Above: Three-dimensional reconstruction of the maxillary bone of a fetal Tarsius syrichta (center) with depositional (red) and resorptive (yellow) surfaces coded. On the left are soft tissue structures that influence bone growth remodeling, such as the eye (green), nasal mucosa (orange) infraorbital nerve (brown) and dental follicles (blue). On the right, these structures are ghosted against the bone. (image (c) Timothy D. Smith, 2011). Related to current research: "Making Monkey Faces:  An Investigation of Bone Growth Patterns that Shape the Forward-Facing Eyes of Our Primate Relatives" (PASSHE grant)

Timothy D. Smith, PhD
Associate Professor, School of Physical Therapy,
Slippery Rock University, Slippery Rock, PA 16057
Phone: 724 738 2885
Fax: 724 738 2113
Comments: E-mail: timothy.smith@sru.edu
or click here


I received a Ph.D. in physical anthropology from the University of Pittsburgh in 1995. My dissertation concerned growth and development of the human vomeronasal organ in fetuses with and without cleft lip and palate. My current teaching foci include gross anatomy, embryology, and histology. Other teaching interests include fetal development and congenital disorders and  comparative anatomy. For further information, see links to the histology atlas and current research interests.



1985-1989 B.F.A. (Drawing)  Carnegie-Mellon University, Pittsburgh, PA.


1990-1993 M.A. (Physical Anthropology)    University of Pittsburgh, Pittsburgh, PA.

1994-1995 Ph.D. (Physical Anthropology)    University of Pittsburgh, Pittsburgh, PA.

Teaching Responsibilities

PT 630, PT 635 Gross Anatomy I & II

PT 627 Human Development

PT 633 Histology (Link to Histology Atlas)

Research Interests

I am devoted to fostering student involvement in research. My research interests include comparative histology and histochemistry of the chemical sense organs, functional anatomy and histology of the shoulder and hip, and craniofacial development and growth. A special emphasis of my research has focused on evolution of the vomeronasal complex in primates. My graduate students and I are investigating functional correlates of bony articular surfaces and limb muscle masses in mammalian species that exhibit differing locomotor behaviors. Finally, I have an interest in functional anatomy and locomotion of river dolphins. Most of my published work (see below) is a collaborative effort with students from Slippery Rock University (in red) and scientists from numerous other institutions. 


       1998 Guest Editor

                            1. Microscopy Research & Technique (Vol. 41, No. 6), spec. issue: Histology and Histochemistry of the Vomeronasal Organ, Pt I

            2. Microscopy Research & Technique (Vol. 43, No. 6), spec.  issue: Histology and Histochemistry of the Vomeronasal Organ, Pt II  

        2004 Guest Editor (with CF Ross and NJ Dominy)

The Anatomical Record (Vol. 281A, no. 1, 2004), special issue: Evolution of the Special Senses in Primates

        2004 Co-Chair/Co-Organizer (with CF Ross, NJ Dominy)

Symposium: Evolution of the Special Senses in Primates. 73nd annual meeting of the American Association of Physical Anthropologists, April 2004, Tampa FLA.  

   2010 Guest Editor (with JM Organ, VB DeLeon, and Q Wang)  

                          The Anatomical Record (Vol. 293, no. 4, 2010, special issue: From Head to Tail: New Models and Approaches in Primate Functional Anatomy and  Biomechanics

Selected Publications

Smith TD, JB Rossie, GM Cooper, RM Schmieg, CJ Bonar, MP Mooney, and MI Siegel. 2011. Comparative micro CT and histological study of maxillary pneumatization in four species of New            World monkeys: The perinatal period. Am J Phys Anthropol, 144, no. 3: 392-410. e-mail for a PDF  

Smith, TD, JC Dennis, KP  Bhatnagar, EC Garrett, CJ Bonar,  and EE Morrison. 2011. Olfactory marker protein expression in the vomeronasal neuroepithelium of tamarins (Saguinus spp). Brain Research 1375: 7-18. e-mail for a PDF  

Smith TD, Burrows AM, Dumont ER. 2010. Microanatomical assessment of nasomaxillary suture patency. Anatomical Record, 293:651-657. e-mail for a PDF

Smith TD, JB Rossie, GM Cooper, KA Carmody, RM Schmieg, CJ Bonar, MP Mooney, MI Siegel. 2010. The maxillary sinus in three genera of New World monkeys: Factors that constrain secondary pneumatization. Anatomical Record, 203:91-107. e-mail for a PDF

Reinholt LE, AM Burrows, TP Eiting, ER Dumont, and TD Smith. 2009. Brief communication: Histology and microCT as methods for assessing facial suture patency. American Journal of   Physical Anthropology, 138:499-506. e-mail for a PDF

Smith TD, JB Rossie. 2008. Nasal Fossa of Mouse and Dwarf Lemurs (Primates, Cheirogaleidae). Anat Rec, 291:895-915.e-mail for a PDF  

