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258 DISCUSSION.—Considering that the distal end of this tarsometatarsus seems a strange mosaic, had the proximal end been preserved, it might well have resembled that of some other bird altogether, not to mention what the rest of the skeleton may have resembled. Because the preserved part of this bone is, however, a mosaic of exclusively charadriiform characters, the Graculavidae is the best present assignment for this specimen. This fragment would seem, at least in the absence of other Paleocene fossils of more modem aspect, to represent a bird older than the divergence time of any of the extant charadriiform families, as does Dakotornis. On body size, the present specimen could be referable to either of the monotypic genera Dakotornis or Graculavus. The specimens of Dakotornis cooperi are about one million years younger than this specimen, and those of Graculavus velox Marsh are about four million years older (see Olson, 1994, for the probable early Paleocene age of G. velox and the other New Jersey graculavids). GRACULAVIDAE, gen. et sp. indescript. FIGURE 4 MATERIAL.—Distal 6 mm of a right tarsometatarsus, with the outer trochlea broken away, SMM P96.9.3; collected by Bmce R. Erickson and field crew, summer 1976. LOCALITY.—North Dakota, Billings County, -10 mi (16 km) NW of Medora; Wannagan Creek Quarry, field map quadrant 0-19. HORIZON AND AGE.—Late Paleocene, early Tiffanian 4, Bullion Creek Formation, Wannagan Creek Quarry, Bed 2 (lignitic shale); absolute age, -60 Ma. DESCRIPTION.—This tarsometatarsal fragment is from a bird about the size of a Wilson's Plover {Charadrius wilsonia) and has the following characters: (1) the distal foramen is moderately large and oval and occupies the distal end of a shallow anterior tendinal groove; (2) in posterior view, two tiny foramina occur just proximal to the distal foramen; (3) the metatarsal facet is shallow and oval; (4) the trochlear arch in distal view is relatively low, with the outer trochlea not posteriorly retracted, but with the inner trochlea retracted so that its central point is posterior to the posteriormost extent of the middle trochlea; the trochlear arch in posterior view (as is common in Charadriiformes) forms a subsymmetrical basin between the three trochleae; (5) the inner trochlea is bulbous (as in most Charadriiformes) and is oriented distomedially; (6) the inner trochlea in distal view bears a very slight, posteriorly oriented wing on its medial side (a smaller wing than in modem Charadriiformes); (7) the inner trochlea is elevated so that its distal extent is about level with the proximal extent of the middle trochlea's digital groove; (8) the middle trochlea extends considerably farther distally than the other two; its lateral rim is slightly greater in distal extent and in anterior extent than is the medial rim; and (9) the outer trochlea would seem to have a slightly greater distal extent than the inner trochlea (the broken-off outer trochlea is preserved, but its contact with the adjacent part of the bone at the breakage is lost). SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY FIGURE 4.—Distal end of right tarsometatarsus of Graculavidae gen. et sp. indescript., SMM P96.9.3 (x8): a, anterior view; b, medial view; c, posterior view; d, distal view. COMPARISONS.—Fossil tarsometatarsi of small, spurless galliforms can be easily mistaken for those of charadriiforms (Olson and Farrand, 1974). Indeed, none of the nine characters listed in the previous paragraph would necessarily be inconsistent with an assignment of the specimen to Galliformes. Olson and Farrand (1974), however, list 10 other tarsometatarsal characters, all relating to the trochleae, that distinguish galliforms from charadriiforms. The present specimen differs from the Galliformes in all 10 of these trochlear characters. Specimen SMM P96.9.3 shares seven of the nine characters listed above (1-5, 8, 9) with the plover family, Charadriidae, a family not known in the fossil record earlier than the early Miocene (Unwin, 1993). The specimen differs from Charadriidae in two characters of the inner trochlea (6, 7), in which it more closely resembles the Paleocene Telmatomis and modem avocets (Recurvirostridae), respectively. It also resembles the Recurvirostridae in characters 3 and 9. DISCUSSION.—There is no evidence that the families Charadriidae and Recurvirostridae had diverged prior to the date of this fossil (Unwin, 1993), which may represent part of the "graculavid" stock prior to the divergence of these families. Of course, the presence of primitive charadriiform characters in this specimen (and in the Brisbane tarsometatarsus described above), perhaps by retention from much earlier forms, can tell us nothing certain about the divergence times of the extant charadriiform families. The very small body size of the bird represented by specimen P96.9.3, perhaps the smallest currently known Paleocene bird, need not exclude it from the Graculavidae (which are otherwise much larger birds), as this is only a form-family in any case
