擬態生物學 - 多態型 (polymorphism)

Fig. 1 本研究定序的目標物種的分佈與其相關類群的親緣關係

標題：Butterfly genome reveals promiscuous exchange of mimicry adaptations among species

摘要

The evolutionary importance of hybridization and introgression has long been debated. Hybrids are usually rare and unfit, but even infrequent hybridization can aid adaptation by transferring beneficial traits between species. Here we use genomic tools to investigate introgression in Heliconius, a rapidly radiating genus of neotropical butterflies widely used in studies of ecology, behaviour, mimicry and speciation. We sequenced the genome of Heliconius melpomene and compared it with other taxa to investigate chromosomal evolution in Lepidoptera and gene flow among multiple Heliconius species and races. Among 12,669 predicted genes, biologically important expansions of families of chemosensory and Hox genes are particularly noteworthy. Chromosomal organization has remained broadly conserved since the Cretaceous period, when butterflies split from the Bombyx (silkmoth) lineage. Using genomic resequencing, we show hybrid exchange of genes between three co-mimics, Heliconius melpomene, Heliconius timareta and Heliconius elevatus, especially at two genomic regions that control mimicry pattern. We infer that closely related Heliconius species exchange protective colour-pattern genes promiscuously, implying that hybridization has an important role in adaptive radiation.

Fig. 2 a. 毒蝶染色體與家蠶染色體同源基因的比較圖，染色體間的連線代表同源基因的來源
b. 毒蝶化學感受蛋白與大樺斑蝶、家蠶間的關係，顯示出蝴蝶獨有與毒蝶獨有的擴張

Fig. 4 這個分析顯示控制擬態斑紋的基因座B/D的在族群間的基因交換證據

這篇是「毒蝶是否真的交換了控制擬態斑紋的基因？」的主要基礎，作者群是很威猛的毒蝶基因體協會 (The Heliconius Genome Consortium)，透過定序數個Heliconius melpomene族群的全基因體，比較毒蝶的基因體變化與鱗翅目基因體的演化。作者群從基因體中挑出12669基因出來作各種分析，首先比較毒蝶與家蠶 (Bombyx mori)在染色體學上的差異，大部分的基因都能在家蠶上發現，但染色體位置有所差異。其次將化學感受蛋白(Chemosensory protein, CSP)的序列，加上家蠶與大樺斑蝶 (Danaus plexippus)，以系統發生學的方式分析，發現蝴蝶特有與毒蝶特有的基因擴張。毒蝶內部基因體分析則發現族群間或是共擬態者間的基因交換頻繁，特別是兩個控制擬態斑紋的基因，顯示這樣的交換可能對於毒蝶的迅速種化與斑紋的多樣化有關。

Wrote by Chia-Hsuan Wei

Fig. 1 實驗區域與實驗中所使用的表型
Hom. 同型合子
Het. 異型合子
exotic 非本地型

標題：Crossing fitness valleys: empirical estimation of a fitness landscape associated with polymorphic mimicry

摘要

Characterizing fitness landscapes associated with polymorphic adaptive traits enables investigation of mechanisms allowing transitions between fitness peaks. Here, we explore how natural selection can promote genetic mechanisms preventing heterozygous phenotypes from falling into non-adaptive valleys. Polymorphic mimicry is an ideal system to investigate such fitness landscapes, because the direction of selection acting on complex mimetic colour patterns can be predicted by the local mimetic community composition. Using more than 5000 artificial butterflies displaying colour patterns exhibited by the polymorphic Mullerian mimic Heliconius numata,we directly tested the
role of wild predators in shaping fitness landscapes.We compared predation rates on mimetic phenotypes (homozygotes at the supergene controlling colour pattern), intermediate phenotypes (heterozygotes), exotic morphs (absent from the local community) and palatable cryptic phenotypes. Exotic morphswere significantly more attacked than local morphs, highlighting predators’ discriminatory capacities. Overall, intermediates were attacked twice asmuch as local homozygotes, suggesting the existence of deep fitness valleys promoting strict dominance and reduced recombination between supergene alleles. By including information on predators’ colour perception, we also showed that protection on intermediates strongly depends on their phenotypic similarity to homozygous phenotypes and that ridges exist between similar phenotypes, which may facilitate divergence in colour patterns.

在先前的導讀中提到，控制穆氏擬態的毒蝶 (Heliconius) 斑紋基因的顯性性狀比較佔有優勢，中間型或隱性性狀能控制斑紋的機會相對減弱，而顯隱性與中間型又分別代表同型合子 (homozygous) 與異型合子 (heterozygous)，雖說遺傳結構支持顯性性狀比較容易表現，可這畢竟是實驗室內觀測到天擇的結果，在已經知道超基因的控制與顯隱性的優勢後，再次回到野外的話，這些斑紋的適存度 (fitness)又會有什麼樣的故事？

這篇研究就在討論我們已經知道穆氏擬態的毒蝶具有多態型，而且不同區域的多態型的遺傳結構各有不同，顯隱性表現也有各自的差異，但這是因為遺傳結構的限制，或是自然環境中天擇的影響，控制某些基因型的選汰還不清楚，在已經知道這麼多的基因體結構與其跟天擇連結的推測後，作者群再次把故事拉回野外實驗室中，測試天擇比較偏好同型或異型合子。

作者群依據Heliconius numata不同區域的型態，分別放置同型、中間型與非本地型的人工蝴蝶到野外環境中，總共放了超過5000隻的樣本到野外中，經過三天後回收樣本，檢查是否有U型或V型的攻擊痕跡，必且根據對應的形狀判斷是來自哪種捕食者的攻擊。

Fig. 2 不同表型的被攻擊次數，顯示中間型與非當地型被攻擊次數顯著較高
BB aa 同型合子
Ba 異型合子
exotic 非本地型

Fig. 3 不同表型在各種視覺模式下的關聯性

綜合各區域的結果顯示，同型合子個體受到的攻擊率顯著較低，異型合子（中間型）個體與非本地型的攻擊率顯著較高至同型合子的兩倍，支持這些斑紋的在不同場域的適存度有深度的分化。最後，若結合捕食者的視覺模型來看的話，發現中間型的適存度與當地的擬態型相似度有密切的關係，結合之前的研究來看，這個現象也會貢獻在斑紋的分化上。

<<相關閱讀>>

Wrote by Chia-Hsuan Wei

Heliconius numata的斑紋多樣性

photo: Keith. S. Brown Jr. (1979)

