臺灣博碩士論文加值系統

喜楠瘿蚋屬(Daphnephila)是台灣闊葉林常見的癭蚋類群。這些喜樟瘿蚋屬的昆蟲常在相同寄主上造不同外形的蟲癭，並且共享相似的生態區位與資源。我使用寄主為紅楠的莖癭喜樟癭蚋、台灣喜樟癭蚋、淑燕喜樟癭蚋、窄癭喜樟癭蚋與鳥頭喜樟癭蚋的蟲癭作為主要材料，分別進行下列探討：(1)持續多年野外調查，瞭解使用相同資源的蟲癭是否有生活史差異；(2)使用穩定性碳同位素技術，測試蟲癭的營養適應假說；(3)進行蟲癭形態測量，測試蟲癭是否為造癭者的延伸表徵；(4)分離不同發育時期的蟲癭瘿室內的真菌；(5)觀察與造癭者相關的寄生蜂與食瘿者，瞭解蟲癭在該地生態環境中的上一階層消費者，建構初階的蟲癭群聚資訊。
主要結果如下：(1)在2004到2011年間觀察台灣陽明山地區大屯山脈兩側105棵紅楠上造葉瘿的四種喜樟瘿蚋屬族群，顯示這四種喜樟瘿蚋屬的成蟲有冬季與春季兩次羽化時間，分別對應寄主植物的兩次抽芽時期。喜樟瘿蚋屬的生活史可分為一年一世代與兩年一世代兩群，差異在於一齡滯育時期的延長。這樣的兩種生活史策略經常來自相同母蟲所產下的同批卵，顯示後代羽化的時間並不總是與母蟲一致。(2)穩定性同位素的分析顯示瘿組織的碳同位素高於寄主植物，因此當植物被昆蟲造瘿之後，會形成類似食物鏈關係的碳同位素累積。(3)根據蟲癭外形的長寬比值，紅楠上的喜樟瘿蚋屬的蟲癭可分為圓胖型(台灣喜樟癭蚋、淑燕喜樟癭蚋)與瘦長型(莖癭喜樟癭蚋、窄癭喜樟癭蚋、鳥頭喜樟癭蚋)。蟲癭內部的儲存細胞層數顯示圓胖型的蟲癭較瘦長型蟲癭多層，只有屬於瘦長型蟲癭的莖癭喜樟癭蚋蟲癭與圓胖型蟲癭更為相似。蟲癭組織部分保有寄主植物組織的特性，如在莖癭裡似輪限薄壁組織的薄壁組織與葉癭裡的維管束鞘。(4)根據蟲室分離的真菌，優勢真菌可分為成熟期蟲癭的B. dothidea與發育期蟲癭的Fusarium spp，並分布於台灣各地的喜樟瘿蚋屬蟲癭內，但是其他非優勢菌種的種類會隨地區不同而有所改變。(5)喜樟瘿蚋屬的寄生蜂主要是Platygaster sp. 與 Eupelmus sp.。我發現五種未曾紀錄過的野生動物食癭行為，分別是臺灣獼猴、臺灣藍鵲、赤腹松鼠與兩種夜蛾科的幼蟲。這五種野生動物平常就會取食紅楠的葉、芽與果實，推測食癭行為乃偶發性，但在冬季或早春食物稀少時會食瘿作為食物補充來源。
結論：(1) 這是第一個在紅楠多年枝條上確認造葉瘿的喜樟瘿蚋屬昆蟲生活史的研究，發現這四種昆蟲同步使用相同資源與有相似的生活史。(2) 穩定性碳同位素的結果顯示喜樟瘿蚋屬的演化適應是朝向較小蟲癭而投資越少能量於蟲癭的方向發展。(3)確認無論蟲癭形狀、蟲癭組織結構、蟲癭能量累積平均、真菌相均為造瘿昆蟲的表徵，並且顯示喜樟瘿蚋屬在紅楠上的適應性演化。(4) Botryosphaeria dothidea是主要的蟲癭內共生菌，但來自相同族群蟲癭內不同的真菌相顯示喜樟瘿蚋屬可以使用其他真菌作為次級共生菌。(5)此研究首度紀錄兩種特有種脊椎動物的食瘿行為。

