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274 America. Science, 214(4526): 1236-1237. Iordanski, N.N. 1990. Evolution of Complex Adaptations: Feeding Apparatus in Amphibians and Reptiles. 310 pages, 65 figures. Moscow: Nauca. [In Russian.] Kuular, U.S., and F.Ya. Dzerzhinsky 1994. Trophical Adaptations of Cranes (Gruidae) as Seen from the Comparative and Functional Morphology of the Jaw Apparatus. 62 pages, 11 figures. Deposited Paper in VINITI, Russian Academy of Sciences, N 2904, Moscow State University, Moscow. [In Russian.] McDowell, S. 1948. The Bony Palate of Birds, Part 1: The Palaeognathae. Auk, 65(4):520-549. 1978. Homology Mapping of the Primitive Archosaurian Reptile Palate on the Palate of Birds. Evolutionary Theory, 4:81-94. Moller, W. 1930. Uber die Schnabel—und Zungenmechanik blutenbesuchender Vogel, Teil I. Biologia Generalis, 6:651-726. 1931. Uber die Schnabel—und Zungenmechanik blutenbesuchender Vogel, Teil II. Biologia Generalis, 7:99-154. Muller, H.J. 1963. Die Morphologie und Entwicklung des Craniums von Rhea americana Linne, II: Viszeralskelett, Mittelohr und Osteocranium. Zeitschrift fur die Wissenschaftliche Zoologie, 168(l/2):35-l 18. Parker, T.J. 1891. Observations on the Anatomy and Development of Apteryx. Philosophical Transactions of the Royal Society of London, series B, 182:25-134. Parker, W.K. 1866. On the Structure and Development of the Skull in the Ostrich Tribe. Philosophical Transactions of the Royal Society of London, SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY 156:113-183, 15 plates. Pycraft, W.P. 1900. On the Morphology and Phylogeny of the Palaeognathae (Ratitae and Crypturi) and Neognathae (Carinatae). Transactions of The Zoological Society, London, 15(5, 6): 149-290. Sibley, C.G., and B.L. Monroe 1990. Distribution and Taxonomy of Birds of the World. 1111 pages. New Haven: Yale University Press. Webb, M. 1957. The Ontogeny of the Cranial Bones, Cranial Peripheral and Cranial Parasympathetic Nerves, Together with a Study of the Visceral Muscles of Struthio. Acta Zoologica (Stockholm), 38:81-203. Yudin, K.A. 1961. On the Mechanismus of Mandible in Charadriiformes, Procellariiformes, and Some Other Birds. Trudy Zoologicheskogo Instituta Akademii Nauk SSSR [Proceedings of the Zoological Institute of the Academy of Sciences of USSR], 29:257-302, 30 figures. [In Russian.] 1970. Biological Significance and Evolution of the Cranial Kinetics in Birds. Trudy Zoologicheskogo Instituta Akademii Nauk SSSR [Proceedings of the Zoological Institute of the Academy of Sciences of USSR], 47:32-66. [In Russian.] 1978. Classical Morphological Characters and Recent Avian Systematics. Trudy Zoologicheskogo Instituta Akademii Nauk SSSR [Proceedings of the Zoological Institute of the Academy of Sciences of USSR], 76:3-8. [In Russian.] Zusi, R.L. 1984. A Functional and Evolutionary Analysis of Rhynchokinesis in Birds. Smithsonian Contributions to Zoology, 395: 40 pages, 20 figures.
