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Smithsonian National Museum of Natural History

Department of Paleobiology

  • Matthew T. Carrano
  • Curator of Dinosauria
  • Phone:   202-633-1314
  • Fax:   202-786-2832
  • E-mail Address:   carranom
  • Mailing Address:
    Smithsonian Institution
    PO Box 37012, MRC 121
    Washington, DC 20013-7012
  • Shipping Address:
    Smithsonian Institution
    National Museum of Natural History
    10th & Constitution NW
    Washington, DC 20560-0121


Ph.D. University of Chicago, 1998
M.S. University of Chicago, 1995
B.S. Brown University, 1991

Research Interests

I. Dinosaur Evolution and the Fossil Record
I am particularly interested in large-scale evolutionary patterns within the Dinosauria. This diverse, long-lived vertebrate group (even excluding avian members) dominated terrestrial habitats worldwide from the Late Triassic through to the end of the Cretaceous. Thus they provide an interesting and instructive parallel for the Cenozoic radiation of mammals, permitting exploration of such evolutionary patterns as Cope's Rule (increasing body size), the cursorial-graviportal locomotor gradient, and biogeographic scenarios associated with continental fragmentation. My work focuses on using dinosaur phylogeny to track these evolutionary changes, both throughout the group and within specific clades (e.g., sauropods). Numerous similarities between mammalian and dinosaurian patterns are evident, particularly with regard to size and locomotion, but exceptions highlight potentially important biological differences between these two major clades. I also integrate information from the vast dinosaur ichnological record into these studies.

Associated with this work, I participate in the Vertebrate Paleontology Working Group, which is assembling the vertebrate component of the Paleobiology Database (PBDB). The PBDB will ultimately provide full internet access to all published vertebrate fossil localities for any researcher worldwide. Because the database also includes records of marine invertebrates, taphonomy, and fossil plants, it holds great potential for the exploration of innumerable macroevolutionary patterns. To date, more than 4000 dinosaur-bearing localities have been entered into the PBDB, which can be analyzed and downloaded by any visitor or researcher. This dataset has allowed me to study and quantify many aspects of the dinosaur fossil record in detail, including the group’s diversity and geographic history, as well as the effects of sampling biases.

Collaborators: John Alroy and the Vertebrate Paleontology Working Group (National Center for Ecological Analysis and Synthesis, Santa Barbara, CA); Jeffrey Wilson (University of Michigan, Ann Arbor)

II. Dinosaur Phylogeny and Exploration
Currently I am completing a major reevaluation of the phylogeny of basal (non-coelurosaur) theropods, with an emphasis on the relationships of ceratosaurs. This work is based on first-hand observations of nearly all basal theropod specimens worldwide, and provides an opportunity to employ a variety of additional data (including stratigraphic) in testing the robustness of the theropod tree. In addition, the diversity and temporal longevity of ceratosaurs affords them relevance for understanding biogeographic and other trends in dinosaur evolution.

My phylogenetic work includes a substantial component dedicated to understanding new theropod taxa from Madagascar (e.g., Majungasaurus and Masiakasaurus) and their relatives (abelisaurids and noasaurids, respectively). I have participated in two Stony Brook University/Université d'Antananarive expeditions to the Late Cretaceous Maevarano Formation in the Mahajunga Basin of northwestern Madagascar, led by Dr. David Krause. In addition, in 2004 I led a Smithsonian expedition to look for underlying Early Cretaceous terrestrial deposits. These formations hold the potential to greatly clarify our understanding of Gondwanan dinosaur evolution, because they represent a currently unsampled interval in the Malagasy record.

Collaborators: Scott Sampson and Mark Loewen (University of Utah, Salt Lake City), Oliver Rauhut (Bayerische Staatssammlung für Paläontologie und Geologie, Munich)

III. Functional Morphology and Locomotion
I have a longstanding interest in the functional morphology and biomechanics of vertebrate locomotion. In the past, I used experimental data gathered from extant birds and mammals to draw conclusions about locomotor habits in extinct vertebrates, particularly dinosaurs. This included a reappraisal of the role of torsion in vertebrate limbs, as well as a new and more explicit evaluation of the relationship between limb morphology and locomotor habit. In summary, data from extant taxa suggest that most dinosaurs were broadly similar in general locomotor habit, and that the unique hindlimb locomotor system seen in modern birds probably evolved after the origin of flight, in conjunction with a major reduction in body size. Similarities in macroevolutionary patterns of locomotor morphology between dinosaurs and mammals may shed light on the underlying selective pressures driving these trends.

