Lawrence M. Witmer, PhD
Professor of Anatomy
Chang Professor of Paleontology

Dept. of Biomedical Sciences
Heritage College of Osteopathic Medicine
Life Science Building, Rm 123
Ohio University
Athens, Ohio 45701 USA

Email: witmerL@ohio.edu

All publications listed in order.  For a breakdown based on type of publication (e.g., Research Articles, Books , Abstracts) see Curriculum Vitae.

1. Sues, H.-D., A. O. Averianov, R. C. Ridgely, and L. M. Witmer. 2015. Titanosauria (Dinosauria, Sauropoda) from the Upper Cretaceous Bissekty Formation of Uzbekistan. Journal of Vertebrate Paleontology 35(1):e889145-1–e889145-14.
   • Download supplemental figures of the braincase and endocast
   • Download a 3D PDF of the braincase, brain endocast, and labyrinth: small (5 MB), medium (16 MB), large (30 MB) Note: save 3D PDFs to your computer before running.
   • Download the original CT scan data in DICOM format (51 MB): WitmerLab site, figshare

2. Lautenschlager, S., L. M. Witmer, Perle A., L. E. Zanno, and E. J. Rayfield. 2014. Cranial anatomy of Erlikosaurus andrewsi (Dinosauria, Therizinosauria): new insights based on digital reconstruction. Journal of Vertebrate Paleontology 34(6):1263–1291.
   • Download a 3D PDF of the restored fossil skull of Erlikosaurus
   • Download a 3D PDF of the original fossil skull of Erlikosaurus

3. Bourke, J. M., Wm. R. Porter, R. C. Ridgely, T. R. Lyson, E. R. Schachner, P. R. Bell, and L. M. Witmer. 2014. Breathing life into dinosaurs: tackling challenges of soft-tissue restoration and nasal airflow in extinct species. Anatomical Record 297:2148–2186.
   • Visit the Project Page for a Common Language Summary, PDF download of the article, and other content
   • Check out the YouTube video

4. Eastman, J. T., L. M. Witmer, R. C. Ridgely, and K. L. Kuhn. 2014. Divergence in skeletal mass and bone morphology in Antarctic notothenioid fishes. Journal of Morphology 275:841–861. DOI 10.1002/jmor.20258

5. Lautenschlager, S., L. M. Witmer, Perle A., and E. J. Rayfield. 2013. Edentulism, beaks, and biomechanical innovations in the evolution of theropod dinosaurs. Proceedings of the National Academy of Sciences 110:20657–20662. doi:10.1073/pnas.1310711110.

6. Wroe, S., C. Uphar, W. C. H. Parr, P. Clausen, R. C. Ridgely, and L. M. Witmer. 2103. Comparative biomechanical modeling of metatherian and placental saber-tooths: A different kind of bite for an extreme pouched predator. PLOS ONE 8(6): e66888. doi:10.1371/journal.pone.0066888.

7. Hurlburt, G. R., R. C. Ridgely, and L. M. Witmer. 2013. Relative size of brain and cerebrum in tyrannosaurid dinosaurs: an analysis using brain-endocast quantitative relationships in extant alligators. Pp. 134–154 In J. M. Parrish, R. E. Molnar, P. J. Currie, and E. B. Koppelhus (eds.) Tyrannosaurid Paleobiology. Indiana University Press, Bloomington.

8. Snively, E., J. R. Cotton, John R., R. C. Ridgely, and L. M. Witmer. 2013. Multibody dynamics model of head and neck function in Allosaurus (Dinosauria, Theropoda), Palaeontologia Electronica Vol. 16 (2) 11A 29p.
   • Visit the Project Page for a Common Language Summary, PDF download of the article, and other content
   • Check out the YouTube video
   • Visit the Palaeontologia Electronica site

9. Knoll, F., R. C. Ridgely, F. Ortega, J. L. Sanz, and L. M. Witmer. 2013. Neurocranial osteology and neuroanatomy of a Late Cretaceous titanosaurian sauropod from Spain (Ampelosaurus sp.). PLOS ONE 8(1): e54991. doi:10.1371/journal.pone.0054991.

10. Lautenschlager, S., E. J. Rayfield, Perle A., L. E. Zanno, and L. M. Witmer. 2012. The endocranial anatomy of Therizinosauria and its implications for sensory and cognitive function. PLOS ONE 7(12): e52289. doi:10.1371/journal.pone.0052289.

11. Young, M. T., E. J. Rayfield, C. M. Holliday, L. M. Witmer, D. J. Button, P. Upchurch, and P. M. Barrett. 2012. Cranial biomechanics of Diplodocus (Dinosauria, Sauropoda): testing hypotheses of feeding behaviour in an extinct megaherbivore. Naturwissenschaften 99(8):637–643.

12. Dilkes, D. W., J. R. Hutchinson, C. M. Holliday, and L. M. Witmer. 2012. Reconstructing the musculature of dinosaurs. Pp. 150–190 In M. K. Brett-Surman, T. R. Holtz, and J. O. Farlow (eds.) The Complete Dinosaur, 2nd Edition. Indiana University Press, Bloomington.

13. Knoll, F., L. M. Witmer, F. Ortega, R. C. Ridgely, and D. Schwarz-Wings. 2012. The braincase of the basal sauropod dinosaur Spinophorosaurus and 3D reconstructions of the cranial endocast and inner ear. PLoS ONE 7(1): e30060. doi:10.1371/journal.pone.0030060

14. Miyahsita, T., V. M. Arbour, L. M. Witmer, and P. J. Currie. 2011. The internal cranial morphology of an armoured dinosaur Euoplocephalus corroborated by X-ray computed tomographic reconstruction. Journal of Anatomy 219:661–675.
    • Find out the back story on Witmer's blog post on Pick-and-Scalpel

15. Witmer, L. M. 2011. Palaeontology: Archaeopteryx: an icon knocked from its perch. Nature 475:458–459.

16. Tsuihiji, T., M. Watabe, K. Togtbaatar, T. Tsubamoto, R. Barsbold, S. Suzuki, A. H. Lee, R. C. Ridgely, Y. Kawahara, and L. M. Witmer. 2011. Cranial osteology of a juvenile specimen of Tarbosaurus bataar from the Nemegt Formation (Upper Cretaceous) of Bugin Tsav, Mongolia. Journal of Vertebrate Paleontology 31:497–517.

