Research Themes
The Zanno Lab studies the history of life in deep time, from microstructure to macroevolution.
Reproductive Biology
Selected Funding
2024 European Synchrotron Radiation Facility (ESRF): 3D Architecture of Medullary Bone in Extinct and
Extant Avemetatarsalia: testing controversial reports of avian reproductive tissues in non-avian dinosaurs and
pterosaurs. (financial equivalent $110,000, PIs A Canoville, LE Zanno)
2024 – 2025 Stanford Synchrotron Radiation Lightsource (SSRL): Evolution of medullary bone: Identification
of reproductive tissues in the fossil record. (financial equivalent $333,400, PIs J Anne, A Canoville, Co-PIs
LE Zanno, N Edwards)
​
2021 Canadian Light Source (CLS): 3D histology and life history of Tyrannosaurus rex. (financial equivalent
$42,000, PIs E Snively, M Gilbert, Co-PIs H Woodward, LE Zanno)
​
2020 – 2022 Stanford Synchrotron Radiation Lightsource (SSRL): Testing new approaches to diagnose reproductive
bone tissues in birds and non-avian dinosaurs. (financial equivalent $480,000, PIs J Anne, A Canoville, Co-PIs
LE Zanno, MH Schweitzer, N Edwards, A Van Veelen)
2020 – 2023 Triangle Center for Evolutionary Medicine: A new approach for identifying pathologic bone in living
and extinct species. ($12,000, PI LE Zanno, Co-PIs E Hyland and C Walker)
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2019 American Ornithological Society Research Award: Do Female Penguins Deposit Medullary Bone? ($1,644,
PI A Canoville, Co-PIs LE Zanno, MH Schweitzer)
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2016 – 2020 National Science Foundation, Integrative Organismal Systems, Early-concept Grants for
Exploratory Research (EAGER, award #1552328): Experimental techniques for discerning female-specific
phenotypes in non-avian theropod dinosaurs. ($185,690, PI LE Zanno; w/Co-PI M Schweitzer)
Selected Publications
Hedge J, Bender E, Zanno LE. 2025. Novel Quantification of Eggshell Surfaces in Dromaius
novaehollandiae with Implications for the Fossil Eggshells of Oviraptorosauria (Dinosauria).
Ecology and Evolution. 15(5), e71260
Hedge J, Tucker RT, Makovicky PJ, Zanno LE. 2025. Fossil Eggshell Diversity of the Mussentuchit
Member, Cedar Mountain Formation, Utah. PLOS ONE 20(2): e0314689.
https://doi.org/10.1371/journal.pone.0314689
​
Anné J, Canoville A†, Edwards NP, Schweitzer MH, Zanno LE. 2023. Independent evidence for the
preservation of endogenous bone biochemistry in a specimen of Tyrannosaurus rex. Biology 12:264
​
Chinzorig T, Beguesse KA, Canoville A, Phillips G, Zanno LE. 2022. Chronic fracture and osteomyelitis in a large-bodied ornithomimosaur with implications for the identification of unusual endosteal bone in the fossil record. Anat Rec, special issue Dinosaurs: New Ideas From Old Bones 2022; 1–16 https://doi.org/10.1002/ar.25069
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Canoville A†, Zanno LE, Zheng M, Schweitzer M. 2021. Keratan sulfate as a marker for medullary bone. Journal of Anatomy, First published: 04 January 2021. https://doi.org/10.1111/joa.13388​
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Canoville A, Schweitzer M, Zanno LE. 2020 (Invited). Identifying medullary bone in extinct avemetatarsalians: challenges, implications, and perspectives. Philosophical Transactions B 375: 20190133. doi.org/10.1098/rstb.2019.0133
​​
Zanno LE, Avrahami HM, Gates TA, Tucker R, Makovicky PJ. 2019. Preliminary Data On Elongatoolithid Clutches And Associated Fauna From The Cenomanian Mussentuchit Member, Cedar Mountain Formation Of Utah. VII International Symposium on Dinosaur Eggs and Babies, Shiyan, China
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Canoville A, Schweitzer M, Zanno LE. 2019. Systemic distribution of medullary bone in the avian skeleton: ground truthing criteria for the identification of reproductive tissues in extinct Avemetatarsalia. BMC Evolutionary Biology 2019 19:71 DOI: 10.1186/s12862-019-1402-7​
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Schweitzer MH, Zheng W, Zanno LE, Sugiyama T. 2016. Chemistry supports the identification of gender- specific reproductive tissue in Tyranosaurus rex. Scientific Reports 6, Article number: 23099 (2016) doi:10.1038/srep23099
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Canoville A, Zanno LE, Zheng W, Schweitzer MH. 2017. Paleohistology Of A Gravid Oviraptorosaurian Dinosaur From The Upper Cretaceous Nanxiong Formation, China, With An Enigmatic Endosteal Tissue. J Vert Paleontol Progr Abstr, 2017, 93​
Mid-Cretaceous Thermal Maximum
Selected Funding
2025 – 2030 National Science Foundation, Biodiversity on a Changing Planet:
Collaborative Research: BoCP-Implementation: US-South Africa: Surviving A
Super Greenhouse: Terrestrial Biodiversity Dynamics During The Cretaceous
Thermal Maximum. ($3,280,488 PI LE Zanno)
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2023 – 2025 National Geographic Society (NGS-100601R-23): Evaluating the Impact of Climate Change on Asian Ecosystems during the mid-Cretaceous Thermal Maximum. ($99,995, PI LE Zanno, Co-PIs T Chinzorig, R Tucker, C Suarez, J Yoshida, K Tsogtbaatar)
2019 – 2025 National Science Foundation, Frontier Research in Earth Science (DES, FRES, award #1925973): Collaborative Research: Time of Transformation: integrating the dynamic geologic, climatic, and biotic systems of North America during the Early to Late Cretaceous transition. ($2,499,950, PIs LE Zanno, C Suarez, R Cifelli, M Suarez, P Makovicky)
Selected Publications
Chinzorig T, Takasaki R, Yoshida J, Tucker RT, Buyantegsh B, Mainbayar B, Tsogtbaatar K, Zanno LE. 2025.
