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Feeding Ecology, Gull Harassment and Reproductive Success of Female

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Feeding Ecology, Gull Harassment and Reproductive Success of Female FEEDING ECOLOGY, GULL HARASSMENT AND REPRODUCTIVE SUCCESS OF FEMALE SOUTHERN RIGHT WHALES by Carina Flavia Marón A dissertation submitted to the faculty of The University of Utah in partial fulfillment of the requirements for the degree of Doctor of Philosophy Department of Biology The University of Utah August 2015 Copyright © Carina Flavia Marón 2015 All Rights Reserved The University of Utah Graduate School STATEMENT OF DISSERTATION APPROVAL The dissertation of Carina Flavia Marón has been approved by the following supervisory committee members: Jon Seger , Chair 05/04/15 Date Approved Victoria Rowntree , Member 05/04/15 Date Approved M. Denise Dearing , Member 05/04/15 Date Approved Colleen G. Farmer , Member 05/04/15 Date Approved Robert Ward , Member 05/04/15 Date Approved and by M. Denise Dearing , Chair/Dean of the Department of Biology and by David B. Kieda, Dean of The Graduate School. ABSTRACT Many factors can affect reproductive success in animals. In marine mammals, both nutritional and environmental stresses have been shown to influence maternal fertility and offspring survival. The southern right whales (Eubalaena australis) that calve off Península Valdés, Argentina, have experienced increased rates of calf death since 2003, but the cause(s) of this elevated mortality remain unknown. My research addresses the possible effects of diet and Kelp Gull harassment on the reproductive success of the whales. This study also provides novel insights into the diet composition, patterns of fattening, behavior, and reproductive life histories of right whales. I first asked whether dead calves show a reduced proportion of fatty acids that are essential for calf growth, but I found no evidence to support this hypothesis. Then, I asked whether mothers of dead calves feed in different feeding grounds than mothers of living calves, but I found no significant differences among the two groups. Third, I investigated the diet composition of whales, and found that some mothers rely more than others on certain copepod species. Next, I asked whether the body fat condition of calves worsens in years with high calf mortality, or in calves with more gull-inflicted lesions. I found no evidence to suggest a decline in the body fat condition of calves. Then, I evaluated how blubber thickness changes with calf growth and found that calves acquire a more streamlined body shape as they age. I asked whether the incidence of gull-inflicted lesions in mothers and calves has increased in the 2000s (coincidentally with calf deaths). The numbers and areas of lesions in calves have increased greatly compared to mothers, suggesting that the health of nursing calves could be compromised if wounding continues to increase. Finally, I investigated the occurrence of abnormal calving intervals and found that they increase significantly in years of high calf mortality compared to years of low mortality. I modeled the potential effects of a sustained increase in calf deaths and found that if elevated rates of calf mortality continue for some years, the population’s growth could be slowed substantially. iv To my family, friends and the whales. TABLE OF CONTENTS ABSTRACT ....................................................................................................................... iii LIST OF TABLES ............................................................................................................. ix LIST OF FIGURES .......................................................................................................... xii ACKNOWLEDGEMENTS ............................................................................................. xvi Chapters 1. INTRODUCTION .........................................................................................................1 References ............................................................................................................................4 2. DIETARY DIVERSITY IN THE SOUTHERN RIGHT WHALE POPULATION OFF ARGENTINA INFERRED FROM FATTY ACIDS AND STABLE ISOTOPES .....7 Abstract ................................................................................................................................7 Introduction ..........................................................................................................................8 Materials and Methods .......................................................................................................12 Blubber and Skin Samples from Dead Calves ...........................................................12 Blubber and Skin Samples from Living Whales ........................................................13 Lipid Analysis ............................................................................................................13 Stable Isotope Analysis ..............................................................................................14 Fatty Acid Data for Zooplankton ...............................................................................15 Statistical Analyses .....................................................................................................16 Results ................................................................................................................................17 Fatty Acid Composition in Southern Right Whales ...................................................17 Fatty Acid Composition of Calves Changes with Growth .........................................18 Large Dead Calves Do Not Show Signs of Malnutrition ...........................................19 Fatty Acid Profiles of Living Calves Show Evidence of Nursing .............................20 Mothers Feed on a Varied Diet ..................................................................................21 Feeding Grounds Do Not Differ Among Mothers of Living and Dead Calves .........22 Discussion ..........................................................................................................................23 References ..........................................................................................................................31 3. NO EVIDENCE OF MALNUTRITION IN DEAD SOUTHERN RIGHT WHALE CALVES OFF ARGENTINA AS INFERRED FROM BLUBBER THICKNESS MEASUREMENTS ...........................................................................................................54 Abstract ..............................................................................................................................54 Introduction ........................................................................................................................55 Materials and Methods .......................................................................................................59 Necropsies ..................................................................................................................59 Blubber Thickness and Length Measurements ..........................................................59 Carcass Decomposition ..............................................................................................60 Stranding Time in the Season .....................................................................................61 High and Low Mortality Years ..................................................................................61 Gull-Inflicted Lesions .................................................................................................62 Statistical Analyses .....................................................................................................62 Results ................................................................................................................................63 Blubber Thickness Increases with Calf Length ..........................................................63 Blubber Thickness Often Does Not Change with Carcass Decay ..............................64 Blubber Thickness Differs Between Stranding Locations but Not Between Sexes ...64 Calves that Died Late in the Season Have Thicker Blubber ......................................65 Calves that Died in High Mortality Years Have Thicker Blubber .............................65 Gull-Inflicted Lesions Appear Not to Affect Blubber Thickness ..............................65 Calves Acquire a Streamlined Body Shape with Growth ..........................................66 Discussion ..........................................................................................................................67 References ..........................................................................................................................72 4. INCREASED WOUNDING OF SOUTHERN RIGHT WHALE CALVES BY KELP GULLS AT PENÍNSULA VALDÉS ................................................................................85 Abstract ..............................................................................................................................85 Introduction ........................................................................................................................86 Materials and Methods .......................................................................................................89 Presence and Number of Lesions on Living Mother-Calf Pairs .................................89 Presence and Number of Lesions on Dead Calves .....................................................90 Size of Lesions on Living and Dead Whales ..............................................................91
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