西北太平洋两种鲐属鱼类的分子系统地理学研究
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摘要
鲐属（Scomber）为暖水性中上层鱼类，在海洋捕捞业中占有重要的地位，但其系统发育关系与分类、物种划分尚存争议。本研究基于形态学、线粒体DNA和核基因分析了鲐属鱼类的分类与系统发育关系，并探讨了鲐属鱼类的起源进化历史。同时我们以在西北太平洋分布的日本鲐（Scomber japonicus）和澳洲鲐（Scomber australasicus）为研究对象，分别采用形态学、线粒体DNA和微卫星三种标记来检测两种鲐属鱼类的群体遗传结构和系统地理格局，并探讨了其群体动态历史的演化过程，揭示历史因素、当前环境因素及鱼类自身的生态习性对海洋鱼类群体遗传结构和系统地理分布模式的综合作用。主要研究结果如下：
1.鲐属鱼类系统发育关系与进化历史
（1）采用传统形态和耳石形态两种研究方法，分析了三种鲐属鱼类（大西洋鲐、日本鲐和澳洲鲐）的系统发育关系，结果表明：在外观上大西洋鲐（Scomberscombrus）矢耳石与其他两种鲐鱼完全不同，日本鲐与澳洲鲐矢耳石形状相似；判别分析、主成分分析与聚类分析结果一致，均表明大西洋鲐与其他两种鲐鱼的形态差异明显，日本鲐和澳洲鲐的形态相似，亲缘关系紧密。
（2）本文以线粒体DNA（COI、Cyt b和控制区）和核基因（5S rDNA）为分子标记，结合线粒体基因和核基因各自的优点，在每个基因片段独立分析的基础上，利用贝叶斯联合模型分析方法重建鲐属鱼类的系统发育关系。结果显示：所有的系统发育分析均解决和支持鲐属鱼类的单系性；不同分区策略的贝叶斯分析获得了拓扑结构一致且各节点后验概率均较高的系统发育树，日本鲐与澳洲鲐聚为姐妹群，再与圆鲐聚类，大西洋鲐位于基部位置。
形态与分子数据均支持澳洲鲐的物种有效性及大西洋鲐与日本鲐和澳洲鲐存在较大差异。由于圆鲐整鱼样品的缺乏，未能从形态学角度对鲐属鱼类进行全面地分析。但本研究的分子证据强烈支持圆鲐的物种有效性，并将圆鲐置于日本鲐和澳洲鲐聚类分支的姐妹群位置上，进一步澄清了鲐属鱼类之间系统发育关系的疑问。此外，我们还将鲐属鱼类的地理分布与其系统关系相结合综合探讨了鲐属鱼类的起源进化历史，指出地理阻碍导致的分类阶元的隔离分化（vicariance）过程与各阶元在扩散分化的生态适应（adaption）过程相互补充交叉，促进了鲐属鱼类分化并逐渐形成现在的地理分布格局。
2.日本鲐种群遗传结构与分子系统地理学研究
（1）运用形态学方法分析了西北太平洋日本鲐群体之间的形态差异。结果显示，在传统形态和耳石形态上日本鲐各群体间均存在差异，但不同海域之间没有形成明显的分化。此外，对日本鲐不同体长组个体的耳石形态特征进行比较分析发现，日本鲐耳石形态特征随着鱼体生长而发生显著变化。
（2）应用线粒体DNA控制区对西北太平洋14个日本鲐地理群体开展了分子系统地理学研究，结果显示在日本鲐种内检测到了2个明显分化的系统发育类群，但两个类群在空间上的分布频率并不存在显著差异，即不存在地理结构，未检测到显著的群体遗传结构。我们推测更新世冰期的隔离很可能是日本鲐产生系统分化的主要原因，而冰川消退后的群体间的重新混合可能消除先前因隔离而积累的遗传差异。此外，核苷酸不配对分布和中性检验分析的结果表明西北太平洋日本鲐经历了近期群体扩张事件。
（3）应用微卫星分子标记开展了西北太平洋日本鲐的群体遗传结构分析。结果显示日本鲐在微卫星DNA上展现了较高的遗传多样性水平；通过检测8个微卫星多态位点差异，在日本鲐群体间存在显著的遗传分化，推测地理距离、海流以及日本鲐自身的生态特征很可能是导致群体间产生遗传差异的原因。
3.澳洲鲐种群遗传结构与分子系统地理学研究
（1）采用传统的形态度量与框架结构度量相结合的研究方法，分析了中日澳洲鲐群体之间的形态差异。多种多元统计方法结果基本一致，珠海群体与日本相模湾群体间存在形态差异。我们推测海流对澳洲鲐不同群体间的形态差异产生了重要影响。
（2）基于线粒体DNA控制区的分子系统地理学研究结果表明，澳洲鲐种内无谱系结构，在所研究范围内没有检测到明显的系统地理分化，一方面可能是因为间冰期澳洲鲐群体发生栖息地大规模扩张所引起的；另一方面可能是由澳洲鲐本身的生活史特征造成的，澳洲鲐成鱼进行长距离洄游，扩散能力较强，补充大且存在较长的浮游生活期，从而在洋流的作用下使群体间存在较高频率的基因交流。
（3）通过检测8个微卫星多态位点差异，在日本太平洋沿岸4个澳洲鲐地理群体中检测到微弱但显著的遗传结构，北太平洋中部群体与其他群体存在明显的遗传分化，距离隔离及勘察加寒流可能对澳洲鲐群体间基因交流产生了阻碍。同样，澳洲鲐在微卫星DNA上具有较高的遗传多样性水平。
