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標題: Gall and colony size of gall-inducing social aphid Colophina clematis and its biology
造癭的社會性蚜蟲Colophina clematis數量與癭大小的關係及其生物學
作者: Chen, Bo-Fei
出版社: 昆蟲學系所
引用: Aoki, S. 1976. Occurrence of dimorphism in the first instar larva of Colophina clematis (Homoptera, Aphidoidea). Kontyû 44: 130-137. Aoki, S. 1977. Colophina clematis (Homoptera, Pemphigidae), an aphid species with soldiers. Kontyû 45: 276-282. Aoki, S. 1980. Life cycles of two Colophina aphids (Homoptera, Pemphigidae) producing soldiers. Kontyû 48: 464-476. Aoki, S. 1983. A new Taiwanese species of Colophina (Homoptera, Aphidoidea) producing large soldiers. Kontyû 51: 282-288. Aoki, S. 2003. Soldiers, altruistic dispersal and its consequences for aphid societies. pp. 201-215. In: T. Kikuchi, N. Azuma, and S. Higashi, eds. Genes, Behaviors and Evolution of Social Insects. Hokkaido University Press, Sapporo. Aoki, S., and S. Makino. 1982. Gall usurpation and lethal fighting among fundatrices of the aphid Epipemphigus niisimae (Homoptera, Pemphigidae). Kontyû 50: 365-376. Blackman, R. L., and V. F. Eastop. 2000. Aphid on the world's crops 2nd ed. John Wiley and Sons Limited, West Sussex. Crespi, B. J. 1992. Eusociality in Australian gall thrips. Nature 359:724-726. Dixon, A. F. G. 1998. Aphid ecology, 2nd ed. Blackie and Son Limited, Glasgow. Hamilton, W. D. 1964. The genetical evolution of social behaviour. I, II. J. Theor. Biol. 7: 1-52. Hori, K. 1992. Insect secretions and their effect on plant growth, with special reference to hemepterans. pp. 157-170. In: J. D. Shorthouse, and O. Rohfritsch. eds. Biology of insect-induced galls. Oxford University Press, New York. Huang, T. C. et al. eds. 1996. Flora of Taiwan, 2nd ed. vol. 2. Editorial Committee of the Flora of Taiwan, Second Edition, Taipei. Jarvis, J. U. M. 1981. Eusociality in a mammal: Cooperative breeding in naked mole-rat colonies. Science 212: 571-573. Kent, D. S., and J. A. Simpson. 1992. Eusociality in the beetle Austroplatypus incompertus (Coleoptera: Curculionidae). Naturwissenschaften 79: 86-87. Lattin, J. D. 2000. Minute pirate bugs (Anthocoridae). pp. 607-637. In: C. W. Schaefer, and A. R. Panizzi. eds. Heteroptera of economic importance. CRC Press LLC, Florida. Mani, M. S. 1992. Introduction to cecidology. pp. 3-7. In: J. D. Shorthouse, and O. Rohfritsch. eds. Biology of insect-induced galls. Oxford University Press, New York. Meyer, J. 1987. Plant galls and gall inducers. Gebrüder Borntraeger, Berlin. Meyling, N. V., A. Enkegaard, and H. Brødsgaard. 2003. Two Anthocoris bugs as predators of glasshouse aphids - voracity and prey preference. Entomol. Exp. Appl. 108: 59-70. Price, P. W. 1984. Insect ecology. John Wiley & Sons, Inc. USA. Price, P. W., G. L. Waring, and G. W. Fernandes. 1986. Hypotheses on the adaptive nature of gall. Proc. Entomol. Soc. Wash. 88: 361-363. Price, P. W., G. W. Fernandes and G. L. Waring. 1987. Adaptive nature of insect galls. Environ. Entomol. 16: 15-24. Rohfritsch, O. 1992. Patterns in gall development. pp. 60-86. In: J. D. Shorthouse, and O. Rohfritsch. eds. Biology of insect-induced galls. Oxford University Press, New York. Sakata, K., and Y. Itô. 1991. Life history characteristics and behaviour of the bamboo aphid, Pseudoregma bambucicola (Hemiptera: Pemphigidae), having sterile soldiers. Insectes Soc. 38: 317-326. Stern, D. L., and W. A. Foster. 1996. The evolution of solders in aphids. Biol. Rev. 71: 27-79. Su, H. J. 1984. Studies on the climate and vegetation type of the natural forest in Taiwan (Ⅱ). Altitudinal vegetation zones in relation to temperature gradient. Quart. Journ. Chin. Forest. 17: 57-73. Wool, D. 2005. Gall-inducing aphids: biology, ecology, and evolution. pp. 73-132. In: A. Raman, C. W. Schaefer, and T. M. Withers, eds. Biology, Ecology, and Evolution of Gall-inducing Arthropods. Science Publishers Inc., New Hampshire. Wu, Z. Y., P. H. Raven & D. Y. Hong, eds. 2001. Flora of China vol. 6 (Caryophyllaceae through Lardizabalaceae). Science Press and Missouri Botanical Garden Press, Beijing and St. Louis.
