The DOI matches its Crossref record; 78 of the 78 references it cites were found in a registry or an index.
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.
Overview
Integrity signals
Registry
Cites
Statistics
Verification
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?
In the citing paper
“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]
In the citing paper
“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]
Citations
Cites
Phylogenies and the Comparative Method
Felsenstein, J. (1985) · The American Naturalist · doi:10.1086/284325 · Cited by 9,793
History, A. M. O. N. (2019) · American Museum Novitates · doi:10.1206/novi · Cited by 4,144
Smithsonian Miscellaneous Collections
(1924) · Nature · doi:10.1038/114085a0 · Cited by 2,336
Procedures for the Analysis of Comparative Data Using Phylogenetically Independent Contrasts
Garland, T., Harvey, P. H., & Ives, A. R. (1992) · Systematic Biology · doi:10.1093/sysbio/41.1.18 · Cited by 2,151
(2013) · Chinese Science Bulletin · doi:10.1007/11434.1861-9541 · Cited by 1,955
Saurischian monophyly and the origin of birds
Gauthier, J. (1986) · The Academy · doi:10.5281/zenodo.16171435 · Cited by 1,294
The Bulletin of the American Museum of Natural History
Gratacap, L. P. (1907) · Science · doi:10.1126/science.25.629.99 · Cited by 1,258
Functional morphology in vertebrate paleontology
(1995) · Choice Reviews Online · doi:10.5860/choice.32-6223 · Cited by 1,162
Predatory Dinosaurs from the Sahara and Late Cretaceous Faunal Differentiation
Sereno, P. C., Dutheil, D. B., Iarochène, M., Larsson, H. C. E., Lyon, G. H., Magwene, P. M., Sidor, C. A., Varricchio, D. J., & Wilson, J. A. (1996) · Science · doi:10.1126/science.272.5264.986 · Cited by 417
A new carnosaur (Dinosauria, Theropoda) from the Jurassic of Xinjiang, People's Republic of China
Currie, P., & Zhao, X.-J. (1993) · Canadian Journal of Earth Sciences · doi:10.1139/e93-179 · Cited by 397
SOME LOCOMOTORY ADAPTATIONS IN MAMMALS.
Smith, J. M., & Savage, R. J. (1956) · Journal of the Linnean Society of London Zoology · doi:10.1111/j.1096-3642.1956.tb02220.x · Cited by 370
Archaeopteryx and the origin of birds
OSTROM, J. H. (1976) · Biological Journal of the Linnean Society · doi:10.1111/j.1095-8312.1976.tb00244.x · Cited by 361
Discordance and concordance between morphological and taxonomic diversity
Foote, M. (1993) · Paleobiology · doi:10.1017/s0094837300015864 · Cited by 351
Caudofemoral musculature and the evolution of theropod locomotion
Gatesy, S. M. (1990) · Paleobiology · doi:10.1017/s0094837300009866 · Cited by 312
Self-citationGilmore, C. W. (1920) · Bulletin of the United States National Museum · doi:10.5479/si.03629236.110.i · Cited by 311
A classification for the birds of the world
Wetmore, A. B. (1960) · DSpace Repository (Smithsonian) · Cited by 290
Locomotor modules and the evolution of avian flight
Gatesy, S. M., & Dial, K. P. (1996) · Evolution · doi:10.1111/j.1558-5646.1996.tb04496.x · Cited by 290
Self-citationTheoretical Aspects of Cursorial Adaptations in Dinosaurs
Coombs, W. P. (1978) · The Quarterly Review of Biology · doi:10.1086/410790 · Cited by 289
Does metatarsal/femur ratio predict maximal running speed in cursorial mammals?
