西北太平洋五种海洋鱼类的分子系统地理学研究
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摘要
本文以西北太平洋小黄鱼(Larimichthys polyactis)、高眼鲽(Cleisthenes herzensteini)、斯氏美首鲽(Glyptocephalus stelleri)、长鲽(Tanakius kitaharai)、白氏文昌鱼(branchiostoma belcheri)和带鱼(Trichiurus japonicus)为研究对象,采用线粒体DNA分子标记技术对这六种海洋物种的分子系统地理学进行了研究,系统研究了这五种鱼类和文昌鱼的分子系统地理分布模式,阐述了五种海洋鱼类和文昌鱼的遗传多样性水平和遗传结构现状,探讨了其群体动态历史的演化过程,揭示历史因素(更新世冰期等)和当前环境因素(洋流等)对塑造海洋鱼类群体遗传结构和分子系统地理分布模式的重要作用。主要研究结果如下：
1、研究了小黄鱼群体遗传多样性水平、遗传结构现状及其群体动态历史,对小黄鱼分布区内16个群体共298个个体的线粒体DNA控制区第一高变区序列进行了测定和分析。在小黄鱼控制区序列上检测到了高水平的基因多样度,揭示小黄鱼存在较高水平的遗传多样性。分子方差分析(AMOVA)和两两群体相比较的FST结果显示小黄鱼在所研究范围内存在显著的遗传结构。在小黄鱼群体内,我们检测到两个单倍型类群,这两个单倍型类群间的分化发生于更新世晚期,更新世冰期海平面的下降使西北太平洋边缘海之间产生隔离,这可能是两个单倍型类群之间发生分化的原因。两个单倍型类群在地域上的分布频率具有明显差异。最小跨度树(MST)结果和无限突变位点模型中性检验及核苷酸不配对分布表明小黄鱼经历了更新世的群体扩张(61kya-245kya)事件。
2、种内系统进化关系为探讨历史因素和当前因素如何塑造群体当前分布格局提供了可能。为检测高眼鲽的群体遗传结构、群体动态历史及其殖化进程,我们对其分布范围内6个群体共121个个体的线粒体控制区序列进行了测定和分析。在高眼鲽控制区序列上检测到了较高水平的基因多样度,揭示高眼鲽可能存在较高水平的遗传多样度。群体遗传结构研究结果显示：高眼鲽在其分布范围内不存在IBD模式；分子方差分析(AMOVA)和FST结果显示高眼鲽在其分布范围内不存在显著的遗传分化；群体分化的确切检验结果显示不同地理群体个体间是随机交配的。中性检验及核苷酸不配对分布表明高眼鲽经历了更新世的群体扩张(94kya～376kya)事件；嵌套分支系统地理学分析(NCPA)结果显示,高眼鲽栖息地连续扩散是造成其当前的地理分布模式的主要原因。更新世冰期高眼鲽扩散后新建立的群体尚未在迁移与遗传漂变之间取得平衡,这是造成其在所研究范围内缺少与地理相对应的系统地理结构的原因。
3、采用线粒体DNA控制区部分序列对日本沿海的斯氏美首鲽群体遗传结构和群体动态历史进行了研究。采集了日本沿海斯氏美首鲽5个群体共143个个体。研究结果显示线粒体DNA控制区序列检测到高的单倍型多样度(h=0.99±0.004),揭示斯氏美首鲽具有较高的种内多态。单倍型最小跨度树(MST)、中性检验和核苷酸不配对分布(Mismatch)分析结果显示斯氏美首鲽群体经历了近期的群体扩张事件(124.1kya-413.4kya)。分子方差分析和两两群体相比较的Fst分析结果显示在日本沿海分布的斯氏美首鲽群体间未检测到显著的遗传分化。
4、更新世冰期海平面的下降对物种当前的分布格局具有重要的影响,为检测更新世冰期对分布于日本海和外太平洋物种群落遗传格局的影响,我们采用线粒体DNA控制区部分序列对日本海和外太平洋的长鲽群体展开研究。在3个地理群体66个个体中共检测到35个单倍型。三个群体的单倍型多样性指数和核苷酸多态性指数分别在0.93和0.95,0.009和0.011之间。群体内和群体间的序列差异不显著,揭示长鲽群体的同质性。分子方差分析和两两群体相比较的Fst分析结果显示日本海群体和外太平洋群体之间无显著的遗传分化。最小跨度树(MST)和NJ关系树的结果显示,长鲽群体中存在微弱的谱系分化,35个单倍型明显的分成四个小支(与地理不相对应),这可能是由于更新世冰期海平面反复波动导致长鲽群体发生了近期隔离,同时研究结果还揭示长鲽群体间存在较强的基因交流。
5、以往的研究显示分布于中国南北的文昌鱼为白氏文昌鱼亚种和白氏文昌鱼青岛亚种,为了确定中国北方和日本白氏文昌鱼的分类地位,本研究采集了中国南方、北方的白氏文昌鱼样本,并结合日本文昌鱼的线粒体基因组信息展开了研究。基于线粒体DNA的COI,Cytb和16S rRNA片段研究结果显示,中国南北的白氏文昌鱼遗传差异分别达到了0.19,0.21和0.17,远远高于其他种的种内差异水平；中国北方白氏文昌鱼和日本白氏文昌鱼的遗传差异皆小于0.01。三个片段的系统发育研究结果显示,白氏文昌鱼明显的分为两支：中国南方支系和中国北方及日本支系。本研究的结果显示中国南北白氏文昌鱼达到了种间分化水平,基于动物命名法优先原则和本研究的结果,认为中国北方和日本的白氏文昌鱼应命名为青岛文昌鱼；白氏文昌鱼和青岛文昌鱼的分化事件发生于39.90Mya-43.24 Mya.