The Relationships of the Early Cretaceous Ambiortus and Otogornis (Aves: Ambiortiformes) ABSTRACT Ambiortus from the Khurilt beds (Neocomian) of central Mongolia shows a combination of characters that confirms the assignment of this fossil to a separate order, Ambiortiformes. Otogornis genghisi Hou, 1994, of the Yijinhouluo Formation (earliest Cretaceous or latest Jurassic) of Ordos, China, was first described as Aves incertae sedis. Ambiortus and Otogornis share specialized characters, such as a thickened, three-edged acrocoracoid with an acute top; a flat, wide humeral articular facet of the scapula; ventral position of a small, oval humeral articular head; and a thin, long intermediate phalanx of the major wing digit. The generic status of Otogornis is supported by some other diagnostic characters. Several advanced characters demonstrate the assignment of Ambiortus and Otogornis to the Palaeognathae. These two forms show the occurrence of paleognathous birds in the Early Cretaceous of Central Asia. Introduction The Early Cretaceous Ambiortus dementjevi Kurochkin, 1982, was described as a member of the new family Ambiortidae and order Ambiortiformes, which was assigned to the infraclass Carinatae (Kurochkin, 1982). Ambiortus was based on an associated portion of the skeleton, including the vertebrae, the shoulder girdle, and some wing bones preserved on the main slab and counterslab, and also on the distal portion of the wing bones and feather imprints, which are preserved on an associated slab. Two papers containing more extensive description and comparison of this fossil were published later (Kurochkin, 1985a, 1985b). The surprising appearance of this Early Cretaceous tme bird in the paleontological record made comparison with other birds very difficult. In these first papers I attempted to compare Ambiortus with members of the living Gmidae, Rallidae, Strigidae, Alcedinidae, Momotidae, and, su- Evgeny N. Kurochkin, Paleontological Institute of the Russian Academy of Sciences, 123 Profsouznaya Street, 117868 Moscow, GSP-7, Russia. Evgeny N. Kurochkin 275 perficially, with the Archaeopterygiformes, Ichthyornithiformes, Paleogene paleognaths (later described as Lithornithiformes Houde, 1988), and Wyleyia Harrison and Walker, 1973. However, further study and a possibility of a direct comparison with the Enantiornithes, Archaeopteryx, Ichthyornis, Wyleyia, Paracathartes, Lithomis, Palaeotis, and living Palaeognathae, and also additional preparation of the holotype of Ambiortus, showed the published anatomical descriptions and morphological comparison of this fossil to be incomplete and partly erroneous. Evidence for the relationships of Ambiortus with the Palaeognathae was published beginning in 1985 (Kurochkin, 1985a, 1988, 1995a, 1995b). This mainly was based on comparison with the Paleogene Lithornithiformes studied by Houde and Olson (1981) and Houde (1988). Olson (1985) emphasized that Ambiortus shows some decided similarities with the Paleogene paleognathous birds and also may share some common characters of the humems with Ichthyornis. Martin (1987, 1991) united Ambiortus with Apatomis and the Ichthyomithiformes. Cracraft (1986) concluded that Ambiortus can be assigned to the Carinatae, in which he included the Palaeognathae, Neognathae, and Ichthyornis. Sereno and Rao (1992) have placed Ambiortus outside the Ornithurae without character evidence. Chiappe (1995) considered Ambiortus to be the oldest probable ornithurine, yet one of unclear relationships. Elzanowski (1995) assigned Ambiortus to the Carinatae and primitive Neornithes, close to Ichthyornis; however, his cladistic analysis of the skeletal characters also placed the Enantiornithes, Cathayomis, and Concomis among the Carinatae. Obviously, Ambiortus has nothing in common with the Enantiornithes. In that paper, Elzanowski (1995) made mainly mistaken interpretations of skeletal characters in Ambiortus that are discussed below. Feduccia (1995, 1996) placed Ambiortus in the basal Ornithurae together with Gansus, the Hesperornithiformes, and Ichthyornithiformes. Thus, controversial phylogenetic conclusions exist concerning Ambiortus, and it has remained isolated in the avian phylogenetic tree as the result of incomplete and questionable descriptions (Kurochkin, 1982, 1985a, 1985b), especially those
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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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16 Cor. Hum. Uln. Cpm. Fern. Tbt. T
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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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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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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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