Social Context of Gray Whale Eschrichtius Robustus Sound Activity
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SOCIAL CONTEXT OF GRAY WHALE Eschrichtius robustus SOUND ACTIVITY A Thesis by SARAH MARIE CHARLES Submitted to the Office of Graduate Studies of Texas A&M University in partial fulfillment of the requirements for the degree of MASTER OF SCIENCE May 2011 Major Subject: Wildlife and Fisheries Sciences SOCIAL CONTEXT OF GRAY WHALE Eschrichtius robustus SOUND ACTIVITY A Thesis by SARAH MARIE CHARLES Submitted to the Office of Graduate Studies of Texas A&M University in partial fulfillment of the requirements for the degree of MASTER OF SCIENCE Approved by: Chair of Committee, Bernd G. Würsig Committee Members, Robert H. Benson Roel R. Lopez Head of Department, Thomas E. Lacher, Jr. May 2011 Major Subject: Wildlife and Fisheries Sciences iii ABSTRACT Social Context of Gray Whale Eschrichtius robustus Sound Activity. (May 2011) Sarah Marie Charles, B.S., Xavier University Chair of Advisory Committee: Dr. Bernd G. Würsig This thesis examines sound production of eastern gray whales (Eschrichtius robustus) in the wintering lagoons to determine whether sound use is a function of social context. Proportions of sounds used, parameters of each sound class, and rates of sounds were compared among social contexts. Data revealed the strong possibility of context- specific use for particular sound classes. Additionally, sound parameters and rates of production varied by social context. These results reflect similar variations in gray whale repertoire throughout their range that may be due to changes in social and behavioral contexts. Gray whale sounds are classified into several classes based on aural and visual characteristics. This study verifies the classification system determined in previous studies, with the exception of class 8, and supports the division of class 1 into subclasses 1a and 1b. Class 1 appeared to be critical during sexual contexts and all highly social contexts, regardless of age and sex class. Although highly recognizable, its parameters exhibited much variation among social contexts; therefore class 1 may communicate graded emotional states in short-range interactions. Other classes of sounds may be utilized for long-distance communication, as startle responses, or ―precursors‖ to the adult repertoire. Frequency-related parameters of all sound classes showed variation among social contexts, but duration demonstrated very little variation. Calf-containing contexts exhibited greatest and most varied frequencies; this is to be expected if gray whale‘s sound mechanism is related to body and tracheal length. Variation also may indicate that physical maturity or learning play a role in the repertoire development. The lowest and least varied frequencies were observed in adult contexts. Sound production rates also varied by social context. Active adults produced sounds at high rates during iv short intervals; mixed/unknown contexts were often silent. Calf-containing contexts produced sounds at intermediate rates and were never silent. The correlations demonstrated here between social context and use of sounds will allow for acoustics to be an indicator of group composition, seasonal movements, and social patterns, thus relieving dependency on difficult visual observation. Additionally, such correlations provide preliminary information for determining sound functions. v ACKNOWLEDGMENTS I thank God first and foremost for the blessings He has provided me; the opportunity to study gray whale sounds is just one of many! I also thank Bernd Würsig for the opportunity to study under his tutelage and glean from his experiences and knowledge. Bernd‘s patience with my ―long term‖ study has been greatly appreciated. I also extend thanks to Bernd and Mel for travel funds, housing, and their warm hospitality. I am extremely grateful to the staff, pangueros and students of the School for Field Studies / Center for Coastal Studies in Puerto San Carlos. Without their resources and hard work, this study would not have been possible. Francisco (Paco) Ollervides and SFS created this tremendous opportunity by generously providing boats and fuel, acoustic equipment, student assistants, and room and board during data collection. Hector Pérez-Cortés extended the school‘s generosity for a second season, and he was also an important source of gray whale knowledge. The students were invaluable in data collection and management, and additionally provided plenty of fun adventures and comical diversions! The ‗sound‘ advice (pun intended!) of Manuel Castellote and Shannon Rankin has been priceless. Thanks to my committee members Robert Benson and Roel Lopez. I am grateful for Dr. Benson‘s instruction in sound analysis and Dr. Lopez‘s advice in data management and analysis. I appreciate Robin Vaughn for being a helpful long-distance lab mate. The help received at the Texas A&M Statistics Department Statistical Consulting Center was invaluable. I am extremely grateful to May Boggess and graduate students Jing Li, Randall Gil, and Mir Usman Ali for their uphill battle with my data. Without their expertise and instruction, I would still be drowning in my data! The Cornell Lab of Ornithology and Macauley Library Audio Workshop provided basic instruction in audio recording techniques and analysis with Raven software. The Marine Mammal Research Program (Texas A&M University at vi Galveston) and the Center for Bioacoustics (Texas A&M University at Corpus Christi) provided resources and labs for advanced sound analysis and organization. There are countless others who have contributed to this study and for their help I am grateful: Jane Packard, Sheyna Wisdom, Nicole Crane, Christopher Clark for loan of acoustic instruments, Steven Swartz for advice, Molly Meserve, and Robert McCleery. I greatly appreciate the grants and funding that made this study possible: Erma Lee and Luke Mooney travel grant; Texas A&M Office of Graduate Studies Merit Fellowship; MARB travel grant; National Fish and Wildlife Foundation; Office of Graduate Studies Research and Presentation Grant; and Graduate Program Enhancement Fund (GPEF) Travel and Presentation Grant. Finally, I cannot thank enough my loved ones for their unconditional and endless supply of love and support. My parents, Louis and Regina Rohrkasse, provided me with a fantastic education and the opportunities to pursue my dreams. Thank you mom for sending my hometown chili and Girl Scout cookies to México! My husband Eddie patiently endured several years of out-of-state and out-of-country research. I am so grateful to be so blessed! vii TABLE OF CONTENTS Page ABSTRACT .......................................................................................................... iii ACKNOWLEDGMENTS ...................................................................................... v TABLE OF CONTENTS ....................................................................................... vii LIST OF FIGURES ............................................................................................... ix LIST OF TABLES ................................................................................................. xii I INTRODUCTION ........................................................................................... 1 Determining Sound Function by Social Context ......................................... 2 Characteristics of Social Context ................................................................ 5 Products of Determining Social Context and Function of Sounds ............... 6 Gray Whale Natural History ....................................................................... 9 Gray Whale Sound Repertoire .................................................................... 11 II METHODS ...................................................................................................... 15 Behavior Analysis ...................................................................................... 17 Acoustic Analysis ....................................................................................... 18 Statistical Analysis ..................................................................................... 19 III RESULTS ........................................................................................................ 22 Classes of Sounds and Their Parameters ..................................................... 22 Parameter Variations by Class .................................................................... 35 Parameter Variations between Class 1 Subclasses....................................... 40 Parameter Variations by Social Context ...................................................... 46 Class 1 Subclass Parameter Variations by Social Context ........................... 51 Relative Use of Classes by Social Context .................................................. 62 Relative Use of Class 1 Subclasses ............................................................. 68 Relative Use of Types ................................................................................ 69 Rates of Sound Production ......................................................................... 71 Use of Four Social Contexts ....................................................................... 75 IV DISCUSSION .................................................................................................. 83 Class Parameters ........................................................................................ 86 viii Page Class Parameters by Social Context ...........................................................