Functional space analyses reveal the function and evolution of the most bizarre theropod manual unguals

Maniraptoran dinosaurs include the ancestors of birds, and most used their hands for grasping and in flight, but early-branching maniraptorans had extraordinary claws of mysterious function. Alvarezsauroids had short, strong arms and hands with a stout, rock-pick-like, single functional finger. Therizinosaurians had elongate fingers with slender and sickle-like unguals, sometimes over one metre long. Here we develop a comprehensive methodological framework to investigate what the functions of these most bizarre bony claws are and how they formed. Our analysis includes finite element analysis and a newly established functional-space analysis and also involves shape and size effects in an assessment of function and evolution. We find a distinct functional divergence among manual unguals of early-branching maniraptorans, and we identify a complex relationship between their structural strength, morphological specialisations, and size changes. Our analysis reveals that efficient digging capabilities only emerged in late-branching alvarezsauroid forelimbs, rejecting the hypothesis of functional vestigial structures like T. rex. Our results also support the statement that most therizinosaurians were herbivores. However, the bizarre, huge Therizinosaurus had sickle-like unguals of such length that no mechanical function has been identified; we suggest they were decorative and lengthened by peramorphic growth linked to increased body size.