桉樹枝癭釉小蜂（Leptocybe invasa）是一種來自澳洲桉樹上的新興蟲害，雌蜂會將卵產於寄主的幼嫩組織，形成之蟲癭可供幼蟲營養及庇護處，對於被寄生的桉屬（Eucalyptus）植物的生長與發育影響甚鉅。過去對桉樹枝癭釉小蜂雌蟲的產卵選擇性，以及蟲癭組織發育過程的研究甚少。經由顯微觀察，有助於我們了解蟲癭組織的外觀形態與內部結構的改變。本實驗觀察比較三種對桉樹枝癭釉小蜂具不同抗性的桉屬植物上，蟲癭發育結構特徵的差異性，探究影響桉樹枝癭釉小蜂的產卵偏好性與蟲癭發育的原因。研究結果顯示，赤桉-玫瑰桉雜交品系（E. camaldulensis x E. grandis）和赤桉（E. camaldulensis）葉柄的蟲癭，蟲室外的厚壁細胞層的增生與較厚的皮層細胞層，推測應和幼蟲抵禦天敵的策略有關；成熟期蟲癭呈現顏色對比較強烈的紅色，或許是寄主植物用來吸引寄生蜂攻擊蟲癭的策略。雌蟲產卵偏好性的實驗中，尾葉桉被感染的比例較高，但赤桉-玫瑰桉雜交品系上之蟲癭敗育比例較低，後者可能與其表皮層較薄及單寧等次級代謝物的含量較低有關。本研究也發現三種桉樹在不同抽芽季節，具不同表型的幼葉。因其幼葉組織結構特徵的差異性，造成對桉樹枝癭釉小蜂的抗性有所不同。蟲癭在不同桉屬植物上的形態與內部結構相似，但在不同抗性的桉樹以及環境溫度條件下，桉樹枝癭釉小蜂的生活史周期有所差異。此篇研究以南台灣地區桉屬植物上的桉樹枝癭釉小蜂為對象，描述其蟲癭各個發育時期的特徵，有助於掌握桉樹枝癭釉小蜂的生長狀況，提前做蟲害防治上的因應。此外，富含抑制幼蟲生長之次級代謝物，或具不易遭雌蟲寄生之幼葉表型的桉樹品系，可提供抗病性樹種的選植，以降低桉樹枝癭釉小蜂的為害。

The blue gum chalcid, Leptocybe invasa (Hymenoptera: Eulophidae) is a gall-forming pest of Eucalyptus trees that originated from Australia. The gall provides nutrients and protection for the developing larvae, while it also causes growth retardation and esthetic damage of the host. This study focused on comparing the differences in morphology and histology on the development of L. invasa galls on three Eucalyptus species, invesgating female chalcid’s oviposition preference and the success rate of gall formation, and assessing their resistance to L. invasa. The results indicate sclerenchyma cell layer outside chamber and the formation of thicker cortex with tannin near infection site both correlates with larva defense mechanism. In the oviposition choice experiment, L. invasa prefer laying eggs on E. urophylla （U）, but the survival rate of larvae is higher on E. camaldulensis x E. grandis （CG） with thinner epidermis and less secondary compounds. However, different types of young leaves might be a factor relates to host’s resistance capability. L. invasa gall structure on different species is similar, but the gall development (especially at the pupa stage) pattern varies: except for hosts’ inter-species difference, L. invasa’s growth rate are likely to be temperature-dependent. This study describes L. invasa’s life cycle on Eucalyptus trees in Southern Taiwan, discussing host’s physical and chemical properties relates to their resistance to L. invasa, and providing information for pest management of L. invasa in enhancing insect control of Eucalyptus trees.

