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Geospatial Variation of Ribbed Mussel (Geukensia demissa) Ecosystem Services across the Salt Marsh Landscape A Thesis Submitted to the Faculty of Drexel University by Joshua Andrew Moody in partial fulfillment of the requirements for the degree of Doctor of Philosophy December 2017 ©Copyright 2017 Joshua Andrew Moody. All Rights Reserved ii Acknowledgements I would like to dedicate this work to my beautiful and wonderful partner Kat Buchanan and our two amazing children, Cooper Kathleen and Liam James. Kat, without your love and support I never could have completed this work. Your endless patience, understanding, and sacrifice are deeply appreciated and unwaveringly acknowledged. I love you as big as the sky and deep as the ocean; you are my everything. I would like to thank my parents for their ability to see “the long game”, and for their support and love as I searched for my path. I would like to thank my committee: Dr. Susan Kilham; Dr. Danielle Kreeger, Dr. David Bushek; Dr. Tracy Quirk; Dr. Elizabeth Watson; and Dr. Robert Chant. Thank you for all of the time you have invested in my training, the comments you have provided on this work, and the encouragement to see this endeavor through. I would especially like to thank Dr. Danielle Kreeger and Dr. David Bushek, my greatest professional champions, for taking a chance on me so many years ago. Thank you for seeing something in me that gave you both the confidence to invest so many resources in my development. You have deeply changed my life, and I am lucky to be able to work with you both. Words cannot convey the breadth of my gratitude. I would also like to thank my colleagues Angela Padeletti, Kurt Cheng, and LeeAnn Haaf, who have helped me in so many ways since the beginning of this effort. Whether it was field support, intellectual motivation, questions that birthed new ideas, or simply being there when I needed you, thank you. I am lucky to know and work with you all, hopefully for many years to come. Additionally, thank you to the entire staff of the Partnership for the Delaware Estuary (past and present) who have provided support, especially: Sarah Bouboulis; Spencer Roberts; Sandra Demberger; Ryan Flannary; Emily Baumbach; Pricilla Cole; Laura Whalen; and many others. Finally I would like to thank the Environmental Protection Agency Region II (Irene Purdy and Kathleen Drake) as well as the EPA Office of Research and Development Atlantic Ecology Division (Marty Chintala and Suzanne Ayvazian) for their support of this work via Regionally Applied Research Effort (RARE) Grant Number 83523301. iii Table of Contents List of Tables ................................................................................................................................... vi List of Figures .................................................................................................................................. ix Abstract .......................................................................................................................................... xi Chapter 1: Introduction ................................................................................................................... 1 1.1 Mid-Atlantic Salt Marshes ................................................................................................ 1 1.2 Ecosystem Services of Salt Marshes ................................................................................ 2 1.3 Impacts of Sea Level Rise on Mid-Atlantic Salt Marshes ................................................. 4 1.4 The Life History of the Ribbed Mussel (Geukensia demissa) ........................................... 7 1.5 Ecology of the Ribbed Mussel (Geukensia demissa) ........................................................ 8 1.6 Spatial Variation of the Ribbed Mussel (Geukensia demissa) in Salt Marshes .............. 10 1.7 The Ribbed Mussel (Geukensia demissa) and the Nitrogen Cycle ................................. 10 1.8 Research Questions and Dissertation Structure ............................................................ 13 Chapter 2: Temporal and Spatial Variation of the Gross Removal of Total Suspended Solids and Particulate Nutrients by Ribbed Mussels (Geukensia demissa) in Salt Marshes of the Mid- Atlantic Region, USA ..................................................................................................................... 15 2.1 Abstract ................................................................................................................................ 15 2.2 Introduction ......................................................................................................................... 16 2.3 Methods ............................................................................................................................... 18 2.3.1 Study Sites and Habitat Delineation ............................................................................. 18 2.3.2 Ribbed Mussel and Seston Collection ........................................................................... 19 2.3.3 Experimental Approach ................................................................................................ 20 2.3.4 Mussel Body Metrics ..................................................................................................... 21 2.3.5 Clearance Rate Calculation ........................................................................................... 21 2.3.6 Seston Composition ...................................................................................................... 22 2.3.7 Filtration Rate Calculation............................................................................................. 23 2.3.8 Allometric Scaling of Physiological Rates ...................................................................... 23 2.3.9 Statistical Analyses ........................................................................................................ 23 2.4 Results .................................................................................................................................. 24 2.4.1 Mussel Body Metrics ..................................................................................................... 24 2.4.2 Seston Composition ...................................................................................................... 25 2.4.3 Clearance Rates ............................................................................................................. 26 iv 2.4.4 Filtration Rates .............................................................................................................. 26 2.5 Discussion............................................................................................................................. 27 Chapter 3: Ecosystem Services of the Ribbed Mussel (Geukensia demissa) Across the Salt Marsh Landscape ........................................................................................................................... 39 3.1 Abstract ................................................................................................................................ 39 3.2 Introduction ......................................................................................................................... 40 3.3 Methods ............................................................................................................................... 44 3.3.1 Site Selection ................................................................................................................. 44 3.3.2 Habitat Delineation ....................................................................................................... 44 3.3.3 Transect and Sampling Plot Layout ............................................................................... 45 3.3.4 Physiological Rates, Total Suspended Solids (TSS) and Particulate Nitrogen (PN) Concentrations ....................................................................................................................... 46 3.3.5 Ribbed Mussel Density and Size Demographics ........................................................... 47 3.3.6 Ribbed Mussel Biomass ................................................................................................ 48 3.3.7 Inundation time per Habitat ......................................................................................... 49 3.3.8 Habitat Delineation and Percent Habitat Area ............................................................. 51 3.3.9 Calculation of Filtration Rate of Total Suspended Solids and Particulate Nitrogen ..... 51 3.3.10 Contribution to Vertical Accretion .............................................................................. 52 3.3.11 Net Removal of Particulate Nitrogen (PN) .................................................................. 53 3.3.12 Statistical Analyses ...................................................................................................... 54 3.4 Results .................................................................................................................................. 55 3.4.1 Physiological Rates, Total Suspended Solids (TSS) and Particulate Nitrogen (PN) Concentrations ....................................................................................................................... 55 3.4.2 Ribbed Mussel Biomass ................................................................................................ 56 3.4.3 Inundation time and Elevation