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264 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY minate on a prominent, raised linear scar 7 mm long, extending from within the tricipital fossa near the base of the ventral tubercle distally in the axis of the shaft. The scar may be the central vane of a partly pinnate M. humerotriceps. Similar, although shallower, striations occur in the tricipital fossa of the Western Gull, Larus occidentalis. A similar long scar is present in a slightly different position in several modem larids examined and in the Burhinidae. A short, lower crest or ridge extends from close to the proximal end of the long scar proximad and deeper into the triceps fossa. Dorsal to the longer scar there is a small, rough, irregular surface that may be the impression for M. scapulohumeralis cranialis. The caudal margin of the humems is very robust and distinct. It is far dorsal to the ventral tubercle. At the level of the base of the ventral tubercle the caudal margin bends abmptly dorsad to terminate at the proximal articular surface of the humerus about halfway between the apex and the dorsal tubercle. The area between the ventral tubercle and the caudal margin of the humems is broader than that in Graculavus velox. Discussion Cretaceous and Paleogene neornithines with the proximal end of the humems similar to that of Graculavus are Telmatornis, Presbyornis, Telmabates, and Zhylgaia. Olson and Parris (1987) compared Graculavus most closely to the Burhinidae in the Charadriiformes. Telmatomis Marsh, 1870, has been referred to the Charadriiformes (Shufeldt, 1915; Cracraft, 1972; Olson and Parris, 1987) and is most similar to the Scolopacidae. Presbyornis Wetmore, 1926, is referable to the Anseriformes based on associated cranial material (Olson and Feduccia, 1980). Telmabates Howard, 1955, evidently is referable to the Presbyornithidae (Feduccia and McGrew, 1974). Zhylgaia Nesov, 1988, was referred originally to the Presbyornithidae in Charadriiformes but later (Nesov, 1992) was referred, only tentatively, to the Presbyornithidae. Zhylgaia has a very steeply angled ventral tubercle (almost 90° to the axis of the shaft), a distinct although shallow impression for M. coracobrachialis cranialis, the head not much undercut, the capital incisure broad and shallow, and the caudal margin of the humems well defined but not dorsal to the ventral tubercle. These conditions suggest that it does not belong with either the Anseriformes or the Charadriiformes. Thus, the humems in all of these early waterbirds is very similar, but the birds are not all referable to the same modem higher-level taxon. In general, the proximal end of the humems in these birds differs from that of most modem Charadriiformes as follows: 1. Among the articular head and associated structures, the head is smaller, and the dorsal tubercle is farther from the head and is smaller but more protmdent. 2. Impressions for ligaments and muscles on the cranial surface are much less pronounced, including a shallow impression for M. coracobrachialis cranialis and a short sulcus for the transverse ligament. 3. Attachments of the muscles in and around the tricipital fossa are more robust, especially in Graculavus itself, but there is no dorsal (second) tricipital fossa. These differences suggest a distinctive flight mechanism. Graculavus, Telmatomis, Presbyornis, Telmabates, and Zhylgaia from the Late Cretaceous and the Paleogene are more similar to each other than to any modern bird. Many of the characters of Graculavus are not unique to the Charadriiformes but occur also among extant waterbirds in a mosaic pattern. The double-basined tricipital fossa occurs in virtually identical form in the Charadriiformes and the Oceanitidae (Procellariiformes). Extreme elongation of the ventral tubercle, very deeply undercut articular head, and very prominent caudal margin of the humerus are present in many Charadriiformes and in most Procellariiformes. A small articular head occurs in many Procellariiformes and in the Pelecaniformes. A small articular head, wide distance between the dorsal and ventral tubercles, and