Skull morphology of the enigmatic Genyornis newtoni Stirling and Zeitz, 1896 (Aves, Dromornithidae), with implications for functional morphology, ecology, and evolution in the context of Galloanserae

The presence of Dromornithidae in the Australian Cenozoic fossil record was first reported in 1872, yet although eight species and hundreds of specimens are known, key information on their morphology remains elusive. This is especially so for their skulls, which contributes to a lack of resolution regarding their relationships within Galloanserae. The skull of the Pleistocene dromornithid, Genyornis newtoni, was initially described in 1913. Additional fossils of this species have since been discovered and understanding of avian skull osteology, arthrology, and myological correlates has greatly advanced. Here we present a complete redescription of the skull of Genyornis newtoni, updating knowledge on its morphology, soft-tissue correlates, and palaeobiology. We explore the diversity within Dromornithidae and make comprehensive comparisons to fossil and extant galloanserans. Furthermore, we expand on the homologies of skull muscles, especially regarding the jaw adductors and address the conflicting and unstable placement of dromornithids within Galloanserae. Findings support generic distinction of Genyornis newtoni, and do not support the close association of Dromornithidae and Gastornithidae. We thus recommend removal of the dromornithids from the Gastornithiformes. Considering character polarities, the results of our phylogenetic analyses, and palaeogeography, our findings instead support the alternative hypotheses, of dromornithids within, or close to, the Suborder Anhimae with Anseriformes.