17. Zelenitsky, D. K., F. Therrien, R. C. Ridgely, A. R. McGee, and L. M. Witmer. 2011. Evolution of olfaction in non-avian theropod dinosaurs and birds. Proceedings of the Royal Society B.

18. Witmer, L. M. and R. C. Ridgely. 2010. The Cleveland tyrannosaur skull (Nanotyrannus or Tyrannosaurus): new findings based on CT scanning, with special reference to the braincase. Kirtlandia 57:61–81.

19. Holliday, C. M., R. C. Ridgely, J. C. Sedlmayr, and L. M. Witmer. 2010. Cartilaginous epiphyses in extant archosaurs and their implications for reconstructing limb function in dinosaurs. PLoS ONE 5(9): e13120.
    • Download a PDF of the article with Supporting Information
    • Find out the back story on Witmer's blog post on Pick-and-Scalpel
    • Visit Casey Holliday's site at the University of Missouri for even more resources

20. Degrange, F. J., C. P. Tambussi, K. Moreno, L. M. Witmer, and S. Wroe. 2010. Mechanical analysis of feeding behavior in the extinct "terror bird" Andalgalornis steulleti (Gruiformes: Phorusrhacidae). PLoS ONE 5(8): e11856. doi:10.1371/journal.pone.001185
    • Visit the Project Page for a Common Language Summary, PDF download of the article, and other content
    • Visit a page with Additional Resources, such as news releases, images, movies, animations, and more
    • Visit the WitmerLab Facebook page for a photo album

21. Hieronymus, T. L., and L. M. Witmer. 2010. Homology and evolution of avian compound rhamphothecae. Auk 127:590–604.

22. Witmer, L. M. 2009. Palaeontology: feathered dinosaurs in a tangle. Nature 461:601–602.

23. Holliday, C. M. and L. M. Witmer. 2009. The epipterygoid of crocodyliforms and its significance for the evolution of the orbitotemporal region of eusuchians. Journal of Vertebrate Paleontology 29:715–733.

24. Witmer, L. M. and R. C. Ridgely. 2009. New insights into the brain, braincase, and ear region of tyrannosaurs, with implications for sensory organization and behavior. Anatomical Record 292:1266–1296.

25. Evans, D., L. M. Witmer, and R. C. Ridgely. 2009. Endocranial anatomy of lambeosaurine dinosaurs: a sensorineural perspective on cranial crest function. Anatomical Record 292:1315–1337.

26. Hieronymus, T. L., L. M. Witmer, D. H. Tanke, and P. J. Currie. 2009. The facial integument of centrosaurine ceratopsids: morphological and histological correlates of novel skin structures. Anatomical Record 292:1370–1396.

27. Witmer, L. M. 2009. Dinosaurs: Fuzzy origins for feathers. Nature 458:293–295.

28. Walsh, S. A., P. M. Barrett, A. C. Milner, G. Manley, and L. M. Witmer. 2009. Inner ear anatomy is a proxy for deducing auditory capability and behaviour in reptiles and birds. Proceedings of the Royal Society B 279:1355–1360.
    • Additional information and downloads on the WitmerLab site

29. Witmer, L. M., and R. C. Ridgely. 2008. The paranasal air sinuses of predatory and armored dinosaurs (Archosauria: Theropoda and Ankylosauria) and their contribution to cephalic architecture. Anatomical Record 291:1362–1388.
    • Additional information and downloads on the WitmerLab site

30. Witmer, L. M., and R. C. Ridgely. 2008. Structure of the brain cavity and inner ear of the centrosaurine ceratopsid Pachyrhinosaurus based on CT scanning and 3D visualization. Pp. 117–144 in P. J. Currie (ed.), A New Horned Dinosaur From an Upper Cretaceous Bone Bed in Alberta. National Research Council Research Press, Ottawa..
    • Additional information and downloads on the WitmerLab site

31. Holliday, C. M. and L. M. Witmer. 2008. Cranial kinesis in dinosaurs: intracranial joints, protractor muscles, and their significance for cranial evolution and function in diapsids. Journal of Vertebrate Paleontology 28:1073–1088.
    • Additional information and downloads on the Casey Holliday's site

32. Hurlburt, G. R., R. C. Ridgely, and L. M. Witmer. In press. Relative size of brain and cerebrum in Tyrannosaurus rex: an analysis using brain-endocast quantitative relationships in extant alligators. in Origin, Systematics, and Paleobiology of the Tyrannosauridae, J. M. Parrish, M Henderson, P. J. Currie, E. Koppelhus (Eds.), Northern Illinois University Press.

33. Witmer, L. M., R. C. Ridgely, D. L. Dufeau, and M. C. Semones. 2008. Using CT to peer into the past: 3D visualization of the brain and ear regions of birds, crocodiles, and nonavian dinosaurs. Pp. 67–88 in H. Endo and R. Frey (eds.), Anatomical Imaging: Towards a New Morphology. Springer-Verlag, Tokyo.

34. Witmer, L. M., and R. C. Ridgely. 2008. Air spaces within the heads of dinosaurs and their contribution to cephalic structure. 68^th Annual Meeting of the Society of Vertebrate Paleontology, Cleveland, OH. Journal of Vertebrate Paleontology 28(Supplement to 3):161A.

