四足动物起源于以脊索为主导的轴柱结构的肉鳍鱼类1–3。在这一过渡过程中，头骨与肩带分离4–6，形成了颈部区域。这种头-肩分离现象引发了假说，认为在四足动物起源期间颈部活动性增加，从而促进了向更陆生栖息地的生态转变4,5,7–9。然而，椎体成分的不完全骨化10给这些早期四足动物颈部活动性的建模带来了挑战。

Tetrapods arose from lobe-finned fishes with a notochord-dominated axial column1–3. During this transition, the skull became separated from the shoulder girdle4–6, giving rise to a neck region. This head–shoulder separation has generated hypotheses suggesting that neck mobility increased during the origin of tetrapods, facilitating the ecological transition towards more terrestrial habitats4,5,7–9. However, incompletely co-ossified vertebral components10 have posed a challenge to modelling the mobility of these early tetrapod necks.