Daphnephila is a common group of Cecidomyiidae in the evergreen broadleaf forests of Taiwan. Diverse galls induced by different species of Daphnephila often concurrently exist on the same host species demonstrating a striking niche divergence and resource sharing. The galls induced by D. truncicola, D. taiwanensis, D. sueyenae, D. stenocalia, and D. ornithocephala on Machilus thunbergii were examined : (1) to know whether the life cycle differs among various Daphnephila gall midge species accompanying the same host M. thunbergii; (2) to test the nutritional adaptation hypothesis by stable isotope; (3) to examined the galls to verify the hypothesis that the morphology of galls is an expression of the extended phenotype of the respective gall-inducing insect; (4) to establish the fungus flora in different gall midge development stage by isolating the fungi inside galls from the immature to mature stage; (5) to record the parasitoids and cecidophagous predators associated with Daphnephila galls.
Results are as following. (1) Populations of four leaf-gall inducing Daphnephila species were tracked on 105 M. thunbergii trees at four sites on Mt. Datun in Yangmingshan National Park in Northern Taiwan for seven years (2004–2011). Daphnephila populations recognized as either the winter or the spring emergent group could take advantages of two bud bursting seasons, either winter (January–February) or spring (March–May), based upon the availability of their oviposition time and location and they use either the vegetative leaves of the winter mixed buds with inflorescence and vegetative shoots or the spring vegetative buds producing no reproductive organs. Daphnephila populations show two types of life cycles: one-year type and two-year type, the latter with an extended diapause to the second year of the first-instar larva. Tracing the development of eggs from the same batches, I found the offspring did not always follow the life strategies of their parents. (2) Gall tissues had higher carbon isotope contents than the galled leaves. Consequently, a relocation of carbon isotope composition occurred when gall formed and a typical food chain relationship is revealed. (3) Based on their length―width ratio, the materials were grouped into either fleshy (those induced by D. taiwanensis and D. sueyenae) or slim (those induced by D. truncicola, D. stenocalia, and D. ornithocephala) types of galls. The numbers of reserve and nutritive cell layers in galls were greater in the stem galls induced by D. truncicola, similar to those in the fleshy leaf galls, but were fewer in other slim leaf galls. The gall tissues retained some property characteristic from plant tissues, such as the boundary-parenchyma-like parenchyma in stem galls and the bundle sheath in leaf galls. (4) Based on the fungal taxa isolated from the larval chambers, the dominant fungi in mature galls was B. dothidea while in immature galls were Fusarium spp. Regardless sampled localities, the above dominant status of fungal groups remained the same, whereas the non-dominant fungal taxa varied among different areas. (5) The major parasitoids are Platygaster sp. and Eupelmus sp. Five species of animals are newly recorded to be cecidophages on Daphnephila galls, i.e. the Formosan rockmonkey (Macaca cyclopis), the Taiwan blue magpie (Urocissa caerulea), the Formosan red-bellied tree squirrel (Callosciurus erythraeus), and two species of Noctuidae larvae. These animals are facultative cecidophages and usually feed on the leaves, buds or fruits of host plants. They selectively nibble at the gall tissues in winter or early spring, indicating that they may use galls as a supplemental food when their usual food resource is low.
In conclusion: (1) this is the first survey on the cohort of M. thunbergii associated with Daphnephila galls in Taiwan. Synchronizing utilization of the host resources and similar life cycles were found among the four leaf-galling Daphnephila species; (2) the results of stable carbon isotope suggests that the selection forces may toward smaller sizes for galls and decreasing carbon isotope contents with an optimal relationship; (3) the shapes, structure, nutritive tissues, energy levels, and multiple coexisting fungal taxa within galls reinforce that they are extended phenotypes of the respective gall-inducing Daphnephila species and they represent adaptive evolution of Daphnephila on M. thunbergii; (4) Botryosphaeria dothidea is the primary associate and possibly a symbiont. The different fungal taxa isolated from mature galls induced by the same Daphnephila populations suggest that these Cecidomyiidae may use different species of secondary fungal associates to possibly enhance their nutrition; and (5) this study newly recorded the cecidophages behavior of two endemic vertebrate.