Recently, I have collaborated on more disparate functional projects. These included finite-element analyses of the mechanics and morphology of dinosaur phalanges, and phylogenetic and morphometric studies of locomotor evolution in plesiosaurs.

Collaborators: Karen Moreno (University of New South Wales, Sydney), Robin O'Keefe (Marshall University, Huntington, WV)

Current and Former Research Assistants, Post-doctoral Fellows and Students


  • Robert V. Hill, Ph.D. (2004), Stony Brook University, external committee member
  • Keith Metzger, Ph.D. (2005), Stony Brook University, external committee member
  • Margaret I. Hall, Ph.D. (2005), Stony Brook University, external committee member
  • Karen Moreno, Ph.D. (2005), University of Bristol, co-advisor
  • Mark A. Loewen Ph.D. (expected 2007), University of Utah, external committee member

Undergraduate Interns:

  • Jorge Velez-Juarbe, B.S. (expected 2007), University of Puerto Rico
  • Kaitlin Maguire, B.A. (2005), The George Washington University
  • Jessica Walden, B.S. (expected 2009), The George Washington University/Cornell University
  • Matthew Oreska, B.A. (expected 2007), The College of William & Mary


Positions Held :

Recent Grants

  • 2006: Smithsonian Collections Care and Preservation Fund Award: "Conservation of the Fossil Reptile and Amphibian Collections"
  • 2003-2004: Smithsonian Walcott Fund: "Evolutionary Patterns in Late Jurassic Dinosaurs of North America"
  • 2002-2004: National Geographic Society: "Medial Cretaceous Vertebrate Faunas of Madagascar"
  • 2001-2003: Jurassic Foundation (with J. A. Wilson, co-PI): "From Sauropoden to Sauropods: Translating the Works of Werner Janensch"
  • 1999-2003: National Science Foundation Grant, Systematic Biology (with S. D. Sampson, co-PI): "Osteology and Phylogeny of Basal Theropod Dinosaurs"
  • 1999-2003: Jurassic Foundation: "The Role of Subdivided Muscles in the Archosaur Hind Limb"

Editorial Service

  • 2006-present: Content Review Board, Smithsonian Institution Scholarly Press
  • 2003-2006: Technical Editor, Journal of Paleontology
  • 2000-2003: Associate Editor, Journal of Vertebrate Paleontology
  • 2000: Guest Editor, "Dinosaurs" issue, Odyssey Magazine, Cobblestone Press

Field Work

  • 2003-present: Late Jurassic–Early Cretaceous interval of the Western Interior, North America [view the 2003 and 2004 field seasons]
  • 1999-2004: Cretaceous deposits of the Mahajanga Basin, northwestern Madagascar [view the 2004 field season]


Evans, David C., Barrett, Paul M., Brink, Kirstin S. and Carrano, Matthew T. 2015. Osteology and bone microstructure of new, small theropod dinosaur material from the early Late Cretaceous of Morocco, Gondwana Research, 27(3):1034-1041

Ksepka, Daniel T., Parham, James F., Allman, James F., Benton, Michael J., Carrano, Matthew T., Cranston, Karen A., Donoghue, Philip C. J., Head, Jason J., Hermsen, Elizabeth J., Irmis, Randall B., Joyce, Walter G., Kohli, Manpreet, Lamm, Kristin S., Leehr, Dan, Patané, José S. L., Polly, P. David, Phillips, Matthew J., Smith, N. Adam, Smith, Nathan D., van Tuinen, Marcel, Ware, Jessica L. and Warnock, Rachel C. M. 2015. The Fossil Calibration Database, A New Resource for Divergence Dating, Systematic Biology, 64(5):853-859

Aiello, B. R., Iriarte-Diaz, J., Blob, R. W., Butcher, M. T., Carrano, Matthew T., Espinoza, N. R., Main, R. P. and Ross, C. F. 2015. Bone strain magnitude is correlated with bone strain rate in tetrapods: implications for models of mechanotransduction, Proceedings of the Royal Society of London B: Biological Sciences, 282(1810)

Mannion, Philip D., Benson, Roger B. J., Carrano, Matthew T., Tennant, Jonathan P., Judd, Jack and Butler, Richard J. 2015. Climate constrains the evolutionary history and biodiversity of crocodylians, Nature Communications, 6

