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A Revision of the Fossil Swifts (Vertebrata, Aves, Suborder, Apodi), with Descriptions of Three New Genera and Two New Species

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A Revision of the Fossil Swifts (Vertebrata, Aves, Suborder, Apodi), with Descriptions of Three New Genera and Two New Species Meded. Werkgr. Tert. Kwart. Geol. vol. 21 (4) pp. 157-177 2 tabs, 8 figs Leiden, December 1984 A revision of the fossil swifts (Vertebrata, Aves, Suborder, Apodi), with descriptions of three new genera and two new species by C.J.O. Harrison Harrow, Middlesex, U.K. Harrison, C. J. 0. A revision of the fossil swifts (Vertebrata, Aves, suborder with of three two Apodi), descriptions new genera and new species. - Meded. Werkgr. Tert. Kwart. Geol., 21 (4): 157-177, 2 tabs, 8 figs. Leiden, December 1984. Humeri are available for all fossil swifts, and other forelimb elements for all but one, making osteological comparisons with other fossil and fossil humeral Recent species possible. Six genera are recognized, diag- for and - noses provided each, two new genera species being described Eocypselus vincenti of the Early Eocene and Scaniacypselus wardi of the Middle Eocene. In addition a new genus Procypseloides is erected for with which intermedius Cypseloides ignotus, Cypselavus is syno- nymized. Genera are tentatively assigned to families and subfamilies, new taxa being erected for Eocypselus and Primapus. The morphological divergencies that characterize the Recent major taxa - Hemiprocnidae, Cypseloidinae and Apodinae - are already apparent in the Eocene swifts. C. J. 0. Harrison, 48 Earl's Crescent, Harrow, Middlesex HA1 1XN, England. Contents: Samenvatting, p. 158 Introduction, p. 158 159 Diagnoses and descriptions, p. 159 Order Apodiformes, hummingbirds and swifts, p. Suborder 161 Trochili, hummingbirds, p. Suborder 161 Apodi, swifts, p. 161 Genera included in this study, p. 161 Genus Eocypselus gen. nov., p. Genus 163 Aegialornis Lydekker, 1891, p. Genus 163 Cypselavus Gaillard, 1908, p. Genus Hemiprocne Nitzsch, 1829, p. 164 Genus Primapus Harrison & Walker, 1975, p. 164 Genus Scaniacypselus gen. nov., p. 166 Genus Procypseloides gen. nov., p. 168 Genus 170 Cypseloides Streubel, 1848, p. Genus 170 Apus Scopoli, 1777, p. Families and subfamilies, p. 170 Measurements, p. 175 177 References, p. 158 Samenvatting Een revisie van de fossiele gierzwaluwen (Vertebrata, Aves, suborde Apodi), met beschrijvingen van drie nieuwe geslachten en twee nieuwe soorten. Van alle bekende fossiele gierzwaluwen zijn de opperarmbeenderen beschikbaar en andere vleugelelementen voor alle soorten behalve één, hetgeen osteologische vergelijkingen met andere fossiele en recente soorten mogelijk maakt. Er worden zes fossiele geslachten erkend. Voor elk daarvan worden de Twee nieuwe diagnoses gegeven van opperarmbeenderen. soorten en twee nieuwe geslachten worden beschreven, nl. Eocypselus vincenti gen. et sp. nov. uit het Vroeg Eoceen het en Scaniacypselus wardi gen. et sp. nov. uit Midden Eoceen. Verder wordt Procypseloides gen. nov. beschreven voor de soort Cypseloides ignotus, waarmede Cypselavus intermedius wordt gesyno- nimiseerd. De geslachten worden voorlopig tot families en subfamilies gerekend, waarbij nieuwe taxa worden opgesteld voor Eocypselus en Primapus. De morphologische divergenties die de belang- rijkste recente taxa (Hemiprocnidae, Cypseloidinae en Apodinae) karakteriseren zijn al duidelijk aanwezig bij de Eocene gierzwaluwen. Introduction Bird bones are mostly hollow and fragile and unlikely to survive intact in most of the geological strata that provide palaeontological material. Those that are preserved completely tend to be short, stout and solid. The humeri of Apodiformes possess these useful characters. The hummingbirds and swifts that constitute this order have long slender wings. The latter are formed from long forces feathers attached to a relatively short bony structure on which strong muscular are exerted and which has become modified into a series of short, stout bones. The humerus in particular has changed from the elongated and often slender structure present in most birds to a short, squarish bone in the more highly evolved species. 