A Domed Pachycephalosaur From the Early Cretaceous of Mongolia. Nature. 2025 Sep 17:1-8
​
Tucker RT, Venter KE, Lana C, Roberts EM, Chinzorig T, Tsogtbaatar K, Chiarenza AA, Zanno LE. 2025.
U-Pb calcite age dating of fossil eggshell, an accurate deep-time geochronometer. Communications
Earth & Environment.
Cilliers CD, Contreras D,Tucker, RT, Roychoudhury A, Zanno LE. Submitted. A new floral assemblage from
the Turonian–Coniacian Moreno Hill Formation, New Mexico, USA. GEOBIOS.
Zanno LE. Accepted. Mid-Cretaceous Earth. In: Zanno LE, Arbour VM, Holtz T. editors. The Complete Dinosaur, 3rd Edition: Indiana University Press.
​
Tucker RT, King MR, Delgerzaya P, Freimuth WJ, Chinzorig T, Tsogtbaatar K, Takasaki R, Saneyoshi M, Chiarenza AA, Zanno LE. Accepted. Geological and paleontological reconstruction of the Bayanshiree Formation, Eastern Gobi Basin, Mongolia.
van Niekerk JB, Tucker RT, Delgerzaya P, Freimuth WJ, Chinzorig T†, Tsogtbaatar K, Yoshida J, Zanno LE. 2025. Geological reassessment of syn-rift extensional sequences in the Shine Usny Tolgod and Dzun Shakhai fossil localities, Eastern Gobi Basin, Mongolia. Sedimentology 72, 1275–1315. https://doi.org/10.1111/sed.70003
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Avrahami HM, Makovicky PJ, Tucker, RT, Zanno LE. 2024. A New Semi-Fossorial Thescelosaurine Dinosaur from The Cenomanian-age Mussentuchit member of the Cedar Mountain Formation, Utah. The Anatomical Record. https://doi.org/10.1002/ar.25505
Tucker RT, King MR, Mohr MT, Renault RK, Crowley JI, Fekete JW, Makovicky PJ, Zanno LE. 2024. Tectono-sedimentation history of the upper Cedar Mountain Formation, Central Utah, USA. Sedimentology. https://doi.org/10.1111/sed.13211
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Cilliers CD, Tucker RT, Freimuth WJ^, Beguesse KA, Zanno LE. 2023. Geological assessment of Turonian-Coniacian terrestrial sedimentation records during climatic recovery, Moreno Hill Formation, Zuni Basin. The Mountain Geologist 60: 103–135. https://doi.org/10.31582/rmag.mg.60.3.103
Tucker RT, Crowley JI, Renault RK, Makovicky PJ, Cifelli RL, Zanno LE. 2023. Exceptional age constraint on a fossiliferous sedimentary archive preceding the Cretaceous Thermal Maximum. Geology 51: 962–967. https://doi.org/10.1130/G51278.1
Zanno LE, Gates TA, Avrahami HM, Tucker RT, Makovicky PJ. 2023. An early-diverging iguanodontian (Dinosauria: Rhabdodontomorpha) from the Late Cretaceous of North America. PLOS ONE 18(6): e0286042. https://doi.org/10.1371/journal.pone.0286042
Renault RK, Tucker RT, King MR, Crowley J, Hyland EG, Zanno LE. 2023. Dating the Greenhorn transgression and OAE2 of the Western Interior Seaway in central Utah: Implications for the absolute age of the Cenomanian-Turonian boundary. Cretaceous Research 2023: 105464.
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Chinzorig T , Cullen T, Phillips G, Lamb J, Larson PL, Rolke R, Zanno LE. 2022. Large-bodied ornithomimosaurs inhabited Appalachia during the Late Cretaceous of North America. PLOS ONE 17(10), e0266648.