標題：Evolution of dominance mechanisms at a butterfly mimicry supergene

摘要

Genetic dominance in polymorphic loci may respond to selection; however, the evolution of dominance in complex traits remains a puzzle. We analyse dominance at a wing-patterning supergene controlling local mimicry polymorphism in the butterfly Heliconius numata. Supergene alleles are associated with chromosomal inversion polymorphism, defining ancestral versus derived alleles. Using controlled crosses and the new procedure, Colour Pattern Modelling, allowing whole-wing pattern comparisons, we estimate dominance coefficients between alleles. Here we show strict dominance in sympatry favouring mimicry and inconsistent dominance throughout the wing between alleles from distant populations. Furthermore, dominance among derived alleles is uncoordinated across wing-pattern elements, producing mosaic heterozygous patterns determined by a hierarchy in colour expression. By contrast, heterozygotes with an ancestral allele show complete, coordinated dominance of the derived allele, independently of colours. Therefore, distinct dominance mechanisms have evolved in association with supergene inversions, in response to strong selection on mimicry polymorphism.

求學的過程中大概多少都學過孟德爾（遺傳學之父，不知道他是誰的請面壁）的豌豆實驗，從那時候開始我們學過棋盤格方法，推演顯性與隱性間性狀在子代身上出現的比例。孟德爾的遺傳實驗讓我們知道兩件事：1）如果要知道性狀的顯隱性，要各種自交雜交回交子代間雜交再雜交 2）如果實驗對象沒辦法做到容易飼養這件事，想知道性狀的顯隱性簡直天方夜譚。到了現在，對於某些物種性狀的顯隱性我們可以容易的察覺，像人類、果蠅這類知之甚詳物種，但對於更多那些只知道現象，但對性狀的顯隱性卻很難推斷，而且性狀的變化對物種的演化有何優勢，我們瞭解的還只是表層的現象而已。

在開始之前這篇前，先釐清一件事：可能有人覺得顯性比起隱性在天擇上更有優勢，因為在遺傳上比較容易表現出來，但實際上不是如此，隱性基因的頻率也可能比顯性基因來的高，一出現顯性性狀就被篩選，當然反過來也有可能。知道這件事後，這篇的概念就是想瞭解一種在南美洲參與穆氏擬態的毒蝶，Heliconius numata，的超基因(supergene)（註）如何控制斑紋顯隱性在翅紋上的分佈，以及這在演化上哪種更具有優勢。毒蝶很符合好操作好生好養的條件，首先透過很多的雜交取得同型合子(homozygote)的個體(AA, aa)，然後再度雜交，就能取得異型合子的個體(Aa)，透過三種基因型的比較，我們就知道哪些斑紋是顯性或隱性，再跟實際上野外捕捉的個體相比，就能知道顯性或隱性在自然環境哪種比較佔有優勢。

（註：超基因指數個控制某些性狀的基因座，因為天擇與分子機制的調控而集中到某處，看起來就像一個很大的基因座控制與微調全部的性狀，以這個例子來說，這個超基因控制的就是擬態斑紋的表現，而實際上如何透過基因倒位/插入/反轉來調控，又是另一個故事）

Fig. 2 上半部是共域個體雜交結果，下半部是半共域個體雜交結果，黑色背景圖為基因表現相對顯性程度，可以看出共域個體的顯性基因所佔比例較高

Fig. 3 異型合子的個體顯示參與擬態的顏色相對表現程度

作者群藉由比較不同族群的擬態斑紋型態間的雜交，找出斑紋的顯隱性，發現共域的個體間雜交，會出現與顯性基因比較相近的斑紋，而異域雜交的個體就不一定，顯示在某個區域內，顯性遺傳是比較佔有優勢的。作者群還做了一個有趣的分析，毒蝶的斑紋主要由黑、黃、橘所組成，那麼斑紋的變化與不同區域的顏色比例有關，這些顏色間是否有相對的顯隱性？結果發現這些透過六個基因座所控制的顯性性狀，其顏色的顯隱性排序是黑色>橘色>黃色。透過這些遺傳學的結果，讓我們對於擬態的多態性斑紋變化，以及族群間的優勢斑紋的維持機制又有更深一層的認識。

最後要說的是，這樣的研究是透過長期累積的基因體研究、野外的資料與無數的養蟲雜交才能有這樣的成果，而且還有物種本身單態性的優勢，讓推測遺傳性狀上不那麼困難。若是像大鳳蝶這種雄性單態性，雌性多態性的物種，光是要知道雄性是不是同型合子就很困難，推測顯隱性更是一大挑戰。

Wrote by Chia-Hsuan Wei

Wood tiger moth (Parasemia plantaginis) (wiki)

標題：Changes in predator community structure shifts the efficacy of two warning signals in Arctiid moths

摘要

1. Polymorphism in warning coloration is puzzling because positive frequency-dependent selection by predators is expected to promote monomorphic warning signals in defended prey.

2. We studied predation on the warning-coloured wood tiger moth (Parasemia plantaginis) by using artificial prey resembling white and yellow male colour morphs in five separate populations with different naturally occurring morph frequencies.

3. We tested whether predation favours one of the colour morphs over the other and whether that is influenced either by local, natural colour morph frequencies or predator community composition.

4. We found that yellow specimens were attacked less than white ones regardless of the local frequency of the morphs indicating frequency-independent selection, but predation did depend on predator community composition: yellows suffered less attacks when Paridae were abundant, whereas whites suffered less attacks when Prunellidae were abundant.

5. Our results suggest that spatial heterogeneity in predator community composition can generate a geographical mosaic of selection facilitating the evolution of polymorphic warning signals. This is the first time this mechanism gains experimental support. Altogether, this study sheds light on the evolution of adaptive coloration in heterogeneous environments.

