A taxonomic and anatomic assessment of the extinct Zygodactylidae (Aves) from the Green River Formation of Wyoming and placement of Zygodactylidae within Aves

Birds are the most diverse extant group of terrestrial vertebrates, and relationships amongst major extant and extinct avian lineages remain hotly debated. A clade of Aves which has received limited attention is the extinct Zygodactylidae, a species-rich group of perching birds that possess a foot with a retroverted fourth toe, an elongate tarsometatarsus and a large intermetacarpal process in the wing. Specimens currently included within Zygodactylidae previously were thought to be sister taxa to songbirds (Passeriformes) or woodpeckers and allies (Piciformes). Zygodactylids were most abundant during the Eocene in North America and Europe and persisted to the Early Miocene. Five exceptionally preserved fossils from the Early Eocene Green River Formation of Wyoming are described, and provide insights into the interrelationships of zygodactylid taxa and the position of the clade within Aves. In an attempt to resolve systematic relationships within zygodactylids, and the position of the clade within Aves, I conducted two sets of phylogenetic analyses. The first focused on clarifying relationships within Zygodactylidae. Each taxon was evaluated for 37 morphological characters. Resulting strict consensus cladograms yield topologies in which two of the new Green River specimens are positioned in a clade within Zygodactylus, a taxon previously known only from the Early Oligocene and Early Miocene of Europe. The second set of analyses sought to assess which extant avian lineage is most closely allied with Zygodactylidae. Those analyses used a dataset of 135 characters evaluated for 57 species and a supraspecific terminal, Zygodactylidae. Scoring of Zygodactylidae was based on morphological observations from all described taxa within Zygodactylidae. The extant species sample was chosen to evaluate previously proposed hypotheses of relationships between Zygodactylidae and other avian clades and included songbirds, parrots and 43 species from the coraciiform-piciform clade (e.g., woodpeckers, galbulids, rollers and motmots). Outgroup species were iteratively swapped to determine if outgroup choice affected recovered estimates of zygodactylid relationships within Aves. Zygodactylidae is the sister taxon to songbirds in the resultant tree topologies. These results forward our understanding of the relationship between Zygodactylidae and Passeriformes within Aves.