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The Pennsylvania State University The Pennsylvania State University The Graduate School Ecology VOCAL NOISE COMPENSATION IN NONHUMAN MAMMALS: MODIFICATION TYPES AND USAGE PATTERNS A Dissertation in Ecology by Cara F. Hotchkin 2012 Cara F. Hotchkin Submitted in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy December 2012 The dissertation of Cara F. Hotchkin was reviewed and approved* by the following: Susan E. Parks Associate Professor of Acoustics and Ecology Senior Research Associate, Penn State Applied Research Laboratory Dissertation Adviser Chair of Committee Victoria A. Braithwaite Professor of Fisheries and Biology Thomas B. Gabrielson Professor of Acoustics Senior Scientist, Penn State Applied Research Laboratory Tracy L. Langkilde Associate Professor of Biology Jennifer L. Miksis-Olds Assistant Professor of Acoustics Senior Research Associate, Penn State Applied Research Laboratory David M. Eissenstat Professor of Woody Plant Physiology Chair, Intercollege Graduate Degree Program in Ecology *Signatures are on file in the Graduate School iii ABSTRACT Vocal noise compensation and vocal flexibility in response to increased noise are important evolutionary adaptations which allow signalers to successfully communicate in highly variable acoustic environments. While many species of mammals use multiple types of noise- induced modifications, the details of vocal responses to noise are unclear. Open questions include whether specific noise parameters elicit certain modification types, whether animals can adjust different vocal characteristics independently, and what effects behavioral contexts have on vocal noise compensation. The goals of this dissertation were to evaluate these questions, and to characterize the effects of noise amplitude and bandwidth (including degree of spectral overlap) on the acoustic structure of vocalizations produced by two acoustically dependent non-human mammal species. Methods included passive acoustic recordings of vocalizations, investigation of the subjects’ acoustic habitats, and controlled playback experiments. Amplitude and spectral characteristics of acoustic habitats were examined for two (one captive and one wild) beluga whale (Delphinapterus leucas) populations. In the wild habitat, noise was measured during beluga encounters at two sites. Noise sources and levels varied between the two recording locations, with higher overall levels and a higher vessel/beluga encounter rate at Beluga River, and more noise from weather events at Kenai River. In the captive habitat, at Mystic Aquarium, noise levels were substantially higher than noise in Cook Inlet, and noise varied spatially within the exhibit and over both short- and long temporal scales. Comparison of noise levels with published beluga audiograms suggests that due to the differences in noise levels, belugas at Mystic Aquarium would be likely to hear noise at and above 2 kHz, and that in Cook Inlet, belugas are more likely to hear only noise above 4 kHz. Noise-induced vocal modifications were studied in both the captive and wild beluga populations during increased noise from exhibit maintenance or vessel passages, respectively. In both populations, noise-induced changes to vocalization structure were observed, primarily in the spectral content of calls. In captive belugas, minimum call frequencies were significantly related to narrowband noise levels, while in the wild population peak frequency was more likely to be related to noise levels. There were no consistent relationships between the duration of calls and noise levels in either population. Behavioral or environmental contexts also had significant relationships with vocalization structure, indicating a possible interaction between the signalers’ communicative motivation and vocal noise compensation strategies, as observed for human speech. Modifications observed from the captive whales appeared unlikely to increase iv communication success during increased exhibit noise, but those produced by wild whales may serve a noise compensation function. Experimental tests of the onset order of vocal modification types and subjects’ attention to specific noise parameters were performed with a captive group of cotton-top tamarins (Sagunius oedipus), an acoustically dependent social New World primate. Observed vocal modifications included the Lombard effect, frequency shifts, and temporal changes, but differed between the two examined call types. Changes to the spectral content and amplitude of vocalizations appeared to occur non-simultaneously for both call types. Data in this chapter represent the first observations of short-term changes to the spectral content of non-human primate vocalizations, and the first empirical demonstration of shifts in spectral tilt in non-human mammal calls. Similarity in responses of humans, cotton-top tamarins, and other non-human mammals indicate relative consistency of selection for signalers able to compensate for short- term increases in noise level. The work presented in this dissertation expands current knowledge of the effects of noise on acoustic communication by nonhuman animals, and advances both theoretical understanding of the evolution of vocal noise compensation and practical considerations of the use of vocal modifications for monitoring and conservation of vulnerable species. It provides substantial evidence for the importance of communicative motivation in vocal noise compensation, and demonstrates previously unknown vocal flexibility in a well-studied non-human primate species. This dissertation advances the study of acoustic communication by non-human mammals, with relevance to the evolution of communication and applied conservation biology. v TABLE OF CONTENTS LIST OF FIGURES ................................................................................................................. ix LIST OF TABLES ................................................................................................................... xvi ACKNOWLEDGEMENTS ..................................................................................................... xix Chapter 1 Introduction ............................................................................................................. 1 Acoustic communication by animals ............................................................................... 1 Relevance of this study .................................................................................................... 3 Summary of chapters and appendices .............................................................................. 4 References ........................................................................................................................ 5 Chapter 2 The Lombard Effect and other noise-induced vocal modifications: insights from mammalian communication systems ....................................................................... 7 Abstract ............................................................................................................................ 7 Introduction ...................................................................................................................... 7 Historical context and terminology .......................................................................... 10 Effects of noise on human speech .................................................................................... 11 The Lombard effect in humans ................................................................................ 12 Other NIVMs ........................................................................................................... 16 Lombard effect – NIVM linkage (“Lombard speech”) ............................................ 18 Effects of noise on non-human mammal vocalizations ................................................... 18 Vocal noise compensation in non-human mammals ................................................ 19 The Lombard effect in mammals ............................................................................. 19 Other NIVMs in non-human mammals .................................................................... 23 Lombard effect – NIVM Linkage in non-human mammals ..................................... 27 Future research directions ................................................................................................ 28 Future directions for human research ....................................................................... 29 Future directions for non-human mammal research ................................................. 30 Conclusions ...................................................................................................................... 33 APPENDIX: Glossary of terms ....................................................................................... 35 Acknowledgements .......................................................................................................... 38 References ........................................................................................................................ 38 Chapter 3 Characterization of acoustic habitat in captive and wild beluga environments ...... 44 Abstract ............................................................................................................................ 44 Introduction ...................................................................................................................... 45 Acoustic Habitats ....................................................................................................
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