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Tucker RT, Hyland EG, Gates TA, King MR, Roberts EM, Foley EK, Berndt D, Hanta R, Khansubha S-O, Aswasereelert W, Zanno LE. 2022. Age, depositional history, and paleoclimatic setting of the Early Cretaceous dinosaur assemblages from the Sao Khua Formation (Khorat Group), Thailand. Palaeogeography, Palaeoclimatology, Palaeoecology 111107.
Krumenacker LJ*, Zanno LE*, Sues HD. 2022. A partial tyrannosauroid femur from
the mid-Cretaceous Wayan Formation of eastern Idaho, USA. Journal of Paleontology
1–10 *these authors contributed equally.
Tucker R, Suarez CA, Makovicky PJ, Zanno LE. 2022. Paralic sedimentology of the
Mussentuchit Member Coastal Plain, Cedar Mountain Formation, Central Utah, USA.
Journal of Sedimentary Research, 92(6), 546–569.
Cilliers CD, Tucker RT, Crowley JL, Zanno LE. 2021. Age constraint for the Moreno
Hill Formation (Zuni Basin) by CA-TIMS and LA-ICP-MS detrital zircon
geochronology. PeerJ 9:e10948.
King MR, La Croix AD, Gates, TA, Anderson P, Zanno LE. 2021. Glossifungites gingrasi sp. nov. a probable subaqueous insect domicile from the Cretaceous Ferron Sandstone, Utah. Journal of Paleontology, published online 1/25/2021. https://doi.org/10.1017/jpa.2020.115
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Tucker RT, Zanno LE, Huang H-Q, Makovicky PJ. 2020. A refined temporal framework for newly discovered fossil assemblages of the upper Cedar Mountain Formation (Mussentuchit Member), Mussentuchit Wash, Central Utah. Cretaceous Research. https://doi.org/10.1016/j.cretres.2020.104384
extinct Avemetatarsalia. BMC Evolutionary Biology 2019 19:71. https://doi.org/10.1186/s12862-019-1402-7
Zanno LE, Gates TA, Canoville A ,Tucker RT, Avrahami H, Makovicky PJ. 2019. Diminutive, fleet-footed tyrannosauroid narrows the 70-million-year gap in the fossil record of North American “tyrants.” Communications Biology 2: 64 (2019). https://doi.org/10.1038/s42003-019-0308-7§¶*
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Avrahami HM, Heckert AB, Gates TA, Makovicky PJ, Zanno LE. 2018. A new microvertebrate assemblage from the Mussentuchit Member, Cedar Mountain Formation: insights into the paleobiodiversity and paleobiogeography of early Late Cretaceous ecosystems in western North America. PeerJ 6:e5883. https://doi.org/10.7717/peerj.5883
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Gates TA, Tsogtbaatar K, Zanno LE, Chinzorig T, Watabe M. 2018. A new iguanodontian (Dinosauria: Ornithopoda) from the Early Cretaceous of Mongolia. PeerJ 6:e5300 https://doi.org/10.7717/peerj.5300
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King MR, Gates TA, Gingras MK, Zanno LE, Pemberton G. 2018. Transgressive erosion expressed as a Rhizocorallium-dominated xylic firmground: an example from the Blackhawk Formation, Utah. PALAIOS.
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McDonald AT, Gates TA, Zanno LE, Makovicky PJ. 2017. Anatomy, taphonomy, and phylogenetic implications of a new specimen of Eolambia caroljonesa (Dinosauria: Ornithopoda) from the Cedar Mountain Formation, Utah, USA. PLOS ONE 12(5): e0176896. https://doi.org/10.1371/journal.pone.0176896
Arbour VM, Zanno LE, Gates TA. 2016. Ankylosaurian dinosaur palaeoenvironmental associations are influenced by sea level fluctuation, extirpation, and geodispersal. Palaeogeography, Palaeoclimatology, Palaeoecology. https://doi.org/10.1016/j.palaeo.2016.02.033
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Zanno LE, Makovicky PJ. 2013. Neovenatorid theropods are apex predators in the Late Cretaceous of North America. Nature Comm. 4:2827. https://doi.org/10.1038/ncomms3827
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Zanno LE, Makovicky PJ. 2011. On the earliest record of Cretaceous tyrannosauroids in western North America: implications for an Early Cretaceous Laurasian interchange event. Hist Biol 23:317– 325. First published on: 24 February 2011 (iFirst). https://doi.org/10.1080/08912963.2010.543952.
New Species
Selected Publications
Zanno LE and Napoli JG. 2025. Nanotyrannus and Tyrannosaurus co-existed at the close of
the Cretaceous. Nature https://doi-org.prox.lib.ncsu.edu/10.1038/s41586-025-09801-6
​
Chinzorig T, Takasaki R, Yoshida J, Tucker RT, Buyantegsh B, Mainbayar B, Tsogtbaatar K,
Zanno LE. Accepted. A Domed Pachycephalosaur From the Early Cretaceous of Mongolia.
