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282 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY LCR CTR PPR LRD LUL DMB FIGURE 7.—Otogornis genghisi Hou, 1994. Caudal view. ACR=acromion, APR=acrocoracoid process, CTR=carpal trochlea, DMB=damaged bones, HAF= humeral articular facet, IPH=intermediate phalanx of major wing digit, LCM=left carpometacarpus, LCR=left coracoid, LRD=left radius, LUL=left ulna, MAM=major metacarpal, MIM=minor metacarpal, PPR=procoracoid process, RHM=right humerus, RRD=right radius, RSC=right scapula, SGR=scapular groove, TRF=tricipital fossa, UPH=ungual phalanx of major wing digit, VTB= ventral tubercle. (Scale bar=lcm.) mentjevi, and they also look more robust. Comparison of the separate morphological characters, however, shows that these two Early Cretaceous birds have a close relationship. The char 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. Character Cervical vertebrae Ventral caudal transverse process Furcula Hypocleideum Acromion Acromial dorsal tubercle Coracoidal cotyla Acrocoracoid Scapular cotyla Humeral articular head Bicipital crest Pneumotricipital fossa Ventral edge, proximal end of humerus Cranial tubercle Transverse ligamental depression Intermediate phalanx of major wing digit acters and their conditions in Ambiortus, Otogornis, and some higher avian taxa are shown in Table 1. The Ichthyomithes and Neognathae are accepted as outgroups for determining polarity (advanced or primitive condition). The Enantiornithes are used only for general comparison because they have no close relationship with the Ornithurae. Ambiortus and Otogornis share the combination of the following characters: a thickened, three-edged acrocoracoid with an acute top (character 8); a flat, wide humeral articular facet of the scapula; ventral position of a small, short, and oval humeral articular head (character 10); and a long, thin intermediate phalanx of the major wing digit (character 16). These characters provide evidence for a close relationship between Ambiortus and Otogornis, and for the assignment of Otogornis to the Ambiortiformes. A convex coracoidal cotyla in the scapula (character 7) and concave scapular cotyla in the coracoid (character 9) unite Ambiortus and Otogornis with the Ornithurae. Two advanced characters support the assignment of Ambiortus to the Neornithes. These are the heterocoelous cervical vertebrae (character 1) and the U-shaped furcula (character 3). With the Palaeognathae, Ambiortus and Otogornis share the advanced condition of a projecting, dorsoventrally compressed scapular acromion (character 5) with a tubercle or prominence on its dorsal side (character 6); a projecting ventral edge of the humeral proximal end (character 13); and a remarkable cranial tubercle with a pit in the center of the cranial surface of this projecting edge (character 14). Ambiortus also shares with the Palaeognathae the strong, ventral, caudal transverse processes of the cervical vertebrae (character 2; unknown for Otogornis). Ambiortus and Otogornis also have a number of generalized characters that are common to the Palaeognathae and/or Lithornithiformes and are primitive in respect to the Neognathae. TABLE 1.—Distribution of some characters among Ambiortus, Otogornis, and other birds. Ambiortus heterocelous present U-shaped absent projecting, blunt present convex three-edged, acute concave small absent depression strongly projecting present fossa long, thin ? 7 7 ? Otogornis projecting present? convex three-edged, acute concave small absent depression strongly projecting present fossa long, thin Ichthyomithes amphicelous absent U-shaped absent attenuated absent convex rounded concave large absent tricipital fossa projecting absent absent short Neognathae heterocelous absent U, V-shaped absent or pesent attenuated or short absent convex rounded or elongate concave or flat large present fossa or foramen rounded or projecting absent furrow long, flat Palaeognathae heterocelous present U-shaped absent projecting present convex short, rounded concave small absent fossa or depression strongly projecting present? shallow furrow long, three-edged Lithomithiformes Enantiornithes heterocelous present U-shaped absent projecting, acute present convex rounded concave small absent fossa strongly projecting present fossa ? opisthocelous absent V-shaped present short absent concave stick-like boss small absent fossa rounded absent present long, flat
