Luo Lab | University of Chicago


Stilson K. T., Luo Z.-X., Li, P., Olson, S., and C. F. Ross. 2023. Three-dimensional mandibular kinematics of mastication in the marsupial Didelphis virginiana. Philosophical Transactions of Royal Society, Series B. 378: 20220548 (

Li, P., Ross, C. F., Luo, Z.-X., and N. J. Gidmark. 2023. Head posture impacts mammalian hyoid position and suprahyoid muscle length: implication for swallowing biomechanics. Philosophical Transactions of Royal Society, Series B. 378: 20220552 (

Lautenschlager, S., Fagan, M.J., Luo, Z.X., Bird, C.M., Gill, P. and E.J. Rayfield. 2023. Functional reorganisation of the cranial skeleton during the cynodont–mammaliaform transition. Communications Biology 6, 367 (2023). (

Luo, Z.-X. and T. Martin. 2023. Mandibular and Dental Characteristics of the Late Jurassic Mammal Henkelotherium guimarotae(Paurodontidae, Dryolestida).  PalZ (Paläontologische Zeitschrift) 97: 569-619. ( 

Bendel, E.‑M., Kammerer, C. F., Luo, Z.X., Smith, R. M. H. and J. Fröbisch. 2022. The earliest segmental sternum in a Permian synapsid and its implications for the evolution of mammalian locomotion and ventilation. Scientific Reports (2022) 12:13472 (DOI:10.1038/s41598-022-17492-6).

Araújo, R., David, R., Benoit, J., Lungmus, J. K., Stoessel, A., Barrett, P. M., Maisano, J. A., Ekdale, E., Orliac, M., Luo, Z.-X,Martinelli, A. G., Hoffman, E. A., Sidor, C. A., Martins, R. M. S., Spoor, F., and K. D. Angielczyk. 2022. Inner ear biomechanics reveals a Late Triassic origin for mammalian endothermy. Nature 607:726-731.

Luo, Z.-X., Bhullar, B.-A.S., Crompton, A.W., Neander, A.I., and T.B. Rowe. 2022. Reexamination of the mandibular and dental morphology of the Early Jurassic mammaliaform Hadrocodium wui. Acta Palaeontologica Polonica. 67 (1): 95-113.

Pevsner, S. K., Grossnickle, G. M. and Z.-X. Luo. 2022. The functional diversity of marsupial limbs is influenced by both ecology and developmental constraint.  Biological Journal of the Linnean Society. 135(3): 569-585. (

Sulser, R. B., Patterson, B. D., Urban, D. J., Neander, A. I. and Z.-X. Luo. 2022. Evolution of Inner ear neuroanatomy of bats and implications for echolocation. Nature. 602: 449-454 ( (

Li, P., Ross, C. F., and Z.-X. Luo. 2022. Morphological disparity and evolutionary transformations in the primate hyoid apparatus. Journal of Human Evolution. 162 (2022): 103094 Published online 2021 (

Panciroli, E. L., Benson, R. B. J., Fernandez, V., Humage, M., Martin-Serra, A., Walsh, S., Luo, Z.-X. and N. C. Fraser, 2021b. Postcrania of Borealestes (Mammaliformes: Docodonta) and the emergence of ecomorphological diversity in early Mammals. Palaeontology. 2021: e12577 (pp. 1–36) (

Panciroli, E. L., Benson, R. B. J., Fernandez, V., Butler, R. J., Fraser, N. C., Luo, Z.-X. and S. Walsh. 2021a. New species of mammaliaform and the cranium of Borealestes (Mammaliformes: Docodonta) from the Middle Jurassic of the British Isles. Zoological Journal of the Linnean Society. 192(4): 1323–1362 (

Grossnickle, D. M., Chen, M., Wauer, J. G. A., Pevsner, S. K., Weaver, L. N., Meng, Q.-J., Liu, D., Zhang, Y.-G., and Z.-X. Luo. 2020.  Incomplete convergence of gliding-mammal skeletons.  Evolution. 74: 2662-2680. (