Nature. 2025 Sep 17:1-8
​
Avrahami HM, Makovicky PJ, Tucker, RT, Zanno LE. 2024. A New Semi-Fossorial
Thescelosaurine Dinosaur from The Cenomanian-age Mussentuchit member of the Cedar Mountain Formation, Utah. The Anatomical Record. https://doi.org/10.1002/ar.25505
​
Zanno LE, Gates TA, Avrahami HM^, Tucker RT, Makovicky PJ. 2023. An early-diverging iguanodontian (Dinosauria: Rhabdodontomorpha) from the Late Cretaceous of North America. PLOS ONE 18(6): e0286042. https://doi.org/10.1371/journal.pone.0286042
​
King MR, La Croix AD, Gates, TA, Anderson P, Zanno LE. 2021. Glossifungites gingrasi sp. nov. a probable subaqueous insect domicile from the Cretaceous Ferron Sandstone, Utah. Journal of Paleontology, published online 1/25/2021. https://doi.org/10.1017/jpa.2020.115
​
Zanno LE, Gates TA, Canoville A† ,Tucker RT, Avrahami H^, Makovicky PJ. 2019. Diminutive, fleet-footed tyrannosauroid narrows the 70-million-year gap in the fossil record of North American “tyrants.” Communications Biology 2: 64 (2019)
​
Gates TA, Tsogtbaatar K, Zanno LE, Chinzorig T, Watabe M. 2018. A new iguanodontian (Dinosauria: Ornithopoda) from the Early Cretaceous of Mongolia. PeerJ 6:e5300 https://doi.org/10.7717/peerj.5300
​​
Zanno LE, Drymala SD, Nesbitt SJ, Schneider VP. 2015. Early crocodylomorph increases top predator diversity during rise of dinosaurs. Sci. Rep. 5:9276. https://doi.org/10.1038/srep09276
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Zanno LE, Makovicky PJ. 2013. Neovenatorid theropods are apex predators in the Late Cretaceous of North America. Nature Comm. 4:2827. https://doi.org/10.1038/ncomms3827
​
Zanno LE, Varricchio DJ, O’Connor PM, Titus AL, Knell MJ. 2011. A new troodontid theropod, Talos sampsoni gen. et sp. nov., from the Upper Cretaceous Western Interior Basin of North America. PLOS ONE 6(9): e24487. https://doi.org/10.1371/journal.pone.0024487
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Zanno LE, Gillette DD, Albright LB, Titus AL. 2009. A new North American therizinosaurid and the role of herbivory in “predatory” dinosaur evolution. Proc R Soc B 276:3505–3511
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Gates TA, Sampson SD, Delgado de Jesus CR, Zanno LE, Eberth D, Hernandez-Rivera R, Martinez MA, Kirkland JI. 2008. Velafrons coahuilaensis, a new lambeosaurine hadrosaur (Dinosauria: Ornithopoda) from the late Campanian Cerro del Pueblo Formation, Coahuila, Mexico. J Vertebr Paleontol 27:917–930
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Kirkland JI, Zanno LE, Sampson SD, Clark JM, DeBlieux DD. 2005. A primitive therizinosauroid dinosaur from the Early Cretaceous of Utah. Nature 435:84–87
Zanno LE, Sampson SD. 2005. A new oviraptorosaur (Theropoda, Maniraptora) from the Late Cretaceous (Campanian) of Utah. J Vertebr Paleontol 25:897–904​​
End-Cretaceous Ecosystems
Selected Publications
Zanno LE. 2025. 2025. Dinosaur diversity before the asteroid.
Science Oct 23;390(6771):332-333. doi: 10.1126/science.aeb5725.
Epub 2025 Oct 23. PMID: 41129655
Zanno LE and Napoli JG. 2025. Nanotyrannus and Tyrannosaurus
co-existed at the close of the Cretaceous. Nature
https://doi-org.prox.lib.ncsu.edu/10.1038/s41586-025-09801-6
​
Roberts EM, Hendrix MS, Ramezani J, Clyde WC, Zippe P,
Hodgson S, Yuleridge V, Zanno LE. 2025. High precision
CA-ID-TIMS U-Pb zircon age for the Dueling Dinosaur locality, with implications for the basal age and duration of the Hell Creek Fm, Montana. bioRxiv 2025-07
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Wilkinson R, Cullen TM, Avrahami HM^, Braman D, Zanno LE, Currie PJ, Evans DC. Accepted. Sedimentology and paleontology of the Acheroraptor holotype locality, a multi-taxic bonebed in the Hell Creek Formation of Montana, USA
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Titus A, Irmis R.B, Sampson SD, Zanno LE, Albright LB, Sertich JJW, Roberts EM, Farke AA. 2025. Twenty-Five Years of Paleontological Research in Grand Staircase-Escalante National Monument, Utah. In Parks Stewardship Forum (Vol. 41, No. 3, pp. 402-424). George Wright Society
Carr TD*, Napoli JG*, Brusatte SL, Holtz TR, Hone DWE, Williamson TE, Zanno LE. 2022. Insufficient Evidence for Multiple Species of Tyrannosaurus in the Latest Cretaceous of North America: A Comment on “The Tyrant Lizard King, Queen and Emperor: Multiple Lines of Morphological and Stratigraphic Evidence Support Subtle Evolution and Probable Speciation Within the North American Genus Tyrannosaurus. Evolutionary Biology 2022: 1–15 *these authors contributed equally
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​Cullen TM, Zanno LE, Larson DW, Currie PJ, Evans DC. 2021. Theropod biodiversity patterns in the Dinosaur Park Formation (Late Cretaceous: Campanian) of Alberta revealed through morphometrics and biostratigraphy. Canadian Journal of Earth Sciences 30 June 2021. https://doi.org/10.1139/cjes-2020-0145