Scomber species are epipelagic and pelagic fishes with gregarious behavior.Scomber mackerel fisheries are very important in many regions throughout theirdistributions. Despite its economic importance for international fisheries, thetaxonomy and phylogenetic relationships of Scomber have long been controversial. Inthe present study, phylogenetic analysis of Scomber was conducted usingmorphological, mitochondrial and nuclear DNA sequence data to clarify the currenttaxonomic classification, and to assess phylogenetic relationships and theevolutionary history of this genus. Additionally, we estimated the population structureand phylogeographic pattern of Scomber japonicus and Scomber australasicus inNorthwestern Pacific using morphological, mitochondrial DNA sequence andmicrosatellite DNA markers to infer the relative role of historical vicariant events,environmental factors and life history traits in shaping and maintaining the populationstructure pattern of these two species.
1. The taxonomy, and phylogeny and evolution of Scomber
(1) The phylogenetic relationships of three Scomber species was inferred byusing morphological and otolith shape analysis. The shape of Scomber scombrusotolith is totally different from those of S. japonicus and S. australasicus. The resultsof multivariate analysis consistently showed S. japonicus is morphologically similarto S. australasicus, whereas large difference was observed between S. scombrus andthe other two Scomber species based on morphological characters and otolith shapevariable.
(2) A molecular phylogenetic analysis of Scomber was conducted based onmitochondrial (COI, Cyt b and control region) and nuclear (5S rDNA) DNA sequencedata in multiple perspective. The present study produced a well-resolved phylogenythat strongly supported the monophyly of Scomber. We confirmed that S. japonicusand S. colias are genetically distinct. Although morphologically and ecologicallysimilar to S. colias, the molecular data showed that S. japonicus has a greatermolecular affinity with S. australasicus, which conflicts with the traditional taxonomy.This phylogenetic pattern was corroborated by the morphological data. Thepresent-day geographic range of each species was compared with the resultantphylogeny to evaluate possible hypotheses for its diversification and range expansion.In view of the unique geographic distributions of Scomber species, we hypothesizedthat speciation in the genus was primarily based on historical vicariance and adaption.
