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208 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY sia helenae (1). Muscle dissections were stained by the method of Bock and Shear (1972), and magnifications from x8 to x50, mainly x 16 and x25, were used. The analysis of forelimb muscles also utilizes data from Cohn (1968) and from Zusi and Bentz (1982, 1984). This paper presents a brief summary of the unpublished results of the investigation of the forelimb muscles in apodiforms (Karhu, 1992a) (see "Forelimb Muscles: Tendencies of Transformation," below). Anatomical terminology used in the descriptions generally follows Baumel et al. (1993). ACKNOWLEDGMENTS.—I am very grateful to A.F. Bannikov (Paleontological Institute, Moscow) for donation of the specimen described in the present paper. I wish to acknowledge C. Mourer-Chauvire (Universite Claude Bernard, Lyon) for the opportunity to study the coracoid specimens of Aegialornis gallicus Lydekker. For the loan of skeletal and spirit specimens of Apodiformes I am very indebted to the following: CT. Collins (California State University, Long Beach), F.Y. Dzerzhinsky (Moscow State University, Moscow), V.M. Loskot (Zoological Institute, St. Petersburg), G.F. Mees (Rijksmuseum van Natuurlijke Historie, Leiden), S.L. Olson, R.L. Zusi, J.P. Angle, and J. Dean (National Museum of Natural History, Smithsonian Institution, Washington, D.C), and F. Vuilleumier (American Museum of Natural History, New York). The manuscript was substantially improved by the thoughtful comments of R.L. Zusi and S.L. Olson. I am very thankful to R.L. Zusi and CT. Collins for their reviews. This research was supported by grants from the Russian Foundation for Basic Researches (number 96-04-50822) and the Frank M. Chapman Memorial Fund of the American Museum of Natural History. Systematic Paleontology Order APODIFORMES Suborder APODI Family JUNGORNITHIDAE TYPE GENUS.—Jungornis Karhu, 1988. EMENDED DIAGNOSIS.—Apex carinae moderately developed. Facies articulares coracoidei widely spaced and separated from rostri sterni. Sulci carinae well pronounced. Proc. acrocoracoideus claviculae placed along dorsal margin of scapus, being considerably narrower than scapus in dorsoventral dimension; concave facies articularis acrocoracoidea claviculae oriented caudolaterally. Facies articularis humeralis scapulae directed cranioventrally. Proc. acrocoracoideus between facies articularis clavicularis and impressio lig. acrocoracohumeralis stretched mediolaterally. Proc. lateralis coracoidei well pronounced, protruding noticeably laterad beyond level of angulus lateralis. Facies articularis sternalis coracoidei wide, with distinctly outlined angulus medialis projecting sternally approximately to same level as angulus lateralis. Caput humeri directed caudally. Crista deltopectoralis high and proximally placed. Proc. supracondylaris dorsalis distally placed. Proximoventral border of cotyla ventralis ulnae not pronounced. Phalanx proximalis digiti majoris bifenestrated. INCLUDED GENERA.—Jungornis Karhu, 1988; Palescyvus Karhu, 1988; Argornis, new genus. STRATIGRAPHIC AND GEOGRAPHIC DISTRIBUTION.—Upper Eocene and lower Oligocene, northern Caucasus, Russia; upper Eocene, Phosphorites du Quercy, France. REMARKS.—In comparison with the former diagnosis of the family Jungornithidae (Karhu, 1988), the emendation omits the presence of the distal enlargement of the middle of the caput humeri and the proximity of the tuberculum supracondylare ventrale and tuberculum M. pronator superficialis. It adds points concerning the development of proc. lateralis coracoidei, orientation of the caput humeri, position of proc. supracondylaris dorsalis, development of the proximoventral border of the cotyla ventralis ulnae, and fenestration of the proximal phalanx of the major digit. Argornis, new genus TYPE SPECIES.—Argornis caucasicus, new species. DISTRIBUTION.—Upper Eocene; northern Caucasus, Russia. ETYMOLOGY.—From the Greek argos, swift, and ornis, bird; the gender is masculine. DIAGNOSIS.