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204 is about 80% of the external tarsal length of females of Megapodius laperouse laperouse (50-55 mm), the smallest living species of megapode. The proximal and distal widths of the tarsometatarsus of Ngawupodius are proportionally great compared to its total length: about 21% for both measurements, at the top of the range among megapodes (15%-21%). Likewise, Ngawupodius has a proportionally wide middle shaft (10.6% of total length, range of megapodes 6%-ll%; 45% of distal width, 38%-45% in other species). In these proportions Ngawupodius resembles only Leipoa (Figure ID). Comparisons with Progura are not possible from existing material. The structure and length of the tarsometatarsus in Leipoa may be related to its more open habitat, and the resemblances to Ngawupodius could be coincidental. Given what is known of living species, a general range of size can be proposed for Ngawupodius. If it had disproportionally short legs, as in Leipoa, then it may have been similar in body length and weight to the smallest species of Megapodius. If the leg proportions were more typical of the other megapodes, then estimates of about 225-235 mm overall length and 230-330 g are reasonable. The small, undescribed species of Megapodius from 'Eua, Tonga, appears to have been of comparable size to Ngawupodius minya (D. Steadman, pers. comm., 1996). On the basis of pollen of grasses and subtropical rainforest flora from the lower member of the Namba Formation, Tedford et al. (1977:56) suggested a paleohabitat of "riparian forests with savannas on better drained fluviatiles." Some related but specifically distinct mammals in the faunas of the two members were inferred by Tedford et al. (1977) to indicate that the division between them recorded both a change in the depositional environment and a significant time gap. If the general vegetation in the upper member remained much the same as that suggested by the pollen from the lower member, then it is tempting to envisage a situation much like that in northern Australia today. The Orange-footed Scrubfowl, Megapodius reinwardt, frequents riparian forests bordered by tropical savanna woodland. Ngawupodius may have occupied a similar habitat in the Oligocene environment of Lake Pinpa. Balouet, Jean Christophe, and Storrs L. Olson 1989. Fossil Birds from Late Quaternary Deposits in New Caledonia. Smithsonian Contributions to Zoology, 469:38 pages, 16 figures. Baumel, Julian J., and Lawrence M. Witmer 1993. Osteologia. In J.J. Baumel, A.S. King, J.E. Breazile, H.E. Evans, and C. Vanden Berge, editors, Handbook of Avian Anatomy: Nomina Anatomica Avium. Publications of the Nuttall Ornithological Club, 23:45-132, figures 4.1—4.18. Boles, Walter E., and Brian Mackness 1994. Birds from the Bluff Downs Local Fauna, Allingham Formation. Literature Cited SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY The extinction of Ngawupodius was neither through competition with pheasants nor predation by felids. The only phasianids known from the Australian fossil record or occurring in a natural state in Australia today are small quail of the genus Coturnix (up to 120 g). These are too small to compete actively with megapodes, even Ngawupodius. Coturnix is known from other late-Oligocene-aged central Australian sites, although not Lake Pinpa. Pheasants, partridges, or other galliforms of a size comparable to any of the megapodes are not known from Australia. Likewise, the Felidae and Viverridae do not occur in Australia, and there is no evidence that marsupial carnivores have a particularly deleterious effect on megapode populations. Marsupial carnivores were well represented in Australia during the Oligocene and also have been found in the Ericmas Fauna (Dasyuridae). If it left no descendants, the eventual extinction of Ngawupodius may be related to a changing environment. With the late Miocene drying of Australia, the wet forest vegetation that apparently was its habitat was lost from the center of the continent. Similar habitats are now restricted to parts of eastern and northern Australia. Other central Australian birds that were lost after these climatic alterations were aquatic forms (e.g., flamingos, Phoenicopteridae). It is not yet possible to verify the loss of other terrestrial birds because these are not well represented, and existing material has yet to be studied in any depth. The morphology of Ngawupodius holds no clues to its relationships within the family; it shares osteological characters with most genera of megapodes. Until a complete phylogenetic analysis of the Megapodiidae is performed, the polarities of these characters will not be known. Likewise, whether Ngawupodius was ancestral to any of the living forms, and if so, how directly, cannot be determined. Similarities in proportions and its occurrence in central Australia raise the possibility that Ngawupodius may have been a direct ancestor of Leipoa. Conversely, Ngawupodius may have represented a distinct lineage within the family. There is general agreement that, regardless of their center of origin, megapodes were isolated in Australo-Papua for an extended period, which we know now to extend at least to the late Oligocene, Ngawupodius being the oldest known member of the Megapodiidae. Records of the South Australian Museum, 27(2): 139-149,2 figures. Brom, T.G., and R.W.R.J. Dekker 1992. Current Studies on Megapode Phylogeny. In R.W.R.J. Dekker and D.N. Jones, editors, Proceedings of the First International Megapode Symposium, Christchurch, New Zealand, December 1990. Zoologische Verhandelingen, 278:7-17, figures 1-5. Callen, Roger A., and Richard H. Tedford 1976. New Late Cainozoic Rock Units and Depositional Environments, Lake Frome Area, South Australia. Transactions of the Royal Society of South Australia, 100(3): 125-168, 24 figures.