如果警戒性斑紋有用的話，我們期待這個物種在生態與演化上有什麼反應？是不是這個訊號要很穩定，就像走在鋼索上，如果不小心出現什麼有點跟原本不一樣的，立刻遭受攻擊的機會就上升了？是不是這個訊號要很明顯，能夠很有效率的傳達給捕食者，最好是看到就覺得這東西危險，連碰都別碰？

如果有這麼簡單就好了。自然界的運作原則永遠比我們想的複雜，因為我們總是會發現某些警戒性的物種具有兩種以上的斑紋，如果理論預測是對的，那麼實際上就是某些還不瞭解的運作機制參與其中，多態型對警戒性的運作對象會產生什麼影響？

Fig. 1 作者所參考的標本（上排）與人工假蛾（下排）

本篇討論的主角：wood tiger moth (Parasemia plantaginis) (有些翻譯為車前燈蛾)是一種廣部於舊北區的燈蛾，食草廣泛，在大英的寄主資料庫中查到取食杜鵑花科、蓼科、車前草科、樺木科與菊科，而這種類隨著食草不同，體內的防禦與免疫程度也會隨之變動。這種蛾主要有兩個顏色：黃色與白色，配上黑色的條紋顯示出極度明顯的對比，因此被認為是警戒性的物種。

作者群想要知道不同的顏色在不同區域的比例差異從何而來，是因為捕食者的族群結構差異，或是受限於當地族群顏色出現的比例（出現的越多次，捕食者越容易學到，稱為frequency-dependent selection）？作者用塑像用黏土製成身體，以數位拍攝真正的標本後印出，將翅膀裁切下來黏在黏土身體上，製成與真蛾標本同樣大小的人工假蛾進行實驗。實驗在五個地方進行：愛沙尼亞、芬蘭中部、芬蘭南部、蘇格蘭與奧蘭，每個地方選取15條穿越線（奧蘭16條），各自放置以下10隻不同外觀的假蛾：白色/黃色普通型、白色/黃色較黑色型以及控制組（共50隻），每隻樣本的距離為20公尺，並且記錄U型與V型的鳥攻擊痕跡。

山雀科 (Paridae)
photo: Handbook of the birds of the world

岩鷚科 (Prunellidae)
photo: Ren Hathway in Handbook of the Birds of the World. V. 10

結果發現無論是哪個區域，黃色被攻擊的頻率都比起白色來的高，這似乎與當地色型出現的頻率無關。但作者群發現另一個有趣的現象：當山雀科 (Paridae) 的鳥類比較多的時候，黃色的被攻擊率就比較低；當岩鷚科 (Prunellidae) 的鳥類比較多的時候，白色的被攻擊率就比較低。這是第一個支持空間的異質性讓捕食者結構不同時，對於警戒性的斑紋有不同程度的選汰壓力的證據。

簡單說個總結，其實就像人類不同的飲食文化一樣，不同的物種，不同的族群喜歡吃的東西不同，因此造成當地產出的差異。可是我認為這個研究有些邏輯上的衝突：如果這個燈蛾警戒性本身很有用，那麼捕食者應該能不吃就不吃，也就是捕食者應該已經「被教育」的好好的，如果是這樣，作者群放出去的假蛾應該會完完整整的沒有任何攻擊痕跡；但作者群的數據來源卻必須基於攻擊燈蛾的咬痕，也就是這個燈蛾本身就是捕食者的食物之一，才有可能蒐集到這樣的數據。那麼，這個燈蛾的警戒效率是不是還真值得懷疑？

至於這些鳥的視覺模型的重要性自然更不在話下。

Wrote by Chia-Hsuan Wei

photo from Papilio dardanus (wiki)

標題：Characterising the phenotypic diversity of Papilio dardanus wing patterns using an extensive museum collection.

摘要

The history of 20th Century evolutionary biology can be followed through the study of mimetic butterflies. From the initial findings of discontinuous polymorphism through the debates regarding the evolution of mimicry and the step-size of evolutionary change, to the studies on supergene evolution and molecular characterisation of butterfly genomes, mimetic butterflies have been at the heart of evolutionary thought for over 100 years. During this time, few species have received as much attention and in-depth study as Papilio dardanus. To assist all aspects of mimicry research, we present a complete data-derived overview of the extent of polymorphism within this species. Using historical samples permanently held by the NHM London, we document the extent of phenotypic variation and characterise the diversity present in each of the subspecies and how it varies across Africa. We also demonstrate an association between ‘‘imperfect’’ mimetic forms and the transitional race formed in the area where Eastern and Western African populations meet around Lake Victoria. We present a novel portal for access to this collection, www.mimeticbutterflies.org, allowing remote access to this unique repository. It is hoped that this online resource can act as a nucleus for the sharing and dissemination of other collections databases and imagery connected with mimetic butterflies.

非洲白鳳蝶的多態型，左上方框內是雄性，線的上方是各種斑蝶與細蝶，線的下方是各型雌性白鳳蝶
photo from mimeticbutterflies.org

非洲的白鳳蝶(Papilio dardanus)是知名的貝氏擬態的鳳蝶，其豐富的多態型各自擬態非洲各地的細蝶、斑蝶等不好吃的種類。白鳳蝶的擬態實際上只有雌性參與其中，而雄性只有一種型態，在多態型的雌性中，除了擬態各種不同的蝴蝶外，神奇的也存在雄性型的雌性。

1960年代，透過孟德爾式的遺傳學雜交試驗瞭解不同族群間的斑紋變化，而且結果顯示白鳳蝶的斑紋變化是來自一個超基因；到了2008年，基於現代的分生技術，終於找到超基因的位置。即使如此，雌性的型態實在太多了，除了上面這些看起來精準擬態的斑紋外，也有更多看起來是中間型的種類，因此雖然已經研究到基因體層級的情況下，基礎的斑紋與地理分佈還沒有整體性的調查過。

雌性各種不同的型與名稱
photo from Fig. 1

這篇透過英國倫敦自然史博物館(the Natural History Museum, London)的極大量鱗翅目館藏，蒐集標籤上的採集資訊，將採集的地點轉換成現代的地點並查出座標，把各個不同的斑紋分類後，再轉換為二維座標系統輸入地理資訊系統 (GIS)，把各個不同的斑紋分類後，就能每個型態的分佈呈現在現代的地圖上，並且透過採集點的連線，推估分佈範圍。

取樣密度與地點
photo from Fig. 3

各型的分佈範圍
photo from Fig. 6

結果就如上圖，把大部分的斑紋的分佈範圍推估出來，而那些看起來不完美擬態的斑紋分佈於維多利亞湖附近，支持這是西部與東部族群相遇的雜交區。

中間型的分佈區域
photo from Fig. 7

這分析的概念很簡單，但卻是瞭解一個擬態群的多樣性最快的方式，而且最重要的，這仰賴保存完整與大量的標本館藏。這樣的研究說明透過良好的想法與完整的分析，即使基礎的研究也能從中得到大量的資訊，也更凸顯標本蒐藏的重要性。