—Facies articularis acrocoracoidei claviculae lengthened mediolaterally. Acromion scapulae with cranial margin beveled laterally and crista lig. acrocoraco-acromiale well developed. Dorsal side of medial part of proc. acrococoracoideus forms high, caudally projecting crest, with mediolaterally narrow base. Facies articularis sternalis coracoidei saddleshaped, with only medial part of crista ventralis protruding ventrad. Angulus lateralis of sternal facet projecting a little more distally than angulus medialis. Ventral part of caput humeri oriented perpendicularly to long axis of bone, whereas dorsal part oriented obliquely to it, being placed more distally relative to ventral part; distal border of caput humeri clearly outlined. Tuberculum M. tensor propatagialis pars brevis well pronounced and placed just distal to proc. supracondylaris dorsalis humeri. Tuberculum M. pronator superficialis clearly detached from tuberculum supracondylare ventrale humeri. Tuberculum supracondylare ventrale adjoins condylus ventralis humeri. Epicondylus ventralis markedly prominent ventrad. Proc. flexorius projects slightly distally beyond condylus ventralis. Tuberculum lig. collateralis ventralis ulnae relatively small and weakly protruding ventrad. Tuberculum bicipitale brachii radii with impression of M. biceps brachii. COMPARISON.—Clavicula: In Argornis the facies articularis acrocoracoidei is more elongated mediolaterally, narrower dorsoventrally, and directed more caudally than it is in Jungornis, in which it is a rounded, caudolaterally orientated facet. The overall configuration of the facies articularis acrocoracoidei is very unusual in both Jungornis and Argornis in compari-
NUMBER 89 209 son with other known Apodiformes, and it appears to be closest to that in the Caprimulgidae. Scapula: The cranial margin of the acromion is strongly beveled laterally in Argornis, whereas in Jungornis the margin is thickened. The crista lig. acrocoraco-acromiale is well developed in Argornis, but it is not pronounced in Jungornis. In Argornis the facies articularis humeralis is relatively longer craniocaudally and wider dorsoventrally in comparison with Jungornis. Coracoid: The overall configuration is close to that in Jungornis. In these genera the shaft is relatively slender, and the mediolateral width of the processus acrocoracoideus exceeds the distance between the angulus medialis and the angulus lateralis of the sternal facet (Figure 3c,D), whereas in Palescyvus the shaft is stouter, and the acrocoracoid is narrower than the sternal facet in frontal aspect (Figure 3E). In Argornis the dorsal crest of the medial portion of the acrocoracoid protrudes caudally almost to the level of the middle of the dorsal aperture of the canalis triosseus, and the base of this crest does not reach laterally to the level of the medial border of the impressio lig. acrocoracohumerale. In Jungornis the crest projects much less caudad, and its base is relatively wider, extending laterad to the level of the medial edge of the impression mentioned above. In Palescyvus the medial portion of the acrocoracoid curves strongly caudally but lacks a pronounced dorsal crest. The base of the proc. procoracoideus is relatively wider in Argornis in comparison with either Jungornis or Palescyvus. The facies articularis sternalis is saddle-shaped in Argornis but is concave in Jungornis. In Argornis only the medial part of the crista ventralis of the sternal facet protrudes ventrad, whereas in Jungornis the entire crista ventralis forms a ventral convexity. The ratio of the greatest dorsoventral width of the sternal facet to the distance between the angulus medialis and angulus lateralis is smaller in Argornis than it is in Jungornis. The angulus medialis is rounded in Argornis and in Palescyvus, whereas in Jungornis it is moderately sharpened. The angulus lateralis projects slightly distad beyond the level of the angulus medialis in Argornis; in Jungornis and Palescyvus the angulus medialis and angulus lateralis are placed approximately on the same level (Figure 3C-E). Humerus: In Argornis the humeral shaft is more slender, and both the proximal and distal ends are relatively narrower in comparison with Jungornis (Figure 2E-G,R). The smaller, ventral part of the caput humeri is situated perpendicularly to the long axis of the bone in Argornis, unlike in Jungornis, in which the entire caput humeri is transversely placed (Figure 