35. Daniel, J. C., and L. M. Witmer. 2008. What can sediment patterns tell us about soft tissue?: an actualistic taphonomic study of ostrich heads during and pos-burial. 68^th Annual Meeting of the Society of Vertebrate Paleontology, Cleveland, OH. Journal of Vertebrate Paleontology 28(Supplement to 3):69A.

36. Evans, D., L. M. Witmer, R. C. Ridgely, and J. R. Horner. 2008. Endocranial anatomy of lambeosaurine dinosaurs: implications for cranial crest function and evolution. 68^th Annual Meeting of the Society of Vertebrate Paleontology, Cleveland, OH. Journal of Vertebrate Paleontology 28(Supplement to 3):75A.

37. Dufeau, D. L., and L. M. Witmer. 2008. Morphological patterns and phylogenetic trends in theropod braincase pneumaticity. 68^th Annual Meeting of the Society of Vertebrate Paleontology, Cleveland, OH. Journal of Vertebrate Paleontology 28(Supplement to 3):73A

38. Tsuihiji, T., L. M. Witmer, M. Watabe, R. Barsbold, and K. Tsogtbaatar. 2008. New information on the cranial anatomy of Avimimus portentosus (Dinosauria: Theropoda) including virtual endocasts of the brain and inner ear. 68^th Annual Meeting of the Society of Vertebrate Paleontology, Cleveland, OH. Journal of Vertebrate Paleontology 28(Supplement to 3):153A.

39. Ridgely, R. C., and L. M. Witmer. 2008. Gross Anatomical Brain Region Approximation (GABRA): a new technique for assessing brain structure in dinosaurs and other fossil archosaurs. 68^th Annual Meeting of the Society of Vertebrate Paleontology, Cleveland, OH. Journal of Vertebrate Paleontology 28(Supplement to 3):131A

40. Walsh, S. A, A. C. Milner, P. M. Barrett, G. Manley, and L. M. Witmer. 2008. Can hearing and vocalization capacities be estimated from cochlear duct endocasts? 68^th Annual Meeting of the Society of Vertebrate Paleontology, Cleveland, OH. Journal of Vertebrate Paleontology 28(Supplement to 3):157A.

41. Smith, D., D. L. Dufeau, R. K. Sanders, R. C. Ridgely, and L. M. Witmer. 2008. The cranial endocast of Eutretauranosuchus delfsi (Crocodyliformes, Goniopholididae) and its relationship to other cephalic spaces. 68^th Annual Meeting of the Society of Vertebrate Paleontology, Cleveland, OH. Journal of Vertebrate Paleontology 28(Supplement to 3):144A.

42. Degrange, F., K. Moreno, S.  Wroe, C. Tambussi, and L. M. Witmer. 2008. A computational biomechanical approach to the reconstruction of predatory behavior in the terror bird Andalgalornis steulleti. 68^th Annual Meeting of the Society of Vertebrate Paleontology, Cleveland, OH. Journal of Vertebrate Paleontology 28(Supplement to 3):71A.

43. Hieronymus, T. L., and L. M. Witmer. 2008. The facial skin of Majungasaurus crenatissimus (Abelisauridae: Saurischia): pronounced dermal metaplasia as the cause of rugosity in abelisaurid skulls. 68^th Annual Meeting of the Society of Vertebrate Paleontology, Cleveland, OH. Journal of Vertebrate Paleontology 28(Supplement to 3):90A.

44. Iwaniuk, A. N, H. F. James, S. L. Olson, D. R. W. Wylie, R. C. Ridgely, and L. M. Witmer. 2008. Evidence of extreme somatosensory specialization in an extinct duck. Program No. 79.13. 2008 Neuroscience Meeting Planner. Washington, DC: Society for Neuroscience, 2008.

45. Degrange, F. J., C. P. Tambussi, K. Moreno, S. Wroe, and L. M. Witmer. 2008. Andalgalornis steulleti (Aves, Phorusrhacidae): respuesta biomecánica del cráneo ante acción de fuerzas extrínsecas. III Latin American Congress of Vertebrate Paleontology, Neuquén, Argentina.

46. Young, M. T., E. J. Rayfield, P. M. Barrett, P. Upchurch, and L. M. Witmer. 2007. Elucidating the feeding mechanics of Diplodocus longus using the Finite-element method. 51^st Annual Meeting of the Palaeontological Association, Uppsala, Sweden. Palaeontological Association Newsletter 66:101.

47. Sereno, P. C., J. A. Wilson, L. M. Witmer, J. A. Whitlock, A. Maga, O. Ide, and T. A. Rowe. 2007. Structural extremes in a Cretaceous dinosaur. PLoS ONE 2(11): e1230. doi:10.1371/journal.pone.0001230.
    • PLoS Supplementary Information
    • Additional information and downloads on the WitmerLab site
    • Additional information and downloads on the Project Exploration site
    • Additional information and downloads on the DigiMorph site

48. Hieronymus, T. L. and L. M. Witmer. 2007. Turtle beaks, bird beaks, croc beaks? Parallel evolution of rhamphothecae in Sauropsida. Journal of Morphology 268(12):1083–1084.

49. Holliday, C. M. and L. M. Witmer. 2007. Cranial kinesis in dinosaurs: significance for functional inferences and evolution. Journal of Morphology 268(12):1085–1086.

50. Ridgely, R. C. and L. M. Witmer. 2007. Gross Anatomical Brain Region Approximation (GABRA): a new technique for assessing brain structure in dinosaurs and other fossil archosaurs. Journal of Morphology 268(12):1124.

51. Tickhill, J. and L. M. Witmer. 2007. The Virtual Pig Head: digital imaging in cephalic anatomy. Journal of Morphology 268(12):1141.

52. Witmer, L. M. and R. C. Ridgely. 2007. Evolving an on-board flight computer: brain, ears, and exaptation in the evolution of birds and other theropod dinosaurs. Journal of Morphology 268(12):1150.