Wrote by Chia-Hsuan Wei

photo from Ecuatoria: Dendrobatids

標題：Number of genes controlling a quantitative trait in a hybrid zone of the aposematic frog Ranitomeya imitator

摘要

The number of genes controlling mimetic traits has been a topic of much research and discussion. In this paper, we examine a mimetic, dendrobatid frog Ranitomeya imitator, which harbours extensive phenotypic variation with multiple mimetic morphs, not unlike the celebrated Heliconius system. However, the genetic basis for this polymorphism is unknown, and not easy to determine using standard experimental approaches, for this hard-to-breed species. To circumvent this problem,we first develop a newprotocol for automatic quantification of complex colour pattern phenotypes from images. Using this method, which has the potential to be applied in many other systems, we define a phenotype associated with differences in colour pattern between different mimetic morphs. We then proceed to develop a maxi- mum-likelihood method for estimating the number of genes affecting a quantitative trait segregating in a hybrid zone. This method takes advantage of estimates of admixture proportions obtained using genetic data, such as microsatellite markers, and is applicable to any other system where a phenotype has been quantified in an admixture/introgression zone. We evaluate the method using extensive simulations and apply it to the R. imitator system. We show that probably one or two, or at most three genes, control the mimetic phenotype segregating in a R. imitator hybrid zone identified using image analyses.

photo from content

用來研究擬態系統的分類群中，除了毒蝶 (Heliconius)目前有系統的從生態到基因體，很少有例子能夠做到這種程度，主要是因為毒蝶好抓好生好養，可以容易的透過孟德爾式遺傳的方式找出性狀的變化，進而推估控制斑紋的基因數量、是否聯會(linkage)等遺傳上的資訊。很多其他的生物沒辦法做到這樣的研究，就是因為數量少難生難養，或是生活史世代太長，沒辦法進一步的探索這些資訊，進而推估背後的演化來源等等相關理論。

但科學家總是有辦法的。精靈箭毒蛙 (Ranitomeya imitator)是一種有多態性的物種，像毒蝶一樣，每一個不同區域的樣式都擬態當地的model，但箭毒蛙就是一個難生難養，難以取得大量子代的物種，這個研究就假設「如果一個基因控制能控制一種變異，那麼從透過分析表型的變異程度，應該可以知道到底有幾個基因控制這些表型」。作者透過把高解析度的照片弄成如上圖展示擬態的樣式，透過文中所寫的likelihood model，配上一些微衛星體的分子資料的比較，模擬究竟有幾種基因牽涉其中。

photo from content

模擬的結果顯示看起來是兩個，至多三個基因控制這些擬態的斑紋。這個方式有很多好處：

1) 只要透過照片就能分析，不需大量採集野外個體

2) 省去飼養的時間

3) 不需要花大錢全基因體定序就能有結果

這種模擬的結果，有點類似前測試，或是只能知道虛無假說的方向，提供一個推估同樣難生難養的分類群的遺傳資訊的方法，可是基因實際上有幾個，分佈在哪些染色體上，是否形成超基因 (supergene) ，對於演化的歷程有沒有影響，是無法從這個方法得知的。

Wrote by Chia-Hsuan Wei

photo from J. P. Lawrence

標題：Colour and Escape Behaviour in Polymorphic Populations of an Aposematic Poison Frog [pdf]

摘要

The phenomenon of aposematism, or the pairing of antipredator defence with conspicuous or distinctive signals, serves as an excellent example of how traits act in concert to shape fitness. Not only does this complex phenotype require the integration of multiple traits, it alters the fitness pay-offs of yet others. The protection offered by aposematism may, for example, reduce the costs associated with foraging or sexual display. Thus, well-protected aposematic lineages should be bolder, more active and less likely to respond to perceived threats of predation than more cryptic lineages. Comparisons of differently coloured morphs of the poly-typic strawberry poison frog (Oophaga pumilio) have supported the predicted behavioural correlates of aposematism, with the exception of those regarding responsiveness to simulated predators. We tested the key prediction that aposematic coloration will be associated with reduced sensitivity to predators in two polymorphic O. pumilio populations. The novel approach of studying polymorphic populations allowed us to assess the effect of colour in the absence of potentially confounding habitat differences. We found that colour was associated with the probability that a frog would attempt escape and the distance at which it fled, but only in one population, and not in the predicted direction. An overall comparison of the two populations superficially supported our predictions, but this pattern actually arose because frogs occupying higher perches were less likely to respond, a pattern that may reflect the value of high perches and the costs associated with returning to them after attempted escape. These results highlight the complexity of the relationship between predators and prey, the challenges associated with understanding how and why traits are correlated, and the intimate ties between behaviour and morphological evolution.

警戒色是生來嚇阻警告捕食者，告訴牠們「我不好吃別吃我」的視覺訊號，這對擁有警戒色的物種的好處是避免被攻擊與提高生存率。既然如此，是不是就代表這些物種在外出覓食或是求偶的時候，可以花費比較少的能量躲避捕食者呢？（就是我夠兇不用躲的意思）

這個研究用草莓箭毒蛙 (Oophaga pumilio)做為研究對象，草莓箭毒蛙本身就是具有多態性的物種，從看起來警戒到看起來隱蔽的樣式都有，因此可以降低不同種間遺傳結構所產生的行為差異，很適合做為討論這個問題的標的物種。作者選定兩個地方的族群 Isla Bsastimentos與Dolphin bay （都在巴拿馬），作者實際到這兩處的雨林中蒐集資料，透過穿越線的方式找尋野生的個體，然後以人類做為捕食者慢慢靠近的方式進行實驗。首先選定兩個類型的體色：有警戒色的與比較黯淡的，評估族群間是否有體色上的差異（沒差，有差就不用做了），接著評估兩種行為1) 個體看到捕食者時是否企圖要逃跑 2) 捕食者與個體開始逃跑間的最短距離 (flight initiation distance, FID)。結果顯示兩個區域間企圖逃跑的比例不一樣，在Bastimentos的族群有企圖逃跑行為的比Dolphin Bay的要少，如果與顏色有關的話，那麼兩者間的比例應該會很接近，這邊顯示只有其中一個族群反應作者的假說。另一個分析顯示這些個體要不要討跑與所停的高度(perch height)比較有關，與作者的假說相關度比較低，顯示停棲高度可能與對能不能發現捕食者，進而引發逃跑反應比較有影響。

flight initiation distance (FID) wiki

其實這個研究的前提有點弔詭，如果一個物種有防禦+有亮眼的顏色＝警戒性，那麼有防禦+黯淡的顏色就不等於警戒性了嗎？因為個體還是有毒，因此這個「黯淡的顏色」還是有可能被天敵當成某種避忌訊號，只是以人類的視覺角度來看的話，就會變成沒有要不要逃跑取決於其他因子而不是因為有沒有警戒色。

