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形态系统发育中的趋同与噪声镶嵌:类似灵猫科的食肉动物包含了什么?

Mosaics of convergences and noise in morphological phylogenies: what's in a viverrid-like carnivoran?

作者信息

Gaubert Philippe, Wozencraft W Chris, Cordeiro-Estrela Pedro, Veron Géraldine

机构信息

Unité Origine, Structure et Evolution de la Biodiversité, CNRS UMR 5202, Département Systématique et Evolution, Muséum National d'Histoire Naturelle, Zoologie: Mammifères et Oiseaux, CP 51, 57 rue Cuvier, 75231 Paris Cedex 05, France.

出版信息

Syst Biol. 2005 Dec;54(6):865-94. doi: 10.1080/10635150500232769.

Abstract

Adaptive convergence in morphological characters has not been thoroughly investigated, and the processes by which phylogenetic relationships may be misled by morphological convergence remains unclear. We undertook a case study on the morphological evolution of viverrid-like feliformians (Nandinia, Cryptoprocta, Fossa, Eupleres, Prionodon) and built the largest morphological matrix concerning the suborder Feliformia to date. A total of 349 characters grouped into four anatomical partitions were used for all species of Viverridae and viverrid-like taxa plus representatives of the Felidae, Hyaenidae, Herpestidae, and one Malagasy mongoose. Recent molecular phylogenetic analyses suggest that viverrid-like morphotypes appeared independently at least three times during feliformian evolution. We thus used a synthetic molecular tree to assess morphological evolutionary patterns characterizing the viverrid-like taxa. We examined phylogenetic signal, convergence and noise in morphological characters using (a) tree-length distribution (g1), (b) partitioned Bremer support, (c) RI values and their distribution, (d) respective contributions of diagnostic synapomorphies at the nodes for each partition, (e) patterns of shared convergences among viverrid-like taxa and other feliformian lineages, (f) tree-length differences among alternative hypotheses, and (g) the successive removal of convergent character states from the original matrix. In addition, the lability of complex morphological structures was assessed by mapping them onto the synthetic molecular tree. The unconstrained morphological analysis yielded phylogenetic groupings that closely reflected traditional classification. The use of a synthetic molecular tree (constraint) combined with our thorough morphological investigations revealed the mosaics of convergences likely to have contributed to part of the historical uncertainty over viverrid classification. It also showed that complex morphological structures could be subjected to reversible evolutionary trends. The morphological matrix proved useful in characterizing several feliformian clades with diagnostic synapomorphies. These results support the removal from the traditionally held Viverridae of several viverrid-like taxa into three distinct families: Nandiniidae (Nandinia), Prionodontidae (Prionodon), and the newly defined Eupleridae (including Cryptoprocta, Fossa, Eupleres plus all "mongoose-like" Malagasy taxa). No clearly "phylogenetically misleading" data subsets could be identified, and the great majority of morphological convergences appeared to be nonadaptive. The multiple approaches used in this study revealed that the most disruptive element with regards to morphological phylogenetic reconstruction was noise, which blured the expression of phylogenetic signal. This study demonstrates the crucial need to consider independent (molecular) phylogenies in order to produce reliable evolutionary hypotheses and should promote a new approach to the definition of morphological characters in mammals. [Constrained analysis; convergence; evolutionary scenario; Feliformia; morphology; noise; phylogenetic signal; phylogeny; Viverridae.].

摘要

形态特征的适应性趋同尚未得到充分研究,形态趋同可能误导系统发育关系的过程仍不清楚。我们对类灵猫科猫型亚目动物(南氏灵猫、隐肛狸、马岛獴、窄纹獴、缟灵猫)的形态进化进行了案例研究,并构建了迄今为止关于猫型亚目的最大形态矩阵。总共349个特征被分为四个解剖部分,用于所有灵猫科物种、类灵猫类群以及猫科、鬣狗科、獴科的代表物种和一种马达加斯加獴。最近的分子系统发育分析表明,类灵猫形态型在猫型亚目进化过程中至少独立出现了三次。因此,我们使用一个综合分子树来评估表征类灵猫类群的形态进化模式。我们使用以下方法检查形态特征中的系统发育信号、趋同和噪声:(a)树长分布(g1),(b)分区布雷默支持率,(c)RI值及其分布,(d)每个分区节点处诊断共衍征的各自贡献,(e)类灵猫类群与其他猫型亚目谱系之间的共享趋同模式,(f)替代假设之间的树长差异,以及(g)从原始矩阵中连续去除趋同特征状态。此外,通过将复杂形态结构映射到综合分子树上,评估了它们的不稳定性。无约束形态分析产生的系统发育分组与传统分类密切相关。使用综合分子树(约束)并结合我们全面的形态学研究,揭示了趋同镶嵌现象,这可能是造成灵猫科分类历史不确定性的部分原因。研究还表明,复杂形态结构可能经历可逆的进化趋势。形态矩阵被证明有助于用诊断共衍征表征几个猫型亚目分支。这些结果支持将几个类灵猫类群从传统的灵猫科中分离出来,归入三个不同的科:南氏灵猫科(南氏灵猫)、缟灵猫科(缟灵猫)和新定义的马岛獴科(包括隐肛狸、马岛獴、窄纹獴以及所有“獴样”马达加斯加类群)。未发现明显“误导系统发育”的数据子集,并且绝大多数形态趋同似乎是非适应性的。本研究中使用的多种方法表明,在形态系统发育重建方面,最具干扰性的因素是噪声,它模糊了系统发育信号的表达。这项研究表明,为了得出可靠的进化假设,迫切需要考虑独立的(分子)系统发育,并且应该推动一种新的方法来定义哺乳动物的形态特征。[约束分析;趋同;进化情景;猫型亚目;形态学;噪声;系统发育信号;系统发育;灵猫科。]

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