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242 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY The ilia appear to be fully fused to the sacral vertebrae, as in all living Anseriformes, which makes the unfused innominates of Presbyornis seem all the more anomalous. PECTORAL APPENDAGE HUMERUS (Figure 8).—The humems is characterized by the extremely short, robust shaft with markedly sigmoid curvatures both anteroposteriorly and lateromedially. The total length of the bone is 33% shorter than in Anseranas, yet the shaft is slightly wider, indicating very different proportions and presumably a much different flight pattern. The pectoral crest is very broad and rounded, unlike Anseranas or the Anatidae, but is somewhat similar to Chauna. The head and internal tuberosity are massive, and the capital groove is extremely wide and deep, unlike other anseriforms. The orientation of the internal tuberosity may have been distorted by compression and perhaps pushed distally; regardless, it overhangs the bicipital fossa, which is small and may not have been pneumatic (obscured by pyrite). The capital ridge is very well defined. The distal end is expanded and flattened compared with most Anatidae but is generally similar to that of Anseranas except that the brachial depression is longer and narrower, the radial condyle is longer, and the olecranal and tricipital fossae are slightly deeper. The twisting of the shaft is remarkable because it appears as though the distal end has been rotated clockwise about the long axis of the bone by perhaps 15° or more. RADIUS AND ULNA.—In their incomplete condition there is little to be said about these elements. Compared with Anser, the fossil radius agrees with Anseranas in the more expanded distal end and more slender shaft, which is more sharply angular in cross section, with more flattened surfaces than in Anseranas. CARPOMETACARPUS (Figure 9).—This element agrees with that of Anseranas in being short and stout, although it is more robust even than that in Anseranas. It is intermediate in length between the male and female specimens compared. The alular metacarpal is more vertically oriented than in Anseranas and is more like that in the Anatidae. It also is much blunter, with a larger digital facet. ALAR PHALANGES (Figure 9).—Phalanx 1 of the major digit is relatively short and stout, as in Anseranas, but the proximal articulation is wider and not as deep. Phalanx 2 of the major digit and the minor digit are each relatively shorter, the former much more so, than in Anseranas. Discussion The two species of Anatalavis provide the only recognized occurrence of the family Anseranatidae in the fossil record. Anatalavis rex, from the Hornerstown Formation of New Jersey, whether Late Cretaceous or early Paleocene in age, also provides the earliest certain record of the entire order Anseriformes. The material of Anatalavis oxfordi, from the lower Eocene London Clay, is more complete than that of any early Tertiary anseriform yet described, apart from Presbyornis, and provides us with a new set of clues regarding early evolution in waterfowl. The skull in Anatalavis oxfordi indicates that it was most likely an obligate filter feeder. The bill is very broad yet is thin and weak. The retroarticular process of the mandible is quite well developed, although not nearly to the extent observable in the more extreme members of the Anatidae, yet the massive development of M. depressor mandibulae shows it to have been more adapted for straining, as opposed to grasping, in which the mandibular adductors play a greater role (Goodman and Fisher, 1962). This is practically the opposite of its nearest presumed relative, the Australian Magpie Goose {Anseranas semipalmata), in which the bill is strong, deep, and hooked and is used in digging out tubers and other plant material (Frith, 1967). Although the palate is very poorly preserved in the holotype of A. oxfordi, the fundamentally different morphology of the pterygoid compared with modem Anseriformes suggests that aspects of the organization and function of the skull in Anatalavis may have differed considerably from that in living waterfowl. In the shoulder girdle, the peculiar structure of the furcula and the pneumatic foramen in the dorsal surface of the sternal end of the coracoid are considered to be derived characters uniting Anatalavis and Anseranas in the family Anseranatidae. The two otherwise have very little else in common that is not generally present in most of the rest of the order. The proportionately short and very robust, twisted humems of Anatalavis is unique in the order and bespeaks a different mode of flight that probably was very strong and rapid. Many extant waterfowl are strong, fast fliers without having such a robust humems, however. The overall proportions of the humems are more like those of a falcon {Falco, Falconidae), although why a filter feeder would need such a wing is not easily envisioned. The pelvis of Anatalavis is likewise peculiar for an anseriform in the short, expanded anterior portions of the ilia, whereas in all other waterfowl, including the Anhimidae, the preacetabular part of the pelvis is longer and narrower. From its resemblance to such wading birds as ibises (Plataleidae) and other Ciconiiformes, the pelvis of Anatalavis is presumably primitive within the order. The innominate bones, however, are fused to the sacrum, which is a more derived condition also found in Ciconiiformes and other Anseriformes except Presbyornis. In the lack of fusion of the innominates, Presbyornis more nearly resembles the Charadriiformes. With the recognition of Anatalavis as a member of the Anseranatidae, we can trace each of the three major modem lineages of Anseriformes back to the early Eocene, or earlier in the case of Anatalavis. According to the phylogeny developed by Ericson (1997), the Presbyornithidae, which were probably world-wide in distribution in the early Tertiary, are on the lineage leading to the Anatidae proper. As yet unpublished early
NUMBER 89 243 Eocene records of the Anhimidae from Wyoming and England establish that this group was in existence at the same time and occurred outside of South America, the modem members of the family evidently being highly derived relicts. Anseranas in Australia likewise now appears to be a rather specialized relict of a once more diverse family Anseranatidae. Although the Anatidae proper probably existed in the Paleogene, they do not appear with any certainty or regularity in the Andors, Allison Victor 1988. Giant Groundbirds of North America (Aves, Diatrymidae). xvi+ 577 pages. Doctoral dissertation, Columbia University, New York. [Available from University Microfilms International, Ann Arbor, Michigan, order no. DA8815650.] 1992. Reappraisal of the Eocene Groundbird Diatryma (Aves: Anserimorphae). In Kenneth E. Campbell, Jr., editor, Papers in Avian Paleontology Honoring Pierce Brodkorb. Science Series, Natural History Museum of Los Angeles County, 36:109-125. Ericson, Per 1996. The Skeletal Evidence for a Sister-Group Relationship of Anseriform and Galliform Birds—A Critical Evaluation. Journal of Avian Biology, 27:195-202. 1997. Systematic Relationships of the Paleogene Family Presbyornithidae (Aves: Anseriformes). Zoological Journal of the Linnean Society, London,\2\A29-
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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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Page 211 and 212: NUMBER 89 201 lis hypotarsi along t
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Page 217 and 218: A New Genus and Species of the Fami
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Page 221 and 222: NUMBER 89 211 FIGURE 1.—Argornis
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Page 233 and 234: A Fossil Screamer (Anseriformes: An
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Page 245 and 246: NUMBER 89 235 mal was found under t
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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 298 and 299: 288 Palaeontological Institute. [Sp
Page 300 and 301: 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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