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Publications of Bruce A. Young, PhD



Vedurmudi AP, Young BA, van Hemmen JL. Internally coupled ears: mathematical structures and mechanisms underlying ICE. Biol Cybern. 2016;110(4-5):359-382. PubMed
Nasoori A, Shahbazzadeh D, Tsubota T, Young BA. The defensive behaviour of Naja oxiana,with comments on the visual displays of cobras. Herpetological Bulletin. 2016; 138: 13-17. Link
Jackson TNW, Young B, Underwood G, McCarthy CJ, Kochva E, Vidal N, van der Weerd L, Nabuurs R, Dobson J, Whitehead D, Vonk FJ, Hendrikx I, Hay C, Fry BG. (2016) Endless forms most beautiful: the evolution of ophidian oral glands, including the venom system, and the use of appropriate terminology for homologous structures. Zoomorphology, 1-24; DOI 10.1007/s00435-016-0332-9. Link

Young BA. Anatomical influences on internally coupled ears in reptiles. Biol Cybern. 2016;110(4-5):255-261. PubMed

Young BA, Dumais J, John N, Lyons B, Macduff A, Most M, Reiser NA, Reiser PJ. Functional segregation within the muscles of aquatic propulsion in the Asiatic Water Monitor Lizard (Varanus salvator).Front Physiol. 2016;7:380 PubMed
Han D, Young BA. Anatomical Basis of Dynamic Modulation of Tympanic Tension in the Water Monitor Lizard, Varanus salvator. Anat Rec (Hoboken). 2016 Jun 17;299(9):1270-1280. doi: 10.1002/ar.23382. PubMed
Vedurmudi AP, Goulet J, Christensen-Dalsgaard J, Young BA, Williams R, van Hemmen JL. How Internally Coupled Ears Generate Temporal and Amplitude Cues for Sound Localization. Phys Rev Lett. 2016 Jan 15;116(2):028101. PubMed
Bierman HS, Thornton JL, Jones HG, Koka K, Young BA, Brandt C, Christensen-Dalsgaard J, Carr CE, Tollin DJ. Biophysics of directional hearing in the American alligator (Alligator mississippiensis). J Exp Biol. 2014 Apr 1;217(Pt 7):1094-107. doi: 10.1242/jeb.092866. PubMed
Most MG, Grace KM, Young BA. Terrestrial locomotion in the marine filesnake, Acrochordus granulatus (Schneider, 1799). Hamadryad 2014 36(2):142-8.
Kohl T, Colayori SE, Westhoff G, Bakken GS, Young BA. Directional sensitivity in the thermal response of the facial pit in western diamondback rattlesnakes (Crotalus atrox). J Exp Biol. 2012 Aug 1;215(Pt 15):2630-6. doi:10.1242/jeb.065896. PubMed
Fry BG, Casewell NR, Wüster W, Vidal N, Young B, Jackson TN. The structural and functional diversification of the Toxicofera reptile venom system. Toxicon. 2012 Sep 15;60(4):434-48. doi: 10.1016/j.toxicon.2012.02.013. Epub 2012 Mar 14. PubMed
Burnell A, Collins S, Young BA. On the postpulmonary septum of Varanus and it’s implications for Mosasaurine ventilation and physiology. 2012 Bulletin of the Geological Society of France. 183: 159-169.
Burnell A, Collins S, Young BA. Vertebral morphometrics in Varanus. Bulletin of the Geological Society of France. 2012 183: 151-158.
Young BA, Herzog F, Friedel P, Rammensee S, Bausch A, van Hemmen JL. Tears of venom: hydrodynamics of reptilian envenomation. Phys Rev Lett. 2011 May 13;106(19):198103. Epub 2011 May 12. PubMed
Lindgren J, Polcyn M, Young BA. Landlubbers to leviathans: modular evolution and swimming in mosasaurine mosasaurs. Paleobiology 2010 37: 445-469.