丁小余、施國新、陳維培、徐祥生。1993。茭白冬芽的發育及抗寒性的形態學研究。武漢植物學研究11(2)：104-112。
王維洋。1992。台灣桉樹病害調查報告。林業試驗所研究報告季刊7(2): 179-194。
甘偉航、胡大維、楊政川。1990。五種桉樹在東台地區造林適應性初步試驗。林業試驗所研究報告季刊5(2)：99-109。
吳耀軍、李德偉、常明山、黃華艷、陳尚文。2010a。桉樹枝癭姬小蜂生物學特性研究。中國森林病蟲29(5)：1-10。
吳耀軍、常明山、李德偉、羅基同、秦元麗、黃華艷、葉建仁。2010b。桉樹枝癭姬小蜂為害對桉樹縮合單寧含量的影響。南京林葉大學學報（自然科學版）34(6)： 96-100。
吳耀軍、奚福生、羅基同、李德偉、常明山。2009。桉樹枝癭姬小蜂生物學特性及控制技術。廣西植保22(2)：5-9。
馬英玲、韋春義、龐正轟。2009。桉樹枝癭姬小蜂的風險評估。廣西農業科學37(21)：10052-10053。
徐國華、陳維培、魏錦城、施國新。1999。莼菜冬芽越冬階段葉的解剖學和生理學研究。武漢植物學研究17(2)：97-100。
張四美、陳維培。1988。莼菜(Brasenia schreberi Gmel.)冬芽的形態結構與發育。武漢植物學研究6(1)：25-31。
陸長梅、吳國榮、周長芳、顧龔平。2001。懸鈴木越冬芽在冬季的生理適應。西北植物學報21(4)：650-655。
陳正豐、楊政川、張添榮、洪富文。1995。二十種桉樹類在台灣造林適應性試驗。林業試驗所研究報告季刊10(3)：283-292。
常潤磊、周旭東。2010。我國桉樹枝癭姬小蜂研究現狀。桉樹科技27(1)：75-78。
黃馨瑩、吳宜穗、董景生。2011。刺桐釉小蜂(Quadrastichus erythrinae Kim)的產卵選擇與造癭偏好。台灣昆蟲81：67-78。
楊曼妙。1999。造癭昆蟲生物學與進化。昆蟲分類及進化研討會專刊113-125。
董景生。2010。危害桉樹造林木的造癭釉小蜂。林業研究專訊17(2)：35-36。
董景生、楊恩誠、林雅玲。2011。桉樹枝癭釉小蜂緊急防治試驗。行政院農業委員會林務局委託研究計畫系列編號00-00-5-01。
趙丹陽、徐家雄、林明生、邱煥秀、鍾鎮奎、陳沐榮、陳瑞屏。2008。桉樹枝癭姬小蜂為害後桉樹損失量測定。廣東林業科技24(6)：58-60。
趙若帆。2010。香楠葉部的四種癭蚋蟲癭之形態比較。國立成功大學生命科學系碩士論文。
錢軍、羅湘粵、吳彪 。2009。桉樹枝癭姬小蜂在海南的發生與防治。熱帶林業37(1)：40-42。
Abrahamson, W. G., G. Melika, R. Scrafford, and G. Csóka. 1998. Gall-inducing insects provide insights into plant systematic relationships. American Journal of Botany 85: 1159-1165.
Arduin, M., G. W. Fernandes, and J. E. Kraus. 2005. Morphogenesis of galls induced by Baccharopelma dracunculifoliae (Hemiptera: Psyllidae) on Baccharis dracunculifolia (Asteraceae) leaves. Brazil journal of biology 65(4): 559-571.
Askew, R. R. 1961. On the biology of the inhabitants of oak galls of Cynipidae (Hymenptera) in Britain. Transactions of the society for British entomology 14: 237-268.
Austin, A. D., D. K.Yeates, G. Cassis, M. J. Fletcher., J. La Salle, J. F. Lawrence, P. B. McQuillan, L. A. Mound, D. J. Bickel, P. J. Gullan, D. F. Hales, and G. S. Taylor. 2004. Insects 'Down Under' - Diversity, endemism and evolution of the Australian insect fauna: examples from select orders. Australian Journal of Entomology 43: 216-234.
Bequaert, J. 1924. Galls that secrete honeydew: a contribution to the problem as to whether galls are altruistic adaptions. Bulletin of the Brooklin entomological society 19(4): 101-124.