2. Population genetic structure and molecular phylogeography of S. japonicus
(1) Morphological characters and otolith shape variable were compared bymultivariate analysis for S. japonicus populations in Northwestern Pacific. Significantdifferences were found in many characters among eight S. japonicus populations.However, no geographical trend was observed. In addition, comparative analysis ofotolith shape variable from different length groups indicated that otolith shape of S.japonicus varied with growth rate.
(2) Molecular phylogeography of S. japonicus in Northwestern Pacific wasinvestigated using mitochondrial control region sequences. Two significantly distinctlineages were detected, which might be isolated during glacial periods of Pleistocene.However, the relative frequency of individuals occupying the two major lineages didnot differ significantly among sample locations. It is supposed that the presentdistribution of haplotypes is indicative of secondary contact with subsequent highgene flow over the sampled range. Both mismatch distribution analyses and neutralitytests suggested a late Pleistocene population expansion for S. japonicus.
(3) Genetic differentiation of S. japonicus in Northwestern Pacific was surveyedat eight microsatellite loci. Microsatellite analyses revealed relative high genetic diversity for S. japonicus due to high mutation rate of microsatellite DNA as well aslarge effective population size of this species. Significant population genetic structurewas detected among S. japonicus populations, which might be caused by thecomplicated interaction between marine currents and biological characteristics of S.japonicus.
3. Population genetic structure and molecular phylogeography of S. australasicus
(1) Morphological traits and truss network characters were measured toinvestigate morphological difference among S. australasicus populations.Multivariate analysis consistently found significant differences between China andJapan populations, which may be caused by marine currents in these two areas.
(2) The first hypervariable region (HVR-1) of the mitochondrial DNA controlregion was analyzed for samples collected from five locations in Northwestern Pacific.NJ tree and median-joining network revealed no significant phylogeographic structure.No significant genetic structure was detected among populations of S. australasicusanalyzed. It is likely a consequence of extensive gene flow caused by passive larvaltransport and/or extremely high vagility in adult as well as population expansionduring late Pleistocene.
(3) Weak but significant genetic structure was found in four S. australasicuspopulations in Japan’s Pacific coastal waters based on microsatellite analyses. Smallbut significant genetic differentiation between the middle of north Pacific populationand other populations might be attributed to the limited gene flow due to Kamchatka.In accordance with mtDNA result, microsatellite diversities were relatively high anddid not show any geographical trends.
