Phylogenetic classification of the Drosophilidae Rondani (Diptera): the role of morphology in the postgenomic era
Phylogenetic classification of the Drosophilidae Rondani (Diptera): the role of morphology in the postgenomic era
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DOI：
10.1111/j.1365-3113.2012.00665.x
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发表时间：
2013-04-01
影响因子：
4.8
通讯作者：
Yassin, Amir
中科院分区：
文献类型：
--
作者：
Yassin, Amir
现在，分子序列在系统发育推断中压倒了形态学。尽管如此，大多数分子研究都是针对有限数量的类群进行的，因为很少可以从旧博物馆类型或化石中分析 DNA。在过去的 20 年里，超过 150 项分子研究对目前基于形态特征的果蝇科 Rondani 的系统发育分类提出了挑战。大多数研究涉及单个果蝇属，即堕落果蝇属，并且仅包括该科 78 个属中 17 个属中的少数代表性物种。因此，这些分子研究无法提供替代的分类方案。使用来自 1 个帽状体和 4 个 eHydroid 科的外类群对 33 个属的 7 个核基因和 1 个线粒体基因 (8248 bp) 进行了超级矩阵分析。贝叶斯系统发育与之前的分子研究（包括全基因组序列）一致，并将果蝇科分为四个单系进化枝。然后对四个进化枝之间的形态特征（主要是雄性生殖器）进行彻底比较，并鉴定出同源特征状态。然后对这些状态进行了 70 个属的检查，并提出了果蝇科的修订后的系统发育、科群分类。 Cladochaetini Grimaldi 部落的两个 Cladochaeta Coquillett 属和 Diathoneura Duda 属被转移到 EpHydridae 科。果蝇科分为两个亚科：Steganinae Hendel（30 属）和 Drosophilinae Rondani（43 属）。另外两个属，Apacrochaeta Duda 和 Sphyrnoceps de Meijere，是 incertae sedis，而 Palmophila Grimaldi 是 Drosophilasyn.n 的同义词。果蝇亚科又分为两个族：重新进化的Colocasiomyini Okada（九属）和Drosophilini Okada。果蝇属并系未解决，以免影响重要实验室模式物种的二名；然而，其亚属分类根据分子和形态学数据进行了修订。三个亚属，即 Chusqueophila Brncic、Phloridosa Sturtevant 和 Psilodorha Okada，与果蝇亚属 (Drosophila) Fallen syns.n 同义。在果蝇 (Drosophila) 的 45 个种群和 5 个种复合体中，有 22 个类群和 1 个复合体被转移到果蝇 (Siplodora) Patterson & Mainland 亚属，而 6 个类群、2 个种亚群和 3 个复合体被认为是果蝇属内的不确定物种。不同的形态特征在不同的系统发育尺度上提供不同的信号：胸部特征（翅脉和胸骨前形状）区分科；与抓握和勃起相关的特征将亚科与部落区分开来；而阴茎副骺，即辅助插入器官，区分果蝇属和果蝇亚属。该研究表明有必要在分子系统发育框架内分析形态特征，以将分子系统发育转化为分类学综合分类。
Molecular sequences now overwhelm morphology in phylogenetic inference. Nonetheless, most molecular studies are conducted on a limited number of taxa, as DNA rarely can be analysed from old museum types or fossils. During the last 20 years, more than 150 molecular studies have challenged the current phylogenetic classification of the family Drosophilidae Rondani based on morphological characters. Most studies concerned a single genus, Drosophila Fallen, and included only few representative species from 17 out of the 78 genera of the family. Therefore, these molecular studies were unable to provide an alternative classification scheme. A supermatrix analysis of seven nuclear and one mitochondrial genes (8248 bp) for 33 genera was conducted using outgroups from one calyptrate and four ephydroid families. The Bayesian phylogeny was consistent with previous molecular studies including whole genome sequences and divided the Drosophilidae into four monophyletic clades. Morphological characters, mostly male genitalia, then were compared thoroughly between the four clades and homologous character states were identified. These states were then checked for 70 genera and a revised phylogenetic, family-group classification for the Drosophilidae is proposed. Two genera Cladochaeta Coquillett and Diathoneura Duda of the tribe Cladochaetini Grimaldi are transferred to the family Ephydridae. The Drosophilidae is divided into two subfamilies: Steganinae Hendel (30 genera) and Drosophilinae Rondani (43 genera). A further two genera, Apacrochaeta Duda and Sphyrnoceps de Meijere, are incertae sedis, and Palmophila Grimaldi, is synonymized with Drosophilasyn.n. The Drosophilinae is subdivided into two tribes: the re-elevated Colocasiomyini Okada (nine genera) and Drosophilini Okada. The paraphyly of the genus Drosophila was not resolved to avoid affecting the binomina of important laboratory model species; however, its subgeneric classification was revised in light of molecular and morphological data. Three subgenera, namely Chusqueophila Brncic, Phloridosa Sturtevant and Psilodorha Okada, were synonymized with the subgenus Drosophila (Drosophila) Fallen syns.n. Among the 45 species groups and 5 species complexes of Drosophila (Drosophila), 22 groups and 1 complex were transferred to the subgenus Drosophila (Siphlodora) Patterson & Mainland and 6 groups, 2 species subgroups and 3 complexes are considered incertae sedis within the genus Drosophila. Different morphological characters provide different signals at different phylogenetic scales: thoracic characters (wing venation and presternal shape) discriminate families; grasping and erection-related characters discriminate subfamilies to tribes; whereas phallic paraphyses, i.e. auxiliary intromittent organs, discriminate genera and Drosophila subgenera. The study shows the necessity of analysing morphological characters within a molecular phylogenetic framework to translate molecular phylogenies into taxonomically-comprehensive classifications.