53. Daniel, J. and L. M. Witmer. 2007. The role of soft tissues in sediment infilling and patterning: an actualistic study with ostrich heads. Journal of Vertebrate Paleontology 27(Supplement to 3):65A.

54. Dufeau, D. L. and L. M. Witmer. 2007. Ontogeny and phylogeny of the tympanic pneumatic system of crocodyliform archosaurs. Journal of Vertebrate Paleontology 27(Supplement to 3):70A.

55. Hieronymus, T. L. and L. M. Witmer. 2007. Skinning dinosaurs: bony correlates and patterns of cephalic skin evolution in Archosauria. Journal of Vertebrate Paleontology 27(Supplement to 3):89A.

56. Holliday, C. M. and L. M. Witmer. 2007. The epipterygoid of crocodyliforms and its significance in the evolution of the orbitotemporal region of eusuchians. Journal of Vertebrate Paleontology 27(Supplement to 3):90A.

57. Tsuihiji, T., M. Watabe, L. M. Witmer, T. Tsubamoto, K. Tsogtbaatar. 2007. A juvenile skeleton of Tarbosaurus with a nearly complete skull and its implications for ontogenetic change in tyrannosaurids. Journal of Vertebrate Paleontology 27(Supplement to 3):160A.

58. Sampson, S. D. and L. M. Witmer. 2007. Craniofacial anatomy of Majungasaurus crenatissimus (Theropoda: Abelisauridae) from the Late Cretaceous of Madagascar. Memoirs of the Society of Vertebrate Paleontology 8, Journal of Vertebrate Paleontology 27(Supplement to 2):32–102.
    • Additional information, downloads, and 3D digital visualizations on the WitmerLab site

59. Holliday, C. M. and L. M. Witmer. 2007. Archosaur adductor chamber evolution: integration of musculoskeletal and topological criteria in jaw muscle homology. Journal of Morphology 268:457–484.
    • Additional information and downloads on Casey Holliday's site

60. Holliday, C. M., R. C. Ridgely, A. M. Balanoff, and L. M. Witmer. 2006. Cephalic vascular anatomy in flamingos (Phoenicopterus ruber) based on novel vascular injection and computed tomographic imaging analyses. Anatomical Record 288A(10)1031–41.
    • Additional information and downloads on the WitmerLab site

61. Hieronymus, T. L., R. C. Ridgely, and L. M. Witmer. 2006. Structure of white rhinoceros (Ceratotherium simum) horn investigated by x-ray computed tomography and histology with implications for growth and external form. Journal of Morphology 267:1172–1176.
    • Additional information and downloads on the WitmerLab site.

62. Dufeau, D. L. and L. M. Witmer. 2006b. Tympanic pneumaticity in Archosauria: Recognizing patterns of organization and homology. 66^th Annual Meeting of the Society of Vertebrate Paleontology, Ottawa, ON. Journal of Vertebrate Paleontology 26(Suppl. to 3):57A.

63. Holliday, C. M. and L. M. Witmer. 2006b. Critical appraisal of cranial kinesis in dinosaurs. 66^th Annual Meeting of the Society of Vertebrate Paleontology, Ottawa, ON. Journal of Vertebrate Paleontology 26(Suppl. to 3):77A.

64. Hieronymus, T. L., D. Tanke, P. J. Currie, and L. M. Witmer. 2006b. Horn morphology of Pachyrhinosaurus and horn evolution in centrosaurine dinosaurs. 66^th Annual Meeting of the Society of Vertebrate Paleontology, Ottawa, ON. Journal of Vertebrate Paleontology 26(Suppl. to 3):75A.Holliday, C. M. and L. M. Witmer. 2006b. Critical appraisal of cranial kinesis in dinosaurs. 66^th Annual Meeting of the Society of Vertebrate Paleontology, Ottawa, ON. Journal of Vertebrate Paleontology 26(Suppl. to 3):77A.

65. Ridgely, R. C. and L. M. Witmer. 2006b. Dead on arrival: optimizing CT data acquisition of fossils using modern hospital CT scanners. 66^th Annual Meeting of the Society of Vertebrate Paleontology, Ottawa, ON. Journal of Vertebrate Paleontology 26(Suppl. to 3):115A.

66. Daniel, J. and L. M. Witmer. 2005. Heads and skulls as sediment sorters: an actualistic, CT-based study in taphonomy. 65^th Annual Meeting of the Society of Vertebrate Paleontology, Mesa, AZ. Journal of Vertebrate Paleontology 25(Suppl. to 3):13A.

67. Hieronymus, T. L. and L. M. Witmer. 2005. Revised horn reconstructions for extinct rhinocerotid taxa: separating the contributions of dermis and epidermis to horn rugosities. 65^th Annual Meeting of the Society of Vertebrate Paleontology, Mesa, AZ. Journal of Vertebrate Paleontology 25(Suppl. to 3):29A.

68. Witmer, L. M. and R. C. Ridgely. 2005. Tyrannosaur brain and ear structure: ontogeny and implications for sensory function and behavior. 65^th Annual Meeting of the Society of Vertebrate Paleontology, Mesa, AZ. Journal of Vertebrate Paleontology 25(Suppl. to 3):10A.

69. Clifford, A. B. and L. M. Witmer.  2004.  Case studies in novel narial anatomy: 3. Structure and function of the nasal cavity of saiga (Artiodactyla: Bovidae: Saiga tatarica). Journal of Zoology 264:217–230.
    • Digital Visualizations on DigiMorph.org.

70. Norman, D. B., L. M. Witmer, and D. B. Weishampel.  2004.  Basal Ornithischia. Pp. 325–334  in The Dinosauria, 2nd Edition, D. B. Weishampel, P. Dodson, and H. Osmólska (Eds.), University of California Press, Berkeley.