Westhoff G, Boetig M, Bleckmann H, Young BA. Target tracking during venom 'spitting' by cobras. J Exp Biol. 2010 Jun 1;213(11):1797-802. doi:10.1242/jeb.037135. PubMed

Young BA, Kardong KV. The functional morphology of hooding in cobras. J Exp Biol. 2010 May;213(Pt 9):1521-8. doi: 10.1242/jeb.034447. PubMed
Young BA. How a heavy-bodied snake strikes quickly: high-power axial musculature in the puff adder (Bitis arietans). J Exp Zool A Ecol Genet Physiol. 2010 Feb 1;313(2):114-21. doi: 10.1002/jez.579. PubMed
Young,BA, Boetig M, Westhoff G. Spitting behaviour of hatchling red spitting cobras (Naja pallida). Herpetological Journal 2009 19: 185-191.
Young BA, Boetig M, Westhoff G. Functional bases of the spatial dispersal of venom during cobra "spitting". Physiol Biochem Zool. 2009 Jan-Feb;82(1):80-9.doi: 10.1086/595589. PubMed
Young BA, Boetig M, Fahey A, Lawrence A. The diversity of aquatic locomotion in extant varanoid lizards. Proc. 2008 Fort Hayes St. Univ. 3: 159-167.
Friedel P, Young BA, van Hemmen JL. Auditory localization of ground-borne vibrations in snakes. Phys Rev Lett. 2008 Feb 1;100(4):048701. Epub 2008 Jan 28. PubMed
Fry BG, Scheib H, van der Weerd L, Young B, McNaughtan J, Ramjan SF, Vidal N, Poelmann RE, Norman JA. Evolution of an arsenal: structural and functional diversification of the venom system in the advanced snakes (Caenophidia). Mol Cell Proteomics. 2008 Feb;7(2):215-46. Epub 2007 Sep 12. PubMed
Parker MR, Young BA, Kardong KV. The forked tongue and edge detection in snakes (Crotalus oreganus): an experimental test. J Comp Psychol. 2008 Feb;122(1):35-40. doi: 10.1037/0735-7036.122.1.35. PubMed
Young BA, Aguiar A, Lillywhite H. Foraging cues used by Cottonmouth snakes (Agkistrodon piscivorus conanti) on Sea Horse Key, Florida. South Amer. J. Herpetol. 2008 3(2): 135-144.
Young BA, Kardong KV. Mechanisms controlling venom expulsion in the western diamondback rattlesnake, Crotalus atrox. J Exp Zool A Ecol Genet Physiol. 2007 Jan 1;307(1):18-27. PubMed
Young BA, Harris J. Auditory response of the Northern Pacific rattlesnake (Crotalus viridis oreganus): Does behavior match physiology? Northwest Sci. 2006 80:218-233.
Young BA. Snake sounds. Nature Austr. 2005 28(4): 34-41.

Young BA, O’Shea M. Analyses of venom spitting in African cobras (Elapidae: Serpentes). Afr. Zool. 2005 40(1): 71-76.

