Department of Musculoskeletal Biology, Institute of Aging and Chronic Disease, University of Liverpool, Liverpool, UK.
J Anat. 2012 Mar;220(3):212-32. doi: 10.1111/j.1469-7580.2011.01469.x. Epub 2012 Jan 3.
Ornithischia (the 'bird-hipped' dinosaurs) encompasses bipedal, facultative quadrupedal and quadrupedal taxa. Primitive ornithischians were small bipeds, but large body size and obligate quadrupedality evolved independently in all major ornithischian lineages. Numerous pelvic and hind limb features distinguish ornithischians from the majority of other non-avian dinosaurs. However, some of these features, notably a retroverted pubis and elongate iliac preacetabular process, appeared convergently in maniraptoran theropods, and were inherited by their avian descendants. During maniraptoran/avian evolution these pelvic modifications led to significant changes in the functions of associated muscles, involving alterations to the moment arms and the activation patterns of pelvic musculature. However, the functions of these features in ornithischians and their influence on locomotion have not been tested and remain poorly understood. Here, we provide quantitative tests of bipedal ornithischian muscle function using computational modelling to estimate 3D hind limb moment arms for the most complete basal ornithischian, Lesothosaurus diagnosticus. This approach enables sensitivity analyses to be carried out to explore the effects of uncertainties in muscle reconstructions of extinct taxa, and allows direct comparisons to be made with similarly constructed models of other bipedal dinosaurs. This analysis supports some previously proposed qualitative inferences of muscle function in basal ornithischians. However, more importantly, this work highlights ambiguities in the roles of certain muscles, notably those inserting close to the hip joint. Comparative analysis reveals that moment arm polarities and magnitudes in Lesothosaurus, basal tetanuran theropods and the extant ostrich are generally similar. However, several key differences are identified, most significantly in comparisons between the moment arms of muscles associated with convergent osteological features in ornithischians and birds. Craniad migration of the iliofemoralis group muscles in birds correlates with increased leverage and use of medial femoral rotation to counter stance phase adduction moments at the hip. In Lesothosaurus the iliofemoralis group maintains significantly higher moment arms for abduction, consistent with the hip abduction mode of lateral limb support hypothesized for basal dinosaurs. Sensitivity analysis highlights ambiguity in the role of musculature associated with the retroverted pubis (puboischiofemoralis externus group) in ornithischians. However, it seems likely that this musculature may have predominantly functioned similarly to homologous muscles in extant birds, activating during the swing phase to adduct the lower limb through lateral rotation of the femur. Overall the results suggest that locomotor muscle leverage in Lesothosaurus (and by inference basal ornithischians in general) was more similar to that of other non-avian dinosaurs than the ostrich, representing what was probably the basal dinosaur condition. This work thereby contradicts previous hypotheses of ornithischian-bird functional convergence.
鸟臀目(“有鸟类臀部的恐龙”)包含双足、 facultative 四足和四足类群。原始的鸟臀目恐龙是小型双足动物,但大型身体大小和强制性四足性在所有主要的鸟臀目谱系中独立进化。许多骨盆和后肢特征将鸟臀目与大多数其他非鸟类恐龙区分开来。然而,其中一些特征,特别是反向耻骨和延长的髂骨前髋臼突,在手盗龙类兽脚亚目中趋同出现,并被它们的鸟类后代继承。在手盗龙类/鸟类进化过程中,这些骨盆的变化导致与相关肌肉功能的显著变化,涉及到对力矩臂和骨盆肌肉激活模式的改变。然而,这些特征在鸟臀目动物中的功能及其对运动的影响尚未经过测试,仍然知之甚少。在这里,我们使用计算模型对最完整的基础鸟臀目动物——莱索托龙(Lesothosaurus diagnosticus)进行了双足鸟臀目动物肌肉功能的定量测试,以估计 3D 后肢力矩臂。这种方法可以进行敏感性分析,以探索对灭绝分类群肌肉重建的不确定性的影响,并允许与其他双足恐龙的类似构建模型进行直接比较。这项分析支持了一些以前提出的关于基础鸟臀目动物肌肉功能的定性推论。然而,更重要的是,这项工作强调了某些肌肉的作用的模糊性,特别是那些靠近髋关节插入的肌肉。比较分析表明,莱索托龙、基础坚尾龙类兽脚亚目恐龙和现生鸵鸟的后肢力矩臂极性和大小通常相似。然而,确定了几个关键差异,最重要的是在与鸟臀目动物和鸟类中趋同的骨骼特征相关的肌肉的力矩臂之间的比较。鸟类中腰股骨群肌肉的头侧迁移与增加杠杆作用和利用股骨内侧旋转来抵消髋关节在支撑阶段的内收力矩有关。在莱索托龙中,腰股骨群保持着更高的外展力矩臂,这与对基础恐龙假设的侧肢支撑的髋关节外展模式一致。敏感性分析强调了在鸟臀目动物中与反向耻骨(耻骨坐骨股骨外侧群)相关的肌肉作用的模糊性。然而,这种肌肉很可能主要与现生鸟类中的同源肌肉类似,在摆动阶段通过股骨的外侧旋转来使下肢内收,从而发挥作用。总体而言,研究结果表明,莱索托龙(以及一般意义上的基础鸟臀目动物)的运动肌肉杠杆作用与其他非鸟类恐龙更相似,而不是鸵鸟,代表了可能是基础恐龙的状态。这一结果与之前关于鸟臀目动物-鸟类功能趋同的假设相矛盾。