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56 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY radiocarpalis (not so in Pelagodroma). Humeri of P. grallaria are longer, and the tuberculum dorsale is broader, not as raised above the margo caudalis, and not as elongate distally as those of P. marina. In F. grallaria the tuberculum ventrale is as high as wide rather than higher than wide as in P. marina. On the cranial surface, the sulcus at the base of the crista deltopectoralis is shallower than in P. marina. The sulcus ligamentis transversus ends ventrally beside a shallow sulcus on the ventral margin of the crista bicipitalis in F. grallaria, but not in P. marina, where there is no shallow sulcus. Oceanites sp. A few bones of a very small storm-petrel are smaller than those found in any genus except Oceanites. Comparisons were made with Grey-backed Storm-petrels {Oceanites nereis) and Wilson's Storm-petrels {O oceanicus). The few measurements available suggest O nereis is smaller, but although lengths of the tarsometatarsi overlap {O. nereis, mean=33.47 mm, range=31.1-34.6 mm, n=4; O oceanicus, mean=35.21 mm, range=33.0-37.2 mm, n=5), those of the ulnae do not {O. nereis, mean= 17.67 mm, range= 16.7-18.4 mm, «=4; O oceanicus, mean= 19.73 mm, range= 19.1-20.4 mm, n-5). Ulnae of two of the fossil specimens are 18.1 mm and 18.7 mm long, suggesting that they may belong to the smaller taxon. Oceanites oceanicus is regularly seen offshore, and a specimen of O o. parvus was collected ashore on Amsterdam (Roux and Martinez, 1987). In 1995 a small colony of O. oceanicus was found breeding on St. Paul (Micol, 1995). Because the specimens of O. oceanicus measured were mainly from Heard Island, they may be bigger than birds breeding on more northern islands (see discussion in Marchant and Higgins, 1990), so it is possible that O oceanicus from the Amsterdam group could be similar in size to O nereis. The fossils are referred only to Oceanites sp. Pelecanoides urinatrix Bones of a diving-petrel are present in many sites. At present, two diving-petrels are found in the Indian Ocean. The smaller South Georgian Diving-petrel {Pelecanoides georgicus) breeds on subantarctic islands, and the larger Common Diving-petrel (P. urinatrix) has a more widespread distribution, between 35°S and 55°S (Marchant and Higgins, 1990). Both exhibit size variation that is to some extent clinal, with southern populations being larger (Marchant and Higgins, 1990), but this is not the case in New Zealand (Appendix 7). The fossils, particularly the ulnae and carpometacarpi, are larger than bones of P georgicus from Heard Island, smaller than bones of P. urinatrix exsul from both Auckland and Heard islands, and smaller than bones of P. u. urinatrix from beaches around Cook Strait in New Zealand (Appendix 7). They are, however, similar in size to P. u. chathamensis from Southeast Island in the Chatham Group, and so they are referred to P. uri natrix. There are two records of P. urinatrix from Amsterdam (Roux and Martinez, 1987; Micol and Jouventin, 1995). Catharacta skua Skua bones were recovered from several sites, but most were in poor condition. They are much smaller than those of the Subantarctic (Brown or Southern Great) Skua {Catharacta skua lonnbergi) from the New Zealand subantarctic islands and are bigger than than those of the Antarctic Skua (C. maccormicki) (Appendix 8). The carpometacarpus is longer than the tarsometatarsus in C. maccormicki, in contrast to C. skua lonnbergi, in which the opposite is true. Unfortunately, none of the fossils were complete enough to use this feature. The Tristan Skua (C. skua hamiltoni), however, for which no comparative material was available, is smaller than C. skua lonnbergi and breeds on Gough Island and in small numbers on Amsterdam (Micol, 1995). Because the fossils are markedly smaller than bones of C. skua lonnbergi but are bigger than bones of C. maccormicki, it seems probable that the Amsterdam skua fossils are referable to C. skua hamiltoni. Anas marecula The bones of a small duck found in the collection were obviously missed when duck material was extracted for the study that resulted in the description of a new species, Anas marecula, by Olson and Jouvenin (1996) (see Table 1). The series they analyzed, now in the USNM, was composed of at least 33 individuals. Those listed herein are almost certainly parts of the same individuals. Discussion COMPOSITION OF THE FOSSIL FAUNA We consider some records based on fossils from Amsterdam Island to be of questionable validity; these are as follows: 1. Wandering Albatross {Diomedea exulans). The record of this species is of bones of a single individual identified by Jouanin and Paulian (1960) before D. amsterdamensis was described, and which Jouventin et al. (1989) reported to be of similar size to the bones they referred to D. amsterdamensis. 2. Kermadec Petrel {Pterodroma neglecta). Jouanin and Paulian (1960) identified this species from a few bones that were smaller than those herein described of Pterodroma macroptera and bigger than those of P. mollis. The given lengths for the bones are in the size range of P. baraui and P. arminjoniana. Pterodroma baraui was only described in 1964, and neither of these two species was compared to the fossils. Pending reexamination of the bones, the record of P. neglecta from Amsterdam is suspect. 3. Broad-billed Prion {Pachyptila vittata). Jouanin and Paulian (1960) identified the prion bones they had as this species and thought it probable that they were of the subspecies P.