6、为检测带鱼的群体遗传多样性水平、遗传结构现状及其群体动态历史,我们采集了带鱼分布区内3个群体共54个个体的线粒体DNA控制区序列进行了测定和分析。在带鱼控制区序列上检测到了较高水平的基因多样度,揭示带鱼存在较高水平的遗传多样性。最小跨度树(MST)结果和中性检验及核苷酸不配对分布表明带鱼经历了更新世的群体扩张(49.3 kya-197 kya)事件。分子方差分析和FST结果显示带鱼在所研究范围内不存在显著的遗传结构。确切检验结果显示带鱼在所研究的范围内为一个未分化的群体。带鱼群体在所研究范围内没有检测到显著的系统地理分化,一方面可能是因为间冰期带鱼群体在中国大陆架区发生大范围栖息地扩张所引起的；另一方面带鱼产浮性卵从而在洋流的作用下使群体间存在较高频率的基因交流。
In this study, the molecular phylogeography of Larimichthys polyactis, Cleisthenes herzensteini, Glyptocephalus stelleri, Tanakius kitaharai, Branchiostoma, Trichiurus japonicus in the Northwestern Pacific were studied by mitochondrial DNA marker. The phylogeographic patterns, genetic diversities and population structures of the six marine fishes were carried out and the population dynamics of historical evolution were also reconstructed in the present studies. We discussed the role of historical and contemporary factors in shaping the phylogeographic pattern and genetic structures of marine species.
1、The genetic diversity and population genetic structure of the small yellow croaker (Larimichthys polyactis) were investigated.298 individuals were sampled from 16 localities ranged its distribution. Genetic variation in DNA sequences were examined from the first hypervariable region (HVR-1) of the mitochondrial DNA control region. High levels of haplotype diversity (h=0.992±0.002) in the HVR-1 region were detected, indicating a high level of genetic diversity. Hierarchical molecular variance analysis (AMOVA) and conventional population Fst comparisons revealed significant genetic structure throughout its range, which is inconsistent with previous findings based on the morphological and ecological studies. Two distinct lineages were found, which might be isolated and diverged in different marginal seas of the Northwestern Pacific during Pleitocene low sea level stands. There were strong geographical differences in haplotype frequencies of the 2 lineages. The star burst structure of the minimum spanning tree also suggested a very recent origin for most haplotypes. The demographic history of L. polyactis was examined by using neutrality tests and mismatch distribution analysis, which indicated a Pleistocene population expansion at about 61,000～245,000 years ago.
