【作者】 李隽；

【导师】 傅萃长；

【作者基本信息】 复旦大学 ， 生态学， 2011， 博士

【副题名】兼论裂腹鱼属鱼类与两栖动物小鲵属物种多样性空间格局的成因

【摘要】 真骨鱼次亚纲Teleostei,俗称真骨鱼类,约占全世界已记录28000种鱼类的96%左右。真骨鱼类下设海鲢亚部Elopomorpha、骨舌鱼亚部Osteoglossomorpha、骨鳔鱼亚部Ostarioclupeomorpha (= Otocephala)和真真骨鱼亚部Euteleostei。真真骨鱼亚部,俗称真真骨鱼类,是真骨鱼类中物种多样性最丰富的类群,大约17500种。真真骨鱼类可划分为原棘鳍总目Protacanthopterygii和新真骨鱼类Neoteleostei两大类群。原棘鳍总目,俗称低等真真骨鱼类,包括鲑形目Salmoniformes、狗鱼目Esociformes、水珍鱼目Argentiniformes与胡瓜鱼目Osmeriformes (sensu Nelson,2006)。低等真真骨鱼类的系统发育关系是鱼类系统学中长期存在争议的科学问题之一。基于线粒体全基因组与核基因多位点的分子证据,本论文重建低等真真骨鱼类的系统发育关系,聚焦于解决三个问题：1)‘迷一样’的低等真真骨鱼类鳞南乳鱼科Lepidogalaxiidae的系统发育位置；2)低等真真骨鱼类目间分子系统发育关系；3)胡瓜鱼目鱼类分子系统发育关系。另外,本论文亦开展了物种多样性空间格局的成因研究。主要聚焦于物种多样性海拔分布格局与间断分布格局的成因：1)选择青藏高原及其邻近地区特有的裂腹鱼属Schizothorax鱼类作为模式类群,分析气候、面积、分化速率及定居时间与物种数之间的关系,探讨鱼类多样性海拔分布格局的生态与进化成因；2)选择东亚地区两栖动物小鲵属Hynobius作为模式类群,探讨新生代东亚边缘地质构造演化在小鲵属大陆—岛屿间断分布格局形成中的作用。主要研究结果如下：1、基于线粒体基因组证据的鳞南乳鱼科系统发育位置基于线粒体基因组,重建了低等真真骨鱼类系统发育关系。结果表明：1)原先放在胡瓜鱼目中的鳞南乳鱼科位于真真骨鱼类的基部,是其余所有真真骨鱼类的姊妹群；2)原先放在胡瓜鱼目中的南乳鱼科Galaxiidae作为单系群,建议提升为南乳鱼目Galaxiiformes; 3)重新定义胡瓜鱼目仅包括青瓜鱼科Retropinnidae、胡瓜鱼科Osmeridae、银鱼科Salangidae与香鱼科Plecoglossidae。2、基于核基因证据的低等真真骨鱼类目间系统发育关系基于10个核基因的部分序列,重建了低等真真骨鱼类目间系统发育关系。结果表明：1)原先放在水珍鱼目中的黑头鱼亚目Alepocephaloidei和骨鳔鱼亚部鱼类形成姊妹群关系,同意基于线粒体基因组证据提出的提升黑头鱼亚目为黑头鱼目Alepocephaliformes的建议；2)同意基于线粒体基因组证据提出的将鳞南乳鱼科放在真真骨鱼类基部的观点,并建议提升为鳞南乳鱼目Lepidogalaxiiformes;3)同意基于线粒体基因组证据提出的水珍鱼目、鲑形目和狗鱼目鱼类组成单系类群,以及胡瓜鱼目和巨口鱼目Stomiiformes形成姊妹群关系的观点；4)南乳鱼目是新真骨鱼类Neoteleostei(减去巨口鱼目)的姊妹群。