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Because he was so gracious to bring it up....
Habib, M. B. and C. B. Ruff. 2008. The effects of locomotion on the structural
characteristics of avian limb bones. _Zoological Journal of the Linnaean
Society_ 153(3):601â624.
Abstract:
"Despite the wide range of locomotor adaptations in birds, little detailed
attention has been given to the relationships between the quantitative
structural characteristics of avian limb bones and bird behaviour. Possible
differences in forelimb relative to hindlimb strength across species have been
especially neglected. We generated cross-sectional, geometric data from
peripheral quantitative computed tomography scans of the humerus and femur of
127 avian skeletons, representing 15 species of extant birds in 13 families.
The sample includes terrestrial runners, arboreal perchers, hindlimb-propelled
divers, forelimb-propelled divers and dynamic soarers. The hindlimb-propelled
diving class includes a recently flightless island form. Our results
demonstrate that locomotor dynamics can be differentiated in most cases based
on cross-sectional properties, and that structural proportions are often more
informative than bone length proportions for determining
behaviour and locomotion. Recently flightless forms, for example, are more
easily distinguished using structural ratios than using length ratios. A proper
phylogenetic context is important for correctly interpreting structural
characteristics, especially for recently flightless forms. Some of the most
extreme adaptations to mechanical loading are seen in aquatic forms. Penguins
have forelimbs adapted to very high loads. Aquatic species differ from
non-aquatic species on the basis of relative cortical thickness. The
combination of bone structural strength and relative cortical area of the
humerus successfully differentiates all of our locomotor groups. The methods
used in this study are highly applicable to fossil taxa, for which morphology
is known but behaviour is not. The use of bone structural characteristics is
particularly useful in palaeontology not only because it generates strong
signals for many locomotor guilds, but also because analysing such
traits does not require knowledge of body mass, which can be difficult to
estimate reliably for fossil taxa."