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Bipedalism, flight, and the evolution of theropod locomotor diversity
Abstract
The evolution of theropod flight has been characterized as a shift from one to three locomotor modules. Basal theropods, which were terrestrial bipeds, had a single locomotor module composed of the hind limb and tail. In birds, aerial locomotion was acquired with the origination of the wing module and a decoupling of the hind limb and tail into separate pelvic and caudal modules. This increase in modularity is thought to have granted birds more locomotor “options” than non-avian theropods. More specifically, an aerial locomotor system could have eased constraints on the hind limb and allowed specialization for habitats and lifestyles unavailable to non-birds. If so, bird hind limbs should be more disparate than those of non-avian theropods. We addressed this hypothesis by visualizing one aspect of limb design, the proportions of the three main segments, using ternary diagrams. Our results show that avian hind limb proportions are much more disparate than those of non-avian theropods. This broad range of limb design correlates with a radiation in locomotor diversity founded on three locomotor modules. We propose that birds have reached regions of proportion morphospace that are off limits to bipeds with only one locomotor module. In comparison, the limbs of non-avian theropods are conservatively proportioned. Despite great variation in body size, theropods other than birds do not exhibit specializations for locomotion other than terrestrial bipedalism. Although other aspects of size and shape need to be analyzed, the relationship between modular flexibility and morphological disparity appears to play an important role in theropod locomotor evolution.
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“The proportions of the manual phalanges are consistent with a specialized grasping function present in deinonychosaurs [65].”
From The ecology of Archaeopteryx (2026) · cites this paper as [65]
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“Pedal phalanges are shorter distally within each digit and pedal proportions plot intermediate between arboreal and terrestrial species, close to taxa like extant columbiforms, suggesting broad and generalized locomotive capabilities [65].”
From The ecology of Archaeopteryx (2026) · cites this paper as [65]
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To Hop or Not to Hop? The Answer Is in the Bird Trees
Provini, P., & Höfling, E. (2020) · Systematic Biology · doi:10.1093/sysbio/syaa015 · Cited by 22
Eliason, C. M., Proffitt, J. V., & Clarke, J. A. (2022) · Evolution · doi:10.1093/evolut/qpac039 · Cited by 22
Nudds, R. L., Atterholt, J., Wang, X., You, H., & Dyke, G. J. (2012) · Journal of Evolutionary Biology · doi:10.1111/jeb.12036 · Cited by 21
Forearm Posture and Mobility in Quadrupedal Dinosaurs
VanBuren, C. S., & Bonnan, M. F. (2013) · PLoS ONE · doi:10.1371/journal.pone.0074842 · Cited by 21
A juvenile anuran from the Lower Cretaceous Jiufotang Formation, Liaoning, China
Wang, Y., Jones, M. E. H., & Evans, S. E. (2007) · Cretaceous Research · doi:10.1016/j.cretres.2006.07.003 · Cited by 20
Falk, A. R., Lamsdell, J. C., & Gong, E. (2021) · Paleobiology · doi:10.1017/pab.2020.39 · Cited by 20
Metadata
- Authors
- Gatesy, S. M., & Middleton, K. M.
- Affiliations
- Brown University (US)
- Venue
- Journal of Vertebrate Paleontology
- Publisher
- Informa UK Limited
- Published
- 1997-06-19
- Volume
- 17
- Issue
- 2
- Pages
- 308-329
- ISSN
- 0272-4634 (ISSN-L)
- 1937-2809
- OpenAlex
- W2028889824
- Cited by
- 209
- References
- 78
- Citations by year
- 2026: 7 · 2025: 7 · 2024: 9 · 2023: 18 · 2022: 11 · 2021: 11 · 2020: 12 · 2019: 7 · 2018: 17 · 2017: 7 · 2016: 11 · 2015: 4 · 2014: 7 · 2013: 9 · 2012: 9
- FWCI
- 4.81
- Type
- journal article
- Language
- en
- Open access
- closed
- Locations
- Journal of Vertebrate Paleontology (publishedVersion)
- Topics
- Physical Sciences › Earth and Planetary Sciences › Paleontology › Paleontology and Evolutionary Biology
- Life Sciences › Agricultural and Biological Sciences › Ecology, Evolution, Behavior and Systematics › Morphological variations and asymmetry
- Physical Sciences › Earth and Planetary Sciences › Paleontology › Evolution and Paleontology Studies
- Keywords
- theropod dinosaurs, bipedalism, morphological disparity, flight evolution, morphospace
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