Exceptional dinosaur fossils reveal early origin of avian-style digestion
Open accessAbstract
Birds have a highly specialized and efficient digestive system, but when this system originated remains uncertain. Here we report six gastric pellets attributable to the recently discovered 160-million-year-old troodontid dinosaur Anchiornis, which is among the key taxa for understanding the transition to birds. The gastric pellets contain lightly acid-etched lizard bones or fish scales, and some are associated with Anchiornis skeletons or even situated within the oesophagus. Anchiornis is the earliest and most basal theropod known to have produced gastric pellets. In combination with other lines of evidence, the pellets suggest that a digestive system resembling that of modern birds was already present in basal members of the Paraves, a clade including troodontids, dromaeosaurids, and birds, and that the evolution of modern avian digestion may have been related to the appearance of aerial locomotion in this lineage.
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These differences may reflect the elevated energetic cost associated with the appearance of flight in Aves.
“These differences may reflect the elevated energetic cost associated with the appearance of flight in Aves [85].”
From The ecology of Archaeopteryx (2026) · cites this paper as [85]
Archaeopteryx preserves three feeding-related structures that are not observed in nonvolant theropods and two of which are currently only recognized in birds: a primitive bill-tip organ, oral papillae, and an ossified basihyal.
“Archaeopteryx preserves three feeding related structures that are not observed in nonvolant theropods and two of which are currently only recognized in birds: a primitive bill-tip organ, oral papillae, and an ossified basihyal [85] (Fig.3).”
From The ecology of Archaeopteryx (2026) · cites this paper as [85]
The Results describe gastric pellets in Anchiornis, and the Discussion addresses digestive features in theropods; neither addresses the claimed structures in Archaeopteryx. Searches for “bill-tip organ,” “oral papillae,” and “ossified basihyal” found no supporting passage in our copy. The paper’s silence on these structures is not a contradiction. (LLM, conf=0.75)
Experimental — accuracy not yet measuredThe bill-tip organ in Archaeopteryx and other toothed Mesozoic birds differs from that observed in living birds, and is considered an early morphotype representing an intermediate evolutionary step in the evolution of the modern bill-tip organ.
“The bill-tip organ in Archaeopteryx and other toothed Mesozoic birds differs from that observed in living birds, and is considered an early morphotype representing an intermediate evolutionary step in the evolution of the modern bill-tip organ [85].”
From The ecology of Archaeopteryx (2026) · cites this paper as [85]
Split into 2 atoms — 0 confirmed, 0 partially supported, 0 disputed, 0 misattributed, 2 unverifiable. Expand the claim to see per-atom verdicts.
Experimental — accuracy not yet measuredThe Chicago specimen of Archaeopteryx preserves soft tissue traces along the medial margin of the left palatine, interpreted as remnants of choanal oral papillae.
“This specimen also preserves soft tissue traces along the medial margin of the left palatine interpreted as remnants of choanal oral papillae [85] (Fig.3).”
From The ecology of Archaeopteryx (2026) · cites this paper as [85]
The Results describe Anchiornis specimens and their preserved gastric pellets, not the Chicago specimen of Archaeopteryx. Searches of our copy for the Chicago specimen, left palatine, soft-tissue traces, and choanal oral papillae found no passage addressing the claim. (LLM, conf=0.76)
Experimental — accuracy not yet measuredAnchiornis lacks a bill-tip organ, ossified basihyal, and potentially oral papillae.
“It also lacks a bill-tip organ, ossified basihyal, and potentially oral papillae [85].”
From The ecology of Archaeopteryx (2026) · cites this paper as [85]
The Results describe Anchiornis specimens and gastric pellets, while the Discussion describes its diet, digestion, and relatively short snout; no page numbers are available in our copy. Searches for bill-tip organ, ossified basihyal, and oral papillae found no passage addressing whether Anchiornis had those structures. (LLM, conf=0.75)
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Cites
A pre-Archaeopteryx troodontid theropod from China with long feathers on the metatarsus
Hu, D., Hou, L., Zhang, L., & Xu, X. (2009) · Nature · doi:10.1038/nature08322 · Cited by 365 · see citing works
Self-citationAn integrative approach to understanding bird origins
Xu, X., Zhou, Z., Dudley, R., Mackem, S., Chuong, C.-M., Erickson, G. M., & Varricchio, D. J. (2014) · Science · doi:10.1126/science.1253293 · Cited by 319 · see citing works
Self-citationAn Archaeopteryx-like theropod from China and the origin of Avialae
Xu, X., You, H., Du, K., & Han, F. (2011) · Nature · doi:10.1038/nature10288 · Cited by 314 · see citing works
Self-citationBrusatte, S. L., Lloyd, G. T., Wang, S. C., & Norell, M. A. (2014) · Current Biology · doi:10.1016/j.cub.2014.08.034 · Cited by 313 · see citing works
Evolutionary origins of the avian brain
Balanoff, A. M., Bever, G. S., Rowe, T. B., & Norell, M. A. (2013) · Nature · doi:10.1038/nature12424 · Cited by 195 · see citing works
Linking the evolution of body shape and locomotor biomechanics in bird-line archosaurs
Allen, V., Bates, K. T., Li, Z., & Hutchinson, J. R. (2013) · Nature · doi:10.1038/nature12059 · Cited by 163 · see citing works
Flagged first, then by citations
Research works printed but not linked: 32
- [1]Duke GE Gastrointestinal physiology and nutrition in wild birds Proceedings of the Nutrition Society 1997 56 1049 1056 10.1079/PNS19970109 9483670 · DOI
- [2]McLelland, J. In Form and Function in Birds (eds King, A. S. & McLelland, J.) 69–181 (Academic Press, 1979).