NUMBER 89 57 vittata macgillivrayi. Data herein show that P. vittata is distinct from P. macgillivrayi, and both are distinct from P. salvini, so the Amsterdam prion should not be listed as Pachyptila vittata. The 20 species of seabird recorded herein as fossils from Amsterdam Island underestimate the total because the Sooty Albatross {Phoebetria fusca) and the Antarctic Tem {Sterna vittata) presently breed on Amsterdam (Jouventin et al.,1984) but are not represented among the fossils. Micol and Jouventin (1995) noted that at least one Phoebetria fusca had been identified in fossil material, but no material was seen by THW to substantiate this. Jouventin (1994) and Micol and Jouventin (1995) reported the supposed presence of two extinct species of Pterodroma and two extinct storm-petrels. This study finds no evidence for any extinct procellarid having previously existed on Amsterdam Island. Procellaria cinerea, Pterodroma macroptera, and P. mollis are common as fossils and undoubtedly bred there. The large number of P. baraui bones from site 8 suggests this species also was breeding on Amsterdam; in contrast, the few bones of P. arminjoniana could be from nonbreeding visitors, or possibly from skua kills. SIZE RANGES OF SPECIES Pterodroma macroptera: Several species of seabirds represented in the fossil fauna of Amsterdam Island are smaller than conspecific populations in the New Zealand region. Data in Appendix 1 show that P. macroptera from Amsterdam are smaller than Australasian and South Atlantic specimens. The specimens available from Eclipse Island in Western Australia are of similar size to the Amsterdam specimens, but the one specimen from Coffin Island is larger than any Amsterdam specimen and is within the size range for P. macroptera gouldi. Measurements given in Marchant and Higgins (1990) show that P. m. macroptera is a little smaller than P. m. gouldi. They also show that males are slightly larger than females in most measurements, but there is no detectable dimorphism in the fossil sample, which has apparently normal, unimodal, size distributions. Pterodroma mollis: The limited evidence suggests that P. mollis from Amsterdam is smaller than Australasian and Atlantic birds. Measurements in Marchant and Higgins (1990), however, indicate no geographical or sexual size variation. The fossil samples have mainly unimodal size distributions (Figure 8) except for humeri, where the distribution is bimodal, suggesting that there may be some sexual dimorphism. Procellaria cinerea: The Amsterdam Procellaria cinerea have bone lengths 4%-6% smaller than birds from the New Zealand region. Measurements of birds from the Crozet and Kerguelen islands and from New Zealand (Marchant and Higgins, 1990) also indicate that Indian Ocean birds are smaller than New Zealand ones. Some sexual dimorphism is apparent in external measurements in Marchant and Higgins (1990), although their two data sets are contradictory: in one the males are larger and in the other the females are larger. Measurements of sexed skeletons in our comparative series (3 males, 7 females) suggest that females are smaller, although the small sample size precludes meaningful statistical comparison. The fossil bones have a length distribution (Figure 12) that trends in most cases toward bimodality, compared to the unimodal (apparently normal) distributions of Pterodroma macroptera and Pachyptila macgillivrayi (Figures 7, 10). If the modem sample is representative, then the larger bones probably represent males. Pachyptila macgillivrayi: The differences detailed above show that the Amsterdam prion is a distinct species, P. macgillivrayi, that is endemic to the Amsterdam-St. Paul group. The very large samples of long bones are apparently normally distributed and unimodal, suggesting there is no sexual size dimorphism. Sexual dimorphism in prions is generally slight, although Genevois and Bretagnolle (1995) found male Thin-billed Prions (P. belcheri) to be larger overall and to have larger bills than do females. Puffinus assimilis: There are several subspecies of P. assimilis in the southern oceans region. The nominate race from Norfolk Island (P. assimilis assimilis) is the smallest (Appendix 5). The two New Zealand subspecies, P. a. haurakiensis and P. a. kermadecensis, are of similar size and are larger than P. a. assimilis but are smaller than P. a. elegans, from farther south in the Antipodes (Appendix 5). None of the beach-cast specimens from New Zealand west-coast beaches are large enough to be P. a. elegans, and they are considered to be P. a. kermadecensis (J.A. Bartle, pers. comm., 18 June 1995; verified in many cases by the species of lice present). Size variation therefore appears to be clinal, with larger birds in the south. Puffinus assimilis bones from Amsterdam are on average slightly smaller than those of P. a. elegans from the Antipodes but are much larger than those of the other subspecies considered above, and so they are probably referable to P. assimilis elegans. This subspecies ranges from the Antipodes and Chatham islands in the New Zealand region to Tristan da Cunha and Gough Island (Marchant and Higgins, 1990). Pelecanoides urinatrix: As discussed above, there is considerable variation in mean size of individuals between populations in this species. Although some of the variation may be clinal, with more southern populations comprising larger individuals, this is not the case in New Zealand, where populations in Cook Strait and farther north are distinctly larger than those around Stewart Island and on the Chatham Islands. The limited data presented herein suggest that the Amsterdam birds are smaller than those from Heard Island, significantly smaller than those from the lower latitude populations in the Cook Strait-Wellington region of New Zealand, and similar to those from slightly higher latitudes on Southeast Island, in the Chatham Group. Size variation in Pelecanoides is thus not well explained by clinal factors.
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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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Page 44 and 45: 34 ability in such a way that it ca
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Page 54 and 55: Site 13 Research Base ENTRANCE Lowe
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Page 81 and 82: NUMBER 89 71 Mallorca, probably in
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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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A Fossil Screamer (Anseriformes: An
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NUMBER 89 FIGURE 3.—Chaunoides an
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NUMBER 89 227 B C D FIGURE 6.—The
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NUMBER 89 229 FIGURE 9.—Right tib
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The Anseriform Relationships of Ana
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NUMBER 89 233 Subfamily ANATALAVINA
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NUMBER 89 235 mal was found under t
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NUMBER 89 237 tion, with retroartic
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NUMBER 89 FIGURE 7.—Sternum and p
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NUMBER 89 241 der. The bone is very
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NUMBER 89 243 Eocene records of the
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Presbyornis isoni and Other Late Pa
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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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