Comparative Thermal Ecology of the Dusky Pipefish, Syngnathus
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COMPARATIVE THERMAL ECOLOGY OF THE DUSKY PIPEFISH, SYNGNATHUS FLORIDAE AND THE GULF PIPEFISH, SYNGNATHUS SCOVELLI by Lois Anne O’Boyle B.S. Bloomsburg University, 2003 A thesis submitted to the Department of Biology College of Arts and Sciences The University of West Florida In partial fulfillment of the requirements for the degree of Master of Science 2011 The thesis of Lois Anne O’Boyle is approved: ____________________________________________ _________________ Anne A. Boettcher, Ph.D., Committee Member Date ____________________________________________ _________________ Christopher L. Pomory, Ph.D., Committee Member Date ____________________________________________ _________________ Wayne A. Bennett Jr., Ph.D., Committee Chair Date Accepted for the Department/Division: ____________________________________________ _________________ George L. Stewart, Ph.D., Chair Date Accepted for the University: ____________________________________________ _________________ Richard S. Podemski, Ph.D., Dean, Graduate School Date ii ACKNOWLEDGEMENTS I would like to thank my committee members, Dr. Wayne Bennett, Dr. Christopher Pomory, and Dr. Anne Boettcher for their insight and advice on this project. I acknowledge the staff of the biology department for providing research space and necessary equipment, particularly Jim Hammond for never hesitating to answer the phone when I needed him, along with countless hours of equipment construction, maintenance, and a laugh or two. I acknowledge the Scholarly and Creative Activities Committee as well as the Marine Ecology Research Society for funding. I would like to thank my fellow lab members, past and present, particularly Ryan Saylor and Justin Speaks for assistance collecting and maintaining pipefish as well as the memories made in the lab and outside of the lab. Finally I would like to sincerely thank my family and friends for their never- ending support and love. Without their patience I would not have had the courage to complete this degree. Most importantly I thank my husband, Bob, who never lost faith in me and always encouraged me to pursue my dream. iii TABLE OF CONTENTS ACKNOWLEDGEMENTS ............................................................................................... iii LIST OF TABLES ...............................................................................................................v LIST OF FIGURES ........................................................................................................... vi ABSTRACT ...................................................................................................................... vii CHAPTER I. INTRODUCTION ...........................................................................1 CHAPTER II. MATERIALS AND METHODS .....................................................6 A. Pensacola Bay Thermal Profile ..................................................6 B. Pipefish Collection Data ............................................................7 C. Chronic Thermal Limits .............................................................8 D. Critical Thermal Limits..............................................................9 E. Thermal Tolerance Polygons ...................................................11 F. Statistical Analyses ..................................................................13 CHAPTER III. RESULTS ......................................................................................14 A. Thermal Profile of Santa Rosa Sound .......................................14 B. Chronic Thermal Analysis ........................................................15 C. Critical Thermal Analysis .........................................................17 D. Thermal Tolerance Polygons ....................................................19 CHAPTER IV. DISCUSSION ................................................................................23 A. Pipefish Thermal Physiology ....................................................23 B. Geographic Zone Interpretations ...............................................24 C. Comparison of Polygon Areas ..................................................25 D. Distributional Differences in Thermal Tolerance .....................26 E. Behavioral Responses to Water Temperature ...........................26 F. Direct Climactic Changes in Pipefish Ecology .........................27 G. Indirect Climactic Impacts ........................................................29 REFERENCES ..................................................................................................................31 APPENDIX ........................................................................................................................38 