2、Intraspecific phylogenies can provide useful insights into how populations have been shaped by historical and contemporary processes. To determine the population genetic structure and the demographic and colonization history of Cleisthenes herzensteini in the Northwestern Pacific, one hundred and twenty-one individuals were sampled from six localities along the coastal regions of Japan and the Yellow Sea of China. Mitochondrial DNA variation was analyzed using DNA sequence data from the 5'end of control region. High levels of haplotype diversity (>0.96) were found for all populations, indicating a high level of genetic diversity. No pattern of isolation by distance was detected among the population differentiation throughout the examined range. Analyses of molecular variance (AMOVA) and the conventional population statistic Fst revealed no significant population genetic structure among populations. And according to the exact test of differentiation among populations, the null hypothesis that C. herzensteini within the examined range constituted a panmictic mtDNA gene pool was accepted. The demographic history of C. herzensteini was examined using neutrality test and mismatch distribution analyses and results indicated Pleistocene population expansion (about 94kya～376kya) in the species, which was consistent with the inference result of nested clade phylogeographical analysis (NCPA) showing contiguous range expansion for C. herzensteini. The lack of phylogeographical structure for the species may reflect a recent range expansion after the glacial maximum and insufficient time to attain migration-drift equilibrium.
3、The demographic history and population genetic structure of blackfin flounder (Glyptocephalus stelleri) along coastal regions of Japan were investigated. Genetic variation in DNA sequences were examined from the first hypervariable region (HVR-1) of the mitochondrial DNA control region. High level of haplotypic diversity (h=0.99±0.004) was detected, indicating a high level of intrapopulation genetic diversity. The starburst structure of the minimum spanning tree (MST) suggested a very recent origin for most haplotypes. The demographic history of G stelleri was examined by using neutrality tests and mismatch distribution analysis, which also indicated a Pleistocene population expansion at about 124,100-413,400 years ago. Hierarchical molecular variance analysis (AMOVA) and conventional population Fst comparisons revealed no significant genetic differentiation existed throughout the examined range.
4、The first hypervariable region (HVR-1) of the mitochondrial DNA control region was utilized for determination of genetic variation and population structure in willowy flounder (Tanakius kitaharai) collected from Aomori, Ibaraki and Niigata. A total of 35 haplotypes were detected among 66 individuals with a total of 30 variable sites out of 387 bp sequenced. Average sequence differences between populations (1.0 1.1%) were comparable to those within populations (0.9-1.2%), suggesting no genetic heterogeneity among samples. The pattern of distribution of genetic variability with high level of haplotype diversity (h=0.94) and moderate nucleotide diversity (n=1.0%) was also detected in the HVR-1 region of the mitochondrial DNA control region. AMOVA tests and the conventional populationΦst comparisons revealed no significant genetic structure among the populations. Partitioning populations into coherent geographic groups divided willowy flounder samples (Φct=-0.007, P>0.05) into two major groups:a Sea of Japan group composed of Aomori and Niigata populations; a Pacific Ocean group composed of Ibaraki populations. The minimum spanning tree constructed with 35 haplotypes showed four low-divergent clades, corresponding to those defined in the NJ tree. However, these clades did not appear to have geographic structure. Altogether, the results indicate that willowy flounder is panmictic throughout the examined range in Aomori, Ibaraki and Niigata.