Boucek, Z. 1988. Australia Chalcidoidea (Hymenoptera): A biosystematic revision of genera of fourteen families, with a reclassification of species. Centre for Agricultural Bioscience International, Wallingford, England, pp. 83.
Byers, J. A., J. W. Brewer, and D. W. Denna. 1976. Plant growth hormones in pinyon insect galls. Marcellia 39: 125-134.
Chalker-Scott, L. 1999. Environmental significance of anthocyanins in plant stress responses. Photochemistry and photobiology 70(1): 1-9.
Chittka, L., and T. F. Döring. 2007. Are autumn foliage colors red signals to alphids? PLoS biology 5(8): 1640-1644.
Choinski Jr., J. S., and J. M. Johnson. 1993. Changes in photosynthesis and water status of developing leaves of Brachystegia spiciformis Benth. Tree physiology 13: 17-27.
Cockerell, T. D. A. 1890. The evolution of insect galls. Entomologist 23: 73-76.
Cornell, H. V. 1983. The secondary chemistry and complex morphology of galls formed by the Cynipinae (Hymenoptear): Why and how? American midland naturalist 110: 225-234.
Csóka, G. 1997. Gubacsok - Plant Galls. Agroinform Kiadó, Budapest, Hungary.
DeBruyn, L., I. Vandevyvere, D. Jaminé, and E. Prinsen. 1998. The effects of gall formation by Lipara lucens (Diptera, Choropidae) on its host Phragmites australis (Poaceae). In: G. Csóka, W. J. Mattson, G. N. Stone, and P. W. Price (eds.), The Biology of Gall-Inducing Arthropods. Forest Service USDA, St. Paul, America, pp. 173-187.
Dobson, S. L., and M. A. Tanouye. 1998. Evidence for a genomic imprinting sex determination mechanism in Nasonia vitripennis (Hymenoptera; Chalcidoidea). Genetics 149: 233-242.
Doganlar, O. 2005. Occurrence of Leptocybe invasa Fisher & La Salle, 2004 (Hymenoptera: Chalcidoidea) on Eucalyptus camaldulensis in Turkey, with a description of the male sex. Zoology in the Middle East 35: 112-114.
Dreger-Jauffret, F., and J. D. Shorthouse. 1992. Diversity of gall-inducing insects and their galls. In: J. D. Shorthouse, and O, Rohfritsch, eds. Biology of insect-induced galls. Oxford University Press, New York, America, pp. 8-33.
El-Akkad, S. S. 2004. Biochemical changes induced in Populus nigra leaves by galling aphids Pemphigous populi. International Journal of Agricultural and Biology 6: 659-664.
Espirito-Santo, M. M., and G. W. Fernandes, 2007. How many species of gall-inducing insects are there on earth, and where are they? Annals of the Entomological Society of America 100(2): 95-99.
Felt, E. P. 1940. Plant galls and gall makers. Comstock, Ithaca, America, pp. 3-9.
Fernandes, G. W., P. W. Price, S. J. Gonçalves-Alvim, T. P. Craig, and D. Yanega. 2000. Response of the galling insect Aciuruna trixa Curran (Diptera: Tephritidae) to host plant quality. Annals of the entomological society of Brazil 29(3): 423-431.
Gates, M. W., and M. E. Schauff. 2005. The first report of Oncastichus goughi (Hymenoptera: Eulophidae): an introduced pest of waxflower (Myrtaceae; Chamaelaucium unicnatum) from South America. Entomological News 116(2): 115-116.
Gibson, G. A. P., J. T. Huber, and J. B. Wolley. 1997. Annotated keys to the genera of Nearctic Chalcidoidea (Hymenoptera). Ottawa, Canada, pp. 329.
Gil, L., W. Tadesse, E. Tolosana, and R. López. 2010. Eucalyptus species management, history, status and trends in Ethiopia. Proceedings from the Congress held Addis Ababa. September 15th-17th, 2010.
