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118 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY over, they did so in much the same environment as is now there (Worthy, 1989). The dominant moa in subalpine sites is the Upland Moa {Megalapteryx didinus), with the Crested Moa {Pachyornis australis) the only other emeid. The dinomithids are most commonly represented by the Slender Moa {Dinornis struthoides) and the Large Bush Moa {D. novaezealandiae); the Giant Moa {D. giganteus) has never been found at these altitudes. Associated birds included Finsch's Duck {Euryanas finschi), Greatspotted Kiwi {Apteryx haastii), Little-spotted Kiwi {Apteryx owenii), New Zealand Coot {Fulica prisca), South Island Takahe {Porphyrio hochstetteri), Weka {Gallirallus australis), Eyles's Harrier {Circus eylesi), Haast's Eagle {Harpagornis moorei), New Zealand Falcon {Falco novaeseelandiae), Kakapo {Strigops habroptilus), Kea {Nestor notabilis), New Zealand Pipit {Anthus novaeseelandiae), Rock Wren {Xenicus gilviventris), and Stephens Island Wren {Traversia lyalli) (Worthy, 1989; unpublished data). Downslope from the subalpine zone there is a gradual change in the species composition of moa faunas. For example, in sites of Holocene age in northwest Nelson near Mt. Arthur, Megalapteryx didinus dominates assemblages between 700 m and 900 m, but Anomalopteryx didiformis also is present. Below 700 m, A. didiformis is the only emeid present. Similar altitudinal changes in species composition are known for Fiordland (Worthy, 1989). On Takaka Hill, the Holocene moa fauna is dominated by A. didiformis, whereas M. didinus is unknown (Worthy and Holdaway, 1994a). In deposits of the last glacial age, however, M. didinus is present, and A. didiformis is absent, a difference best explained as the result of altitudinal depression of the subalpine ecosystems. The deposits in Honeycomb Hill Cave, to the west, record a similar pattern: M. didinus and Pachyornis australis dominate deposits 14,000-20,000 years old. REGIONAL CHANGES IN AVIFAUNAS The most significant result of the recent studies of the South Island Quaternary avifaunas is that faunas from sites separated by as little as a few meters may differ markedly in species composition because of different ages. As a result, where in the past such associations were used as evidence for the coexistence of various species (e.g., Atkinson and Millener, 1991), they are now known to be the result of deposition at different times with markedly different environments. Graham and Lundelius (1984) expressed the opinion that most individual stratigraphic units are deposited over too short a time period for them to have accumulated through periods of environmental change. In New Zealand, unconformities separating deposits of glacial, late glacial, and Holocene age are the exception rather than the rule, and many sites have faunal remains essentially on the cave floor that range in age from modern to 20,000-30,000 years old, such as Hawkes Cave (Worthy and Holdaway, 1994a). Articulated skeletons of all ages indicate continuous deposition throughout this time. In the caves where many dates on individual bones are available, such as Madonna Cave (Worthy and Holdaway, 1993), Hawkes Cave, Kairuru Cave, Irvines Cave (Worthy and Holdaway, 1994a), and Honeycomb Hill Cave (Worthy, 1993a), the association of the moas Pachyornis elephantopus and Euryapteryx geranoides with Anomalopteryx didiformis is shown to be the result of deposition at different time periods. Many other undated talus accumulations beneath cave entrances have essentially unstratified deposits, with these same species found together, indicating that the deposits were accumulated and mixed over a significant time period, for example, Ngarua Cave and Commentary Cave (Worthy and Holdaway, 1994a). Graham (1993) described deposits such as these as time-averaged sequences and detailed numerous methods, with examples, whereby disharmonious associations could form by various time-averaging processes. In New Zealand, the factors that promote time-averaged sequences are constant humidity (deposits are always wet), low temperatures (most sites average
NUMBER 89 119 TABLE 3.—Lists of characteristic species of the Anomalopteryx and Euryapteryx assemblages that respectively characterize the closed-canopy wetter forests typical of western areas in the Holocene and the forestshrubland-grassland mosaics of the drier eastern areas, when these species occur together in abundance. Anomalopteryx assemblage Euryapteryx assemblage Anomalopteryx didiformis Dinornis novaezealandiae Apteryx australis Strigops habroptilus Anas chlorotis Gallirallus australis Callaeas cinerea Philesturnus carunculatus Pachyplichas jagmi/yaldwyni Xenicus longipes Petroica australis Euryapteryx geranoides Euryapteryx curtus Emeus crassus Dinornis giganteus Pachyornis mappini/elephantopus Pachyornis australis (uplands only) Megalapteryx didinus (uplands only) Cnem iorn is gracilis/calcitrans Euryanas finsch i Aptornis otidiformis/defossor Gallinula hodgenorum Fulica prisca Harpagornis moorei Coturnix novaezelandiae tered. Acanthisittid wrens of several species are abundant in deposits (when conditions of preservation permit), and the New Zealand Robin {Petroica australis) and Saddleback {Philesturnus carunculatus) are abundant. It is important to note that these species are found in faunas deposited under vegetational mosaics but are relatively rare and infrequent, in contrast to their numerical abundance and dominance of faunas from areas where the vegetation was a closed-canopy forest. It is therefore not presence or absence so much as relative frequency that is important in the definition of this assemblage. In contrast, the mere presence of species listed in the Euryapteryx assemblage indicates the presence of open habitat. NORTH CANTERBURY.