2H,R). There is no distal enlargement of the middle of the caput humeri on the facies caudalis in Argornis as there is in Jungornis. In Argornis the clearly pronounced tuberculum of M. tensor propatagialis pars brevis adjoins distally the base of the proc. supracondylaris dorsalis. This process in Jungornis is adjoined distally by a high thin crest, and the insertion of M. tensor propatagialis pars brevis is not marked. Tuberculum M. pronator superficialis is low and is clearly separated from the tubercu lum supracondylare ventrale in Argornis, whereas in Jungornis the former protrudes strongly proximad and fuses with the proximal part of the tuberculum supracondylare ventrale. In Argornis the tuberculum supracondylare ventrale adjoins the base of the condylus ventralis ventroproximally, but in Jungornis it is placed more proximally and is well separated from the base of the condylus ventralis. In Argornis the condylus ventralis projects distad beyond the condylus dorsalis, whereas the distal borders of both condyli are placed on the same level in Jungornis. In Argornis the epicondylus ventralis protrudes strongly ventrad, is widened proximodistally, and is fused distally with the proc. flexorius, unlike the much smaller and detached epicondylus ventralis of Jungornis. The proc. flexorius projects distad less in Argornis than in Jungornis. Ulna: In Argornis the tuberculum lig. collaterale ventrale is smaller and protrudes less ventrad in comparison with Jungornis. Radius: In contrast to Jungornis, there is an impression of M. biceps brachii on the ventrocranial surface of the tuberculum bicipitale brachii in Argornis. REMARKS.—An extremely peculiar morphology of the sternocoracoidal articulation is among the most characteristic features of the Apodiformes sensu Wetmore, 1960 (Lucas, 1893; Lowe, 1939; Cohn, 1968; Karhu, 1988, 1992a). In the Apodiformes, the sternum possesses a weakly saddle-shaped or convex facies articularis coracoidei instead of the coracoidal sulcus of most birds. Consequently, the coracoid has a more or less dorsoventrally widened facies articularis sternalis that is slightly saddle-shaped, or concave, and placed on the whole perpendicularly to the long axis of the bone. There is a single exception: in Aegialomis the coracoid has the sternal facet ventrally widened near the angulus medialis, whereas its greater part is wedge-shaped in dorsoventral section, as is typical for most birds. The structure of the sternal facet of the coracoid is considerably more generalized in Argornis in comparison with Jungornis. In Jungornis the overall configuration of the facies articularis sternalis is apodid-like, with the entire ventral margin convex ventrad but to a lesser degree than in true swifts. In Argornis the coracoid possesses a facies articularis sternalis that is widened by a ventral prominence in its medial part only, which is similar to the most generalized type within Apodiformes as demonstrated by Aegialomis. Argornis differs from Aegialomis, however, in having the dorsal edge of the sternal facet situated much more sternally (Figure 3A,C). The other relatively generalized character of the coracoid in Argornis is the placement of the angulus lateralis slightly beyond the level of the angulus medialis (Figure 3C). In Jungornis and Palescyvus the angulus lateralis and angulus medialis are on the same level relative to the long axis of the bone (Figure 3D,E), which was pointed out as a distinctive familial character in the former diagnosis of the Jungornithidae (Karhu, 1988). In the Hemiprocnidae and Apodidae, as well as in the Eocene genus Aegialomis, the angulus lateralis protrudes con-
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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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Page 199 and 200: NUMBER 89 189 carpal trochlea relat
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Page 203 and 204: NUMBER 89 193 We compared the parat
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Page 241 and 242: The Anseriform Relationships of Ana
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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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