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A new Mongolian tyrannosauroid and the evolution of Eutyrannosauria

Nature volume 642, pages 973–979 (2025)Cite this article

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Abstract

Eutyrannosaurians were large predatory dinosaurs that dominated Asian and North American terrestrial faunas in latest Cretaceous times. These apex predators arose from smaller-bodied tyrannosauroids during the ‘middle’ Cretaceous that are poorly known owing to the paucity of fossil material1,2,3. Here we report on a new tyrannosauroid, Khankhuuluu mongoliensis gen. et sp. nov., from lower Upper Cretaceous deposits of Mongolia that provides a new perspective on eutyrannosaurian origins and evolution. Phylogenetic analyses recover Khankhuuluu immediately outside Eutyrannosauria and recover the massive, deep-snouted Tyrannosaurini and the smaller, gracile, shallow-snouted Alioramini as highly derived eutyrannosaurian sister clades. Khankhuuluu and the late-diverging Alioramini independently share features related to a shallow skull and gracile build with juvenile eutyrannosaurians, reinforcing the key role heterochrony had in eutyrannosaurian evolution. Although eutyrannosaurians were mainly influenced by peramorphosis or accelerated growth4,5,6,7,8,9,10, Alioramini is revealed as a derived lineage that retained immature features through paedomorphosis and is not a more basal lineage as widely accepted11,12,13,14,15,16,17,18,19. Our results reveal that Asian tyrannosauroids (similar to Khankhuuluu) dispersed to North America, giving rise to Eutyrannosauria in the mid-Late Cretaceous. Eutyrannosauria diversified and remained exclusively in North America until a single dispersal to Asia in the latest Cretaceous that established Alioramini and Tyrannosaurini. Stark morphological differences between Alioramini and Tyrannosaurini probably evolved due to divergent heterochronic trends—paedomorphosis versus peramorphosis, respectively—allowing them to coexist in Asia and occupy different ecological niches.

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Fig. 1: Skeletal elements and anatomy of K. mongoliensis (MPC-D 100/50 and MPC-D 100/51).
Fig. 2: Skeletal evidence for peramorphic influences on the origin of eutyrannosaurian traits.
Fig. 3: Evolution of Tyrannosauroidea.

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Data availability

All data files relevant to our analyses and results are provided in the Article, its Supplementary Information and at Dryad59 (https://doi.org/10.5061/dryad.fj6q5744h). These files include our phylogenetic character dataset (.xlsx and .nex), phylogenetic characters illustrated graphically (.pdf), rationalization of cut-off values used for proportional characters (.xlsx), resultant tree files (.tre) and a file (log.txt) providing Templeton’s test results. The holotype and referred specimens of K. mongoliensis are catalogued and available for study to qualified researchers at the Institute of Paleontology, Mongolian Academy of Sciences in Ulaanbaatar, Mongolia.

Code availability

All R scripts and corresponding data files for replication of our time-calibration and ancestral state estimation used to reproduce our results are provided in the Supplementary Information and in the associated source data files located at Dryad59 (https://doi.org/10.5061/dryad.fj6q5744h).

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Acknowledgements

We thank U. Sanjaadash and I. Damdinsuren for specimen access and assistance; P. Makovicky, X. Xing and J. Shu’an for photographs of specimens. D.K.Z. thanks the musician/guitarist Slash for his unwavering support and encouragement. Funding was provided by the Natural Sciences and Engineering Research Council Discovery Grant (RGPIN 04854) to D.K.Z., the Japan Society for the Promotion of Science KAKENHI (JP23K03557) to Y.K., and by an Izaak Walton Killam Memorial Research Scholarship to J.T.V.

Author information

Authors and Affiliations

  1. Department of Earth, Energy and Environment, University of Calgary, Calgary, Alberta, Canada

    Jared T. Voris, Darla K. Zelenitsky & François Therrien

  2. Hokkaido University Museum, Hokkaido University, Sapporo, Japan

    Yoshitsugu Kobayashi

  3. Department of Biology, Cape Breton University, Sydney, Nova Scotia, Canada

    Sean P. Modesto

  4. Royal Tyrrell Museum of Palaeontology, Drumheller, Alberta, Canada

    François Therrien

  5. Department of Natural History Sciences, Graduate School of Science, Hokkaido University, Sapporo, Japan

    Hiroki Tsutsumi

  6. Department of Biological Sciences, North Carolina State University, Raleigh, NC, USA

    Tsogtbaatar Chinzorig

  7. Institute of Paleontology, Mongolian Academy of Sciences, Ulaanbaatar, Mongolia

    Tsogtbaatar Chinzorig & Khishigjav Tsogtbaatar

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Contributions

J.T.V. and D.K.Z. conceived and designed the project. J.T.V. and D.K.Z. collected and compiled data and performed analyses. Y.K., S.P.M., F.T, H.T., T.C. and K.T. provided data and support for analyses. The original manuscript draft was written by J.T.V. and D.K.Z. All of the authors contributed to manuscript revisions. J.T.V., D.K.Z., Y.K., H.T., T.C. and K.T. examined key specimens.