5、It is commonly accepted that the Branchiostoma lancelets in South and North China belong to Branchiostoma belcheri belcheri (Gray,1847) and Branchiostoma belcheri tsingtauense (Tchang and Koo,1934), respectively. Three partial mitochondrial DNA (mtDNA) fragments of cytochrome oxidase c subunit I (COI), cytochrome b (Cytb), and 16S ribosomal RNA (16S rRNA) genes were sequenced to analyze phylogenetic relationships of the Branchiostoma lancelets from South (Xiamen) and North (Qingdao and Rizhao) China, and phylogenetic trees constructed also included the existing data from Japanese waters. The genetic distances of the lancelets between South and North China averaged 0.19,0.21, and 0.17 based on partial sequences analysis of COI, Cytb, and 16S rRNA genes, respectively, which were much higher than those were observed in other intraspecific variations. However, the value between North China and Japanese waters was only 0.01 based on partial sequences analysis of three mtDNA genes, which indicated low intraspecific genetic divergence existed in the two areas. The results also clearly indicated two monophyletic clades (clade A (North China and Japanese waters), clade B (South China)) existed in the specimens, corresponding to the South and North China, respectively. Above all, our results indicate that the Branchiostoma lancelets in South and North China should belong to different species, and the subspecies B. belcheri tsingtauense together with the lancelets in most Japanese waters is an independent species. According to the rule of priority and present studies, the Branchiostoma lancelets in North China and most Japanese waters should be revised to B. tsingtauense. The divergence time between B. belcheri and B. tsingtauense was estimated at about 39.90-43.24 million years ago.
6、The genetic diversity and population genetic structure of Trichiurus japonicus were investigated. Fifty-four individuals were sampled from 3 localities of East China Sea and South China Sea. Genetic variation in DNA sequences were examined from the mitochondrial DNA control region. High levels of haplotype diversity (h=0.98±0.01) in the control region were detected, indicating a high level of genetic diverstiy. A total of 42 polymorphic sites were found, and 40 haplotypes were defined. The pairwise nucleotide differences between samples ranged from 4.65±2.38 to 4.87±2.48. The demographic history of T. japonicus was examined by using neutrality tests and mismatch distribution analysis, which indicated a Pleistocene population expansion