First published online November 2, 2007
Journal of Experimental Biology 210, 3955-3961 (2007)
Published by The Company of Biologists 2007
doi: 10.1242/jeb.008953
Functional significance of the uncinate processes in birds
Peter G. Tickle1,
A. Roland Ennos1,
Laura E. Lennox1,
Steven F. Perry2 and
Jonathan R. Codd1,*
1 Faculty of Life Sciences, University of Manchester, Jackson's Mill, PO Box
88, Sackville Street, Manchester M60 1QD, UK
2 Institute for Zoology, Bonn University, Germany

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Fig. 1. Representative skeletons showing the morphological differences in the rib
cage associated with different forms of locomotion in (A) a walking species,
cassowary (Casuaris casuaris); (B) a non-specialist, eagle owl
(Bubo bubo); and (C) a diving species, razorbill (Alca
torda). Uncinate processes are short in walking species, of intermediate
length in non-specialists and long in diving species. In all photographs
cranial is to the left; scale bar, 5 cm.
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Fig. 2. Geometric model of uncinate function. (A) The situation in birds without an
uncinate process. The length of the Mm. appendicocostales, L, changes
with the rib angle, , depending on the distance down the rib,
P, of the posterior attachment. (B) The situation with an uncinate
process of perpendicular length Q behind the anterior rib. Cranial is
to the left.
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Fig. 3. (A) Changes in length of the Mm. appendicocostales muscle with rib angle,
, for various relative values of uncinate length, Q, and
distance of posterior attachment, P. (B) Changes in mechanical
advantage of the Mm. appendicocostales muscle with rib angle, , for
various relative values of uncinate length, Q, and distance of
posterior attachment, P. It can be seen that mechanical advantage
increases with , and with higher values of Q.
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Fig. 4. Mechanical advantage (corrected for muscle length L) for
representative species calculated with (solid line) and without (broken line)
the uncinate processes. (A) A diving bird, the razorbill Alca torda;
(B,C) non-specialist birds, (B) barnacle goose Branta leucopsis and
(C) kestrel Falco tinnunculus; and a walking bird (D) the red-legged
partridge Alectoris rufa.
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Fig. 5. Canonical variate analysis (CVA) of skeletal morphology in birds. Function
1 against function 2 for walking species (squares, N=10);
non-specialists (circles, N=66); diving birds (triangles,
N=24). Functions 1 and 2 were primarily functions of relative
uncinate length and width and rib length, respectively. Solid black squares
represent significantly different group centroids. Letters highlight
borderline species of respective groups: the fulmar ( ), the green
woodpecker (ß) and the swallow (µ).
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© The Company of Biologists Ltd 2007