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题名	鮡科及纹胸鮡属鱼类分类、系统进化与生物地理学研究
作者	蒋万胜
学位类别	博士
答辩日期	2012-06
授予单位	中国科学院研究生院
授予地点	北京
导师	杨君兴,陈小勇
关键词	鮡科 纹胸鮡属 系统分类学 系统发育 生物地理学
其他题名	Systematics, phylogenetics and biogeography of catfishes in Sisoridae and Glyptothorax (Teleostei: Siluriformes)
学位专业	动物学
中文摘要	鮡科 (Sisoridae) 是亚洲鲇形目 (Siluriformes) 鱼类主要分布在山川河流中的小型鱼类代表类群。有200多种物种分布在西起两河流域的幼发拉底河和底格里斯河，东至中国的长江及其以南的沿海省份，南至大巽他群岛，和这其中包括的南亚次大陆、东南亚和中国南部的广大地区。本研究通过广泛采样和合作，从分子的角度探讨了鮡科鱼类的系统发育关系和生物地理学演化过程。结果显示：(1)、在广泛包含鮡科几乎所有属的情况下，鮡科鱼类为一单系。原来具有争议的骨鮡科 (Erethistidae) 从属于鮡科下的一个分支。结合前人基于形态学的研究结果，骨鮡科的科级地位应当废除，而降为鮡科下的骨鮡族Erethistini。(2)、鮡科鱼类包含三大分支，分别是Clade A，Clade B和Clade C。首先Clade A+Clade B与Clade C形成姐妹群，然后发生Clade A与Clade B的分化。Clade A包含真鮡族Sisorini (Gagata, Nangra, Gogangra, Sisor) 和骨鮡族Erethistini (Hara, Ayarnangra, Caelatoglanis, Erethistoides, Erethistes, Pseudolaguvia, Conta)。Clade B包含纹胸鮡族Glyptothoracini (Glyptothorax) 和魾族Bagariini (Bagarius)。Clade C包含褶鮡族Pseudecheneidini (Pseudecheneis)和鰋鮡族Glyptosternini (Oreoglanis, Creteuchiloglanis, Pseudexostoma, Pareuchiloglanis, Euchiloglanis, Exostoma, Glyptosternon, Glaridoglanis, Myersglanis)。(3)、生物地理学研究结果表明：鮡科鱼类约起源于中新世晚期约11.38 Ma的东喜马拉雅区域 (恒河-布拉马普特拉河-伊洛瓦底江)，在较短的时间内分化成为Clade A+Clade B与Clade C两大支系。这两个支系的分化对应着青藏高原第三次快速隆升的阶段 (13-8 Ma)，高原隆升伴随着河流的深切和峡谷的形成，导致环境剧烈改变，从而分化出较为适应激流生境的Clade C。Clade C又进一步分化为褶鮡族和高度适应激流生境的鰋鮡族 (鰋鮡鱼类)。Clade A和Clade B在9.2 Ma分化，以及Clade C中的凿齿鮡Glaridoglanis、原鮡Glyptosternon和鰋Exostoma与其他鰋鮡鱼类在9.14 Ma和8.39 Ma分化可能对应着高原隆升水系袭夺导致的祖先区域的东西隔离和随后的扩散。Clade A中真鮡族和骨鮡族的分化跟印度河袭夺恒河上游的部分水系有关，从而使部分类群得以向印度河及其以西的水系扩散。Clade B分化为广泛而且同域分布纹胸鮡族和魾族可能是由于生态位分化的结果。Clade C中金沙江水系的种类与其他种类隔离，支持长江袭夺原本作为古红河支流金沙江上游的假说，这种袭夺大概发生在4.82 Ma。 纹胸鮡属 (Glyptothorax) 是鮡科里面种类最多，分布最广的类群。除青藏高原部分高海拔区域外，鮡科鱼类的分布边缘，皆由纹胸鮡属占据。至今约有有效种类80余种。同样通过广泛的样品收集，探讨了纹胸鮡属的单系性和系统发育关系。结果表明：(1)、在以鮡科其他属特别是同样具有胸附着器的其他类群作为外群的情况下，纹胸鮡属形成一单系，与魾属互为姐妹群。胸附着器在纹胸鮡属内具有同源性。(2)、在包含约三分之二的物种情况下，纹胸鮡属形成9大支系。其中基部类群Clade A由胸附着器不甚发达和善游的种类组成，这意味着原始类群对底栖附着生活的依赖性不强。(3)、对各支系的具体情况进行了详细的探讨。整体而言，各支系与水系之间没有呈现出良好的对应关系，这说明该类群有着较为久远的进化历史。但个别分支的类群具有相似的体色而形成一定的单系结构，这可能与其分布区域水系的最终隔离事件有关。(4)、胸附着器在物种鉴定方面展现出良好的应用价值，但在系统发育关系上指示意义不大，而表现出较大程度的环境可塑性。(5)、部分类群，特别是跨国境分布和跨水系分布的广布种的分类地位亟需要进一步的系统整理。 对扎那纹胸鮡G. zanaensis和以往被认为是扎那纹胸鮡的种类进行了形态和分子两方面的系统研究。重新整理和描述了扎那纹胸鮡，并限定其仅分布于怒江上游 (从西藏昌都到云南泸水段)。重新恢复了长须纹胸鮡G. longinema的有效种地位，其代表原来分布在怒江扎那纹胸鮡的部分种群和澜沧江的扎那纹胸鮡。并描述了怒江水系的两个新种——粒线纹胸鮡Glyptothorax granosus和异色纹胸鮡Glyptothorax fucatus。对这四个物种分别进行了详细的描述和比较。 对中华纹胸鮡G. sinensis及其姐妹种进行了谱系地理学研究。首先，分子系统发育结果支持福建纹胸鮡G. fokiensis应为中华纹胸鮡的同物异名。四个研究物种均有着较为久远的进化历史，同时也具有较高的遗传多样性。通过澜沧江水系的大斑纹胸鮡G. macromaculatus的最先分化，推测澜沧江水系与红河水系隔离的时间大概在上新世3.3-4.15 Ma。通过红河水系红河纹胸鮡G. honghensis的进一步分化，表明红河水系与其以东的水系隔离时间约在3.02-3.82 Ma。其中李仙江和元江长时间内都有着广泛的交流，直到0.28-0.63 Ma左右才发生隔离，当前下游之间的水系连通很难造成被隔离的上游种群之间的基因交流。广西沿海独立水系与海南岛之间关系较近，十万大山和云开山脉有效地隔离了海南纹胸鮡G. hainanensis和中华纹胸鮡之间的交流，这种隔离发生在2.39-3.05 Ma；中更新世以来琼州海峡陆桥连接是海南种群与大陆种群交流的主要渠道。中华纹胸鮡的谱系地理学研究显示：虽然长江上游在早上新世即可能与古红河发生隔离，但此时长江并没有全线贯通，而是通过一些南北流向的河流汇入西江，从西江入海，长江和西江鱼类之间有着长时间的交流和共通。直到约1.7-2.23 Ma，随着云贵高原隆升导致一系列山脉的形成，阻断了这种交流后，长江也即深切三峡与下游连接，从此全线贯通，流入东海。之前作为南北通道的河流继而分别变成长江和西江的支流。长江下游省份与广东和福建之间由于地势较为低平，鱼类之间也一直有着广泛的联系。后来由于南岭和武夷山的升高有效地阻隔了这种联系，而珠江的三条水系之间鱼类的交流从此增多。但这种交流可能并不太适合山川激流起源的纹胸鮡种类，以至于西江与北江和东江之间没有发生二次接触的痕迹。 对怒江水系纹胸鮡属鱼类的整理过程中，先后发现两个新种。其中一种胸附着器向前延伸至鳃颊部以及颏部 (第一次记录到我国有这种特殊胸附着器的种类)，和身体具有不规则的斜向排列的斑块，命名为斜斑纹胸鮡—Glyptothorax obliquimaculatus；另一种具有特殊的皮肤颗粒，和似长矛矛头的胸附着器，命名为矛形纹胸鮡—Glyptothorax lanceatus。综合跨境分布的种类的形态特征，给出了整个萨尔温江水系纹胸鮡属鱼类的分类检索表。
