岩虾属及其邻近属（甲壳动物亚门：隐虾亚科）分子系统发育研究
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摘要
岩虾属(Periclimenes Costa,1844)隶属于甲壳动物亚门，十足目，长臂虾科，隐虾亚科(Pontoniinae)，是隐虾亚科中最大的属。尽管已经有很多物种被从该属移出建立新的属，但是该属迄今仍包括155个种。岩虾属形态多样，与其它海洋生物的共栖关系复杂，其起源、亚群间的系统学关系不清晰，是一个明显的多系群而非自然的单系类群。本研究通过筛选，选取了线粒体基因组中的16S rDNA作为遗传标记，利用长臂虾亚科的沼虾属(Macrobrachium Bate,1868)的物种作为外群，对世界分布的岩虾属及其邻近属的系统发育关系进行探讨。综合最大简约法，最大似然法以及贝叶斯法构建的系统发育树的结果显示：
本研究表明，隐虾亚科的岩虾属及其邻近属构成一个单系群；Crinotonia属最先从岩虾属及其邻近属的物种中独立出来，是该单系群中较原始的类群；其后，岩虾属的邻近属明显分为两大谱系，即“Philarius+Cuapetes+Palaemonella+Vir+Exoclimenella+Sandimenes”和“Harpiliopsis+Brucecaris+Ancylomenes+Unguicaris+Laomenes+Leptomenaeus+Manipontonia+Blepharocaris+Patonia+Phycomenes”；而岩虾属本身的种不规则地分布于这两个谱系中，从分子系统学上证实了该属是一个多系群；拟长臂虾属Palaemonella Dana，1852和曼隐虾属Vir Holthuis,1952聚在一起，并具有强的自展值和贝叶斯后验概率值的支持，但是不支持拟长臂虾属Palaemonella Dana，1852和曼隐虾属Vir Holthuis,1952是隐虾亚科中最原始的类群；岩虾属的Periclimenes foresti种组，Periclimenesalcocki种组及Periclimenes hertwigi种组的物种分别聚在一起，但是不支持从形态学上对Periclimenes brevicarpalis种组的划分，该种组的有效性还需要进一步的研究；Cuapetes属并不是一个单系群，Cuapetes anacanthus在Cuapetes属中的系统发育地位有待进一步研究；支持基于形态学特征将Periclimenes aesopius从广义的岩虾属中独立出来，建立的新属Ancylomenes Okuno&Bruce,2010，并且该属是一个单系群；对与棘皮动物海百合共生的岩虾属及其邻近属的种的系统发育学研究表明，它们与海百合形成的共生关系及形态特征上的适应是由于趋同进化和平行进化的结果。
本论文丰富了利用线粒体基因进行分子系统学研究的内容，揭示了岩虾属及其邻近属的系统发育关系，并对岩虾属进一步的形态分类学研究提供了初步的分子系统学资料。
The genus Periclimenes Costa,1844has become the most speciose genus of thePontoniinae, Palaemonidae, Decapoda, Crustacea. Up to date, there are still155species included in this large genus despite many species having been separated fromit and given generic status over recent years. It has been suggested that Periclimenesis a polyphyletic taxon, however, the morphological study couldn’t resolve well theorigin and phylogenetic relationships of the Periclimenes and its related genera as thegenus Periclimenes owns a great diversity of morphological characters and themajority of the species are considered as the associates of an variety of marineinvertebrates.
Nucleotide sequences of the mitochondrial16S rRNA gene from extant genera ofthe Pontoniinae were used to reconstruct the phylogeny of the Periclimenes and itsrelated genera, and Macrobrachium japonicum was selected as the outgoup.According to the Phylogenetic trees constructed based on the three methods,parsimony (MP), maximum likelihood (ML) and Bayesian inference (BI), the mainconclusions of the present study were shown as follows:
According to this study, the genus Periclimenes and its related genera clustertogether and form a monophyletic group within the Pontoniinae; Crinotonia Marin,2006, the first genus separated from the truck of the phylogenetic trees ofPericlimenes and its related genera of the Pontoniinae, should be the most primitivegenus; the remaining genera excluding Periclimenes are divided into two distinctclades, with the first clade consisting of Philarius, Cuapetes, Palaemonella, and Vir,and the second clade including Harpiliopsis, Brucecaris, Ancylomenes, Unguicarisand Laomenes; the species of Periclimenes Costa,1844are scattered in the two clades,implying this genus Periclimenes is a polyphyletic group; Palaemonella Dana，1852and Vir Holthuis,1952are clustered together in the first clade supported strongly bythe bootstrap (BP) and posterior probabilities (PP), but the analysis result does notsupport that they are the most primitive group in the subfamily Pontoniinae; species of Periclimenes foresti species group， Periclimenes alcocki species complex andPericlimenes hertwigi species group are clustered together separately, but whetherPericlimenes brevicarpalis species group is a valid species group still needs furtherstudy; Ancylomenes Okuno&Bruce,2010, which was established to cover thePericlimenes aesopius species group of the genus Periclimenes Costa, is a valid genus,and the monophyly of this genus was supported by the molecular analysis; thepolyphyly of the crinoid-associated fauna of the Periclimenes and its related generademonstrates that the very diverse morphology are the consequence of the convergentand parallel evolutions.