71. Norman, D. B., L. M. Witmer, and D. B. Weishampel.  2004.  Basal Thyreophora. Pp. 335-342 in The Dinosauria, 2nd Edition, D. B. Weishampel, P. Dodson, and H. Osmólska (Eds.), University of California Press, Berkeley.

72. Norman, D. B., H.-D. Sues, L. M. Witmer, and R. A. Coria.  2004.  Basal Ornithopoda. Pp. 393–412 in The Dinosauria, 2nd Edition, D. B. Weishampel, P. Dodson, and H. Osmólska (Eds.), University of California Press, Berkeley.

73. Witmer, L. M., R. C. Ridgely, H. Mayle, and D. Adams.  2004.  The best of both worlds: integrating CT and MR in a study of pig knees. RT Image 17(32):16–19.
    • RT Image web site
    • Digital Visualizations on DigiMorph.org

74. Witmer, L. M.  2004.  Inside the oldest bird brain.  Nature 430:619–620.

75. Clifford, A. B. and L. M. Witmer.  2004.  Case studies in novel narial anatomy: 2. The enigmatic nose of moose (Artiodactyla: Cervidae: Alces alces). Journal of Zoology 262:339–360. 
    • Accompanying Nature "News & Views"
    • Digital visualizations on DigiMorph.org

76. L. M. Witmer, R. C. Ridgely, and S. D. Sampson.  2004.The ear region, cerebral endocast, and cephalic sinuses of the abelisaurid theropod dinosaur Majungatholus. 64^th Annual Meeting of the Society of Vertebrate Paleontology, Denver, CO. Journal of Vertebrate Paleontology 24(Suppl. to 3):131A.

77. Ridgely, R. C. and L. M. Witmer.  2004.  New applications of CT scanning and 3D modeling for dinosaur visualization. 64^th Annual Meeting of the Society of Vertebrate Paleontology, Denver, CO. Journal of Vertebrate Paleontology 24(Suppl. to 3):103A–104A.

78. Holliday, C. M. and L. M. Witmer.  2004.  Anatomical domains within the heads of archosaurs and their relevance for functional interpretation. 64^th Annual Meeting of the Society of Vertebrate Paleontology, Denver, CO. Journal of Vertebrate Paleontology 24(Suppl. to 3):71A

79. Hieronymus, T. L. and L. M. Witmer.  2004.  Cranial rugosity and dinosaur “horns:” rhino and giraffe as model systems for skin reconstruction in fossil taxa. 64^th Annual Meeting of the Society of Vertebrate Paleontology, Denver, CO. Journal of Vertebrate Paleontology 24(Suppl. to 3):70A.

80. L. M. Witmer.  2004.  Invited Plenary Lecture—Fleshing out fossils: the present as the key to a very different past. 7^th International Congress of Vertebrate Morphology, Boca Raton, FL. Journal of Morphology 260(3):340.

81. Ridgely, R. C. and L. M. Witmer.  2004.  Beyond imaging: Using 3D-datasets in comparative morphology . 7^th International Congress of Vertebrate Morphology, Boca Raton, FL. Journal of Morphology 260(3):321–322.

82. Clifford, A. B. and L. M. Witmer.  2004.  Rules of construction in proboscis building in mammals. 7^th International Congress of Vertebrate Morphology, Boca Raton, FL. Journal of Morphology 260(3):283.

83. Hieronymus, T. L. and L. M. Witmer.  2004.  Rhinoceros horn attachment: anatomy and histology of a dermally influenced bone rugosity. 7^th International Congress of Vertebrate Morphology, Boca Raton, FL. Journal of Morphology 260(3):298.

84. Witmer, L. M., S. Chatterjee, J. Franzosa, T. Rowe, and R. C. Ridgely. 2004. Neuroanatomy and vestibular apparatus of pterosaurs: Implications for flight, posture, and behavior. Annual Meeting of the Society of Integrative and Comparative Biology, New Orleans, LA. Integrative and Comparative Biology 43(6):832.

85. Hieronymus, T. L. and L. M. Witmer. 2003. Dermal enthesis: anatomy and histology of rhinoceros horn attachment. Annual Meeting of the Society of Integrative and Comparative Biology, New Orleans, LA. Integrative and Comparative Biology 43(6):874

86. Witmer, L. M., S. Chatterjee, J. Franzosa, and T. Rowe.  2003.  Neuroanatomy of flying reptiles and implications for flight, posture and behaviour. Nature 425:950–953.
    • Accompanying Nature "News & Views"
    • Digital visualizations on DigiMorph.org: Anhanguera, Rhamphorhynchus

87. You H.-L., Luo Z.-X.,N. H. Shubin, L. M. Witmer, Tang A.-L, and Tang F.  2003.  The earliest-known duck-billed dinosaur from deposits of late Early Cretaceous age in northwestern China and hadrosaur evolution. Cretaceous Research 24:346–355.

88. Hill, R. V., L. M. Witmer, M.A. Norell. 2003.  A new specimen of Pinacosaurus grangeri (Dinosauria: Ornithischia) from the Late Cretaceous of Mongolia: ontogeny and phylogeny of ankylosaurs. American Museum Novitates 3395:1–29.

89. Witmer, L. M. 2003. Narial anatomy of ankylosaurian dinosaurs: osteology and soft-tissue reconstruction. 63^rd Annual Meeting of the Society of Vertebrate Paleontology, St. Paul, MN. Journal of Vertebrate Paleontology 23(Suppl. to 3):111A–112A.

90. Hieronymus, T. L. and L. M. Witmer. 2003. Rhinoceros horn attachment: anatomy and histology. 63^rd Annual Meeting of the Society of Vertebrate Paleontology, St. Paul, MN. Journal of Vertebrate Paleontology 23(Suppl. to 3):61A.

