Please use this identifier to cite or link to this item: http://hdl.handle.net/11455/89113
標題: Host-plant diversity and systematics of oak gallwasps (Hymenoptera: Cynipidae) in Taiwan
台灣橡樹癭蜂(膜翅目:癭蜂科)之寄主多樣性與系統分類
作者: 唐昌廸
Chang-Ti Tang
關鍵字: oak gallwasps;Fagaceae;oak;Quercus;gall;cyclical parthenogenesis;Asia;taxonomy;evolution;Dryocosmus kuriphilus;橡樹癭蜂;殼斗科;橡樹;麻櫟屬;蟲癭;循環孤雌生殖;亞洲;分類;演化;板栗癭蜂
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摘要: 
Oak gallwasps lie in the tribe Cynipini of Cynipidae, with about 1,000 known species and predominantly distribute in temperate zone of north hemisphere. The vast majority species of oak gallwasps induce galls on oaks (Quercus subgenus Quercus) of Fagaceae, and possess cyclical parthenogenesis in life cycles that alternate between sexual generation and asexual generation. A portion of species the generation alternation even involving host alternation. Galls of oak gallwasps are complex and diverse and are species-specific or generation-specific. This character add difficulties in pairing of different generations and taxonomic treatment. Europe and North America are the long-standing centers of oak gallwasp faunal studies, and most species were recorded from the two regions. In contrast, the Asian fauna was little studied and just 34 valid specie on Quercus subgenus Quercus recorded from Japan and Russian Far East. However, recent studies in Taiwan and Japan found that genera Castanopsis, Lithocarpus, and Quercus subgenus Cyclobalanopsis of Fagaceae endemic in Asia are also potential host-plants of oak gallwasps, suggesting that global richness of oak gallwasps is much diverse than previously known. The island Taiwan is situated at southeast Asia and harbors 44 native species in four genera. In addition, there are two cultivated species in the genus Castanea of Fagaceae. Thus, the known diversity of Fagaceae in Taiwan reflects the diversity of Asian Fagaceae. Establishment of oak gallwasp taxonomy and biology in Taiwan will be conducive to clarifying diversity and host-plant ranges of oak gallwasps in Asia, and complement the key information that has been long-scarce in global gallwasp fauna. This study sampled all plant Fagaceous species in Taiwan and extensively surveyed gall flora of oak gallwasps. The following results were achieved by collaborated with European researchers: (i) Castanopsis, Lithocarpus, and Quercus subgenus Cyclobalanopsis are new host-plant records to oak gallwasps; (ii) one hundred and fifteen gall morphotypes were recorded from 28 plant species in four genera, and more than half types were from Quercus subgenus Cyclobalanopsis; (iii) galls show organ specificity on their host-plants; (iv) forty-four oak gallwasp adult morphospecies were obtained from the 115 gall morphotypes. By specimen examination and DNA barcoding (the mitochondrial gene cytochrome b) to identify species, 17 new species (include two new genera) and one new record were described and published. Remaining oak gallwasp morphospecies are under descriptions; (v) analyses of DNA barcoding sequences has paired the sexual generation and asexual generation of the new record Cerroneuroterus vonkuenburgi (Dettmer), while the rest remain only sexual generation (35 spp.) or asexual generation (7 spp.) were recorded. Samples for DNA barcoding have been preserved for future analyses by sequences to pair the different generations of known species; (vi) inclusion of the host-plant records from this study and previous references, range of oak gallwasp host-plants increased from four Fagaceous genera to six genera. During the study period, damages of a new invasive pest Dryocosmus kuriphilus on chestnuts were recorded in Chungpu Township, Chiayi County. In order to propose appropriate strategies for management of this new pest in Taiwan, investigation of biology was conducted at chestnut orchards in Chiayi County. The established information of gall development and phenology of host-plants will serve as a reference for pesticide control. Because of the results in this study, some point of views in the past are challenged and waiting for further investigations. For example: from the point of view of Systematics, extant Western Palaearctic oak gallwasps were extremely conservative to sections Cerris and Quercus s.s. oaks in the history of diversification. These two oak sections support discrete groups of gallwasps and forming two monophyletic groups. Results of morphology-based taxonomy in this study, however, found that congeneric oak gallwasps associate with multiple host-plant groups. Whether congeneric oak gallwasps that associate various host-plant groups is a monophyly need further tests. Reconstruction of a phylogeny by mitochondrial cytochrome b gene included oak gallwasps from the Western Palaearctic, nominated species in this study, and additional unnamed species from Taiwan and China. Result revealed that morphologically congeneric species use multiple host-plant groups were polyphyletic and convergence in morphology rather than forming a monophyly. In addition, most gallwasp lineages induce galls on ancestral Castanopsis, Lithocarpus, and Quercus subgenus Cyclobalanopsis are basal to the phylogeny, with derived lineages on Quercus subgenus Quercus. However, support to these clades about above relationships were low. Multiple gene fragments should be adopted to further test the oak gallwasp phylogeny and evolution of oak gallwasp host associations. Taken together, diversity of oak gallwasps in Asia is rich and waiting for exploration. This is important for understanding of global oak gallwasp richness. The achievement of this study confirmed that endemic Fagaceae in Asia harbors oak gallwasps and provides a piece of key puzzle in understanding of the evolution of global oak gallwasp diversity.