1.鲐属鱼类系统发育关系与进化历史
（1）采用传统形态和耳石形态两种研究方法，分析了三种鲐属鱼类（大西洋鲐、日本鲐和澳洲鲐）的系统发育关系，结果表明：在外观上大西洋鲐（Scomberscombrus）矢耳石与其他两种鲐鱼完全不同，日本鲐与澳洲鲐矢耳石形状相似；判别分析、主成分分析与聚类分析结果一致，均表明大西洋鲐与其他两种鲐鱼的形态差异明显，日本鲐和澳洲鲐的形态相似，亲缘关系紧密。
（2）本文以线粒体DNA（COI、Cyt b和控制区）和核基因（5S rDNA）为分子标记，结合线粒体基因和核基因各自的优点，在每个基因片段独立分析的基础上，利用贝叶斯联合模型分析方法重建鲐属鱼类的系统发育关系。结果显示：所有的系统发育分析均解决和支持鲐属鱼类的单系性；不同分区策略的贝叶斯分析获得了拓扑结构一致且各节点后验概率均较高的系统发育树，日本鲐与澳洲鲐聚为姐妹群，再与圆鲐聚类，大西洋鲐位于基部位置。
形态与分子数据均支持澳洲鲐的物种有效性及大西洋鲐与日本鲐和澳洲鲐存在较大差异。由于圆鲐整鱼样品的缺乏，未能从形态学角度对鲐属鱼类进行全面地分析。但本研究的分子证据强烈支持圆鲐的物种有效性，并将圆鲐置于日本鲐和澳洲鲐聚类分支的姐妹群位置上，进一步澄清了鲐属鱼类之间系统发育关系的疑问。此外，我们还将鲐属鱼类的地理分布与其系统关系相结合综合探讨了鲐属鱼类的起源进化历史，指出地理阻碍导致的分类阶元的隔离分化（vicariance）过程与各阶元在扩散分化的生态适应（adaption）过程相互补充交叉，促进了鲐属鱼类分化并逐渐形成现在的地理分布格局。
2.日本鲐种群遗传结构与分子系统地理学研究
（1）运用形态学方法分析了西北太平洋日本鲐群体之间的形态差异。结果显示，在传统形态和耳石形态上日本鲐各群体间均存在差异，但不同海域之间没有形成明显的分化。此外，对日本鲐不同体长组个体的耳石形态特征进行比较分析发现，日本鲐耳石形态特征随着鱼体生长而发生显著变化。
（2）应用线粒体DNA控制区对西北太平洋14个日本鲐地理群体开展了分子系统地理学研究，结果显示在日本鲐种内检测到了2个明显分化的系统发育类群，但两个类群在空间上的分布频率并不存在显著差异，即不存在地理结构，未检测到显著的群体遗传结构。我们推测更新世冰期的隔离很可能是日本鲐产生系统分化的主要原因，而冰川消退后的群体间的重新混合可能消除先前因隔离而积累的遗传差异。此外，核苷酸不配对分布和中性检验分析的结果表明西北太平洋日本鲐经历了近期群体扩张事件。
（3）应用微卫星分子标记开展了西北太平洋日本鲐的群体遗传结构分析。结果显示日本鲐在微卫星DNA上展现了较高的遗传多样性水平；通过检测8个微卫星多态位点差异，在日本鲐群体间存在显著的遗传分化，推测地理距离、海流以及日本鲐自身的生态特征很可能是导致群体间产生遗传差异的原因。
3.澳洲鲐种群遗传结构与分子系统地理学研究
（1）采用传统的形态度量与框架结构度量相结合的研究方法，分析了中日澳洲鲐群体之间的形态差异。多种多元统计方法结果基本一致，珠海群体与日本相模湾群体间存在形态差异。我们推测海流对澳洲鲐不同群体间的形态差异产生了重要影响。
（2）基于线粒体DNA控制区的分子系统地理学研究结果表明，澳洲鲐种内无谱系结构，在所研究范围内没有检测到明显的系统地理分化，一方面可能是因为间冰期澳洲鲐群体发生栖息地大规模扩张所引起的；另一方面可能是由澳洲鲐本身的生活史特征造成的，澳洲鲐成鱼进行长距离洄游，扩散能力较强，补充大且存在较长的浮游生活期，从而在洋流的作用下使群体间存在较高频率的基因交流。
（3）通过检测8个微卫星多态位点差异，在日本太平洋沿岸4个澳洲鲐地理群体中检测到微弱但显著的遗传结构，北太平洋中部群体与其他群体存在明显的遗传分化，距离隔离及勘察加寒流可能对澳洲鲐群体间基因交流产生了阻碍。同样，澳洲鲐在微卫星DNA上具有较高的遗传多样性水平。
Scomber species are epipelagic and pelagic fishes with gregarious behavior.Scomber mackerel fisheries are very important in many regions throughout theirdistributions. Despite its economic importance for international fisheries, thetaxonomy and phylogenetic relationships of Scomber have long been controversial. Inthe present study, phylogenetic analysis of Scomber was conducted usingmorphological, mitochondrial and nuclear DNA sequence data to clarify the currenttaxonomic classification, and to assess phylogenetic relationships and theevolutionary history of this genus. Additionally, we estimated the population structureand phylogeographic pattern of Scomber japonicus and Scomber australasicus inNorthwestern Pacific using morphological, mitochondrial DNA sequence andmicrosatellite DNA markers to infer the relative role of historical vicariant events,environmental factors and life history traits in shaping and maintaining the populationstructure pattern of these two species.
