Abstract
Source–filter coupling is the rarest acoustic phenomenon not only in Iberian red deer, but in any mammal. In most mammals, sound production can be well described in the framework of source–filter theory. The vocal output is the result of combined work of the larynx (the source) and of the supralaryngeal vocal tract (the filter). The source–filter theory suggests the independence of source and filter. Thus, vocal tract filtering should not affect the fundamental frequency (f0) of the sound created in the larynx. Spectrographically, the source is mostly characterized by the f0 and its harmonics, while the filter by the vocal tract resonances, i.e., formant frequencies. Nevertheless, a non-independent (coupled) source and filter can be proposed when the vocal folds start oscillating at one of the formant frequencies. Coupling between source and filter has been found in human singers and predicted for red deer Cervus elaphus by a computer modeling approach. This study describes different modes of phonation in a natural bout of rutting calls of Iberian red deer Cervus elaphus hispanicus and the transition from a chaotic mode to a probable source–filter coupling mode. This phenomenon might be involved in the production of extremely high-frequency bugles of North American and Asian subspecies of C. elaphus.
References
Boersma P, Weenink D (2009) Praat: Doing Phonetics by Computer. http://www.praat.org
Efremova KO, Volodin IA, Volodina EV, Frey R, Lapshina EN, Soldatova NV (2011) Developmental changes of nasal and oral calls in the goitred gazelle Gazella subgutturosa, a nonhuman mammal with a sexually dimorphic and descended larynx. Naturwissenschaften 98:919–931
Fant G (1960) Acoustic theory of speech production. Mouton & Co, Hague
Feighny JJ, Williamson KE, Clarke JA (2006) North American elk bugle vocalizations: male and female bugle call structure and context. J Mammal 87:1072–1077
Fitch WT, Hauser MD (2002) Unpacking “honesty”: vertebrate vocal production and the evolution of acoustic signals. In A. Simmons A, Fay RR, Popper AN (eds) Acoustic communication, Springer handbook of auditory research. Springer, Berlin, pp 65–137
Fitch WT, Reby D (2001) The descended larynx is not uniquely human. Proc Royal Soc B 268:1669–1675
Frey R, Volodin I, Volodina E, Carranza J, Torres-Porras J (2012) Vocal anatomy, tongue protrusion behaviour and the acoustics of rutting roars in free-ranging Iberian red deer stags (Cervus elaphus hispanicus). J Anat 220:271–292
Hatzikirou H, Fitch WT, Herzel H (2006) Voice instabilities due to source–tract interactions. Acta Acustica 92:468–475
Herzel H, Reuter R (1997) Whistle register and biphonation in a child’s voice. Folia Phoniat Logop 49:216–224
Kidjo N, Cargnelutti B, Charlton BD, Wilson C, Reby D (2008) Vocal behaviour in the endangered Corsican deer: description and phylogenetic implications. Bioacoustics 18:159–181
Ludt CJ, Schroeder W, Rottmann O, Kuehn R (2004) Mitochondrial DNA phylogeography of red deer (Cervus elaphus). Molec Phylogen Evol 31:1064–1083
Mahmut H, Masuda R, Onuma M, Takahashi M, Nagata J, Suzuki M, Ohtaishi N (2002) Molecular phylogeography of the red deer (Cervus elaphus) populations in Xinjiang of China: comparison with other Asian, European, and North American populations. Zool Sci 19:485–495
Mergell P, Herzel H (1997) Modelling biphonation—the role of the vocal tract. Speech Communicat 22:141–154
Nikol’skii AA (1975) Basic patterns of male Bactrian red deer (Cervus elaphus bactrianus) rutting calls. Zool Zh 54:1897–1900 (in Russian)
Nikol’skii AA (2011) The effect of amplitude modulation on the spectrum structure of the red deer sound signal. Doklady Biol Sci 437:107–109 (in Russian)
Nikol’skii AA, Pereladova OB, Rutovskaja MV, Formozov NA (1979) The geographical variability of rut calls in red deer males. Bull Moscow Soc Natur, Biol Ser 84(6):46–55, in Russian
Peters G, East ML, Herzel H, Henschel JR, Mills MGL, Wilhelm K, Hofer H (2004) Spotted hyaena whoops: frequent incidence of vocal instabilities in a mammalian loud call. Bioacoustics 14:99–109
Reby D, McComb K (2003) Anatomical constraints generate honesty: acoustic cues to age and weight in the roars of red deer stags. Anim Behav 65:519–530
Riede T, Suthers RA (2009) Vocal tract motor patterns and resonance during constant frequency song: the white-throated sparrow. J Comp Physiol A 195:183–192
Riede T, Titze IR (2008) Vocal fold elasticity of the Rocky Mountain elk (Cervus elaphus nelsoni)—producing high fundamental frequency vocalization with a very long vocal fold. J Exp Biol 211:2144–2154
Riede T, Suthers RA, Fletcher NH, Blevins WE (2006) Songbirds tune their vocal tract to the fundamental frequency of their song. PNAS 103:5543–5548
Struhsaker TT (1968) The behavior of the elk (Cervus canadensis) during the rut. Z Tierpsychol 24:80–114
Taylor AM, Reby D (2010) The contribution of source–filter theory to mammal vocal communication research. J Zool (L) 280:221–236
Titze IR (1994) Principles of voice production. Prentice-Hall, Englewood Cliffs
Titze IR (2008) Nonlinear source–filter coupling in phonation: theory. J Acoust Soc Am 123:2733–2749
Titze IR, Riede T (2010) A cervid vocal fold model suggests greater glottal efficiency in calling at high frequencies. PLoS Comp Biol 6(8):e1000897. doi:10.1371/journal.pcbi.1000897
Titze IR, Riede T, Popollo P (2008) Nonlinear source–filter coupling in phonation: vocal exercises. J Acoust Soc Am 123:1902–1915
Volodin IA, Lapshina EN, Volodina EV, Frey R, Soldatova NV (2011) Nasal and oral calls in juvenile goitred gazelles (Gazella subgutturosa) and their potential to encode sex and identity. Ethology 117:294–308
Wilden I, Herzel H, Peters G, Tembrock G (1998) Subharmonics, biphonation, and deterministic chaos in mammal vocalization. Bioacoustics 9:171–196
Zachos FE, Hartl GB (2011) Phylogeography, population genetics and conservation of the European red deer Cervus elaphus. Mammal Rev 41:138–150
Acknowledgments
We would like to thank David Reby, Tobias Riede, and Hansjoerg Kunc for discussion and Robert Huber and the two anonymous reviewers for their kind and instructive comments. During our work, we adhered to the “Guidelines for the treatment of animals in behavioural research and teaching” (Anim. Behav., 2006, 71, 245–253) and to the laws of Spain, Germany, and the Russian Federation, the countries where the research was conducted. This study was supported by the Russian Foundation for Basic Research, grant 12-04-00260 (for IV and EV) and Spanish projects CGL2007-63594 and CGL2010-17163 (for JC and JT).
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Volodin, I., Volodina, E., Frey, R. et al. Spectrographic analysis points to source–filter coupling in rutting roars of Iberian red deer. acta ethol 16, 57–63 (2013). https://doi.org/10.1007/s10211-012-0133-1
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DOI: https://doi.org/10.1007/s10211-012-0133-1