Young BA, Dunlap K, Koenig K, Singer M. The buccal buckle: the functional morphology of venom spitting in cobras. J Exp Biol. 2004 Sep;207(Pt 20):3483-94. PubMed
Grande T, Young BA. The ontogeny of the Weberian apparatus in the zebrafish Danio rerio (Ostariophysi: Cypriniformes) with comments on the homology of otophysan Weberian ossicles. Zool. J. Linn. Soc. 2004 140: 241-254.
Hartdegen RW, Russell MJ, Young BA, Reams RD. Krokotiiliskinkin Tribolonotus gracilis (DE ROOY, 1909) ääntely ja todisteita poikasten hoidosta. Herpetomania 2003 12(4): 5-9.
Young BA. Snake bioacoustics: toward a richer understanding of the behavioral ecology of snakes. Q Rev Biol. 2003 Sep;78(3):303-25. Review. PubMed
Young BA, Morain M. The mechanics of vertical burrowing in the Saharan sand vipers (Cerastes). Copeia 2003: 132-138.
Young, BA, Phelan M, Morain M, Ommundsen M, Kurt R. Venom injection in rattlesnakes (Crotalus): Peripheral resistance and the pressure-balance hypothesis. Can. J. Zool. 2003 81: 313-320.
Young BA, Aguiar A. Response of western diamondback rattlesnakes Crotalus atrox to airborne sounds. J Exp Biol. 2002 Oct;205(Pt 19):3087-92. PubMed
Young BA, Lee CE, Daley KM. Do snakes meter venom? Biosciences 2002 52:1121-1126.
Young BA, Lee CE, Daley KM. On a flap and a foot: Aerial locomotion in the "Flying" gecko, Ptychozoon kuhli. J. Herpetol. 2002 36(3): 412-418.
Young BA, Morain M. The use of ground-borne vibrations for prey localization in the Saharan sand vipers (Cerastes). J Exp Biol. 2002 Mar;205(Pt 5):661-5. PubMed
Berkhoudt H, Wilson P, Young BA. Taste buds in the palatal mucosa of snakes. Afr. Zool. 2001 36(2): 185-188.
Hartdegen RW, Russell MJ, Young BA, Reams RD. Vocalization of the Crocodile skink, Tribolonotus gracilis, and evidence of parental care. Cont. Herpetol. 2001(on-line publication).
Young BA, Blair M, Zahn K, Marvin J. Mechanics of venom expulsion in Crotalus,with special reference to the role of the fang sheath. Anat Rec. 2001 Dec 1;264(4):415-26. PubMed
Young BA, Jaggers J, Nejman N, Kley N. Defensive buccal expansion in Bitis arietans. Copeia 2001(1): 270-273.
Young BA, Phelan M, Jaggers J, Nejman N. Kinematic modulation of the strike of the western diamondback rattlesnakes (Crotalus atrox). Hamadryad 2001 26(2): 316-349.
Young BA, Zahn K. Venom flow in rattlesnakes: mechanics and metering. J Exp Biol. 2001 Dec;204(Pt 24):4345-51. PubMed
Young BA. The comparative morphology of the ophidian larynx. Acta Zool. 2000 81:177-193.
Young BA, Marvin J, Marosi K. The potential significance of groundborne vibration to predator-prey relationships in snakes. Hamadryad 2000 25(2): 164-174
Young BA, Nejman N, Jaggers J. Snakes that inflate: The functional morphology of a defensive behavior. Zoology 2000 103: 25-30.
Young BA, Zahn K, Blair M, Lalor J. Functional subdivision of the venom gland musculature and the regulation of venom expulsion in rattlesnakes. J Morphol. 2000 Dec;246(3):249-59. PubMed
Cascardi J, Young BA, Husic HD, Sherma J. Protein variation in the venom spat by the red spitting cobra, Naja pallida (Reptilia: Serpentes). Toxicon. 1999 Sep;37(9):1271-9. PubMed
Young BA. The comparative morphology of the mandibular midline raphe in snakes (Reptilia: Squamata). Zool. Anz. 1999 237: 217-241.
Young BA, Lalor J, Solomon J. The comparative biomechanics of an ophidian defensive behavior: Head triangulation in hognose snake (Heterodon) and egg-eating snake (Dasypeltis). J. Zool. Lond. 1999 248: 169-177.
Young BA, Marsit C, Meltzer K. Comparative morphology of the cloacal scent gland in snakes (Serpentes: Reptilia). Anat Rec. 1999 Oct 1;256(2):127-38. PubMed
Young BA, Meltzer K, Marsit C. Scratching the surface of mimicry: Sound production through scale abrasion in snakes. Hamadryad 1999 24: 29-38.
Young BA, Meltzer K, Marsit C, Abishahin G. Cloacal popping in snakes. J. Herpetol. 1999 33: 557-566.
Young BA, Nejman N, Meltzer K, Marvin J. The mechanics of sound production in the puff adder bitis arietans (Serpentes: viperidae) and the information content of the snake hiss. J Exp Biol. 1999 Sep;202 (Pt 17):2281-9. PubMed