- [3]Wang Min Zhou Zhonghe Sullivan Corwin A Fish-Eating Enantiornithine Bird from the Early Cretaceous of China Provides Evidence of Modern Avian Digestive Features Current Biology 2016 26 9 1170 1176 10.1016/j.cub.2016.02.055 27133872 · DOI
- [4]Zheng X Fossil evidence of avian crops from the Early Cretaceous of China Proceedings of National Academy of Sciences USA 2011 108 15904 15907 10.1073/pnas.1112694108 PMC3179114 21896733 · DOI
- [5]O’Connor JK Zhou ZH Early evolution of the biological bird: perspectives from new fossil discoveries in China Journal of Ornithology 2015 156 333 342 10.1007/s10336-015-1222-5 · DOI
- [6]Naish D The fossil record of bird behaviour Journal of Zoology 2014 292 268 280 10.1111/jzo.12113 · DOI
- [7]Chin K Tokaryk TT Erickson GM Calk LC A king-sized theropod coprolite Nature 1998 393 680 682 10.1038/31461 · DOI
- [8]Hone DWE Rauhut OWM Feeding behaviour and bone utilization by theropod dinosaurs Lethaia 2010 43 232 244 10.1111/j.1502-3931.2009.00187.x · DOI
- [9]Varricchio DJ Gut contents from a Cretaceous tyrannosaurid; implications for theropod dinosaur digestive tracts Journal of Paleontology 2001 75 401 406 10.1017/S0022336000018199 · DOI
- [11]Liu YQ Timing of the earliest known feathered dinosaurs and transitional pterosaurs older than the Jehol Biota Palaeogeography Palaeoclimatology Palaeoecology 2012 323–325 1 12 10.1016/j.bbr.2011.03.031 · DOI
- [12]Xu X A new feathered maniraptoran dinosaur fossil that fills a morphological gap in avian origin Chinese Science Bulletin 2009 54 430 435
- [14]Li QG Plumage color patterns of an extinct dinosaur Science 2010 327 1369 1372 10.1126/science.1186290 20133521 · DOI · Paper page
- [17]Xing L Abdominal contents from two large Early Cretaceous compsognathids (Dinosauria: Theropoda) demonstrate feeding on confuciusornithids and dromaeosaurids PLoS ONE 2012 7 e44012 10.1371/journal.pone.0044012 22952855 PMC3430630 · DOI
- [19]Sanz JL A Early Cretaceous pellet Nature 2001 409 998 999 10.1038/35059172 11234054 · DOI
- [20]Myhrvold NP A call to search for fossilised gastric pellets Historical Biology 2013 24 505 517 10.1080/08912963.2011.631703 · DOI
- [22]Northwood C Early Triassic coprolites from Australia and their palaeobiological significance Palaeontology 2005 48 49 68 10.1111/j.1475-4983.2004.00432.x · DOI
- [23]Zatoń M Coprolites of Late Triassic carnivorous vertebrates from Poland: An integrative approach Palaeogeography, Palaeoclimatology, Palaeoecology 2015 430 21 46 10.1016/j.palaeo.2015.04.009 · DOI
- [25]O’Connor J Zhou ZH Xu X Additional specimen of Microraptor provides unique evidence of dinosaurs preying on birds Proceedings of the National Academy of Science, USA 2011 108 19662 19665 10.1073/pnas.1117727108 PMC3241752 22106278 · DOI · Paper page
- [26]Xing L-D Piscivory in the feathered dinosaur Microraptor Evolution 2013 67 2441 2445 10.1111/evo.12119 23888864 · DOI
- [27]Larsson H Hone D Dececchi T Sullivan C Xu X The winged non-avian dinosaur Microraptor fed on mammals: Implications for the Jehol Biota ecosystem Journal of Vertebrate Paleontology 2010 30 114A
- [28]Xu X A Jurassic ceratosaur from China helps clarify avian digit homologies Nature 2009 459 940 944 10.1038/nature08124 19536256 · DOI
- [29]Makovicky, P. J., Kobayashi, Y. & Currie, P. J. In The Dinosauria (second edition) (eds Weishampel, D. B., Dodson, P. & Osmolska, H.) 137–150 (University of California Press, 2004).