英文摘要	The catfish family Sisoridae is the largest Asian catfish family in Siluriformes with more than 200 species. Most of them are small-sized catfishes and preferring to inhabit hill-streams or fast-flowing large rivers. There are widely distributed in throughout the entire southern arc of the Asia continent, from Asia Minor (in the Tigris and Euphrates River drainages) eastward to East Asia (in the Yangtze River drainage) and southward to Southeast Asia (in the Greater Sunda Islands), including the broad area of Indian subcontinent, Southeast Asia and South of China. We studied the molecular phylogenetic relationships and biogeography of Sisoridae by broad sampling that ensuring at least one representive in each genus. The results showed that: (1), The Sisoridae was a monophyletic group only when including both the previous Erethistidae and other Sisorid fishes. It supported the previous morphological result that Sisoridae should be expanded to include members previously placed in the family Erethistidae. The previous family Erethistidae was downgraded to a tribe (Erethistini) in the family Sisoridae. (2), Sisoridae was recovered as three major subclades as Clade A，Clade B and Clade C. Clade A grouped with Clade B and both together as the sister group of Clade C. Clade A contained Sisorini (Gagata, Nangra, Gogangra, Sisor) and Erethistini (Hara, Ayarnangra, Caelatoglanis, Erethistoides, Erethistes, Pseudolaguvia, Conta). Clade B contained Glyptothoracini (Glyptothorax) and Bagariini (Bagarius). Clade C contained Pseudecheneidini (Pseudecheneis) and Glyptosternini (Oreoglanis, Creteuchiloglanis, Pseudexostoma, Pareuchiloglanis, Euchiloglanis, Exostoma, Glyptosternon, Glaridoglanis, Myersglanis). (3), Biogeography analysis suggested that Sisoridae was possibly originated from east Himalaya (Ganges-Brahmaputra-Irrawaddy) ca. 11.38 Ma in the late Miocene, and then divided into two lineages as Clade A+Clade B and Clade C. This division was corresponding to the third rapid uplift of Qinghai-Tibet Plateau (13-8 Ma), following the rapid river incision and gorge formation, which pushed forward the habitat changes and promoted the evolution of torrent adaptive group Clade C. And then the Clade C was evolved into a group of Glyptosternini (Glyptosternoid fishes) with high adaption to the torrent habitat. The division of Clade A and Clade B (ca. 9.2 Ma), as well as the Clade C members Glaridoglanis, Glyptosternon and Exostoma divided from the remainders (9.14 Ma and 8.39 Ma) was possibly triggered by large-scale drainage captures and changes, which leaded the ancestor area west-east isolation and then dispersal. The division of Sisorini and Erethistini in Clade A was related to the Indus captured the upper Ganges and lead into the dispersal of some Sisorini to Indus and then to farther west. Both the two tribes Bagariini and Glyptothoracini in Clade B were wide and in a way of sympatric distributions. The division of these two tribes was likely caused by the ecological niche separation. The species in Jinshajiang