1. The taxonomy, and phylogeny and evolution of Scomber
(1) The phylogenetic relationships of three Scomber species was inferred byusing morphological and otolith shape analysis. The shape of Scomber scombrusotolith is totally different from those of S. japonicus and S. australasicus. The resultsof multivariate analysis consistently showed S. japonicus is morphologically similarto S. australasicus, whereas large difference was observed between S. scombrus andthe other two Scomber species based on morphological characters and otolith shapevariable.
(2) A molecular phylogenetic analysis of Scomber was conducted based onmitochondrial (COI, Cyt b and control region) and nuclear (5S rDNA) DNA sequencedata in multiple perspective. The present study produced a well-resolved phylogenythat strongly supported the monophyly of Scomber. We confirmed that S. japonicusand S. colias are genetically distinct. Although morphologically and ecologicallysimilar to S. colias, the molecular data showed that S. japonicus has a greatermolecular affinity with S. australasicus, which conflicts with the traditional taxonomy.This phylogenetic pattern was corroborated by the morphological data. Thepresent-day geographic range of each species was compared with the resultantphylogeny to evaluate possible hypotheses for its diversification and range expansion.In view of the unique geographic distributions of Scomber species, we hypothesizedthat speciation in the genus was primarily based on historical vicariance and adaption.
2. Population genetic structure and molecular phylogeography of S. japonicus
(1) Morphological characters and otolith shape variable were compared bymultivariate analysis for S. japonicus populations in Northwestern Pacific. Significantdifferences were found in many characters among eight S. japonicus populations.However, no geographical trend was observed. In addition, comparative analysis ofotolith shape variable from different length groups indicated that otolith shape of S.japonicus varied with growth rate.
(2) Molecular phylogeography of S. japonicus in Northwestern Pacific wasinvestigated using mitochondrial control region sequences. Two significantly distinctlineages were detected, which might be isolated during glacial periods of Pleistocene.However, the relative frequency of individuals occupying the two major lineages didnot differ significantly among sample locations. It is supposed that the presentdistribution of haplotypes is indicative of secondary contact with subsequent highgene flow over the sampled range. Both mismatch distribution analyses and neutralitytests suggested a late Pleistocene population expansion for S. japonicus.
(3) Genetic differentiation of S. japonicus in Northwestern Pacific was surveyedat eight microsatellite loci. Microsatellite analyses revealed relative high genetic diversity for S. japonicus due to high mutation rate of microsatellite DNA as well aslarge effective population size of this species. Significant population genetic structurewas detected among S. japonicus populations, which might be caused by thecomplicated interaction between marine currents and biological characteristics of S.japonicus.
3. Population genetic structure and molecular phylogeography of S. australasicus
(1) Morphological traits and truss network characters were measured toinvestigate morphological difference among S. australasicus populations.Multivariate analysis consistently found significant differences between China andJapan populations, which may be caused by marine currents in these two areas.
(2) The first hypervariable region (HVR-1) of the mitochondrial DNA controlregion was analyzed for samples collected from five locations in Northwestern Pacific.NJ tree and median-joining network revealed no significant phylogeographic structure.No significant genetic structure was detected among populations of S. australasicusanalyzed. It is likely a consequence of extensive gene flow caused by passive larvaltransport and/or extremely high vagility in adult as well as population expansionduring late Pleistocene.
(3) Weak but significant genetic structure was found in four S. australasicuspopulations in Japan’s Pacific coastal waters based on microsatellite analyses. Smallbut significant genetic differentiation between the middle of north Pacific populationand other populations might be attributed to the limited gene flow due to Kamchatka.In accordance with mtDNA result, microsatellite diversities were relatively high anddid not show any geographical trends.
引文
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