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292 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY D B FIGURE 4.—Comparisons of the first phalanx of the second manual digit in dorsal view: A, Ornitholestes hermanni Osborn (modified from Osborn, 1917); B, Deinonychus antirrhopus (modified from Ostrom, 1976); c, Archaeopteryx bavarica Wellnhofer (modified from Wellnhofer, 1993); D, Confuciusomis sanctus Hou et al. (from a cast of the holotype of Confuciusomis); E, Cathayornis yandica Zhou et al. (modified from Zhou et al., 1992); F, Meleagris gallapavo Linnaeus. Drawings not to scale. shorter in the theropods. In addition to the differences mentioned by Ostrom, Archaeopteryx has many advanced avian attributes of the wrist and hand, with the complex and peculiar pattern of the avian carpometacarpus already formed at this early stage of avian evolution. Without these specializations, the attachment of the primary feathers to the hand would be hardly imaginable, let alone flight. Vazquez (1992) discussed the functional morphology of the avian wrist and stated that Archaeopteryx lacks many of the features of modem birds and was probably incapable of executing all the kinematics of modern avian powered flight. Although we might agree that modem birds have a wrist better designed for powered flight, Archaeopteryx is not so deficit in those features as Vazquez (1992) supposed. As shown by Martin (1991), Archaeopteryx has a basically avian wrist, with all of the bones found in modern birds, including an L- or Vshaped cuneiform (ulnare) to glide on the articular ridge of the carpometacarpus. This fact has been missed by most workers, including Martin (1983), because the preservation in Archaeopteryx usually shows only the dorsal or ventral side. Fortunately, the Eichstatt specimen (Wellnhofer, 1974, figs. 8, 9) presents palmar and anconal (ventral and dorsal) views of the same specimen. In palmar view the ulnare is large and elongate; in anconal view it is small and round. It could therefore be either pyramid-shaped, which would make it unlike the shape of any known relative, including other birds or dinosaurs, or L-shaped as in modem birds but not in dinosaurs. We accept the L-shaped interpretation. The perching capability of Archaeopteryx has recently been argued with strong evidence (Feduccia, 1993). The wing claws in Archaeopteryx are long and curved. The first digit diverges from the others (Zhou, 1995). The manual digits are relatively slender. All these characters, in combination, seem to show an overwhelmingly avian pattern and show that the wings could not have been used for predation (Ostrom, 1974). It seems more reasonable to suggest that the oldest bird, although limited in flying power, lived an arboreal life just as do most modem birds, with its wings used for both flight and climbing. The appearance of feathers was the critical point in avian evolution, and the modern appearance of the feathers in Archaeopteryx has often been noted (Feduccia and Tordoff, 1979; Norberg, 1995). The close match of the bones of the wrist and manus with modem birds suggests that flight played a vital role in the early evolution of birds. Furthermore, there was coevolution of the skeleton and feathers as two inseparable parts of the flight mechanism. The recognition of many avian characters in Archaeopteryx is important not only for identifying more fragmentary fossils but also for recognizing potential protobirds from even older strata. Because more and more people believe that Archaeop-
NUMBER 89 293 teryx is a side branch in avian evolution (Martin, 1983; Feduccia, 1995), the oldest ancestor of birds might have existed in the Early or Middle Jurassic or even Late Triassic. The recent Chinese finding of a Late Jurassic-Early Cretaceous beaked bird (Hou et al, 1995) seems to lend further credibility to this proposal. Many of the characters discussed above may appear to be subtle, but their importance and evolutionary implication are probably no less than many superficially significant morphological changes. Although many shared features have been de Beer, G. 1954. Archaeopteryx lithographica: A Study Based on the British Museum Specimen. 68 pages. London: British Museum (Natural History). Feduccia, A. 1993. Evidence from Claw Geometry Indicating Arboreal Habits of Archaeopteryx. Science, 259:790-793. 1995. Explosive Evolution in Tertiary Birds and Mammals. Science, 267:637-638. Feduccia, A., and H.B. Tordoff 1979. Feathers of Archaeopteryx: Asymmetric Vanes Indicate Aerodynamic Function. Science, 203:1021-1022. Fisher, H. 1957. Bony Mechanism of Automatic Flexion and Extension in the Pigeon's Wing. Science, 126:446. Heilmann, G. 1926. The Origin of Birds, iii+208 pages. London: H.F. and G. Whitherby. Hinchliffe, J.R. 1985. "One, Two, Three" or "Two, Three, Four": an Embryologist's View of the Homologies of the Digits and Carpus of Modern Birds. In M.K. Hecht, J. K. Ostrom, G. Viohl, and P. Wellnhofer, editors, The Beginning of Birds, pages 141-147. Eichstatt: Freunde des Jura-Museums Eichstatt, Willibaldsburg. Hogg, D.A. 1980. A Re-investigation of the Centers of Ossification in the Avian Skeleton at and After Hatching. Journal of Anatomy, 130(4):725-743. Holmgren, N. 1955. Studies on the Phylogeny of Birds. Acta Zoologica, 36:1085. Hou, L., Z. Zhou, L. Martin, and A. Feduccia 1995. A Beaked Bird from the Jurassic of China. Nature, 377:616-618. Martin, L.D. 1983. The Origin and Early Radiation of Birds. In A.H. Bush and G.A. Clark, editors, Perspectives in Ornithology: Essays Presented for the Centennial of the American Ornithologist's Union, pages 291-338. Cambridge: Cambridge University Press. 1991. Mesozoic Birds and the Origin of Birds. In H.-P. Schultze and L. Trueb, editors, Origins of the Higher Groups ofTetrapods, pages 485-540. Ithaca: Cornell University Press. Literature Cited suggested between theropod dinosaurs and Archaeopteryx, they often lack the detailed similarity we should expect in homologous characters. Ostrom (1985) recognized only two uniquely avian characters in Archaeopteryx: an ossified furcula and feathers. The disclosure of eight uniquely avian characters in the wrist and manus of Archaeopteryx provides further evidence for mosaic evolution in the vertebrate history, and encourages us to examine the anatomy of these unique fossils more closely. Norberg, R.A. 1995. Feather Asymmetry in Archaeopteryx (Response to J.R. Speakman and S.C Thomson in Vol. 370, p. 514). Nature, 374(6519):221. Osborn, H.F. 1917. Skeletal Adaptations of Ornitholestes, Struthiomimus, Tyrannosaurus. Bulletin of the American Museum of Natural History, 35: 733-771. Ostrom, J.H. 1974. Archaeopteryx and the Origin of Flight. Quarterly Review of Biology, 49:21-41. 