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256 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY ca, although Olson (1994) noted that the large-bodied P. isoni might well be assignable to a different genus were more of its skeleton known. The individual represented by SMM P96.9.2 was slightly larger than the one represented by the holotype of Presbyornis isoni (see Table 1). Their relative difference in size is not greater than that often seen within avian species (pers. obs.), including Presbyornis pervetus (P.G.P. Ericson, pers. comm., 1996). Order CHARADRIIFORMES Form-Family GRACULAVIDAE Dakotornis cooperi Erickson, 1975 FIGURE 2 MATERIAL.—Holotype: A complete right humems, SMM P74.24.106 (figured in Erickson, 1975), collected by Bmce R. Erickson and field crew, July 1974. Referred Material: A complete right humems, SMM P75.22.7 (Figure 2), collected by Bmce R. Erickson, summer 1975; a mostly complete left tibiotarsus, SMM P75.22.25 (not figured), collected by Tim Mc- Cutcheon, 6 July 1975. LOCALITY.—North Dakota, Billings County, -10 mi (16 km) NW of Medora, Wannagan Creek Quarry. Field map quadrants: holotype humems, G-5; other humems, G-2; tibiotarsus, K-6 (quadrants are 5 ft (1.5 m) on a side). HORIZON AND AGE.—Late Paleocene, early Tiffanian 4, Bullion Creek Formation, Wannagan Creek Quarry, Bed 2 (lignitic shale); absolute age, -60 Ma. DISCUSSION.—Dakotornis cooperi Erickson is the only bird from Wannagan Creek to have been previously published. It was originally described as a "primitive ibis-like bird" representing the extinct family Dakotomithidae within the suborder Plataleae, although its resemblance to thick-knees was mentioned (Erickson, 1975). Although the humems of Dakotornis shares numerous characters with the Plataleidae (ibises), especially in its stoutness and general outline, Dakotornis more closely resembles the Graculavidae, as Olson and Parris (1987) pointed out, and even some of the modem charadriiforms such as Burhinidae (thick-knees), Haematopodidae (oystercatchers), Recurvirostridae (avocets), Laridae (gulls), and Scolopacidae (godwits, phalaropes, etc.). These charadriiforms resemble Dakotornis in having a shallow proximal end of the humems that is not pneumatic, a median crest at a nearly right angle to the shaft, and a well-developed central ridge. In all of these characters these modem charadriiforms and Dakotornis differ from ibises. In two other characters of the humems—the proximally produced external tuberosity and the lack of a typically charadriiform ectepicondylar spur—Dakotornis is as similar to thick-knees as to ibises. The rounded ectepicondylar prominence of Dakotornis closely resembles that of Burhinus, whereas in other charadriiforms a spur extends from the proximal rim of this prominence. Dakotornis would thus seem to be a good charadriiform. Peters (1983) and Olson (1985) noted the similarities between ibises and charadriiforms (and gruiforms). The two known humeri of Dakotornis cooperi, holotype and referred, discovered in different field seasons, have until now been separated from each other through loans to different persons; I am the first person to see both original specimens side by side. The referred humerus (Figure 2) agrees in all characters with the holotype. The only apparent difference is that the external tuberosity of the holotype (as figured in Erickson, 1975) appears much narrower and sharper than the broad, rounded external tuberosity of the referred humems. The apparent narrowness of the tuberosity in the holotype, however, is due only to breakage. These two right humeri of Dakotornis cooperi are of nearly identical size; the holotype is 87.1 mm long, and the other is 89.0 mm long. -^» :'/;•• 4 FIGURE 2.—Referred right humerus of Dakotornis cooperi, SMM P75.22.7 (x 1), anconal view. The tibiotarsus (SMM P75.22.25) was tentatively assigned to Dakotornis cooperi on the basis of size and provenance. Olson and Parris (1987) have noted the graculavid nature of this spec- GRACULAVIDAE, gen. et sp. probabiliter indescript. MATERIAL.—A mostly complete third cervical vertebra, 18.5 mm long, -15 mm wide (both anteriorly and posteriorly), SMM P77.7.159 (not figured); collected by Richard C. Holtzman, summer 1977. LOCALITY.—North Dakota, Morton County, -8 mi (13 km) S of Judson; Judson Locality. HORIZON AND AGE.—Late Paleocene, early Tiffanian 4, Bullion Creek Formation, Judson Locality; absolute age, -60 Ma. COMPARISONS.—This specimen is referred to the Graculavidae due to its Paleocene age and its close resemblance to modem charadriiform cervical vertebrae. This cervical vertebra is identified as the third on the grounds of its caudally oriented, long-bottomed cariniform hypapophysis, gracile neural spine, and reduced pleurapophyses. The third cervical is typically a short vertebra, unlike this specimen, which is 1.2 times longer than its width, although this degree of elongation is common among charadriiforms. In its general outline and proportions between its parts, the specimen most closely resembles the cer-