3、胡瓜鱼目鱼类的分子系统发育关系基于9个核基因的部分序列,重建了胡瓜鱼目鱼类的分子系统发育关系。结果表明：1)青瓜鱼科位于胡瓜鱼目基部；2)银鱼科和香鱼科形成姊妹群关系,它们一起构成胡瓜鱼科的姊妹群；3)在胡瓜鱼科内部,毛鳞鱼属Mallotus占据基部位置,胡瓜鱼属Osmerus和油胡瓜鱼属+异胡瓜鱼属+蜡鱼属Spirinchus+ Allosmerus+Thaleichthys形成姊妹群关系,这两个类群共同构成公鱼属Hypomesus的姊妹群；4)在银鱼科内部,同意本实验室先前的结果,即大银鱼属Protosalanx和新日本银鱼属Neosalangichthys构成大银鱼亚科Protosalanginae,银鱼属Salanx、白肌银鱼属Leucosoma和日本银鱼属Salangichthys构成银鱼亚科Salanginae; 5)在青瓜鱼科内部,南茴鱼属Prototroctes占据基部位置,青瓜鱼属Retropinna存在并系发生现象,新岛青瓜鱼(Stokellia anisodon)和青瓜鱼(Retropinna retropinna)形成姊妹群。4、裂腹鱼属鱼类物种多样性海拔分布格局的成因选择裂腹鱼属鱼类为模式类群,对鱼类物种多样性海拔分布格局的生态与进化成因进行了探讨。结果表明：1)裂腹鱼属鱼类的物种数在中海拔(1700-2200m)出现峰值；2)年均降水、面积、Mid-domain效应和分化速率均不能很好地预测物种多样性的海拔格局；3)年均温度和定居时间是物种多样性海拔格局最重要的影响因子。5、东亚地区两栖动物小鲵属间断分布格局的成因选择两栖动物小鲵属为模式类群,对新生代东亚边缘地质构造演化在小鲵属大陆—岛屿间断分布格局形成中的作用进行了探讨。结果表明：1)现存小鲵属物种起源于早始新世的西南日本和北海道岛；2)中始新世西南日本和北海道岛的隔离导致了北海道特有的滞育小鲵与其余所有小鲵的分化；3)从晚始新世到晚中新世,分布于西南日本的小鲵祖先种通过陆地连接或大陆桥在不同时间段分别进入台湾岛、中国大陆中部、朝鲜半岛一中国东北与日本本州岛东北部。综上所述,基于线粒体基因组与核基因的分子证据,本论文提出了低等真真骨鱼类系统发育关系的新观点,即该类群可分为四个谱系：1)鳞南乳鱼科占据真真骨鱼类的基部,建议提升为鳞南乳鱼目；2)水珍鱼目、鲑形目和狗鱼目鱼类一起组成单系；3)重新定义胡瓜鱼目仅包括青瓜鱼科、胡瓜鱼科、银鱼科与香鱼科,与巨口鱼目形成姊妹群关系；4)建议将南乳鱼科提升为南乳鱼目,并与新真骨鱼类(减去巨口鱼目)形成姊妹群关系。本论文亦清楚地提出胡瓜鱼目鱼类的相互关系为：(青瓜鱼科,(胡瓜鱼科,(银鱼科,香鱼科)))。研究结果极大地推进了对真真骨鱼类系统发育关系的理解,为今后真真骨鱼类的历史生物地理学研究奠定了基础。同时,本论文通过物种多样性空间格局的成因研究,取得以下两方面研究成果：1)首先指出“物种形成时间效应”与“生态位保守”在鱼类多样性海拔格局的形成过程中扮演重要角色；2)首次阐明新生代始新世到中新世东亚边缘地质构造演化在两栖类小鲵属大陆—岛屿间断分布格局形成过程中扮演了重要作用,支持台湾岛部分区域是始新世末期钓鱼岛隆起残遗的地质假说,并提出了“走出西南日本”(Out of southwestern Japan)生物地理假说,对理解日本、朝鲜半岛、与我国大陆及台湾地区之间生物的扩散提出了新观点。更多还原