Gullan, P. F., D. R. Miller, and L. G. Cook. 2005. Gall-inducing scale insects (Hemiptera: Sternorrhyncha: Coccoidea). In: A. Raman, C. W. Schaefer, and T. M. Withers (eds.), Biology, ecology, and evolution of gall-Inducing arthropods. Science Publishers, Inc., Enfield, New Hampshire, America, pp. 507-537.
Häring, D. A., Huber, M. J., D. Suter, P. J. Edwards, and A. Lüscher. 2008. Plant enemy-derived elicitors increase the folira tannin concentration of Onobrychis viciifolia without a trade-of to growth. Annuals of botany 102: 979-987.
Harper, L. J., K. Schonrögge, K. Y. Lim, P. Francis, and C. P. Lichtenstein. 2004. Cynipid galls: insect-induced modifications of plant development creates novel plant orgns. Plant, cell and environment 27: 327-335.
Hartley, S. E. 1998. The chemical composition of plant galls: are levels of nutrients and secondary compounds controlled by the gall-former. Oecologia 113: 492-501.
Inbar, M., R. T. Mayer, and H. Doostdar. Induced activity of pathogenesis related (PR) proteins in aphid galls. 2003. Symbiosis 34: 293-300.
Inbar, M., I. Izhaki, A. Koplovich, I. Lupo, N. Silanikove, T. Glasser, Y. Gerchman, A. Perevolotsky, and S. Lev-Yadun. 2010. Why do many galls have conspicuous colors? A new hypothesis. Arthropod-plant interactions 4: 1-6.
Jacob, J. P., and A. R. Kumar, 2009. Incidence of galls induced by Leptocybe invasa on seedlings of Eucalyptus camaldulensis and E. tereticornis from different seed sources in southern India. International journal of ecology and environmental sciences 35(2-3): 187-198.
Javaregowda, J., and S. T. Prabhu. 2010. Susceptibility of eucalyptus species and clones to gall wasp, Leptocybe invasa Fisher and La Salle (Eulophidae: Hymenoptera) in Karnataka. Karnataka Journal of Agricultural Sciences 23: 220-221.
Kavitha Kumari, N. 2009. Bioecology and management of Eucalyptus gall wasp, Leptocybe invasa Fisher & La Salle (Hymenoptera: Eulophidae). M.S. Thesis, University of Agricultural Sciences, Dharwad, India.
Kavitha Kumari, N., H. Kulkarni, A. S. Vastrad, and K. Goud Basavana. 2010. Biology of eucalyptus gall wasp, Leptocybe invasa Fisher and La Salle (Hymenoptera: Eulophidae). Karnataka Journal of agriculture science 23(1): 211-212.
Kim, I. K., Z. Mendel, A. Protasov, D. Blumberg, and J. La Salle. 2008. Taxonomy, biology, and efficacy of two Australian parasitoids of the eucalyptus gall wasp, Leptocybe invasa Fisher & La Salle (Hymenoptera: Eulophidae: Tetrastichinae). Zootaxa 1910: 1-20.
Koncz, N. K., L. J. Szabó, C. Máthé, K. Jámbrik, and M. M-Hamvas, 2011. Histological study of quercus galls of Neuroterus quercusbaccarum (Linnaeus, 1758) (Hymenoptera: Cynipidae). Acta Biologica Szegediensis 55: 247 - 253.
Küster, E. 1911. Die Gallen der Pflanzen. Ein Lehrbuch der Botanikerund Entomologen. S. Hirzel, Leipzig, Germany, pp. 85-160.
Labandeira, C. C., and T. L. Phillips. 1996. A Carboniferous insect gall: Insight into early ecologic history of the Holometabola. Proceedings of National Academy Sciences U.S.A. 93: 8470-8474.
Larew, H. G. 1982. A comparative anatomical study of galls caused by the major cecidogenetic groups, with special emphasis on the nutritive tissue. Oregon State University, Oregon, America.
La Salle, J. 2005. Biology of gall inducers and evolution of gall induction in Chalcidoidea (Hymenoptera: Eulophidae, Eurytomidae, Pteromalidae, Tanaostigmatidae, Torymidae). In: A. Raman, C. W. Schaefer, and T. M. Withers (eds.), Biology, ecology, and evolution of gall-Inducing arthropods. Science Publishers, Inc., Enfield, New Hampshire, America, pp. 507-537.
