A Review of the Lower Miocene Swifts (Aves: Apodidae) Charles T. Collins Natural History for support and I am also indebted ABSTRACT to Dr. J. P. Lehman of the Institut de Pal^ontol- ogie, Museum National D’Histoire Naturelle, Paris, Three nominal species of swifts have been de¬ for lending the types of Cypselus ignotus and scribed from lower Miocene (Aquitanian) depos¬ Collocalia incerta. its of France. Re-examination of these forms, Cypselus [ = Apus] ignotus Milne-Edwards, Collo- calia incerta Milne-Edwards, and Cypselavus inter- Family APODIDAE medius Gaillard, indicates that they are attrib¬ utable to a single species, ignotus, referable to the Subfamily CYPSELOIDINAE modern genus Cypseloides. This provides the first occurrence of the Cypseloidinae in the fossil record and indicates a possible origin in the Old World Cypseloides ignotus (Milne-Edwards, 1871) for this primitive group of swifts, presently re¬ stricted to the New World. Cypselus ignotus Milne-Edwards ,1871:394 ,pi .177 :figs .9-13. Apu signotu s.—Paris 1,912:286. Collocalia incerta Milne-Edwards ,1871:394 ,pi .177 :figs .1-8. Cypselavus intermedius Gaillard ,1939:42 ,fig .20. Introduction From the Aquitanian deposits at St.-Gerand-le- Up to now, five species of fossil swifts have been Puy, Departement de Allier, France, Milne- described, all coming from Tertiary deposits in Edwards (1871) described a new species of swift, France. The present paper is aimed at reviewing Cypselus [=Apus~\ ignotus. This was based on a the three nominal species from deposits of early complete right carpometacarpus and a left ulna Miocene age, with comparisons being made with a with the proximal end badly chipped. When I ex¬ much wider array of skeletal material of modern amined these specimens, a second left ulna, excel¬ swifts than were available to the original describers lently preserved, had somehow been associated of the fossil forms. Recent swifts examined in this with the two syntypes. This is identical to the first study included Cypseloides rutilus, C. cherriei, ulna and I therefore refer it to the species ignotus C. niger, and Streptoprocne zonaris in the Cypse¬ also. From the same deposits, Milne-Edwards loidinae; from one to several species in the genera (1871) named a second species of swift, Collocalia Apus, Aeronautes, Cypsiurus, Tachornis, Reinarda, incerta, based on a single well-preserved left tibio- and Panyptila in the Apodinae; and Chaetura, Col- tarsus. This was characterized as being much too localia, and Hirundapus in the Chaeturinae. small to have come from the same species as the Acknowledgments. —I thank the Frank M. Chap¬ wing elements assigned to Apus ignotus. Consider¬ man Memorial Fund of the American Museum of ably later, Gaillard (1939) reported a left humerus of a swift from Aquitanian deposits at Chavroches, Charles T. Collins, Department of Biology, California State also in'the Departement de Allier, which he de¬ University ,Long Beach ,California 90840. scribed as a new species, ihtermedius, in the 129 130 SMITHSONIA NCONTRIBUTION TS OPALEOBIOLOGY Eocene-Oligocene genus Cypselavus (Gaillard, ternal cotyla appears slightly more undercut at its 1908). palmar edge and the distal ligamental attachment I have studied the original material of Apus of the carpal tuberosity is less laterally flared. ignotus and Collocalia incerta as well as Gaillard’s The carpometacarpus of swifts shows less marked (1989) description and illustrations of Cypselavus distinctions between the subfamilies than does the intermedins. While all these fossils clearly belong ulna. That of ignotus (Figure 1 b) is somewhat to the Apodidae, it is equally apparent that none is longer and stockier than in Cypseloides rutilus or referable either to Apus or to Collocalia. C. cherriei, but it does have the more widely flared In size and conformation, the two ulnae of pollical facet of metacarpal I and the more pro¬ ignotus (Figure lfl) are very similar to those of nounced tendinal groove on metacarpal II typical some of the smaller species of the genus Cypse- of the Cypseloidinae. The fossil also has a broader loides. They lack the well-developed olecranal and more flared tuberosity