Young BA, Saunders M. Direct visualization of blood flow through the interaortic foramen of the eastern diamondback rattlesnake, Crotalus adamanteus, using echocardiography and color Doppler imaging. J Exp Zool. 1999 Dec 1;284(7):742-745. PubMed
Young BA, Solomon J, Abishahin G. How many ways can a snake growl? The morphology of sound production in Ptyas mucosus and its potential mimicry of Ophiophagus. Herpetol. J. 1999 9(3): 89-94.
Kinney C, Abishahin G, Young BA. Hissing in rattlesnakes: Redundant signaling or inflationary epiphenomenon? J. Exp. Zool. 1998 280: 107-113.
Young BA. How to hiss loudly: Narial flaring and acoustic amplification in Russell’s viper (Daboia russelii). Hamadryad 1998 23(2): 103-110.
Young BA. On the acoustic biology of snakes. [Invited paper] Lit. Serpent. 1998 18(4): 84-91.
Young BA. The comparative morphology of the intermandibular connective tissue in snakes (Reptilia: Squamata). Zool. Anz. 1998 237: 59-84.
Young BA, Abishahin G, Bruther M, Kinney C, Sgroi J. Acoustic analysis of the defensive sounds of Varanus salvator with notes on sound production in other varanid species. Hamadryad 1998 23: 1-14.
Young B, Frazer B, Fried B, Lee M, Lalor J, Sherma J. HPTLC determination of cloacal scent gland lipids from two sympatric snakes: The eastern diamondback rattlesnake (Crotalus adamanteus) and the Florida cottonmouth (Agkistrodon piscivorus conanti). J.P. C. 1998 12: 196-201.
Young B, Lalor J. Sound production in the eastern hognose snake, Heterodon platyrhinos (Serpentes: Colubridae): Does it snore? Amph.-Rept. 1998 19:407-418.
Young BA, Wallach V. Description of a papillate tactile organ in the Typhlopidae. S. Afr. J. Zool. 1998 33: 249-253.
Grande T, Young B. Morphological development of the opercular apparatus in Kneria wittei (Ostariophysi: Gonorynchiformes) with comments on its possible function. Acta Zool. 1997 78: 145-162.
Young BA. A review of sound production and hearing in snakes, with a discussion of intraspecific acoustic communication in snakes. J. Penn. Acad. Sci. 1997 71: 39-46.
Young BA. On the absence of taste buds in monitor lizards (Varanus) and snakes. J. Herpetol. 1997 31: 130-137.
Young BA, Wassersug RJ, Pinder A. Gravitational gradients and blood flow patterns in specialized arboreal (Ahaetulla nasuta) and terrestrial (Crotalus adamanteus) snakes. J Comp Physiol B. 1997;167:481-93. PubMed
Young BA, Kardong KV. Dentitional surface features in snakes (Lepidosauria: Serpentes). Amph.-Rept. 1996 17: 261-276.
Rasmussen S, Young BA, and Krimm H. On the “spitting” behavior in cobras. J. Zool. Lond. 1995 237(1): 27-36.
Young BA, Brown IP. The physical basis of the rattling sound in the rattlesnake, Crotalus viridis oreganus. J. Herpetol. 1995 29(1): 80-85.
Young BA, Sheft S, Yost W. Sound production in Pituophis melanoleucus (Serpentes: Colubridae) with the first description of a vocal cord in snakes. J Exp Zool. 1995 Dec 15;273(6):472-81. PubMed
Young BA. Cartilago cordis in serpents. Anat Rec. 1994 Oct;240(2):243-7. PubMed
Young BA, Street SL, Wassersug RJ. Anatomical and gravitational influences on cardiac displacement in snakes (Lepidosauria, Serpentes). Zoomorphology.1994;114:169-75. PubMed
Young BA, Brown IP. On the acoustic profile of the rattlesnake rattle. Amph.-Rept. 1993 14: 373-380.
Young BA, Lillywhite HB, Wassersug RJ. On the structure of the aortic valves in snakes (Reptilia: Serpentes). J. Morphol. 1993 216(2): 141-160.
Young BA. On the necessity of an archetypal concept in morphology: With special reference to the concepts of “structure” and “homology”. Biol. and Phil. 1992 8:225-248.
Young BA. Tracheal diverticula in snakes: Possible functions and evolutionary origins. J. Zool. Lond. 1992 227: 567-583.
Young BA. Evaluating hypotheses for the transfer of stimulus particles to Jacobson’s organ in snakes. [invited paper] Brain Behav. and Evol. 1991 41: 203-209.
Young BA. Morphological basis of "growling" in the king cobra, Ophiophagus hannah. J Exp Zool. 1991 Dec;260(3):275-87. PubMed
Young BA. Is there a direct link between the ophidian tongue and Jacobson’s organ? Amph.-Rept. 1990 11: 263-276.