Carrano, Matthew T. and D'Emic, Michael D. 2015. Osteoderms of the titanosaur sauropod dinosaur Alamosaurus sanjuanensis Gilmore, 1922, Journal of Vertebrate Paleontology, 35(1)

Benson, Roger B. J., Campione, Nicolás E., Carrano, Matthew T., Mannion, Philip D., Sullivan, Corwin, Upchurch, Paul and Evans, David C. 2014. Rates of Dinosaur Body Mass Evolution Indicate 170 Million Years of Sustained Ecological Innovation on the Avian Stem Lineage, PLoS Biology, 12(5):1-11

Brusatte, Stephen L., Butler, Richard J., Barrett, Paul M., Carrano, Matthew T., Evans, David C., Lloyd, Graeme T., Mannion, Philip D., Norell, Mark A., Peppe, Daniel J., Upchurch, Paul and Williamson, Thomas E. 2014. The extinction of the dinosaurs, Biological Reviews, 90(2):628-642

Conrad, Jack L., Head, Jason J. and Carrano, Matthew T. 2014. Unusual Soft-Tissue Preservation of a Crocodile Lizard (Squamata, Shinisauria) From the Green River Formation (Eocene) and Shinisaur Relationships, The Anatomical Record, 297(3):545-559

Campione, Nicolas E., Evans, David C., Brown, Caleb M. and Carrano, Matthew T. 2014. Body mass estimation in non-avian bipeds using a theoretical conversion to quadruped stylopodial proportions, Methods in Ecology and Evolution, 5(9):913-923

Thomas, Daniel B., McGraw, Kevin J., Butler, Michael W., Carrano, Matthew T., Madden, Odile and James, Helen F. 2014. Ancient origins and multiple appearances of carotenoid-pigmented feathers in birds, Proceedings of the Royal Society B: Biological Sciences, 281(1788)

Uhen, Mark D., Barnosky, Anthony D., Bills, Brian, Blois, Jessica, Carrano, Matthew T., Carrasco, Marc A., Erickson, Gregory M., Eronen, Jussi T., Fortelius, Mikael, Graham, Russell W., Grimm, Eric C., O'Leary, Maureen A., Mast, Austin, Piel, William H., Polly, P. David and Saila, Laura K. 2013. From Card Catalogs to Computers: Databases in Vertebrate Paleontology, Journal of Vertebrate Paleontology, 33(1):13-28

Oreska, Matthew P. J., Carrano, Matthew T. and Dzikiewicz, Katherine M. 2013. Vertebrate Paleontology of the Cloverly Formation (Lower Cretaceous), I: Faunal Composition, Biogeographic Relationships, and Sampling, Journal of Vertebrate Paleontology, 33(2):264-292

Thomas, Daniel B., James, Helen F., Carrano, Matthew T. and Madden, Odile. 2012. Searching for Evidence of Fossil Feather Color with Spectroscopy, Journal of Vertebrate Paleontology, 32:183-184

Fitzgerald, Erich M. G., Carrano, Matthew T., Holland, Timothy, Wagstaff, Barbara E., Pickering, David, Rich, Thomas H. and Vickers-Rich, Patricia. 2012. First ceratosaurian dinosaur from Australia, Naturwissenschaften, 99(5):397-405

Carrano, Matthew T., Benson, Roger B. J. and Sampson, Scott D. 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda), Journal of Systematic Palaeontology, 10(2):211-300

Benson, Roger B. J., Butler, Richard J., Carrano, Matthew T. and O'Connor, Patrick M. 2012. Air-filled postcranial bones in theropod dinosaurs: physiological implications and the 'reptile'–bird transition, Biological Reviews, 87(1):168-193

Burch, Sara H. and Carrano, Matthew T. 2012. An Articulated Pectoral Girdle and Forelimb of the Abelisaurid Theropod Majungasaurus Crenatissimus from the Late Cretaceous of Madagascar, Journal of Vertebrate Paleontology, 32(1):1-16

Carrano, Matthew T., Loewen, Mark A. and Sertich, Joseph J. W. 2011. New materials of Masiakasaurus knopfleri Sampson, Carrano, and Forster, 2001, and implications for the morphology of the Noasauridae (Theropoda:Ceratosauria), Smithsonian Contributions to Paleobiology, (95):1-53

