The Cochlear Labyrinth of Krapina Neandertals

........................................................................................................iii Acknowledgements...........................................................................................iv Chapter I: Overview of Anatomy and Physiology of the Ear..........................................1 The external ear.......................................................................................1 The middle ear.........................................................................................2 The inner ear...........................................................................................3 The vestibular system ......................................................................4 The cochlea...................................................................................4 Cochlear mechanics and physiology of sound transmission....................................6 Chapter II: Comparative Hearing among Mammals....................................................9 Basic measures ........................................................................................9 Mammalian hearing sensitivity and frequency processing ....................................11 Hearing sensitivity in primates....................................................................11 Generalized vs. specialized cochleae..............................................................12 Predictions from gross cochlear morphology...................................................14 Chapter III: Evolution of Hearing.........................................................................17 Fossil evidence: primates...........................................................................17 Fossil evidence: hominids..........................................................................18 Auditory structures in Neandertals...............................................................20 Chapter IV: The Krapina Site..............................................................................23 Discovery and excavation of Krapina............................................................23 Stratigraphy and chronology......................................................................25 vi Chapter V: The Current Study............................................................................28 Materials and methods..............................................................................29 Specimens and CT scanning..............................................................29 Delimitation of the cochlea................................................................32 Segmentation, thresholding, and 3D reconstruction...................................34 Error and validation........................................................................37 Statistical analyses.........................................................................37 Chapter VI: Results..........................................................................................40 Qualitative analysis..................................................................................40 Quantitative Analysis...............................................................................42 Descriptive statistics.......................................................................42 Nonparametric analyses....................................................................43 Absolute cochlear volume........................................................43 Scaled cochlear volume...........................................................44 Linear regression...........................................................................46 Chapter VII: Discussion....................................................................................48 Potential mechanisms for the relationship between cochlear volume and hearing ability.........................................49 Implications for ecology and communication of Neandertals................................50 Future directions for the Krapina scans ..........................................................52 Summary.............................................................................................53 Bibliography........................................................................................54 Appendices....................................................................................................59

[1]  J. Moggi-Cecchi,et al.  The mixed dentition and associated skull fragments of a juvenile fossil hominid from Sterkfontein, South Africa. , 1998, American journal of physical anthropology.

[2]  E. Carbonell,et al.  Auditory capacities in Middle Pleistocene humans from the Sierra de Atapuerca in Spain. , 2004, Proceedings of the National Academy of Sciences of the United States of America.

[3]  Jean-Jacques Hublin,et al.  The bony labyrinth of Neanderthals. , 2003, Journal of human evolution.

[4]  F. Spoor,et al.  Comparative review of the human bony labyrinth. , 1998, American journal of physical anthropology.

[5]  J. J. Flynn,et al.  The bony labyrinth of the early platyrrhine primate Chilecebus. , 2010, Journal of human evolution.

[6]  Richard R. Fay,et al.  Comparative Hearing: Mammals , 1994, Springer Handbook of Auditory Research.

[7]  M. Wolpoff.  The Krapina dental remains. , 1978, American journal of physical anthropology.

[8]  J. H. Elder.  Auditory acuity of the chimpanzee. , 1934 .

[9]  M. Collard,et al.  A fossil stapes from Sterkfontein, South Africa, and the hearing capabilities of early hominids. , 2002, Journal of human evolution.

[10]  C. Snowdon,et al.  Environmental correlates of vocal communication of wild pygmy marmosets, Cebuella pygmaea , 2002, Animal Behaviour.

[11]  R. J. Smith,et al.  Body mass in comparative primatology. , 1997, Journal of human evolution.

[12]  C. Ruff,et al.  Body mass and encephalization in Pleistocene Homo , 1997, Nature.

[13]  Stanley A. Gelfand,et al.  Hearing: An Introduction to Psychological and Physiological Acoustics, Fourth Edition , 1998 .

[14]  H. Uylings,et al.  Bivariate linear models in neurobiology: problems of concept and methodology , 1986, Journal of Neuroscience Methods.

[15]  F. Spoor,et al.  Prenatal growth and development of the modern human labyrinth , 2004, Journal of anatomy.

[16]  K. Gorjanović-Kramberger.  Der diluviale Mensch von Krapina in Kroatien : ein Beitrag zur Paläoanthropologie , 1906 .

[17]  E. Carbonell,et al.  Three new human skulls from the Sima de los Huesos Middle Pleistocene site in Sierra de Atapuerca, Spain , 1993, Nature.

[18]  A. Popper,et al.  The Evolutionary biology of hearing , 1992 .

[19]  P. Iiyambo,et al.  Statistics for life sciences I , 2012 .

[20]  Charles H. Brown,et al.  Habitat acoustics and primate communication , 1986, American journal of primatology.

