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Function of body coloration of Taiwanese crab spiders
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|摘要:||蟹蛛是不結網而坐等在花上捕捉傳粉昆蟲的捕食者，有些種類的成熟雌性個體能依據停棲花的顏色改變體色，且可在白色和黃色間進行轉換。蟹蛛體色的功能已有許多研究探討，有學者提出四種假說: (1)對獵物或捕食者具隱蔽性、(2)吸引獵物、(3)蜘蛛會被避開、(4)蜘蛛不被在乎。有許多因素可能影響體色功能，如蟹蛛體色的反射光譜特性、蟹蛛與花顏色對比產生的光訊號、獵物或捕食者視覺系統對顏色的感知，及牠們對蟹蛛的行為反應等。起初有些學者根據色差模型提出蜜蜂獵物與鳥類捕食者在遠距離無法輕易察覺有歐洲蟹蛛在花上；之後有研究指出許多澳洲種類蟹蛛的體表會反射UV光，但歐洲種類則否。行為實驗也發現蜜蜂會偏好有澳洲蟹蛛存在的花，卻避開有歐洲蟹蛛棲息的花。此研究探討了台灣兩種常見蟹蛛：三角蟹蛛(Thomisus labefactus)和三突花蛛(Misumenops tricuspidatus)在日間和夜間的體色功能。近年有許多研究指出許多結圓網蜘蛛的體表具有明亮色斑，不論是在日間或夜間皆可增加獵物攔截率。雖然蟹蛛普遍被認為是日行性蜘蛛，當我在台中大坑調查野外蟹蛛的24小時活動情形，發現其實蟹蛛在夜間的活動量相當大。在白色大花咸豐草上，蟹蛛在日夜間都會出來活動及覓食；且在黃色馬纓丹上的蟹蛛族群在夜間的出現量與捕捉到的獵物量及生物量皆顯著高於日間。在此觀察中，我記錄到蟹蛛在日間與夜間的主要獵物是鱗翅目昆蟲。接著我使用光譜儀測量蟹蛛與花的反射光譜，再利用日間蜜蜂及夜間天蛾的視覺模型去量化蟹蛛體色在昆蟲眼中的情形。色差計算結果顯示，兩種顏色蟹蛛在日夜間其體色與花之顏色可被區分，只有白蟹蛛在被蜜蜂以遠距離觀看時可能不易與白色花瓣區分。最後我在野外進行操控性實驗，利用夜視攝影機監測日間和夜間的獵物及捕食者對蟹蛛的行為反應。此研究使用相似於真實蜘蛛體色的黏土及顏料製作白色與黃色之蜘蛛模型，將其用蟲針固定於花上，並與空花進行獵物和捕食者造訪率之比較。結果顯示，日間獵物在有蟹蛛模型花之降落率顯著低於空花，夜間獵物些微偏好空花，但無顯著差異。潛在捕食者只在日間被記錄到，捕食性膜翅目些微偏好審視有蟹蛛模型之花，但無顯著差異。然而，我也發現黃花之捕食者審視率為於白花的兩倍，推測可能是白蟹蛛在白花上的單色色差值較低，兩者之顏色較接近進而降低了捕食者的注意。本研究結果揭露了日行性及夜行性的鱗翅目昆蟲對花上存有蟹蛛時的反應，台灣蟹蛛體色可能對日間視覺性捕食者具隱蔽效果，但會付出被日間獵物避開的代價；然而蟹蛛轉而依賴於夜間大量捕食以彌補日間獵物的損失。|
Crab spiders (Thomisidae) are sit-and-wait predators which can change body colors according to the color of flowers they sit on. The function of crab spiders’ body coloration has been studied over one century and four hypotheses had been proposed: (1) crypsis, (2) prey attraction, (3) spider avoidance and (4) indifference to spiders. There are many factors influencing the function of crab spiders’ body coloration, such as reflecting properties of spider’s body color, color contrast between spider and flower, color perception in vision systems and responding behavior of signal receivers such as prey or predator. Several researches demonstrated that the body color of European crab spiders could not be distinguished from that of flowers when viewed by honeybees or birds under achromatic vision. Previous studies also revealed that reflectance properties of crab spiders’ body coloration may vary with different geographic regions. For example, many species of Australian crab spiders reflect UV, but European species do not. Furthermore, behavioral experiments showed that honeybees landed more on flowers with Australian crab spiders than spider-free flowers, but landed less on flowers with European crab spiders. In this study, two species of Taiwanese crab spiders, Thomisus labefactus and Misumenops tricuspidatus were used to evaluate the function of East Asian crab spiders’ body colorations. Since recent studies revealed that many orb-web spiders’ conspicuous body bands can visually attract prey during day and night, I first surveyed the temporal activity patterns of wild crab spiders in Daken, Taichung city, Taiwan. In white Biden pilosa habitat, crab spiders actively hunted both at day and night. In yellow Lantana camara habitat, they mainly hunted during nighttime, and the number and biomass of their nocturnal prey were significantly higher than those of diurnal prey. The recorded major diurnal and nocturnal prey were Lepidoptera insects. Next, I used spectrometer to measure reflectance spectra of spiders and flowers, and used diurnal and nocturnal vision models to quantify how these crab spiders were viewed by insects. The results of color contrast calculations showed that two color morphs of crab spiders were conspicuous to diurnal hymenopterans and nocturnal lepidopterans. Only white crab spiders may not be easily perceived from white flowers when viewed by hymenopterans in long distance. Finally, I performed field experiments by using infrared video cameras to monitor the responses of insect prey and predators to crab spiders during day and night. I used clay and paint with spectral properties similar to the real crab spiders to manufacture dummy spiders. The insect visitation rates of flowers with dummy were compared to that of vacant flowers. The results showed that significantly more diurnal prey landed on vacant flower. Slightly more nocturnal prey landed on vacant flowers but such difference was not statistically significant. The potential predators were only recorded at daytime. Hymenopteran predators seem to be more likely to scan flowers with dummy, but such difference was not statistically significant. However, the predator scanning rates in yellow flowers were two times higher than that in white flowers. A lower achromatic color contrast value of white spiders seen against white flowers might be responsible for such results. This study demonstrated how diurnal and nocturnal lepidopterans responded to crab spiders. Results of my study suggested that body color of Taiwanese crab spiders may function to protect spiders against visual predators but at a cost of deterring diurnal prey. However, such cost could be balanced by higher prey intake of these spiders’ nocturnal hunting.
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