drainages separated from others in Clade C supported the hypothesis of middle Yangtze River captured the upper Yangtze which was original one of the upper reaches of paleo-Red River. We estimated this capture might have happened at ca. 4.82 Ma. Glyptothroax is the most species-diverse genus and widely distributed group in Sisoridae, with more than 80 valid species spanning in the whole distribution area of Sisoridae except some high altitude reaches in Qinghai-Tibet plateau. We used the DNA sequences of almost two-thirds of its known species to examine monophyly of Glyptothorax and phylogenetic relationships with the genus. The results showed that: (1), The monophyly of Glyptothorax was strongly supported when the other genera in Sisoridae were treated as outgroups. The presence of thoracic adhesive apparatus in Glyptothorax was confirmed as homology, even though it is not unique within the Sisoridae. Bagarius was recovered as the sister group of Glyptothorax. (2), We recovered nine well supported subclades (labeled as Clade A-I). The basic subclade Clade A included species with fusiform bodies and less strongly-developed thoracic adhesive apparatus which implied that the ancestor of Glyptothorax was less benthic than its current descendant. (3), We discussed each subclades in detail. Generally speaking, subclades were not corresponding with the current drainages pattern, which implied a long and complex evolution progress of these fishes. However, some geographic separated cluster of subclades present possible synapomorphy by body color and plaque hinted us that they might be speciated in a phase related to the current drainages finally formation. (4), The thoracic adhesive apparatus is useful in diagnosing species but not in elucidating phylogenetic relationships, which is an environmental plastic characteristic in its morphological evolution for the most part. (5), The current taxonomy of some species, especially these cross-countries and trans-drainages required reexamination. We re-examined the taxonomic status of the Glyptothorax catfishes usually identified as G. zanaensis using a combination of morphometric and molecular data. Our study restricted the distribution of G. zanaensis to the upper Salween River (Nujiang) in Tibet and Yunnan, and revalidated G. longinema for populations from the upper Salween (Nujiang) and upper Mekong (Lancanjiang) drainages in Yunnan previously identified as G. zanaensis. We also described two previously unnamed species as G. granosus and G. fucatus in Nujiang drainages. All four species were diagnosed and described in detail. The phylogeography of G. sinensis and its sister species was studied using mitochondrial DNA Cyt b gene sequences. Phylogenetic analysis confirmed that the G. fokiensis should be junior synonym of G. sinensis. The four species involved into this study divided and then evolved respectively at long time ago, and all of them with high genetic diversity. Glyptothorax macromaculatus divided first and it inferred that Lancangjiang drainages separated from Red River drainages would happen at ca. 3.3-4.15 Ma in the Pliocene. Glyptothorax honghensis divided from the remainders which inferred that