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Correspondence to Jared T. Voris or Darla K. Zelenitsky.

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Extended data figures and tables

Extended Data Fig. 1 Diagnostic and phylogenetically significant characters of Khankhuuluu. Autapomorphies of Khankhuuluu in yellow.

Synapomorphies of Khankhuuluu+Eutyrannosauria or characters restricted to Eutyrannosauria in magenta. Nasals of Khankhuuluu MPC-D 100/50 (1a, 2a) and Gorgosaurus ROM 1247 (1b, 2b) in dorsal (1a, b) and ventral (2a, b) views. Quadratojugals of Khankhuuluu (3a) and Daspletosaurus (3b). Scapulocoracoid of Khankhuuluu MPC-D 100/50 (4a) and Albertosaurus TMP 1986.064.0001 (4b). Quadrates of Khankhuuluu MPC-D 100/50 (5a) and MPC-D 100/51 (6a) and Gorgosaurus TMP 1986.144.0001 (5b and 6b) in lateral (5) and posterior (6) views. Distal tibia of Khankhuuluu MPC-D 100/51 (7a) and Albertosaurus TMP 1986.064.0001 (7b) in anterior view. Scale bars = 5 cm. Abbreviations: aco, astragalar contact orientation changes across angle (unambiguous synapomorphy); edp, expanded dorsal process of quadratojugal; lcp, lacrimal cornual process; mfp, medial frontal process; npr, small nasal pneumatic recess present; nr, nasal rugosities; qf, quadrate foramen; qjc, quadratojugal contact medial margin; qr, quadrate recess; qrwa, quadrate recess aperture medial wall absent; qrwp, quadrate recess aperture medial wall present; sad, deep subacromial depression; sfp, supernumerary frontal processes present medial frontal process absent; sgf, subglenoid fossa ventral face present.

Extended Data Fig. 2

Cladogram with examples of Tyrannosaurinae skulls, including: a, Bistahieversor, b, Daspletosaurus, c, Alioramus (Alioramini), and d, Tyrannosaurus (Tyrannosaurini). Alioramini is supported as both a highly derived member of Tyrannosaurinae and as a sister taxon to the Tyrannosaurini by numerous unambiguous and exclusive synapomorphies. See Supplementary Information 8 for a full list of synapomorphies.

Extended Data Fig. 3 Phylogeny of Tyrannosauroidea following a priori pruning of taxa scoring for less than 20% of characters.

Taxa excluded from the comprehensive analysis depicted in Fig. 3 are Kileskus, Juratyrant, Sinotyrannus, Stokesososaurus, Timurlengia, Alectrosaurus, Dryptosaurus, Nanuqsaurus, Thanatotheristes, and Zhuchengtyrannus. Values adjacent to nodes denote node support values: Bremer supports (left values < 20), bootstrap indices (right values > 50). Bremer support values and bootstrap indices increase significantly for most clades relative to those clades in the comprehensive analysis in Fig. 3. Among derived tyrannosaurine taxa (e.g., Alioramini+Tyrannosaurini), higher node values are found here relative to similar positions in the comprehensive tree in Fig. 3. This reveals that lower overall support values in the comprehensive tree is primarily due to ambiguity incurred in character state transformations with the inclusion of highly fragmentary taxa, particularly in late-diverging tyrannosauroids (e.g., Nanuqsaurus [8.1% of characters scored], Thanatotheristes [19.0%] Zhuchengtyrannus [10.3%]).

Supplementary information

Supplementary Information

Supporting information for main text includes measurements and ontogenetic assessment for Khankhuuluu mongoliensis, supplementary discussions for the phylogenetic position of Alioramini, phylogenetic protocols including taxon and character omissions, character list, clade synapomorphies, Templeton’s test results, and additional tests of time calibration.

Reporting Summary

Supplementary Data

Character scoresheet: phylogenetic character scoresheet.

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Voris, J.T., Zelenitsky, D.K., Kobayashi, Y. et al. A new Mongolian tyrannosauroid and the evolution of Eutyrannosauria. Nature 642, 973–979 (2025). https://doi.org/10.1038/s41586-025-08964-6

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