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Burgener L, Hyland E, Griffith E, Mitášová H, Zanno LE, Gates TA. 2021. An extreme climate gradient-induced ecological regionalization in the Upper Cretaceous Western Interior Basin of North America. GSA Bulletin. v. 133. https://doi.org/10.1130/B35904.1
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​Woodward HN, Tremaine K, Williams SA, Zanno LE, Horner JR, Myhrvold N. 2020. Growing up Tyrannosaurus rex: osteohistology refutes the pygmy ‘Nanotyrannus’ and supports ontogenetic niche partitioning in juvenile Tyrannosaurus. Science Advances 2020; 6: eaax6250
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Arbour VM, Zanno LE, Larson DW, Sues H-D. 2016. The furculae of the dromaeosaurid dinosaur Dakotaraptor steini are trionychid turtle entoplastra. PeerJ. 4:e1691. https://doi.org/10.7717/peerj.1691
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Gates TA, Zanno LE, Makovicky PJ. 2015. Theropod teeth from the upper Maastrichtian Hell Creek Formation “Sue” Quarry: new morphotypes and faunal comparisons. Acta Paleontol Pol
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Zanno LE, Loewen MA, Farke AA, Kim G-S, Claessens LPAM, McGarrity CTA. 2013. Late Cretaceous theropod dinosaurs of southern Utah. In: Titus A, Loewen MA. editors. At the Top of the Grand Staircase: the Late Cretaceous of southern Utah. Bloomington: Indiana University Press. Pp. 504–525
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Gates TA, Prieto-Márquez A, Zanno LE. 2012. Mountain Building Triggered Late Cretaceous North American Megaherbivore Dinosaur Radiation. PLOS ONE 7(8): e42135. (published 02 August 2012)
​
Gates TA, Sampson SD, Zanno LE, Roberts EM, Eaton JG, Nydam RL, Hutchison JH, Smith JA, Loewen MA, Getty MA. 2010. Biogeography of terrestrial and freshwater vertebrates from the Late Cretaceous (Campanian) Western Interior of North America. Palaeogeogr Paleoclimatol Paleoecol 291:371–387
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Zanno LE, Wiersma JP, Loewen MA, Sampson SD, Getty MA. 2010. Preliminary report on the theropod dinosaur fauna of the late Campanian Kaiparowits Formation, Grand Staircase- Escalante National Monument, Utah. Learning from the Land: Grand Staircase-Escalante Natl Monument Sci Symp Proc 2006 2:204–217
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Sampson SD, Gates TA, Roberts EM, Getty MA, Zanno LE, Loewen MA, Smith JA, Lund EK, Sertich J, Titus AL. 2010. Grand Staircase-Escalante National Monument: a new and critical window into the world of dinosaurs. Learning from the Land: Grand Staircase-Escalante Natl Monument Sci Symp Proc 2006 2:171–188​
Dinosaur Anatomy, Paleobiology, & Macroevolution
Selected Publications
Avrahami HA, Zanno LE. Submitted. Intraspecific variation in the skull of the thescelosaurid Fona
herzogae from The Cenomanian-age Mussentuchit Member of the Cedar Mountain Formation, Utah.
Freimuth WJ, Zanno LE. Accepted. New craniodental materials of Falcarius utahensis (Theropoda:
Therizinosauria) reveal patterns of intraspecific variation and cranial evolution in early
coelurosaurians. The Anatomical Record
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Choiniere JN, Chapelle KEJ, Ford DP, Balanoff A, Hendrickx C, Napoli JG, Radermacher VJ,
Zanno LE, Norell MA. Submitted. CT scanning and a revised phylogenetic analysis reveal
Ornitholestes hermanni as the earliest-branching oviraptorosaur
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Arbour V, Zanno LE, Holtz T. Accepted. Dinosaur diversity. In: Zanno LE, Arbour VM, Holtz T. editors. The Complete Dinosaur, 3rd Edition: Indiana University Press​
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Choiniere J, Zanno LE, Balanoff A. Accepted (Invited). Basal Maniraptora. In: Makovicky PJ, Carrano M, Barrett P, Weishampel D. editors. The Dinosauria, 3rd Edition: Cambridge University Press
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Button DJ, Zanno LE. 2023. Neuroanatomy of the late Cretaceous Thescelosaurus neglectus (Neornithischia: Thescelosauridae) reveals novel ecological specialisations within Dinosauria. Scientific Reports 13, 19224 (2023). https://doi.org/10.1038/s41598-023-45658-3
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Arbour VA, Zanno LE, Evans DC. 2022. Paleopathological evidence for intraspecific combat in ankylosaurid dinosaurs. Biology Letters. Published:07 December 2022 https://doi.org/10.1098/rsbl.2022.0404
Benson RJ, Brown CM, Campione NE, Cullen TM, Evans DC, Zanno LE. 2022. Comment on “The influence of juvenile dinosaurs on community structure and diversity.” Science 375(6578), eabj5976.
