Cross sectional geometry of the forelimb skeleton and flight mode in pelecaniform birds
Abstract
Avian wing elements have been shown to experience both dorsoventral bending and torsional loads during flapping flight. However, not all birds use continuous flapping as a primary flight strategy. The pelecaniforms exhibit extraordinary diversity in flight mode, utilizing flapping, flap‐gliding, and soaring. Here we (1) characterize the cross‐sectional geometry of the three main wing bone (humerus, ulna, carpometacarpus), (2) use elements of beam theory to estimate resistance to loading, and (3) examine patterns of variation in hypothesized loading resistance relative to flight and diving mode in 16 species of pelecaniform birds. Patterns emerge that are common to all species, as well as some characteristics that are flight‐ and diving‐mode specific. In all birds examined, the distal most wing segment (carpometacarpus) is the most elliptical (relatively high Imax/Imin) at mid‐shaft, suggesting a shape optimized to resist bending loads in a dorsoventral direction. As primary flight feathers attach at an oblique angle relative to the long axis of the carpometacarpus, they are likely responsible for inducing bending of this element during flight. Moreover, among flight modes examined the flapping group (cormorants) exhibits more elliptical humeri and carpometacarpi than other flight modes, perhaps pertaining to the higher frequency of bending loads in these elements. The soaring birds (pelicans and gannets) exhibit wing elements with near‐circular cross‐sections and higher polar moments of area than in the flap and flap‐gliding birds, suggesting shapes optimized to offer increased resistance to torsional loads. This analysis of cross‐sectional geometry has enhanced our interpretation of how the wing elements are being loaded and ultimately how they are being used during normal activities. J. Morphol., 2011. © 2011 Wiley‐Liss,Inc.
Integrity signals
Registry
Cites
Statistics
Claims the paper makes
Claims other papers make about this one
Each is checked against this paper’s own text: does it say what the citing paper says it does?
Mass-normalised torsional resistance retains a small yet significant allometric effect that reflects the inherent proportional relation between flight adaptations and body size, and should thus not be removed when investigating locomotory affinities of extinct taxa.
“Furthermore, mass-normalised torsional resistance retains a small yet significant allometric effect that reflects the inherent proportional relation between flight adaptations and body size, and should thus not be removed when investigating the locomotory affinities of extinct taxa12,20.”
From Wing bone geometry reveals active flight in Archaeopteryx (2018) · cites this paper as [12]
The inverse scaling of relative torsional resistance with reliance on flapping may reflect either a multidirectional stress regime or elevated torsional loading experienced during gliding and soaring as opposed to the directionally confined stresses associated with flapping.
“This may reflect either a multidirectional stress regime or elevated torsional loading experienced during gliding and soaring as opposed to the directionally confined stresses associated with flapping12.”
From Wing bone geometry reveals active flight in Archaeopteryx (2018) · cites this paper as [12]
The ratio of Imax over Imin provides a reliable measure for the ellipticity of the transverse bone shaft.
“The ratio of Imax over Imin provides a reliable measure for the ellipticity of the transverse bone shaft and has been considered as such in biomechanical explorations12,60,66,67.”
From Wing bone geometry reveals active flight in Archaeopteryx (2018) · cites this paper as [12]
Values for J obtained from cross sections with an Imax/Imin >1.50 are typically overestimated but remain informative when considered proportionally rather than quantitatively.
“Values for J obtained from cross sections with an Imax/Imin >1.50 are typically overestimated12,60,61, but remain informative when considered proportionally rather than quantitatively (as is its derivative Zp41,61).”
From Wing bone geometry reveals active flight in Archaeopteryx (2018) · cites this paper as [12]
Cites
Phylogenies and the Comparative Method
Felsenstein, J. (1985) · The American Naturalist · doi:10.1086/284325 · Cited by 9,793 · see citing works
Carlson, K. J. (2005) · American Journal of Physical Anthropology · doi:10.1002/ajpa.20124 · Cited by 112 · see citing works
Flagged first, then by citations
Cited by
Sorted by citations
Wing bone geometry reveals active flight in Archaeopteryx
Voeten, D. F. A. E., Cubo, J., de Margerie, E., Röper, M., Beyrand, V., Bureš, S., Tafforeau, P., & Sanchez, S. (2018) · Nature Communications · doi:10.1038/s41467-018-03296-8 · Cited by 77 · see citing works
“Both referred avian flight classifications were complemented with the following locomotor categories: (6) long-tailed pterosaurian flight, (7) short-tailed pterosaurian flight, (8) (avian) non-volant wing-propelled diving, (9) ratite bipedal, (10) (non-avian) dinosaurian bipedal, (11) (non-avian) dinosaurian omnipedal…”
BackgroundFunctional Morphometric Analysis of the Furcula in Mesozoic Birds
Close, R. A., & Rayfield, E. J. (2012) · PLoS ONE · doi:10.1371/journal.pone.0036664 · Cited by 44 · see citing works
Morphological Disparity of the Humerus in Modern Birds
Serrano, F. J., Costa-Pérez, M., Navalón, G., & Martín-Serra, A. (2020) · Diversity · doi:10.3390/d12050173 · Cited by 30 · see citing works
- Authors
- Simons, E. L. R., Hieronymus, T. L., & O'Connor, P. M.
- Affiliations
- Midwestern University (US)
- Northeast Ohio Medical University (US)
- Corresponding author
- Simons, Erin L. R.
- Venue
- Journal of Morphology
- Publisher
- Wiley
- Published
- 2011-05-12
- Volume
- 272
- Issue
- 8
- Pages
- 958-971
- ISSN
- 0362-2525 (ISSN-L)
- 1097-4687
- OpenAlex
- W2121164226
- Cited by
- 55
- References
- 2
- Citations by year
- 2026: 1 · 2025: 6 · 2024: 1 · 2022: 2 · 2021: 2 · 2020: 6 · 2019: 3 · 2018: 6 · 2017: 2 · 2016: 3 · 2015: 8 · 2014: 6 · 2013: 7 · 2012: 2
- FWCI
- 5.77
- Type
- journal article
- Language
- en
- Open access
- closed
- Locations
- Journal of Morphology (publishedVersion)
- PubMed (publishedVersion)
- Topics
- Health Sciences › Medicine › Urology › Comparative Animal Anatomy Studies
- Physical Sciences › Earth and Planetary Sciences › Paleontology › Paleontology and Evolutionary Biology
- Physical Sciences › Engineering › Aerospace Engineering › Biomimetic flight and propulsion mechanisms
- Keywords
- flight modes, flapping flight, soaring flight, diving, humerus, ulna, carpometacarpus, bending loading, torsional loading, beam theory, cormorants, pelicans, gannets
- MeSH terms
- Animals, Birds, Bone and Bones, Diving, Flight, Animal, Humerus, Wings, Animal
About these checks
Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.
Statistics and arithmetic are recomputed, not judged.
Confidence bands are shown only for fields calibrated on our annotated test set. None is calibrated yet, so no band is shown.
What counts as a claim ›How we measure confidence ›How we check this ›
Are you an author, or do you see a problem with this page? Request a review