NUMBER 89 283 These are the absence of the bicipital crest and intumescence (character 11); the absence of the pneumatic foramen in the pneumotricipital fossa, being expressed only as an undivided tricipital depression (character 12); a small ventral tubercle on the proximal end of the humems; the presence of a ligamental fossa instead of a transversal ligamental furrow (character 15); a rounded cross-section of the shaft of the radius; and the presence of an ungual phalanx on the major wing digit. These characters demonstrate that the Ambiortiformes have a common origin with other orders of paleognathous birds. Otogornis differs from Ambiortus in having a smaller procoracoid process; a deep groove on the lateral side of the shoulder end of the scapula; a wide scapular blade (narrow in Ambiortus); a flat, elongated excavation along the cranial side of the deltopectoral crest; and the presence of a capital groove, which is divided into two furrows (Figures 6, 7). Differences in the detailed morphology of the scapula and humems support their separate generic status, although it could be argued that they are only two species of a single genus. Ambiortus dementjevi is smaller than Otogornis genghisi. The maximum width of the proximal end of the humems of A. dementjevi is 13.0 mm, and the maximum width across the most projecting edge of the deltopectoral crest is 11.2 mm. The same measurements in O. genghisi are 15.8 mm and 12.2 mm, respectively. Chen, Peiji, and Zhenlu Chang 1994. Nonmarine Cretaceous Stratigraphy of Eastern China. Cretaceous Research, 15:245-257, 4 figures. Chiappe, Luis M. 1995. First 85 Millions Years of Avian Evolution. Nature. 378:349-355, 6 figures. Cracraft, J. 1986. The Origin and Early Diversification of Birds. Paleobiology, 12:383-399, 1 table, 4 figures. Dmitriev, V.Yu., and V.V. Zherikhin 1988. [The Changes of Diversity in the Insect Families on the Data by the Method of Accumulated Occurrences.] In A.G. Ponomarenko, editor, Cretaceous Biocenotic Crisis and the Evolution of Insects, pages 208-215, 1 table, 1 figure. Moscow: Nauka. [In Russian.] Dong, Zhi-Ming 1993. A Lower Cretaceous Enantiornithine Bird from the Ordos Basin of Inner Mongolia, People's Republic of China. Canadian Journal of the Earth Sciences, 30:2177-2179, 1 table, 2 figures. Elzanowski, A. 1995. Cretaceous Birds and Avian Phylogeny. Courier Forschungsinstitut Senckenberg, 181:37-53, 4 figures. Fan, Jin 1996. New Advances in the Late Mesozoic Stratigraphic Research of Western Liaoning, China. Vertebrata Palasiatica, 34(2): 102-122, 3 tables. [In Chinese, with English summary.] Literature Cited Conclusions Ambiortus from central Mongolia and Otogornis from the Ordos Basin, China, show a close relationship based on the shared, specialized characters 8, 10, and 16 in the structure of the forelimb and shoulder girdle (Table 1). At the same time, Ambiortus and Otogornis show some differences in the shoulder girdle and the forelimb that support their separate generic status. The relationships of Ambiortus and Otogornis with other birds are determined by comparison with the Ichthyomithes, Neornithes, Palaeognathae, and Neognathae. Ambiortus and Otogornis share an advanced condition of characters 7 and 9, which are common to the Ornithurae. Ambiortus shares with the Neornithes an advanced condition of characters 1 and 3, which are unknown for Otogornis. At the same time, the Ambiortiformes share with the Palaeognathae (including Lithomithiformes) such specialized characters as 5, 6, 13, and 14, which suggests their assignment to the parvclass Palaeognathae, sensu Kurochkin (1995b). No common advanced characters were found for the Ambiortiformes, Ichthyomithes, and Enantiornithes. This study confirms that the Ambiortiformes are not closely related to the Ichthyomithes or the Neognathae and are totally unrelated to the Enantiornithes. The Early Cretaceous Ambiortiformes were flying palaeognathous birds. Thereby, they document an early diversification of ornithurine birds into two main evolutionary branches: Palaeognathae and Neognathae. Feduccia, Alan 1995. Explosive Evolution in Tertiary Birds and Mammals. Science, 267:637-638, 2 figures. 1996. Origin and Evolution of Birds, x+420 pages. New Haven and London: Yale University Press. Harland, W.B., R.L. Armstrong, A.V. Cox, L.E. Craig, A.G. Smith, and D.G. Smith 1989. A Geologic Time Scale 1989. xvi+263 pages. Cambridge: Cambridge University Press. Harrison, C.J.D., and CA. Walker 1973. Wyleyia: A New Bird Humerus from the Lower Cretaceous of England. Palaeontology, 16:721-728, 2 figures. Hou, Lianhai 1994. A Late Mesozoic Bird from Inner Mongolia. Vertebrata Palasiatica, 32(4):258-266, 3 figures, 1 plate. Houde, P. 1988. Paleognathous Birds from the Early Tertiary of the Northern Hemisphere. Publications of the Nuttall Ornithological Club, 22: vii+148 pages, 27 tables, 41 figures. Houde, P., and S.L. Olson 1981. Paleognathous Carinate Birds from the Early Tertiary of North America. Science, 214:1236-1237, 2 figures. Krassilov, V.A. 1982. Early Cretaceous Flora of Mongolia. Palaeontographica, series B, Paldophytologie, 181:1-43, 1 table, 11 figures, 20 plates.
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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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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 294 and 295: 284 Kurochkin, E.N. 1982. [New Orde
Page 298 and 299: 288 Palaeontological Institute. [Sp
Page 300 and 301: 290 1991). In this paper we use a "
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338 evolved independently twice." M
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