[21]  J. Radovčić,et al.  New reconstruction of Krapina 5, a male Neandertal cranial vault from Krapina, Croatia. , 2006, American journal of physical anthropology.

[22]  Richard R. Fay,et al.  Structure and Function in Sound Discrimination Among Vertebrates , 1992 .

[23]  Charles H. Brown,et al.  Old world monkey vocalizations: adaptation to the local habitat? , 1995, Animal Behaviour.

[24]  Y. Rak,et al.  Ear ossicle of Australopithecus robustus , 1979, Nature.

[25]  R. Fay.  Comparative Auditory Research , 1994 .

[26]  D. D. Greenwood.  A cochlear frequency-position function for several species--29 years later. , 1990, The Journal of the Acoustical Society of America.

[27]  Gunnar Fant,et al.  Speech sounds and features , 1973 .

[28]  Mark N. Coleman.  What Do Primates Hear? A Meta-analysis of All Known Nonhuman Primate Behavioral Audiograms , 2009, International Journal of Primatology.

[29]  M. Colbert,et al.  Auditory Morphology and Hearing Sensitivity in Fossil New World Monkeys , 2010, Anatomical record.

[30]  R. Heffner.  Primate hearing from a mammalian perspective. , 2004, The anatomical record. Part A, Discoveries in molecular, cellular, and evolutionary biology.

[31]  C D West,et al.  The relationship of the spiral turns of the cochlea and the length of the basilar membrane to the range of audible frequencies in ground dwelling mammals. , 1985, The Journal of the Acoustical Society of America.

[32]  J. Mitani,et al.  Geographic variation in the calls of wild chimpanzees: A reassessment , 1999, American journal of primatology.

[33]  G. Békésy,et al.  Experiments in Hearing , 1963 .

[34]  E. Kirk,et al.  Cochlear Labyrinth Volume and Hearing Abilities in Primates , 2009, Anatomical record.

[35]  B Masterton,et al.  The evolution of human hearing. , 1969, The Journal of the Acoustical Society of America.

[36]  M. Colbert,et al.  Correlations between auditory structures and hearing sensitivity in non‐human primates , 2009, Journal of morphology.

[37]  Mark N. Coleman.  Functional Morphology and Evolution of the Primate Auditory System , 2007 .

[38]  James D. Miller.  Sex differences in the length of the organ of Corti in humans. , 2007, The Journal of the Acoustical Society of America.

[39]  Richard J. Smith.  Use and misuse of the reduced major axis for line-fitting. , 2009, American journal of physical anthropology.

[40]  N. Dominy,et al.  Primate communication in the pure ultrasound , 2012, Biology Letters.

[41]  John J. Rosowski,et al.  Outer and Middle Ears , 1994 .

[42]  F. Smith.  The Neandertal remains from Krapina : a descriptive and comparative study , 1976 .

[43]  M. Westoby,et al.  Bivariate line‐fitting methods for allometry , 2006, Biological reviews of the Cambridge Philosophical Society.

[44]  D. D. Greenwood.  Critical Bandwidth and the Frequency Coordinates of the Basilar Membrane , 1961 .

[45]  W. T. Peake,et al.  Experiments in Hearing , 1963 .

[46]  F. Spoor,et al.  Semicircular canal system in early primates. , 2009, Journal of human evolution.

[47]  R. Fay,et al.  Structure of the Mammalian Cochlea , 1994 .

[48]  R. Quam,et al.  Auditory ossicles from southwest Asian Mousterian sites. , 2008, Journal of human evolution.

[49]  W. Howells,et al.  The Fate of the "Classic" Neanderthals: A Consideration of Hominid Catastrophism [and Comments and Reply] , 1964, Current Anthropology.

[50]  C. Niemitz.  Chapter 14 – Outline of the Behavior of Tarsius bancanus , 1979 .

[51]  Daphne Manoussaki,et al.  The influence of cochlear shape on low-frequency hearing , 2008, Proceedings of the National Academy of Sciences.

[52]  Timothy M. Ryan,et al.  The primate semicircular canal system and locomotion , 2007, Proceedings of the National Academy of Sciences.

[53]  H. Schwarcz,et al.  ESR ages for Krapina hominids , 1995, Nature.

[54]  M. Wolpoff,et al.  The Krapina hominids : an illustrated catalog of skeletal collection , 1988 .

[55]  M. Silcox,et al.  Cochlear Labyrinth Volume in Euarchontoglirans: Implications for the Evolution of Hearing in Primates , 2011, Anatomical record.

[56]  Richard J. Smith.  Regression models for prediction equations , 1994 .

[57]  F Zonneveld,et al.  Morphometry of the primate bony labyrinth: a new method based on high-resolution computed tomography. , 1995, Journal of anatomy.