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Liao C-C, Zanno LE, Wang S, Xu X. 2021. Postcranial osteology of Beipiaosaurus inexpectus (Theropoda: Therizinosauria). PLoS ONE 16(9): e0257913​
Ksepka DT, Balanoff AM, Smith NA, Bever GS, Bhullar B-AS, Bourdon E, Braun EL, J. Burleigh JG, Clarke JA, Colbert MW, Corfield JR, Degrange FJ, De Pietri VL, Early CM, Field DJ, Gignac PM, Gold MEL, Kimball RT, Kawabe S, Lefebvre L, Marugán-Lobón J, Mongle CS, Morhardt A, Norell MA, Ridgely RC, Scofield RP, Tambussi CP, Torres CR, van Tuinen M, Walsh SA, Watanabe A, Witmer LM, Wright AK, Zanno LE, Jarvis ED, Smaers JB. 2020. Tempo and pattern of avian brain size evolution. Current Biology (2020). https://doi.org/10.1016/
j.cub.2020.03.060
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Button D, Zanno LE. 2020. Repeated Evolution of Divergent Modes of Herbivory in Non-avian Dinosaurs. Current Biology 30:1–11. https://doi.org/10.1016/j.cub.2019.10.050
Arbour V, Zanno LE. 2020. Tail weaponry in ankylosaurs and glyptodonts: An example of a rare but strongly convergent phenotype (invited contribution). The Anatomical Record. Published online 05 03 2019. https://doi.org/10.1002/ar.24093
Hedrick BP, Cordero SA, Zanno LE, Noto C, Dodson P. 2019. Quantifying shape and ecology in avian pedal claws: the relationship between the bony core and keratinous sheath. Ecology and Evolution. First published: 30 September 2019. https://doi.org/10.1002/ece3.5507
Arbour V, Zanno LE. 2018. The Evolution of Tail Weaponization in Amniotes. Proc R Soc B. Published 17 January 2018. https://doi.org/10.1098/rspb.2017.2299
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Button K, Hailu Y, Kirkland JI, Zanno LE. 2017. Incremental Growth of Therizinosaurian Dental Tissues:
implications for dietary transitions in Theropoda. PeerJ 5:e4129. https://doi.org/10.7717/peerj.4129
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Gates TA, Organ C, Zanno LE. 2016. Ornamentation and the rapid evolution of gigantic theropod dinosaurs.
Nature Comm 7:12931. https://doi.org/10.1038/ncomms12931
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Zanno LE, Tsogtbaatar K, Chinzorig T, Gates TA. 2016. Specializations of the Mandibular Anatomy and
Dentition of Segnosaurus galbinensis (Dinosauria: Theropoda). PeerJ 4:e1885.
https://doi.org/10.7717/peerj.1885
Hedrick BP, Zanno LE, Wolfe DG, Dodson P. 2015. The Slothful Claw: Osteology and Taphonomy of
Nothronychus mckinleyi and N. graffami (Dinosauria: Theropoda) and Anatomical Considerations for
Derived Therizinosaurids. PLOS ONE 10(6): e0129449. https://doi.org/10.1371/journal.pone.0129449
Lautenschlager S, Witmer LM, Altangerel P, Zanno LE, Rayfield EJ. 2014. Cranial anatomy of Erlikosaurus andrewsi (Dinosauria: Therizinosauria): new insights based on digital reconstruction. J Vert Paleontol 34: 1263–1291
​Zanno LE, Makovicky PJ. 2013. No evidence for directional evolution of body mass in herbivorous theropod dinosaurs. Proc R Soc B. 280(1751), 20122526. https://doi.org/10.1098/rspb.2012.2526 (published 28 November 2012)
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​Lautenschlager S, Rayfield EJ, Altangerel P, Zanno LE, Witmer LM. 2012. The Endocranial Anatomy of Therizinosauria and its Implications for Sensory and Cognitive Function. PLOS ONE 7(12): e52289. (published 19 December 2012). https://doi.org/10.1371/journal.pone.0052289
Zanno LE, Makovicky PJ. 2011. Herbivorous ecomorphology and specialization patterns in theropod dinosaur evolution. Proc Natl Acad Sci U S A 108:232–237; published ahead of print December 20, 2010. https://doi.org/10.1073/pnas.1011924108
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Makovicky PJ, Zanno LE. 2011. Theropod diversity and the refinement of avian characteristics. In Dyke G, Kaiser G, editors. Living Dinosaurs: The Evolutionary History of Modern Birds. Berkeley: University of California Press. pp. 9–29
Smith DK, Zanno LE, Sanders RK, Kirkland JI, DeBlieux DD. 2011. New information on the braincase of the North American therizinosaurian Falcarius utahensis. J Vertebr Paleontol 31:387–404
Zanno LE. 2010. A taxonomic and phylogenetic re-evaluation of Therizinosauria. J Syst Palaeontol 8:503–543
Zanno LE. 2010. Osteology of Falcarius utahensis: characterizing the anatomy of basal therizinosaurs. Zool J Linn Soc 158:196–230