1. General introduction 1
1.1. Gall maker and the organisms associated with galls 1
1.2. Gall midges and the tribe Asphondyliini 2
1.3. Gall midges and their hosts in Taiwan 5
1.3.1. Daphnephila 6
1.3.2. Machilus hosts 7
1.4. Galling Hypothesis 10
1.4. Long-term study area 11

2. Host phenology and life cycles of Daphnephila species (Diptera: Cecidomyiidae) on the leaves of Machilus thunbergii (Lauraceae) in Northern
Taiwan 14
2.1. Introduction 14
2.2. Materials and Methods 17
2.2.1. Study sites 17
2.2.2. Leaf longevity and host plant phenology 18
2.2.3. Observation of the life cycle of Daphnephila species 19
2.2.4. Life span of gall midges 20
2.3. Results 21
2.3.1. Host phenology and leaf age 21
2.3.2. Daphnephila life cycle 23
2.3.2.1. Matting and oviposition 23
2.3.2.2. First instar and difference of diapause 25
2.3.2.3. Mature gall and offspring emergence 26
2.3.2.4. The adult life span of Daphnephila 27
2.4. Discussion 27

3. Using stable isotopes to investigate the nutritional use of galling insects in Yangmingshan National Park 32
3.1. Introduction 32
3.1.1. Hypotheses of gall adaptation 32
3.1.2. Tracing the nutritional use 33
3.2. Material and methods 34
3.2.1. Study region and galls 34
3.2.2. Stable isotope analysis 35
3.3. Results 36
3.4. Discussion 40

4. Structure of five Daphnephila galls on Machilus thunbergii in northeastern
Taiwan 42
4.1. Introduction 42
4.2. Material and methods 45
4.3. Results 48
4.3.1. The structure of host tissues 48
4.3.2. The gall structure 50
4.3.2.1. Nutritive tissue 51
4.3.2.2. Sclerenchyma zone 55
4.3.2.3. Transitional ground tissue inside vascular zone 56
4.3.2.4. The structure and ratio at vascular zone 57
4.3.2.5. Cell composition of cortex 59
4.3.2.6. Epidermal characteristics of galls 60
4.4. Discussion 63

5. Associated fungi in Machilus host and Daphnephila galls 68
5.1. Introduction 68
5.2. Materials and methods 70
5.2.1. Study sites and sampling 70
5.2.2. Sampling and fungal isolation 73
5.2.3. Assessment of isolation frequency 74
5.3. Results 74
5.3.1. Isolation frequency of fungi varied in host plants
in different areas 74
5.3.2. Isolation frequency of fungi varied in galls at different
life cycle stages 76
5.3.3. Dominant fungi in D. taiwanensis galls 80
5.3.4. Phenological survey of host plant in different areas 82
5.4. Discussion 83

6. Other organisms associated with Daphnephila galls 86
6.1. Introduction 86
6.2. Materials and methods 87
6.3. Results 88
6.3.1. Associated parasitic wasps in YMSNP 88
6.3.2. Associated parasitic wasps throughout Taiwan 89
6.3.2.1. Parasitic wasps of the family Platygastridae 90
6.3.2.2. Parasitic wasps of the family Pteromalidae 91
6.3.2.3. Parasitic wasps of the family Eulophidae 91
6.3.2.4. Parasitic wasps of the family Braconidae 92
6.3.2.5. Parasitic wasps of the family Eupelmidae 93
6.3.2.6. Parasitic wasps of the family Torymidae 93
6.3.2.7. Parasitic wasps of the family Eurytomidae 93
6.3.2.8. Parasitic wasps of the family Ormyridae 94
6.3.2.9. Parasitic wasps of the family Ceraphronidae 94
6.3.2.10. Parasitic wasps of the family Encyrtidae 94
6.3.2.11. Parasitic wasps of the family Ichneumonidae 95
6.3.3. Cecidophagous animals 90
6.4. Discussion 99

7. Conclusion 101

8. References 104

9. Appendix 122

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