Garland, T., & Janis, C. M. (1993) · Journal of Zoology · doi:10.1111/j.1469-7998.1993.tb02626.x · Cited by 285
Dilophosaurus Wetherilli (Dinosauria, Theropoda). Osteology and comparisons
Welles, S. P. (1984) · Palaeontographica Abteilung A · Cited by 266
The phylogenetic position of the Tyrannosauridae: implications for theropod systematics
Holtz, T. R. (1994) · Journal of Paleontology · doi:10.1017/s0022336000026706 · Cited by 255
Novas, F. E. (1994) · Journal of Vertebrate Paleontology · doi:10.1080/02724634.1994.10011523 · Cited by 247
The first 85 million years of avian evolution
Chiappe, L. M. (1995) · Nature · doi:10.1038/378349a0 · Cited by 240
Gould, S. J. (1991) · Paleobiology · doi:10.1017/s0094837300010745 · Cited by 233
Ostrich Dinosaurs from the Late Cretaceous of Western Canada
Russell, D. A. (1972) · Canadian Journal of Earth Sciences · doi:10.1139/e72-031 · Cited by 216
NOTES ON THE PRINCIPLES OF QUADRUPEDAL LOCOMOTION AND ON THE MECHANISM OF HE LIMBS IN HOOFED ANIMALS
Gregory, W. K. (1912) · Annals of the New York Academy of Sciences · doi:10.1111/j.1749-6632.1912.tb55164.x · Cited by 207
(1926) · University of Michigan Library Repository · Cited by 200
Holtz, T. R. (1995) · Journal of Vertebrate Paleontology · doi:10.1080/02724634.1995.10011574 · Cited by 183
Russell, D. A., & Dong, Z.-M. (1993) · Canadian Journal of Earth Sciences · doi:10.1139/e93-187 · Cited by 181
Early Cretaceous Dinosaurs from the Sahara
Sereno, P., Wilson, J. A., Larsson, H., Dutheil, D., & Sues, H. (1994) · Science · doi:10.1126/science.266.5183.267 · Cited by 180
Osmólska, H., Roniewicz, E., & Барсболд, Р. (1972) · Cited by 178
Papers in avian paleontology, honoring Pierce Brodkorb
Brodkorb, P., & Campbell, K. E. (1992) · Medical Entomology and Zoology · Cited by 171
The Osteology of Compsognathus longipes WAGNER
Ostrom, J. H. (1978) · Biodiversity Heritage Library (Smithsonian Institution) · doi:10.5281/zenodo.16246150 · Cited by 169
Odontornithes: a Monograph of the Extinct Toothed Birds of North America
Marsh, O. C. (1880) · Nature · doi:10.1038/022457a0 · Cited by 159
Early Evolution of Avian Flight and Perching: New Evidence from the Lower Cretaceous of China
Sereno, P. C., & Chenggang, R. (1992) · Science · doi:10.1126/science.255.5046.845 · Cited by 159
A beaked bird from the Jurassic of China
Hou, L., Zhou, Z., Martin, L. D., & Feduccia, A. (1995) · Nature · doi:10.1038/377616a0 · Cited by 150
Norberg, U. M. (1979) · Philosophical Transactions of the Royal Society of London. B, Biological Sciences · doi:10.1098/rstb.1979.0054 · Cited by 150
Feduccia, J. A. (1980) · Medical Entomology and Zoology · Cited by 147
Butler, P. J. (1991) · Journal of Experimental Biology · doi:10.1242/jeb.160.1.233 · Cited by 147
The titanotheres of ancient Wyoming, Dakota, and Nebraska
Osborn, H. F. (1929) · Monographs of the United States Geological Survey · doi:10.5962/bhl.title.36431 · Cited by 146
Testing evolutionary constraint hypotheses with early Paleozoic gastropods
Wagner, P. J. (1995) · Paleobiology · doi:10.1017/s0094837300013294 · Cited by 138
The Titanotheres of ancient Wyoming, Dakota, and Nebraska
Osborn, H. F. (1929) · doi:10.3133/m55 · Cited by 138
Adaptations for Locomotion and Feeding in the Anhinga and the Double-Crested Cormorant
Owre, O. T. (1967) · Ornithological Monographs · doi:10.2307/40166666 · Cited by 137
Adaptations and Comparative Anatomy of the Locomotor Apparatus of New World Vultures