Luo, Z.-X. and G. A. Manley. 2020. Origins and Evolution of Mammalian Ears and Hearing Function. Pp. 207-252 (Chapter 2). In: The Senses – A Comprehensive Reference (Volume 2) (Second Edition) (Fritzsch, B. Editor; Grothe, B. volume editor); Elsevier Academic Press. ( (pdf)

Panciroli, E. L., Benson, R. B. J., and Z.-X. Luo. 2019. The mandible and dentition of Borealestes serendipitus (Docodonta) from the Middle Jurassic of Skye, Scotland.  Journal of Vertebrate Paleontology. 39: e1621884 (17 pages) (

Jäger, K. R. K., Luo, Z.X. and T Martin. 2019. Postcranial skeleton of Henkelotherium guimarotae (Cladotheria, Mammalia) and locomotor adaptation. Journal of Mammalian Evolution, 3: 349-372 (

Zhou, C.-F. Bhullar, B.-A. S., Neander, A. I., Martin, T. and Z.-X, Luo. 2019. New Jurassic mammaliaform sheds light on early evolution of mammal-like hyoid bones. Science. 365: 276-279. (

Panciroli, E. L., Schultz, J. A., and Z.-X. Luo. 2019. Morphology of the petrosal and stapes of Borealestes (Mammaliaformes, Docodonta) from the Middle Jurassic of Skye, Scotland. Papers in Palaeontology. 5:139-156 (

Lautenschlager, S., Gill, P., Luo, Z.-X., Fagan, M. J., and E. J. Rayfield. 2018. The role of miniaturisation in the evolution of the mammalian jaw. Nature. 561:533-537 (

Huttenlocker, A. K., Grossnickle, D. M., Kirkland, J. I., Schultz, J. A., and Z.-X. Luo. 2018. Late-surviving stem mammal links the lowermost Cretaceous of North America and Gondwana. Nature. 558:109-112 (

Chen, M., Z.-X. Luo, and G. P. Wilson. 2017. The postcranial skeleton of Yanoconodon allini from the Early Cretaceous of Hebei, China and its implications for locomotor adaptation in eutriconodontan mammals. Journal of Vertebrate Paleontology. 37(3): e1315425 (23 pages) (

Schultz, J. A, Bhullar, B. A. S., and Z.-X. Luo. 2017 (2019). Re-examination of the Jurassic mammaliaform Docodon victor by computed tomography and occlusal functional analysis. Journal of Mammalian Evolution. 26: 9-38. (published online 2017; in print 2019) (

Meng, Q.-J., Grossnickle, D. M., Liu, D., Zhang, Y.-G., Neander, A. I., Ji, Q., and Z.-X. Luo. 2017.  New gliding mammaliaforms from the Jurassic.  Nature. 548: 291-296. (

Luo, Z.-X.,  Meng, Q.-J., Grossnickle, D. M., Liu, D., Zhang, Y.-G., Neander, A. I., and Ji, Q. 2017.  New evidence for mammaliaform ear evolution and feeding adaptation in a Jurassic ecosystem.  Nature. 548: 326-329 (

Anthwal, N.  Urban, D. J., Luo, Z.-X., Sears, K. E., and Abigail S. Tucker. 2017. Meckel’s cartilage breakdown offers clues to mammalian middle ear evolution.  Nature Ecology & Evolution. 1:0093. (

Urban, D. J., Anthwal, N., Luo, Z.-X., Maier, J. A., Sadier, A., Tucker, A. S., and K. E. Sears.. 2017. A new developmental mechanism for the separation of the mammalian middle ear ossicles from the jaw.  Proceedings of Royal Society (London) B – Biological Sciences. 284: 20162416. (Published online in 2016) (

Schultz, J. A., U. Zeller and Luo, Z.-X. 2017.  Inner ear labyrinth anatomy of monotremes and  implications for mammalian inner ear evolution. Journal of Morphology. 278:236–263. (Published online in 2016) (