10-15 be These humeri are usually small, little more than mm long, and may easily overlooked, but tend to persist relatively intact in fossil deposits. A number of early swift-like birds have been described from such specimens, and since it is usually the humerus that is available it is possible to of which compare these and to establish some degrees similarity or divergence may provide possible clues to affinities and evolutionary trends. Of particular importance are the relative length and stoutness of the shaft, length of the crista pectoralis, and the development of the attachment M. radialis and M. brevis. processes of extensor metacarpi tensor propatagialis pars Several species of fossil swift were identified from deposits in France during the past century. More recently a species was described from the Early Eocene of south-east England. The discovery of another British Early Eocene species differing from the last and of an additional species from the Middle Eocene of Denmark, has made it possible to provide a broader re-examination of these taxa. Both the last two specimens are humeri with a few associated bones of the pectoral girdle, making a more complete comparison possible. available indicate their The present paper identifies and defines the taxa and attempts to place In it been describe within the pattern of evolutionary trends. doing so has necessary to two new for and genera and two new species, provide a new genus an already-described species a lectotype for another. In the following descriptions and definitions the various forms within the suborder 159 Apodi are listed by genus, and the family groupings are introduced only within the subsequent discussions. The terminology used is that of the Nomina Anatomica Avium (Baumel, 1979). DIAGNOSES AND DESCRIPTIONS this In section the diagnoses have been primarily based on the humeri which provide reasonably discrete characters for this purpose. Order APODIFORMES, hummingbirds and swifts These are birds having short, stout humeri with a well-developed process of M. extensor meta- carpi radialis proximally elevated on the lateral side of the shaft, and a more distal tubercle of M. brevis often and the tensor propatagialis pars apparent showing same tendency to proximal elevation. Some of the earlier fossil forms show similarities in some characters of the humerus to those of the nightjars (Caprimulgiformes). Collins (1976a) in studying the humeri of the more divergent species of the Eocene/Oligocene genus Aegialornis referred them to the Caprimulgiformes, and influenced Brodkorb (1971) in this respect, although this has not been subsequently accepted Chordeiles (Harrison, 1975; Mourer-Chauvire, 1978). Collins cited nightjars of the genus as providing evidence of affinity. With the humerus of a very early, generalised species Eocypselus vincenti (fig. IE) now available it is possible to compare the most generalised and most derived forms of swifts with nightjars such as the recent common nighthawk Chordeiles minor (fig. 1 A). those In general nightjar humeri are larger and more slender than of swifts. In swift humeri of the more slender type there is a tendency towards a ventral (medial) arching of the whole distal end. One nightjar family, the owlet-frogmouths (Aegothaledidae) show a superficially similar dis- placement but this is achieved by a distinct medial bend of the shaft, and in most nightjars the proximal head is aligned with the shaft. The caudally-curving arch of the crista bicipitalis and tuberculum ventrale over the pneumotricipital fossa is more distal in the swifts and the bicipital the area more caudoventrally curved. The main cranial projection of crista pectoralis is more distally sited in nightjars. The tuberculum dorsale is small and proximally sited, whereas those of swifts, end the although variable, are much larger, distally elongated and caudally tilted. The distal of it humerus is broad in swifts, and even more so in hummingbirds, but in nightjars is narrower, with a smaller and more deeply defined and proximally placed M. brachialis fossa. In the latter the is condylus ventralis more rounded and the fossa olecrani less deep and more ventrally placed, and in addition the processus flexorius is a simple caudal projection and is not dorsocaudally elongated as in the swifts. Although Chordeiles shows papillae for both M. extensor metacarpi radialis and lateral distal M. tensor propatagialis pars brevis, these are tiny and distally sited on the side near the end. similarities Collins's that Although some general osteological may justify (1976a) suggestion nightjars and swifts have some affinity and possibly a common evolutionary origin later than that of other ordinal taxa, the structures present show sufficient characters to maintain the separation of the taxa involved. 160 Fig. 1. Left humeri, standardizedfor comparison. The first the others swifts is a nightjar (Caprimulgiformes), are (Apodiformes). b = d = = f = distal. Views (left to right): a = cranial, = caudal, c lateral, medial, e proximal, A. Common nighthawk, Chordeiles minor, Recent; x 2/3. B. Lesser treeswiftHemiprocne comata, Recent; x 2. C. Cypselavus gallicus, Late Eocene/Early Oligocene (after Mourer-Chauviré, 1978); x 2¼. D. Aegialornis gallicus, Late Eocene/Early Oligocene ;x 1. E. Eocypselus vincenti, Early Eocene, holotype; x 2. 161 Suborder TROCHILI, hummingbirds These small with the humerus short and the distal end are to tiny birds, very broad, comparable in width with the proximal end, the shaft forming only about a third of the total length, and a large, broad-ended process of M. extensor metacarpi radialis about halfway along the
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