Fisher, H. I. (1946) · The American Midland Naturalist · doi:10.2307/2421553 · Cited by 135
Acrocanthosaurus atokensis, a New Genus and Species of Lower Cretaceous Theropoda from Oklahoma
Stovall, J. W., & Langston, W. (1950) · The American Midland Naturalist · doi:10.2307/2421859 · Cited by 129
Skeletal Adaptations of Ornitholestes, Struthiomimus, Tyrannosaurus
Osborn, H. F. (1917) · Zenodo (CERN European Organization for Nuclear Research) · doi:10.5281/zenodo.1040384 · Cited by 127
The Cretaceous Theropodous dinosaur Gorgosaurus
Lambe, L. (1917) · doi:10.4095/101672 · Cited by 124
Genera of Tertiary and recent rhynchonelloid brachiopods
Wetmore, A. B. (1959) · Miscellaneous Publications · Cited by 121
The affinities of a new theropod from the Alxa Desert, Inner Mongolia, People's Republic of China
Russell, D. A., & Dong, Z.-M. (1993) · Canadian Journal of Earth Sciences · doi:10.1139/e93-183 · Cited by 120
Hind limb scaling in birds and other theropods: Implications for terrestrial locomotion
Gatesy, S. M. (1991) · Journal of Morphology · doi:10.1002/jmor.1052090107 · Cited by 120
Self-citationCzerkas, S. M., & Olson, E. C. (1987) · Medical Entomology and Zoology · Cited by 114
(1979) · PubMed · Cited by 112
Perimeter-based Fourier analysis: a new morphometric method applied to the trilobite cranidium
Foote, M. (1989) · Journal of Paleontology · doi:10.1017/s0022336000036556 · Cited by 93
FLIGHTLESSNESS IN STEAMER‐DUCKS (ANATIDAE:TACHYERES): ITS MORPHOLOGICAL BASES AND PROBABLE EVOLUTION
Livezey, B. C., & Humphrey, P. S. (1986) · Evolution · doi:10.1111/j.1558-5646.1986.tb00506.x · Cited by 92
Allometry of the leg bones of moas (Dinornithes) and other birds
Alexander, R. M. (1983) · Journal of Zoology · doi:10.1111/j.1469-7998.1983.tb05785.x · Cited by 90
A juvenile specimen of Saurornithoides mongoliensis from the Upper Cretaceous of northern China
Currie, P., & Peng, J.-H. (1993) · Canadian Journal of Earth Sciences · doi:10.1139/e93-193 · Cited by 79
PRELIMINARY REPORT ON A MESOZOIC BIRD FROM LIAONING, CHINA
周忠和, 金帆, & 张江永 (1992) · Chinese Science Bulletin · doi:10.1360/sb1992-37-16-1365 · Cited by 74
ON A NEW SPECIMEN OF THE LOWER CRETACEOUS THEROPOD DINOSAUR DEINONYCHUS ANTIRRHOPUS.
Ostrom, J. H. (1976) · Biodiversity Heritage Library (Smithsonian Institution) · doi:10.5962/bhl.part.216 · Cited by 74
LIVEZEY, B. C. (1992) · Zoological Journal of the Linnean Society · doi:10.1111/j.1096-3642.1992.tb01229.x · Cited by 68
The Skeleton of Baptornis advenus (Aves: Hesperornithiformes)
MARTIN, L. D., & Tate, J. (1976) · Biodiversity Heritage Library (Smithsonian Institution) · doi:10.5281/zenodo.16342628 · Cited by 68
The Skeleton of Baptornis advenus (Aves: Hesperornithiformes)
Martin, L. D., & Tate, J. (1976) · Smithsonian contributions to paleobiology · doi:10.5479/si.00810266.27.35 · Cited by 66
A radiation of coly-like birds from the Eocene of North America (Aves: Sandcoleiformes new order)
Houde, P., & Olson, S. L. (1992) · Smithsonian Digital Repository (Smithsonian Institution) · Cited by 62
Altangerel, P., Chiappe, L. M., Барсболд, Р., Clark, J. M., Norell, M. A., & Akademi., M. S. U. (1994) · American Museum Novitates · Cited by 61
Mechanical constraints on the evolution of flight
Pennycuick, C. J. (1986) · Bristol Research (University of Bristol) · Cited by 54
DAS FUENFTE SKELETTEXEMPLAR VON ARCHAEOPTERYX.
