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288 Palaeontological Institute. [Special issue.] Kurten, Bjom 1967-1970. Continental Drift and the Palaeogeography of Reptiles and Mammals. Commentationes Biologicae, Societas Scientiarum Fennica, 31(1): 1-8, 2 figures. [Volume 31 number 1 published in 1967.] Lacasa Ruiz, A. 1989. Nuevo genero de ave fosil del yacimiento neocomiense del Montsec (provincia de Lerida, Espana). Estudios Geologicos, Instituto de Geologia (Madrid), 45(5-6):417-425, 5 figures. Lamb, J.P, L.M. Chiappe, and P.G.P. Erickson 1993. A Marine Enantiornithine from the Cretaceous of Alabama. [Abstract.] Journal of Vertebrate Paleontology, 13(3):45. Martin, L. 1995. The Enantiornithes: Terrestrial Birds of the Cretaceous. In D.S. Peters, editor, Acta Palaeomithologica, 3 Symposium SAPE; 5 Internationale Senckenberg-Konferenz 22-26 Juni 1992. Courier Forschungsinstitut Senckenberg, 181:23-36, 10 figures. Molnar, R.E. 1986. An Enantiornithine Bird from the Lower Cretaceous of Queensland, Australia. Nature, 322:736-738, 4 figures. Nesov, L.A., and A.V. Panteleev 1993. [On the Similarity of the Cretaceous Ornithofaunas between South America and Western Asia.] Russian Academy of Sciences, Proceedings of the Zoological Institute, St. Petersburg, 252:84-94. [In Russian, with English summary.] Rich, P.V. 1976. The History of Birds on the Island Continent Australia. Proceedings of the 16th International Ornithological Congress, Canberra, pages 53-65, 6 figures. Sanz, J.L., and J.F. Bonaparte 1992. A New Order of Birds (Class Aves) from the Lower Cretaceous of Spain. In K.E. Campbell, Jr., editor, Papers in Avian Paleontology SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY Honoring Pierce Brodkorb. Science Series, Natural History Museum of Los Angeles County, 36:39-49, 8 figures. Sanz, J.L., and A.D. Buscalioni 1992. A New Bird from the Early Cretaceous of Las Hoyas, Spain, and the Early Radiation of Birds. Palaeontology, 35(4):829-845, 11 figures. Sanz, J.L., L.M. Chiappe, P.B. Perez-Moreno, A.D. Buscalioni, J.J. Moratalla, F. Ortega, and F.J. Poyato-Ariza 1996. An Early Cretaceous Bird from Spain and Its Implications for the Evolution of Avian Flight. Nature, 382:442-445. Sereno, PC, and C.G. Rao 1992. Early Evolution of Avian Flight and Perching: New Evidence from the Lower Cretaceous of China. Science, 255:845-848, 5 figures. Smith, A.G., D.G. Smith, and B.M. Funnell 1995. Atlas of Mesozoic and Cenozoic Coastlines, ix+99 pages, 2 figures, 31 numbered maps, 1 unnumbered map. Cambridge: Cambridge University Press. [First published in 1994, reprinted in 1995.] Walker, CA. 1981. New Subclass of Birds from the Cretaceous of South America. Nature, 292:51-53. Zhou, Z. 1995a. The Discovery of Early Cretaceous Birds in China. In D.S. Peters, editor, Acta palaeomithologica, 3 Symposium SAPE; 5 Internationale Senckenberg-Konferenz 22-26 Juni 1992. Courier Forschungsinstitut Senckenberg, 181:9-22, 10 figures. 1995b. Discovery of a New Enantiornithine Bird from the Early Cretaceous of Liaoning, China. Vertebrata Palasiatica, 33(2):99-l 13,4 figures, 1 plate. 1995c. New Understanding of the Evolution of the Limb and Girdle Elements in Early Birds—Evidence from Chinese Fossils. In A. Sun and Y. Wang, editors, Sixth Symposium on Mesozoic Terrestrial Ecosystems and Biota; Short Papers, pages 209-214. Beijing: China Ocean Press.
