BREVIA

Such variation is expected because extant birds

Feather Quill Knobs in the display variable counts even within species (3).
Known coelurosaurs with wing feathers of

Dinosaur Velociraptor modern aspect are small basal members of their

respective clades. Some have been considered
possibly volant (4, 5), and it has been suggested
Alan H. Turner,1* Peter J. Makovicky,2 Mark A. Norell1 that the large-bodied, derived members of the
feathered theropod clades may not have retained
ome nonavian theropod dinosaurs were at of the caudal margin of the ulna (Fig. 1). These feathers or only retained feathers while juve-

S least partially covered in feathers or fila- are regularly spaced about 4 mm apart. Topo-
mentous protofeathers (1). However, a graphically, these papillae correspond to the quill
complete understanding of feather distribution knobs in living birds. Given their spacing in
niles (6). This Velociraptor specimen indicates
this is not the case for at least one lineage of
dromaeosaurids. An examination of the living
among theropod dinosaurs is limited because IGM 100/981, we estimated that there is space families of birds shows a significant correlation
feathers are typically preserved only in lager- for eight additional secondary feathers. This between the absence of ulnar papillae and the
stätten like that of Solnhofen, Germany or suggests that 14 secondaries were present in loss and/or reduction in volancy, even though
Liaoning, China. Such deposits possess clear Velociraptor, which compares well with the 12 or some strong flyers lack papillae (7). This raises
taphonomic biases toward small-bodied animals, more secondaries in Archaeopteryx (3). About 18 the possibility that ulnar papillar reduction or
limiting our knowledge regarding feather pres- secondaries are suggested for the dromaeo- absence in large-bodied derived dromaeosaurids

Downloaded from http://science.sciencemag.org/ on April 11, 2017

ence in larger members of feathered clades. saurid Microraptor (4), whereas its close relative reflects loss of aerodynamic capabilities from the
We present direct evidence of feathers in Rahonavis appears to have possessed just 10 (5). clade’s ancestral members. Quill knobs in
Velociraptor mongoliensis Velociraptor could reflect retention of feathers
based on the presence from smaller possibly volant ancestors, but such
of quill knobs on the pos- feathers may have had other functions. Although
terior forearm. In many thermoregulatory effects of secondaries on the
living birds, raised knobs ulna would be negligible, such feathers could
along the caudal margin have been used for display (1), in shielding nests
of the ulna reveal where for thermal control (8), or for creating negative
the quills of the second- lift during incline running (9). Whether this fea-
ary feathers are anchored ture represents retention of an ancestral function
to the bone by follicular or the cooption for other purposes, the presence
ligaments. Quill knobs of quilled feathers on the posterior of the arms
are variably present in in a medium-sized derived, clearly nonvolant
extant bird species and dromaeosaur can now be established.
are present in only a References and Notes
few basal taxa such as 1. M. A. Norell, X. Xu, Annu. Rev. Earth Planet. Sci. 33, 277
Ichthyornis (2), so their (2005).
absence does not nec- 2. J. A. Clarke, Bull. Am. Mus. Nat. Hist. 286, 1 (2004).
3. A. Elzanowski, in Mesozoic Birds, Above the Heads of
essarily indicate a lack Dinosaurs, L. M. Chiappe, L. M. Witmer, Eds. (Univ.
of feathers. Their pres- California Press, Berkeley, CA, 2002), pp. 129–159.
ence, however, is a di- 4. X. Xu et al., Nature 421, 335 (2003).
5. C. A. Forster, S. D. Sampson, L. M. Chiappe, D. W. Krause,
rect indicator of feathers Science 279, 1915 (1998).
of modern aspect (e.g., 6. X. Xu et al., Nature 431, 680 (2004).
feathers composed of a 7. Materials and methods are available on Science Online.
rachis and vanes formed 8. T. P. Hopp, M. J. Oren, in Feathered Dinosaurs, P. J. Currie,
E. B. Koppelhus, M. A. Shugar, J. L. Wright, Eds. (Indiana
by barbs). Univ. Press, Bloomington, IL, 2004), pp. 234–250.
The specimen IGM 9. K. P. Dial, Science 299, 402 (2003).
(Geological Institute 10. We thank S. Nesbitt for comments, L. Barber and A. Balcarcel
of Mongolia) 100/981 for preparation and casting, M. Ellison for photography,
P. Sweet, P. Capainolo, and the 1998 Gobi field crew. This
was collected at the study was supported by NSF Division of Earth Sciences
Gilvent Wash locality (M.A.N. and P.J.M.) and a NSF Doctoral Dissertation
near Ukhaa Tolgod Improvement grant (A.H.T.).
(Campanian Djadokhta Supporting Online Material
Formation). The speci- www.sciencemag.org/cgi/content/full/317/5845/1721/DC1
Materials and Methods
men is estimated to have
Fig. S1
been 1.5 m long and to References
have weighed roughly Fig. 1. (A) Dorsal view of right ulna of Velociraptor IGM 100/981. (B) Detail 14 May 2007; accepted 30 July 2007
15 kg. It possesses sev- of red box in (A), with arrows showing six evenly spaced feather quill knobs. 10.1126/science.1145076
eral characteristics found In (B), a cast of IGM 100/981 was used. (C) Dorsal view of right ulna of a
in V. mongoliensis, a turkey vulture (Cathartes). (D) Same view of Cathartes as in (C) but with soft 1
common dromaeosaurid tissue dissected to reveal placement of the secondary feathers and greater Division of Paleontology, American Museum of Natural History,
Central Park West at 79th Street, New York, NY 10024–5192,
in the Djadokhta For- secondary coverts relative to the quill knobs. (E) Detail of Cathartes, with USA. 2Department of Geology, The Field Museum, 1400 South
mation. IGM 100/981 one quill completely removed to reveal quill knob. (F) Same view as in (E) Lake Shore Drive, Chicago, IL 60605–2496, USA.
preserves six low papil- but with quill reflected to the left to show placement of quill, knob, and *To whom correspondence should be addressed. E-mail:
lae on the middle third follicular ligament. Follicular ligament indicated with arrow. [email protected]

www.sciencemag.org SCIENCE VOL 317 21 SEPTEMBER 2007 1721

 Feather Quill Knobs in the Dinosaur Velociraptor
Alan H. Turner, Peter J. Makovicky and Mark A. Norell (September
21, 2007)
Science 317 (5845), 1721. [doi: 10.1126/science.1145076]

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