The scapula shares developmental and functional relationships with traits of the basicranium, vertebral column, humerus, and clavicle. As a limb girdle, it also shares analogous characteristics with the pelvis. Despite these relationships, studies of primate shoulder evolution often focus on traits of the scapula in isolation. Such analyses may lead to spurious conclusions, as they implicitly model the scapula as evolving independent of other anatomical regions. Traits of the shoulder girdle share genetic covariances with each other, as well as potential covariances with dimensions of other skeletal elements. To create accurate models of shoulder evolution, it is imperative to account for the constraints imposed by these sources of covariance. Here, we use evolutionary quantitative methods to test a model in which shoulder morphological evolution is influenced by its developmental and functional covariances with the basicranium in the Colobus genus. This evolutionary relationship is also assessed with morphology of the pelvis to provide context to the evolutionary covariance among traits of the basicranium and shoulder girdle. Our results indicate potential evolutionary implications arising from covariances among the basicranium, shoulder, and pelvis. We further propose that the shoulder and basicranium may be examples of developmental, functional, and genetic covariances among traits that manifest an evolutionary suite of mutually constrained morphologies. We demonstrate novel evolutionary relationships among the shoulder girdle and basicranium that affect not only models of primate shoulder evolution but have broader implications for modeling trait evolution across the skeleton.

中文翻译：

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灵长类动物基础，肩部和臀部的可进化性和约束性以及多性状进化的重要性

肩cap骨与碱性肌层，椎骨，肱骨和锁骨的性状共享发育和功能关系。作为肢带，它也具有与骨盆相似的特征。尽管有这些关系，但是关于灵长类动物肩部进化的研究通常只关注孤立的肩the骨特征。这样的分析可能导致虚假结论，因为它们隐式地将肩cap骨建模为独立于其他解剖区域的演变。肩带的性状彼此之间具有遗传协方差，也与其他骨骼元素的尺寸具有潜在的协方差。要创建肩部进化的准确模型，必须考虑这些协方差源所施加的约束。这里，我们使用进化定量方法来测试一个模型，在该模型中，肩部形态演化受其与Colobus属中的碱性花的发育和功能协方差的影响。还用骨盆的形态评估了这种进化关系，以提供基础花兰和肩带特征之间进化协方差的背景。我们的结果表明，潜在的进化影响是由基础胎层，肩膀和骨盆之间的协方差引起的。我们进一步提出，肩和碱土可能是性状之间发展，功能和遗传协方差的例子，这些性状表现出相互制约的形态演化套件。