The intersection of paleontology and biomechanics can be reciprocally illuminating, helping to improve paleobiological knowledge of extinct species and furthering our understanding of the generality of biomechanical principles derived from study of extant species. However, working with data gleaned primarily from the fossil record has its challenges. Building on decades of prior research, we outline and critically discuss a complete workflow for biomechanical analysis of extinct species, using locomotor biomechanics in the Triassic theropod dinosaur Coelophysis as a case study. We progress from the digital capture of fossil bone morphology to creating rigged skeletal models, to reconstructing musculature and soft tissue volumes, to the development of computational musculoskeletal models, and finally to the execution of biomechanical simulations. Using a three-dimensional musculoskeletal model comprising 33 muscles, a static inverse simulation of the mid-stance of running shows that Coelophysis probably used more upright (extended) hindlimb postures and was likely capable of withstanding a vertical ground reaction force of magnitude more than 2.5 times body weight. We identify muscle force-generating capacity as a key source of uncertainty in the simulations, highlighting the need for more refined methods of estimating intrinsic muscle parameters such as fiber length. Our approach emphasizes the explicit application of quantitative techniques and physics-based principles, which helps maximize results robustness and reproducibility. Although we focus on one specific taxon and question, many of the techniques and philosophies explored here have much generality to them, so they can be applied in biomechanical investigation of other extinct organisms.

中文翻译：

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如何建造恐龙：已灭绝动物运动生物力学的肌肉骨骼建模和模拟

古生物学和生物力学的交叉可以相互启发，有助于提高对已灭绝物种的古生物学知识，并加深我们对从现存物种研究中得出的生物力学原理的一般性的理解。然而，使用主要从化石记录中收集的数据有其挑战。基于数十年的先前研究，我们概述并批判性地讨论了灭绝物种生物力学分析的完整工作流程，使用三叠纪兽脚类恐龙的运动生物力学腔骨骺作为案例研究。我们从化石骨骼形态的数字捕捉到创建装配骨骼模型，再到重建肌肉组织和软组织体积，再到开发计算肌肉骨骼模型，最后到执行生物力学模拟。使用包含 33 块肌肉的 3D 肌肉骨骼模型，对跑步中间站姿的静态反向模拟表明腔骨骺可能使用更直立（伸展）的后肢姿势，并且可能能够承受超过体重 2.5 倍的垂直地面反作用力。我们将肌肉力量产生能力确定为模拟中不确定性的关键来源，强调需要更精细的方法来估计内在肌肉参数，如纤维长度。我们的方法强调定量技术和基于物理的原理的明确应用，这有助于最大限度地提高结果的稳健性和可重复性。虽然我们专注于一个特定的分类单元和问题，但这里探讨的许多技术和哲学对它们有很多普遍性，因此它们可以应用于其他已灭绝生物的生物力学研究。