Colophina clematis (Shinji, 1922) 是第一種被發現會產生兵蚜的蚜蟲,為目前全世界已知約50種的社會性蚜蟲之一。Colophina clematis在第一寄主櫸木(Zelkova serrata (Thunb.))的葉片上造球狀的蟲癭,在第二寄主Clematis apiifolia DC.上則不造癭但是卻會產生形態特化的不生殖兵蚜。有關Colophina clematis的研究,目前已發表的資料仍侷限於第二寄主之特化兵蚜的形態及行為的描述,以及簡單的整體生活史敘述。為深入瞭解C. clematis的生物學,本研究針對台灣中部的C. clematis族群進行記錄並分析蟲癭生長情形、癭內蚜蟲族群數量變化、癭內發現的其他生物種類與蚜蟲的關係。並且調查其第二寄主,確認後將其上的C. clematis族群連同寄主一起帶回實驗室飼養,每日記錄各齡期所需的發育時間、成蟲的存活天數及其他未被描述的行為。結果顯示C. clematis於2月末開始造癭,蟲癭於5月成熟開裂,此時有翅型幹子成熟,而癭內蚜蟲數量也在此月份達到高峰。蚜蟲族群的個體在7月完全離開蟲癭,遷移到第二寄主串鼻龍(Clematis grata Wall.)。研究期間發現,成熟的蟲癭體積具有明顯變異,體積依 計算,範圍由188.94 mm3到1607.04 mm3,分析結果顯示成熟蟲癭的體積大小與癭內蚜蟲數量有顯著的正相關(r=0.65829, p<0.01),即越大的癭內,容納有越多的蚜蟲個體。因此蟲癭體積可能為影響癭內蚜蟲個體數的密度制約因子(density-dependent factor)。癭內亦伴隨有其他非造癭生物,包括薊馬、長斑蚜若蟲、花蝽若蟲、雙翅目幼蟲和瓢蟲幼蟲等5類昆蟲。前三者應為偶發性的入侵,並非真正的客居生物(inquiline);而雙翅目幼蟲和瓢蟲幼蟲應為Colophina clematis造癭時期的主要天敵。另外,生活於第二寄主上的一齡若蟲發育天數約為其他齡期若蟲的兩倍(7.6±1.9天),與其他二種同科蚜蟲的發育情形類似。在飼養的77隻個體,僅有一隻完成完整的生命週期,其成蟲壽命共40天並且產生了109隻的一般若蟲和15隻兵蚜。本研究還發現一些未被描述過的行為,包括生活於第二寄主的兵蚜在抓住騷擾物之後行為反應顯示應可以分辨騷擾物為同種蚜蟲或是其他生物,以及成蚜可能會分泌化學物質防禦天敵等。

Colophina clematis (Shinji, 1922) is the first species among nearly 50 aphid species which were already discovered producing sterile soldiers. C. clematis was reported producing morphologically specialized sterile soldiers on the secondary host, Clematis apiifolia DC. in Japan. On its primary host, Zelkova serrata (Thunb.), Colophina clematis induces rolling-sphere galls on the leaves. The reported studies of C. clematis was restricted to the description of morphology of sterile soldiers and brief life cycle. To understand batter about the biology of C. clematis, further survey and analyses were conducted in central Taiwan, including gall developments, variations of individual numbers inside galls, other organisms associate with galls, and their relationships with the gallers. In addition, life history information and behavior observation based on laboratory-reared colonies on the secondary host were recorded, including length of each stadium, longevity of adult life, and some undescribed behaviors. The results showed that C. clematis induced galls in late February. Galls matured and dehisced in May when the progenies of stem mothers were mature, and the numbers of individuals inside the galls also reached the highest peak in the same month. The aphids left the galls completely in July and migrated to the secondary host, Clematis grata. The volume of matured galls varied obviously from 188.94 mm3 to 1607.04 mm3, and the analytic results showed that the gall volume was positively correlated with the individual numbers inside the galls (r=0.65829, p<0.01). In other words, bigger galls hosted more aphids. Therefore, it indicates that the gall volume is a density-dependent factor and, hence, affect the colony sizes inside the galls. Five other taxa were found associated with galls, including a thrips, a Tinocallis aphid nymph, an anthocorid nymph, a dipteran larva, and a coccinellid larva. The former three were considered as occasional invaders but not real inquilines. The latter two should be the main predators of Colophina clematis on Clematis. In addition, the length of 1st stadium of the nymphs which lived on the secondary host was almost twice longer than the lengths of other stadia (7.91.9 days), and the developmental strategy was similar to another two aphid species of the same family. Only one individual completed the whole life span and totally produced 109 normal nymphs and 15 soldiers. Two undescribed behaviors were also recorded in this study. The soldiers which lived on Clematis seemed to be able to identify whether the disturbances are con-specific or not when they hugged the disturbances, and adult Colophina clematis which lived on Clematis seemed that they also perform defensing behaviors by secreting waxy chemical materials.
其他識別: U0005-2108200717212500
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