the Red River drainages separated from its eastern river at ca. 3.02-3.82 Ma in the middle Pliocene. The fish populations in Lixianjiang River and Yuanjiang River could free communicate until the final geographic isolation happened at ca. 0.28-0.63 Ma. Although the lower reaches of these two drainages are linking at present that couldn’t provide effective access between the populations in the isolated upper reaches. The Shiwandashan Mountain and Yunkai Mountain served as the geographic isolation of G. hainanensis and G. sinensis at ca. 2.39-3.05 Ma. The close relationships of fishes between South Guangxi and Hainan Island would be arisen from free communications by the frequent sea regressions in Pleistocene ice age. However, in populations of G. hainanensis, the foremost communication was taken place at the land bridge on Qiongzhou strait. Phylogeography of G. sinensis showed that: Although upper Yangtze reaches diverted to middle Yangtze from paleo-Red River might be happened in early Pliocene, it would not connect the lower reaches immediately but flow to Xijiang River (West Pearl River) and then to South China Sea. Fishes of Yangtze River and Xijiang River could free communicate through these North-South channels in a very long time. These communications would be interrupted by the orogenesis following the uplift of Yunnan-Guizhou Pleatau. It became the impetus of Yangtze run across the Three Gorges and then link up the lower reaches flowing to the East China Sea. The North-South channels thus became the different tributaries of Yangtze and Xijiang River. The low plain crossing Lower Yangtze River basin and Southeast China (Guangdong and Fujian province etc.) at that time would provide broad communications of fishes in these areas. Subsequent uplift of Nanling Mountain and Wuyishan Mountain interrupted these connections, and then the communications among three major tributaries of Pearl River (Xijiang, Beijiang and Dongjiang River) increased afterward. However, the communications between Xijiang River and the other two tributaries (Beijiang and Dongjiang River) would not be fit for hill-streams originated species G. sinensis because there was no sign of second contact of these populations. When the systematic review to Glyptothorax in Nujiang drainages carried out in these years, two new species were described. One with unculiferous ridges of thoracic adhesive apparatus extending anteriorly onto the gular region easily distinguished from all the other congeners (without unculiferous ridges on gular region) in China, which was named as G. obliquimaculatus in having irregular oblique blotches along lateral surface. The other one distinguished from other congeners in having unusual elongate, ovoid tubercles on dorsal head and narrow thoracic adhesive apparatus like the tip of a lance, which was named as G. lanceatus. The key to Glyptothorax in the Salween River drainages was also given in this thesis.
语种	中文
公开日期	2013-04-25
内容类型	学位论文
源URL	[http://159.226.149.42:8088/handle/152453/7408]
专题	昆明动物研究所_系统进化与生物地理学 昆明动物研究所_东南亚野生动物多样性
推荐引用方式 GB/T 7714	蒋万胜. 鮡科及纹胸鮡属鱼类分类、系统进化与生物地理学研究[D]. 北京. 中国科学院研究生院. 2012.

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