Luo, Z.-X., Schultz, J. A. and E. G. Ekdale. 2016. Evolution of the middle and inner ears of mammaliaforms: the approach to mammals.  In J. A. Clack, R. R. Fay and A. N. Popper (editors) Evolution of the Vertebrate Ear: Evidence from the Fossil Record.  Springer Handbooks for Auditory Research. 59:139-174. (

Lautenschlager, S., Gill, P., Luo, Z.-X., Fagan, M. J. and E. J. Rayfield. 2017.  Morphological evolution of the mammalian jaw adductor complex.  Biological Reviews. 92: 1910–1940 (

Rowe, T. B., Luo, Z.-X., Ketcham, R. A., Maisano, J. A. and M. W. Colbert. 2016. Data Descriptor: X-ray computed tomography datasets for forensic analysis of vertebrate fossils. Scientific Data. 3:160040 (

Girard, R., Zeineddine, H. A., Orsbon, C., Tan, H., Moore, T., Hobson, N., Shenkar, R., Lightle, R., Shi, C., Fam, M. D., Cao, Y., Shen, L., Neander, A. I., Rorrer, A., Gallione, C., Tang, A. T., Kahn, M. L., Marchuk, D. A., Luo, Z.-X., Awad, I. A. 2016.  Micro-computed tomography in murine models of cerebral cavernous malformations as a paradigm for brain disease. Journal of Neuroscience Methods. 271:14–24. (

Luo, Z.-X. Meng, Q.-J., Liu, D., Y-G. Zhang, and Yuan, C.-X. 2016. Cruro-pedal structure of the paul-choffatiid multituberculate Rugosodon eurasiaticus and evolution of the multituberculate ankle. Palaeontologia Polonica 67: 149-169. (download pdf) (doi: 10.4202/pp.2016.67_149)

Luo, Z.X., Gatesy, S.M., Jenkins, F.A., Amaral, W.W. and Shubin, N.H. 2015. Mandibular and dental characteristics of Late Triassic mammaliaform Haramiyavia and their ramifications for basal mammal evolution. Proceedings of the National Academy of Sciences USA 112(51): E7101-E7109. (

Martin, T., Marugán-Lobón, J., Vullo, R., Martín-Abad, H., Luo, Z.-X., and A. D. Buscalioni. 2015. A Cretaceous eutriconodont and integument evolution of early mammals. Nature. 526(7573): 380-384. (

Luo, Z.-X. 2015. Origin of the Mammalian Shoulder. Pp.167-187. In: K. P. Dial, N. H. Shubin & E. L. Brainerd (eds.) Great Transformations: Major Events in the History of Vertebrate Life. The University of Chicago Press. (pdf)

Luo, Z.-X., Q.-J. Meng, Q. Ji, D. Liu, Y.-G. Zhang, and A. I. Neander. 2015. Evolutionary development in basal mammaliaforms as revealed by a docodontan. Science347: 760-764 (

Meng, Q.-J., Q. Ji, Y.-G. Zhang, D., Liu, D. M. Grossnickle, and Z.-X. Luo. 2015. An arboreal docodont from the Jurassic and mammaliaform ecological diversification. Science347: 764-768 (

Hughes, E. M., J. R. Wible, M. Spaulding, and Z.-X. Luo. 2015. Mammalian petrosal from the Upper Jurassic Morrison Formation of Fruita, Colorado. Annals of Carnegie Museum. 83: 1-17 (

Yuan, C.-X., Ji, Q., Meng, Q.-J. and Z.-X. Luo. 2013. Earliest evolution of multituberculate mammals revealed by a new Jurassic fossil. Science. 341: 779-783. (

Zhou, C.-F., Wu, S., Martin, T. and Z.-X. Luo. 2013. A Jurassic mammaliaform and the earliest mammalian evolutionary adaptations. Nature. 500: 163-168 (

