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From the Late Mesozoic of Northeastern China

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From the Late Mesozoic of Northeastern China ANATOMY AND SYSTEMATICS OF THE CONFUCIUSORNITHIDAE (THEROPODA: AVES) FROM THE LATE MESOZOIC OF NORTHEASTERN CHINA LUIS M. CHIAPPE Research Associate, Department of Ornithology American Museum of Natural History Current address: Associate Curator, Vertebrate Paleontology Natural History Museum of Los Angeles County 900 Exposition Boulevard, Los Angeles, CA 90007 JI SHU'AN National Geological Museum of China Yangrou Hutong 15, Xisi, 100034 Beijing, China JI QIANG National Geological Museum of China Yangrou Hutong 15, Xisi, 100034 Beijing, China MARK A. NORELL Chairman Division of Paleontology American Museum of Natural History BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY Number 242, 89 pages, 70 ®gures, 4 tables Issued November 10, 1999 Price: $8.60 a copy Copyright q American Museum of Natural History 1999 ISSN 0003-0090 CONTENTS Abstract ....................................................................... 3 Introduction .................................................................... 4 Anatomical Abbreviations ..................................................... 4 Institutional Abbreviations ..................................................... 5 Geological Setting .............................................................. 5 Systematic Paleontology ......................................................... 9 Anatomy of Confuciusornis sanctus .............................................. 17 Skull and Mandible .......................................................... 17 Vertebral Column ............................................................ 25 Thoracic Girdle ............................................................. 29 Sternum, Ribs, and Gastralia .................................................. 29 Thoracic Limb .............................................................. 33 Pelvic Girdle ................................................................ 41 Pelvic Limb ................................................................. 43 Plumage .................................................................... 48 Anatomy of Changchengornis hengdaoziensis ..................................... 50 Skull and Mandible .......................................................... 50 Vertebral Column ............................................................ 51 Thoracic Girdle ............................................................. 53 Sternum, Ribs, and Gastralia .................................................. 58 Thoracic Limb .............................................................. 60 Pelvic Girdle ................................................................ 63 Pelvic Limb ................................................................. 63 Plumage .................................................................... 67 Forgeries ..................................................................... 68 Systematics of the Confuciusornithidae ........................................... 68 Implications for the Evolution of Avian Cranial Kinesis ............................ 72 Differential Plumage and Sexual Dimorphism ..................................... 75 Life-style of the Confuciusornithidae ............................................. 77 Acknowledgments ............................................................. 80 References .................................................................... 80 Index ......................................................................... 87 2 1999 CHIAPPE ET AL.: FOSSIL BIRDS FROM NORTHEASTERN CHINA 3 ABSTRACT The osteology and plumage of Confuciusornis sanctus and Changchengornis hengdaoziensis from the Chaomidianzi Formation (previously referred to as the lower section of the Yixian Formation) of western Liaoning Province (China) are described in detail. Confuciusornis sanctus and Changchengornis hengdaoziensis have toothless, beaked skulls (the tomial crest is straight in the former species and strongly curved in the latter) and retain the dorsal portion of the nasal process of the maxilla. The skull of Confuciusornis sanctus is of typical diapsid plan. It possesses a triradiate postorbital that with the squamosal forms a complete supratemporal arcade. Furthermore, a robust jugal±postorbital contact completely separates the infratemporal fenestra from the orbit. Although the postorbital region is not preserved in Changchengornis hengdaoziensis, it likely resembles that of Confuciusornis sanc- tus. Both species have abbreviated tails with long pygostyles, not the long, bony tail originally reconstructed in Confuciusornis sanctus. Scapulae and coracoids are fused to form scapulo- coracoids. The coracoids are strutlike but much shorter than the scapulae. The