林地領域性蝶種以視覺偵測配偶及從事求偶飛行與太陽光斑大小之相關性

photo by Zéza Lemos

Visual mate detection and mate flight pursuit in relation to sunspot size in a woodland territorial butterfly

Animal Behaviour Volume 78, Issue 1, July 2009, Pages 17-23
Martin BergmanCorresponding ^a ,Christer Wiklunda
^a Department of Zoology, Stockholm University, Sweden

據領域具有相當可觀的利益。雄性的斑點木蝶（Pararge aegeria）在開放森林棲地的太陽光斑上飛行，勝利的一方成為光斑的佔有者，而落敗的一方則沒有領地且只能在小的光斑中坐等雌蝶。先前研究發現,雖然雌性並未特別受到領地佔有者或者光斑領域本身的吸引，但是雄性佔有領地者的交配成功率較沒有領地者高。 作者在此測試以下假說：(1)雄性佔有領地者有較高交配成功率的原因在於在大型光斑中的配偶偵測比在小型光斑中更有效率 (2) 只有在某個特定大小以上的光斑能被界定為領域。領域光斑的田野評估顯示，被守禦的光斑大小顯著大於森林底層光斑的平均尺寸。作者以實驗測試雄蝶以視覺偵測一隻模型蝴蝶經過光斑時的能力，結果顯示在光斑中飛行長距離飛行的情形下,雄蝶能較為成功地追求並攔截經過的模型。因此，作者提出光的狀況、視覺配偶偵測與完整的配偶飛行追求可解釋雄性的斑點木蝶（Pararge aegeria）在開放森林棲地守禦大型光斑的理由。

Territory residency is associated with considerable benefits. In the speckled wood butterfly, Pararge aegeria, males fight over ownership of large sunspots in open forest habitats; winners become sunspot residents, and losers become nonterritorial and sit and wait for females in small sunspots. A previous study has shown that residents have higher mating success than nonterritorial males, although females are not more attracted to territorial males or sunspot territories per se. Here we tested the hypotheses (1) that the higher success of resident males is caused by visual mate detection being more efficient in a large than in a small sunspot, and (2) that only sunspots above a certain size are defended as territories. Field assessment of territorial sunspot size showed that defended sunspots were significantly larger than ‘average sunspots’ on the forest floor. Experimental tests of male ability to detect visually a model butterfly passing through a sunspot showed that males were more successful in pursuing and intercepting a passing model when flown a longer distance in the sunspot. Hence, we conclude that light conditions and associated visual mate detection and ability to complete mate flight pursuit can explain why P. aegeria males defend territories in large sunspots in forest habitats.

Keywords: butterfly vision; Lepidoptera; light gap; mate searching; Pararge aegeria; sexual selection; speckled wood butterfly; territoriality

蝴蝶的固定眼紋展示可恫嚇攻擊它的鳥類

Fixed eyespot display in a butterfly thwarts attacking birds

Article References - selected

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Animal Behaviour
Volume 77, Issue 6, June 2009, Pages 1415-1419

Ullasa Kodandaramaiah, Adrian Vallina and Christer Wiklunda

Department of Zoology, Stockholm University, Sweden
Available online 10 April 2009.

眼紋出現在許多鱗翅目昆蟲中，被猜測具有適應上的價值。所提出的假說中其中一個功能是捕食者會被獵物身上大而醒目的眼紋驚嚇，接著避免攻擊這類獵物。近期實驗顯示，眼紋與恫嚇行為的結合可提升存活率。作者測試僅僅存在明顯的眼紋時、是否可以嚇退攻擊的鳥類，意即不斷地展示眼紋而不表現恫嚇行為。作者將孔雀蛺蝶 Junonia almana 的翅膀黏貼在一片卡紙上以模擬成一隻蝴蝶展翅的樣貌，同時將ㄧ隻麵包蟲黏在雙翅中央身體的位置。此實驗中以山雀 Parus major 作為捕食者，在”具有眼紋”與”塗去眼紋”兩種模式獵物中做出選擇。具有眼紋的獵物顯著地比塗去眼紋者較少受到攻擊，首次攻擊與第二次攻擊之間間隔的時間也比被塗去眼紋者更長。以上結果支持天然產生的蝴蝶眼紋可提升存活率，即便是在持續展示且靜止不動的的情形下。