Wrote by Chia-Hsuan Wei

photo from Lorquin's admiral (wiki)

photo from Adelpha californica (wiki)

標題：A test of fundamental questions in mimicry theory using long-term datasets

摘要

Since the phenomenon of mimicry was first described by Bates in 1862 it has become one of the foundational examples of adaptive evolution. Numerous subcategories of mimicry and dozens of hypotheses pertaining to its evolution and maintenance have been proposed. Many of these hypotheses, however, are difficult to test in experimental settings, and data from natural observations are often inadequate. Here we use data from a long-term survey of butterfly presence and abundance to test several hypotheses pertaining to Batesian and female-limited polymorphic mimicry (FPM; a special case of Batesian mimicry). We found strong evidence that models outnumber mimics in both mimicry systems, but no evidence for an increase in relative abundance of FPM mimics to their Batesian counterparts. Tests of the early-emergence/model first hypothesis showed strong evidence that the Batesian mimic routinely emerges after the model, while emergence timing in the FPM system was site specific, suggesting that other ecological factors are at play. These results demonstrate the importance of long-term field observations for testing evolutionary and ecological hypotheses.

如果擬態群在自然的情況下是有效的，那麼這個擬態群可能符合一些基本的前提，像是model比起mimic來說數量較多、比較難吃、時序上比較早出現等等，都是這個擬態群「應該」要符合的。雖說如此，能夠支持這些基本假設的證據卻是少之又少，也就代表這些基本的假設究竟是科學家自己的想像，還是真正發生在自然界呢？

這個研究透過累積十年的蝴蝶記錄資料來探討這個問題，資料記錄的地點在北美洲，分別是Gates Canyon、Washington與Lang Crossing（詳細資訊見butterfly.ucdavis.edu），記錄的時間大約是每14天一次，累積了1999-2010間的資料。

他們想要討論的問題在於如果貝氏擬態的mimic太多，會導致整個系統的保護作用降低或失效，因此mimic絕對不能比model還多，因此一個可能性就是「model在自然界中絕對數量比mimic多」，另一個可能性是如果mimic只有其中一個性別擬態，那麼相對來說mimic的數量也會減少，因此他們選了兩個系統，其中一個是代表貝氏擬態的系統，Adelpha californica (model)與Limenitis lorquinii (mimic)（見上圖），另一個是雌性擬態的系統（見下圖）。

結果顯示兩個系統中，model的數量都遠多於mimic。發生的時序上來說，貝氏擬態的mimic出現時序都晚於其model，而雌性擬態的物種出現則局現在某幾個地方，可能有其他的生態因子限制其分佈。

photo from Euphydryas chalcedona (wikimedia) (left) and Butterflies of America (right)

photo from Wild Utah (left) and Butterflies of America (right)

上排是model，Euphydryas chalcedona，是一個有兩型的物種；下排是mimic，Chlosyne palla，是一個雌性多態型的物種，作者認為左下的黑色型擬態左上的黑色型

這個例子是個以溫帶物種來說明的例子，但在熱帶/亞熱帶區域卻總有些例子是相反的（mimic早於model出現），雖然已經有些行為學上的證據支持這樣的現象發生，若有可能的話，透過類似的觀察與紀錄累積，或許熱帶/亞熱帶區域的結果會不一樣。（第一個問題：去哪找人合作觀察十年？）

Wrote by Chia-Hsuan Wei

photo from Elias Marianne

標題：Diversity in Müllerian mimicry: The optimal predator sampling strategy explains both local and regional polymorphism in prey

摘要

The convergent evolution of warning signals in unpalatable species, known as Müllerian mimicry, has been observed in a wide variety of taxonomic groups. This form of mimicry is generally thought to have arisen as a consequence of local frequency-dependent selection imposed by sampling predators. However, despite clear evidence for local selection against rare warning signals, there appears an almost embarrassing amount of polymorphism in natural warning colors, both within and among populations. Because the model of predator cognition widely invoked to explain Mullerian mimicry (Muller’s “fixed nk” model) is highly simplified and has not been empirically supported; here, we explore the dynamical consequences of the optimal strategy for sampling unfamiliar prey. This strategy, based on a classical exploration–exploitation trade-off, not only allows for a variable number of prey sampled, but also accounts for predator neophobia under some conditions. In contrast to Müller’s “fixed nk” sampling rule, the optimal sampling strategy is capable of generating a variety of dynamical outcomes, including mimicry but also regional and local polymorphism. Moreover, the heterogeneity of predator behavior across space and time that a more nuanced foraging strategy allows, can even further facilitate the emergence of both local and regional polymorphism in prey warning color.

在開始前，我們先思考一件事：如果擬態是十分有用的禦敵策略，那麼物種在演化的初期發展出一個紋路後，是否應該其他的物種都加入擴大這個擬態群，讓這個紋路變的廣泛的有用結果大家都長的一樣，更增強這個紋路的作用才對？

但現實中明顯不是這樣，各種不同區域的擬態群各自林立，物種內出現各種不同的多態型，這代表上面那個假設必然出了什麼狀況，才會讓這個假設與實際上差異這麼大。

這篇所使用的方法是大家都頭痛的數理模型，考慮不同的參數後進行模擬，通常這類型的文章考慮的不外乎有幾個因子：防禦/非防禦獵物的比例，捕食者攻擊的頻率，記憶力，一些學習參數等等，而這個研究考量的是「捕食者的最佳取食策略」，也就是透過計算攻擊防禦/非防禦獵物的比率，來達到利益的最大化，然後跟傳統的穆氏擬態的數學模型（捕食者一定要透過攻擊固定數量的獵物才能讓利益最大化）比較。結果顯示這個新的模型比較能夠解釋現實上的狀況，也就是解釋為何擬態中的多態型能夠穩定的存在。捕食者透過挑選防禦/非防禦能讓捕食效益產身變化外，其本身的恐新症（neophobia）在某種情況下也會成為影響挑選獵物的因子之一。

結論上來說，就是用數理的方式告訴大家捕食者如何維持多態型的穩定，不過這個穩定在之前的研究中也有些討論，可能是來自微棲地的分隔所造成。

老話一句，電腦上看起來是這樣，那在自然環境中看起來又是怎樣呢？

Wrote by Chia-Hsuan Wei

（圖片選自內文）

原文標題：Cryptic differences in colour among Müllerian mimics: how canthe visual capacities of predators and prey shape the evolution of wing colors?