Hall, Margaret I., Kirk, E. C., Kamilar, Jason M. and Carrano, Matthew T. 2011. Comment on "Nocturnality in Dinosaurs Inferred from Scleral Ring and Orbit Morphology", Science, 334(6063):1641-1641

Upchurch, P., Mannion, P. D., Benson, R. B. J., Butler, R. J. and Carrano, Matthew T. 2011. Geological and anthropogenic controls on the sampling of the terrestrial fossil record: a case study from the Dinosauria, Geological Society, London, Special Publications, 358(1):209-240

Butler, Richard J., Benson, Roger B. J., Carrano, Matthew T., Mannion, Philip D. and Upchurch, Paul. 2011. Sea level, dinosaur diversity and sampling biases: investigating the 'common cause' hypothesis in the terrestrial realm, Proceedings of the Royal Society (London) B, 278(1709):1165-1170

Wilson, Jeffrey A., Barrett, Paul M. and Carrano, Matthew T. 2011. An associated partial skeleton of Jainosaurus cf. septentrionalis (Dinosauria: Sauropoda) from the Late Cretaceous of Chhota Simla, Central India, Palaeontology, 54(5):981-998

Ksepka, Daniel T., Benton, Michael J., Carrano, Matthew T., Gandolfo, Maria A., Head, Jason J., Hermsen, Elizabeth J., Joyce, Walter G., Lamm, Kristin S., Patané, José S. L., Phillips, Matthew J., Polly, P. David, Van Tuinen, Marcel, Ware, Jessica L., Warnock, Rachel C. M. and Parham, James F. 2011. Synthesizing and databasing fossil calibrations: divergence dating and beyond, Biology Letters, 7(6):801-803

Mannion, Philip D., Benson, Roger B. J., Upchurch, Paul, Butler, Richard J., Carrano, Matthew T. and Barrett, Paul M. 2011. A temperate palaeodiversity peak in Mesozoic dinosaurs and evidence for Late Cretaceous geographical partitioning, Global Ecology and Biogeography, 21(9):898-908

Mannion, Philip D., Upchurch, Paul, Carrano, Matthew T. and Barrett, Paul M. 2011. Testing the effect of the rock record on diversity: a multidisciplinary approach to elucidating the generic richness of sauropodomorph dinosaurs through time, Biological Reviews, 86(1):157-181

Benson, Roger B. J., Carrano, Matthew T. and Brusatte, Stephen L. 2010. A new clade of archaic large-bodied predatory dinosaurs (Theropoda: Allosauroidea) that survived to the latest Mesozoic, Naturwissenschaften, 97(1):71-78

Carrano, Matthew T., Wilson, Jeffrey A. and Barrett, Paul M. 2010. "The history of dinosaur collecting in central India, 1828-1947". Pp. 161-173 in Dinosaurs and other Extinct Saurians: a Historical Perspective (Moody, R. T. J., Buffetaut, E., Naish, D. and Martill, D. M.). Geological Society

Hunt, Gene and Carrano, Matthew T. 2010. "Models and methods for analyzing phenotypic evolution in lineages and clades". Pp. 245-269 in Quantitative Methods in Paleobiology (Alroy, John and Hunt, Gene). The Paleontological Society

Carrano, Matthew T., Krause, D. W., O'Connor, P. M. and Sampson, S. D. 2009. Case 3487: Megalosaurus crenatissimus Depéret, 1896 (currently Majungasaurus crenatissimus; Dinosauria, Theropoda): proposed replacement of the holotype by a neotype, Bulletin of Zoological Nomenclature, 66(3):261-264

Carrano, Matthew T. and Sampson, S. D. 2008. The phylogeny of Ceratosauria (Dinosauria : Theropoda), Journal of Systematic Palaeontology, 6(2):183-236

Moreno, K., Carrano, Matthew T. and Snyder, R. 2007. Morphological changes in pedal phalanges through ornithopod dinosaur evolution: a biomechanical approach, Journal of Morphology, 268(1):50-63

Carrano, Matthew T. 2007. "Walking and running dinosaurs". Pp. 308 in Dinosaurs (Holtz, Jr, Thomas Richard). Random House Children's Books

Carrano, Matthew T. 2007. The appendicular skeleton of Majungasaurus crenatissimus (Theropoda: Abelisauridae) from the Late Cretaceous of Madagascar, Society of Vertebrate Paleontology Memoir 8.Journal of Vertebrate Paleontology, 27(2):163-179