Abstract
Eyespots have evolved in many lepidopteran insects, which suggests their adaptive value. One of their hypothesized functions is that predators are intimidated by prey with large and conspicuous eyespots and hence refrain from attacking them. Recent experiments have shown that a combination of eyespots and intimidating behaviour can increase survival. We tested whether the mere presence of conspicuous eyespots can thwart attacking birds, that is, when the eyespots are displayed constantly, without any intimidating behaviour. We used prey that consisted of wings of the peacock pansy butterfly, Junonia almana, glued onto a piece of cardboard so as to resemble a butterfly with its wings open. A mealworm was placed between the wings in place of the body. Great tits, Parus major, were used as the predator in the study and were offered a choice between two model prey, one with intact eyespots and one without. Prey with eyespots were attacked significantly fewer times than those without. The time between the first and second attack was longer when the prey without eyespots was attacked first. These results support the hypothesis that naturally occuring butterfly eyespots can increase survival even when they are constantly displayed and motionless.

Keywords: eyespot; intimidation hypothesis; Junonia almana; peacock pansy

綜合現生與化石分類群的形態與分子證據推測喙蝶亞科的親緣關係

文獻來源: Kawahara A. 2009. Phylogeny of snout butterflies (Lepidoptera: Nymphalidae: Libytheinae): combining evidence from the morphology of extant, fossil, and recently extinct taxa. Cladistics 25(3): 263-278. [全文下載]

Abstract
Snout butterflies (Nymphalidae: Libytheinae) are morphologically one of the most unusual groups of Lepidoptera. Relationships among libytheines remain uncertain, especially in the placement of the recently extinct Libythea cinyras and two fossils, L. florissanti, and L. vagabunda. The aim of this study is to present the first phylogenetic hypothesis of Libytheinae utilizing all available morphological data from extant and extinct species. Forty-three parsimony-informative characters were coded, and the all-taxa analysis resulted in six most parsimonious trees (length 92 steps, CI = 0.66, RI = 0.82). The subfamily was resolved as monophyletic and was split into Old World and New World clades. Inclusion of extinct species with considerable missing data had little effect on relationships of extant taxa, although Bremer support values and jackknife frequencies generally decreased if extinct species were included. In order to preserve the monophyly of extant genera, two fossils are assigned to Libytheana for the first time (L. florissanti comb. n. and L. vagabunda comb. n.). This study demonstrates the value of morphological data in phylogenetic analysis, and highlights the contribution that can be made by scoring extinct taxa and including them directly into the analysis.

帶蛾科之學名的正確作者與歷史上曾出現同物異名的有效性問題

文獻來源: Nässig WA, Oberprieler RG. 2007. The nomenclature of the family-group names of Eupterotidae (Bombycoidea). Nota lepidopterologica 30(2): 315—327. [全文下載]

Abstract
The origin and historical use of all family-group names pertaining to the bombycoid family Eupterotidae are determined. The family-group name Eupterotidae takes Swinhoe, 1892 as authorship, although the name was evidently originally coined by Hampson, 1893. However, three older family-group names are available for the taxon: Striphnopterygidae Wallengren, 1858, Phialidae Wallengren, 1865 and Janidae Aurivillius, 1892. Striphnopterygidae has been used as the valid name for the family by at least ten authors until at least 1965 and again in 2006, and as a valid subfamily name throughout, but the younger Eupterotidae gained almost universal acceptance as the name of the family in the literature since about 1928. Application of Article 35.5, introduced in the current, 4th edition of the Code (ICZN 1999), allows the retention of the younger name Eupterotidae as the valid name of the family. The authorship of another bombycoid family name, Lemoniidae, is corrected from Hampson, 1918 to Neumoegen & Dyar, 1894.