at about 49,300-197,000 yeas ago. The star burst structure of the minimum spanning tree also suggestted a very recent origin for most haplotypes. Hierarchical molecular variance analysis (AMOVA) and conventional population Fst comparisons revealed no significant genetic structure throughout the examined range. And according to the exact test of differentiation among populations, the null hypothesis that T. japonicus within the examined range constituted a panmictic mtDNA gene pool was accepted. Recent population expansion and larval dispersal likely have contributed to the genetically homogeneous population structure of the species. The knowledge on genetic diversity and genetic structure will be crucial to establish appropriate fishery management stocks for the species.
1、研究了小黄鱼群体遗传多样性水平、遗传结构现状及其群体动态历史,对小黄鱼分布区内16个群体共298个个体的线粒体DNA控制区第一高变区序列进行了测定和分析。在小黄鱼控制区序列上检测到了高水平的基因多样度,揭示小黄鱼存在较高水平的遗传多样性。分子方差分析(AMOVA)和两两群体相比较的FST结果显示小黄鱼在所研究范围内存在显著的遗传结构。在小黄鱼群体内,我们检测到两个单倍型类群,这两个单倍型类群间的分化发生于更新世晚期,更新世冰期海平面的下降使西北太平洋边缘海之间产生隔离,这可能是两个单倍型类群之间发生分化的原因。两个单倍型类群在地域上的分布频率具有明显差异。最小跨度树(MST)结果和无限突变位点模型中性检验及核苷酸不配对分布表明小黄鱼经历了更新世的群体扩张(61kya-245kya)事件。
2、种内系统进化关系为探讨历史因素和当前因素如何塑造群体当前分布格局提供了可能。为检测高眼鲽的群体遗传结构、群体动态历史及其殖化进程,我们对其分布范围内6个群体共121个个体的线粒体控制区序列进行了测定和分析。在高眼鲽控制区序列上检测到了较高水平的基因多样度,揭示高眼鲽可能存在较高水平的遗传多样度。群体遗传结构研究结果显示：高眼鲽在其分布范围内不存在IBD模式；分子方差分析(AMOVA)和FST结果显示高眼鲽在其分布范围内不存在显著的遗传分化；群体分化的确切检验结果显示不同地理群体个体间是随机交配的。中性检验及核苷酸不配对分布表明高眼鲽经历了更新世的群体扩张(94kya～376kya)事件；嵌套分支系统地理学分析(NCPA)结果显示,高眼鲽栖息地连续扩散是造成其当前的地理分布模式的主要原因。更新世冰期高眼鲽扩散后新建立的群体尚未在迁移与遗传漂变之间取得平衡,这是造成其在所研究范围内缺少与地理相对应的系统地理结构的原因。
3、采用线粒体DNA控制区部分序列对日本沿海的斯氏美首鲽群体遗传结构和群体动态历史进行了研究。采集了日本沿海斯氏美首鲽5个群体共143个个体。研究结果显示线粒体DNA控制区序列检测到高的单倍型多样度(h=0.99±0.004),揭示斯氏美首鲽具有较高的种内多态。单倍型最小跨度树(MST)、中性检验和核苷酸不配对分布(Mismatch)分析结果显示斯氏美首鲽群体经历了近期的群体扩张事件(124.1kya-413.4kya)。分子方差分析和两两群体相比较的Fst分析结果显示在日本沿海分布的斯氏美首鲽群体间未检测到显著的遗传分化。
4、更新世冰期海平面的下降对物种当前的分布格局具有重要的影响,为检测更新世冰期对分布于日本海和外太平洋物种群落遗传格局的影响,我们采用线粒体DNA控制区部分序列对日本海和外太平洋的长鲽群体展开研究。在3个地理群体66个个体中共检测到35个单倍型。三个群体的单倍型多样性指数和核苷酸多态性指数分别在0.93和0.95,0.009和0.011之间。群体内和群体间的序列差异不显著,揭示长鲽群体的同质性。分子方差分析和两两群体相比较的Fst分析结果显示日本海群体和外太平洋群体之间无显著的遗传分化。最小跨度树(MST)和NJ关系树的结果显示,长鲽群体中存在微弱的谱系分化,35个单倍型明显的分成四个小支(与地理不相对应),这可能是由于更新世冰期海平面反复波动导致长鲽群体发生了近期隔离,同时研究结果还揭示长鲽群体间存在较强的基因交流。
5、以往的研究显示分布于中国南北的文昌鱼为白氏文昌鱼亚种和白氏文昌鱼青岛亚种,为了确定中国北方和日本白氏文昌鱼的分类地位,本研究采集了中国南方、北方的白氏文昌鱼样本,并结合日本文昌鱼的线粒体基因组信息展开了研究。基于线粒体DNA的COI,Cytb和16S rRNA片段研究结果显示,中国南北的白氏文昌鱼遗传差异分别达到了0.19,0.21和0.17,远远高于其他种的种内差异水平；中国北方白氏文昌鱼和日本白氏文昌鱼的遗传差异皆小于0.01。三个片段的系统发育研究结果显示,白氏文昌鱼明显的分为两支：中国南方支系和中国北方及日本支系。本研究的结果显示中国南北白氏文昌鱼达到了种间分化水平,基于动物命名法优先原则和本研究的结果,认为中国北方和日本的白氏文昌鱼应命名为青岛文昌鱼；白氏文昌鱼和青岛文昌鱼的分化事件发生于39.90Mya-43.24 Mya.
