複雜的動態構成悌蛺蝶「不精確擬態」的趨同演化基礎
Adelpha iphiclus photo from learnaboutbutterflies.com |
Adelpha serpa photo from learnaboutbutterflies.com |
標題:Complex dynamics underlie the evolution of imperfect wing pattern convergence in butterflies
摘要
Adaptive radiation is characterized by rapid diversification that is strongly associated with ecological specialization. However, understanding the evolutionary mechanisms fueling adaptive diversification requires a detailed knowledge of how natural selection acts at multiple life-history stages. Butterflies within the genus Adelpha represent one of the largest and most diverse butterfly lineages in the Neotropics. Although Adelpha species feed on an extraordinary diversity of larval hosts, convergent evolution is widespread in this group, suggesting that selection for mimicry may contribute to adaptive divergence among species. To investigate this hypothesis, we conducted predation studies in Costa Rica using artificial butterfly facsimiles. Specifically, we predicted that nontoxic, palatable Adelpha species that do not feed on host plants in the family Rubiaceae would benefit from sharing a locally convergent wing pattern with the presumably toxic Rubiaceae-feeding species via reduced predation. Contrary to expectations, we found that the presumed mimic was attacked significantly more than its locally convergent model at a frequency paralleling attack rates on both novel and palatable prey. Although these data reveal the first evidence for protection from avian predators by the supposed toxic, Rubiaceae-feeding Adelpha species, we conclude that imprecise mimetic patterns have high costs for Batesian mimics in the tropics.
悌蛺蝶屬 (Adelpha) 主要分佈於新熱帶區,是個物種多樣性極高的複雜類群,目前包含大約85個已知種。這麼高多樣性的類群當然吸引科學家的目光,除了高多樣性的快速演化與起源在近年的相關研究,另一個關注的焦點在其中有一個翅紋樣式在整個屬內多次演化出現,雖然一直被認為是屬內的擬態關係,但相關研究其實不多。
悌蛺蝶屬 (Adelpha)的翅紋多樣性與分布 photo from Lynsey Bunnefeld |
photo from Finkbeiner et al. 2012 |
這個研究在哥斯大黎加進行,在野外針對應該是被擬態者,專食於茜草科的 Adelpha iphiclus ,與另一種廣食種 Adelpha serpa,測試兩者間的翅紋是否存在擬態關係,亦或是單純的趨同。為何是茜草科呢?數個研究指出茜草科的物種含有會讓草食動物避忌的植物鹼化合物,因此專食於茜草科的物種也很有可能在體內存有這類化合物,讓天敵避開 A. iphiclus。實驗用的蝴蝶是將紙翅膀印出後,黏在黏土做的身體上 (如右圖),並且以光學的方法確定其反射光譜與真實的蝴蝶相似後,再放至野外進行實驗,也就是如果在環境中的捕食者已經被「教育」過了,那麼應該會避免 A. iphiclus,而如果與 A. serpa 間有擬態關係的話,那麼兩者間的被攻擊率應該會十分接近。
結果相當出人意料:A. iphiclus 的被攻擊率相較做為控制組的 Junonia 與另一種 A. leucopthalma來說是顯著的最低,但居然也顯著的低於與斑紋十分相似的 A. serpa!難道兩者間沒有擬態關係嗎?
這個結果代表幾件十分重要的事:1) 雖然這個相似的斑紋一直被當成是擬態的例子,但從結果來看很明顯並非如此,而是A. iphiclus這個物種的斑紋對捕食者是警戒色,但 A. serpa 的斑紋則沒有類似的作用;2) 這也是個很好的例子,即使從人類的角度來看十分相似 ,對捕食者 (在本研究中是鳥類) 來說還是能夠分辨其差異,這跟其他的不精確擬態群有點相似,在環境中可以接受不小的差異 (如食蚜蠅),但這個例子是看起來精確,但可能是極度不精確的貝氏擬態關係;3) 雖然這個研究不支持兩者間的擬態關係,但可能是某些環境因素造成這樣的結果,像是兩者間的頻率或是防禦程度在族群內變動等,也就是可能在其他區域的結果可能不同。
本實驗中所使用的物種與統計結果 |
相關文獻
- Mullen SP, Savage WK, Wahlberg N & Willmott KR. 2011. Rapid diversification and not clade age explains high diversity in neotropical Adelpha butterflies. Proceedings of the Royal Society B: Biological Sciences 278: 1777–1785
- Ebel ER, Dacosta JM, Sorenson MD, Hill RI, Briscoe AD, Willmott KR & Mullen SP. 2015. Rapid diversification associated with ecological specialization in Neotropical Adelpha butterflies. Molecular Ecology 24: 2392–2405
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