Phylogenetic relationships of the Australian Oligo-Miocene ratite Emuarius gidju Casuariidae

In Australia, ratites (Aves: Palaeognathae) are represented in the extant fauna by the family Casuariidae with 1 species of emu Dromaius novaehollandiae and 1 cassowary Casuarius casuarius. The Australian fossil record reveals no other extinct ratite families but there are a number of other casuariid species. Most significant of these, due to its Oligo-Miocene age and because it is known from abundant material, is Emuarius gidju. Here, we describe additional material and confirm that the taxon had a temporal range of Late Oligocene to Middle Miocene (approximately 24-15 Ma). We reveal new morphological details, including notably that the species had relatively much smaller eyes than D. novaehollandiae, in addition to a less well-developed cursorial ability, as inferred from its pelvic limb. In these respects, Emuarius is similar to Casuarius and suggest that it was adapted to denser vegetation than the open woodlands and grasslands that characterise much of Australia today and to which D. novaehollandiae, with its large eyes and enhanced cursorial ability, is strongly adapted. Emuarius was compared to and found to be distinct from the poorly provenanced Australian fossil species C. lydekkeri. We conducted a phylogenetic analysis of morphological data that robustly shows that E. gidju is the sister taxon of Dromaius and together these taxa form a clade that is sister to Casuarius. This indicates that the evolution towards enhanced cursorality that characterises Dromaius took place after the divergence of the emu-cassowary lineages and was likely not the driving mechanism of this divergence. Comparisons between D. novaehollandiae and D. baudinianus revealed no qualitative skeletal differences and we suggest that the latter taxon is best considered to be an island dwarf that should be taxonomically recognized at a subspecific level only.