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木蝨(半翅目:木蝨總科)的發聲行為:功能形態與機制

By Zong-Ze Wu and 吳宗澤

Abstract

Sound production in psyllids has been described but to date, the exact mechanism has remained unclear. Several sound production mechanisms in psyllids have been proposed, with most suggesting stridulation between the anal area of forewing and axillary cord as the mode of action. The aim of this study is to determine the specific sound producing structures and mechanisms of the psyllids. Extensive tests were implemented on the species Macrohomotama gladiata Kuwayama belonging to Homotomidae. Through cutting off possible sound producing organs and observing wing-beat frequency with a high-speed camera, acoustic signals were recorded to examine six possible hypotheses. These hypotheses include wing-beating, wing-wing rubbing, wing-thorax rubbing, wing-leg rubbing, leg-abdomen rubbing, and axillary sclerite rubbing. In order to confirm whether a particular mechanism is common in psyllids, other species belonging to different families were also examined, including Trioza sozanica (Boselli) (Triozidae), Mesohomotama camphorae Kuwayama (Carsidaridae), Psylla oluanpiensis Yang and Psylla tobirae Miyatake (Psyllidae). Scanning electron microscopy (SEM) was used to observe the existence of special features that might serve as sound producing structures. Results indicate that psyllids were not able to make any signal after the entire forewing was removed, leading to a rejection of the wing-leg rubbing hypothesis. The wing-beating frequency did not match the dominant frequency of sound, resulting in the rejection of wing-beating hypothesis. Psyllids are still able to produce signals following the removal of the anal area of the forewing so the wing-thorax rubbing hypothesis is also rejected. Psyllids with the forewing membrane area removed can also still make signals, so the hypotheses of wing-wing rubbing and wing-leg rubbing are rejected. Therefore, the axillary sclerite-rubbing hypothesis is accepted due to the fact that the remaining axillary sclerites can not rub each other or rub legs. However, removal of the anal and membrane area of forewing produces sound with weaker amplitude and the modification is more obvious in the former than in the latter. Because to keep anal area of forewing without axillary sclerite is impossible, the hypothesis of wing-thorax rubbing was rejected due to the ablility of sound production with the axillary sclerite present. Our work suggests that psyllids likely make sound by rubbing axillary cord and anal area of forewing. We also further inspected the surface of the axillary sclerite because of the known rough surface on the axillary cord of psyllids. SEM photographs show that the secondary axillary sclerite is stronger than other axillary sclerites and also bears many protuberances that would be suitable for stridulation. In conclusion, the sound producing mechanism of psyllids involves two groups of morphological structures. The first group of stridulation structures is the anal area of forewing and the axillary cord, and the second one is the axillary sclerite of forewing and the mesothorax.木蝨發聲行為已在許多文獻中記載,但對於發聲機制仍停留於推測階段。多數文獻推測木蝨是藉由前翅翅臀區 (anal area of forewing) 摩擦翅基索 (axillary cord) 而產生聲音。本研究以榕木蝨科的高背木蝨 (Macrohomotama gladiata Kuwayama) 為主要材料,透過切除可能發聲位置與利用高速攝影機觀察翅震動頻率,配合錄音與觀察實驗,以瞭解木蝨發聲結構與機制,並驗證「翅震發聲假說」、「翅翅摩擦發聲假說」、「翅胸摩擦假說」、「足腹摩擦假說」、「翅足摩擦假說」以及「翅鍵骨 (axillary sclerite) 發聲假說」之可能性。同時檢視不同科的物種以確定發聲機制的普遍性,包括三叉木蝨科的虎皮楠木蝨 (Trioza sozanica (Boselli))、錦葵木蝨科的黃槿木蝨 (Mesohomotama camphorae Kuwayama)、與木蝨科的台灣桐木蝨 (Psylla oluanpiensis Yang) 及海桐木蝨 (Psylla tobirae Miyatake)。進一步透過掃描式電子顯微鏡 (SEM) 之檢視,瞭解其發聲結構是否具有特殊構造用於發聲。結果顯示,切除完整前翅的組別之木蝨無法產生聲音訊號,因此排除足腹摩擦假說與後翅涉及發聲的可能性;翅震頻率與聲音主頻率不一致,排除翅震發聲假說;移除前翅臀區的木蝨依然可以發出聲音訊號,排除翅胸摩擦假說;在切除前翅膜質區的組別中,所有的木蝨皆可產生聲音訊號,由於僅剩的左右兩側之翅鍵骨無法相互摩擦,亦無法進行翅足摩擦,因此拒絕翅翅摩擦與翅足摩擦假說並接受翅鍵骨假說。切除翅臀區與切除翅膜質區後,2組實驗皆可錄得聲音訊號但明顯為弱,且發現翅臀區切除後聲音訊號的改變量大於移除翅鍵骨後聲音訊號的改變量。由於實驗無法移除翅鍵骨而保留翅臀區,在翅鍵骨存在可發聲的情況下造成翅胸摩擦假說被拒絕。以上實驗支持木蝨可能藉由翅基索與前翅翅臀區摩擦發聲。目前已知木蝨翅基索具有粗糙的表面結構,本研究進一步檢視翅鍵骨之表面結構。SEM結果顯示,木蝨之第二翅鍵骨較其他翅鍵骨發達,且表面具有凸起結構適合用於摩擦發聲。本研究認為木蝨發聲涉及2組形態構造,第一組為翅臀區摩擦翅基索,第二組為翅鍵骨摩擦中胸。致謝 i 摘要 iii Abstract iv 目次 vi 表目次 viii 圖目次 ix 前言 1 前人研究 2 一、聲音在昆蟲扮演的角色 2 二、昆蟲的發聲機制 4 三、半翅目的發聲機制 5 四、木蝨的發聲研究 5 五、木蝨的可能發聲構造─胸與翅的形態 6 材料與方法 8 一、設立實驗假說 8 二、假說驗證流程 9 三、飼養方式與環境 9 四、切除與錄音測試 10 五、木蝨聲音特性定義 11 六、高速攝影機 11 七、掃描式電子顯微鏡檢視 11 八、統計分析 12 結果 13 一、翅區與木蝨發聲音量之關係 13 二、木蝨振翅頻率 14 三、木蝨發聲結構表面構造 14 四、剪翅後木蝨的行為差異 15 討論 16 一、假說測試 16 (一)翅震發聲假說: 16 (二)翅翅摩擦假說: 16 (三)翅胸摩擦假說: 17 (四)足腹摩擦假說: 17 (五)翅足摩擦假說: 17 (六)翅鍵骨發聲假說: 18 二、發聲機制 18 三、木蝨的發聲行為 19 四、音量與交配成功率的生態意義 20 五、發聲結構與分類學 21 六、結論 21 引用文獻 23 表 28 圖 3

Topics: 木蝨, 發聲機制, 翅鍵骨, 前翅臀區, 翅基索, Psyllids, Sound producing mechanism, Axillary sclerite, Anal area of forewing, Axillary cord
Year: 2017
OAI identifier: oai:ir.lib.nchu.edu.tw:123456789/85978
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