[摘要] [原文網址]

Antagonistic interactions between predators and prey often lead to co-evolution. In the case of toxic prey, aposematic colours act as warning signals for predators and play a protective role. Evolutionary convergence in colour patterns among toxic prey evolves due to positive density-dependent selection and the benefits of mutual resemblance in spreading the mortality cost of educating predators over a larger prey assemblage. Comimetic species evolve highly similar colour patterns, but such convergence may interfere with intraspecific signalling and recognition in the prey community, especially for species involved in polymorphic mimicry. Using spectrophotometry measures, we investigated the variation in wing coloration among comimetic butterflies from distantly related lineages. We focused on seven morphs of the polymorphic species Heliconius numata and the seven corresponding comimetic species from the genus Melinaea. Significant differences in the yellow, orange and black patches of the wing were detected between genera. Perceptions of these cryptic differences by bird and butterfly observers were then estimated using models of animal vision based on physiological data. Our results showed that the most strikingly perceived differences were obtained for the contrast of yellow against a black background. The capacity to discriminate between comimetic genera based on this colour contrast was also evaluated to be higher for butterflies than for birds, suggesting that this variation in colour, likely undetectable to birds, might be used by butterflies for distinguishing mating partners without losing the benefits of mimicry. The evolution of wing colour in mimetic butterflies might thus be shaped by the opposite selective pressures exerted by predation and species recognition.

在一些之前的研究中，我們已經知道毒蝶的穆氏擬態中，不同類群的斑紋是由不同的基因調控而來（簡單的說就是趨同），那麼理論上不同的物種間的斑紋顏色應該會多少有差異，若是顏色的差異太大的話，就可能脫離擬態群被捕食者攻擊，因此這些顏色的差異是在一個可容忍的「範圍」內的，事實上來說，我們也的確認為這些擬態者的顏色幾乎沒有差異，但若是顏色幾乎沒有差異的話，各物種是如何辨別同或不同種呢？

這篇研究用光譜儀測試了翅面上的黃、橘與黑色，發現在兩個不同屬間，雖然看起來很像，但在亮度、色調與濃度上各有顯著的差異，那麼，如果顏色是有差異的，穆氏擬態或種內辨識是否就會有所衝突？

這從生理學上得到解釋，捕食者的視覺系統無法分辨這麼細微的差異，而蝴蝶的視覺可分辨這些差異，因此「我們」看起來幾乎無差異的顏色，經由天擇的選汰形成擬態群，蝴蝶又可藉由這些差異分辨種類。（註）

註：在早期的研究中，已發現毒蝶可經由視覺選擇與自身斑紋較類似的個體交尾，因此毒蝶是可藉由視覺分辨個體間差異的。

Wrote by Chia-Hsuan Wei

原文標題：Wing patterning gene redefines the mimetic history of Heliconius butterflies

摘要 [原文網址]

The mimetic butterflies Heliconius erato and Heliconius melpomene have undergone parallel radiations to form a near-identical patchwork of over 20 different wing-pattern races across the Neotropics. Previous molecular phylogenetic work on these radiations has suggested that similar but geographically disjunct color patterns arose multiple times independently in each species. The neutral markers used in these studies, however, can move freely across color pattern boundaries, and therefore might not represent the history of the adaptive traits as accurately as markers linked to color pattern genes. To assess the evolutionary histories across different loci, we compared relationships among races within H. erato and within H. melpomene using a series of unlinked genes, genes linked to color pattern loci, and optix, a gene recently shown to control red color-pattern variation. We found that although unlinked genes partition populations by geographic region,optix had a different history, structuring lineages by red color patterns and supporting a single origin of red-rayed patterns within each species. Genes closely linked (80–250 kb) to optix exhibited only weak associations with color pattern. This study empirically demonstrates the necessity of examining phenotype-determining genomic regions to understand the history of adaptive change in rapidly radiating lineages. With these refined relationships, we resolve a long-standing debate about the origins of the races within each species, supporting the hypothesis that the red-rayed Amazonian pattern evolved recently and expanded, causing disjunctions of more ancestral patterns.

在系統發生學中，如果我們想要知道一個特徵的演化起源，也就是大家最想看的「怎麼演化」，通常會有個樹，上面有很多顏色很漂亮，然後你追著那個你想看的顏色，就會知道這個特徵發生了幾次，然後你在看圖說故事（譬如多年前有人探討竹節蟲翅的演化來源一樣）。重點就在於通常我們重建一棵樹的時候，理論告訴我們要選中性或偏中性的分子標記，不能選那些受到強烈天擇作用的標記，這樣才能純粹的反應物種間的關係，然後再用你想看的特徵去追溯，才會知道演化的來源。

而這如果用在擬態的演化這件事上會怎樣？通常你會看到不同擬態樣子的物種混在同一個支系裡，然後你用這個樹去追溯擬態的特徵，九成九會告訴你「所有的擬態特徵都是每個物種獨立演化多次來的」，所以哪個特徵先出現會變得模糊不清，原因就在於這些中性的分子標記是可以在各個物種間傳來傳去的，不一定會跟著強烈天擇的翅紋走，因此基於這些標記重建的樹無法反應擬態特徵的演化歷史。

所以為了知道這個特徵的演化來源與方向，這篇文章就用了控制斑紋的基因重建親緣關係，結果發現不同物種間但同樣斑紋的種類親緣關係較近，也就是在分子的層級上，擁有相同擬態翅紋的物種是趨同的，這就可以用來解釋很多事情，像是提供穆氏擬態的演化假說證據（註）等等。

當然要做到這個層級的前提有很多，像是要先知道各個控制斑紋的基因是哪個，先花大錢把序列全部定出來，實驗動物本身要能夠雜交容易飼養繁殖等等，相較於其他地區的擬態群來說，毒蝶提供很多這類的優勢，自然是個很好的模式動物。

註：穆氏擬態的演化假說是不同物種間的斑紋會越來越像，以求達最適化。

Wrote by Chia-Hsuan Wei

Wrote by Chia-Hsuan Wei

圖片來源：http://www.dendrobase.hu/wp-content/uploads/2010/07/OophagaHistrionica018800_mark_aartse-tuyn.jpg