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Zanno LE, Gillette DD, Albright LB, Titus AL. 2009. A new North American therizinosaurid and the role of herbivory in “predatory” dinosaur evolution. Proc R Soc B 276:3505–3511
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Zanno LE. 2006. The pectoral girdle and forelimb of the primitive therizinosauroid Falcarius utahensis (Theropoda, Maniraptora): analyzing evolutionary trends within Therizinosauroidea. 2006. J Vertebr Paleontol 26:636–650
Other Vertebrate Systems & Research Methods
Selected Publications
Smith, J.A., Dowding, E.M., Abdelhady, A.A., Abondio, P., Araújo, R., Aze, T., Balisi, M., Buatois,
L.A., Carvajal-Chitty, H., Chattopadhyay, D., Coiro, M., Dietl, G.P., González Arango, C., Kevrekidis,
C., Kimmig, J., Mychajliw, A.M., Pimiento, C., Regalado Fernández, O.R., Schroeder. K.M., Warnock,
R.C.M., Yang, T.R., Yasuhara, M., Akita, L.G., Allen, B.J., Anderson, B.M., Andréoletti, J., Archuby,
F.M., Ballen, G.A., Bari, M.I., Benton, M.J., Bergh, E.W., Brambilla, L., Brombacher, A., Chan, Y.K.S.,
Chiarenza, A.A., Chinzorig, T.†, Coates, K.M., Cordie, D.R., Cortés-Sánchez, M., Cruz-Vega, E.J.,
Cybulski, J.D., De Baets, K., De Entrambasaguas, J., Dillon, E.M., Du, A., Dunhill, A.M., Erlandson,
J.M., Forel, M.B., Foster, W.J., Gates, T.A., Gavryushkina, A., Grace, M.K., Grossart, H.P., Hänsel, P.,
Harnik, P.G., Hopkins, M.J., Hopkins, S., Hu, K., Huang, H.H.M., Irmis, R.B., Jaques, V.A.J., Jenkins,
X.A., Jukar, A.M., Kelley, P.H., Kihn, R.G., Klompmaker, A.A., Kocsis, Á.T., Kriwet, J., Lazarus, D.,
Liao, C.C., Lin, C.H., Louys, J., Lozano-Fernandez, J., Lozano-Francisco, M.C., Lueders-Dumont, J.A.,
Malvè, M.E., Martindale, R.C., Mazzini, I., Modenini, G., Mondal, S., Mondini, M., Monferran, M.D.,
Mulvey, L.P.A., Nanglu, K., Nguyen, J.M.T., Norris, R., O’Dea, A., Ollendorf, A.L., Orihuela, J.,
Pandolfi, J.M., Pereira, T., Piro, A., Plotnick, R.E., Plaza-Torres, S.M., Porto, A., Prieto-Márquez, A.,
Punyasena, S.W., Quental, T.B., Raja, N.B., Ranaivosoa, V., Ribas-Deulofeu, L., Rivals, F., Roden, V.J.,
Rosso, A., Saleh, F., Salvador, R.B., Saupe, E.E., Schneider, S., Sclafani, J.A., Smith, M.R., Souron, A.,
Steinbauer, M.J., Stewart, M., Tambussi, C.P., Thomas, E., Tschopp, E., Tütken, T., Varela, S., Vezzosi, R.I., Villaseñor, A., Weinkauf, M.F.G., Zanno, L.E., Zhang, C., Zhao, Q., Kießling, W. 2025. Identifying the big questions in paleontology: a community-driven project. Paleobiology 1-24. https://doi.org/10.1017/pab.2025.10042​
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​Young MT, Schwab JA, Dufeau D, Cowgill T, Witmer LM, Herrera Y, Walsh S., Dollman K., Choiniere J, Zanno LE, Xu X, Clark J, Brusatte SL. 2024. Skull sinuses precluded extinct crocodile relatives from cetacean-style deep diving as they transitioned from land-to-sea. Royal Society Open Science. 11(10): 241272
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Gates TA, Hengrui C, Hu Y, Han X, Griffith E, Burgener L, Hyland E, Zanno LE. 2022. Estimating ancient biogeographic patterns with statistical model discrimination. Anat Rec, special issue Dinosaurs: New Ideas From Old Bones 2022;1–16 https://doi.org/10.1002/ar.25067
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Miller* H, Avrahami HM, Zanno* LE. 2022. Dental pathology in lamniform and carcharhiniform sharks with comments on the classification and homology of double tooth pathologies in vertebrates. Peer J 10, e12775 *these authors contributed equally
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Kosch J, Zanno LE. 2020. Sampling impacts tooth growth and replacement rate calculations in Archosauria: implications for paleontological studies. PeerJ 8:e9918. https://doi.org/10.7717/peerj.9918
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Schwab JA, Young MT, Neenan JM, Walsh S, Witmer LM, Herrera Y, Zanno LE, Brusatte SL. 2020. Inner Ear Sensory System Changes as Extinct Crocodylomorphs Transitioned from Land to Water. Proc Natl Acad Sci USA; PNAS first published April 20, 2020. https://doi.org/10.1073/pnas.2002146117
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Hoffman DK, Heckert AB, Zanno LE. 2018. Disparate growth strategies within Aetosauria: novel histologic data from the aetosaur Coahomasuchus chathamensis. Anat Rec First Published 08 November 2018. 302.9 (2019): 1504–1515. https://doi.org/10.1002/ar.24019
