對唱、雄蟲搜尋及交尾的階段；至於植物介質如何影響聲學行為，針對木蝨在不同植物上的實驗結果顯示其訊號在非寄主上確實有改變，但木蝨雌雄蟲無論在寄主或非寄主上仍呈現相同的求偶行為且配對分數並無顯著差異，顯示這些植物影響不足以造成物種辨識，木蝨的物種辨識是以對唱行為所確立。
前述實驗顯示木蝨聲學訊號具種間差異且為物種辨識之依據，進一步探討木蝨聲學特徵在系統分類學上所具有的應用性，分析並比較四組不同科的同屬近緣種之聲音訊號，分別是Psyllidae的Cacopsylla屬五種、Homotomidae的Macrohomotoma屬兩種、Triozidae的Trioza屬兩種及Stenopsylla屬三種，結果皆顯示這些形態近緣的木蝨物種在聲學訊號上有分化的現象，五種Cacopsylla可以由唧聲數量、唧聲長度、唧聲間隔、長唧聲長度及主頻率依據判別分析而區分出來；兩種Macrohomotoma在唧聲特徵及主頻率上則依據主成分分析可區分，而兩種Trioza在唧聲長度上具有顯著差異且三種Stenopsylla也可由唧聲長度、主頻率、唧聲數量等特徵以主座標分析有所區別；在高階分類群的聲學特徵之歸群分析上，依據科級並無明顯的歸群現象而依屬級則同屬物種具歸群現象，顯示聲學特徵具有屬級特性，且應可有效應用於系統分類學，為木蝨分類系統之穩定性提供有用特徵。

Psylloids are plant sucking insects which possess high specificity on hosts. Previous studies of psylloids covered not only the basic taxonomy and pest control, but also extended to the issue of cospeciation with host-plant. Acoustic signals of psylloids play an important role in courtship behavior between males and females. They establish specific recognition via acoustic signals before mating. However, the acoustic related study of psylloids was scarce. This study attempts to establish the acoustic database of Psylloidea and has recorded and described acoustic signals and behavior among 46 species which belong to 22 genera of 8 families of Psylloidea. Males and females establish duet during courtship behavior. Usually, male emits the calling signal and female responds to it. Male then search for the female for mating based on response of female. Duet is important in courtship behavior of psylloids. According to the independence test, duet and mating have significant association. The interspecific variation of psylloid signals is high and different signal patterns were performed by males and females.
As the signals of psylloids possess species-specific differences, I further conducted experiments to explore whether acoustic signals function as specific recognition. Results of specific recognition showed that closely related species maintain reproductive isolation via acoustic behavior in both systems of two species of Cacopsylla and two species of Trioza. Based on the quantization of observation, mating scores of conspecific pairs were significantly higher than that of heterospecific pairs. It indicates that conspecific pairs are readily entering steps of duet, male searching, and mating; results of plant effects on acoustic behavior showed that acoustic signals have changed on nonhost plant. Nevertheless, males and females presented the typical courtship behavior and mating scores had no significant difference between different set of plant species. This suggests that plant effects did not affect specific recognition which was established by duet behavior.
Taking further on the result that signals of psylloids possess species-specific difference and work as specific recognition, I investigated whether acoustic characteristics have potential in applying to systematics of Psylloidea. This study further analyzed and compared the acoustic signals of four groups of closely related species belonging to different genera and families respectively. They are the five species of Cacopsylla (Psyllidae); the two species of Macrohomotoma (Homotomidae); and the two species of two Trioza and the three species of Stenopsylla (Triozidae). Results showed that acoustic divergence occurs in these morphologically similar species. The five species of Cacopsylla were distinguished by chirp number, chirp duration, chirp interval, trill duration, and dominant frequency based on discriminant analysis; the two species of Macrohomotoma were clarified by principle component analysis based on chirp characters and dominant frequency; chirp duration of the two species of Trioza also had significant difference and the three species of Stenopsylla were also determined by principle coordinate analysis based on chirp characters and dominant frequency. The ordination analysis of acoustic characters show that there is no clear grouping based on family category. However, it presents a clearer grouping on genus category. This phenomenon indicates that acoustics could be used as a generic characters and may facilitate the classification of Psylloidea.