6、为检测带鱼的群体遗传多样性水平、遗传结构现状及其群体动态历史,我们采集了带鱼分布区内3个群体共54个个体的线粒体DNA控制区序列进行了测定和分析。在带鱼控制区序列上检测到了较高水平的基因多样度,揭示带鱼存在较高水平的遗传多样性。最小跨度树(MST)结果和中性检验及核苷酸不配对分布表明带鱼经历了更新世的群体扩张(49.3 kya-197 kya)事件。分子方差分析和FST结果显示带鱼在所研究范围内不存在显著的遗传结构。确切检验结果显示带鱼在所研究的范围内为一个未分化的群体。带鱼群体在所研究范围内没有检测到显著的系统地理分化,一方面可能是因为间冰期带鱼群体在中国大陆架区发生大范围栖息地扩张所引起的；另一方面带鱼产浮性卵从而在洋流的作用下使群体间存在较高频率的基因交流。
In this study, the molecular phylogeography of Larimichthys polyactis, Cleisthenes herzensteini, Glyptocephalus stelleri, Tanakius kitaharai, Branchiostoma, Trichiurus japonicus in the Northwestern Pacific were studied by mitochondrial DNA marker. The phylogeographic patterns, genetic diversities and population structures of the six marine fishes were carried out and the population dynamics of historical evolution were also reconstructed in the present studies. We discussed the role of historical and contemporary factors in shaping the phylogeographic pattern and genetic structures of marine species.
1、The genetic diversity and population genetic structure of the small yellow croaker (Larimichthys polyactis) were investigated.298 individuals were sampled from 16 localities ranged its distribution. Genetic variation in DNA sequences were examined from the first hypervariable region (HVR-1) of the mitochondrial DNA control region. High levels of haplotype diversity (h=0.992±0.002) in the HVR-1 region were detected, indicating a high level of genetic diversity. Hierarchical molecular variance analysis (AMOVA) and conventional population Fst comparisons revealed significant genetic structure throughout its range, which is inconsistent with previous findings based on the morphological and ecological studies. Two distinct lineages were found, which might be isolated and diverged in different marginal seas of the Northwestern Pacific during Pleitocene low sea level stands. There were strong geographical differences in haplotype frequencies of the 2 lineages. The star burst structure of the minimum spanning tree also suggested a very recent origin for most haplotypes. The demographic history of L. polyactis was examined by using neutrality tests and mismatch distribution analysis, which indicated a Pleistocene population expansion at about 61,000～245,000 years ago.
2、Intraspecific phylogenies can provide useful insights into how populations have been shaped by historical and contemporary processes. To determine the population genetic structure and the demographic and colonization history of Cleisthenes herzensteini in the Northwestern Pacific, one hundred and twenty-one individuals were sampled from six localities along the coastal regions of Japan and the Yellow Sea of China. Mitochondrial DNA variation was analyzed using DNA sequence data from the 5'end of control region. High levels of haplotype diversity (>0.96) were found for all populations, indicating a high level of genetic diversity. No pattern of isolation by distance was detected among the population differentiation throughout the examined range. Analyses of molecular variance (AMOVA) and the conventional population statistic Fst revealed no significant population genetic structure among populations. And according to the exact test of differentiation among populations, the null hypothesis that C. herzensteini within the examined range constituted a panmictic mtDNA gene pool was accepted. The demographic history of C. herzensteini was examined using neutrality test and mismatch distribution analyses and results indicated Pleistocene population expansion (about 94kya～376kya) in the species, which was consistent with the inference result of nested clade phylogeographical analysis (NCPA) showing contiguous range expansion for C. herzensteini. The lack of phylogeographical structure for the species may reflect a recent range expansion after the glacial maximum and insufficient time to attain migration-drift equilibrium.
3、The demographic history and population genetic structure of blackfin flounder (Glyptocephalus stelleri) along coastal regions of Japan were investigated. Genetic variation in DNA sequences were examined from the first hypervariable region (HVR-1) of the mitochondrial DNA control region. High level of haplotypic diversity (h=0.99±0.004) was detected, indicating a high level of intrapopulation genetic diversity. The starburst structure of the minimum spanning tree (MST) suggested a very recent origin for most haplotypes. The demographic history of G stelleri was examined by using neutrality tests and mismatch distribution analysis, which also indicated a Pleistocene population expansion at about 124,100-413,400 years ago. Hierarchical molecular variance analysis (AMOVA) and conventional population Fst comparisons revealed no significant genetic differentiation existed throughout the examined range.
