Bill disparity and feeding strategies amongst fossil and modern penguins

One of the most remarkable differences between Paleogene penguins and their living relatives is the shape and length of their beaks. Many of the Eocene and Oligocene penguins have a thin and elongated spear-like bill, which contrasts with the proportionally shorter and more robust bill of most living species. These differences suggest an important shift on their feeding strategies. This study explores the morphological disparity on the skull of penguins, emphasizing bill morphology and it relationship with feeding habits. For this, the skull of 118 species of aquatic birds including 21 fossil and living penguins were analysed using two-dimensional geometric morphometric. The results show that, unlike what has been reported for modern birds overall, in penguins and Aequornithes bill elongation is related to a reduction of the braincase. The discriminant analysis shows that there are significant differences between penguins that feed near or far from the coast and between those that consume nectonic and planktonic prey, identifying Madryniornis as the only extinct form with possible planktonic diet. Additionally, it is clear that Paleogene penguins occupy a region of morphospace unexplored by most diving birds, with the western grebe being its closest modern analogue. This is consistent with the hypothesis that giant penguins hunted by harpooning and not by biting as living forms do, signalling a significant change in the habits of these birds leading to the emergence of its crown group.