Functional constraints on the number and shape of flight feathers
Open accessAbstract
As a fundamental ecological aspect of most organisms, locomotor function significantly constrains morphology. At the same time, the evolution of novel locomotor abilities has produced dramatic morphological transformations, initiating some of the most significant diversifications in life history. Despite significant new fossil evidence, it remains unclear whether volant locomotion had a single or multiple origins in pennaraptoran dinosaurs and the volant abilities of individual taxa are controversial. The evolution of powered flight in modern birds involved exaptation of feathered surfaces extending off the limbs and tail yet most studies concerning flight potential in pennaraptorans do not account for the structure and morphology of the wing feathers themselves. Analysis of the number and shape of remex and rectrix feathers across a large dataset of extant birds indicates that the number of remiges and rectrices and the degree of primary vane asymmetry strongly correlate with locomotor ability revealing important functional constraints. Among these traits, phenotypic flexibility varies reflected by the different rates at which morphological changes evolve, such that some traits reflect the ancestral condition, whereas others reflect current locomotor function. While Mesozoic birds and Microraptor have remex morphologies consistent with extant volant birds, that of anchiornithines deviate significantly providing strong evidence this clade was not volant. The results of these analyses support a single origin of dinosaurian flight and indicate the early stages of feathered wing evolution are not sampled by the currently available fossil record.
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The primaries of Archaeopteryx have an asymmetry value of 3.47.
“The primaries have an asymmetry value of 3.47, within the range of volant neornithines (2.5-6.63) [5, 14].”
From The ecology of Archaeopteryx (2026) · cites this paper as [5]
The Results section on primary feather asymmetry places Archaeopteryx and other stem birds within the asymmetry range of volant crown birds, and gives that range as 2.50–6.63. It does not give 3.47 as Archaeopteryx’s value; the reported 2.72–4.50 range covers multiple stem-bird taxa. (LLM, conf=0.86)
Experimental — accuracy not yet measuredThe primary feathers of Archaeopteryx exhibit a degree of vane asymmetry within the range of extant volant birds, whereas these feathers are nearly symmetrical in non-avian pennaraptorans with the exception of the volant Microraptor.
“the primary feathers exhibit a degree of vane asymmetry that is within the range of extant volant birds, whereas these feathers are nearly symmetrical in non-avian pennaraptorans with the exception of the volant Microraptor [5, 20]”
From The ecology of Archaeopteryx (2026) · cites this paper as [5]
Anchiornis was non-volant.
“it is increasingly clear from the morphology of the skeleton, forelimb feathers, molt pattern, and ontogenetic changes in proportions that this taxon was non-volant [2, 4-6]”
From The ecology of Archaeopteryx (2026) · cites this paper as [5]
The Archaeopteryx wing is formed by 11 primaries.
“As in volant neornithines (9-11 primaries), the Archaeopteryx wing is formed by 11 primaries, although the 11th specimen appears to have 12 [5].”
From The ecology of Archaeopteryx (2026) · cites this paper as [5]
The 11th specimen of Archaeopteryx appears to have 12 primaries.
“As in volant neornithines (9-11 primaries), the Archaeopteryx wing is formed by 11 primaries, although the 11th specimen appears to have 12 [5].”
From The ecology of Archaeopteryx (2026) · cites this paper as [5]
The primary asymmetry value of 3.47 is within the range of volant neornithines (2.5-6.63).
“The primaries have an asymmetry value of 3.47, within the range of volant neornithines (2.5-6.63) [5, 14].”
From The ecology of Archaeopteryx (2026) · cites this paper as [5]
Interpretations regarding flight ability are additionally supported by bone density and cross-sectional geometry, and brain morphology.
“Interpretations regarding flight ability are additionally supported by bone density and cross-sectional geometry, and brain morphology [2, 5, 16, 22].”
From The ecology of Archaeopteryx (2026) · cites this paper as [5]
The Abstract links flight ability to flight-feather number and vane asymmetry, while Methods > Statistical Analysis examines feather traits and ulna length. Searches for bone density, cross-sectional geometry, and brain morphology found no passage in our copy saying that those features support flight-ability interpretations. (LLM, conf=0.76)
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Self-citationFlight, symmetry and barb angle evolution in the feathers of birds and other dinosaurs
Wang, X., Tang, H. K., & Clarke, J. A. (2019) · Biology Letters · doi:10.1098/rsbl.2019.0622 · Cited by 5 · see citing works
Zhao, T., Li, Z., Zhang, H., & Pan, Y. (2022) · Journal of Palaeogeography · doi:10.1016/j.jop.2022.08.002 · Cited by 4 · see citing works
Flagged first, then by citations
Research works printed but not linked: 4
- [66]L. Jenni, R. Winkler, The Biology of Moult in Birds (Bloomsbury Publishing, 2020).
- [68]W. Wang, J. K. O’Connor, Morphological coevolution of the pygostyle and tail feathers in Early Cretaceous birds. Vertebr. Palasiat. 55, 1 (2017).