Smith TD, JB Rossie, KP Bhatnagar. 2007. Evolution of the nose and nasal skeleton in primates. Evolutionary Anthropology, 16:132-146.e-mail for a PDF

Smith TD, KP Bhatnagar, JB Rossie, BA Docherty, AM Burrows, MP Mooney, MI Siegel. 2007. Scaling of the first ethmoturbinal in nocturnal strepsirrhines: olfactory and respiratory surfaces. Anatomical Record, 290:215-237.e-mail for a PDF

Rossie JB, TD Smith. 2007. Ontogeny of the nasolacrimal duct in Haplorhini (Mammalia: Primates): functional and phylogenetic significance. Journal of Anatomy 210:195-208.

Atzeva M,  B Demes, ML Kirkbride, AM Burrows, and TD Smith. 2007. Comparison of hind limb muscle mass in neonate and adult prosimian primates. Journal of Human Evolution, 52:231-242.e-mail for a PDF

Bhatnagar KP, and TD Smith. 2006. The vomeronasal organ and its evolutionary loss in catarrhine primates. In: The Evolution of Primate Nervous Systems. Vol. 5. TM Preuss and JH Kaas (editors). p 141-148.

Smith TD, Rossie JB. 2006. Primate olfaction: Anatomy and evolution. In:  Olfaction and the brain: Window to the mind, Brewer W, Castle D, Pantelis C (eds), Cambridge University Press, p 135-166. e-mail for a PDF

Smith, TD, LJ Alport, AM Burrows, KP Bhatnagar, JC Dennis, P Tuladhar, EE Morrison. 2007. Perinatal size and maturation of the olfactory and vomeronasal neuroepithelia in lorisoids and lemuroids. American Journal of Primatology 69:74-85. in PDF format.

Smith TD, KP Bhatnagar, JC Dennis, EE Morrison, TJ Park. 2007. Growth deficient vomeronasal organs in the naked mole-rat (Heterocephalus glaber). Brain Research, 1132:78-83. e-mail for a PDF

Smith, TD, KP Bhatnagar, AM Burrows, KL Shimp, JC Dennis, MA Smith, L Maico-Tan, and EE Morrison. 2005. The vomeronasal organ of greater bushbabies (Otolemur spp.): species, sex, and age differences. Journal of Neurocytology 34:135-147. in PDF format.

Smith, TD, JB Rossie, GM Cooper, MP Mooney, and MI Siegel. 2005. Secondary pneumatization in the maxillary sinus of callitrichid primates: insights from immunohistochemistry and bone cell distribution.  Anatomical Record 285A:677-689. See ancillary material related to this articlein PDF format.

Kinzinger, JH, EW Johnson, KP Bhatnagar, CJ Bonar, AM Burrows, MP Mooney, MI  Siegel, TD Smith. 2005. Comparative study of lectin reactvity in the vomeronasal organ of human and non-human primates. Anatomical Record, Part A, 284A:550-560. in PDF format.

Smith TD, and KP Bhatnagar. 2004. Microsmatic primates: Reconsidering how and when size matters. Anatomical Record, 279B:24-31.in PDF format

Smith TD, KP Bhatnagar, P Tuladhar, and AM Burrows. 2004. Distribution of olfactory epithelium in the primate nasal cavity: Are “microsmia” and “macrosmia” valid morphological concepts? Anatomical Record, 281A:1173-1181. in PDF format

Smith, TD,  JC Dennis, KP Bhatnagar, CJ Bonar, AM Burrows, and EE Morrison. 2004. Ontogenetic observations on the vomeronasal organ in two species of tamarins using neuron-specific beta III tubulin. Anatomical Record, 278A:409-414. in PDF format

Bhatnagar, KP and TD Smith. 2003. The Human Vomeronasal Organ. Part V. An interpretationof its discovery by Ruysch, Jacobson, or Kölliker, with an English translation of Kölliker (1877). Anatomical Record, 270B:4-15.in PDF format

Smith, TD, KP Bhatnagar, CJ Bonar, KL Shimp, MP Mooney, and MI Siegel. 2003. Ontogenetic characteristics of the vomeronasal organ in Saguinus geoffroyi and Leontopithecus rosalia with comparisons to other primates. American Journal of Physical Anthropology, 121:342-353. in PDF format (and please note Erratum, regarding the incorrect date of publication printed on p. 342)

Shimp, KL, KP Bhatnagar, C Bonar, TD Smith. 2003. Ontogeny of the nasopalatine duct in primates. Anatomical Record, 274A: 862-869.in PDF format

Smith TD, MI Siegel, KP Bhatnagar. 2003. Observations on the vomeronasal organ of prenatal Tarsius bancanus borneanus with implications for ancestral characteristics. Journal of Anatomy, 203:473-481.in PDF format*