- [30]Kobayashi Y Herbivorous diet in an ornithomimid dinosaur Nature 1999 402 480 481 10.1038/44999 · DOI
- [31]Osmólska, H., Currie, P. J. & Barsbold, R. In The Dinosauria 2nd edn (eds Weishampel, D. B., Dodson, P. & Osmólska, H.) 165–183 (University of California Press, 2004).
- [33]Farmer C Uriona T Olsen D Steenblik M Sanders K The right-to left shunt of crocodilians serves digestion Physilogical and Biochemical Zoology 2008 81 125 137 10.1086/524150 18194087 · DOI
- [35]Chen PJ Dong ZM Zhen SN An exceptionally well-preserved Theropod dinosaur from the Yixian Formation of China Nature 1998 391 147 152 10.1038/34356 · DOI
- [37]Zanno LE Makovicky PJ Herbivorous ecomorphology and specialization patterns in theropod dinosaur evolution Proceedings of the National Academy of Sciences of the United States of America 2011 108 232 237 10.1073/pnas.1011924108 21173263 PMC3017133 · DOI
- [38]Choiniere JN A basal alvarezsauroid theropod from the early Late Jurassic of Xinjiang, China Science 2010 327 571 574 10.1126/science.1182143 20110503 · DOI · Paper page
- [39]Choiniere J Clark JM Norell MA Xu X Cranial osteology of Haplocheirus sollers Choiniere et al., 2010 (Theropoda: Alvarezsauroidea) American Museum Novitates 2014 3816 1 44 10.1206/3816.1 · DOI
- [40]Senter P Function in the stunted forelimbs of Mononykus olecranus (Theropoda), a dinosaurian anteater Paleobiology 2005 31 373 381 10.1666/0094-8373(2005)031[0373:FITSFO]2.0.CO;2 · DOI
- [44]Li Q Melanosome evolution indicates a key physiological shift within feathered dinosaurs Nature 2014 507 350 353 10.1038/nature12973 24522537 · DOI
- [46]Hunt AP Lucas SG Classification of vertebrate coprolites and related trace fossils New Mexico Museum of Natural History and Science Bulletin 2012 57 137 146
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Pittman, M., Bell, P. R., Miller, C. V., Enriquez, N. J., Wang, X., Zheng, X., Tsang, L. R., Tse, Y. T., Landes, M., & Kaye, T. G. (2022) · Nature Communications · doi:10.1038/s41467-022-35039-1 · Cited by 34 · see citing works
Self-citationO’Connor, J. K., & Clark, A. D. (2026) · Discover Ecology · doi:10.1007/s44396-026-00026-z
- Authors
- Zheng, X., Wang, X., Sullivan, C., Zhang, X., Zhang, F., Wang, Y., Li, F., & Xu, X.
- Affiliations
- Linyi University (CN)
- University of Alberta (CA)
- Shandong Pingyi Tianyu Museum of Natural History (CN)
- Chinese Academy of Sciences (CN)
- Corresponding author
- Wang, Xiaoli
- Xu, Xing
- ORCID
- Venue
- Scientific Reports
- Published
- 2018-09-21
- Volume
- 8
- Issue
- 1
- Pages
- 14217-14217
- ISSN
- 2045-2322 (ISSN-L)
- PMC
- PMC6155034
- OpenAlex
- W2890462859
- Cited by
- 34
- References
- 6
- Citations by year
- 2026: 2 · 2025: 7 · 2024: 2 · 2023: 7 · 2022: 4 · 2021: 4 · 2020: 6 · 2019: 2
- FWCI
- 2.21
- Type
- journal article
- Language
- en
- Open access
- gold
- doi.org
- Locations
- Scientific Reports (publishedVersion, cc-by)
- Europe PMC (PubMed Central) (publishedVersion, cc-by)
- PubMed Central (publishedVersion, cc-by)
- PubMed (publishedVersion)
- DOAJ (DOAJ: Directory of Open Access Journals) (submittedVersion)
- Funders
- University of Alberta
- National Natural Science Foundation of China (41402017, 41120124002, 41472023, 41372014, 91514302, #41688103)
- Natural Sciences and Engineering Research Council of Canada (RGPIN 2017 ?, Discovery Grant (RGPIN–2017–06246), 06246, rgpin-2017-06246)
- Topics
- Physical Sciences › Earth and Planetary Sciences › Paleontology › Paleontology and Evolutionary Biology
- Physical Sciences › Earth and Planetary Sciences › Paleontology › Evolution and Paleontology Studies
- Life Sciences › Agricultural and Biological Sciences › Ecology, Evolution, Behavior and Systematics › Fossil Insects in Amber
- Keywords
- troodontids, digestive system evolution, Paraves, theropod dinosaurs, dinosaur-bird transition, aerial locomotion, esophagus, fish scales
- MeSH terms
- Animals, Birds, Bone and Bones, Digestive System, Biological Evolution, Fishes, Fossils, Lizards, Phylogeny, Dinosaurs
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