Science 21 September 2007:
Vol. 317. no. 5845, p. 1721
DOI: 10.1126/science.1145076Brevia<?xml:namespace prefix = o ns = "urn:schemas-microsoft-com:office:office" />
Feather Quill Knobs in the Dinosaur Velociraptor
Alan H. Turner,1* Peter J. Makovicky,2 Mark A. Norell1
Some nonavian theropod dinosaurs were at least partially coveredin feathers or filamentous protofeathers. However, a completeunderstanding of feather distribution among theropod dinosaursis limited because feathers are typically preserved only inlagerstätten like that of <?xml:namespace prefix = st1 ns = "urn:schemas-microsoft-com:office:smarttags" />Solnhofen, Germany or Liaoning, China.Such deposits possess clear taphonomic biases toward small-bodiedanimals, limiting our knowledge regarding feather presence inlarger members of feathered clades. We present direct evidenceof feathers in Velociraptor mongoliensis based on the presenceof quill knobs on the posterior forearm. This report of secondariesin a larger-bodied, derived, and clearly flightless member ofa nonavian theropod clade represented by feathered relativesis a substantial contribution to our knowledge of the evolutionof feathers.
1 Division of Paleontology, American Museum of Natural History, Central Park West at 79th Street, New York, NY100245192, USA.
2 Department of Geology, The Field Museum, 1400 South Lake Shore Drive, Chicago, IL 606052496, USA.
* To whom correspondence should be addressed. E-mail: firstname.lastname@example.org
Some nonavian theropod dinosaurs were at least partially coveredin feathers or filamentous protofeathers (1). However, a completeunderstanding of feather distribution among theropod dinosaursis limited because feathers are typically preserved only inlagerstätten like that of Solnhofen, Germany or Liaoning,China. Such deposits possess clear taphonomic biases towardsmall-bodied animals, limiting our knowledge regarding featherpresence in larger members of feathered clades.
We present direct evidence of feathers in Velociraptor mongoliensisbased on the presence of quill knobs on the posterior forearm.In many living birds, raised knobs along the caudal margin oftheulnarevealwhere the quills of the secondary feathers areanchored to the bone by follicular ligaments. Quill knobs arevariably present in extant bird species and are present in onlya few basal taxa such as Ichthyornis (2), so their absence doesnot necessarily indicate a lack of feathers. Their presence,however, is a direct indicator of feathers of modern aspect(e.g., feathers composed of a rachis and vanes formed by barbs).
The specimen IGM (Geological Institute of Mongolia) 100/981was collected at the Gilvent Wash locality near Ukhaa Tolgod(Campanian Djadokhta Formation). The specimen is estimated tohave been 1.5 m long and to have weighed roughly 15 kg. It possessesseveral characteristics found in V. mongoliensis, a common dromaeosauridin the Djadokhta Formation. IGM 100/981 preserves six low papillaeon the middle third of the caudal margin of the ulna (Fig. 1).These are regularly spaced about 4 mm apart. Topographically,these papillae correspond to the quill knobs in living birds.Given their spacing in IGM 100/981, we estimated that thereis space for eight additional secondary feathers. This suggeststhat 14 secondaries were present in Velociraptor, which compareswell with the 12 or more secondaries in Archaeopteryx (3). About18 secondaries are suggested for the dromaeosaurid Microraptor(4), whereas its close relative Rahonavis appears to have possessedjust 10 (5).
Fig. 1. (A) Dorsal view of right ulna of Velociraptor IGM 100/981. (B) Detail of red box in (A), with arrows showing six evenly spaced feather quill knobs. In (B), a cast of IGM 100/981 was used. (C) Dorsal view of right ulna of a turkey vulture (Cathartes). (D) Same view of Cathartes as in (C) but with soft tissue dissected to reveal placement of the secondary feathers and greater secondary coverts relative to the quill knobs. (E) Detail of Cathartes, with one quill completely removed to reveal quill knob. (F) Same view as in (E) but with quill reflected to the left to show placement of quill, knob, and follicular ligament. Follicular ligament indicated with arrow
Such variation is expected because extant birds display variablecounts even within species (3).
Known coelurosaurs with wing feathers of modern aspect are smallbasal members of their respective clades. Some have been consideredpossibly volant (4, 5), and it has been suggested that the large-bodied,derived members of the feathered theropod clades may not haveretained feathers or only retained feathers while juveniles(6). This Velociraptor specimen indicates this is not the casefor at least one lineage of dromaeosaurids. An examination ofthe living families of birds shows a significant correlationbetween the absence of ulnar papillae and the loss and/or reductionin volancy, even though some strong flyers lack papillae (7).This raises the possibility that ulnar papillar reduction orabsence in large-bodied derived dromaeosaurids reflects lossof aerodynamic capabilities from the clade's ancestral members.Quill knobs in Velociraptor could reflect retention of feathersfrom smaller possibly volant ancestors, but such feathers mayhave had other functions. Although thermoregulatory effectsof secondaries on the ulna would be negligible, such featherscould have been used for display (1), in shielding nests forthermal control (8), or for creating negative lift during inclinerunning (9). Whether this feature represents retention of anancestral function or the cooption for other purposes, the presenceof quilled feathers on the posterior of the arms in a medium-sizedderived, clearly nonvolant dromaeosaur can now be established.
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· 7. Materials and methods are available on Science Online.
· 8. T. P. Hopp, M. J. Oren, in Feathered Dinosaurs, P. J. Currie, E. B. Koppelhus, M. A. Shugar, J. L. Wright, Eds. (Indiana Univ. Press, Bloomington, IL, 2004), pp. 234250.
· 9. K. P. Dial, Science 299, 402 (2003).[Abstract/Free Full Text]
· 10. We thank S. Nesbitt for comments, L. Barber and A. Balcarcel for preparation and casting, M. Ellison for photography, P. Sweet, P. Capainolo, and the 1998 Gobi field crew. This study was supported by NSF Division of Earth Sciences (M.A.N. and P.J.M.) and a NSF Doctoral Dissertation Improvement grant (A.H.T.).
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