Ruf, I., Z.-X. Luo, and T. Martin. 2013. Re-investigation of the basicranium of Haldanodon exspectatus (Docodonta, Mammaliaformes). Journal of Vertebrate Paleontology. 33:382-400. (

O’Leary, M. A., Bloch, J. I., Flynn, J. J., Gaudin, T. J., Giallombardo, A., Giannini, N. P., Goldberg, S. L., Kraatz, B. P., Luo Z.-X., Meng, J., Ni, X., Novacek, M. J., Perini, F. A., Randall, Z. S., Rougier, G. W., Sargis, E. J., Silcox, M. T., Simmons, N. B., Spaulding, M., Velazco, P. M., Weksler, M., Wible, J. R., and A. L. Cirranello. 2013. The placental mammal ancestor and the post-K-Pg radiation of placentals. Science 339: 662-667. (

Luo, Z.-X., I. Ruf and T. Martin. 2012. The petrosal and inner ear of the Late Jurassic cladotherian mammal Dryolestes leiriensis and implications for evolution of ear in therian mammals. Zoological Journal of Linnean Society (London) 166:433-463.(

Chen, M. and Z.-X. Luo. 2012 (2013). Postcranial skeleton of the Cretaceous mammal Akidolestes cifellii and its locomotor adaptations. Journal of Mammalian Evolution. 20: 159-189 (published online 2012; in print 2023) (

Older Publications

Luo, Z.-X. 2011. Developmental patterns in Mesozoic evolution of mammal ears. Annual Review of Ecology, Evolution and Systematics 42:355-80 (

Luo, Z.-X., Yuan, C.-X., Q.-J. Meng, and Q. Ji. 2011. A Jurassic eutherian mammal and the divergence of marsupials and placentals. Nature 476: 442-445 (

Rowe, T. B., Macrini, T. E., and Z.-X. Luo. 2011. Fossil evidence on origin of the mammalian brain. Science 332: 955-957 (

Luo, Z.-X., Ruf, I., J. A. Schultz, and Thomas Martin. 2011. Fossil evidence on evolution of inner ear cochlea in Jurassic mammals. Proceedings of Royal Society Series-B (Biological Sciences) 278: 28-34 (

Luo, Z.-X. 2010. Evolutionary Development and Homoplasy of the Middle Ear in Early Mammal Evolution. Pp. 163-179. In: Darwin’s Heritage Today (Manyuan Long, Zhonghe Zhou and Hongya Gu, editors). China Higher Education Press. Beijing, China. (link to pdf)

Ji, Q., Luo, Z.-X., Zhang, Z.-L., Yuan, C.-X. and L. Xu. 2009. Evolutionary development of the middle ear in Mesozoic therian mammals. Science 326:278-231 (

Gao, C.-L., Wilson, G. P., Luo, Z.-X., Maga, A. M., Meng, Q.-J., and Wang, X.-R. 2009. A new mammal skull from the Lower Cretaceous of China with implications for the evolution of obtuse angled molars and amphilestid eutriconodonts. Proceedings of Royal Society (London) Series B (Biological Sciences). 277:237-246 (

Ruf, I., Z.-X. Luo, J. R. Wible, and T. Martin. 2009. Petrosal Anatomy and Inner Ear Structure of the Late Jurassic Mammal Henkelotherium And the Ear Region Characters of Basal Therian Mammals. Journal of Anatomy. 214: 679-693. (

Chen, M. and Z.-X. Luo. 2008. Morphology of dentition and postcranial skeleton of AkidolestesActa Geologica Sinica (Chinese Edition) 82(2): 155-164 (with English Abstract).

You, H.-L. and Z.-X. Luo. 2008. Dinosaurs from the Lower Cretaceous Gonpoquan Basin in Jiuquan Area, Gansu Province, China. Acta Geologica Sinica (Chinese Edition) 82(1): 139-144. (with English Abstract).