Young BA. The influences of the aquatic medium on the prey capture system of snakes. J. Nat. Hist. 1990 25: 519-531.
Young BA, Magon DK, Goslow GE. Length-tension and histochemical properties of selected shoulder muscles of the savannah monitor lizard (Varanus exanthematicus): Implications for function and evolution. J. Exp. Zool. 1990 256: 63-74.
Young BA. Ontogenetic changes in the feeding system of the red-sided garter snake, Thamnophis sirtalis parietalis. I. Allometric analysis. J. Zool. Lond. 1989 218(3): 365-382.
Young BA. The anatomy of the head of the red-sided garter snake, Thamnophis sirtalis parietalis. Part I - Osteology. Zool. Jb. Anat. 1989 118: 251-272.
Young BA. The anatomy of the head of the red-sided garter snake, Thamnophis sirtalis parietalis. Part II - Myology. Zool. Jb. Anat. 1989 118: 325-354.
Young BA. Ashley MA. A mathematical model of relative muscle force with a perturbation analysis of selected muscle parameters. J. Theor. Biol. 1989 138:213-233.
Young BA. A model for the evolution of functional-morphological systems. Evol. Theory. 1988  8(3): 125-150.
Young BA. The arthrology of the head of the red-sided garter snake, Thamnophis sirtalis parietalis. Neth. J. Zool. 1988 38(2-4): 166-205.
Young BA. The cephalic vascular anatomy of three species of sea snakes. J Morphol. 1988 May;196(2):195-204. PubMed
Young BA. The subclavian loop of Varanus salvator. Copeia. 1988(4): 1029-1034.
Young BA. The cephalic vascular anatomy of three species of sea snakes. J. Morphol. 1988 196:195-204.
Young BA. The comparative cephalic osteology and myology of three species of sea snakes. Zool. Jb. Anat. 1987 116: 421-452.
Young BA. The cranial nerves of three species of sea snakes. Can J. Zool. 1987 65(9):2236-2240.
Books, Book Chapters, Reviews Return to top
9. Young BA. 2015. The evolution of the snake ear and its restricted frequency response range. In: All animals are interesting, A Festschrift in honour of Anthony P. Russell. (O.R.P. Bininda-Edwards, G. Powell, H. Jamniczky, and A. Bauer, eds.) Universitat Oldensburg Press. pp. 423-438.
8. Fry B, Sunagar K, Casewell N, Kochva E, Scheib H, Wüster W, Vidal N, Young B, Burbrink F, Pyron A, Vonk F, Jackson, T. 2015. The origin and evolution of the Toxicofera reptile venom system.In B. G. Fry (Ed.). Venomous Reptiles and Their Toxins. Evolution, Pathophysiology and Biodiscovery. Oxford University Press , 2015. p. 1-576.
Young BA, Mathevon N, Tang Y. 2014. Reptile auditory neuroethology: What do reptiles do with their hearing. In: Insights from Comparative Hearing Research, Springer Handbook for Auditory Research (R. Fay, C. Koppel, G. Manley, A. Popper, eds.) Springer, New York. 49: 323 – 346.
Young BA. 2010. Vibration detection in snakes. Pp. 85 – 98 In: The Use of Vibrations in Communication: properties, mechanisms and function across taxa. (O’Connell-Rodwell C.E., ed.) Research Signpost.
Young BA. 2008. Perspectives on the regulation of venom expulsion in snakes. Pp. 181 – 190 In: The Biology of Rattlesnakes, W. Hayes, M. Cardwell, K. Beaman, and S. Bush, eds., Loma Linda University Press.
Young BA, Jackson K. 2008. Functional specializations of the extrinsic venom gland musculature in crotaline snakes: What is the role of the M. Pterygoideus Glandulae?;” Pp. 47 – 54 In: The Biology of Rattlesnakes, W. Hayes, M.Cardwell, K. Beaman, and S. Bush, eds., Loma Linda University Press.
Young BA. 2007. Response of the yellow anaconda (Eunectes notaeus) to underwater pressure waves. Pp. 199 – 206 In: Biology of Boas and Pythons, R. Henderson,Ed., Eagle Mountain Press.
Young BA. 2006. Auditory atavism and integrated pathways for hearing in snakes. Pp. 229 – 232 in: Herpetological Bonnensis II (M. Vences, J. Köhler, T. Ziegler, W. Böhme, eds.) European Society of Herpetology, Bonn.
Young BA. 1997. Hearing, taste, tactile reception, and olfaction. Pp. 195 – 223 In: The Biology, Husbandry, and Health Care of Reptiles, Vol. 1. (Ackerman, L., ed.), T.F.H. Publications, New Jersey.
Poster Presentations/Published Abstracts Return to top