La Salle, J., M. Ramadan, and B. Kumashiro. 2009. A new parasitoid of the Erythrina gall wasp, Quadrastichus erythrinae Kim (Hymenoptera: Eulophidae). Zootaxa 2083: 19-26.
Lalonde, R. G., and J. D. Shorthouse. 1983. Developmental morphology of the gall of Urophora cardui (Diptera, Tephritidae) in the stem of Canada thistle (Cirisium arvense). Canadian entomology 114: 873-878.
Mani, M. S. 1964. Ecology of plant galls. W. Junk Publishers, Hague, Netherland, pp. 1-434.
Mapes, C. C., and P. J. Davies. 1998. Plant hormones and gall formation by Eurosta solidaginis on Solidago altissima. In: G. Csóka, W. J. Mattson, G. N. Stone, and P. W. Price (eds.), The biology of gall-inducing arthropods. Forest Service USDA, St. Paul, America, pp. 161-172.
Martel, V., and G. Boivin. 2004. Premating dispersion in the egg parasitoid Trichgramma (Hymenoptera: Trichogrammatidae). Environmental entomology 33(4): 855-859.
McManus, J. F. A. 1948. Histological and histochemical use of periodic acid. Stain Technology 23: 99-108.
Mendel, Z., A. Protasov, N. Fisher, and J. La Salle. 2004. Taxonomy and biology of Leptocybe invasa gen. & sp. n. (Hymenoptera: Eulophidae), and invasive gall inducer on Eucalyptus. Australian Journal of Entomology 43(2): 101-113.
Meyer, J. 1957. Cécidogenêse comparée de quelques galles d'arthropodes et évolution cytologique des tissus nouriciers. Thesis, University of Strasbourg, France.
Miller, D. G., C. T. Ivey, and J. D. Shedd. 2009. Support for the microenvironment hypothesis for adaptive value of gall induction in the California gall wasp, Andricus quercuscalifornicus. Entomologia experimentalis et applicate 132: 126-133.
Minsheng, Y. 2003. Present situation and prospects for eucalypt plantation in China. In: J. W., Turnbull（eds.）, Eucalyptus in Asia. Proceedings of an international conference held in Zhanjiang, Guangdong, pp. 9-15.
Morrow, P. A., and L. R. Fox. 1980. Effects of variation in Eucalyptus essential oil yield on insect growth and grazing damage. Oecologia 45: 209-219.
Narendran, T. C., S. Santhosh, and K. Sudheer. 2007. Biosystematics and biogeography of oriental Chalcidoidea (Hymenoptera) associated with plant galls. Oriental Insects 41: 141-167.
Nyeko, P., K. E. Mutitu, B. Otieno, G. Ngae, and R. Day. 2010. Variations in Leptocybe invasa (Hymenoptera: Eulophidae) population and infestation on eucalyptus germplasms in Uganda and Kenya. International Journal of Pest Management 56(2): 137-144.
Olenichenko, N. A., and N. V. Zagoskina. 2005. Response of winter wheat to cold: production of phenol compounds and L-phenylalanine ammonia lyase activity. Applied biochemistry and microbiology 41(6): 681-685.
Paine, T. D., M. J. Steinbauer, and S. A. Lawson. 2011. Native and exotic pests of Eucalyptus: a worldwide perspective. Annual Review of Entomology 56: 181-201.
Price, P. W. 2005. Adaptive radiation of gall-inducing insects. Basic and Applied Ecology 6: 413-421.
Price, P. W., G. L. Waring, and G. W. Fernandes. 1986. Hypothesis on the adaptive nature of galls. Proceedings of the Entomological Society of Washington 88(2): 361-363.
Raman, A. 2011. Morphogenesis of insect-induced plant galls: facts and questions. Flora 206: 517-533.