Luo, Z.-X. 2007. Transformation and diversification in early mammal evolution. Nature 450: 1011-1019.(

Luo, Z.-X., Q. Ji and C.-X. Yuan. 2007. Convergent dental adaptations in pseudo-tribosphenic and tribosphenic mammals. Nature 450: 93-97 (

Luo, Z.-X., P.-J. Chen, G. Li and M. Chen. 2007. A new eutriconodont mammal and evolutionary development of early mammals. Nature 446: 288-293 (

Luo, Z.-X., and T. Martin. 2007. Analysis of molar structure and phylogeny of docodontan genera. Bulletin of Carnegie Museum of Natural History 39: 27-47.([27:AOMSAP]2.0.CO;2)

Luo, Z.-X. 2007. Successive diversifications in early mammalian evolution. Pp. 337-391. In: Major Transitions In Vertebrate Evolution (J. S. Anderson and H.-D. Sues, eds.). Indiana University Press, Bloomington and Indianapolis, Indiana.

Ji, Q., Z.-X. Luo, C.-X. Yuan, and A. R. Tabrum. 2006. A swimming mammaliaform from the Middle Jurassic and ecomorphological diversification of early mammals. Science 311: 1123-1127 (

Li, G., and Z.-X. Luo. 2006. A Cretaceous symmetrodont therian with some monotreme-like postcranial features. Nature 439:195-200 (

Hurum, J. H., Z.-X. Luo, and Z. Kielan-Jaworowska. 2006. Were mammals originally venomous? Acta Palaeontologica Polonica 51(1): 1-11. (Acta Palaeontologica Polonica PDF download)

Luo, Z.-X., and J. R. Wible. 2005. A Late Jurassic digging mammal and early mammalian diversification. Science 308:103-107 (

Kielan-Jaworowska, Zofia, Richard L. Cifelli, & Zhe-Xi Luo. 2004. Mammals from the Age of Dinosaurs – Origins, Evolution, and Structure. Columbia University Press, New York. Pp. i-xv; 1-630. (More information is available on the website for Columbia University Press)

Luo, Z.-X., Q. Ji, J. R. Wible and C-X. Yuan. 2003. An Early Cretaceous Tribosphenic mammal and metatherian evolution. Science 302:1934-1940 (DOI: 10.1126/science.1090718)

Luo, Z.-X., Z. Kielan-Jaworowska, R. L. Cifelli. 2002. In Quest for A Phylogeny of Mesozoic Mammals. Acta Palaeontologica Polonica. 47(1): 1-78. (PDF download from Acta Palaeontologica Polonica)

Ji, Q., Z.-X. Luo, C-X. Yuan, J. R. Wible, J.-P. Zhang, & J. A. Georgi. 2002. The earliest known eutherian mammal. Nature 416: 816-822 (

Luo, Z.-X. 2001. Inner ear and its bony housing in tritylodonts and implications for evolution of mammalian ear. Pp. 81-97. In: F. A. Jenkins et al. (eds.), Studies in Organismic and Evolutionary Biology in Honor of A. W. Crompton. Bulletin of Museum of Comparative Zoology, Harvard University. 156(1): 81-97.

Luo, Z.-X., A. W. Crompton and A-L. Sun. 2001. A new mammaliaform from the Early Jurassic of China and evolution of mammalian characteristics. Science 292: 1535-1540 (

Luo, Z.-X., R. C. Cifelli and Z. Kielan-Jaworowska. 2001. Dual evolution of tribosphenic mammals. Nature 409: 53-57 (

Tang, F., Z.-X. Luo, Z.-H. Zhou, H.-L. You, J. A. Georgi, Z.-L. Tang, and X.-Z. Wang. 2001. Biostratigraphy and Paleoenvironment of the Dinosaur-bearing Sediments in Lower Cretaceous of Mazongshan Area, Gansu Province, China. Cretaceous Research 22: 115-129 (

Luo, Z.-X. 2000. Evolution: in search of the whales’ sisters. Nature 404: 235 – 239 (

Luo, Z.-X. and P. D. Gingerich. 1999. Transition from terrestrial ungulates to aquatic whales: transformation of the basicranium and evolution of hearing. Papers on Paleontology (Monograph) 31: 1-98. Museum of Paleontology, University of Michigan, Ann Arbor, Michigan.

