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266 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY Paleocene Aquia Formation of Maryland and Virginia. Proceedings of the Biological Society of Washington, 107(3):429-435. Olson, S.L., and A. Feduccia 1980. Presbyornis and the Origin of the Anseriformes (Aves: Charadriomorphae). Smithsonian Contributions to Zoology, 323: 24 pages, 15 figures. Olson, S.L., and D.C. Parris 1987. The Cretaceous Birds of New Jersey. Smithsonian Contributions to Paleobiology, 63: 22 pages, 11 figures. Shufeldt, R.W. 1915. Fossil Birds in the Marsh Collection of Yale University. Transactions of the Connecticut Academy of Arts and Sciences, 19:1-110, 15 plates. Wetmore, A. 1926. Fossil Birds from the Green River Deposits of Eastern Utah. Annals of the Carnegie Museum, 16(3^1):391^102, plates 36, 37. 1960. A Classification for the Birds of the World. Smithsonian Miscellaneous Collections, 139( 11): 1 -37.
Implications of the Cranial Morphology of Paleognaths for Avian Evolution ABSTRACT In the early evolution of birds, bill formation produced a problem for muscular control of the thin, elongated upper jaw. In particular, it required a relatively high retracting force. Three sources of this force evolved. (1) A powerful M. retractor palatini (especially in Tinamiformes and Apteryx), originating primarily on the vomer and pterygoid, developed to provide direct muscular connection between the dermal palate and the cranial base. It apparently evolved due to a joining of the medial portions of the pterygoid and mandibular depressor muscles, which were aligned by development of the proc. mandibulae medialis (a character unique to birds). (2) The ancestral pseudotemporalis muscle developed into two portions, a large postorbital portion and an almost horizontally oriented intramandibular portion. Each portion seves to increase the retraction ability of the muscle as a whole. (3) The external mandibular adductor muscle developed, which, in neognaths, is larger than either muscle previously mentioned. Its evolutionary development was temporarily retarded by reduction of one of its places of origin—the upper temporal arch. Introduction For more than a century, paleognaths have been subjected to morphological studies in order to ascertain their apparently primitive nature and to discover their position in avian phylogeny (W.K. Parker, 1866; T.J. Parker, 1891; Pycraft, 1900; Mc Dowell, 1948; Hofer, 1945, 1950, 1955; de Beer, 1956; Webb, 1957; Muller, 1963; Bock, 1963; Cracraft, 1974; Yudin, 1970, 1978). I shall try to extract information on avian ancestry from the comparative and functional morphology of the feeding apparatus in paleognaths. Nomenclature for species' binomials and English names of modem birds follows Sibley and Monroe (1990). ACKNOWLEDGMENTS.—In the process of this work I received valuable assistance from the following individuals and Felix Y. Dzerzhinsky, Faculty of Biology, Moscow State University, Moscow 119899, Russia. Felix Y. Dzerzhinsky 267 institutions. F. Vuilleumier and A.V. Andors (American Museum of Natural History, New York, New York) and E.G. Kordicova (Institute of Zoology, Kazakh Academy of Sciences, Almaty, Kazakhstan) granted me access to the alcoholic specimen of Apteryx sp.; K.A. Yudin and V.M. Loskot (Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia) provided me with specimens of Eudromia and Casuarius and with several skulls of paleognaths, respectively. M.V. Bevolskaya (Institute of Cattle-Breeding Askania-Nova, Ukraine) permitted me to dissect the alcoholic heads of Dromaius novaehollandiae (Latham), Rhea americana (Linnaeus), and Struthio camelus (Linnaeus). A. Elzanowski (National Museum of Natural History, Smithsonian Institution, Washington, D.C.) and E.N. Kurochkin kindly assisted me in obtaining literature. A.N. Kuznetsov helped me in editing the manuscript and in translating it into English. S.C. Bennett, L.D. Martin, and S.L. Olson read the English version with fruitful criticism and helped me in editing it. I am sincerely grateful to all these persons. This work was supported by The Cultural Initiative Foundation, Moscow, and by The Russian Foundation of Basic Researches (RFBR, grant N 96-04-50822). Skeleto-muscular Consequences of Bill Formation The adductory force of the mandible is transferred to the upper jaw through a food object. Resistance of the upper jaw to this force is produced (Figure 1) by a combination of tension on the ventral stalk (premaxillary and maxillary bones with palate caudally) and longitudinal compression of the dorsal stalk (frontal projection of premaxillary and premaxillary processes of the nasal bones). The longer the jaw grew, the greater the forces became, and, due to jaw lightening, the stresses became ever greater. The active forces necessary for normal grasping of food items must be supplied by muscles. The muscular force that creates tension in the palate and upper-bill floor also can accomplish ventral movement of the upper jaw by means of re-
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»'' § "S^ iFjwtM . •- .m-i Avia
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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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Page 267 and 268: NUMBER 89 257 vical vertebrae of th
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Page 300 and 301: 290 1991). In this paper we use a "
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Early Evolution of Birds: Roundtabl
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338 evolved independently twice." M
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1 i'~L ,i "^ 1 •vw* HBB!/'' . m
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