91. Clifford, A. B., and Witmer, L. M.  2003. Nasal structures in moose (Cervidae: Alces). Annual Meeting of the Society of Integrative and Comparative Biology, Toronto, Ontario. Integrative and Comparative Biology 42(6):1209.

92. Holliday, C. M. and L. M. Witmer.  2003. Archosaur jaw muscle homology and evolution: extant data. Annual Meeting of the Society of Integrative and Comparative Biology, Toronto, Ontario. Integrative and Comparative Biology 42(6):1244–1245.

93. Bimber, O., S. M. Gatesy, L. M. Witmer, R. Raskar, and L. M. Encarnação.  2002.  Merging fossil specimens with computer-generated information. IEEE Computer 35(9):25–30.

94. Chiappe, L. M. and L. M. Witmer. 2002. Preface. Pp. ix–xii in Mesozoic Birds: Above the Heads of Dinosaurs, L. M. Chiappe and L. M. Witmer (eds.), University of California Press, Berkeley.

95. Witmer, L. M.  2002.  The debate on avian ancestry: phylogeny, function, and fossils. pp. 3–30; In Mesozoic Birds: Above the Heads of Dinosaurs, L. M. Chiappe and L. M. Witmer (eds.), University of California Press, Berkeley.

96. Witmer, L. M., S. Chatterjee, T. Rowe, and J. Franzosa.  2002.  Anatomy of the brain and vestibular apparatus in two pterosaurs: implications for flight, head posture, and behavior. Annual Meeting of the Society of Vertebrate Paleontology, Norman, Oklahoma. Journal of Vertebrate Paleontology 22(Suppl. to 3):120A–121A.

97. Bimber, O., S. M. Gatesy, and L. M. Witmer.  2002.  Augmented paleontology: merging fossil specimens with computer generated information for analysis and education. Annual Meeting of the Society of Vertebrate Paleontology.  Journal of Vertebrate Paleontology 22(Suppl. to 3):36A.

98. Holliday, C. M. and L. M. Witmer.  2002.  Functional anatomy of archosaurian jaw musculature: preliminary findings. Annual Meeting of the Society of Vertebrate Paleontology.  Journal of Vertebrate Paleontology 22(Suppl. to 3):66A.

99. Clifford, A. B. and L. M. Witmer.  2002.  Not all noses are hoses: an appraisal of proboscis evolution in mammals. Annual Meeting of the Society of Vertebrate Paleontology.  Journal of Vertebrate Paleontology 22(Suppl. to 3):45.

100. Sedlmayr, J. C. and L. M. Witmer.  2002.  Rapid technique for imaging the blood vascular system using stereoangiography. Anatomical Record 267(4):330–336.

101. Clifford, A. B., and L. M. Witmer.  2002. Proboscis evolution in Mammalia: preliminary studies. Annual Meeting of the Society of Integrative and Comparative Biology, Anaheim, California. American Zoologist 41(6):1412.

102. Holliday, C. M., R. C. Ridgely, J. C. Sedlmayr, and L. M. Witmer. 2002.  The articular cartilage of extant archosaur limb bones: implications for dinosaur functional morphology and allometry.  Annual Meeting of the Society of Integrative and Comparative Biology, Anaheim, California. American Zoologist 41 (6):1473–1474.  

103. Witmer, L. M.  2001.  Nostril position in dinosaurs and other vertebrates and its significance for nasal function.  Science 293:850–853.
     • Erik Stokstad's accompanying piece in Science 

104. Witmer, L. M.  2001.  The role of Protoavis in the debate on avian origins. pp. 537–548  In New Perspectives on the Origin and Early Evolution of Birds, J. A. Gauthier and L. F. Gall (eds.), Yale Peabody Museum, New Haven.

105. Witmer, L. M.  2001.  A nose for all reasons. Natural History Magazine, June 2001 110(5):64–71.

106. Witmer, L. M.  2001. The position of the fleshy nostril in dinosaurs and other vertebrates and its significance for nasal function. Journal of Vertebrate Paleontology 21 (3:Suppl.).

107. Holliday, C. M., R. C. Ridgely, J. C. Sedlmayr, and Witmer, L. M.  2001.  The articular cartilage of extant archosaur limb bones: implications for dinosaur functional morphology and allometry. Journal of Vertebrate Paleontology 21 (3:Suppl.).

108. Hill, R. V., L. M. Witmer, and M. A. Norell.  2001.  A new juvenile specimen of Pinacosaurus grangeri: ontogeny and phylogeny of ankylosaurs. Journal of Vertebrate Paleontology 21 (3:Suppl.).

109. Witmer, L. M.  2001. Reconstructing the evolutionary history of the nasal apparatus of Sauropsida, with special reference to Archosauria. Journal of Morphology 248(3):301.

110. Sedlmayr, J. C., and L. M Witmer.  2001. Homology and evolution of cephalic vasculature in Archosauria. Journal of Morphology 248(3):282.  

111. Holliday, C. M., J. C. Sedlmayr, R. C. Ridgely, and L. M Witmer.  2001. The articular cartilage of extant archosaur long bones: implications for dinosaur functional morphology and allometry. Journal of Morphology 248(3):241–242.

112. Clifford, A. B., and L. M. Witmer.  2001. The narial anatomy of hooded seals (Cystophora cristata) with respect to other Carnivora.   American Zoologist 40(6):976.

113. Sedlmayr, J. C, and L. M. Witmer.  2001. The avian suborbital air sac and its potential role in brain cooling in birds and other dinosaurs.  American Zoologist 40(6):1206.    

114. Witmer, L. M.  2000.  Science, art, and dinosaurs. Essay and review of Dinosaur Imagery: The Science of Lost Worlds and Jurassic Art, edited by C. R. Crumly. Science 5489:460–461.

115. Witmer, L. M.  2000.  Narial anatomy of extant amniotes and its significance for the interpretation of narial novelty in dinosaurs. American Zoologist 39(5):98A–99A.

