银鱼科鱼类的分子系统发育研究
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摘要
银鱼科鱼类是我国重要经济鱼类,但其属间系统发育关系与属、种分类阶元界定尚存争议。本研究使用4种线粒体基因：12S与16S核糖体RNA基因(12S与16S)、细胞色素c氧化酶亚基1基因(COI)以及NADH脱氢酶亚基1基因(ND1)联合重建银鱼科属间系统发育。进一步选择7种核基因：28S核糖体基因(28S)、重组激活基因1(RAG1)、ZIC家族成员1基因(ZIC1)、外胚层神经皮层1类似蛋白基因(ENCC1)、RNF213基因(RNF213)、糖基转移酶基因(Glyt)、SH3与PX结构域3类似蛋白基因(SH3PX3)联合构建银鱼科属间系统发育。通过整合本研究的分子系统发育分析结果与前人形态学和分子学证据,探讨银鱼的属、种分类阶元界定。
基于4种线粒体基因的系统发育分析结果表明：1)银鱼科是单系群；2)安氏新银鱼与大银鱼互为姊妹群；3)‘短吻-陈氏-太湖-近太湖新银鱼复合群’与‘乔氏-寡齿新银鱼复合群’均为单系类群,但谱系内部的种间关系无法区分；4)短吻间银鱼先与居氏银鱼组成姊妹群,再一起同有明银鱼形成姊妹群关系；5)白肌银鱼为有效属,位于银鱼科基部。基于线粒体基因的分析无法确定日本银鱼属物种的位置。
基于7种核基因的系统发育分析结果支持线粒体分析的主要结论,即：1)银鱼科是单系群；2)‘短吻-陈氏-太湖-近太湖新银鱼复合群’与‘乔氏-寡齿新银鱼复合群’均为单系类群,但谱系内部的种间关系无法区分。同时,基于核基因的分析以很高的统计支持解决了线粒体基因不能很好解析的系统关系：1)银鱼科内部分为两个谱系,即‘大银鱼属+新银鱼属+石川日本银鱼’谱系和‘间银鱼属+银鱼属+白肌银鱼属+小齿日本银鱼’谱系；2)石川日本银鱼和小齿日本银鱼分别占据这两个谱系的基部位置。
综合线粒体和核基因的系统发育分析结果,本研究认为：1)银鱼科是一个单系类群；2)银鱼科内部可划分两大谱系,即‘大银鱼属+新银鱼属+石川日本银鱼’谱系和‘间银鱼属+银鱼属+白肌银鱼属+小齿日本银鱼’谱系,分别称为大银鱼亚科和银鱼亚科；3)间银鱼属不成立,短吻间银鱼归于银鱼属；4)建议依据石川日本银鱼建立新属；5)白肌银鱼属为有效属；6)支持将安氏新银鱼恢复为安氏大银鱼,归于大银鱼属；7)支持陈氏新银鱼、太湖新银鱼、近太湖新银鱼均是短吻新银鱼的同物异名；8)推测寡齿新银鱼和雷氏新银鱼为乔氏新银鱼的同物异名。
Noodle-fishes (Osmeriformes:Salangidae) are important commercial fishery species. However, their phylogenetic relationships are still unresolved and the taxonomy at the genus or species level has been disputed. This study examined the genus-level phylogenetic relationships of salangid fishes using four mitochondrial genes (12S and 16S ribosome RNA gene, cytochrome c oxidase subunit I gene and NADH dehydrogenase subunit 1 gene) and seven nuclear genes (28S ribosomal RNA gene, recombination activating gene 1, ectodermal-neural cortex 1-like protein gene, RNF213 gene, glycosyltransferase gene, SH3 and PX domain-containing 3-like protein gene, Zic family member 1 gene). Integrating the results of molecular phylogenetic analyses in this study with morphological and molecular evidences from previously published studies, we discussed taxonomic status of noodle-fishes at the genus or species level.