圖片連結: Bettaman之flickr相簿

由雄蟲合生毛刷特徵進行斑螟類親緣關係與比較形態學研究

Phylogenetic utility and comparative morphology of the composite scale brushes in male phycitine moths (Lepidoptera, Pyralidae)

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Thomas J. Simonsen and Amanda D. Roe

Zoologischer Anzeiger 248 (2009): 119–136

鱗翅目形態中一般所稱的毛刷 (hair brushes)、毛筆器 (hair pencil)與發香鱗(androconial scales)等特徵，在不同總科類群中多次的出現，其類型與繁複程度、分佈部位皆有所不同，也因而造成此類特徵在命名上的混淆與爭議。

雖然過去不少親緣關係的研究經常選用此類特徵進行形態親緣重建的特徵，然而相較其應用於系統發生學，此類特徵的同源性、組織學與功能形態學的討論顯得少且不完整。

作者Simonsen (註一) 與 Roe是兩位加拿大亞伯達省大學生物科學系的研究員，研究方向著重於鱗翅目螟蛾科類群之系統發生。在此篇文章選用了佔螟蛾科2/3物種數的斑螟亞科(Phycitinae)中的Dioryctria屬(台灣共3種)，來討論高多樣性的第八腹節合生毛刷(composite hair brushes)在超微結構(ultrastructural)、總體形態(gross morphology)與相關且複雜的鄰近腺體組織的同源性，並映像(mapping)此類特徵至Dioryctria屬分子親緣關係樹後，所得到結果的演化意義。

研究方法中，作者使用掃描式電子顯微鏡(SEM)作特徵的觀察與呈現，而非傳統形態所利用的光學或解剖顯微鏡；此外，作者選用了粒線體COI與COII序列，並使用最簡約法(maximum parsimony)進行親緣關係重建，而非一般分子親緣樹重建上所使用的最大概似法(maximum likelihood)或貝葉氏機率(Baysian probability)。

結果方面，作者將此類合生毛叢器特徵再映像至分子樹上，可支持Dioryctria屬的單系性，然而在特徵狀態上具有多次的轉換，也就是屬內特徵狀態具有趨同演化的結果，但當中的樹型分佈卻可支持屬內某些種群(species groups)的單系性。而作者也利用所獲得的結果來闡述結合各種層級特徵的評估可對斑螟亞科的演化關係在未來提供豐富的資訊；最後作者也提到使用掃描式電子顯微鏡而非光學或解剖顯微鏡來分析超微結構，可獲得更多且為3D的立體結構特徵。

註一 Thomas Simonsen的個人介紹

Abstract

Male pyralid moths in the subfamily of Phycitinae are known to possess composite scale brush structures associated with the 8th abdominal sternite, but the histology and the structural morphology of these organs have not been adequately explored. As such, the phylogenetic utility of these structures is unknown. We examine the pre-genitalic abdominal histology of male Dioryctria reniculelloides (Pyralidae: Phycitinae) associated with the composite scale brushes, as well as structural morphology within the genus Dioryctria and two closely related genera. The composite scale brushes are composed of fused scales. The musculature associated with the base of sternum 8 shows considerable modification compared to previously described Lepidoptera. Complex glandular tissue was also found associated with the scale brush structures, suggesting secretory function. Phylogenetic utility of ultrastructure and gross morphology was examined for major Dioryctria species groups. Many characters were homoplasious, but several supported the monophyly of the genus, as well as some internal relationships. In conclusion, the combination of ultrastructural, gross morphological and histological characters can be a rich source of information for elucidating a range of evolutionary relationships within the subfamily.

Keywords: Dioryctria; Phylogeny; Ultrastructure; Histology; Homology

[TaiBNET名錄修訂]Argyarctia拼法訂正

TaiBNET鱗翅目燈蛾科名錄應進行以下的修訂: 資料庫上的拼法Argyractia乃根據Lepidoptera of Taiwan而來, 但此拼法事實上是錯誤的, 應根據江田信豐(KODA Nobutoyo)的原始發表訂正為Argyarctia.

圖片連結: gaga昆蟲網

舊北區捲蛾誌第一卷 - 總論與捲蛾族(Tortricini)

Razowski J. 2008. Tortricidae (Lepidoptera) of the Palaearctic Region Volume 1 General Part and Tortricini. Frantisek Slamka, Bratislaza (Slovakia). [網址]

波蘭籍的捲蛾學者Josef Razowki可能是當今世界上在捲蛾的種級分類工作上最為活躍的學者, 自2002年以來他已經連續發表了四本以上的歐洲產捲蛾科專冊, 而2008年所發行的這本書則是他首次將他過去研究歐洲以外地區的捲蛾成果集結整理, 其中也包含了少數分布於台灣的種類. 此外台灣產捲蛾也曾經被Razowski於2002年簡短地回顧, 從該篇論文以及中國動物誌的捲蛾科問世後, 台灣的捲蛾科基礎分類研究工作就沒有更新的進展了.

