A phylogenetic analysis of basal Anseriformes, the fossil Presbyornis, and the interordinal relationships of waterfowl

A phylogenetic analysis of 123 morphological characters of basal waterfowl (Aves: Anseriformes) and other selected avian orders confirmed that the screamers (Anhimae: Anhitn-idae) are the sister-group of other waterfowl (Anseres), and that the magpie goose (Anseranatidae: Anseranas semipalmata) is the sister group of other modern waterfowl exclusive of screamers (Anatidae sensu stricto). The analysis also supports the traditional hypothesis of the gallinaceous birds (Galliformes) as the sister group of the Anseriformes. Presbyornis, a fossil from the early Eocene of Wyoming and averred by Olson & Feduccia as showing that the Anseriformes were derived from shorebirds (Charadriiformes), was found to represent the sister group of the Anatidae. Associated hypotheses by Olson & Feduccia concerning the implications of Presbyornis for the phylogenetic relationships of flamingos (Phoenicopteriformes), the position of the Anhimidae within the waterfowl, relationships among modern Anatidae, and a plausible evolutionary scenario for waterfowl also are rejected. Analyses revealed that cranial characters were critical to the establishment of the Galliformes as the sister group of the Anseriformes; exclusion of the Anhimidae, especially in combination with Anseranas, also undermined the support for this inference. Placement of Presbyornis as the sister group of the Anatidae casts doubt on the role suggested by Feduccia of ‘transitional shorebirds' in the origin of modern avian orders, and calls into question the concept of ‘fossil mosaics’. The phylogenetic hypothesis is used to reconstruct an evolutionary scenario for selected ecomorphological characters in the galliform-anseriform transition, to predict the most parsimonious states of these characters for Presbyornis, and to propose a phylogenetic classification of the higher-order taxa of waterfowl. This re-examination of Presbyornis also is used to exemplify the fundamental methodological shortcomings of the intuitive approach to the reconstruction of phylogenetic relationships.     

Systematic relationships of the palaeogene family Presbyornithidae (Aves: Anseriformes)

The early Tertiary (Paleocene and Eocene) family Presbyornithidae is one of the most completely known group of fossil birds. Essentially all parts of the skeleton are represented in the fossil record, allowing a thorough analysis of the phylogenetic position of the family. Forty-two families of nonpasserine birds representing the orders Ciconiiformes, Anseriformes, Galliformes, Gruiformes and Charadriiformes, were included in a cladistic analysis of 71 skeletal characters. The previously suggested anseriform affinity of the Presbyornithidae was confirmed. Furthermore, the family proved to be closer to the Anatidae than to the Anhimidae or Anseranatidae. The many postcranial similarities with certain charadriiform birds as the Burhinidae, obviously are plesiomorphies. By this observation, a better undestanding of character evolution in nonpasserine skeletal morphology is gained. The often suggested close relationship of anseriform and galliform birds is not confirmed by osteology. Instead, the Anseriformes and the Phoenicopteridae form a monophyletic clade that is the sister to the remaining ciconiiform birds. This result renders the Ciconiiformes sensu Wetmore (1960) polyphyletic.     

Morphology of the quadrate in the Eocene anseriform Presbyornis and extant galloanserine birds

Despite the notoriety, phylogenetic significance, and large number of available specimens of Presbyornis, its cranial anatomy has never been studied in detail, and its quadrate has been partly misinterpreted. We studied five quadrates of Presbyornis that reveal features hitherto unknown in the anseriforms but otherwise present in galliforms. As a result, we analyzed the variable quadrate characters among all extant galloanserine families and identified synapomorphies and other morphological variation among the major galloanserine clades. In terms of quadrate morphology, Presbyornis is more plesiomorphic than any extant anseriform (including the Anhimidae) and shares ancestral galloanserine characters with the Megapodiidae, the earliest branch of extant galliforms. The quadrate's morphology is inconsistent with the currently accepted anseriform phylogeny that nests Presbyornis within the crown‐group as a close relative of the Anatidae. The presbyornithid quadrates exhibit an unusual variation in the presence of a caudomedial pneumatic foramen, which we interpret as a result of a discontinuous change in the growth path of the pneumatic diverticulum. Another episode of morphogenetic imbalance in the growth path of the pneumatic diverticulum may have accompanied the disappearance of the basiorbital pneumatic foramen (along with the pneumatization of the pterygoid) at the origin of the crown‐group anseriforms. This episode is marked by the striking individual variation in the presence and location of pneumatic foramina in the mandibular part of the quadrate in the Anhimidae. J. Morphol., 2010. © 2009 Wiley‐Liss, Inc.