臺灣產扁蚜亞科之生物系統分類與分子親緣關係研究

本研究於近年來，重新採集、調查扁蚜亞科 (Hormaphidinae: Aphididae: Hemiptera) 的種類，在臺灣地區之物種豐富度；由採集的資料得知，門前氏扁蚜屬 (Monzenia Takahashi, 1962)、鄰日胸扁蚜屬 (Paranipponaphis Takahashi, 1959)、管扁蚜屬 (Tuberaphis Takahashi, 1933) 和三節角扁蚜屬 (Triantennaria gen. nov.) 等 4 屬，為臺灣地區發現之新屬或新記錄屬，多腺扁蚜 (Astegopteryx glandulosa Noordam, 1991)、巴西蝨蚜 (Cerataphis brasiliensis (Ghesuière, 1934))、柚葉藤蝨蚜 (C. aff. pothophila Noordam, 1991)、微小門前氏扁蚜 (Monzenia minuta sp. nov.)、長蝨蚜 (Neothoracaphis elongata (Takahashi, 1958))、櫟蝨蚜 (N. querciphaga (Takahashi, 1958))、高尾蝨蚜 (Paranipponaphis takaoensis Takahashi, 1959)、歪體網體蝨蚜 (Reticulaphis asymmetrica Hille Ris Lambers and Takahashi, 1959)、圓體網體蝨蚜 (R. distylii (van der Goot, 1917))、腫胸網體蝨蚜 (R. inflata Yeh and Hsu, 2008)、輪網體蝨蚜 (R. rotifera Hille Ris Lambers and Takahashi, 1959)、稜果榕網體蝨蚜 (R. septica Yeh and Hsu, 2008)、韓國管扁蚜 (Tuberaphis coreana Takahashi, 1933)、竹之內管扁蚜 (T. takenouchii (Takahashi, 1934)) 和蚊母樹三節角扁蚜 (Triantennaria distylia sp. nov.) 等 15 種，為臺灣地區發現之新種或新記錄種，並有 5 種於第一 (原生) 寄主植物 (primary host) 發現之待鑑定種類。在重新調查之後，臺灣地區記錄之扁蚜亞科種類增加至 23 屬，包含 54 種 (未包含待鑑定種)，此物種豐富度近全世界已知種類之三分之一強。在生活環的研究方面，已知扁蚜亞科之蚜蟲種類於第一 (原生) 寄主植物誘發蟲癭 (gall)，目前於安息香科 (Styracaceae) 之寄主植物，共記錄 8 種堅蚜族 (Cerataphidini) 種類之蟲癭，於烏皮九芎 (Styrax formosana) 記錄貓爪癭蚜 (Ceratovacuna nekoashi (Sasaki, 1910))、韓國管扁蚜和竹之內管扁蚜等 3 種扁蚜之蟲癭，另於紅皮 (S. suberifolia) 記錄竹葉扁蚜 (Astegopteryx bambucifoliae (Takahashi, 1921))、鐘型蝨蚜 (Cerataphis jamuritsu (Takahashi, 1931))、安息香圓尾扁蚜 (Ceratoglyphina styracicola (Takahashi, 1936))、竹莖扁蚜 (Pseudoregma bambucicola (Takahashi, 1921)) 和恆春扁蚜 (P. koshunensis (Takahashi, 1924)) 等 5 種扁蚜之蟲癭。另外，於金縷梅科 (Hamamelidaceae) 之寄主植物，共記錄 8 種日本扁蚜族 (Nipponaphidini) 種類之蟲癭，於蚊母樹 (Distylium racemosum) 記錄微小門前氏扁蚜之蟲癭，於水絲梨 (Sycopsis sinensis) 記錄大顆瘤蝨蚜 (Metanipponaphis cuspidatae (Essig and Kuwana, 1918))、網背蝨蚜 (Neothoracaphis quercicola (Takahashi, 1921)) 和 4 種待鑑定種 (Syc. sinensis-branch-spindlelike sp. 1, Syc. sinensis-branch-spindlelike sp. 2, Syc. sinensis-branch-urchinlike, Syc. sinensis-leaf-pearlike) 等 6 種扁蚜之蟲癭，另於尖葉水絲梨 (Distyliopsis dunnii) 記錄另一待鑑定種 (Dis. dunnii-branch-urchinlike) 之蟲癭。然而，研究中發現，蚊母樹三節角扁蚜可以於蚊母樹和尖葉水絲梨都誘發蟲癭，此為極少數已知的扁蚜種類，能夠在不同的第一 (原生) 寄主植物上誘發蟲癭的例子。本研究另外以 DNA 之分子資料，進行扁蚜亞科之系統分類和親緣關係之探討，研究中選取蚜蟲核 (nuclear) DNA 之延伸因子 1α (elongation factor 1 alpha, EF1α) 和部份粒線體 (mitochondrial) DNA 之基因序列，進行比較分析；於分析後得知，在親緣關係樹的重建方面，核 DNA 之延伸因子 1α 較粒線體 DNA 之基因序列，有較好的解析度，而將兩者序列合併後一起分析，能夠得到更佳的結果，對於合併序列的分析，採用貝氏演算法 (Bayesian algorithm)，比其他演算法有更高的支持度。以核 DNA 之延伸因子 1α 所重建之親緣關係樹中，目前已知的 3 個族，各自成為單系群 (monophyly)，但是蚊母樹三節角扁蚜並不包含於任何一個族之中。最後，根據基因序列所重建的親緣關係，配合其他的演化特徵，對兵蚜階級 (soldier caste) 和扁蚜亞科之演化歷程進行推論。