Feathered Dinosaur or Bird? A New Look at the Hand of Archaeopteryx ABSTRACT A detailed examination of wrist and manus skeletons in Archaeopteryx, and their comparison with those of modem birds, demonstrates an overwhelmingly avian appearance, much more so than has been previously recognized. Many workers have considered feathers to be the only indisputable evidence for the avian identity of this early bird. Although only a few skeletal characters have been used to support its avian identity, we believe that this is due to a lack of detail in previous analyses. We offer a list of eight uniquely derived avian characters or character complexes in the wrist and manus of Archaeopteryx. This further indicates that Archaeopteryx is a bird, with wings used for flying rather than for predation, and provides some fundamental skeletal differences between the oldest birds and their immediate ancestors. We extend our comparisons to the only other bird with Archaeopteryx-like morphology in the manus, Confuciusornis, and show how the wrist and manus may provide useful clues for discerning potentially older and unknown birds in the future. In addition, the large number of uniquely avian characters in the wrist and manus contrasts with a more primitive anatomy in other parts, providing another example of mosaic evolution, as the structure of the wing modernized at a more rapid rate than other anatomical units. Introduction Since the discovery of Archaeopteryx in 1861, extensive studies have been conducted on this genus, and the past two decades marked a new era of study for Jurassic birds. One result has been the resurrection, mainly by Ostrom in the 1970s, of the theory of the dinosaur origin of birds. This hypothesis derives most of its support from comparison between Archaeopteryx and a few theropod dinosaurs, primarily Deinonychus. Although strongly challenged by ornithologists and many pale- Zhonghe Zhou and Larry D. Martin Zhonghe Zhou, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, P.O. Box 643, Beijing 100044, China. Larry Martin, Natural History Museum and Department of Ecology and Evolutionary Biology, University of Kansas, Lawrence, Kansas 66045, United States. 289 ornithologists, this theory has been broadly acclaimed among vertebrate paleontologists. Ostrom even went farther, stating that, were it not for the remarkable feather imprints, both of the early Archaeopteryx specimens (London and Berlin) would have been identified unquestionably as coelurosaurian theropods (Ostrom, 1976). This argument has been echoed in an extensive literature. Less attention has been paid to the significant similarity between Archaeopteryx and modem birds apart from the feathers and claws (Feduccia and Tordoff, 1979; Feduccia, 1993). The wrist and manus bones in Archaeopteryx, when submitted to detailed analysis and comparison with modem birds, illustrate many avian skeletal characters that are important to the flight of birds and that were subject to complex morphological change in early avian evolution. ACKNOWLEDGMENTS.—We thank Desui Miao for reading the manuscript, and we benefited from discussion with Christopher Bennett. We are grateful to Peter Wellnhofer, Burkhard Stephan, Lawrence Witmer, and John Ostrom for their critical reviews and valuable advice and suggestions. We also are indebted to John Chorn for reading the abstract and preparing some of the figures. Character Analyses A total of eight uniquely derived avian characters or character complexes from the wrist and manus skeletons of Archaeopteryx are recognized. We have been able to examine the originals or good casts of all seven known specimens of Archaeopteryx, especially the Berlin and Eichstatt specimens. Ostrom listed several theropod dinosaurs as having the closest wrist and manus structure to Archaeopteryx. Among the genera most frequently used in comparisons are Deinonychus, Velociraptor, Omitholestes, and Chirostenotes; therefore, our comparisons will focus on the similarities between Archaeopteryx and modem birds on one hand, and the difference between Archaeopteryx and these dinosaurs on the other. The homologies of the digits of birds and dinosaurs is still controversial among paleontologists and embryologists (Hinchliffe, 1985; Martin,
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SMITHSONIAN CONTRIBUTIONS TO PALEOB
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ABSTRACT Olson, Storrs L., editor.
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42 2km SMITHSONIAN CONTRIBUTIONS TO
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Site 13 Research Base ENTRANCE Lowe
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Comparison of Paleoecological Patte
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NUMBER 89 69 TABLE 1.—Vertebrate
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NUMBER 89 71 Mallorca, probably in
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NUMBER 89 Alcover, J.A. 1989. Les a
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The History of the Chatham Islands'
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NUMBER 89 87 Species Common Name Pe
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NUMBER 89 89 NZA 797,1930,1934 Unco
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NUMBER 89 91 anoramphus spp.), Chat
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NUMBER 89 93 TABLE 2.—Land snails
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NUMBER 89 95 counterparts, with som
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NUMBER 89 population, if such exist
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NUMBER 89 99 FIGURE 11.—Skulls of
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NUMBER 89 101 FIGURE 13.—Lower ma
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NUMBER 89 the Chatham Island Pied O
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NUMBER 89 Locality 9, Western Maung
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The Middle Pleistocene Avifauna of
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NUMBER 89 Accordi, B. 1962. La grot
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Seabirds and Late Pleistocene Marin
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NUMBER 89 141 METHODS Only strictly
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NUMBER 89 151 FIGURE 10.—Area of
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NUMBER 89 153 FIGURE 12.—Area of
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NUMBER 89 155 the period studied. T
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NUMBER 89 157 Walker, C.A., G.M. Wr
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164 Vl 620 M 570 £ 520 S 470f •
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166 birds, such as the two species
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170 cional Autonoma de Mexico, for
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174 ated with this specimen, see Mi
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The Fossil Record of Condors (Cicon
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NUMBER 89 179 FIGURE 2.—Geographi
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Two New Fossil Eagles from the Late
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NUMBER 89 187 TABLE 1.—Measuremen
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NUMBER 89 191 FIGURE 4.—Holotypic
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NUMBER 89 193 We compared the parat
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A New Genus of Dwarf Megapode (Gall
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NUMBER 89 201 lis hypotarsi along t
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NUMBER 89 203 The fossil is larger
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NUMBER 89 205 Clark, George A., Jr.
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A New Genus and Species of the Fami
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NUMBER 89 209 son with other known
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NUMBER 89 211 FIGURE 1.—Argornis
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NUMBER 89 213 AM AL AM AL AM AL AM
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NUMBER 89 215 caput humeri perpendi
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Selmes absurdipes, New Genus, New S
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NUMBER 89 219 FIGURE 2.—Selmes ab
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A Fossil Screamer (Anseriformes: An
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The Anseriform Relationships of Ana
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NUMBER 89 233 Subfamily ANATALAVINA
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