NUMBER 89 205 Clark, George A., Jr. 1964a. Ontogeny and Evolution in the Megapodes (Aves: Galliformes). Postilla, 78: 37 pages, 11 figures. 1964b. Life Histories and the Evolution of Megapodes. The Living Bird, 3:149-167, 15 figures. Cracraft, Joel 1973. Continental Drift, Paleoclimatology, and the Evolution and Biogeography of Birds. Journal of Zoology, London, 169(4):455-545, 20 figures, del Hoyo, Josep, Andrew Elliot, and Jordi Sargatal, editors 1994. Handbook of the Birds of the World, Volume 2: New World Vultures to Guineafowl. 638 pages, 60 plates. Barcelona: Lynx Edicions. Dekker, Ren£ W.R.J. 1989. Predation and the Western Limits of Megapode Distribution (Megapodiidae; Aves). Journal of Biogeography, 16(4):317-321, 2 figures. Dekker, R.W.R.J., and T.G. Brom 1992. Megapode Phylogeny and the Interpretation of Incubation Strategies. In R.W.R.J. Dekker and D.N. Jones, editors, Proceedings of the First International Megapode Symposium, Christchurch, New Zealand, December 1990. Zoologische Verhandelingen, 278:19-31, figures 6-11. De Vis, C.W. 1888. Australian Ancestry of the Crowned Pigeon of New Guinea. Proceedings of the Royal Society of Queensland, 5(4): 127-131. 1889. Additions to the List of Fossil Birds. Proceedings of the Royal Society of Queensland, 6(l):55-58. Gaillard, C. 1908. Les oiseaux des Phosphorites du Quercy. Annates de I 'Universite de Lyon, new series, 1(23): 1-100. Jones, Darryl N., Rene W.R.J. Dekker, and Cees S. Roselaar 1995. The Megapodes. 262 pages, 8 plates. Oxford: Oxford University Press. Longman, H.A. 1945. Fossil Vertebrates from Gore Quarries. Memoirs of the Queensland Museum, 12(3): 164. Lydekker, Richard 1891. Catalogue of the Fossil Birds in the British Museum (Natural History). 368 pages, 75 figures. London: British Museum (Natural History). Marchant, Stephen, and Peter J. Higgins, editors 1993. Handbook of Australian, New Zealand and Antarctic Birds, Volume 2: Raptors to Lapwings. 984 pages, 68 plates. Melbourne: Oxford University Press. Mourer-Chauvire, Cecile 1982. Les oiseaux fossiles des Phosphorites du Quercy (Eocene superieur a Oligocene superieur): Implications paleobiogeographiques. In E. Buffetaut, P. Janvier, J.C. Rage, and P. Tassy, Phylogenie et Paleobiogeographie: Livre jubilaire en l'honneur de Robert Hoffstetter. Geobios, special memoir, 6:413-426, 3 figures. 1992. The Galliformes (Aves) from the Phosphorites du Quercy (France): Systematics and Biostratigraphy. In K.E. Campbell, Jr., editor, Papers in Avian Paleontology Honoring Pierce Brodkorb. Science Series, Natural History Museum of Los Angeles County, 36:67-95, 14 figures. Olson, Storrs L. 1980. The Significance of the Distribution of the Megapodiidae. Emu, 80(l):21-24, 1 figure. 1985. The Fossil Record of Birds. In D.S. Farner, J.R. King, and K.C. Parkes, editors, Avian Biology, 8:79-238, 11 figures. New York: Academic Press. Peter, J.M., and M.B. Peter 1993. Appendix III: Aboriginal Names. In Stephen Marchant and Peter J. Higgins, editors, Handbook of Australian, New Zealand and Ant arctic Birds, Volume 2: Raptors to Lapwings, pages 971-973. Melbourne: Oxford University Press. Peters, James Lee 1934. Family Megapodiidae. In Check-list of Birds of the World, 2:3-9. Cambridge: Harvard University Press. Poplin, Francois 1980. Sylviomis neocaledoniae n.g. n.sp.; ratite eteint de la Nouvelle Caledonie. Comptes Rendus de I'Academie des Sciences de Paris, series D, 290:489-492, 4 figures. Poplin, Francois, and Cecile Mourer-Chauvire 1985. Sylviomis neocaledoniae (Aves, Galliformes, Megapodiidae), oiseaux g6ant Eteint de l'ile de Pins (Nouvelle-Caledonie). Geobios,\%(\):13-91, 8 figures, 4 plates. Reed, A.W. 1977. Aboriginal Words and Place Names. 286 pages. Adelaide: Rigby. Rich, Thomas H., Michael Archer, Suzanne J. Hand, Henk Godthelp, Jeannette Muirhead, Neville S. Pledge, Timothy F. Flannery, Michael O. Woodburne, Judd A. Case, Richard H. Tedford, William D. Turnbull, Ernest L. Lundelius, Jr., Leallyn S.V. Rich, Michael J. Whitelaw, Anne Kemp, and Patricia V. Rich 1991. Australian Mesozoic and Tertiary Ten-estrial Mammal Localities. In P.V. Vickers-Rich, J.M. Monaghan, R.F. Baird, and T.H. Rich, editors, Vertebrate Palaeontology of Australasia, pages 1005-1070. Melbourne: Thomas Nelson. Roselaar, CS. 1994. Systematic Notes on Megapodiidae (Aves, Galliformes), Including the Description of Five New Subspecies. Bulletin, Zoologisch Museum, Universiteit van Amsterdam, 14(2):9-36, 8 figures. Sibley, Charles G., and Jon E. Ahlquist 1990. Phylogeny and Classification of Birds: A Study in Molecular Evolution. 