Krause, D. W., Sampson, S. D., Carrano, Matthew T. and O'Conner, Patrick M. 2007. Overview of the history of discovery, taxonomy, phylogeny, and biogeography of Majungasaurus crenatissimus (Theropoda: Abelisauridae) from the Late Cretaceous of Madagascar, Society of Vertebrate Paleontology Memoir 8.Journal of Vertebrate Paleontology, 27(2):1-20

Gaudin, T. J. and Carrano, Matthew T. 2006. "James Allen Hopson: a bibliography (1964-2003)". Pp. 507-515 in Amniote Paleobiology: Perspectives on the Evolution of Mammals, Birds, and Reptiles (Carrano, Matthew T., Blob, R. W., Gaudin, T. and Wible, J.). University of Chicago Press

Carrano, Matthew T. and Velez-Juarbe, Jorge. 2006. Paleoecology of the Quarry 9 vertebrate assemblage from Como Bluff Wyoming (Morrison Formation, Late Jurassic), Palaeogeography, palaeoclimatology, palaeoecology, 237(2-4):147-159

Carrano, Matthew T., Gaudin, T. J., Blob, R. W. and Wible, J. R.2006. Amniote Paleobiology: Perspectives on the Evolution of Mammals, Birds, and Reptiles. University of Chicago Press

Gaudin, T. J., Carrano, Matthew T., Blob, R. W. and Wible, J. R. 2006. "Introduction". Pp. 1-17 in Amniote Paleobiology: Perspectives on the Evolution of Mammals, Birds, and Reptiles (Carrano, Matthew T., Blob, R. W., Gaudin, T. and Wible, J.). University of Chicago Press

Carrano, Matthew T. 2006. "Body-size evolution in the Dinosauria". Pp. 225-268 in Amniote Paleobiology: Perspectives on the Evolution of Mammals, Birds, and Reptiles (Carrano, Matthew T., Gaudin, T. J., Blob, R. W. and Wible, J. R.). University of Chicago Press

Crompton, A. W., Jenkins, F. A., Hopson, S., Gaudin, T. J. and Carrano, Matthew T. 2006. "James Allen Hopson: a biography". Pp. 507-515 in Amniote Paleobiology: Perspectives on the Evolution of Mammals, Birds, and Reptiles (Carrano, Matthew T., Blob, R. W., Gaudin, T. and Wible, J.). University of Chicago Press

Krause, D. W., O'Connor, P. M., Rasoamiaramanana, A. H., Buckley, G. A., Burney, D., Carrano, Matthew T., Chatrath, P. S., Flynn, J. J., Forster, C. A., Godfrey, L. R., Jungers, W. L., Rogers, R. R., Samonds, K. E., Simons, E. L. and Wyss, A. R. 2006. Preserving Madagascar's natural heritage: the importance of keeping the island's vertebrate fossils in the public domain, Madagascar Conservation and Development, 1(1):43-47

Carrano, Matthew T. 2005. "The evolution of sauropod locomotion: morphological diversity of a secondarily quadrupedal radiation". Pp. 229-251 in The Sauropods: Evolution and Paleobiology (Curry Rogers, K. A. and Wilson, J. A.). University of California Press

Carrano, Matthew T., Hutchinson, J. R. and Sampson, S. D. 2005. New information on Segisaurus halli, a small theropod dinosaur from the Early Jurassic of Arizona, Journal of Vertebrate Paleontology, 25(4):835-849

Carrano, Matthew T. and O'Conner, Patrick M. 2005. Bird's Eye view, Natural History, 114(4):42-47

O'Keefe, F. R. and Carrano, Matthew T. 2005. Correlated trends in the evolution of the plesiosaur locomotor system, Paleobiology, 31(4):656-675

Carrano, Matthew T. 2004. Why are there no living animals as large as the dinosaurs?, Discover, 25(1):12

Carrano, Matthew T. and Sampson, S. D. 2004. A review of coelophysoids (Dinosauria: Theropoda) from the Lower Jurassic of Europe, with comments on the late history of the Coelophysoidea, Neues Jahrbuch fAr Geologie und PalAontologie Monatshefte, 2004(9):537-558

Carrano, Matthew T., Sampson, S. D. and Forster, C. A. 2002. The osteology of Masiakasaurus knopfleri, a small abelisauroid (Dinosauria: Theropoda) from the Late Cretaceous of Madagascar, Journal of Vertebrate Paleontology, 22(3):510-534