A. Animal Care and Use Committee Approval Letters .................39 iv LIST OF TABLES 1. Pipefish Chronic Thermal Tolerance Values. ...........................................................16 2. Pipefish Critical Thermal Tolerance Values .............................................................18 3. Replicate Thermal Tolerance Polygon Areas ............................................................20 v LIST OF FIGURES 1. Santa Rosa Sound Seagrass habitats selected for water temperature collection. Locations for each site are as follows: Bob Sikes Bridge (BSB) (30°21’ 15.3” N, 87°09’ 30.9” W), Pensacola Beach Side (PBS) (30°21’12.8”N, 87°02’ 10.9”W), Nantahala Road (NAN) (30°23’ 06.5”N, 87° 00’ 48.4”W) .......................................7 2. Ecological thermal tolerance polygon construction. Critical thermal maxima (CTmaxima) and critical thermal minima (CTminima) regression lines form upper and lower polygon boundaries .................................................................................12 3. Monthly (•) maximum and minimum water temperatures plotted with weekly average (solid line) for three sites in Santa Rosa Sound, Florida for the period of September 2008 to August 2009. Sites are labeled and located as follows: (A), Bob Sikes Bridge (30°21’ 15.3” N, 87°09’ 30.9” W), (B), Pensacola Beach Side (30°21’12.8”N, 87°02’ 10.9”W), (C), Nantahala Road (30°23’ 06.5”N, 87° 00’ 48.4”W) ....................................................................................................................15 4. Total thermal tolerance polygons for dusky (A) pipefish (n=4) and gulf (B) pipefish (n=5). Filled circles represent CTM regression values and open circles represent chronic thermal values extrapolated from CTM regression lines. Regression lines are shown with error bars for + 1 SD. Values for thermal tolerance zone areas given as upper acquired (UAZ), lower acquired (LAZ), intrinsic (ITZ) and total, for each species ........................................................................................21 5. Dusky and gulf pipefish comparison polygon. Values represented by open triangles ( ) identify the dusky pipefish polygon while filled circles (•) represent values for gulf pipefish .............................................................................................................22 vi ABSTRACT COMPARATIVE THERMAL ECOLOGY OF THE DUSKY PIPEFISH, SYNGNATHUS FLORIDAE AND THE GULF PIPEFISH, SYNGNATHUS SCOVELLI Lois Anne O’Boyle Across their latitudinal range, pipefish regularly experience large seasonal and diel shifts in water temperature yet little is known about the thermal tolerance of these fishes. In this study, chronic thermal limits determined acclimation temperatures for dusky (Syngnathus floridae) and gulf pipefish (Syngnathus scovelli) inhabiting seagrass beds in the Gulf of Mexico. Critical thermal methodology (CTM) was employed to estimate upper and lower thermal limits for each species as well as construct a thermal tolerance polygon demonstrating each species’ thermal niche. Daily water temperatures collected were used to identify specific variation encountered by pipefish in this study. Chronic thermal ranges were significantly different (p<0.0001) and provided acclimation temperature ranges of 11-33°C for gulf and 12-32°C for dusky pipefish. Critical thermal maxima and minima were significantly correlated with acclimation temperature (p<0.0001 in both species) and accounted for 93-98% of the variability in CTM. Polygons calculated for dusky and gulf pipefish had total areas of 617°C² and 736°C², respectively. Gulf pipefish possess significantly larger intrinsic and total tolerance area than dusky pipefish, which may indicate disparate use of seagrass habitats. Both species vii utilize well developed mechanisms for thermal tolerance as well as behavioral adaptations when temperatures fluctuate. viii CHAPTER I INTRODUCTION Pipefishes (family Syngnathidae) inhabit tidal waters across a wide range of climate zones. The snake pipefish, Entelurus aequoreus, for example, can be found in cold North Atlantic waters that seldom exceed 12°C (Kloppmann and Ulleweit, 2007), whereas scribbled pipefish, Corythoichthys intestinalis and the bellybarred pipefish, Syngnathus spicifer, inhabit tropical areas where temperatures routinely reach 35°C or more (Bennett 2001 unpublished data; Pinto and Punchihewa, 1996, respectively). Even within species, pipefishes can experience wide temperature variations across their geographic range. Gulf pipefish, Syngnathus scovelli, and dusky pipefish, Syngnathus floridae, for example, are common residents of Gulf of Mexico seagrass flats where rapid, and sometimes extreme, water temperature shifts are common. Average temperatures in shallow Gulf of Mexico waters typically exhibit seasonal shifts