2009年2月18日 星期三

Lepidoptera (3) 由分子證據重建尺蛾科高階親緣關係並討論冬尺蛾類群的演化

Phylogeny of the Geometridae


同文載自
台灣鱗翅學簡訊 (Newsletter of Taiwan Lepidopterist)

文獻來源: Yamamoto S, Sota T. 2007. Phylogeny of the Geometridae and the evolution of winter moths inferred from a simultaneous analysis of mitochondrial and nuclear genes. Molecular Phylogenetics and Evolution 44(2): 711-723. [全文連結]

Abstract
Geometridae is one of the most diverse families within the Lepidoptera, comprising nine subfamilies. Winter moths, which have a unique life history, are found in three subfamilies. To examine the phylogeny of the Geometridae at the subfamily level and determine the evolutionary history of winter moths, we constructed phylogenetic trees for all nine geometrid subfamilies using two mitochondrial and two nuclear gene sequences. Specimens of all subfamilies were sampled from Japan. Simultaneous analyses of the combined data from all genes revealed that the Geometridae comprised two major clades: one with subfamilies Larentiinae and Sterrhinae, and the other with the remaining seven subfamilies. The second clade included the largest subfamily, Ennominae, and the subfamily Archiearinae, which is traditionally considered to be an ancestral lineage of the Geometridae. The Larentiinae + Sterrhinae clade contained one winter moth lineage, and the second major clade consisted of three winter moth lineages, including Alsophilinae, which contains winter moths exclusively. Using a Bayesian inference of divergence times, we estimated that geometrids began to diverge 54 Mya (62–48 Mya), whereas winter moth lineages differentiated from non-winter moth lineages 34–12 Mya, during the global cooling events in the Oligocene and the early Miocene. The adaptation to cool climates may have been a preadaptation that facilitated the winter moth life cycle.

簡介
尺 蛾科為鱗翅目具有高多樣性的類群,全世界物種數約為兩萬一千種,除南北兩極以外皆有分佈。據過去十多年內以形態 (Holloway, 1996, 1997)和分子證據(Abraham et al., 2001)進行尺蛾科高階親緣關係的研究,將其內主要劃分為九個亞科 (如下,台灣產其中七亞科,以括號表示台灣所含物種數):
Archiearinae
Alsophilinae
Sterrhinae 姬尺蛾亞科 (大約85種)
Larentinae 波尺蛾亞科 (大約235種)
Geometrinae 青尺蛾亞科 (大約105種)
Ennominae 枝尺蛾亞科 (超過370種)
Oenochrominae 星尺蛾亞科 (4種)
Orthostixinae (1種)
Desmobathrinae (4種)

此文章於2006年投稿並被期刊接受,而此研究為了增加分類群取樣的數量以及其發生期的詳細紀錄(主要用以討論冬尺蛾*的演化),從2004年即開始於日本進行採集計畫,而方法上以較多的分子序列(粒線體DNA(COI-II, 16S)與核DNA(EF-1
α , 28S))與更多的分類群取樣(含括九個尺蛾亞科)來重建尺蛾科高階親緣關係

以下簡述文章結論與重要討論:
  1. 有四組序列合併所得的樹形被分為兩個主要的支系,一是波尺蛾亞科加上姬尺蛾亞科,另一是其餘的七個亞科,當中包含了佔有所有尺蛾種數35%的枝尺蛾亞科,以及過去被認為是尺蛾類基群的Archiearinae亞科,而僅有冬尺蛾類群的Alsophilinae則埋入枝尺蛾亞科支系,並且未與枝尺蛾亞科內之冬尺蛾類群形成單系群;
  2. 過去冬尺蛾類群據形態分類學,被處理於3亞科12屬中,並認為冬尺蛾的出現是為了躲避天敵的捕食;此篇文章取樣了3亞科5屬7種,並與Abraham et al. (2001)皆證實冬尺蛾非單系群,而是獨立演化了至少四次;另此篇文章由分子鐘定年並配合地質年表與事件,推論冬尺蛾的演化源自晚秋自初春之間活動的尺蛾類群,而冬尺蛾的出現可能是由於地質史上的漸新世至中新世早期間,氣候轉趨寒冷的結果,然促進與維持冬尺蛾在僅在冬季活動的選汰作用尚不明。
  3. 雖然Alsophilinae亞科支系在親緣關係樹中被埋入枝尺蛾亞科內,此文章並未對其做任何分類上的處理。
台灣的冬尺蛾類群紀錄共有兩筆,一比是波尺蛾亞科的Operophtera brumata (Linnaeus, 1758)(國外影像連結),一筆是枝尺蛾亞科的Phigalia owadai Nakajima, 1994 (模式標本影像生態影像連結),兩者目前尚未有無翅雌蟲的紀錄。

*冬尺蛾(winter geometrid moths)是一群冬季活動的尺蛾類群通稱,這類尺蛾在親緣上並非單系群,而分類上被歸屬於3亞科(波尺蛾亞科、枝尺蛾亞科與
Alsophilinae亞科)12 屬中,其中Alsophilinae完全由冬尺蛾所組成。冬尺蛾的雌蟲具有無翅(wingless)或短翅(brachypterous)與退化的口器 (brachystomatous)等特殊的形態,由於特殊的型態以及一些種類被認為是害蟲而被加以研究;Niitsu(2001)藉由觀察枝尺蛾亞科的 冬尺蛾屬Nyssiodes lefuarius之蛹期發育,證實此種雌蟲的缺翅形態是由於蛹期的細胞凋零所致。


近期尺蛾與冬尺蛾相關文獻參考:
  • Abraham, et al. (2001). Molecular phylogeny of the subfamilies in geometridae (Geometroidea : Lepidoptera). Molecular Phylogenetics and Evolution 20(1): 65-77.
  • Beljaev, E. A. (2006). A morphological approach to the Ennominae phylogeny (Lepidoptera, Geometridae). Spixiana 29(3): 215-216.
  • Holloway, J. D. (1996). The Moths of Borneo: Part 9; Family Geometridae, subfamily Oenochrominae, Desmobathrinae and Geometrinae. Malayan Nature Journal 49(3-4): 147-326.
  • Holloway, J. D. (1997). The Moths of Borneo: Part 12; Family Geometridae, subfamily Sterrhinae and Larentiinae. Malayan Nature Journal 51(1-4): 1-242.
  • Niitsu, S. (2001). Wing degeneration due to apoptosis in the female of the winter moth Nyssiodes lefuarius (Lepidoptera, Geometridae). Entomological Science 4(1): 1-7.
  • Sihvonen, P. (2005). Phylogeny and classification of the Scopulini moths (Lepidoptera : Geometridae, Sterrhinae). Zoological Journal of the Linnean Society 143(4): 473-530.
  • Sihvonen, P. and L. Kaila (2004). Phylogeny and tribal classification of Sterrhinae with emphasis on delimiting Scopulini (Lepidoptera : Geometridae). Systematic Entomology 29(3): 324-358.
  • Sihvonen, P. and M. Siljander (2005). Species diversity and geographical distribution of Scopulini moths (Lepidoptera : Geometridae, Sterrhinae) on a world-wide scale. Biodiversity and Conservation 14(3): 703-721.
台灣尺蛾科生態影像連結:
台灣的尺蛾/ Geometrid Moths of Taiwan (Geometridae)