Pyraloidea Planet 3已刊出

請由此下載

歐亞大陸與北非夜蛾誌冬夜蛾亞科(I)已出版

相關資訊與訂購方式請見其官方網頁

天擇與性擇在蝴蝶翅膀斑紋演化上的調和

photo by kibuyu

Accommodating natural and sexual selection in butterfly wing pattern evolution

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Jeffrey C. Oliver1,Kendra A. Robertson2 and Antónia Monteiro1,2

Proc. R. Soc. B 7 July 2009 vol. 276 no. 1666 2369-2375

動物的視覺樣式可能具有不同的功能，例如吸引配偶與欺騙天敵。假設一個訊號具有多重的功能，不同功能間便有可能發生衝突，而妨礙任何視覺訊號的最佳化。作者對“Bicyclus屬蝴蝶具有不同視覺訊號功能的空間性區隔“之假說進行驗證。運用特徵演化的親緣關係重建與演化速率之比較，作者發現背面的特徵眼化速率比腹面快速。因此，背面的特徵與作者對交配訊號的預測有關、而腹面特徵則在避免天敵的功能上扮演重要的角色。前翅的特徵亦符合交配訊號的模型、且表現出比後翅特徵更高的演化速率。作者的結果，與前人在行為與發育上的研究資料吻合，均支持“Bicyclus屬蝴蝶具有不同視覺訊號功能的空間性區隔“之假說。本研究是首先以親緣關係架構證明：使用在交配訊息上與用來避免捕食者的訊號空間性區隔是可行的策略，可調和多重的訊號功能。這種訊號策略對於蝴蝶翅膀斑紋元件之發育與分化的演化具有重要的衍生脈絡。

Abstract

Visual patterns in animals may serve different functions, such as attracting mates and deceiving predators. If a signal is used for multiple functions, the opportunity arises for conflict among the different functions, preventing optimization for any one visual signal. Here we investigate the hypothesis that spatial separation of different visual signal functions has occurred in Bicyclus butterflies. Using phylogenetic reconstructions of character evolution and comparisons of evolutionary rates, we found dorsal surface characters to evolve at higher rates than ventral characters. Dorsal characters also displayed sex-based differences in evolutionary rates more often than did ventral characters. Thus, dorsal characters corresponded to our predictions of mate signalling while ventral characters appear to play an important role in predator avoidance. Forewing characters also fit a model of mate signalling, and displayed higher rates of evolution than hindwing characters. Our results, as well as the behavioural and developmental data from previous studies of Bicyclus species, support the hypothesis that spatial separation of visual signal functions has occurred in Bicyclus butterflies. This study is the first to demonstrate, in a phylogenetic framework, that spatial separation of signals used for mate signalling and those used for predator avoidance is a viable strategy to accommodate multiple signal functions. This signalling strategy has important ramifications on the developmental evolution of wing pattern elements and diversification of butterfly species.

以五個核基因重建天蛾科的親緣關係與生物地理

文獻來源: Kawahara AY, Mignault AA, Regier JC, Kitching IJ, Mitter C (2009) Phylogeny and Biogeography of Hawkmoths (Lepidoptera: Sphingidae): Evidence from Five Nuclear Genes. PLoS ONE 4(5): e5719. doi:10.1371/journal.pone.0005719

