The kinetic skull is a key innovation that allowed snakes to capture, manipulate, and swallow prey exclusively using their heads using the coordinated movement of eight bones. Despite these unique feeding behaviors, patterns of evolutionary integration and modularity within the feeding bones of snakes in a phylogenetic framework have yet to be addressed. Here, we use a dataset of 60 μCT‐scanned skulls and high‐density geometric morphometric methods to address the origin and patterns of variation and integration in the feeding bones of aquatic‐foraging snakes. By comparing alternate superimposition protocols allowing us to analyze the entire kinetic feeding system simultaneously, we find that the feeding bones are highly integrated, driven predominantly by functional selective pressures. The most supported pattern of modularity contains four modules, each associated with distinct functional roles: the mandible, the palatopterygoid arch, the maxilla, and the suspensorium. Further, the morphological disparity of each bone is not linked to its magnitude of integration, indicating that integration within the feeding system does not strongly constrain morphological evolution, and that adequate biomechanical solutions to a wide range of feeding ecologies and behaviors are readily evolvable within the constraint due to integration in the snake feeding system.

中文翻译：

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水生觅食蛇多动进食系统的形态整合和模块化

动态头骨是一项关键创新，它使蛇能够通过八块骨头的协调运动完全使用头部捕捉、操纵和吞食猎物。尽管有这些独特的摄食行为，但系统发育框架中蛇的摄食骨骼内的进化整合和模块化模式尚未得到解决。在这里，我们使用 60 μCT 扫描的头骨和高密度几何形态测量方法的数据集来解决水生觅食蛇的摄食骨骼的变化和整合的起源和模式。通过比较替代叠加协议，我们可以同时分析整个动态喂养系统，我们发现喂养骨骼是高度集成的，主要由功能选择压力驱动。最受支持的模块化模式包含四个模块，每个模块都与不同的功能角色相关联：下颌骨、腭弓、上颌骨和悬骨。此外，每块骨骼的形态差异与其整合程度无关，这表明喂养系统内的整合不会强烈限制形态进化，并且针对广泛的喂养生态和行为的适当生物力学解决方案很容易在由于集成在蛇饲养系统中的限制。