Ji, Q., Z.-X. Luo, and S. Ji. 1999. A Chinese triconodont mammal and mosaic evolution of mammalian skeleton. Nature 398:326-330 (

Geisler, J. H., and Z.-X. Luo. 1998. Cranial vascular evolution in mesonychids and cetaceans and the phylogenetic position of cete. Pp. 161-212. In: J. G. M. Thewissen (ed.), Evolutionary Emergence of Whales. Plenum Press, New York.

Hu, Y.-M., Wang, Y.-Q., Z.–X. Luo, and C.-K. Li.  1997.  A new symmetrodont mammal from China and its implications for mammalian evolution. Nature 390: 137-142.(

Geisler, J. H., and Z.–X. Luo1996.  Petrosal and inner ear structures of Herpetocetus, their implications on relationships and hearing function of archaic mysticetes. Journal of Paleontology 70: 1045-1066. (

Luo, Z.-X., and K. Marsh. 1996. Petrosal (periotic) and inner ear of a Pliocene kogiine whale (Kogiinae, Odontoceti): Implications on relationships and hearing evolution of toothed whales. Journal of Vertebrate Paleontology 16:328-348.(

Luo, Z.-X., and E. R. Eastman.  1995.  The petrosal and inner ear structures of a squalodontoid whale: implications for evolution of hearing in odontocetes.  Journal of Vertebrate Paleontology 15: 431-442 (

Luo, Z.-X., A. W. Crompton, and S. G. Lucas.  1995.  Evolutionary origins of the mammalian promontorium and cochlea. Journal of Vertebrate Paleontology 15: 113-121 (

Luo, Z.-X., and A.W. Crompton. 1994. Transformations of the quadrate (incus) through the transition from non-mammalian cynodonts to mammals. Journal of Vertebrate Paleontology 14:341-374 (

Luo, Z.-X. 1994. Sister taxon relationships of mammals and the transformations of the diagnostic mammalian characters. Pp. 98-128 in: In the Shadow of Dinosaurs-Early Mesozoic Tetrapods c dc,dcccv (N. C. Fraser & H.-D. Sues, eds.). Cambridge University Press.

Crompton, A. W., and Z.-X. Luo.  1993.   The relationships of the Liassic mammals  Sinoconodon, Morganucodon oehleri and Dinnetherium. Pp. 30-44,  in: Mammal Phylogeny (F. S. Szalay, M. J. Novacek, and M. C. McKenna , eds.). Springer-Verlag, New York.

Lucas, S. G., and Z.-X. Luo1993Adelobasileus from the Upper Triassic of West Texas: the oldest mammal.  Journal of Vertebrate Paleontology 13: 309-334 (

Luo, Z.-X., and  A.-L. Sun.  1993.  Oligokyphus (Cynodontia: Tritylodontidae) from the Lower Lufeng Formation (Lower Jurassic) of Yunnan, China. Journal of Vertebrate Paleontology 13: 477-482 (DOI:10.1080/02724634.1994.10011526)

Luo, Z.-X., and D. R. Ketten.  1991.   CT scanning and computerized reconstructions of the inner ear structure of multituberculate mammals. Journal of Vertebrate Paleontology, 11: 220-228 (

Luo, Z.-X1991.  The variability of dental morphology and the relationships of the earliest arctocyonid species. Journal of Vertebrate Paleontology 11: 452-471 (

McKenna, M. C., M.-C. Chow, S.-Y. Ting, and  Z.-X. Luo.  1989. Radinskya yupingae, a perissodactyl-like mammal from the late Paleocene of Southern China. Pp. 24-37, in The Evolution of Perissodactyls, (D. R. Prothero, and R. M. Schoch, eds.).  Oxford University Press, Oxford.

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