—The Mt. Cookson study area occupies an intermediate zone between the wetter west and the drier east and, as expected, does not exhibit the same degree of faunal turnover. The drier climate of eastern areas is probably responsible for the delay of the establishment of closed-canopy forest until about 6000 years ago, much later than in the west. During the latter stages of oxygen isotope stage 3 and the early part of stage 2, Pachyornis elephantopus dominated moa faunas on Mt. Cookson but was associated with rare remains of Dinornis giganteus, D. struthoides, Megalapteryx didinus, Emeus crassus, and Euryapteryx geranoides. During the coldest period of the last glacial, however, P. elephantopus was virtually the only moa living in these and other eastern landscapes, where loess was being deposited. But, as in the west, with warming temperatures and forest establishment in the late Holocene, Anomalopteryx didiformis and Dinornis novaezealandiae came to dominate the faunas. Unlike in the west, however, Euryanas finschi and Aptornis defossor remained common in the late Holocene fauna, perhaps because the forests remained much drier. THE TYPICAL EASTERN FAUNAS.—In contrast to the above faunas, those of the lowlands in North and South Canterbury and North Otago provide no evidence for any faunal turnover in the period spanning the last glacial to the late Holocene. The greatest change detected is a reversal of dominance roles within the same suite of species: whereas Emeus crassus, Euryapteryx geranoides, and P. elephantopus were the main emeids throughout this time, Pachyornis elephantopus dominated glacial faunas, but Emeus crassus was the most common during the Holocene. Dinornis giganteus and D. struthoides were the dominant dinornithids during the glacial and the Holocene. In these eastern lowlands, there is no evidence of local extirpation of taxa at the end of the glacial period, as there is for western regions. The lack of glacial faunas containing smaller birds limits comparisons of faunal turnover to moas. Because there are no drastic changes in the moa faunas, however, and because Cnemiornis, Harpagornis, and Aptornis are known to have frequented the Otiran landscapes, and continued to do so in the Holocene, we can speculate on the composition of the associated fauna. These larger birds are part of the Euryapteryx assemblage that frequented Otiran western landscapes. So, as in the west, Euryanas finschi, Gallirallus australis, Nestor notabilis, New Zealand Crow {Corvus moriorum), New Zealand Pipit {Anthus novaeseelandiae), New Zealand Quail {Coturnix novaezelandiae), Piopio {Turnagra capensis), Fulica prisca, and Gallinula hodgenorum were probable associates of Cnemiornis, Harpagornis, and Aptornis in Canterbury. The relative abundance of species in late Holocene faunas of the east differs from that of western areas. This partly results from eastern areas having a much greater diversity resulting from the continued presence of all the presumed Otiran species during the Holocene, when other species typical of forest habitats became established. Such species include Kakapo {Strigops habroptilus), Philesturnus carunculatus, and Petroica australis. These species, however, are all relatively rare compared to their abundance in western faunas. Differences in the frequency of a species can usually be explained by the availability of the preferred habitat of that species. For example, the grassland inhabiting quail, Coturnix novaezelandiae, had little or no habitat in the west during the Holocene and is rare there, but in the east it is common. The most common passerine in late Holocene deposits in eastern areas was Turnagra capensis, which suggests that the preferred habitat of this extinct bird was the shrubland mosaics of drier areas. This is supported by the observation that Turnagra is present in fossil deposits of western areas only in Otiran deposits, when shrubland habitats were widely available. THE CENTRAL OTAGO FAUNAS.—The few data available for the Otiran fauna of central Otago indicate Pachyornis elephantopus and Dinornis giganteus were the most common moas, with Cnemiornis calcitrans being the only known carinate. The abundance of the last species in these deposits and in Otiran loess deposits in Canterbury illustrates its preference for the open, short shrubland/grassland habitats prevailing at that time. The Holocene moa fauna of central Otago is most similar to eastern ones, but it is influenced by altitude. In the broad valleys, Emeus crassus is common, and E. geranoides and P. ele-
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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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50 Si 5 15i 10- 5- 20 J 15 I 10 f 5
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168 SMITHSONIAN CONTRIBUTIONS TO PA
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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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