strong protmsion of the dorsal tubercle occur in some petrels, especially Calonectris (Procellariiformes), in Phaethon, and in some other pelecaniforms (see comparisons in Olson, 1977, figs. 18-20). Judged on varied morphological and behavioral grounds, Phaethon is highly plesiomorphic among Pelecaniformes (Cracraft, 1985; Elzanowski, 1995). This mosaic pattern of sharing characters with Graculavus suggests that the various extant groups of waterbirds have retained different suites of characters that were present in a common ancestor. The occurrence of many characters of Graculavus in the overall very plesiomorphic Phaethon supports this suggestion. Conclusions The new, very large species extends the Cretaceous range of Graculavus from eastern to western North America. Presbyornithids are now reported from the Late Cretaceous through the early Eocene in North and South America, Antarctica, and Mongolia (Howard, 1955; Feduccia and McGrew, 1974; Olson, 1994; Noriega and Tambussi, 1995). Zhylgaia comes from an estuarine habitat in the late Paleocene of Kazakhstan (Nesov, 1988). Graculavus, Telmatomis, Zhylgaia, and the Presbyornithidae show that similar graculavid-like birds were widespread in the Late Cretaceous through the early Tertiary. Reports of graculavid-like birds represented only by other parts of the skeleton are harder to evaluate in this context (e.g., Olson and Parris, 1987; Kurochkin, 1988; Nesov and Jarkov, 1989; Elzanowski and Brett-Surman, 1995). Graculavus augustus was a very large bird and a strong flyer. The bones were delicately sculpted and were not highly pneumatic, resembling contours in flying swimmers and divers; they lacked the extreme inflation of soaring birds. Impressions for tendons and muscles differ sufficiently from those of modem Charadriiformes to suggest distinctive flight mechanics. Graculavus is most similar to the Charadriiformes, but it shows a high proportion of characters that may be plesio-
NUMBER 89 265 morphic among them. Whatever their subsidiary affiliations, the similarity of the humems among these Late Cretaceous and Brodkorb, P. 1963. Birds from the Upper Cretaceous of Wyoming. In Charles G. Sibley, editor, Proceedings of the XIII International Ornithological Congress, Ithaca, 17-24 June, 1962, 1:55-70, 10 figures. Clemens, W.A. 1960. Stratigraphy of the Type Lance Formation. Report of the Twenty- First Session of the International Geological Congress, Norden, 1960, part 5:7-13. 1963. Fossil Mammals of the Type Lance Formation, Wyoming, Part I: Introduction and Multituberculata. University of California Publications in Geological Sciences, 48: 105 pages, 51 figures. Cracraft, J. 1972. A New Cretaceous Charadriiform Family. Auk, 89:36-46, 3 figures. 1985. Monophyly and Phylogenetic Relationships of the Pelecaniformes: A Numerical Cladistic Analysis. Auk, 102:834-853. de Queiroz, K., and J.A. Gauthier 1992. Phylogenetic Taxonomy. Annual Review of Ecology and Systematics, 23:449^180. Dorf, E. 1942. Flora of the Lance Formation at Its Type Locality, Niobrara County, Wyoming, 2: Upper Cretaceous Floras of the Rocky Mountain Region. Publications, Carnegie Institution of Washington, 508:83-144. Elzanowski, A. 1995. Cretaceous Birds and Avian Phylogeny. In D.S. Peters, editor, Acta Palaeomithologica: 3 Symposium SAPE: 5 Internationale Senckenberg-Konferenz, 22-26 Juni 1992. Courier Forschungsinstitut Senckenberg, 181:37-53. Elzanowski, A., and M.K. Brett-Surman 1995. Avian Premaxilla and Tarsometatarsus from the Uppermost Cretaceous of Montana. Auk, 112:762-766. Ericson, P.G.P. 1996. The Skeletal Evidence for a Sister-Group Relationship of Anseriform and Galliform Birds—A Critical Evaluation. Journal of Avian Biology, 27(3): 195-202. Estes, Richard 1964. Fossil Vertebrates from the Late Cretaceous Lance Formation, Eastern Wyoming. University of California Publications in Geological Sciences, 49: 180 pages, 73 figures, 5 plates. Feduccia, A., and P.O. McGrew 1974. A Flamingolike Wader from the Eocene of Wyoming. Contributions to Geology, University of Wyoming, 13(2):49-61, 13 figures. Gallagher, W.B. 1992. Geochemical Investigations of the Cretaceous/Tertiary Boundary in the Inversand Pit, Gloucester County, New Jersey. Bulletin of the New Jersey Academy of Science, 37:19-24. 