Han DYoung BA. Biophysics of the Varanid Tympanic Membrane. Poster presented at the 8th Annual Interdisciplinary Biomedical Research Conference; Kirksville, MO; November 5, 2016.


Nasoori A, Young BA, Tsubata T, Shahbazzadeh D. The defensive behavior of Naja oxiana with comments on the visual displays of cobras. Poster presented at the 3rd International Veterinary Congress, London, August 18-20, 2016.


Young BA. Active Tuning Mechanisms in the Reptile Ear. Invited oral presentation at the Symposium on Biomimetics of Sensory System and Locomotion; The Second Sino-German Symposium on Biomimetics: From Animal Sensory Systems to Locomotion; Garching; Germany; June 29, 2015.


Young BA, Most M, Dumais J, John N, Lyons B, MaCduff A, Reiser P. Multiple Perspectives of the Functional Divisions within Swimming Muscles of the Asiatic Water Monitor (Varanus salvator). Oral presentation at the Society for Integrative and Comparative Biology; Austin, TX; Jan. 3-7, 2014.


Young BA. Building a Better Robotic Snake through the Biophysics of Snake Locomotion. Invited oral presentation at the Sino-German Symposium on Biomimetics: from Animal Systems to Locomotion; The International Society of Bionic Engineering; October 10-15, 2014.


Young BA. Dynamic Modulation of the Middle Ear Cavity in the Water Monitor (Varanus salvator). Invited oral presentation at the International Symposium on Internally Coupled Ears: Evolutionary Origins, Mechanisms, and Neuronal Processing from a Biomimetic Perspective; Technical University of Munich; June 18, 2014.


Young BA. Paleo-Prosthetics, A Surgical Guide to Reconstructing a Mosasaur. Poster presented at 5th Annual Interdisciplinary Biomedical Research Conference; Kirksville, MO; November 2, 2013.