The importance of muscle architecture in biomechanical reconstructions of extinct animals: a case study using Tyrannosaurus rex.

Functional reconstructions of extinct animals represent a crucial step towards understanding palaeocological interactions, selective pressures and macroevolutionary patterns in the fossil record. In recent years, computational approaches have revolutionised the field of 'evolutionary biomechanics' and have, in general, resulted in convergence of quantitative estimates of performance on increasingly narrow ranges for well studied taxa. Studies of body mass and locomotor performance of Tyrannosaurus rex - arguably the most intensively studied extinct animal - typify this pattern, with numerous independent studies predicting similar body masses and maximum locomotor speeds for this animal. In stark contrast to this trend, recent estimates of maximum bite force in T. rex vary considerably (> 50%) despite use of similar quantitative methodologies. Herein we demonstrate that the mechanistic causes of these disparate predictions are indicative of important and underappreciated limiting factors in biomechanical reconstructions of extinct organisms. Detailed comparison of previous models of T. rex bite force reveals that estimations of muscle fibre lengths and architecture are the principal source of disagreement between studies, and therefore that these parameters represents the greatest source of uncertainty in these reconstructions, and potentially therefore extinct animals generally. To address the issue of fibre length and architecture estimation in extinct animals we present data tabulated from the literature of muscle architecture from over 1100 muscles measured in extant terrestrial animals. Application of this dataset in a reanalysis of T. rex bite force emphasises the need for more data on jaw musculature from living carnivorous animals, alongside increased sophistication of modelling approaches. In the latter respect we predict that implementing limits on skeletal loading into musculoskeletal models will narrow predictions for T. rex bite force by excluding higher-end estimates.