116. Sedlmayr, J. C, and L. M. Witmer.  2000. Cephalic vasculature in Archosauria, with special emphasis on the nasal region. American Zoologist 39(5):16A.

117. Witmer, L. M.  1999.  New aspects of avian origins: roundtable report. pp. 327–334 in Avian Paleontology at the Close of the 20^th Century: Proceedings of the 4^th International Meeting of the Society of Avian Paleontology and Evolution, S. L. Olson (ed.), Smithsonian Contributions to Paleobiology 89, Washington.

118. Witmer, L. M., S. D. Sampson, and N. Solounias.  1999. The proboscis of tapirs (Tapirus terrestris): a case study in novel narial anatomy. Journal of Zoology 249:249–267.

119. Solounias, N., J. M. Plavcan, J. Quade and L. M. Witmer. 1999.  The paleoecology of the Pikermian Biome and the savanna myth.  pp. 426-453.  In Hominoid Evolution and Climatic Change in Europe. Volume 1: The Evolution of Neogene Terrestrial Ecosystems in Europe.  J. Agustí, L. Rook, and P. Andrews (eds.).  Cambridge University Press, New York.

120. Witmer, L. M. 1999. The phylogenetic history of paranasal air sinuses. pp. 21–34 in The Paranasal Sinuses of Higher Primates: Development, Function and Evolution, T. Koppe, H. Nagai, and K. W. Alt (eds.), Quintessence, Chicago.

121. Witmer, L. M., and S. D. Sampson.  1999.  Nasal conchae and blood supply in some dinosaurs: physiological implications. Journal of Vertebrate Paleontology 19(3:Suppl.):85A.

122. Sampson, S. D., and L. M. Witmer.  1999.  Novel narial anatomy in ceratopsid dinosaurs. Journal of Vertebrate Paleontology 19(3:Suppl.):72A-73A.

123. Sedlmayr, J. C., and L. M. Witmer.  1999.  Cephalic vasculature in extant and fossil Archosauria, with special emphasis on the nasal region. Journal of Vertebrate Paleontology 19(3:Suppl.)74A.

124. Sampson, S. D., L. M. Witmer, C. A. Forster, D. W. Krause, P. M. O’Connor, P. Dodson, and F. Ravoavy.  1998.  Predatory dinosaur remains from Madagascar: implications for the Cretaceous biogeography of Gondwana. Science 280:1048–1051.

125. Witmer, L. M.  1998.  Dinosaurs for adults.  Review of The Complete Dinosaur by J. O. Farlow and M. K. Brett-Surman.  Science 280:223.

126. Witmer, L. M.  1998.  Aspects of comparative cross-sectional anatomy of dinosaur skulls. Journal of Vertebrate Paleontology 18(3:Suppl.):87A.

127. Witmer, L. M.  1998.  Application of the extant phylogenetic bracket (EPB) approach to the problem of anatomical novelty in the fossil record. Journal of Vertebrate Paleontology 18(3:Suppl.):87A.

128. Papp, M. J., and L. M. Witmer.  1998.  Cheeks, beaks, or freaks: a critical appraisal of buccal soft-tissue anatomy in ornithischian dinosaurs. Journal of Vertebrate Paleontology 18(3:Suppl.):69A.

129. Sedlmayr, J. C., and L. M. Witmer.  1998.  Facial vasculature in extant Crocodylia with special reference to narial cavernous tissue. 58th Annual Meeting of the Society of Vertebrate Paleontology, Snowbird, Utah, October.  Journal of Vertebrate Paleontology 18(3:Suppl.):76A.

130. Sampson, S. D., Witmer, L. M., Forster, C. A., and Krause, D. W.  1998. The evolution and biogeography of Gondwanan theropod dinosaurs: new information from the Late Cretaceous of Madagascar.  Journal of African Earth Sciences 27(1A): 167-168.

131. Witmer, L. M.  1997.  Craniofacial air sinus systems.  pp. 151–159 in The Encyclopedia of Dinosaurs, P. J. Currie and K. Padian (eds.), Academic Press, New York.

132. Witmer, L. M.  1997.  Foreword.  pp. vii–xii in The Rise of Birds: 225 Million Years of Evolution, by S. Chatterjee, Johns Hopkins University Press, Baltimore.

133. Witmer, L. M.   1997.  Flying feathers.  Review of The Origin and Evolution of Birds by Alan Feduccia.  Science 276:1209–1210.

134. Witmer, L. M.  1997.  A new missing link.  Nature 387:349–350.

135. Witmer, L. M.  1997.  The evolution of the antorbital cavity of archosaurs: a study in soft-tissue reconstruction in the fossil record with an analysis of the function of pneumaticity.  Memoirs of the Society of Vertebrate Paleontology, Journal of Vertebrate Paleontology 17(Supplement to 1):1–73.

136. Witmer, L. M., and S. D. Sampson.  1997. The interpretation of narial novelty in dinosaurs.  Journal of Vertebrate Paleontology 17(3:Suppl.):98A.

137. Sampson, S. D., L. M. Witmer, C. A. Forster, D. W. Krause, and P. M. O’Connor.  1997.  Discovery of a complete theropod skull from the Late Cretaceous of Madagascar yields new information on Majungasaurus, Majungatholus, and Abelisauridae.  Journal of Vertebrate Paleontology 17(3:Suppl.):73A.

138. Witmer, L. M.  1997.  Bones, air sacs, and natural selection: a new perspective on the function of pneumatic sinuses. Journal of Morphology 232:340

139. Witmer, L. M., and S. D. Sampson.  1997.  Unusual narial structures in dinosaurs and other vertebrates. Fifth International Congress of Vertebrate Morphology, Bristol, England.  Journal of Morphology 232:340

140. Maxwell W. D. and L. M. Witmer. 1996. The new Deinonychus. Dinosaur Discoveries 2:1–2.

141. Witmer, L. M., and W. D. Maxwell.  1996.  The skull of Deinonychus (Dinosauria: Theropoda): new insights and implications. Journal of Vertebrate Paleontology 16(3:Suppl.):73A.