1993. The Cretaceous/Tertiary Mass Extinction Event in the Northern Atlantic Coastal Plain. The Mosasaur, 5:75-154. Philadelphia: Delaware Valley Palaeontological Society. Gallagher, W.B., and D.C. Parris 1985. Biostratigraphic Succession Across the Cretaceous-Tertiary Boundary at the Inversand Company Pit, Sewell, N.J. In R. Talkington, editor, Geological Investigations of the Coastal Plain of Southern New Jersey: Second Annual Meeting of the Geological Society of New Jersey, Pomona, New Jersey, 1985, pages C1-C16. Literature Cited Paleogene waterbirds probably is due to recent common ancestry. Howard, H. 1955. A New Wading Bird from the Eocene of Patogonia. American Museum Novitates, 1710: 25 pages, 8 figures. Kennedy, W.J., and W.A. Cobban 1996. Maastrichtian Ammonites from the Hornerstown Formation in New Jersey. Journal of Paleontology, 70(5):798-804. Kurochkin, E.N. 1988. [Cretaceous Birds of Mongolia and Their Significance for Study of Phylogeny of Class Aves.] In E.N. Kurochkin, editor, Fossil Reptiles and Birds of Mongolia. Transactions of the Joint Soviet-Mongolian Palaeontological Expedition, 34:33-41. [In Russian.] Lillegraven, J.A., and M.C. McKenna 1986. Fossil Mammals from the "Mesaverde" Formation (Late Cretaceous, Judithian) of the Bighorn and Wind River Basins, Wyoming, with Definitions of Late Cretaceous North American Land- Mammal "Ages." American Museum Novitates, 2840:1-68, 13 figures. Marsh, O.C. 1870. Notice of Some Fossil Birds from the Cretaceous and Tertiary Formations of the United States. American Journal of Science, series 2, 49:205-217. 1872. Preliminary Description of Hesperornis regalis, with Notices of Four Other New Species of Cretaceous Birds. American Journal of Science, series 3, 3:360-365. 1889. Discovery of Cretaceous Mammalia. American Journal of Science, series 3, 38:81-92, 5 plates. 1892. Notes on Mesozoic Vertebrate Fossils. American Journal of Science, series 3, 55:171-175, 3 plates. Nesov, L.A. 1988. [New Cretaceous and Paleogene Birds of Soviet Middle Asia and Kazakhstan and Their Environments]. Proceedings of the Zoological Institute of Leningrad, 182:116—123, 2 plates. [In Russian, with English summary.] 1992. Mesozoic and Paleogene Birds of the USSR and Their Paleoenviroments. In K.E. Campbell, editor, Papers in Avian Paleontology Honoring Pierce Brodkorb; Proceedings of the II International Symposium of the Society of Avian Paleontology and Evolution. Science Series. Natural History Museum of Los Angeles County, 36:465^178. Nesov, L.A., and A.A. Jarkov 1989. [New Birds from the Mesozoic and Paleogene of the USSR and Some Remarks on the Phylogeny of Aves and Origin of Flight.] Proceedings of the Zoological Institute of Leningrad, 197:78-97, 2 plates. [In Russian, with English summary.] Noriega, J.I., and CP Tambussi 1995. A Late Cretaceous Presbyornithidae (Aves: Anseriformes) from Vega Island, Antarctic Peninsula: Paleobiogeographic Implications. Ameghiniana, 32(1):57—61. Olson, S.L. 1977. A Lower Eocene Frigate Bird from the Green River Formation of Wyoming (Pelecaniformes: Fregatidae). Smithsonian Contributions to Paleobiology, 35: 33 pages, 31 figures. 1985. The Fossil Record of Birds. In D.S. Farner, J.R. King, and K.C. Parkes, Avian Biology, 8:79-256, 11 figures. New York: Academic Press. 1994. A Giant Presbyornis (Aves: Anseriformes) and Other Birds from the
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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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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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Page 267 and 268: NUMBER 89 257 vical vertebrae of th
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Page 298 and 299: 288 Palaeontological Institute. [Sp
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Early Evolution of Birds: Roundtabl
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338 evolved independently twice." M
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