Wellnhofer, P. (1974) · Palaeontographica Abteilung A · Cited by 54
Flightlessness in grebes (Aves, Podicipedidae): its independent evolution in three genera
Livezey, B. C. (1989) · Evolution · doi:10.1111/j.1558-5646.1989.tb04205.x · Cited by 51
PRELIMINARY REPORT ON A MESOZOIC BIRD FROM LIAONING, CHINA
Zhou, Z. (1992) · Cited by 50
The Species of Moas (Aves: Dinornithidae)
Cracraft, J. L. (1976) · Smithsonian contributions to paleobiology · doi:10.5479/si.00810266.27.189 · Cited by 43
The Comparative Functional Morphology of the Pelvic Appendage in Three Genera of Cuculidae
Berger, A. J. (1952) · The American Midland Naturalist · doi:10.2307/2422033 · Cited by 43
Palaeotis weigelti restudied : a small middle Eocene Ostrich (Aves : Struthioniformes)
Houde, P., & Haubold, H. (1987) · Palaeovertebrata · doi:10.18563/pv.17.2.27-42 · Cited by 41
Morphology of flightlessness in Chendytes, fossil seaducks (Anatidae: Mergini) of coastal California
Livezey, B. C. (1993) · Journal of Vertebrate Paleontology · doi:10.1080/02724634.1993.10011500 · Cited by 38
A Pterosaur from the Oldman Formation (Cretaceous) of Alberta
Russell, D. A. (1972) · Canadian Journal of Earth Sciences · doi:10.1139/e72-119 · Cited by 34
Variation in bone length and limb proportions in the coot (Fulica americana)
Engels, W. L. (1938) · Journal of Morphology · doi:10.1002/jmor.1050620309 · Cited by 13
Procompsognathus --- Theropod or Thecodont?
Ostrom, J. H. (1981) · Palaeontographica Abteilung A · Cited by 11
The Pleistocene Pied-billed Grebes (Aves: Podicipedidae)
Storer, R. W. (1976) · Smithsonian contributions to paleobiology · doi:10.5479/si.00810266.27.147 · Cited by 8
Tetraonidae and phasianidae of the USSR: ecology and morphology
Kuz'mina, & Siegel-Causey, D. (1992) · Cited by 7
Tetraonidae and phasianidae of the USSR : ecology and morphology
Kuz'mina, M. A., & Causey, D. (1992) · doi:10.5962/bhl.title.46317 · Cited by 5
Research works printed but not linked: 42
- Alexander R. McN., 1983, London, 200, 215
- Barsbold R., 1990, The Dinosauria, 249
- Bidar A., 1972, Annales du Museum D'Histoire Naturelle de Nice, 1, 1
- Camp C. L., 1936, University of California Publications in the Geological Sciences, 24, 39
- Cohn, J. M. W. 1968.The convergent flight mechanism of swifts (Apodi) and hummingbirds (Trochili) (Aves)201 Ph.D. dissertation, University of Michigan, Ann Arbor
- Colbert E. H., 1989, Museum of Northern Arizona Bulletin, 57, 1
- Feduccia, A. 1980.The Age of Birds196Cambridge: Harvard University Press.