furculae are robust and boomerang shaped. The sterna are long and nearly ¯at. A complete basket of gastralia follows the sternum caudally. The wing elements are short, and proportions among individual bones are primitive in that the hand is longer than either the humerus or the ulna and the ulna is shorter than the humerus. The pelvis is opisthopubic. The postacetabular wing of the ilium is much shorter than the preacetabular wing. Differences in plumage, namely the presence or absence of two very long tail feathers, are observed among several well-preserved specimens of Confuciusornis sanctus. This difference is likely the expression of sexual di- morphism, although other biological attributes known for extant populations (e.g., differential molting, correlation between sexual maturity and ornamental plumage) indicate that alternative explanations may also account for the observed plumage variation among specimens of Con- fuciusornis sanctus. Given the latest Jurassic±earliest Cretaceous age of the Chaomidianzi Formation, Confuciu- sornis sanctus and Changchengornis hengdaoziensis are surely among the oldest known birds after the Early Tithonian Archaeopteryx lithographica. Confuciusornis sanctus and Chang- chengornis hengdaoziensis thus furnish the earliest record of beaked birds. The fully diapsid skull of Confuciusornis sanctus, and presumably of Changchengornis hengdaoziensis, and the absence of a bending zone on the base of the snout suggest that earlier interpretations of the skull of Confuciusornis sanctus as prokinetic are incorrect. Confuciu- sornis sanctus probably had very limited cranial kinetic capabilities, if any. Optimization of the postorbital±jugal contact, a character intimately correlated with intracranial kinesis, in a phylogeny of basal avians indicates that the essentially akinetic condition of the skull of Confuciusornis sanctus is a reversal derived from forms possessing kinetic properties. Recent recognition of two additional species of ConfuciusornisÐConfuciusornis chuon- zhous and Confuciusornis suniaeÐare based on anatomical misinterpretations. Thus, Confu- ciusornis suniae and Confuciusornis chuonzhous are regarded as junior synonyms of Confu- ciusornis sanctus. Confuciusornis sanctus is the sister-taxon of Changchengornis hengdao- ziensis, and both are placed within the Confuciusornithidae. Some of the synapomorphies supporting this grouping include the presence of edentolous jaws, a rostrally forked mandibular symphysis, a reduced claw of manual digit II, and a V-shaped caudal margin of the sternum. The Confuciusornithidae is considered to be the sister-group of a clade composed of the Enantiornithomorpha and the Ornithuromorpha. This phylogenetic interpretation is far more parsimonious than previous systematic hypotheses placing the Confuciusornithidae as either within the Enantiornithes or as its sister-group. Purported climbing specializations of Confuciusornis sanctus are evaluated in light of the available anatomical evidence. We conclude that both Confuciusornis sanctus and Chang- chengornis hengdaoziensis were ill suited for tree climbing. The foot of Changchengornis hengdaoziensis, however, suggests a greater grasping ability than that of Confuciusornis sanc- tus. It is argued that both Confuciusornis sanctus and Changchengornis hengdaoziensis were able to ¯y and take off from the ground. The remarkable concentration of specimens of Confuciusornis sanctus from a relatively small quarry near the village of Sihetun (Liaoning Province) suggests several events of mass mortality and, perhaps, a gregarious behavior. 4 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 242 INTRODUCTION The Late Jurassic±Early Cretaceous de- posits of western Liaoning Province (north- eastern China) are famous for the extraordi- nary preservation of fossils (e.g., Zhou Z.-H. et al., 1992; Hou et al., 1995b, c, 1996; Ji Q. and Ji S.-A., 1996, 1997a, b; Hou, 1997; Chen et al., 1998; Ji Q. et al., 1998). These deposits have furnished one of the most im- portant, and certainly the most abundant, col- lections of Mesozoic birds. Taxa discovered in these deposits include Sinornis santensis (Sereno and Rao, 1992), Cathayornis yan- dica (Zhou Z.-H. et al., 1992), Chaoyangia beishanensis (Hou and Zhang, 1993), Bolu- ochia zhengi (Zhou Z.-H., 1995a), Liaoning- ornis longiditris (Hou et al., 1996), Liaox- iornis delicatus (Hou and Chen, 1999), Changchengornis hengdaoziensis (Ji Q. et Fig. 1. Geographic location of the area of Sih- al., 1999), and Confuciusornis sanctus (Hou etun±Jianshangou, Province of Liaoning, north- et al., 1995b, c). eastern China. Confuciusornis sanctus from the Chaomi- dianzi Formation (previously referred to as the lower section of the Yixian Formation) is al., 1999). Additionally, we present a phy- one of the oldest known birds. Hundreds of logenetic discussion
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