- [76]P. Pyle, Identification Guide to North American Birds: Anatidae to Alcidae (Slate Creek Press, 2008).
- [95]J. K. O’Connor, Z. Xiao-Ting, H. U. Han, W. Xiao-Li, Z. Zhong-He, The morphology of Chiappeavis magnapremaxillo (Pengornithidae: Enantiornithes) and a comparison of aerodynamic function in Early Cretaceous avian tail fans. Vertebr. Palasiat. 55, 41 (2017).
Cited by
Sorted by citations
Reorganization of the theropod wrist preceded the origin of avian flight
Napoli, J. G., Fabbri, M., Ruebenstahl, A. A., O’Connor, J. K., Bhullar, B.-A. S., & Norell, M. A. (2025) · Nature · doi:10.1038/s41586-025-09232-3 · Cited by 17 · see citing works
Self-citationThe origin and early evolution of feathers: implications, uncertainties and future prospects
Xu, X., & Barrett, P. M. (2025) · Biology Letters · doi:10.1098/rsbl.2024.0517 · Cited by 10 · see citing works
Wang, R., & Pei, R. (2024) · Historical Biology · doi:10.1080/08912963.2024.2385604 · Cited by 10 · see citing works
Enhanced flight performance and adaptive evolution of Mesozoic giant cicadas
Xu, C., Chen, J., Muijres, F. T., Yu, Y., Jarzembowski, E. A., Zhang, H., & Wáng, B. (2024) · Science Advances · doi:10.1126/sciadv.adr2201 · Cited by 7 · see citing works
Earliest evidence of avian primary feather moult
Wang, X., O'Connor, J., Zheng, X., Wang, Y., & Kiat, Y. (2024) · Biology Letters · doi:10.1098/rsbl.2024.0106 · Cited by 6 · see citing works
Self-citationMorphlight Theory Inspired by Raptor: Arm‐Wing Cooperative Control Method and its Application
Tang, D., Zhao, Y., Wang, K., Chen, K., Liu, Y., Dai, Y., Zheng, C., & Yu, D. (2025) · Journal of Field Robotics · doi:10.1002/rob.22532 · Cited by 5 · see citing works
Avian features of Archaeopteryx feeding apparatus reflect elevated demands of flight
O’Connor, J. K., Clark, A. D., Kuo, P.-C., Wang, M., Shinya, A., Van Beek, C., & Chang, H. (2026) · The Innovation · doi:10.1016/j.xinn.2025.101086 · Cited by 5 · see citing works
Self-citationDinosaur palaeoneurology: an evolving science
Balanoff, A. M. (2024) · Biology Letters · doi:10.1098/rsbl.2024.0472 · Cited by 7 · see citing works
Inferring aerial behavior in Mesozoic dinosaurs: Implications and uncertainties
Xu, X. (2024) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.2401482121 · Cited by 4 · see citing works
Chotard, M., Wang, X., Zheng, X., Kaye, T. G., Grosmougin, M., Barlow, L. A., Kundrát, M., Dececchi, T. A., Habib, M., Zariwala, J., et al. (2025) · BMC Ecology and Evolution · doi:10.1186/s12862-025-02372-0 · Cited by 4 · see citing works
Wing morphology of Anchiornis huxleyi and the evolution of molt strategies in paravian dinosaurs
Kiat, Y., Wang, X., Zheng, X., Wang, Y., & O’Connor, J. (2025) · Communications Biology · doi:10.1038/s42003-025-09019-2 · Cited by 3 · see citing works
Self-citationEvaluating variation in Solnhofen avialans
O’Connor, J., & Marugán-Lobón, J. (2026) · Biology Letters · doi:10.1098/rsbl.2025.0601 · Cited by 3 · see citing works
Self-citationO'Connor, J. K. (2025) · Current Biology · doi:10.1016/j.cub.2025.02.019 · Cited by 3 · see citing works
Self-citationTheoretical morphospace reveals mixed optimisation of the avian wing planform for flight style
Walters, B. D., Liu, Y., Rayfield, E. J., & Donoghue, P. C. J. (2026) · Nature Communications · doi:10.1038/s41467-026-70692-w · Cited by 2 · see citing works
Size of the ulna and taxonomic order affect vane dimensions of secondary remiges in birds
Deeming, D. C., Bayliss, E. P., & Ketley, E. (2025) · Avian Research · doi:10.1016/j.avrs.2025.100245 · Cited by 2 · see citing works
Saitta, E. T., Balaji, L., Mitchell, J. S., & Makovicky, P. J. (2025) · Evolution · doi:10.1093/evolut/qpaf020 · Cited by 1 · see citing works
The effects of annual cycle stages and life-history traits on feather abrasion
Kiat, Y., Dupriez, Q., Pyo, G., Fox, O. J. L., Paz, R., Eilts, H., Talabante, C., & Sapir, N. (2025) · Scientific Reports · doi:10.1038/s41598-025-13458-6 · Cited by 1 · see citing works