Raman, A., C. W. Schaefer, and T. M. Withers. 2005. Morphogenesis of insect-induced plant galls. In: A. Raman, C. Schaefer, T. Withers, (eds.), Biology, ecology, and evolution of gall-inducing Arthropods, Science Publishers, India, pp.1-33.
Rockwood, D. L., A. W. Rudie, S. A. Ralph, J. Y. Zhu, and J. E. Winandy. 2008. Energy product options for Eucalyptus species grown as short rotation woody crops. International journal of molecular sciences 9(8): 1361-1378.
Rohfritsch, O. 1992. Patterns in gall development. In: J. D. Shorthouse, and O. Rohfritsch, (eds.), Biology of insect-induced Galls, Oxford University Press, New York, America, pp. 60-86.
Rohfritsch, O., and J. D. Shorthouse. 1982. Insect galls. In: K. Günter, and J. S. Schell, (eds.), Molecular biology of plant tumors, Academic Press, New York, America, pp. 131-152.
Roskam, J. C. 1992. Evolution of the gall inducing guild. In: J. D. Shorthouse, and O. Rohfritsch, (eds.), Biology of insect-induced galls, Oxford University Press, America, pp. 34-49.
Roubos, C. R., and O. E. Liburd. 2010. Pupation and emergence of blueberry gall midge, Dasineura oxycoccana (Diptera: Cecidomyiidae), under varying temperature conditions. Florida entomologist 93(2): 283-290.
Roux G., A. Roques, and F. Menu. 1997. Effect of temperature and photoperiod on diapauses development in a Douglas fir seed chalcid, Megastigmus spermotrophus. Oecologia 111: 172-177.
Schönrogge, K., L. J., Harper, S. E. Brooks, J. D. Shorthouse, and C. P. Lichtenstein. 1998. Reprogramming plant development: two approaches to study the molecular mechanism of gall formation. In: G. Csóka, W. J. Mattson, G. N. Stone, and P. W. Price (eds.), The Biology of Gall-Inducing Arthropods. Forest Service USDA, St. Paul, America, pp. 173-187.
Schoonhoven, L. M., J. J. A., van Loon, and M. Dicke. 2005. Insect-plant biology, Oxford University Press, America, pp. 440.
Shannon, R. E., and J. W. Brewer. 1980. Starch and sugar levels in three coniferous insect galls. Z. Angew. Entomology 89: 526-533.
Shorthouse J. D, D.Wool, and A. Raman. 2005. Gall-inducing insects -Nature’s most sophisticated herbivores. Basic Applied Ecology 6: 407-411.
Stone, G. N., and K. Schönrogge. 2003. The adaptive significance of insect gall morphology. Trends in Ecology and Evolution 18: 512-522.
Stone, G. N, K, Schönrogge, R. Atkinson, D. Bellido, and J. Pujade-Villar. 2002. The population biology of oak gall wasps (Hymenoptera:Cynipidae). Annual Review of Entomology 47: 633-668.
Taper, M. L., E. M. Zummerman, and T. J. Case. 1986. Sources of mortality for a cynipid gall wasp (Drycocosmus dibuosus) (Hymenoptera, Cynipidae)): the importance of the tannin-fungus interaction. Oecologia 68: 437-445.
Taylor, S. H. 1949. Initiation and development of the gall of Aylax glechome on Nepeta hedercea. American Journal of Botany 36(2): 222-230.
Weis, A. E., W. G. Abrahamson, and K. D. McCrea. 1985. Host gall size and oviposition success by the parasitoid Eurytoma gigantea. Ecological entomology 10: 341-348.
Wells, B. W. 1921. Evolution of zoocecidia. Batanical Gazette 71: 358-377.
Williams, J., and J. Woinarski. 1997. Eucalypt ecology: individuals to ecosystems. Cambridge University Press, Cambridge, UK, pp. 9.
Yang, M. M., and G. S. Tung. 1998. The diversity of insect-induced galls on vascular plants in Taiwan: a preliminary report. In: G. Csóka, W. J. Mattson, G. N. Stone, and P. W. Price (eds.), The Biology of Gall-Inducing Arthropods. Forest Service USDA, St. Paul, America, pp. 173-187.