The phylogenetic analyses based on the four mitochondrial genes showed that:1) the salangids is a monophyletic group; 2) Protosalanx chinensis and Neosalanx anderssoni are sister gourps; 3) two monophyletic clades, 'Neosalanx brevirostris-tangkahkeii-taihuensis-pseudotaihuensis complex' and 'N. jordani-oligodontis complex' are revealed, but the species within the two clades can not be distinguished; 4) Hemisalanx brachyrostralis and Salanx cuvieri are sister groups, and these two species together formed the sister group of S. ariakensis; 5) Leucosoma is a valid genus located at the basal position of Salangidae. However, analysis on the mitochondrial genes could not confidently decide the phylogenetic position of the genus Salangichthys.
The phylogenetic analyses based on the seven nuclear genes confirmed the main conclusions from the study on mitochondrial genes:1) the monphyly of Salangidae; 2) the monophyletic 'N. brevirostris-tangkahkeii-taihuensis-pseudotaihuensis complex' and 'N. jordani-oligodontis complex'with undistinguishable interspecies relationship within the species complex. Moreover, the analyses on nuclear genes resolved the confused problem left by the analyses on mitochondrial genes:1) the family Salangidae could be divided into two clades, i.e. the'Protosalanx+Neosalanx +Salangichthys ishikawae' and the 'Hemisalanx + Salanx + Leucosoma + Salangichthys microdon'groups; 2) Salangichthys ishikawae and Salangichthys microdon occupy the basal positions of the above two groups, respectively.
lintegrating the results from mitochondrial and nuclear genes, we conclude that 1) Salangidae is a monophyletic group; 2) Salangidae could be divided into subfamily Protosalanginae (the Protosalanx + Neosalanx + Salangichthys ishikawae'group) and subfamily Salanginae (the 'Hemisalanx + Salanx + Leucosoma + Salangichthys microdon'group); 3) the genus Hemisalanx is invalid, species of Hemisalanx should be put in the genus Salanx; 4) Salangichthys ishikawae should be put in a new genus; 5) Leucosoma is a valid genus; 6) Neosalanx anderssoni should be reclassified as Protosalanx anderssoni, belonging to the genus Protosalanx; 7) Neosalanx tangkahkeii, N. taihuensis and N. pseudotaihuensis are synonyms of N. brevirostris; 8) Neosalanx oligodontis and N. reganius are synonyms of N. jordani.
基于4种线粒体基因的系统发育分析结果表明：1)银鱼科是单系群；2)安氏新银鱼与大银鱼互为姊妹群；3)‘短吻-陈氏-太湖-近太湖新银鱼复合群’与‘乔氏-寡齿新银鱼复合群’均为单系类群,但谱系内部的种间关系无法区分；4)短吻间银鱼先与居氏银鱼组成姊妹群,再一起同有明银鱼形成姊妹群关系；5)白肌银鱼为有效属,位于银鱼科基部。基于线粒体基因的分析无法确定日本银鱼属物种的位置。
基于7种核基因的系统发育分析结果支持线粒体分析的主要结论,即：1)银鱼科是单系群；2)‘短吻-陈氏-太湖-近太湖新银鱼复合群’与‘乔氏-寡齿新银鱼复合群’均为单系类群,但谱系内部的种间关系无法区分。同时,基于核基因的分析以很高的统计支持解决了线粒体基因不能很好解析的系统关系：1)银鱼科内部分为两个谱系,即‘大银鱼属+新银鱼属+石川日本银鱼’谱系和‘间银鱼属+银鱼属+白肌银鱼属+小齿日本银鱼’谱系；2)石川日本银鱼和小齿日本银鱼分别占据这两个谱系的基部位置。
综合线粒体和核基因的系统发育分析结果,本研究认为：1)银鱼科是一个单系类群；2)银鱼科内部可划分两大谱系,即‘大银鱼属+新银鱼属+石川日本银鱼’谱系和‘间银鱼属+银鱼属+白肌银鱼属+小齿日本银鱼’谱系,分别称为大银鱼亚科和银鱼亚科；3)间银鱼属不成立,短吻间银鱼归于银鱼属；4)建议依据石川日本银鱼建立新属；5)白肌银鱼属为有效属；6)支持将安氏新银鱼恢复为安氏大银鱼,归于大银鱼属；7)支持陈氏新银鱼、太湖新银鱼、近太湖新银鱼均是短吻新银鱼的同物异名；8)推测寡齿新银鱼和雷氏新银鱼为乔氏新银鱼的同物异名。
