The influence of fossils in macroevolutionary analyses of 3D geometric morphometric data: A case study of galloanseran quadrates

In birds and other reptiles, the quadrate acts as a hinge between the lower jaw and the skull and plays an important role in avian cranial kinesis. Though previous studies have qualitatively described substantial variation in quadrate morphology among birds, none have attempted to quantify evolutionary changes in quadrate shape. Here, we investigate geometric evolution of the quadrate in Galloanserae, a major clade of extant birds uniting chicken-like (Galliformes) and duck-like (Anseriformes) fowl. We quantified morphological variation in the quadrate across 50 extant galloanseran species covering all major extant subclades using three-dimensional geometric morphometrics, and performed ancestral shape reconstructions in the context of an up-to-date neornithine phylogeny. We find that our results based only on extant quadrates may overlook plesiomorphic features captured by fossil taxa, resulting in an ancestral state reconstruction for Galloanserae that is seemingly an approximation of the average shape of the extant data set. By contrast, analyses incorporating early fossil galloanseran quadrates (from taxa such as Asteriornis, Presbyornis, and Conflicto) result in ancestral geometric reconstructions more similar to the morphology of extant galliforms, indicating that the quadrate of the last common ancestor of galloanserans may have been more morphologically and functionally similar to those of extant galliforms than to extant anseriforms. These results generally corroborate previous inferences of galloanseran quadrate plesiomorphies and identify several additional plesiomorphic features of the galloanseran quadrate for the first time. Our results illustrate the importance of incorporating fossil taxa into ancestral shape reconstructions and help elucidate important aspects of the morphology and function of the avian feeding apparatus early in crown bird evolutionary history.