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个体发育打乱了系统发育:幼态持续与高级别的蝾螈类关系

Ontogeny discombobulates phylogeny: paedomorphosis and higher-level salamander relationships.

作者信息

Wiens John, Bonett Ronald, Chippindale Paul

机构信息

Department of Ecology and Evolution, Stony Brook University, Stony Brook, New York 11794-5245, USA.

出版信息

Syst Biol. 2005 Feb;54(1):91-110. doi: 10.1080/10635150590906037.

Abstract

Evolutionary developmental biology ("evo-devo") has revolutionized evolutionary biology but has had relatively little impact on systematics. We show that similar large-scale developmental changes in distantly related lineages can dramatically mislead phylogenetic analyses based on morphological data. Salamanders are important model systems in many fields of biology and are of special interest in that many species are paedomorphic and thus never complete metamorphosis. A recent study of higher-level salamander phylogeny placed most paedomorphic families in a single clade based on morphological data. Here, we use new molecular and morphological data to show that this result most likely was caused by the misleading effects of paedomorphosis. We also provide a well-supported estimate of higher-level salamander relationships based on combined molecular and morphological data. Many authors have suggested that paedomorphosis may be problematic in studies of salamander phylogeny, but this hypothesis has never been tested with a rigorous phylogenetic analysis. We find that the misleading effects of paedomorphosis on phylogenetic analysis go beyond the sharing of homoplastic larval traits by paedomorphic adults, and the problem therefore is not solved by simply excluding suspected paedomorphic characters. Instead, two additional factors are critically important in causing paedomorphic species to be phylogenetically "misplaced": (1) the absence of clade-specific synapomorphies that develop during metamorphosis in nonpaedomorphic taxa and allow their "correct" placement and (2) parallel adaptive changes associated with the aquatic habitat of the larval stage. Our results suggest that the effects of paedomorphosis on phylogenetic analyses may be complex, difficult to detect, and can lead to results that are both wrong and statistically well supported by parsimony and Bayesian analyses.

摘要

进化发育生物学(“演化发育生物学”)给进化生物学带来了变革,但对系统分类学的影响相对较小。我们表明,远缘谱系中类似的大规模发育变化会极大地误导基于形态学数据的系统发育分析。蝾螈是生物学许多领域的重要模式生物,特别引人关注的是,许多蝾螈物种具有幼态延续现象,因此从不完成变态发育。最近一项关于蝾螈高级系统发育的研究基于形态学数据将大多数具有幼态延续现象的科归为一个单系类群。在此,我们利用新的分子和形态学数据表明,这一结果很可能是由幼态延续现象的误导作用导致的。我们还基于分子和形态学数据的组合,给出了一个得到充分支持的蝾螈高级分类关系估计。许多作者认为幼态延续现象在蝾螈系统发育研究中可能存在问题,但这一假设从未通过严格的系统发育分析进行检验。我们发现,幼态延续现象对系统发育分析的误导作用超出了具有幼态延续现象的成体共享同塑性幼体特征的范畴,因此,仅仅排除可疑的幼态延续特征并不能解决问题。相反,在导致具有幼态延续现象的物种在系统发育上“位置错误”方面,另外两个因素至关重要:(1)在非幼态延续类群变态发育过程中形成的、能使其得到“正确”定位的类群特异性共衍征缺失;(2)与幼体阶段水生栖息地相关的平行适应性变化。我们的结果表明,幼态延续现象对系统发育分析的影响可能很复杂,难以察觉,并且可能导致在简约分析和贝叶斯分析中都得到错误但统计支持良好的结果。

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