The ‘wishbone’ of birds comprises two clavicles fused into a structure1 known as a furcula. In an influential 1926 book on bird origins by Heilmann2, the furcula's supposed absence in dinosaurs was considered powerful evidence barring them from bird ancestry. A furcula has now been found in several theropod dinosaurs3,4, but its absence in other theropods, and the uncertainty of whether this absence is real or an artefact of preservation, obscures the evolutionary history of this structure5. Here we report the discovery of a furcula in Dromaeosauridae, a group posited to be the closest relative of birds6,7.
Among close bird relatives (non-avialian Maniraptora), a furcula is known only in Oviraptoridae8. Although a furcula was tentatively reported to be present in an articulated skeleton of Velociraptor mongoliensis preserved embracing a Protoceratops skeleton3,6, the only firsthand description of this specimen maintained that it is not present8, and our observations of that specimen confirm this. A furcula has not been identified in any of the other five described species of Dromaeosauridae. Other taxa either lack any evidence of clavicles or, in one troodontid specimen9, clavicles are present but not fused. But the rarity of articulated specimens, the similarity of the furcula to ribs and gastralia, and incomplete preservation conspire to obscure whether this absence is real.
A partial skeleton of Velociraptor mongoliensis (specimen IGM 100/976) was discovered at Tugrugeen Shireh, Mongolia, during the 1991 expedition of the Mongolian Academy of Sciences-American Museum of Natural History Expedition. The specimen consists of a nearly complete skull and the anterior part of the skeleton, preserving derived characters that allow definitive reference to Velociraptor mongoliensis10. The bones are heavily bioturbated, and many elements are punctured by borings similar to those often present in Djadokhta Formation sediments11, but the specimen is otherwise very well preserved and articulated. The block that holds the furcula also preserves a pair of sternal plates, posterior cervical vertebrae, the proximal ends of both humeri, the coracoids and the right scapula. The specimen is large relative to others of this species, and the cervical ribs are nearly fused to the vertebral centra, indicating that the specimen is near full-size.
The furcula is in articulation between the scapulocoracoids anterior to the sternal plates (Fig. 1), in the same position as in oviraptorids and birds. The bone is ‘V’-shaped and very slender, much thinner than in oviraptorids and Archaeopteryx. In cross-section it is nearly circular. A short midline process, the hypocleidium, extends posteroventrally. The proximal, or epicleideal, process sweeps posterodorsally to articulate with the acromion of the scapulocoracoid. The proximal process of the furcula tapers to a point where it contacts the scapulocoracoid, and the posterior articulating surface contacting the acromion is smooth.
The broad distribution of a furcula among the non-flying relatives of birds4,5,8,12 indicates that its origin is not tied to the origin of flight. The furcula's use in powered avian flight is therefore a co-option of a structure already present before flight evolved in the lineage leading to modern birds4. The discovery of a furcula in a taxon previously known from an articulated specimen underscores the difficulty of assessing whether the apparent absence of this feature is real or an artefact of preservation or ontogenetic sampling5.
Baumel, J. J. & Witmer, L. M. Publ. Nuttal Ornithol. Club 23, 45–132 (1993).
Heilmann, G. The Origin of Birds (Witherby, London, 1926).
Ostrom, J. Biol. J. Linn. Soc. 8, 91–182 (1976).
Makovicky, P. J. & Currie, P. J. J. Vert. Paleontol. (in the press).
Bryant, H. & Russell, H. J. Vert. Paleontol. 13, 171–184 (1993).
Gauthier, J. A. Calif. Acad. Sci. Mem. 8, 1–55 (1986).
Holtz, T. J. J. Paleontol. 70, 536–538 (1996).
Barsbold, R. Sovmestnaya Sovetsko-Mongol'skaya Paleontol. Ekspiditsiya, Trudy 19, 5–119 (1983).
Russell, D. A. & Dong, Z.-H. Can. J. Earth Sci. 30, 2163 (1993).
Norell. M. A. et al. J. Vert. Paleontol. 12 Suppl. 45A (1992).
Jerzykiewicz, T. et al. Can. J. Earth Sci. 30, 2180–2195 (1993).
Chure, D. J. & Madsen, J. H. J. Vert. Paleontol. 16, 573–577 (1996).
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The origin and early evolution of birds: discoveries, disputes, and perspectives from fossil evidence