142. Maxwell, W. D., and L. M. Witmer.  1996.  New material of Deinonychus (Dinosauria: Theropoda).  Journal of Vertebrate Paleontology 16(3:Suppl.):51A.

143. Witmer, L. M.  1995.  Homology of facial structures in extant archosaurs (birds and crocodilians), with special reference to paranasal pneumaticity and nasal conchae.  Journal of Morphology 225:269–327.

144. Witmer, L. M.  1995.  The Extant Phylogenetic Bracket and the importance of reconstructing soft tissues in fossils.  pp. 19–33 in Functional Morphology in Vertebrate Paleontology, J. J. Thomason (ed.), Cambridge Univ. Press, New York.

145. Witmer, L. M.  1995.  The Search for the Origin of Birds. Franklin Watts, New York.

146. Witmer, L. M.  1995. Craniofacial pneumaticity and skull evolution in theropod dinosaurs. Journal of Vertebrate Paleontology 15(3:Suppl.):60A.

147. Witmer, L. M.  1994.  A revised terminology of the facial osteology of archosauriforms. Journal of Vertebrate Paleontology 14(3:Suppl.):53A.

148. Baumel, J. J. and Witmer, L. M.  1993.   Osteologia. pp. 45–132 in Handbook of Avian Anatomy: Nomina Anatomica Avium, Second Edition, J. J. Baumel (ed.),  Publications of the Nuttall Ornithological Club No. 23, Cambridge.

149. Witmer, L. M., and D. B. Weishampel. 1993.  Remains of theropod dinosaurs from the Upper Cretaceous St. Mary River Formation of northwestern Montana, with special reference to a new maniraptoran braincase. Journal of Vertebrate Paleontology 13(3:Suppl.):63A.

150. Witmer, L. M.  1992.  The many faces of archosaurs: some trends in archosaur facial evolution. Journal of Vertebrate Paleontology 12(3:Suppl.):60A.

151. Witmer, L. M. and Rose, K. D.  1991.  Biomechanics of the jaw apparatus of the gigantic Eocene bird Diatryma: Implications for diet and mode of life. Paleobiology 17:95–120.   

152. Witmer, L. M.  1991.  Perspectives on avian origins. pp. 427–466 in Origins of the Higher Groups of Tetrapods: Controversy and Consensus, H.-P. Schultze and L. Trueb (eds.), Cornell University Press, Ithaca.

153. Witmer, L. M.  1991.  Homology of facial structures in extant crocodilians and birds: implications for the antorbital fossa of archosaurs. Journal of Vertebrate Paleontology 11(3:Suppl.):62A.

154. Witmer, L. M.  1990.  The craniofacial air sac system of Mesozoic birds (Aves). Zoological Journal of the Linnean Society. 100:327–378.

155. Weishampel, D. B. and Witmer, L. M.  1990.  Lesothosaurus, Pisanosaurus, and Technosaurus. pp. 416–425 in The Dinosauria, D. B. Weishampel, P. Dodson, and H. Osmólska (Eds.), University of California Press, Berkeley.

156. Weishampel, D. B. and Witmer, L. M.  1990.  Heterodontosauridae. pp. 486–497 in The Dinosauria, D. B. Weishampel, P. Dodson, and H. Osmólska (Eds.), University of California Press, Berkeley.

157. Coombs, W. P., Weishampel, D. B., and Witmer, L. M.  1990. Basal Thyreophora. pp. 427–434 in The Dinosauria, D. B. Weishampel, P. Dodson, and H. Osmólska (Eds.), University of California Press, Berkeley.

158. Witmer, L. M.  1990.  A new pleurodiran turtle from the Miocene of east Africa and its implications for the phylogenetics of podocnemine pelomedusids.  Journal of Vertebrate Paleontology 10(3:Suppl.):49A–50A.

159. Martin, L. D., and L. M. Witmer. 1990.  The avifauna of the Niobrara Chalk.  Journal of Vertebrate Paleontology 10(3:Suppl.):34A.

160. Witmer, L. M.  1989.  Review: Peter W. Houde: Paleognathous birds from the early Tertiary of the Northern Hemisphere. Journal of Vertebrate Paleontology 9(3):368–369.

161. Witmer, L. M., K. D. Rose, and T. M. Bown.  1989.  Jaw biomechanics and feeding behavior of the gigantic Eocene bird Diatryma. Journal of Vertebrate Paleontology 9(3:Suppl.):45A.

162. Bühler, P., Martin, L. D., and Witmer, L. M.  1988.  Cranial kinesis in the Late Cretaceous birds Hesperornis and Parahesperornis. Auk: Quarterly Journal of Ornithology 105(1):111–122.

163. Witmer, L. M.  1988.  Mechanisms of sound localization in some fossil archosaurs.  Journal of Vertebrate Paleontology 8(3:Suppl.):29A.

164. Witmer, L. M. and Martin, L. D.  1987.  The primitive features of the avian palate with special reference to Mesozoic birds. Documents des Laboratoires de Géologie, Lyon 99:21–40.

165. Witmer, L. M.  1987.  The nature of the antorbital fossa of archosaurs: shifting the null hypothesis.  pp. 230–235 in Fourth Symposium on Mesozoic Terrestrial Ecosystems, Short Papers (P. J. Currie and E. H. Koster, Eds.), Occasional Paper of the Tyrrell Museum of Palaeontology #3, Drumheller, Alberta.

166. Witmer, L. M.  1979.  Host preference by the Mid-Devonian (Moscow Shale) epizoans Hederella filiformis, Aulopora elleri, Spirorbis sp., and Cornulites sp.  Cornell Journal of Biological Science 1979:21–28.