橡樹癭蜂(oak gallwasps) 屬於癭蜂科橡樹癭蜂族(Cynipidae: Cynipini),主要分布於北溫帶,約1000 種已知種,絕大部分的物種以殼斗科(Fagaceae)之橡樹(Quercus subgenus Quercus,麻櫟屬麻櫟亞屬)為造癭寄主,以循環孤雌生殖進行有性世代與無性世代之交替完成生活史,部分物種的世代交替更涉及寄主轉換。橡樹癭蜂之蟲癭形態複雜多樣,且不同癭蜂物種或世代之蟲癭皆有其獨有的形態,增添不同世代的連結與分類處理的困難度。往昔的橡樹癭蜂相研究主要在歐洲(西舊北區)與北美洲(新北區),因此先前所知的物種紀錄大部分來自此兩大地理區,相較之下,亞洲因缺乏相關研究,過去僅有34種來自日本與蘇聯遠東地區於麻櫟屬麻櫟亞屬造癭的物種紀錄,然而近年來在台灣與日本的相關研究發現亞洲固有之苦櫧屬(Castanopsis)、石櫟屬(Lithocarpus)與麻櫟屬青剛櫟亞屬(Quercus subgenus Cyclobalanopsis)的殼斗科植物也是橡樹癭蜂的潛在寄主,顯示全球的橡樹癭蜂物種多樣性應較以往所知更為豐富。臺灣島位處東南亞,島內之殼斗科植物相包含4個屬(Castanopsis, Fagus, Lithocarpus, Quercus)共44種原生種,另有1屬(Castanea) 2種的栽培種,則台灣已知的殼斗科物種在屬級的豐富度可反映出亞洲殼斗科之多樣性,建立台灣本島之橡樹癭蜂分類與生物學資訊,有助於釐清橡樹癭蜂於亞洲的多樣性與寄主範圍,並增補全球橡樹癭蜂相長期缺乏之關鍵資訊。本研究對台灣全島所有殼斗科植物進行廣泛的癭蜂蟲癭相調查與採集,並與歐洲學者合作進行分類研究探討,取得以下之成果:(1)確認苦櫧屬、石櫟屬與麻櫟屬青剛櫟亞屬為橡樹癭蜂的新記錄寄主;(2)在4屬28 種植物共記錄115種癭蜂蟲癭形態種,超過半數的蟲癭記錄自麻櫟屬青剛櫟亞屬;(3)蟲癭對著生之寄主器官具專一性;(4)從前項樣品中取得44 種橡樹癭蜂成蟲形態種,由標本形態檢查與生命條碼(粒線體DNA之cytb基因)鑑定物種,確認17新種(含2新屬)與1新紀錄種之分類狀態並描述發表,其餘癭蜂形態種刻正著手處理描述中;(5)生命條碼之序列分析與生活史研究已連結新紀錄種癭蜂Cerroneuroterus vonkuenburgi (Dettmer)之有性世代與無性世代,其餘僅記錄有性世代(35種)或無性世代(7種)之癭蜂形態種,也已保存可供生命條碼鑑定的樣品,未來可使用DNA 序列就已知種進行不同世代之配對;(6)整合本研究的調查資料與文獻記載,全球橡樹癭蜂的寄主由先前的4個屬增加至6 個屬。研究期間亦發現新發生之入侵種害蟲板栗癭蜂,在嘉義縣中埔鄉板栗園造成危害,為研擬適合台灣防治此新發生害蟲之策略,於嘉義縣之板栗園進行板栗癭蜂生物學研究,建立蟲癭發生與寄主物候之基礎資料,做為藥劑防治試驗之參考依據。研究結果所顯現事證亦挑戰過去已知的多項看法,提供進一步驗證的見解,包括:就系統發生之觀點,現生西舊北區橡樹癭蜂於分化歷史中對sections Cerris與Quercus s.s.之寄主有高度的保守性,兩類寄主植物支持離散且各自成一單源群之癭蜂類群,然而台灣的癭蜂形態分類之研究結果發現同屬癭蜂於多個寄主類群造癭之紀錄,這些在不同寄主類群造癭的同屬癭蜂是否在親緣關係中成一單源群,有待進一步檢測;藉由粒線體DNA之cytb基因重建西舊北區、本研究已命名之物種、以及來自台灣與中國未定名的橡樹癭蜂樣本之物種的親緣關係,結果顯示形態分類屬於同一屬且利用多類寄主之癭蜂並不屬於一個單源群,而是多次起源且在形態上有趨同演化之現象。此外,在古老之石櫟屬、苦櫧屬與青剛櫟亞屬造癭且分布於亞洲之癭蜂大多數屬於親緣關係中的基群,於麻櫟亞屬造癭之類群為衍生之類群。然而,呈現上述演化結果之演化枝在親緣關係樹中的支持度並不高,未來應採用多段基因進一步驗證橡樹癭蜂之親緣關係以及寄主關係之演化。總括來說,亞洲擁有豐富且待發掘的橡樹癭蜂多樣性,對瞭解全球橡樹癭蜂多樣性至關重要,本研究確認亞洲固有之殼斗科為橡樹癭蜂寄主之成果,對探討全球橡樹癭蜂多樣性演化成因的議題,提供一塊關鍵拼圖。
URI: http://hdl.handle.net/11455/89113
其他識別: U0005-2508201501180900
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