- Garland T., 1993, London, 229, 133
- 10.2307/2410804 · DOI
- Gauthier J., 1986, Memoirs of the California Academy of Sciences, 8, 1
- Gilmore C. W., 1920, Bulletin of the United States National Museum, 110, 1
- Hecht M. K., 1985, The Beginnings of Birds, Proceedings of the International Archae-opteryx Conference, Eichstätt 1984
- Hesse A., 1992, Papers in Avian Paleontology. Honoring Pierce Brodkorb, 171
- Houde P., 1987, Palaeovertebrata, 17, 27
- Houde P., 1992, Papers in Avian Paleontology. Honoring Pierce Brodkorb, 137
- Lambe L. M., 1917, Geological Survey of Canada Memoir, 100, 1
- 10.2307/2409162 · DOI
- 10.2307/2408576 · DOI · Paper page
- Lowe P. R., 1928, 12th Series, 4, 99
- Maleev E. A., 1974, Paleontologicheskaia Ekspeditsiia, 1, 132
- Manion, B. L. 1984.The effects of size and growth on the hindlimb locomotion in the chicken201 Ph.D. dissertation, University of Illinois at Chicago, Chicago
- Marsh O. C., 1880, Report of the United States Geological Exploration of the Fortieth Parallel, 7, 1
- Mattison R. G., 1993, Journal of Vertebrate Paleontology, 13, 49A
- Mattison R. G., 1989, Journal of Vertebrate Paleontology, 9, 32A
- Miller L., 1949, Carnegie Institution of Washington Publication, 584, 201
- Mourer-Chauviré C., 1992, Papers in Avian Paleontology. Honoring Pierce Brodkorb, 68
- Olson S. L., 1992, Papers in Avian Paleontology. Honoring Pierce Brodkorb, 127
- Osborn H. F., 1916, Bulletin of the American Museum of Natura “History, 35, 733
- Osmólska H., 1972, Palaeontologia Polonica, 27, 103
- Ostrom J. H., 1976, Breviora, 439, 1
- Ostrom J. H., 1978, Zitteliana, 4, 73
- Ostrom J. H., 1979, American Scientist, 67, 45
- Ostrom J. H., 1981, Paleontographica A, 175, 179
- Ostrom J. H., 1986, Memoirs of the California Academy of Sciences, 8, 73
- Pennycuick C. J., 1986, Memoirs of the California Academy of Sciences, 8, 83
- Raath M. A., 1969, Arnoldia, 4, 1
- Russell D. A., 1970, Publications in Paleontology, 1, 1
- Russell D. A., 1993, National Geographic Research and Exploration, 9, 70
- Welles S. P., 1984, Palaeontographica A, 185, 85
- Wellnhofer P., 1974, Palaeontographica A, 147, 169
- Wellnhofer P., 1988, Papers in Avian Paleontology. Honoring Pierce Brodkorb, 1
- Wellnhofer P., 1993, Archaeopteryx, 11, 1
Cited by
Sorted by citations
Dyke, G. J., & Norell (2026) · Acta Palaeontologica Polonica · doi:10.4202/app · Cited by 2,807
TESTING FOR DIFFERENT RATES OF CONTINUOUS TRAIT EVOLUTION USING LIKELIHOOD
O’Meara, B. C., Ané, C., Sanderson, M. J., & Wainwright, P. C. (2006) · Evolution · doi:10.1111/j.0014-3820.2006.tb01171.x · Cited by 715
Functional versus morphological diversity in macroevolution
Wainwright, P. C. (2007) · Annual Review of Ecology Evolution and Systematics · doi:10.1146/annurev.ecolsys.38.091206.095706 · Cited by 373
A Review of Dromaeosaurid Systematics and Paravian Phylogeny
Turner, A. H., Makovicky, P. J., & Norell, M. A. (2012) · Bulletin of the American Museum of Natural History · doi:10.1206/748.1 · Cited by 350
Getting a grip on tetrapod grasping: form, function, and evolution
Sustaita, D., Pouydebat, E., Manzano, A. S., Abdala, V. S. L., Hertel, F., & Herrel, A. (2013) · Biological reviews/Biological reviews of the Cambridge Philosophical Society · doi:10.1111/brv.12010 · Cited by 221
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
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