NUMBER 89 259 (Olson and Parris, 1987). The relative difference in size between Graculavus velox and the species represented by P96.9.3 Berggren, William A., Dennis V. Kent, John J. Flynn, and John A. Van Couvering 1985. Cenozoic Geochronology. Bulletin of the Geological Society of America, 96:1407-1418. Boles, Walter E., Henk Godthelp, Suzanne Hand, and Michael Archer 1994. Earliest Australian Non-marine Bird Assemblage from the Early Eocene Tingamurra Local Fauna, Murgon, Southeastern Queensland. Alcheringa. 18:70. Erickson, Bruce R. 1975. Dakotornis cooperi, a New Paleocene Bird from North Dakota. Scientific Publications of the Science Museum of Minnesota, 3(1): 7 pages. 1991. Flora and Fauna of the Wannagan Creek Quarry: Late Paleocene of North America. Scientific Publications of the Science Museum of Minnesota, 7(3): 19 pages. Feduccia, Alan, and Paul O. McGrew 1974. A Flamingolike Wader from the Eocene of Wyoming. Contributions to Geology of the University of Wyoming, 13:49-61. Holtzman, Richard C. 1978. Late Paleocene Mammals of the Tongue River Formation, Western North Dakota. Report of Investigations, North Dakota Geological Survey, 65: ix+88 pages. Kihm, Allen J. 1993. Late Paleocene Mammalian Biochronology of the Fort Union Group in North Dakota: Plesiadapis (Plesiadapiformes) from the Brisbane, Judson, and Wannagan Creek Quarry Local Faunas. In A.J. Kihm and J.H. Hartman, editors, The Marshall Lambert Symposium, pages 26-27. Bismarck: North Dakota Geological Society. Kihm, Allen J., and Joseph H. Hartman 1995. Bird Tracks from the Late Paleocene of North Dakota. Proceedings of the North Dakota Academy of Science, 49:63. Meehan, T.J., and Larry D. Martin 1994. Iterative Community Evolution in the North American Cenozoic. Literature Cited is comparable to that between the largest and smallest modem members of the sandpiper family, Scolopacidae. Journal of Vertebrate Paleontology, 14:3 8 A. Mourer-Chauvire, Cecile 1994. A Large Owl from the Paleocene of France. Palaeontology, 37:339-348. Olson, Storrs L. 1985. The Fossil Record of Birds. In Donald S. Farmer and James R. King, editors, Avian Biology, 8:79-238. New York, Academic Press. 1994. A Giant Presbyornis (Aves: Anseriformes) and Other Birds from the Paleocene Aquia Formation of Maryland and Virginia. Proceedings of the Biological Society of Washington, 107:429-435. Olson, Storrs L., and John Farrand, Jr. 1974. Rhegminornis Restudied: A Tiny Miocene Turkey. Wilson Bulletin, 86:114-120. Olson, Storrs L., and David C. Parris 1987. The Cretaceous Birds of New Jersey. Smithsonian Contributions to Paleobiology, 63: iii+22 pages. Peters, Dieter Stefan 1983. Die "Schnepfenralle" Rhynchaeites messelensis Wittich 1898 ist ein Ibis. Journal fur Ornithologie, 124:1-27. Sibley, Charles G., and Burt L. Monroe, Jr. 1990. Distribution and Taxonomy of Birds of the World. 1111 pages. New Haven: Yale University Press. Sloan, Robert E. 1987. Paleocene and Latest Cretaceous Mammal Ages, Biozones, Magnetozones, Rates of Sedimentation, and Evolution. Geological Society of America Special Paper, 209:165-200, 4 charts. Unwin, David M. 1993. Aves. In Michael J. Benton, editor, The Fossil Record 2, pages 717-737. London: Chapman and Hall. Wetmore, Alexander 1926. Fossil Birds from the Green River Deposits of Eastern Utah. Annals of the Carnegie Museum, 16:391-402, plates 36, 37.
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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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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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Page 298 and 299: 288 Palaeontological Institute. [Sp
Page 300 and 301: 290 1991). In this paper we use a "
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NUMBER 89 309 Jura-Museums, Eichsta
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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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