Self-citationHyperelongate ornamental tail feathers in a new early Cretaceous enantiornithine bird
Clark, A. D., O'Connor, J., Wang, X.-L., Wang, Y., Pruett-Jones, S., Zhang, X.-Y., Wang, X., Zheng, X.-T., & Zhou, Z.-H. (2026) · PLoS ONE · doi:10.1371/journal.pone.0347641 · Cited by 1 · see citing works
Self-citationO’Connor, J. K., & Clark, A. D. (2026) · Discover Ecology · doi:10.1007/s44396-026-00026-z
Self-citationChen, A., van Tuinen, M., & Field, D. J. (2024) · Elsevier eBooks · doi:10.1016/b978-0-443-15750-9.00054-9
Bayesian Least-Squares Supertrees (BLeSS): Flexible Inference of Large Time-Calibrated PhylogeniesPreprint
Černý, D., & Slater, G. J. (2024) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2024.11.29.625936
Reyes-Ramírez, E. D., Díaz-Cháux, J. T., & Valencia, A. V. (2025) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2025.01.28.635237
The potential of fossil primary feather in taxonomic and ecological prediction
Zhang, J., Du, B., Peng, J., Fu, Y., Zhang, M., Cai, J., Wei, M., & Li, A. (2025) · Cretaceous Research · doi:10.1016/j.cretres.2025.106191
Valencia, A. V., Díaz-Cháux, J. T., Bonilla, M. A., Navarro-Morales, A., Giraldo, L. P., & Casanoves, F. (2026) · Biodiversity and Conservation · doi:10.1007/s10531-026-03337-5
Theoretical Morphology of Avian Wing Planform Reveals Variable Optimisation to Flight StylePreprint
Walters, B. D., Liu, Y., Rayfield, E. J., & Donoghue, P. C. J. (2025) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.64898/2025.12.16.692902
McNeil, D. J., Rodewald, A. D., Bennett, R. E., Rosenberg, K. V., Fiss, C. J., Rohwer, V. G., Dhondt, A. A., & Larkin, J. L. (2026) · Ibis · doi:10.1111/ibi.70036
Osváth, G., David, D.-C., Vargancsik, D., Nagy, L. J., Fehér, A. A., Kovács, Z. Z., Lendvai, Á. Z., Vincze, O., Nudds, R. L., Vágási, C. I., et al. (2026) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.64898/2026.06.08.730791
Intraspecific variability in feather moult extent declines under both low and high constraints
Steinberg, A., & Kiat, Y. (2026) · Functional Ecology · doi:10.1111/1365-2435.70397
Self-citationProgress and future directions in dinosaur palaeontology
Xu, X., Upchurch, P., Zanno, L. E., McNamara, M. E., Qin, Z., & Benton, M. (2026) · Nature Reviews Biodiversity · doi:10.1038/s44358-026-00191-9
Bird feathers from a Late Cretaceous coprolite
O'Connor, J. K., DeMar, D. G., Carroll, N. R., Chin, K., Smith, E. A., Holland, M., Clark, A. D., Cooper, C., Kuo, P.-C., Tobin, T. S., et al. (2026) · Current Biology · doi:10.1016/j.cub.2026.08.054
Self-citation
- Authors
- Kiat, Y., & O’Connor, J. K.
- Affiliations
- Field Museum of Natural History (US)
- Negaunee Integrative Research Center, Field Museum of Natural History, Chicago, IL 60605
- Corresponding author
- Kiat, Yosef
- Venue
- Proceedings of the National Academy of Sciences
- Publisher
- National Academy of Sciences
- Published
- 2024-02-12
- Volume
- 121
- Issue
- 8
- Pages
- e2306639121-e2306639121
- ISSN
- 0027-8424 (ISSN-L)
- 1091-6490
- PMC
- PMC10895369
- OpenAlex
- W4391757139
- Cited by
- 30
- References
- 96
- Citations by year
- 2026: 10 · 2025: 13 · 2024: 7
- FWCI
- 9.43
- Type
- journal article
- Language
- en
- Open access
- hybrid
- doi.org
- Licence
- Locations
- Proceedings of the National Academy of Sciences (publishedVersion, cc-by-nc-nd)
- PubMed Central (publishedVersion, cc-by-nc-nd)
- PubMed (publishedVersion)
- Funders
- Field Museum (NA)
- Zuckerman STEM (NA)
- Topics
- Physical Sciences › Earth and Planetary Sciences › Paleontology › Paleontology and Evolutionary Biology
- Physical Sciences › Environmental Science › Ecology › Avian ecology and behavior
- Health Sciences › Medicine › Urology › Comparative Animal Anatomy Studies
- Keywords
- flight feathers, remiges, rectrices, Microraptor, Mesozoic birds, exaptation, phenotypic flexibility, functional constraints
- MeSH terms
- Animals, Birds, Biological Evolution, Feathers, Flight, Animal, Fossils, Locomotion, Phylogeny, Wings, Animal, Dinosaurs
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