4、The first hypervariable region (HVR-1) of the mitochondrial DNA control region was utilized for determination of genetic variation and population structure in willowy flounder (Tanakius kitaharai) collected from Aomori, Ibaraki and Niigata. A total of 35 haplotypes were detected among 66 individuals with a total of 30 variable sites out of 387 bp sequenced. Average sequence differences between populations (1.0 1.1%) were comparable to those within populations (0.9-1.2%), suggesting no genetic heterogeneity among samples. The pattern of distribution of genetic variability with high level of haplotype diversity (h=0.94) and moderate nucleotide diversity (n=1.0%) was also detected in the HVR-1 region of the mitochondrial DNA control region. AMOVA tests and the conventional populationΦst comparisons revealed no significant genetic structure among the populations. Partitioning populations into coherent geographic groups divided willowy flounder samples (Φct=-0.007, P>0.05) into two major groups:a Sea of Japan group composed of Aomori and Niigata populations; a Pacific Ocean group composed of Ibaraki populations. The minimum spanning tree constructed with 35 haplotypes showed four low-divergent clades, corresponding to those defined in the NJ tree. However, these clades did not appear to have geographic structure. Altogether, the results indicate that willowy flounder is panmictic throughout the examined range in Aomori, Ibaraki and Niigata.
5、It is commonly accepted that the Branchiostoma lancelets in South and North China belong to Branchiostoma belcheri belcheri (Gray,1847) and Branchiostoma belcheri tsingtauense (Tchang and Koo,1934), respectively. Three partial mitochondrial DNA (mtDNA) fragments of cytochrome oxidase c subunit I (COI), cytochrome b (Cytb), and 16S ribosomal RNA (16S rRNA) genes were sequenced to analyze phylogenetic relationships of the Branchiostoma lancelets from South (Xiamen) and North (Qingdao and Rizhao) China, and phylogenetic trees constructed also included the existing data from Japanese waters. The genetic distances of the lancelets between South and North China averaged 0.19,0.21, and 0.17 based on partial sequences analysis of COI, Cytb, and 16S rRNA genes, respectively, which were much higher than those were observed in other intraspecific variations. However, the value between North China and Japanese waters was only 0.01 based on partial sequences analysis of three mtDNA genes, which indicated low intraspecific genetic divergence existed in the two areas. The results also clearly indicated two monophyletic clades (clade A (North China and Japanese waters), clade B (South China)) existed in the specimens, corresponding to the South and North China, respectively. Above all, our results indicate that the Branchiostoma lancelets in South and North China should belong to different species, and the subspecies B. belcheri tsingtauense together with the lancelets in most Japanese waters is an independent species. According to the rule of priority and present studies, the Branchiostoma lancelets in North China and most Japanese waters should be revised to B. tsingtauense. The divergence time between B. belcheri and B. tsingtauense was estimated at about 39.90-43.24 million years ago.
6、The genetic diversity and population genetic structure of Trichiurus japonicus were investigated. Fifty-four individuals were sampled from 3 localities of East China Sea and South China Sea. Genetic variation in DNA sequences were examined from the mitochondrial DNA control region. High levels of haplotype diversity (h=0.98±0.01) in the control region were detected, indicating a high level of genetic diverstiy. A total of 42 polymorphic sites were found, and 40 haplotypes were defined. The pairwise nucleotide differences between samples ranged from 4.65±2.38 to 4.87±2.48. The demographic history of T. japonicus was examined by using neutrality tests and mismatch distribution analysis, which indicated a Pleistocene population expansion at about 49,300-197,000 yeas ago. The star burst structure of the minimum spanning tree also suggestted a very recent origin for most haplotypes. Hierarchical molecular variance analysis (AMOVA) and conventional population Fst comparisons revealed no significant genetic structure throughout the examined range. And according to the exact test of differentiation among populations, the null hypothesis that T. japonicus within the examined range constituted a panmictic mtDNA gene pool was accepted. Recent population expansion and larval dispersal likely have contributed to the genetically homogeneous population structure of the species. The knowledge on genetic diversity and genetic structure will be crucial to establish appropriate fishery management stocks for the species.
引文
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