關鍵字

常蚜科；扁蚜亞科；堅蚜族；日本扁蚜族；蟲癭；兵蚜；系統分類

並列摘要

The fauna of the Hormaphidinae was recently resurveyed in Taiwan, and it was found that great biodiversity exists here. Four genera, Monzenia Takahashi, 1962, Paranipponaphis Takahashi, 1959, Tuberaphis Takahashi, 1933, and Triantennaria gen. nov., and 15 species, Astegopteryx glandulosa Noordam, 1991, Cerataphis brasiliensis (Ghesuière, 1934), C. aff. pothophila Noordam, 1991, Monzenia minuta sp. nov., Neothoracaphis elongata (Takahashi, 1958), N. querciphaga (Takahashi, 1958), Paranipponaphis takaoensis Takahashi, 1959, Reticulaphis asymmetrica Hille Ris Lambers and Takahashi, 1959, R. distylii (van der Goot, 1917), R. inflata Yeh and Hsu, 2008, R. rotifera Hille Ris Lambers and Takahashi, 1959, R. septica Yeh and Hsu, 2008, Tuberaphis coreana Takahashi, 1933, T. takenouchii (Takahashi, 1934), and Triantennaria distylia sp. nov., and five unidentified species of gall generations were found to be new to Taiwan from the collections. As the consequence, these increase the total number of species of the Hormaphidinae to 54 species, not including five unidentified species, in 23 genera in Taiwan representing about one-third of the species richness of the Hormaphidinae in the world. Hormaphidine aphids are notable for their complex life cycle on primary hosts on which galls are induced. In this study, eight Cerataphidini species of the gall generation were found on primary hosts, including Styrax spp. (Styracaceae), that is, Ceratovacuna nekoashi (Sasaki, 1910), Tuberaphis coreana Takahashi, 1933, and T. takenouchii (Takahashi, 1934) on the host S. formosana, and Astegopteryx bambucifoliae (Takahashi, 1921), Cerataphis jamuritsu (Takahashi, 1931), Ceratoglyphina styracicola (Takahashi, 1936), Pseudoregma bambucicola (Takahashi, 1921), and P. koshunensis (Takahashi, 1924) on the host S. suberifolia; eight Nipponaphidini species of the gall generation were first reported on various species of the Hamamelidaceae as the primary hosts, that is, Monzenia minuta sp. nov. on Distylium racemosum; Metanipponaphis cuspidatae (Essig and Kuwana, 1918), Neothoracaphis quercicola (Takahashi, 1921), and four unidentified species (Syc. sinensis-branch-spindlelike sp. 1, Syc. sinensis-branch-spindlelike sp. 2, Syc. sinensis-branch-urchinlike, and Syc. sinensis-leaf-pearlike) on the host Sycopsis sinensis, and also one unidentified species (Distyliopsis dunnii-branch-urchinlike) on the host Distyliopsis dunnii. Moreover, Triantennaria distylia sp. nov. can induce similar galls on leaves of Distylium racemosum and Distyliopsis dunnii, and it was interesting to realize that the same aphid species can induce galls on two distinct primary hosts. Records of the primary hosts, Syc. sinensis and Distyliopsis dunii, are both new to hormaphidine aphids. DNA-based approaches were also performed to examine systematic and phylogenetic issues of the Hormaphidinae. From the results of the phylogenetic inferences based on molecular sequences, it was realized that nuclear elongation factor 1 alpha (EF1α) DNA sequences were better than mitochondrial (mt)DNA for inferring the phylogeny of the Hormaphidinae. However, EF1α sequences provide limited resolution within the Hormaphidinae, but increase the proportion of well-supported clades when combined with mtDNA sequences. The Bayesian algorithm was more robust than other algorithms for analyzing the combined dataset. The phylogenetic analysis showed that the three tribes form respective monophyletic clades, except for Triantennaria distylia sp. nov., based on nuclear EF1α sequences. Finally, depending on the phylogenetic trees reconstructed from nuclear EF1α and the combined sequences and referring to certain evolutionary characteristics, the evolution of soldier castes and an evolutionary course of the Hormaphidinae are proposed.

並列關鍵字

Aphididae ； Hormaphidinae ； Cerataphidini ； Nipponaphidini ； galling aphid ； soldier caste ； systematics

參考文獻

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Aoki, S., U. Kurosu, and D. L. Stern. 1991. Aphid soldiers discriminate between soldiers and non-soldiers, rather than between kin and non-kin, in Ceratoglyphina bambusae. Anim. Behav. 42: 865-866.

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臺灣產扁蚜亞科之生物系統分類與分子親緣關係研究

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