Maico, LM, AM Burrows, MP Mooney, MI Siegel, KP Bhatnagar, and TD Smith. 2003. Size of the vomeronasal organ in wild Microtus with different mating strategies. Acta Biologica Hungarica, 54:263-273. in PDF format

Smith, T.D., K.P. Bhatnagar, K.L. Shimp, J.H. Kinzinger, C.J. Bonar, A.M. Burrows, M.P. Mooney, and M.I. Siegel. 2002. Histological definition of the vomeronasal organ in humans and chimpanzees with a comparison to other primates. Anatomical Record, 267:166-176. in PDF format

Smith, T.D., M.I. Siegel, C.J. Bonar, K.P. Bhatnagar, M.P. Mooney, A.M. Burrows, M.A. Smith, and L.M. Maico. 2001. The existence of the vomeronasal organ in postnatal chimpanzees and evidence for its homology to that of humans. Journal of Anatomy, 198:77-82. in PDF format

Smith, T.D., D.L. Roslinski, A.M. Burrows, K.P. Bhatnagar, M.P. Mooney, and M.I. Siegel. 2001. Size of the vomeronasal neuroepithelium in two species of Microtus with differing levels of paternal behavior. Journal of Mammalogy, 82:209-217. in PDF format

Smith, T.D., M.P. Mooney, A.M. Burrows, K.P. Bhatnagar, and M.I. Siegel. 2001. Prenatal Growth and Adult Size of the Vomeronasal Organ in Mouse Lemurs and Humans. Pp. 93-99 In: A.. Marchlewska et al. (eds.), Chemical Signals in Vertebrates IX, Kluwer Academic/ Plenum Press, New York.

Smith, T.D., M.I. Siegel, and K.P. Bhatnagar. 2001. A reappraisal of ontogeny, morphology, functionality, and persisting questions on the vomeronasal system of catarrhine primates. Anatomical Record (New Anatomist), 265:176-192. in PDF format

Bhatnagar, K.P., and T.D. Smith. 2001. The human vomeronasal organ - Part III: postnatal development from infancy through the ninth decade. Journal of Anatomy, 199:289-302. in PDF format

Smith, T.D., T.A. Buttery, K.P. Bhatnagar, A.M. Burrows, M.P. Mooney, and M.I. Siegel. 2001. Anatomical position of the vomeronasal organ in postnatal humans. Annals of Anatomy, 183:475-479.

Buttery, T.A., T.D. Smith, A.M. Burrows, M.P. Mooney, M.I. Siegel, and A.R. Burdi. 2000. Postnatal presence of the paraseptal cartilages in humans: A description of morphology and size. Acta Oto-Laryngologica, 120:77-80.

Roslinski, D.L., K.P. Bhatnagar, A.M. Burrows, and T.D. Smith. 2000. Comparative morphology and histochemistry of glands associated with the vomeronasal organ in humans, mouse lemurs, and voles. Anatomical Record, 260:92-101. in PDF format

Smith, T.D. and K.P. Bhatnagar. 2000. The human vomeronasal organ. Part II: prenatal development. Journal of Anatomy, 197:421-436. in PDF format

Pokorny, M.J., T.D. Smith, S.A. Calus, and E.A. Dennison. 2000. Oral contraceptive use appears to have no effect on peripheral joint laxity. Journal of Orthopaedic & Sports Physical Therapy, 30:683-692.

Smith, T.D., M.I. Siegel, A.M. Burrows, M.P. Mooney, A.R. Burdi, P.A. Fabrizio, and F.R. Clemente. 1998. Searching for the vomeronasal organ of adult humans: Preliminary findings on location, structure, and size. Microscopy Research and Technique, 41:483-491.

Smith, T.D., M.I. Siegel, M.P. Mooney, A.M. Burrows, and J.S. Todhunter. 1997. Formation and enlargement of the paranasal sinuses in normal and cleft lip and palate human fetuses. Cleft Palate-Craniofacial Journal, 34:483-489. in PDF format

Smith, T.D., M.I. Siegel, M.P. Mooney, A.R. Burdi, A.M. Burrows, and J.S. Todhunter. 1997. Prenatal growth of the human vomeronasal organ. Anatomical Record, 248:447-455. in PDF format

Smith, T.D., M.I. Siegel, M.P. Mooney, A.R. Burdi, and J.S. Todhunter. 1996. Vomeronasal organ development in normal and cleft lip and palate fetuses. Cleft Palate Craniofacial Journal, 33:385-394. in PDF format

SRU students indicated in red.

*This is an electronic version of an article published in The Journal of Anatomy: complete citation information for the final version of the paper, as published in the print edition of The Journal of Anatomy, is available on the Blackwell Synergy online delivery service, accessible via the journal's website at http://www.blackwellpublishing.com/journals/joa or http://www.blackwell-synergy.com.