Abstract
Background The 1400 species of hawkmoths (Lepidoptera: Sphingidae) comprise one of most conspicuous and well-studied groups of insects, and provide model systems for diverse biological disciplines. However, a robust phylogenetic framework for the family is currently lacking. Morphology is unable to confidently determine relationships among most groups. As a major step toward understanding relationships of this model group, we have undertaken the first large-scale molecular phylogenetic analysis of hawkmoths representing all subfamilies, tribes and subtribes. Methodology/Principal Findings The data set consisted of 131 sphingid species and 6793 bp of sequence from five protein-coding nuclear genes. Maximum likelihood and parsimony analyses provided strong support for more than two-thirds of all nodes, including strong signal for or against nearly all of the fifteen current subfamily, tribal and sub-tribal groupings. Monophyly was strongly supported for some of these, including Macroglossinae, Sphinginae, Acherontiini, Ambulycini, Philampelini, Choerocampina, and Hemarina. Other groupings proved para- or polyphyletic, and will need significant redefinition; these include Smerinthinae, Smerinthini, Sphingini, Sphingulini, Dilophonotini, Dilophonotina, Macroglossini, and Macroglossina. The basal divergence, strongly supported, is between Macroglossinae and Smerinthinae+Sphinginae. All genes contribute significantly to the signal from the combined data set, and there is little conflict between genes. Ancestral state reconstruction reveals multiple separate origins of New World and Old World radiations. Conclusions/Significance Our study provides the first comprehensive phylogeny of one of the most conspicuous and well-studied insects. The molecular phylogeny challenges current concepts of Sphingidae based on morphology, and provides a foundation for a new classification. While there are multiple independent origins of New World and Old World radiations, we conclude that broad-scale geographic distribution in hawkmoths is more phylogenetically conserved than previously postulated.

蝴蝶翅膀特徵的環境壓力與數量遺傳變異

Environmental stress and quantitative genetic variation in butterfly wing characteristics

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Evol Ecol (2009) 23:473–485 DOI 10.1007/s10682-008-9246-4

W. Talloen AE S. Van Dongen AE H. Van Dyck AE L. Lens

蝴蝶翅膀特徵廣泛地被用作模式系統使用在發育、數量遺傳與表型可塑性的研究上。然而，雖然在演化上有關聯，選汰壓力對遺傳變異本身表現的影響確很少被研究。在這篇文章中，作者探討斑點木蝶（Pararge aegeria）翅紋特徵的遺傳變異如何隨著寄主植物的乾燥程度梯度而變化。測量部位包括前翅區域、基部與末端的黑化程度以及五個黃色斑紋的區域。作者發現表現在黑化的程度的遺傳變異隨著乾燥壓力的提升而增加，且類似地(雖不顯著)，與前翅尺寸相關。結果顯示，狹義“遺傳力(heritability)“的上限、與翅的大小和黑化程度的加成性遺傳變異，都隨著乾燥壓力而上升。不同黃色斑點的樣式則關聯較低，推測存有遺傳變異與環境壓力之間的特徵專一性。

Abstract
Butterfly wing characteristics are extensively used as model system in studies of development, quantitative genetics and phenotypic plasticity. In spite of its evolutionary relevance, however, the effect of stress on the expression of genetic variation itself has only rarely been studied. In this paper, we explore genetic variation of wing characteristics of the Speckled wood Pararge aegeria along a host plant drought stress gradient. Forewing area, basal and distal degree of melanization and the area of five yellow wing spots were measured. We found an increase in (additive) genetic variation in degree of melanization at
higher drought stress, and a similar, yet non-significant, relationship for forewing size. As a result, both the upper limits of the narrow-sense heritability and the coefficient of additive genetic variation of wing size and melanization increased with drought stress. Patterns for the different yellow wing spots were less consistent, suggesting trait-specificity in the relationships between genetic variation and environmental stress.

蝴蝶吸了水菊反應變慢?

這個報導的問題在於什麼呢? 有一點大驚小怪. 首先, 這個植物被誤稱為光葉水菊已經很久, 可是這玩意兒應該稱為光冠水菊才是 ("Gymnocoronis"指的是花冠缺乏絨毛). 其次是, 菊科植物因為可以提供大量的PA植物鹼因此很自然地會成為雄性蝶類(尤其是斑蝶或其它需要PA合成性費洛蒙或成為化學防禦物質的物種)駐留的物種. 所以蝶類吸食這樣的蜜源有什麼問題嗎? 之前有一個說法, 認為外來植物與本土生物沒有交互關係所以不具生態功能, 所以植栽應選擇本土植物以提功人工植栽的生態功能, 可是現在這種水生植物提供這麼棒的蜜源為什麼又是問題了? 曾經成為保育人士或媒體囑目的外來植物在過去難道沒有菊科植物嗎? 例如早期的兩種霍香薊, 近期的大花咸豐草都是菊科植物, 又為什麼沒有引發什麼"蝴蝶變遲鈍", "干擾其它植物授粉"的疑慮呢? 馬鞭草科植物也是校園植栽植物與外來植物的大宗, 例如馬櫻丹, 長穗木, 金露花也都含有高量的PA, 也都是在野外或人工環境下重要的蜜源, 那麼為什麼又沒有人憂慮馬鞭草科的外來植物會干擾其它植物的授粉呢?