NUMBER 89 257 vical vertebrae of the Charadriiformes. The shapes of the facets of the zygapophyses, of the articulations of the centmm, of the ventral pit anterior to the keel, and of other features are like those of most Charadriiformes but are more like recurvirostrids than like burhinids. The unusual feature of its relatively narrow posterior width (which is not appreciably greater than its anterior width) also is found in oystercatchers (Haematopodidae). Another striking feature is the transverse perforation of the centrum. This condition is observed in some of the more caudad cervical vertebrae (but not necessarily in the craniad vertebrae, such as the third) in avocets (Recurvirostridae), gulls (Laridae), skimmers (Rynchopidae), and sandpipers (Scolopacidae). Transverse perforation of the centmm is fairly common in Anseriformes, and in Dendrocygna bicolor even the third cervical is perforated; however, the specimen does not otherwise appear anseriform. DISCUSSION.—Although this specimen most likely represents an unknown species, the size of the vertebra would seem to fall within the upper limits of probable size for Dakotornis cooperi, even if it is somewhat large for a charadriiform body plan. A possible model for the body proportions of the very stout-winged Dakotornis might be the Canvasback {Aythya valisineria), in which the absolute and relative sizes of the humems, vertebrae, tibiotarsus, and tarsometatarsus are very similar to those of the humems of Dakotornis, the present vertebra, the Wannagan Creek tibiotarsus, and the tarsometatarsus described below. GRACULAVIDAE, gen. et sp. probabiliter indescript. FIGURE 3 MATERIAL.—Distal 20 mm of a left tarsometatarsus, lacking most of the outer trochlea, SMM P77.8.210 (Figure 3); collected by Richard C. Holtzman, summer 1977. LOCALITY.—North Dakota, Grant County, -5 mi (8 km) W of Raleigh; Brisbane Locality. HORIZON AND AGE.—Late Paleocene, early Tiffanian 3A, Slope Formation, Brisbane Locality; absolute age, -61 Ma. COMPARISONS.—This specimen is referred to the Graculavidae on the basis of its general resemblance to the tarsometatarsus of charadriiforms, including that of Telmatomis priscus, the only North American graculavid for which the tarsometatarsus was previously known (although T. priscus was a much smaller bird). The posterior surface of the specimen displays what might be called a typical "charadriiform basin," i.e., a deep, fairly symmetrical depression bounded by the longitudinal ridges of the inner and outer trochleae and by the proximal rim of the middle trochlea. The specimen resembles Telmatomis in the following: the distal foramen is moderately large and oval, the metatarsal facet is well developed, the inner trochlea is oriented distomedially, and the inner trochlea is elevated so that its distalmost extent is just proximal to the halfheight of the middle trochlea's digital groove. The inner and outer trochleae, however, are considerably more posteriorly re FIGURE 3.—Distal end of left tarsometatarsus of Graculavidae gen. et sp. prob. indescript., SMM P77.8.210 (x3): a, anterior view; b, medial view; c, posterior view; d, distal view. tracted than in Telmatomis, so that the tarsometatarsus has a greater arch in distal view. The present specimen differs from the Presbyornithidae in having the metatarsal facet oval and distinct rather than long and weak, a relatively smaller distal foramen, and a lesser trochlear arch in distal view. Characters of SMM P77.8.210 resemble those of a number of extant charadriiform families in a mosaic manner. Olson and Parris (1987) noted skua-like (and presbyomithid-like) features in the tibiotarsus of another Paleocene graculavid, Laornis edvardsianus Marsh. SMM P77.8.210 combines tarsometatarsal characters especially of gulls (Laridae), skuas (Stercorariidae), and oystercatchers (Haematopodidae), among others. The general outline of the bone is oystercatcher-like, except that the orientation of the inner trochlea is gull-like, being neither as distally oriented as in oystercatchers nor as medially oriented as in skuas. Its moderately large distal foramen matches that of oystercatchers, unlike the larger foramen in gulls and skuas. The metatarsal facet is skua-like (as seen in Catharacta maccormicki) in both configuration and position (rather than lacking a hind toe as in oystercatchers). The degree of trochlear arch in distal view matches that of gulls or skuas, unlike the very highly arched tarsometatarsus of oystercatchers. The lower elevation of the inner trochlea, unlike that in most of the modem charadriiforms, is similar to that seen in Telmatomis, thick-knees (Burhinidae), or skimmers (Rynchopidae).
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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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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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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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Page 221 and 222: NUMBER 89 211 FIGURE 1.—Argornis
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Page 241 and 242: The Anseriform Relationships of Ana
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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 294 and 295: 284 Kurochkin, E.N. 1982. [New Orde
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 89 307 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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