Assessing the Adequacy of Morphological Models used in Palaeobiology

Laura P. A. Mulvey, Michael R. May, Jeremy M. Brown, Sebastian Hoehna, April M. Wright, Rachel C. M. Warnock
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Abstract

Reconstructing the evolutionary history of different groups of organisms provides insight into how life originated and diversified on Earth. Phylogenetic trees are commonly used to estimate this evolutionary history, providing a hypothesis of the events. Within Bayesian phylogenetics a major step in estimating a tree is in choosing an appropriate model of character evolution. In the case of most extinct species, our only source of information to decipher their phylogenetic relationships is through the morphology of fossils. We therefore use a model of morphological character evolution, the most common of which being the Mk Lewis model. While it is frequently used in palaeobiology, it is not known whether the simple Mk substitution model, or any extensions to it, provide a sufficiently good description of the process of morphological evolution. To determine whether or not the Mk model is appropriate for fossil data we used posterior predictive simulations, a model adequacy approach, to estimate absolute fit of the model to morphological data sets. We first investigate the impact that different versions of the Mk model have on key parameter estimates using tetrapod data sets. We show that choice of substitution model has an impact on both topology and branch lengths, highlighting the importance of model choice. Next, we use simulations to investigate the power of posterior predictive simulations for morphology. Having validated this approach we show that current variations of the Mk model are in fact performing adequately in capturing the evolutionary dynamics that generated our data. We do not find any preference for a particular model extension across multiple data sets, indicating that there is no `one size fits all' when it comes to morphological data and that careful consideration should be given to choosing models of discrete character evolution. By using suitable models of character evolution, we can increase our confidence in our phylogenetic estimates, which should in turn allow us to gain more accurate insights into the evolutionary history of both extinct and extant taxa.
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评估古生物学所用形态学模型的适当性
重建不同生物类群的进化史有助于深入了解生命如何在地球上起源和多样化。系统发生树通常用于估算这种进化历史,提供一种事件假设。在贝叶斯系统发生学中,估计系统树的一个主要步骤是选择一个合适的特征进化模型。就大多数灭绝物种而言,我们解读其系统发育关系的唯一信息来源是化石的形态。因此,我们使用形态特征演化模型,其中最常见的是 Mk Lewis 模型。虽然该模型在古生物学中经常被使用,但简单的 Mk 替换模型或其扩展模型是否能很好地描述形态演化过程还不得而知。为了确定 Mk 模型是否适合化石数据,我们使用了后验预测模拟这种模型充分性方法来估计模型与形态数据集的绝对拟合度。我们首先利用四足动物的数据集研究了不同版本的 Mk 模型对关键参数估计的影响。我们发现,替代模型的选择对拓扑结构和分支长度都有影响,这突出了模型选择的重要性。接下来,我们利用模拟来研究形态学后验预测模拟的能力。在验证了这种方法之后,我们发现 Mk 模型的当前变体在捕捉产生我们数据的进化动态方面的表现是充分的。在多个数据集中,我们没有发现对某一特定模型扩展的偏好,这表明在形态学数据方面没有 "一刀切 "的做法,在选择离散特征演化模型时应慎重考虑。通过使用合适的特征演化模型,我们可以提高系统发生学估计的可信度,进而使我们能够更准确地了解已灭绝类群和现生类群的演化历史。
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