976 pages, 385 figures. New Haven: Yale University Press. Sibley, Charles G., and Burt L. Monroe, Jr. 1990. Distribution and Taxonomy of Birds of the World. 1111 pages. New Haven: Yale University Press. Steadman, David W. 1980. A Review of the Osteology and Paleontology of Turkeys (Aves: Meleagridinae). In K.E. Campbell, Jr., editor, Papers in Avian Paleontology Honoring Hildegard Howard. Contributions in Science, Natural History Museum of Los Angeles County, 330:131-207, 14 figures. 1989. New Species and Records of Birds (Aves: Megapodiidae, Columbidae) from an Archaeological Site on Lifuka, Tonga. Proceedings of the Biological Society of Washington, 102(3):537—552, 7 figures. 1993a. Biogeography of Tongan Birds Before and After Human Impact. Proceedings of the National Academy of Science of the United States of America, 90:818-822, 1 figure. 1993b. Bird Bones from the To'aga Site: Prehistoric Loss of Seabirds and Megapodes. In P.V. Kirch and T.L. Hunt, editors, The To'aga Site: Three Millennia of Polynesian Occupation in the Manu'a Islands, American Samoa. Contributions of the University of California Archaeological Research Facility, Berkeley, 51:217-228, 2 figures. 1995. Prehistoric Extinctions of Pacific Island Birds: Biodiversity Meets Zooarchaeology. Science, 267:1123-1131,4 figures. Tedford, Richard H., Michael Archer, Allan Bartholomai, Michael Plane, Neville S. Pledge, Thomas H. Rich, Patricia Rich, and Rod T. Wells 1977. The Discovery of Miocene Vertebrates, Lake Frome Area, South Australia. BMR Journal of Australian Geology and Geophysics, 2:53-57, 3 figures, van Tets, G.F. 1974. A Revision of the Fossil Megapodiidae (Aves), Including a Description of a New Species of Progura De Vis. Transactions of the Royal Society of South Australia, 98(4):213-224, 5 figures. 1984. A Checklist of Extinct Fossil Australasian Birds. In M. Archer and G. Clayton, editors, Vertebrate Zoogeography and Evolution in
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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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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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Page 165 and 166: NUMBER 89 155 the period studied. T
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Page 209 and 210: A New Genus of Dwarf Megapode (Gall
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Page 241 and 242: The Anseriform Relationships of Ana
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NUMBER 89 255 FIGURE 1.—Referred
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NUMBER 89 257 vical vertebrae of th
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NUMBER 89 259 (Olson and Parris, 19
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274 America. Science, 214(4526): 12
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276 concerning the amphicoelous str
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278 ment of the radius that are con
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280 The left coracoid is represente
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284 Kurochkin, E.N. 1982. [New Orde
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288 Palaeontological Institute. [Sp
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290 1991). In this paper we use a "
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Implantation and Replacement of Bir
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NUMBER 89 297 saurs (Figure 1E-G).
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NUMBER 89 299 would seem unlikely a
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Humeral Rotation and Wrist Supinati
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NUMBER 89 303 apparent. It is now c
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NUMBER 89 305 FIGURE 3.—Dorsal vi
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NUMBER 89 307 FIGURE 4.—Wing of t
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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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