【Abstract】 Teleostean fishes (Teleostei) consist of ca.96% of the 28,000 reported fish species across the world. Teleostei could be further divided into Osteoglossomorpha, Elopomorpha, Ostarioclupeomorpha (= Otocephala) and Euteleostei. Euteleostean fishes (Euteleostei) is the most species-rich fish group of Teleostei with ca.17,500 species. Euteleostei include two subdivisions, i.e., Protacanthopterygii and Neoteleostei. Protacanthopterygii, also named as lower euteleostean fishes, includes four orders (sensu Nelson,2006), i.e., Salmoniformes, Esociformes, Argentiniformes and Osmeriformes. Phylogeny of lower euteleostean fishes is one of the long-debated scientific problems in ichthyological systematics. In this thesis, phylogenetic relationships of lower euteleostean fishes were reconstructed, using mitochondrial genomes and nuclear multiloci, to resolve three problems:i) phylogenetic position of the enigmatic lower euteleostean taxa Lepidogalaxiidae; ii) phylogenetic relationships among all orders of lower euteleostean fishes; iii) phylogenetic relationships of osmeriform fishes.In addition, the processes in shaping spatial patterns of species diversity are also studied in this thesis. Two works were carried out to reveal the underlying causes for elevational patterns and disjunctive distributional patterns in species diversity:i) an endemic fish group in the Tibetan Plateau and its adjacent regions, Schizothorax, is selected as a model taxa to reveal ecological and evolutionary drivers for elevational patterns of fish diversity based on the relationships between species richness and climatic factors, area, diversification rate or time of colonization; ii) an amphibian group in the East Asian margins, Hynobius, is selected as a model taxa to reveal the role of Cenozoic tectonic evolution within East Asian margins in shaping the ’continent-islands’disjunctive distributional pattern of Hynobius.The major results are as follows:1. Phylogenetic position of Lepidogalaxiidae based on mitogenetic evidencePhylogeny of lower euteleostean fishes was reconstructed using mitochondrial genomes. The results show:i) Lepidogalaxiidae that was previously placed in Osmeriformes occupies the basal position of Euteleostei, as the sister group of all the other euteleostean fishes; ii) Galaxiidae that was previously placed in Osmeriformes forms a monophyletic group, and the erection of a new order Galaxiiformes is suggested; iii) Osmeriformes is refined to include only Retropinnidae, Osmeridae, Salangidae and Plecoglossidae.2. Phylogenetic relationships among all orders of lower euteleostean fishe based on nuclear gene evidencePhylogeny of lower euteleostean fishes was reconstructed based on ten nuclear gene fragments. The results show:i) Alepocephaloidei that was previously placed in Argentiniformes is the sister group of Ostarioclupeomorpha, thus the erection of an order Alepocephaliformes based on mitogenomic evidences is supported; ii) the opinion that Lepidogalaxiidae occupies the basal position of Euteleostei revealed from mitogenomic evidence is supported, thus the erection of a new order Lepidogalaxiiformes is suggested; iii) the monophyletic clade consisting of Argentiformes, Salmoniformes and Esociformes, and the sister taxon relationship between Osmeriformes and Stomiiformes revealed from mitogenomic evidences are supported; iv) Galaxiiformes is revealed to be the sister group of Neoteleostei (minus Stomiiformes).3. Molecular phylogeny of OsmeriformesPhylogeny of osmeriform fishes was reconstructed based on nine nuclear gene fragments. The results show:i) Retropinnidae occupies the basal position of Osmeriformes; ii) Salangidae and Plecoglossidae are a sister taxon relationship, and they together form the sister group of Osmeridae; iii) within Osmeridae, Mallotus occupies the basal position; Osmenis and Spirinchns + Allosmerus + Thaleichthys clade are a sister taxon relationship, and they together form the sister group of Hypomesus; iv) the interrelationships of Salangidae revealed by previous works of our laboratory is confirmed, that Protosalanx and Neosalangichthys form the subfamily Protosalanginae, while Salanx, Leucosoma and Salangichthys form the subfamily Salanginae; v) within Retropinnidae, Prototroctes occupies the basal position; Retropinna is found to be paraphyletic that Stokellia anisodon is revealed to be the sister species of Retropinna retropinna.4. The underlying causes for elevational patterns of Schizothorax fishes The genus Schizothorax is selected as a model taxa to reveal ecological and evolutionary drivers for elevational patterns of fish diversity. The results show:i) species richness of Schizothorax fishes peaks at mid-elevation of 1700-2200m; ii) rainfall, mid-domain effect, area and diversification rate are weak predictors of species richness; iii) temperature and time of colonization are the most important variables in explaining the elevational pattern of species richness.5. The underlying causes for disjunctive distributional patterns of East Asian Hynobius salamandersThe amphibian genus Hynobius is selected as a model taxa to reveal the role of Cenozoic tectonic evolution within East Asian margins in shaping the’continent-islands’disjunctive distributional pattern of Hynobius. The results show:i) the extant Hynobius species originated in southwestern Japan and Hokkaido Island in the Early Eocene; ii) a sister taxon relationship between Hynobius retardaius and all remaining species was the results of a vicariance event between Hokkaido Island and southwestern Japan in the Middle Eocene; iii) ancestral Hynobius in southwestern Japan dispersed into the Taiwan Island, central China,’Korean Peninsula and northeastern China’as well as northeastern Honshu during the Late Eocene-Late Miocene.In summary, novel opinions on the phylogeny of lower euteleostean fishes are presented in this thesis on the basis of molecular evidence using mitogenomes and nuclear genes:i) Lepidogalaxiidae occupies the basal position of Euteleostei, and the erection of a new order Lepidogalaxiiformes is suggested; ii) Argentiformes, Salmoniformes and Esociformes form a monophyletic clade; iii) the refined Osmeriformes including Retropinnidae, Osmeridae, Salangidae and Plecoglossidae is the sister group of Stomiiformes; iv) a newly-erected order Galaxiiformes is the sister group of Neoteleostei. Moreover, interrelationships of Osmeriformes are revealed as follows:(Retropinnidae, (Osmeridae, (Salangidae, Plecoglossidae))). The findings from this thesis greatly improve understandings on the phylogeny of euteleostean fishes, which could further aid researches on the historical biogeography of euteleostean fishes.Meanwhile, two major results have been achieved based on studies on the underlying causes for spatial patterns of species diversity:i) this thesis firstly reveals the important roles of the time-for-speciation effect and niche conservatism in shaping elevational patterns of fish diversity; ii) this thesis firstly reveal the important role of the tectonic evolution within East Asia margins during Eocene and Miocene in shaping the’continent-islands’disjunctive distributional pattern of Hynobius salamanders. A geological hypothesis that part of the present-day Taiwan Island is a relict area of the Diaoyudao Uplift in the Late Eocene is supported. A biogeographical hypothesis,’out of southwestern Japan’, is proposed here as a new opinion on explaining the dispersal of biota among Japan, Korea Peninsula, mainland China and Taiwan Island.更多还原