摘要 i
Abstract ii
Contents iv
List of Figures vii
List of Tables xi
1. Overall Introduction 1
1.1 Psylloidea 2
1.2 Substrate-borne signals 3
1.3 Aims of this study 4
2. Duet behavior and acoustic diversity of Psylloidea 9
2.1 Introduction 10
2.2 Materials and methods 11
2.2.1 Preparation of insects 11
2.2.2 Acoustic recording and analysis 12
2.2.3 Statistical analysis 12
2.2.4 Acoustic Terminology 13
2.3 Results 14
2.3.1 Duet behavior of Psylloidea 15
2.3.2 Mating after duet cycle 21
2.3.3 Acoustic diversity of Psylloidea 22
2.3.4 Acoustic description of Psylloidea 29
Aphalaridae 29
Ctenarytaina sp. ‘Syzygium’ 29
Togepsylla takahashii Kuwayama Jr., 1931 29
Calophyidae 29
Calophya meliorata Yang, 1984 29
Cecidopsylla cata Fang & Yang, 1986 30
Carsidaridae 30
Mesohomotoma camphorae Kuwayama, 1908 30
Tenaphalara acutipennis Kuwayama, 1908 30
Homotomidae 31
Homotoma maculata (Yang, 1984) 31
Macrohomotoma gladiata Kuwayama, 1908 31
Macrohomotoma robusta Yang, 1984 31
Liviidae 32
Epipsylla sp. ‘Millettia’ 32
Paurocephala bifasciata Kuwayama, 1931 32
Paurocephala trematos Yang, Yang & Chao, 1986 32
Syntomoza sp. ‘Thespesia’ 33
Phacopteronidae 33
Cornegenapsylla euphoriae (Yang, 1984) 33
Psyllidae 33
Cacopsylla baphicacanthi (Yang, 1984) 33
Cacopsylla deflua (Yang, 1984) 34
Cacopsylla eriobotryacola (Yang, 1984) 34
Cacopsylla evodiae (Miyatake, 1965) 34
Cacopsylla multijuga (Yang, 1984) 35
Cacopsylla oluanpiensis (Yang, 1984) 35
Cacopsylla schefflerae (Yang, 1984) 36
Cacopsylla tobirae (Yang, 1984) 36
Insnesia maculosa Fang & Yang, 1986 37
Paraphyllura micheliae Yang, 1984 37
Psyllidae sp. ‘Eucalyptus’ 37
Triozidae 38
Baeoalitriozus yangi Li, 2011 38
Cecidotrioza epica (Yang, 1984) 38
Leptynoptera sulfurea Crawford, 1919 38
Neotrioza shuiliensis (Yang, 1984) 39
Pauropsylla triozoptera (Crawford, 1913) 39
Stenopsylla nigricornis Kuwayama, 1910 40
Stenopsylla occipitalis Yu, 1956 40
Stenopsylla sp. ‘Symplocos’ 40
Trioza acuminatissima Liao, Huang & Yang 41
Trioza beilschmiediae Yang, 1984 41
Trioza fausta Fang, 1990 41
Trioza neolitseacola Yang, 1984 42
Trioza outeiensis Yang, 1984 42
Trioza resupina Li & Yang, 1984 42
Trioza sozanica (Boselli, 1930) 43
Trioza valida Yang, 1984 43
Trioza sp. ‘Adinandra’ 43
Trioza sp. ‘Castanopsis’ 43
Trioza sp. ‘Cinnamomum’ 44
Trioza sp. ‘Helicia’ 44
Trioza sp. ‘Syzygium’ 45
2.3.5 Species without signals? 92
2.4 Discussion 92
3. Reproduction isolation via acoustic recognition in closely related species 96
3.1 Introduction 97
3.2 Materials and methods 99
3.2.1 Selected species 99
3.2.2 Plant effects 99
3.2.3 Specific recognition 101
3.3 Results 101
3.3.1 Plant effect 101
3.3.2 Specific recognition via duet behavior 110
3.4 Discussion 113
4. Application of acoustic characteristics on systematics of Psylloidea 121
4.1 Introduction 122
4.2 Materials and methods 123
4.2.1 Selected species 123
4.2.2 Acoustic characters 126
4.2.3 Statistical analysis 126
4.3 Results 126
4.3.1 Acoustic divergence of closely related species 126
4.3.2 Acoustic clustering of higher classification 147
4.4 Discussion 155
5. Conclusion 158
6. References cited 160
Appendix 1. Known taxa of insect using substrate-borne signals as mating signals 175
Appendix 2. Information concerning the psyllid species used in this study 200
Appendix 3. The Matlab script for signal plotting. 203

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