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平鲉属(居维叶)岩鱼的起源、演化及多样化

The origin, evolution, and diversification of rockfishes of the genus Sebastes (Cuvier).

作者信息

Hyde John R, Vetter Russell D

机构信息

Scripps Institution of Oceanography, 9500 Gilman Drive, La Jolla, CA 92093-0203, USA.

出版信息

Mol Phylogenet Evol. 2007 Aug;44(2):790-811. doi: 10.1016/j.ympev.2006.12.026. Epub 2007 Jan 12.

Abstract

The evolutionary relationships of the livebearing rockfishes of the genus Sebastes have been a point of interest since their original description. With over 65 species found in the northeast Pacific (NEP), 27 in the northwest Pacific (NWP), seven in the Gulf of California (GC), four in the north Atlantic (NA) and at least two in the southern hemisphere (SH), they represent a fascinating lineage for studies of spatial and temporal patterns of dispersal, vicariance and speciation in the marine environment. Previous studies of Sebastes species have attempted to reconstruct their phylogeny using allozyme patterns or portions of a single mitochondrial gene while incompletely sampling the genus, resulting in a partial picture with low statistical support. In this study, genetic analyses using sequence data (5581 characters) from seven mitochondrial genes (cytochrome b, cytochrome c oxidase subunit 1, 12S rRNA, 16S rRNA, tRNA proline, tRNA threonine and the control region) and two nuclear genes (recombination activating gene 2 and internal transcribed spacer 1), along with a near complete sampling of species, have produced a well supported phylogenetic hypothesis of the relationships between Sebastes species as well as clarifying their position within the scorpaenid subfamily, Sebastinae. Though studies of similar magnitude have been conducted at the family and subfamily level, this represents the most detailed and extensive examination of biogeography and marine speciation within a single, widely distributed marine fish genus. Both Bayesian posterior and maximum parsimony analyses produced highly similar phylogenies suggesting an origin for Sebastes at high-latitudes in the NWP. The majority of previously proposed sub-generic groupings based upon morphology are found to be either para- or polyphletic. Using Bayesian-derived genetic distance measures together with rate smoothing techniques, a molecular clock was applied to the phylogeny. The clock-calibrated data suggest that Sebastes originated in the middle Miocene, concordant with fossil data, and began substantial diversification and dispersal in synchrony with high-latitude cooling and establishment of productive upwelling systems across the north Pacific (NP) in the late Miocene. Contrary to contemporary taxonomic criteria that often group Asian and North American species based on common morphology, the molecular phylogenies tend to indicate geographically circumscribed lineages with no evidence for repeated long distance dispersal between disjunct biogeographic provinces (e.g., Asian species nested within a North American lineage). No examples of large-scale glacial vicariance as would be suggested by Asian and North American sibling species were observed. To the contrary, sibling species tended to be in geographic proximity. While occasional long distance dispersal may occur, such as the single colonization of the SH, and thermal barriers presently exist between the NP, NA, GC, and SH taxa, the observable patterns in Sebastes suggest colonization occurs by stepwise invasion of newly available habitat when temperature conditions permit. Colonization events are spread throughout the sub-generic lineages. Vicariant isolation processes may occur on smaller geographic scales perhaps due to local isolating mechanisms such as glacial advance and retreat, sea level change, and ocean currents. Allopatric differences may be enhanced by a tendency for female mate choice and assortative mating in these livebearing species. The ongoing process of thermal advance and retreat is reflected in contemporary patterns of phylogeographic population genetic structure within species and may be enhanced under climate warming.

摘要

自首次描述以来,平鲉属胎生岩鱼的进化关系一直是人们关注的焦点。在东北太平洋(NEP)发现了65种以上,西北太平洋(NWP)有27种,加利福尼亚湾(GC)有7种,北大西洋(NA)有4种,南半球(SH)至少有2种,它们代表了一个引人入胜的谱系,可用于研究海洋环境中扩散、隔离分化和物种形成的时空模式。先前对平鲉属物种的研究试图利用等位酶模式或单个线粒体基因的部分片段来重建它们的系统发育,但对该属的采样并不完整,导致得到的是一幅不完整且统计支持度较低的图景。在本研究中,利用来自7个线粒体基因(细胞色素b、细胞色素c氧化酶亚基1、12S rRNA、16S rRNA、脯氨酸转运RNA、苏氨酸转运RNA和控制区)和2个核基因(重组激活基因2和内部转录间隔区1)的序列数据(5581个字符)进行遗传分析,并对物种进行了近乎完整的采样,得出了一个得到充分支持的关于平鲉属物种间关系的系统发育假说,同时也明确了它们在鲉科平鲉亚科中的位置。尽管在科和亚科层面已经进行了类似规模的研究,但这是对单个广泛分布的海洋鱼类属内生物地理学和海洋物种形成最详细、最广泛的研究。贝叶斯后验分析和最大简约法分析都产生了高度相似的系统发育树,表明平鲉起源于西北太平洋的高纬度地区。大多数先前基于形态学提出的亚属分类群被发现是并系或多系的。利用贝叶斯推导的遗传距离测量方法并结合速率平滑技术,将分子钟应用于系统发育树。经分子钟校准的数据表明,平鲉起源于中新世中期,与化石数据一致,并在中新世晚期随着北太平洋(NP)高纬度地区的降温以及整个北太平洋生产力上升流系统的建立而开始大量分化和扩散。与当代分类标准通常根据共同形态将亚洲和北美物种归为一类不同,分子系统发育往往表明地理上受限的谱系,没有证据表明在不连续的生物地理省份之间存在反复的长距离扩散(例如,亚洲物种嵌套在北美谱系中)。没有观察到亚洲和北美近缘物种所暗示的大规模冰川隔离分化的例子。相反,近缘物种往往在地理上相邻。虽然偶尔可能会发生长距离扩散,比如南半球的单次定殖,并且目前在北太平洋、北大西洋、加利福尼亚湾和南半球类群之间存在热障,但平鲉中可观察到的模式表明,当温度条件允许时,定殖是通过逐步侵入新的可用栖息地发生的。定殖事件分布在整个亚属谱系中。隔离分化过程可能发生在较小的地理尺度上,这可能是由于局部隔离机制,如冰川进退、海平面变化和洋流。在这些胎生物种中,雌性配偶选择和选型交配的倾向可能会增强异域差异。热进退的持续过程反映在物种内当代的系统发育种群遗传结构模式中,并且在气候变暖的情况下可能会增强。

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