Habitat-Specific Production of a Fall Line River Shoal Macroinvertebrate Assemblage
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HABITAT-SPECIFIC PRODUCTION OF A FALL LINE RIVER SHOAL MACROINVERTEBRATE ASSEMBLAGE by TIMOTHY DAVIS WYNN ARTHUR C. BENKE, COMMITTEE CHAIR ALEXANDER D. HURYN AMELIA K. WARD JONATHAN P. BENSTEAD JACK W. FEMINELLA A DISSERTATION Submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy in the Department of Biological Sciences in the Graduate School of The University of Alabama TUSCALOOSA, ALABAMA 2012 Copyright Timothy Davis Wynn 2012 ALL RIGHTS RESERVED ABSTRACT Fall Line shoals are zones of geomorphic complexity within a river basin, and have been recognized as sites of high aquatic macroinvertebrate diversity and production. The shoals of the free-flowing Cahaba River, in central Alabama, represent some of the most significant remaining examples of this channel feature that was once common throughout many rivers of the southeastern United States prior to widespread river regulation. The goal of this dissertation is to examine how the major habitats of a Cahaba River shoal influence the distribution and secondary production of the macroinvertebrate assemblage. Chapter 2 quantifies the variety of habitat types across the shoal reach and examines the temporal biomass dynamics of the 2 most common in-stream macrophytes, Justicia americana and Podostemum ceratophyllum. Chapter 3 presents a new method for obtaining in situ growth rates of several species of the diverse pleurocerid snail assemblage. The results of this method were later used to estimate production for this family. Chapter 4 describes the distribution, biomass, and production of the nonnative Asiatic clam, Corbicula fluminea, across the shoal reach, highlighting its dependence on Justicia habitat. Finally, Chapter 5 incorporates the preceding chapters into a study of the distribution of macroinvertebrate assemblage production across bare bedrock, Justicia, and Podostemum habitats, as well as the entire shoal reach. Total annual production of all macroinvertebrates was 56.1 g AFDM m-2 y-1 in bedrock, 284.4 in Podostemum, and 177.3 in Justicia habitats. Total habitat-weighted production of the shoals reach was 87.1 g m-2 y-1, with bedrock contributing 24.3%, Podostemum 22.7%, and Justicia 53.0% to this total. This study supports the view that ii Fall Line shoals can support high habitat diversity and production, and that the more complex habitats (e.g., those with macrophytes) enhance benthic invertebrate diversity and production. Also, the influence of a given habitat depends largely on its relative abundance, and this study demonstrated that the Justicia habitat can have a dominant influence on diversity and production of a river reach. This work advances our understanding of the roles of shoal habitats in maintaining the diversity and function of this endangered river channel feature. iii DEDICATION This dissertation is dedicated to my dad, Jeffrey Vernon Wynn, who took the time to show his sons the beauty and mystery of nature. iv LIST OF ABBREVIATIONS AND SYMBOLS Fig. Figure No. Number ind. Individuals spp. Species, plural sp. Species, singular % Percent km Kilometers m Meters cm Centimeters mm Millimeters μm Micrometers AW Aperture width L Shell length bs Body size g Instantaneous growth rate g Grams mg Milligrams AFDM Ash-free dry mass v DM Dry mass # Pound SA Surface area L Liter T Temperature C Degrees Celsius Degrees ‘ Minutes “ Seconds t Time in days y Year mo Month d day min Minute s Second %C Percent cover Ni No. of habitat-specific observations Nt No. of total habitat observations pH Negative logarithmic value of Hydrogen ion conentration NO2-N Nitrite-nitrogen NO3-N Nitrate-nitrogen PO4-P Phosphate-phosphorus CaCO3 Calcium carbonate vi SE Standard error n Number of independent observations F F statistic p Probability of rejecting the null-hypothesis if that hypothesis is true α Significance level r2 Coefficient of determination ANOVA Analysis of variance ln Natural logarithm log Logarithm < Less than << Much less than ≤ Less than or equal to > Greater than ≥ Greater than or equal to + Plus - Minus ± Plus or minus x Multiplication sign x Independent variable = Equal to ~ Approximately ∆ Change in et al. And others vii i.e. That is e.g. For example vs. Versus ® Trademark Inc. Incorporated Co. County AL Alabama MD Maryland UA The University of Alabama USFWS United States Fish and Wildlife Service USGS United States Geological Survey BR Bare bedrock, BRD Dry bedrock POD Podostemum JUS Justicia JUSD Dry Justicia JUStotal Submerged and emergent Justicia tissues JUSemerge Emergent Justicia tissues JUSsub Submerged Justicia tissues HYM Hymenocallis BO Boulder BOD Dry boulder G Gravel viii S Sand M Mud SG Snags DP Deep pool W Ind. mass W Size class mass W Mean mass between