of metacarpal II, pro¬ process found in the subfamilies Apodinae and viding a wider articular facet for digit II, which is Chaeturinae. This condition is typical of the typical of the two smaller species of Cypseloides Cypseloidinae. The fossil ulnae are slightly longer studied, but not of swifts of other subfamilies. and stockier than the ulnae of Cypseloides rutilus As noted by Milne-Edwards (1871) the tibio¬ or C. cherriei, and the prominence for the anterior tarsus of Collocalia incerta (Figure 1 c,d) is indeed articular ligament is more shelf-like. Also, the ex- more delicate than would be expected for any member of the Apodinae or Chaeturinae of the size of Apus ignotus. However, the tibiotarsus in the Cypseloidinae is proportionately more slender than in the other subfamilies of swifts, particularly the Apodinae. There is, in fact, a very close agree¬ ment in overall size and morphology between the type of Collocalia incerta and Recent specimens of Cypseloides rutilus. The posterior intercondylar groove of incerta is not deeply excavated as it is in members of the Apodinae and Chaeturinae, in¬ cluding Apus and Collocalia. The proximal por¬ tion of the shaft is straight, as in Cypseloides, and not distinctly bent laterally as typical of many other swifts. The fossil element is slightly smaller and stockier than in C. rutilus (C. cherriei has a much longer tibiotarsus than C. rutilus in spite of its having wing elements similar in size to both C. rutilus and A. ignotus), and the internal liga¬ mental prominence is less developed but more ex¬ cavated under the lip of the rotular crest. The wing elements of ignotus are clearly those of a small swift belonging to the genus Cypseloides. The tibiotarsus of incerta similarly shows affinities to Cypseloides particularly to C. rutilus. Con¬ trary to Milne-Edwards (1871), it is entirely probable that these fossils, which are from the same locality and horizon, come from the same species. This species should now be known as Ficurf .1.—Bones o fCypseloides ignotus (Milne-Edwards) :a, referred left ulna; b, syntype right carpometacarpus; c, left Cypseloides ignotus (Milne-Edwards) with incerta tibiotarsus (holotypc of Collocalia incerta) ,anterior view; d, becoming a junior synonym, ignotus being chosen same, posterior view. (Approximately X 3.5, c and d at on the basis of line priority. slight ldyiffere nmt agnifications.) In the referred ulna of Cypseloides ignotus, the NUMB E2R7 131 maximum length is 17.9 mm, distal width 3.2 mm, earliest fossil possibly attributable to Apus now be¬ proximal width 3.7 mm, and shaft width 1.6 mm. comes Apus gaillardi (Ennouchi) from the upper No accurate length could be determined for the middle Miocene (Tortonian) of France (Brod¬ chipped ulna in which the distal width is 3.2 mm, korb, 1971). proximal width 3.7 mm, and shaft width 1.8 mm. Although the specimens of Cypselavus gallicus The single carpometacarpus measures 16.4 mm in and Apus gaillardi were not examined in this total length, proximal height 5.2 mm, proximal study, the published illustrations are sufficient to width 2.35 mm, and distal width 3.95 mm. The determine that neither species shows any similari¬ tibiotarsus has a total length of 21.1 mm, width ties to Cypseloides ignotus or the modern Cypselo- across condyles 2.2 mm, width across proximal ar¬ idinae. In fact, Cypselavus gallicus shows a dis¬ ticular surfaces 2.5 mm, and shaft dimensions of tinctly closer resemblance to the Aegialornithidae, 0.9 X 1.0 mm at the narrowest point and 1.0 X 1.5 the humerus agreeing in size with the newly mm at the middle of the fibular crest. described small aegialornithid Primapus lacki, The type-humerus of Cypselavas intermedius from the lower Eocene of Britain (Harrison and Gaillard (1939) was not examined in this study, Walker, 1975). In the published illustrations (Gail¬ but from the original figures it appears to have the lard, 1908), the humerus of Cypselavus gallicus distinctively longer and narrow proportions char¬ appears to lack the prominant ectepicondylar acteristic of the species of Cypseloides. As was process seen in the Aegialornithidae, but this noted by Lowe (1939:324), the