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Hoffman DK, Heckert AB, Zanno LE. 2018 Under the armor: X-ray computed tomographic reconstruction of the internal skeleton of Coahomasuchus chathamensis (Archosauria: Aetosauria) from the Upper Triassic of North Carolina, USA, and a phylogenetic analysis of Aetosauria. PeerJ 6:e4368. https://doi.org/10.7717/peerj.4368
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Drymala SD, Zanno LE. 2016. Osteology of Carnufex carolinensis (Archosauria: Psuedosuchia) from the Pekin Formation of North Carolina and Its Implications for Early Crocodylomorph Evolution. PLOS ONE 11(6): e0157528. https://doi.org/10.1371/journal.pone.0157528
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Zanno LE, Drymala SD, Nesbitt SJ, Schneider VP. 2015. Early crocodylomorph increases top predator diversity during rise of dinosaurs. Sci. Rep. 5:9276. https://doi.org/10.1038/srep09276
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​ Zanno LE, Heckert AB, Kryzyzanowski SE, Lucas SG. 2002. Diminutive Metoposaurid skulls from the Upper Triassic Blue Hills (Adamanian: latest Carnian) of Arizona. In Heckert AB, Lucas SG, editors. Upper Triassic Stratigraphy and Paleontology. Albuquerque: New Mexico Museum of Natural History and Science Bulletin. 21:121–125
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STEM Engagement
Selected Funding
2021 – 2024 Institute of Museum and Library Studies: What do you want to be when you grow up? Using exhibits to
help students see themselves as scientists. ($250,000, Co-PIs W Lovelady, LE Zanno, S Downey, K Chandler)
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​2018 – 2026 Bank of America Charitable Foundation, Anchor Grant: Dueling Dinosaurs Worldwide Education Program.
($1,000,000, PI LE Zanno, Co-PI E Koster, SP L Baird)
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2018 – 2023 NC GlaxoSmithKline Foundation: Dueling Dinosaurs School Education Program. ($500,000, PIs L Baird,
LE Zanno, Co-PI E Koster)
2018 – 2024 Wells Fargo Grant: Dueling Dinosaurs Tactile Experience Exhibit. ($100,000, PIs R. Campbell, LE Zanno,
Co-PI J Stebben)
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2016 – 2019 National Science Foundation Improving Undergraduate STEM Education: Pathways into Geoscience
(IUSE: GEOPATHS-EXTRA, award # 1600545): Collaborative Research: Engaging Diverse Two-Year College
Geoscience Students: Expanding Opportunities Through Undergraduate Research and Mentoring. (PIs LE Zanno, G Miller, A Leinbach, J Fountain)
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2013 – 2019 National Science Foundation Math & Science Partnerships (MSP, award #1319293): Students Discover: Improving Middle School STEM Outcomes through Scaling Citizen Science Projects. ($7,764,481 PI R Dunn, SP LE Zanno (all PIs, CO-PIs, and SP not listed)
Selected Papers
Gates TA, Dunn RR, Thomas J, Bourne NW, Boyd RI, Elliot P, Gandee SR, Argall B, Clark C, Hall K, Shell L, Alford A, Zanno LE. 2025. Shark Tooth Forensics: A Participatory Science Initiative to Unravel Ancient Shark Ecology. The Bulletin of the Ecological Society of America, p.e2213
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Jones E, Kanipe J, Avrahami HA, Cullen T, Zanno LE. 2024. Evaluating The Efficacy Of A Public Science Project To Educate And Engage Students In Identifying Vertebrate Microfossils From The Upper Cretaceous Hell Creek Formation. Vert Paleontol, Progr Abstr, 2024
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Kanipe J, Jones E, Avrahami HA^, Cullen T, Zanno LE. 2024. Public Involvement In The Production of Microfossil Datasets: A Case Study of The Biodiversity and Methodology of The Public Science Project Cretaceous Creatures. Vert Paleontol, Progr Abstr, 2024
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Jones E, Kanipe J^, Avrahami HM^, Cullen T†, Zanno LE. 2023. Assessing Middle School Student Accuracy in Identifying Vertebrate Microfossils from the Upper Cretaceous Hell Creek Formation. J Vert Paleontol, Progr Abstr, 2023: 235–236
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Jones E, Gates TA, Zanno LE. 2023. Paleontology: Involving the public in one of the most popular, yet inaccessible, sciences. Citizen Science Association