size classes N Density B Biomass B Mean annual biomass P Production P/B Production to biomass ratio CPI Cohort production interval X Present JAN January FEB February MAR March APR April JUN June JUL July AUG August SEP September OCT October ix NOV November DEC December x ACKNOWLEDGEMENTS This dissertation would not have been possible without the generous help and encouragement of many people. First, I would like to express my sincere appreciation to all the members of my committee, Drs. Arthur C. Benke, Jonathan P. Benstead, Alexander D. Huryn, Amelia K. Ward, and Jack W. Feminella, for contributing their invaluable time and insight toward the planning and execution of this work. I am especially grateful to my advisor, Dr. Arthur C. Benke, for guiding me through this long and difficult process, and for believing in me even when I doubted myself. Working under his mentorship has been a true honor. I would also like to express my gratitude to my family for their undying support, guidance, and friendship. Special thanks go to my wonderful wife and best friend, Kimberly, who has stood beside me every step of the way, and has given selflessly to help me achieve this goal. Mark Dedmon assisted me through every sampling trip, and was a cheerful companion throughout long exhausting days in the field. Laura Frost was my tireless lab assistant who helped make an impossible task possible. I would also like to thank fellow students Jeffrey Pollock, Michael Venarsky, James Ramsey, Stephanie Parker, and Michael Kendrick for assistance in the field and lab, and for many helpful discussions. Thanks as well to Dr. Christina Staudhammer for helpful advice about statistical methods. Finally, special thanks go to Lori Tolley-Jordan, who first showed me around the Cahaba shoals, and introduced me to its amazing snail fauna. Financial support was provided by the University of Alabama Department of Biological Sciences, the UA Graduate School, and the UA National Alumni Association. xi CONTENTS ABSTRACT.................................................................................................................................... ii DEDICATION............................................................................................................................... iv LIST OF ABBREVIATIONS AND SYMBOLS ............................................................................v ACKNOWLEDGEMENTS........................................................................................................... xi LIST OF TABLES....................................................................................................................... xiii LIST OF FIGURES ..................................................................................................................... xiv LIST OF APPENDICES.............................................................................................................. xvi CHAPTER 1: INTRODUCTION....................................................................................................1 CHAPTER 2: QUANTIFICATION OF BENTHIC HABITATS AND VEGETATION DYNAMICS IN A FALL LINE RIVER SHOAL.....................................................................8 CHAPTER 3: IN SITU ESTIMATION OF PLEUROCERID SNAIL GROWTH USING A TETHERING TECHNIQUE IN A FALL LINE RIVER SHOAL..........................................29 CHAPTER 4: PRODUCTION OF THE NONNATIVE ASIATIC CLAM, CORBICULA FLUMINEA, IN A SOUTHEASTERN RIVER BEDROCK SHOAL....................................45 CHAPTER 5: HABITAT-SPECIFIC PRODUCTION OF A FALL LINE RIVER SHOAL MACROINVERTEBRATE ASSEMBLAGE.........................................................................63 CHAPTER 6: GENERAL CONCLUSIONS...............................................................................116 LITERATURE CITED ................................................................................................................121 xii LIST OF TABLES 2.1 Mean annual velocity and mean annual depth for both macrophyte habitats, Justicia americana and Podostemum ceratophyllum............................................................................21 2.2 Conversion of mean annual biomass per unit habitat to mean annual biomass per unit shoal surface area ..............................................................................................................................22 3.1 Length-mass regression relationships for Pleurocera vestita, Elimia clara, and Elimia showalteri.................................................................................................................................36 3.2 Simple regression model parameters and statistics for each species as well as combined genus and family-level models ................................................................................................39