原文標題：Field but not lab paradigms support generalisation by predators of aposematic polymorphic prey: the Oophaga histrionica complex

摘要 [原文網址]

The persistence of novel aposematic forms, and thereby the evolution of aposematic polymorphism, remain intriguing. Novel and rare forms could be disproportionally attacked by predators that already learned to avoid a pre-existing and more common aposematic form. Alternatively, novel forms could be less frequently attacked if predators are reluctant to attack unknown potential prey (neophobia) or if previous learning allows them to generalise and recognise the novel form as toxic. We used colour variation in polymorphic poison frogs (Oophaga histrionica complex) to test whether predators familiar with one aposematic form do generalise their avoidance behaviour to other aposematic forms. To strengthen our inference, we combined a field test of attack rates to local and non-local models with a lab experiment of generalisation capabilities by newly born chicks. Field predators attacked a significantly lower proportion of 529 aposematic compared to 150 cryptic models. Predators co-occurring with the local aposematic form of O. histrionica equally avoided non-local forms, especially in areas where the species was abundant. Forty-two lab chicks learned to discriminate between an aposematic and a cryptic image, but failed to generalise to other aposematic images, even though we tried with six combinations of aposematic forms. To better mimic the situation in the field, we further tested whether chicks trained with a set of four simultaneous aposematic images would generalise better. They failed to learn the discrimination task. Our data contrast with previous field studies on other poison frogs, and support a role for generalisation, and arguably not neophobia, in predator avoidance of novel aposematic forms.

首先介紹這個箭毒蛙，其實我跟牠不熟，但查了以後發現他真的很多型

圖片來源：http://i795.photobucket.com/albums/yy235/frogandtoadarefriends/Dart%20Frogs/HISTRIONICAGUIDE.jpg

如上圖所示，如果這是真的，那麼這個箭毒蛙至少有40個型，那麼一來這是警戒色，二來捕食者會對這樣的獵物有什麼反應呢？

本文同時進行了野外與實驗室的操作，野外的實驗發現相較於隱蔽的模型而言，捕食者的確較少攻擊具有警戒色的模型，而如果把不是這個區域的型放進來測試的話，當地的捕食者也會避免攻擊這樣的獵物，也就是已經將這些型概括化，而這或許是從捕食者自身的經驗變化而來。另一部分在實驗室的實驗，測試剛出生的小雞對這些模型的反應，結果則顯示雖然小雞能夠分辨警戒色與隱蔽色的差異，但是無法將不同的型概括化。

這個文章很有趣的告訴我們幾件事

1. 野外的情況與實驗室的結果差異甚大，這代表那些用雞做實驗的文章是否需要再探討？

2. 概括化是由經驗累積而來，但出生的小雞的馴化過程是否算是有足夠的經驗累積？或是雞太笨？如果換成其他的鳥或蜥蜴會有一樣的結果嗎？

3. 如果野外的捕食者大多具有概括化能力，那麼就能解釋很多類群具有奇怪的多態型，即使捕食者不是廣域性的也行

4. 獵物本身的每個型似乎還是必須具備警戒性才行，這樣的理論才能成立。

5. 又或者其實族群同時有警戒與隱蔽，但隱蔽性的全被選汰掉了，只有具有警戒性的才能留下來？

好有趣啊，呵呵。

Wrote by Chia-Hsuan Wei

原文標題：Evolution of mimicry patterns in Metriorrhynchus (Coleoptera: Lycidae): The history of dispersal and speciation in Southeast Asia

摘要 [原文網址]

The concept of Müllerian mimicry suggests convergent evolution to an intermediate pattern and does not predict polymorphism in mimicry rings. We examined the evolution of mimicry patterns and the order of divergence of various factors, including the role of aposematic patterns in speciation, in a clade of net-winged beetles with a robust phylogeny that suggests that they dispersed from the Australian to Asian plate. We found strong evidence for the evolution of mimicry via advergence inMetriorrhynchus because older patterns are represented in the Oriental region within more than 100 species of lycids from several lineages. Advergence was likely the cause of the observed intraspecific polymorphism in contrast to the predicted universal monomorphism. Polymorphism was found in populations of two species in Sumatra and Borneo and in populations fine-tuned to subtle variants in various habitats. The advergence is likely to be based on the small population sizes of immigrants. The differences in population sizes result in much higher benefits for dispersing species than native populations. Speciation was trigged by the divergence in aposematic coloration, and the genetic differences accumulated slowly during incomplete isolation. We assumed that the differentiation in genitalia through sexual selection ultimately reinforced speciation initiated by the shift between mimicry patterns.

紅螢是個全世界分布的東西，主要分布在熱帶與亞熱帶，很多的觀察都指出紅螢似乎是個很有用的樣式，諸如一些鱗翅目（如榕蛾）、半翅目（某些星椿）與鞘翅目（如蓬萊紅天牛、金花蟲），都可以看到像是紅螢的影子。紅螢的防禦來自於自體分泌的吡嗪類化合物（pyrazines)與紅螢酸（lycidic acid），這類物質對於大部分的捕食者而言是很有效的避忌物。

這篇文章主要就是在討論紅螢科Metriorrhynchus屬的擬態斑紋的演化，配合生物地理事件加上定年，就可以知道每個節上發生的地理事件，然後利用這些資料推斷擬態的斑紋是否促進種化與限制了分布。這個屬主要的斑紋如下圖，擬態的斑紋就是只1.2/3.4這兩種中段是紅尾段是藍的樣式。利用了系統發生學的方法與定年後，發現擬態的種與其姊妹種（如3.4/5）的遺傳差距其實不高，顯示擬態的斑紋可能促進了族群的分化，而不同區域的擬態群可能來自多態型的分化。

其實這樣的介紹有點太簡單，只把一些比較重要的重點節錄出來，文內討論了種化、斑紋演化、生殖器構造等等，用了很多不同的面象討論這個屬的種化歷程，若要了解的更詳細，還是請看內文吧！

Wrote by Chia-Hsuan Wei

photo from http://www.qwiki.com/q/#!/Peppered_moth

原文標題：Mimicking multiple models: polyphenetic masqueraders gain additional benefits from crypsis

[摘要] [原文網址]

Many prey organisms avoid predation by mimicking inanimate objects: a phenomenon known as masquerade. It is expected that masquerade will show a frequency-dependent advantage such that masquerading species benefit more from their appearance when they are rare in comparison with their models. In such circumstances, selection may favor the coexistence of different phenotypes (polyphenism or polymorphism). The American peppered moth caterpillar Biston betularia cognataria appears to show polyphenetic masquerade: caterpillars found on birch trees look like birch twigs; those on willow trees look like willow twigs. Here, we show in laboratory experiments that the caterpillar does benefit from masquerade and that polyphenism is key to this benefit: avian predators misclassified birch-fed larvae as birch twigs and willow-fed larvae as willow twigs. In a second experiment where the benefits of masquerade were excluded, we show that larvae are less likely to be attacked when located on the host species whose twigs they resemble than when found on an alternative species whose twigs they do not resemble; thus, the polyphenism provides antipredatory benefits through crypsis as well as through masquerade. This is the first time that a species has been demonstrated to have the capacity to benefit both from masquerade and from crypsis, and the first time, polyphenism has been demonstrated to benefit masquerade.