The phylogenetic analysis of Periclimenes and its related genera of thePontoniinae reveals the phylogenetic relationships among these genera, enriches theapplication of the mitochondrial16S rRNA gene sequences on the molecularphylogenetic studies and provides molecular information for the study on thephylogenetic study of the decapod Crustacea.
本研究表明，隐虾亚科的岩虾属及其邻近属构成一个单系群；Crinotonia属最先从岩虾属及其邻近属的物种中独立出来，是该单系群中较原始的类群；其后，岩虾属的邻近属明显分为两大谱系，即“Philarius+Cuapetes+Palaemonella+Vir+Exoclimenella+Sandimenes”和“Harpiliopsis+Brucecaris+Ancylomenes+Unguicaris+Laomenes+Leptomenaeus+Manipontonia+Blepharocaris+Patonia+Phycomenes”；而岩虾属本身的种不规则地分布于这两个谱系中，从分子系统学上证实了该属是一个多系群；拟长臂虾属Palaemonella Dana，1852和曼隐虾属Vir Holthuis,1952聚在一起，并具有强的自展值和贝叶斯后验概率值的支持，但是不支持拟长臂虾属Palaemonella Dana，1852和曼隐虾属Vir Holthuis,1952是隐虾亚科中最原始的类群；岩虾属的Periclimenes foresti种组，Periclimenesalcocki种组及Periclimenes hertwigi种组的物种分别聚在一起，但是不支持从形态学上对Periclimenes brevicarpalis种组的划分，该种组的有效性还需要进一步的研究；Cuapetes属并不是一个单系群，Cuapetes anacanthus在Cuapetes属中的系统发育地位有待进一步研究；支持基于形态学特征将Periclimenes aesopius从广义的岩虾属中独立出来，建立的新属Ancylomenes Okuno&Bruce,2010，并且该属是一个单系群；对与棘皮动物海百合共生的岩虾属及其邻近属的种的系统发育学研究表明，它们与海百合形成的共生关系及形态特征上的适应是由于趋同进化和平行进化的结果。
本论文丰富了利用线粒体基因进行分子系统学研究的内容，揭示了岩虾属及其邻近属的系统发育关系，并对岩虾属进一步的形态分类学研究提供了初步的分子系统学资料。
The genus Periclimenes Costa,1844has become the most speciose genus of thePontoniinae, Palaemonidae, Decapoda, Crustacea. Up to date, there are still155species included in this large genus despite many species having been separated fromit and given generic status over recent years. It has been suggested that Periclimenesis a polyphyletic taxon, however, the morphological study couldn’t resolve well theorigin and phylogenetic relationships of the Periclimenes and its related genera as thegenus Periclimenes owns a great diversity of morphological characters and themajority of the species are considered as the associates of an variety of marineinvertebrates.
Nucleotide sequences of the mitochondrial16S rRNA gene from extant genera ofthe Pontoniinae were used to reconstruct the phylogeny of the Periclimenes and itsrelated genera, and Macrobrachium japonicum was selected as the outgoup.According to the Phylogenetic trees constructed based on the three methods,parsimony (MP), maximum likelihood (ML) and Bayesian inference (BI), the mainconclusions of the present study were shown as follows:
According to this study, the genus Periclimenes and its related genera clustertogether and form a monophyletic group within the Pontoniinae; Crinotonia Marin,2006, the first genus separated from the truck of the phylogenetic trees ofPericlimenes and its related genera of the Pontoniinae, should be the most primitivegenus; the remaining genera excluding Periclimenes are divided into two distinctclades, with the first clade consisting of Philarius, Cuapetes, Palaemonella, and Vir,and the second clade including Harpiliopsis, Brucecaris, Ancylomenes, Unguicarisand Laomenes; the species of Periclimenes Costa,1844are scattered in the two clades,implying this genus Periclimenes is a polyphyletic group; Palaemonella Dana，1852and Vir Holthuis,1952are clustered together in the first clade supported strongly bythe bootstrap (BP) and posterior probabilities (PP), but the analysis result does notsupport that they are the most primitive group in the subfamily Pontoniinae; species of Periclimenes foresti species group， Periclimenes alcocki species complex andPericlimenes hertwigi species group are clustered together separately, but whetherPericlimenes brevicarpalis species group is a valid species group still needs furtherstudy; Ancylomenes Okuno&Bruce,2010, which was established to cover thePericlimenes aesopius species group of the genus Periclimenes Costa, is a valid genus,and the monophyly of this genus was supported by the molecular analysis; thepolyphyly of the crinoid-associated fauna of the Periclimenes and its related generademonstrates that the very diverse morphology are the consequence of the convergentand parallel evolutions.
The phylogenetic analysis of Periclimenes and its related genera of thePontoniinae reveals the phylogenetic relationships among these genera, enriches theapplication of the mitochondrial16S rRNA gene sequences on the molecularphylogenetic studies and provides molecular information for the study on thephylogenetic study of the decapod Crustacea.
引文
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