Noodle-fishes (Osmeriformes:Salangidae) are important commercial fishery species. However, their phylogenetic relationships are still unresolved and the taxonomy at the genus or species level has been disputed. This study examined the genus-level phylogenetic relationships of salangid fishes using four mitochondrial genes (12S and 16S ribosome RNA gene, cytochrome c oxidase subunit I gene and NADH dehydrogenase subunit 1 gene) and seven nuclear genes (28S ribosomal RNA gene, recombination activating gene 1, ectodermal-neural cortex 1-like protein gene, RNF213 gene, glycosyltransferase gene, SH3 and PX domain-containing 3-like protein gene, Zic family member 1 gene). Integrating the results of molecular phylogenetic analyses in this study with morphological and molecular evidences from previously published studies, we discussed taxonomic status of noodle-fishes at the genus or species level.
The phylogenetic analyses based on the four mitochondrial genes showed that:1) the salangids is a monophyletic group; 2) Protosalanx chinensis and Neosalanx anderssoni are sister gourps; 3) two monophyletic clades, 'Neosalanx brevirostris-tangkahkeii-taihuensis-pseudotaihuensis complex' and 'N. jordani-oligodontis complex' are revealed, but the species within the two clades can not be distinguished; 4) Hemisalanx brachyrostralis and Salanx cuvieri are sister groups, and these two species together formed the sister group of S. ariakensis; 5) Leucosoma is a valid genus located at the basal position of Salangidae. However, analysis on the mitochondrial genes could not confidently decide the phylogenetic position of the genus Salangichthys.
The phylogenetic analyses based on the seven nuclear genes confirmed the main conclusions from the study on mitochondrial genes:1) the monphyly of Salangidae; 2) the monophyletic 'N. brevirostris-tangkahkeii-taihuensis-pseudotaihuensis complex' and 'N. jordani-oligodontis complex'with undistinguishable interspecies relationship within the species complex. Moreover, the analyses on nuclear genes resolved the confused problem left by the analyses on mitochondrial genes:1) the family Salangidae could be divided into two clades, i.e. the'Protosalanx+Neosalanx +Salangichthys ishikawae' and the 'Hemisalanx + Salanx + Leucosoma + Salangichthys microdon'groups; 2) Salangichthys ishikawae and Salangichthys microdon occupy the basal positions of the above two groups, respectively.
lintegrating the results from mitochondrial and nuclear genes, we conclude that 1) Salangidae is a monophyletic group; 2) Salangidae could be divided into subfamily Protosalanginae (the Protosalanx + Neosalanx + Salangichthys ishikawae'group) and subfamily Salanginae (the 'Hemisalanx + Salanx + Leucosoma + Salangichthys microdon'group); 3) the genus Hemisalanx is invalid, species of Hemisalanx should be put in the genus Salanx; 4) Salangichthys ishikawae should be put in a new genus; 5) Leucosoma is a valid genus; 6) Neosalanx anderssoni should be reclassified as Protosalanx anderssoni, belonging to the genus Protosalanx; 7) Neosalanx tangkahkeii, N. taihuensis and N. pseudotaihuensis are synonyms of N. brevirostris; 8) Neosalanx oligodontis and N. reganius are synonyms of N. jordani.
引文
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