我認為提出這樣的觀點是需要科學佐證的, 否則隨便講講就會變成"所謂的保育人士"和"解說教育界"口耳相傳的偽科學. 為什麼這樣說呢? 如果斑蝶, 日行性燈蛾等昆蟲需要PA, 他們在台灣的環境中所能夠獲得PA的主要蜜源就是馬鞭草科與菊科植物 (紫草科植物在台灣不是小草本就是只在局部出現的開花期很短的木本, 所以不會成為主要的PA來源). 所以如果水菊植物真的那麼吸引斑蝶, 那麼這種現象倒底會危害什麼植物的授粉呢? 在全球有關外來物種引入造成原生植物授粉作用被干擾的案例研究中, 所有被干擾授粉的植物與原本的授粉動物幾乎都有專一性的授粉關係, 而其花朵與花序的構造都不是開放性的, 也就是說, 對於可進行授粉作用的動物來說是有限制性的. 然而菊科植物, 尤其是在台灣平地分布的大多數菊科植物都不可能具有封閉性的花序與花朵構造. 這也就是說, 它們的授粉動物本來就沒有忠誠度, 那麼就算水菊出現了, 牠們應該還是可以得到其它非蝶類的授粉者才是. 所以水菊的出現真的就會干擾本來就不具專一性授粉媒的授粉關係嗎? 這是一個很值得以科學方法驗證的議題.

另外, 所有嗜食PA的鱗翅目昆蟲對於這些物質本來就會"成癮", 這在許多化學生態研究中都早就被證實, 所以蝴蝶變得比較遲鈍其實只是人類主觀的價值判斷, 這些物質對於蝶類的化學防禦與費洛蒙合成本來就很重要, 所以就算飛起來遲鈍又如何呢?

當然, 水菊是一個外來物種, 不應該被鼓勵放流於台灣的自然環境中, 但是如果水菊被種植在人工植栽環境中, 應該還是有其教育功能, 否則所有的人工植栽中就應該把繁星花, 金露花等外來蜜源植物一併鏟除, 而不要以"本土化"的大帽子危害這項議題的科學論證.

蝴蝶搖頭丸 光葉水菊危害很大 自由 更新日期:"2009/06/01 06:59" 〔記者黃淑莉／斗六報導〕

被視為蝴蝶的搖頭丸的植物─光葉水菊，近幾年普遍成為校園吸引蝴蝶的蜜源植物，由於光葉水菊為外來植物，繁殖快速，生態專家憂心破壞生態。 蝴蝶接觸上癮 反應變慢 光葉水菊原產於美洲，早期引進是為水族箱造景的水生植物，適應力強、生長快速，目前國內許多平地、山區的陸地及河川都可見其蹤跡，尤其光葉水菊的花朵是極佳的蜜源植物，常被學校、生態農場種植以引來蝴蝶。 台灣生態學會雲林工作站主任陳清圳指出，光葉水菊開小白花可能具特殊氣味，特別吸引斑蝶喜好，連許多著名觀光景點，都有栽種光葉水菊招蜂引蝶，不過，蝴蝶接觸過光葉水菊的花朵後，對其他植物就興趣缺缺，甚至反應動作也變慢，因此有人稱光葉水菊是蝴蝶的搖頭丸。 適應強長得快 也能水生 陳清圳表示，光葉水菊適應力極強、生長又快速，日前有人拿幾株到學校生態池種植，沒幾天就長得很茂密，隨手丟棄在一旁空地的根莖也長成一大片，蝴蝶接觸後不願離開，自己曾在國內著名景點看到一小片光葉水菊的上空有好幾百隻蝴蝶盤旋，由於蝴蝶還兼負為許多植物傳播花粉職務，其他植物等不到蝴蝶，久了可能面臨滅種危機。 陳清圳說，光葉水菊不僅在陸地長得很好，它也是水生植物優勢者，排擠到本土水生植物，自然界生態都有其生存食物鏈模式，不免令人憂心光葉水菊對台灣生態的破壞。