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View / Download 4.4 Mb Fine-Scale Foraging Behavior of Humpback Whales Megaptera novaeangliae in the Near- Shore Waters of the Western Antarctic Peninsula by Reny Blue Tyson Marine Science and Conservation Duke University Date:_______________________ Approved: ___________________________ Douglas Nowacek, Supervisor ___________________________ Ari Friedlaender ___________________________ James Hench ___________________________ Daniel Rubenstein Dissertation submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy in Marine Science and Conservation in the Graduate School of Duke University 2014 i v ABSTRACT Fine-Scale Foraging Behavior of Humpback Whales Megaptera novaeangliae in the Near- Shore Waters of the Western Antarctic Peninsula by Reny Blue Tyson Marine Science and Conservation Duke University Date:_______________________ Approved: ___________________________ Douglas Nowacek, Supervisor ___________________________ Ari Friedlaender ___________________________ James Hench ___________________________ Daniel Rubenstein An abstract of a dissertation submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy in Marine Science and Conservation in the Graduate School of Duke University 2014 i v Copyright by Reny Blue Tyson 2014 Abstract High-resolution bio-logging tools were used to examine the fine-scale foraging behaviors of humpback whales (Megaptera novaeangliae) in the coastal waters of the Western Antarctic Peninsula during the austral autumn of 2009 and 2010. Discrete feeding events (i.e., lunges) were inferred from the biologging records of thirteen whales, including a mother and her calf. In general, humpback whales exhibited efficient foraging behaviors that allowed them to maximize energetic gains and minimize energetic costs as predicted by optimal foraging theory. They fed at a continuous and high rate in the upper portion of the water column (< 100 m) from approximately dusk to dawn when their prey (Antarctic krill, Euphausia superba) were most vulnerable and less costly to acquire (i.e., near the surface). When forced to dive to greater depths, they adjusted their behaviors (e.g., descent and ascent rates) so that they could maximize their foraging durations and increase their lunging rates. In addition, humpbacks appeared to accept short term (i.e., dive by dive) costs associated with depleted oxygen stores in favor of maximizing long term (i.e., daily) energetic gains. Such efficient behaviors are particularly beneficial for mother-calf pairs who have additional energetic costs associated with foraging, such as lactation (mother), growth (calf), and maintaining proximity. In addition, because the physiology of humpback whales is poorly understood yet critically important for predicting their behaviors in response to iv fluctuations in their environmental conditions, foraging behaviors inferred from the bio- logging records were used to estimate their metabolic rates, oxygen storage capacities, and oxygen replenishment rates under the framework of optimal foraging theory. This research suggests that the current techniques used to estimate humpback whale oxygen stores is appropriate but that the estimation of metabolic rates of humpbacks while foraging and while traveling need to be addressed further. This work aims to increase the current understanding of humpback whale foraging behaviors along the Western Antarctic Peninsula so that appropriate measures can be taken to aid in their recovery and in the sustainability of the Antarctic marine ecosystem. v Dedication I dedicate this dissertation to my mother, Dr. Gail Tyson, who got mad at me for jumping out of a plane... ....without her. vi Co-Authorship Statement The research presented in this dissertation is part of a collaborate study designed to elucidate the ecological relationships between humpback whale and Antarctic krill in the Western Antarctic Peninsula and aspects it have been previously published elsewhere (e.g., Nowacek et al. 2011, Ware et al. 2011, Espinasse et al. 2012, Johnston et al. 2012, Stimpert et al. 2012, Tyson et al. 2012, Friedlaender et al. 2013). Several colleagues contributed to my research and are (or will be) listed as co-authors on modified versions of each data chapter submitted for peer-review to scientific journals. Douglas Nowacek and Ari Friedlaender were co-principle investigators of the collaborative research project and supplied funding for much of the work. They are co- authors on all of my data chapters. Allison Stimpert and Colin Ware contributed significantly to the analysis of tag data presented in Chapters 2. Finally, James Clark helped me to design the simulated model to estimate humpback whale physiology in Chapter 4 and provided analytical support. vii Contents Abstract ......................................................................................................................................... iv List of Tables ................................................................................................................................. xi List of Figures ............................................................................................................................ xiii Acknowledgements ................................................................................................................. xvii 1. Introduction ............................................................................................................................... 1 1.1 Balaenopterid Foraging Ecology .................................................................................... 2 1.2 Research Objectives .......................................................................................................... 6 1.3 General Materials and Methods ..................................................................................... 8 1.3.1 Study Area .................................................................................................................... 9 1.3.2 Data Collection .......................................................................................................... 12 2. In Synch? Humpback Whale Mother and Calf Foraging Behavior: Insights from Multi-Sensor Suction Cup Tags* ............................................................................................... 20 2.1 Introduction ..................................................................................................................... 20 2.2 Methods ........................................................................................................................... 24 2.3 Results .............................................................................................................................. 30 2.3.1 Foraging Behaviors ................................................................................................... 30 2.3.2 Synchronous Behaviors ............................................................................................ 35 2.4 Discussion ........................................................................................................................ 39 3. How Should a Clever Whale Allocate Foraging Time and Effort .................................... 46 3.1 Introduction ..................................................................................................................... 46 3.2 Methods ........................................................................................................................... 49 viii 3.2.1 Observed Whale Behavior........................................................................................ 49 3.2.2 Optimal Model Based On Time Allocation ........................................................... 52 3.2.3 Statistical Analyses .................................................................................................... 56 3.3 Results .............................................................................................................................. 57 3.3.1 Model Agreement ...................................................................................................... 61 3.3.1.1 Observed Versus Predicted Foraging Behaviors ........................................... 61 3.3.1.2 Observed Versus Predicted Dive Durations .................................................. 65 3.3.1.3 Observed Versus Predicted Post-Dive Surface Durations ........................... 69 3.3.1.4 Individual Variation .......................................................................................... 70 3.3.1.5 Adherence of Data to Model Assumptions .................................................... 74 3.3.2 Proportional Foraging Behaviors ............................................................................ 75 3.4 Discussion ........................................................................................................................ 79 4. Physiology of Foraging Humpback Whales: Insights from an Optimality Approach . 85 4.1 Introduction ..................................................................................................................... 85 4.1.1 Current Understanding of Humpback Whale Metabolic Rates ......................... 87 4.1.2 Current Understanding of Humpback Whale Oxygen Stores and Replenishment Rates .......................................................................................................... 91 4.2 Methods ..........................................................................................................................
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