這篇是由J. Skelhorn與G. Ruxton所寫的文章，文中的出發點很有趣，他們利用北美的胡椒蛾(Biston betularia congataria)的幼蟲做為實驗材料，這個幼蟲有兩型，在樺木上就會像樺木的枝條，而在柳樹上就會像柳樹的枝條，而他們認為這樣在不同的枝條呈現不同的型的隱蔽(crypsis)是一種擬態，而這邊擬態的角色則是樹（註）。

這組人馬同樣用小雞進行實驗，實驗分成兩部分，第一個部分測試鳥類是否會把幼蟲當成枝條，也就是「幼蟲像不像枝條」這件事，實驗結果支持這個理論，但即使這樣，他們還是無法下定論，因為鳥類不攻擊幼蟲也有可能是因為幼蟲本身的隱蔽色而不是因為這樣的擬態，因此他們做了第二部分的實驗，測試幼蟲在枝條上的擬態是否成立，捕食者可能藉由這樣的擬態降低發現幼蟲的機率，而幼蟲成功逃避捕食者的攻擊，實驗結果也支持幼蟲在相似的枝條上的擬態效果是較成功的。

在之前有關擬態的實驗中，其實比較少談到關於多態型，或是擬態多個model之類的文章，可能是因為那些典型的例子比較少出現這樣的狀況，然而近幾年研究擬態的相關學者逐漸發現，多態型與擬態多個model可能比想像中的更常見，而且難以解釋這樣的關係，而這關係通常又牽扯完美與不完美擬態（典型例子中大多認為是完美擬態），因此從兩三年前一篇nature的文章談到multiple model之後，逐漸有文章開始討論這方面的議題，而這是第一篇正式測試多態型(polymorphism)與非遺傳多型性(polyphenism)對於擬態效應的文章，雖然不在討論防禦性擬態，但提供了一些多態型與擬態間的證據。

註：有些學者認為這樣的偽裝(camouflage) 是一種自我擬態(automimicry)，而也有看法認為關係必須局限於物種內才能算是自我擬態，如眼斑這樣的例子。本篇所採用的「擬態」的看法似乎是認為這可能算某種擬態，但在定義上這應該算是偽裝，因此本篇可能採用自我擬態的看法，但個人認為這仍有爭議。

Wrote by Chia-Hsuan Wei

原文標題：Evolution of diversity in warning color and mimicry: polymorphism, shifting balance, and speciation

摘要 [原文網址]
Mimicry and warning color are highly paradoxical adaptations. Color patterns in both Müllerian and Batesian mimicry are often determined by relatively few pattern-regulating loci with major effects. Many of these loci are “supergenes,” consisting of multiple, tightly linked epistatic elements. On the one hand, strong purifying selection on these genes must explain accurate resemblance (a reduction of morphological diversity between species), as well as monomorphic color patterns within species. On the other hand, mimicry has diversified at every taxonomic level; warning color has evolved from cryptic patterns, and there are mimetic polymorphisms within species, multiple color patterns in different geographic races of the same species, mimetic differences between sister species, and multiple mimicry rings within local communities. These contrasting patterns can be explained, in part, by the shape of a “number-dependent” selection function first modeled by Fritz Müller in 1879: Purifying selection against any warning-colored morph is very strong when that morph is rare, but becomes weak in a broad basin of intermediate frequencies, allowing opportunities for polymorphisms and genetic drift. This Müllerian explanation, however, makes unstated assumptions about predator learning and forgetting which have recently been challenged. Today's “receiver psychology” models predict that classical Müllerian mimicry could be much rarer than believed previously, and that “quasi-Batesian mimicry,” a new type of mimicry intermediate between Müllerian and Batesian, could be common. However, the new receiver psychology theory is untested, and indeed it seems to us unlikely; alternative assumptions could easily lead to a more traditional Müllerian/Batesian mimicry divide.

這篇文章的出現，綜合了近100年內發展的擬態理論，解釋擬態與警戒色的形成與演化。

擬態群對於演化學家而言，是件令人驚奇，卻又難以解釋的現象，雖然早在十八世紀就已提出這個現象的解釋，但至今仍然無法以一個全面的結果，說明這個現象如何形成，因此在不同領域的生物學家各持己見，因為沒人真的看到這個現象如何形成，有人用數學模型模擬幾萬個世代，推測斑紋的變化；有人從實驗行為學的角度，測試天擇對於擬態性狀的生成；有人從遺傳學的角度，測得擬態物種間的差異與性狀的變化，但無論哪個領域，都無法完美的解釋擬態的產生，當A理論在A領域生成時，在B領域就無法成立。這篇文章就概括所有領域的理論，提出對於現在擬態生物學的看法。

其中比較不一樣的地方在於，作者表達對於quasi-Batesian mimicry與Mullerian mimicry的看法，他們認為如果不在frequency-dependent 的前提下，只藉著receiver psychology是難以解釋擬態物種多態型的形成，也可能會失去擬態的多樣性，也就是不會有像目前看到這麼多樣化的擬態群，而是會趨於簡單化，就算可食度不同，仍然要回到捕食者是否能遇到這個不好吃的東西，仍然回到frequency-dependent的假設下，大大的質疑quasi-Batesian mimicry的可信度。

雖然如此，quasi-Batesian mimicry仍然無法被推翻，因為我們對於捕食者的反應仍了解的太少，有時甚至無法確定何為捕食者，因此理論跟行為學家對於這樣的推論仍然繼續提出證據，爭論依舊沒有結束。

Wrote by Chia-Hsuan Wei