ectepicondylar could well be the result of damage. The illustra¬ process of intermedius is much more distally po¬ tions of the humerus of Apus gaillardi (Ennouchi, sitioned than in any of the modern forms of the 1930) show it to have the general proportions of Apodinae or Chaeturinae but is only slightly more the modern Apodidae and Chaeturinae. This spe¬ distal than in Cypseloides, a genus that was not cies, and an additional swift from the upper Mio¬ compared by earlier workers. The measurements cene of Italy, are currently under review elsewhere of the type of C. intermedius as given by Gaillard (P. Ballmann, pers. comm.). (1939:43) are: total length 11 mm, proximal width The Cypseloidinae (see Brooke, 1970:14-15 for 4.5 mm, distal width 3 mm. Thus, this specimen use of this term) appears to be the most primitive agrees closely in size with specimens of modern subfamily of the Apodidae. It is therefore not un¬ Cypseloides rutilus and C. cherriei, and it would expected that an extinct species of Cypseloides be therefore also be of the same approximate size as among the earliest known swifts. Like the vultures C. ignotus. The type of Cypselavus intermedius of the family Cathartidae [= Vulturidae], the comes from the same horizon and from a locality modern species of Cypseloidinae are confined to close to that of Cypseloides ignotus. Since it also the New World; but also like the Cathartidae appears to belong to the genus Cypseloides and is (Cracraft and Rich, 1972), they can now be shown of the same size as C. ignotus, I feel that Cypsela¬ to have had a past distribution and possible origin vus intermedius should also be synonymized with in the Old World. Further elucidation of the origin Cypseloides ignotus. As a result, the genus Cypsela¬ and evolution of the Apodidae will have to await a vus Gaillard is reduced to a single species, C. galli- review of additional modern forms and the remain¬ cus, from the upper Eocene or lower Oligocene ing fossil swifts, as well as the swift-like members of (Phosphorites du Quercy) of France; the genus the Aegialornithidae (see Harrison and Walker, Collocalia is deleted from the fossil record; and the 1975; Harrison, 1975, Collins, pp. 121-127, herein). Literature Cited Brodkor bP., families, Tribes, Genera and Subgenera of the 1971. Catalogue of Fossil Birds, Part 4 (Columbiformes Swift s(Aves :Apodidae) D. urban Museum Novitates, through Piciformes). Bulletin of the Florida State 9( 21)3:-24. Museum B, iologica Sl ciences 1, 5(4) 1: 63-266. Cracraft, J., and P. V. Rich 1972. The Systematics and Evolution of the Cathartidae Brooke ,R .K. in the Old World Tertiary. Condor, 74(3):272-283, 1970. Taxonomic and Evolutionary Notes on the Sub- 1fi0gures. 132 SMITHSONIA NCONTRIBUTION TS OPALEOBIOLOGY Ennouch iE,. Harrison, C. J. O., and C. A. Walker 1930. Contribution a I’etude de la faune du Tortonien 1975. A New Swift from the Lower Eocene of Britain. de La Grive-St-Alban (Isere). 135 pages, 6 plates. Ibis ,117(2) :162-164 ,2 plates. Paris L: e sPresse sModemes. Lowe, P. R.1938. On the Systematic Position of the Swifts (Suborder Gaillard C, . Cypseli) and Hummingbirds (Suborder Trochili), 1908. Les oiseaux des phosphorites du Quercy. Annales with Special Reference to their Relation to the du Universite de Lyon, new series, 1(23): 1-178, 37 Orde rPasseriformes T. ransaction so fth eZoological figure s8p, lates. Society o fLondon ,25(4):307-348 ,4 plates. 1939. Contributions a l’etude des oiseaux fossiles. Ar¬ Milne-Edward As,. chives du Museum d’Histoire Naturelle de Lyon, 1871 .Recherches anatomiques e tpaleontologiques pourservir a I’histoire des oiseaux fossiles de la France. 15(m£moir e2) 1: -100 3, 4figures. 4 volumes (1869-1871) .Paris :Victor Masson et Fils. Harrison, C. J. O. Pari sP,. 1975. Ordinal Affinites of the Aegialornithidae. Ibis, 1912. Oiseaux fossiles de France. Revue Frangaise 117(2 )1:64-170 5f,igures. d’Ornitholo g4(ie3,7):283-298. Collins, Charles T. 1976. "A Review of the Lower Miocene Swifts (Aves: Apodidae)." Collected papers in avian paleontology honoring the 90th birthday of Alexander Wetmore 27, 129–132. 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