1976. Archaeopteryx and the Origin of Birds. Biological Journal of the Linnean Society, 8:91-182. 1985. The Meaning of Archaeopteryx. In M.K. Hecht, J.K. Ostrom, G. Viohl, and P. Wellnhofer, editors, The Beginning of Birds, pages 161-176. Eichstatt: Freunde des Jura-Museums Eichstatt, Willibaldsburg. Ostrom, J. 1995. Wing Biomechanics and the Origin of Bird Flight. Neues Jahrbuch fiir Geologie und Paldontologie Abhandlungen, 195(1—3):253—266. Vazquez, R.J. 1992. Functional Osteology of the Avian Wrist and the Evolution of Flapping Flight. Journal of Morphology, 211:259-268. Wellnhofer, P. 1974. Das funfte Skelettexemplar von Archaeopteryx. Palaeontographica, A, 147:169-216. 1988. Ein neues Exemplar von Archaeopteryx. Archaeopteryx, 6:1-30. 1993. Das siebte Exemplar von Archaeopteryx aus den Solnhofener Schichten. Archaeopteryx, 11:1—47. Zhou, Z. 1995. 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. Zhou, Z., F. Jin, and J. Zhang 1992. Preliminary Report on a Mesozoic Bird from Liaoning, China. Chinese Science Bulletin, 37:1365-1368.
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SMITHSONIAN CONTRIBUTIONS TO PALEOB
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ABSTRACT Olson, Storrs L., editor.
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EARLY WATERFOWL (ANSERIFORMES) AND
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v m SMITHSONIAN CONTRIBUTIONS TO PA
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localities, all of them situated in
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Bone of Sus scrofa Linnaeus, introd
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duboisi are larger than the largest
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8 iver (1897) but afterward were tr
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10 SMITHSONIAN CONTRIBUTIONS TO PAL
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16 Cor. Hum. Uln. Cpm. Fern. Tbt. T
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20 sometatarsus are proportionally
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22 Cor. Hum. Uln. Cpm. Fern. Tbt. T
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26 Cor. Hum. Cpm. Fern. Tbt. Tmt. P
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34 ability in such a way that it ca
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42 2km SMITHSONIAN CONTRIBUTIONS TO
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Site 13 Research Base ENTRANCE Lowe
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48 the last has unusually slender w
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50 Si 5 15i 10- 5- 20 J 15 I 10 f 5
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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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NUMBER 89 107 yellowish sand (site
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NUMBER 89 109 ogy. Notornis, supple
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112 SMITHSONIAN CONTRIBUTIONS TO PA
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The Middle Pleistocene Avifauna of
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NUMBER 89 Accordi, B. 1962. La grot
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130 FIGURE 1.—Map showing locatio
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132 Cranium Mandibula Scapula Clavi
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Seabirds and Late Pleistocene Marin
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NUMBER 89 141 METHODS Only strictly
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NUMBER 89 143 FIGURE 2.—Area of s
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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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NUMBER 89 181 FIGURE 5.—Vulturida
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NUMBER 89 183 FIGURE 7.—Referred
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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 89 189 carpal trochlea relat
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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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NUMBER 89 195 FIGURE 6.—Distribut
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NUMBER 89 197 the Florida State Mus
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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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NUMBER 89 221 Costae: Deformed frag
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A Fossil Screamer (Anseriformes: An
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NUMBER 89 FIGURE 3.—Chaunoides an
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NUMBER 89 227 B C D FIGURE 6.—The
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NUMBER 89 229 FIGURE 9.—Right tib
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The Anseriform Relationships of Ana
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NUMBER 89 233 Subfamily ANATALAVINA
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NUMBER 89 235 mal was found under t
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NUMBER 89 237 tion, with retroartic
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NUMBER 89 FIGURE 7.—Sternum and p
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Page 267 and 268: NUMBER 89 257 vical vertebrae of th
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Page 284 and 285: 274 America. Science, 214(4526): 12
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Page 288 and 289: 278 ment of the radius that are con
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Page 298 and 299: 288 Palaeontological Institute. [Sp
Page 300 and 301: 290 1991). In this paper we use a "
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