长臂虾总科(甲壳动物亚门:十足目:真虾下目)分子系统学研究
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摘要
长臂虾总科Palaemonoidea Rafinesque,1815隶于甲壳动物亚门Crustacea,十足目Decapoda,真虾下目Caridea,是真虾下目种类最丰富的总科之一。自从长臂虾总科建立以来,有若干个分类方案被先后提出并加以修订。然而,目前关于长臂虾总科的分类系统仍然存在着争议。本研究通过使用1种线粒体基因(16SrDNA)和3种细胞核基因(histone3,18S rDNA和28S rDNA)探索长臂虾总科内部8个科26个之间的系统发育关系。基于联合数据的最大似然法和贝叶斯法分析,结果支持拟贝隐虾科Anchistioididae、膜角虾科Hymenoceridae和咔咔虾科Kakaducarididae的单系性。与之相对,叶颚虾科Gnathophyllidae被认为是一个并系群,而长臂虾科Palaemonidae则显示为一个复系群。特别地,分析所涉及的长臂虾亚科Palaemoninae的种类可以被近似地分为3个大支,并且鳃甲沟很可能作为一个重要的形态学证据,反映了其生存环境从海洋向淡水变迁的过程。此外,长臂虾总科某些科的分类学地位不很明朗,尤其是隐虾亚科Pontoniinae的系统发育关系还未确定,有必要对长臂虾总科及其下分类单元进行修订和重新评估。
     岩虾属Periclimenes Costa,1844是隐虾亚科种数最多的。近来的研究显示岩虾属很可能是一个复系群,并且可以进一步细分。本研究采用3种核蛋白编码基因(enolase,NaK和PEPCK)和1种线粒体基因(16S rDNA)来分析岩虾属及其邻近之间的系统发育关系,包括了岩虾属及其15个邻近,即Ancylomenes,Brucecaris,Crinotonia,Cuapetes,Harpiliopsis,Harpilius,Laomenes,Leptomenaeus,Manipontonia,Palaemonella,Periclimenella,Philarius,Phycomenes,Unguicaris和Vir。基于独立分析和联合数据的分析,结果均支持岩虾属是一个复系群。本研究涉及的岩虾属的种类可以被划分为几个独立的种组,并且共栖岩虾Periclimenes commensalis、短腕岩虾P. brevicarpalis和细指岩虾P. digitalis的分类学地位需要重新考虑,即是否继续留在岩虾属中还是独立于岩虾属之外的。此外,分析显示大多数岩虾属的邻近都是单系群,并且可以近似地分为两大支。但是,某些深层的系统发育关系仍然不十分明确。
     本研究首次利用多种基因序列对长臂虾总科的系统发育关系进行了较为全面和系统的分析,为其系统演化关系的研究和分类系统的进一步划分界定提供了分子生物学证据。此外,本研究还首次采用多种核蛋白编码基因对隐虾亚科多个之间的系统发育关系进行了探索,扩展了利用核蛋白编码基因进行分子系统学研究的类群范围,为传统的形态分类学提供了新的观点和视角。
Palaemonoidea Rafinesque,1815is one of the most speciose superfamilies ofCaridea, Decapoda, Crustacea. Since it was established, several classification schemesof Palaemonoidea have been proposed and modified. However, the currentclassification of Palaemonoidea is still in dispute. In this study, one mitochondrialgene (16S rDNA) and three nuclear genes (histone3,18S rDNA and28S rDNA) wereused to explore the phylogenetic relationships among the subgroups of thesuperfamily Palaemonoidea, including eight families with26affiliated genera. Basedon the combined data with both maximum likelihood and Bayesian inference analyses,the results support the monophyly of Anchistioididae, Hymenoceridae andKakaducarididae. In contrast, Gnathophyllidae is suggested to be paraphyletic andPalaemonidae is shown to be a polyphyletic group. Especially, our analyses revealthat the studied subfamily Palaemoninae species could be approximately divided intothree clades, and the branchiostegal groove is the probable morphological evidence ofthe environmental transition from sea to fresh water. Besides, for some of thePalaemonoidea families, their taxonomic status is obscure. A revision of theclassification of Palaemonoidea and a reevaluation of its constituent taxa appear to benecessary despite the systematics of the subfamily Pontoniinae is still undetermined.
     The genus Periclimenes Costa,1844is the most species-rich genus in thesubfamily Pontoniinae. Recent studies have suggested that it might be a polyphyletictaxon and could be further subdivided. In this study, three protein-coding nucleargenes (enolase, NaK and PEPCK) and one mitochondrial ribosomal gene (16S rDNA)were used to analyse the phylogenetic relationships among the genera of thePericlimenes complex, includes the genus Periclimenes and15related genera, i.e.Ancylomenes, Brucecaris, Crinotonia, Cuapetes, Harpiliopsis, Harpilius, Laomenes,Leptomenaeus, Manipontonia, Palaemonella, Periclimenella, Philarius, Phycomenes,Unguicaris and Vir. Based on both independent and combined data analyses, theresults support that the genus Periclimenes is a polyphyletic group. Furthermore, thestudied Periclimenes species could be divided into several independent groups, andthe taxonomic status of P. commensalis, P. brevicarpalis and P. digitalis may need tobe reconsidered. Besides, the majority of the related genera of Periclimenes aresuggested to be monophyletic. Our analyses also reveal that these generaapproximately form two main clades, despite some deep relationships are still obscure.
     In this study, a variety of gene sequences were used to analyse the systematicstatus of the superfamily Palaemonoidea for the first time, providing molecularevidences for its phylogenetic studies and further classification. Besides, severalnuclear protein-coding genes were utilized to explore the phylogenetic relationshipsamong multiple genera of the subfamily Pontoniinae for the first time, which enrichesthe application of the nuclear protein-coding genes on the molecular phylogeneticstudies and provide new insights into the classification for the traditionalmorphological taxonomy.
引文
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