Broken gears in the avian molecular clock: new phylogenetic analyses support stem galliform status for Gallinuloides wyomingensis and rallid affinities for Amitabha urbsinterdictensis

Galliformes (landfowl) have been the focus of numerous divergence dating studies that seek a refined understanding of the early radiation of living birds. The Eocene fossil birds Amitabha urbsinterdictensis (Bridger Formation) and Gallinuloides wyomingensis (Green River Formation) have been used extensively in studies dealing with the timing of evolution in crown Galliformes. Divergence estimates from studies incorporating these fossils as calibration points suggest that multiple galliform lineages radiated in the Cretaceous and survived the Cretaceous-Tertiary mass extinction. However, the phylogenetic position of both fossils has been disputed, particularly with regard to crown or stem status. In order to resolve this debate, a new study of A. urbsinterdictensis and G. wyomingensis was undertaken. Further preparation and re-examination of the A. urbsinterdictensis holotype indicates this fossil falls outside both crown and stem Galliformes, and reveals evidence for a relationship with Rallidae (rails). In order to reassess the status of G. wyomingensis, a matrix of 120 morphological characters was constructed by revising and expanding on previous studies. Phylogenetic analyses using this matrix place G. wyomingensis basal to all crown Galliformes. Stem placement of G. wyomingensis is retained and resolution is improved in combined analyses incorporating sequence data from cytochrome b, NADH dehydrogenase subunit 2, mitochondrial control region, 12S rDNA, and nuclear ovomucoid intron G. All evidence indicates that A. urbsinterdictensis and G. wyomingensis are inappropriate internal calibration points for Galliformes and may have contributed to overestimation of divergence event ages. Though stem galliforms existed in the Cretaceous, the divergence of crown lineages in the Cretaceous remains inconclusively demonstrated. Because few galliform fossils have been evaluated phylogenetically, further investigations into the tempo of galliform evolution must await identification of proper fossil calibration points. (C) The Willi Hennig Society 2009.