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(Early Eocene) Lithornithids (Aves) and the Relationships of Lithornithidae Author(S): Sterling J

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(Early Eocene) Lithornithids (Aves) and the Relationships of Lithornithidae Author(S): Sterling J The Anatomy and Taxonomy of the Exquisitely Preserved Green River Formation (Early Eocene) Lithornithids (Aves) and the Relationships of Lithornithidae Author(s): Sterling J. NesbittJulia A. Clarke Source: Bulletin of the American Museum of Natural History, :1-91. Published By: American Museum of Natural History DOI: http://dx.doi.org/10.1206/0003-0090-406.1.1 URL: http://www.bioone.org/doi/full/10.1206/0003-0090-406.1.1 BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the biological, ecological, and environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/terms_of_use. Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder. BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. THE ANATOMY AND TAXONOMY OF THE EXQUISITELY PRESERVED GREEN RIVER FORMATION (EARLY EOCENE) LITHORNITHIDS (AVES) AND THE RELATIONSHIPS OF LITHORNITHIDAE STERLING J. NESBITT Jackson School of Geosciences, the University of Texas at Austin; Division of Paleontology, American Museum of Natural History; Department of Geosciences, Virginia Polytechnic Institute and State University JULIA A. CLARKE Jackson School of Geosciences, the University of Texas at Austin, Division of Paleontology, American Museum of Natural History BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY Number 406, 91 pp., 35 figures, 7 tables Issued June 30, 2016 Copyright © American Museum of Natural History 2016 ISSN 0003-0090 CONTENTS Abstract.............................................................................4 Introduction.........................................................................4 Anatomical abbreviations . .9 Institutional acronyms ...............................................................10 Terminology........................................................................10 Systematic paleontology..............................................................11 Description of Calciavis grandei (AMNH 30578) ........................................11 Skull ............................................................................13 Vertebral column .................................................................16 Ribs.............................................................................18 Pectoral girdle....................................................................19 Forelimb ........................................................................19 Pelvic girdle......................................................................22 Hind limb .......................................................................23 Soft tissue .......................................................................24 Referred specimen of Calciavis grandei AMNH 30560 .................................. 25 Locality .........................................................................25 Preservation .....................................................................25 Skull ............................................................................25 Postcrania .......................................................................27 Other new Green River Formation lithornithid specimens................................29 FMNH PA 729 ...................................................................29 Locality.......................................................................29 Preservation...................................................................29 Description ...................................................................29 Feathering ....................................................................31 Phylogenetic assignment ........................................................31 FMNH PA 739 ...................................................................32 Locality.......................................................................32 Preservation...................................................................32 Description ...................................................................32 Phylogenetic assignment ........................................................34 WSGS U1b-2001 .................................................................34 Locality.......................................................................34 Preservation...................................................................34 Description ...................................................................34 Feathering ....................................................................35 Phylogenetic assignment ........................................................37 Phylogenetic analysis . 37 Taxon sampling ..................................................................37 Character sampling ...............................................................39 Methods.........................................................................40 Results . 41 Alternative hypotheses of the relationships of Lithornithidae ...........................44 2 2016 NESBITT & CLARKE: ANATOMY, TAXONOMY, AND RELATIONSHIPS OF LITHORNITHIDS 3 Alternative hypothesis discussion ................................................51 Discussion .........................................................................53 Species diversity of Green River Formation lithornithids...............................53 Ontogenetic age . .53 Morphological differences.......................................................54 Sexual dimorphism?............................................................56 Monophyly of Lithornithidae and diagnostic features of the clade.......................59 Is Paracathartes howardae part of Lithornithidae?.....................................60 What is the basalmost lithornithid lineage? ..........................................62 Variation within Lithornithidae.....................................................63 Relationships of Lithornithidae .....................................................67 Perspectives ........................................................................70 Acknowledgments...................................................................70 References..........................................................................70 Appendix 1. Phylogenetic characters and data set .......................................75 Appendix 2. Lithornithid terminal taxa ................................................86 ABSTRACT Fossil remains of Paleogene Palaeognathae are poorly documented and are exceedingly rare. One group of palaeognaths, the lithornithids, is well represented in the Paleogene of North America. Nevertheless, few specimens of the same species are known from each of those Paleogene geologic units. Here, we report five new partial skeletons of lithornithids from the Fossil Butte Member of the Green River Formation (Early Eocene) of Wyoming. One spectacularly preserved specimen is identified as the holotype of a new species, Calciavis grandei, gen. et sp. nov., and fully described. Preserved soft tissues (e.g., feathers, pes scales) surround the nearly articulated and complete skel- eton. A second well-preserved but disarticulated skeleton is referred to this new taxon. We conclude that there are only two lithornithid taxa in the Green River Formation after careful comparisons with the other known taxon from the same geological unit, Pseudocrypturus cercanaxius. Morpho- logical data generated from the new taxon and other Green River Formation lithornithid specimens were integrated into a osteology-only phylogenetic data set containing stem avians as outgroups and extinct and extant members of Palaeognathae (Tinamidae, ratites) and Neognathaes (Anseriformes, Galliformes, Neoaves), unnamed lithornithid specimens, and the following named lithornithid taxa: Lithornis plebius, Lithornis promiscuus, Lithornis celetius, Paracathartes howardae. We find a mono- phyletic Lithornithidae (containing Calciavis grandei, Pseudocrypturus cercanaxius, Lithornis plebius, Lithornis promiscuus, Lithornis celetius, Paracathartes howardae) as the sister taxon of Tinamidae at the base of Palaeognathae and also recover a monophyletic Ratitae in the morphology-only analysis. A Lithornithidae-Tinamidae relationship, which could imply a broad Northern Hemisphere distri- bution in the Paleogene for this total group retracted to the present day Neotropical distribution after the Eocene, is weakly supported in our analysis and is also supported by other lines of evidence such as eggshell morphology. Relationships among flightless palaeognaths and assessment of char- acter homology in this group remain problematic. Indeed, when the morphological analyses were constrained to enforce topologies recovered from all recent analyses of molecular sequence data and retroelement insertions, Lithornithidae is no longer recovered with Tinamidae, which is nested within the now paraphyletic ratites, but remains at the base of Palaeognathae. Thus, regardless of the
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