UUnniivveerrssiittyy ooff SSoouutthh FFlloorriiddaa DDiiggiittaall CCoommmmoonnss @@ UUnniivveerrssiittyy ooff SSoouutthh FFlloorriiddaa USF Tampa Graduate Theses and Dissertations USF Graduate Theses and Dissertations 5-3-2012 TThhee EEccoollooggiiccaall RRoollee ooff RRhhiizzoopphhyyttiicc GGrreeeenn AAllggaaee iinn SSoofftt--bboottttoomm HHaabbiittaattss Laura Bedinger University of South Florida, [email protected] Follow this and additional works at: https://digitalcommons.usf.edu/etd Part of the American Studies Commons, and the Ecology and Evolutionary Biology Commons SScchhoollaarr CCoommmmoonnss CCiittaattiioonn Bedinger, Laura, "The Ecological Role of Rhizophytic Green Algae in Soft-bottom Habitats" (2012). USF Tampa Graduate Theses and Dissertations. https://digitalcommons.usf.edu/etd/3972 This Dissertation is brought to you for free and open access by the USF Graduate Theses and Dissertations at Digital Commons @ University of South Florida. It has been accepted for inclusion in USF Tampa Graduate Theses and Dissertations by an authorized administrator of Digital Commons @ University of South Florida. For more information, please contact [email protected]. The Ecological Role of Rhizophytic Green Algae in Soft-bottom Habitats by Laura Ann Bedinger A dissertation submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy Department of Integrative Biology College of Arts and Sciences University of South Florida Major Professor: Susan S. Bell, Ph.D. Clinton J. Dawes, Ph.D. Peter D. Stiling, Ph.D. Kevin S. Beach, Ph.D. Date of Approval: May 3, 2012 Keywords: Bryopsidales, rhizoid, Florida, seagrass bed, biomass, succession, disturbance, vegetative reproduction, regeneration, secondary metabolite, fouling Copyright © 2012, Laura Ann Bedinger DEDICATION This dissertation is dedicated to my amazing husband Matt and my loving parents Gail and Mike. Without their continued support and encouragement this dissertation would not have been possible. It is also dedicated to the ocean and the algae for being a source of inspiration. ACKNOWLEDGMENTS I would like to express my gratitude to my major professor, Susan Bell, for the guidance and support she has provided throughout my doctoral degree and for helping me expand my knowledge of marine ecology. Clinton Dawes graciously shared his knowledge of marine botany, provided helpful feedback on my work, introduced me to scanning electron microscopy, and was very helpful throughout my degree, for all of which I am grateful. Valerie Paul was a great summer fellowship mentor and I thoroughly enjoyed my summer at SMS, Fort Pierce where I learned a lot. Thanks go to Peter Stiling and Kevin Beach for serving on my committee and their input into my research and manuscripts. Financial support was provided by a Link Foundation/Smithsonian Institution Fellowship, a Tharp Summer Fellowship from USF Biology, a Summer Pre-doctoral Fellowship from USF Graduate School, Fern Garden Club, Orange Blossom Garden Club, and Sigma Xi. Thanks go to many graduate students and post docs for their help and friendship as I pursued this doctorate including: Jenny Peterson, Mike Middlebrooks, Justin Krebs, Kerry Bohl, Theresa Meickle, Katie Basiotis, Amy Erickson, Lesley Porter Baggett, Louise Firth, Stacy Villanueva, Alison Meyers, and Michael Meads. Many people at USF deserve thanks for contributing in a variety of ways to my research or other aspects my graduate student career. These people include: Christine Brubaker, Mary Parrish, Betty Loraamm, Skip Pierce, Frederick Essig, Jim Garey, Jason Rohr, and John Lawrence. Many undergraduates helped me complete this research both in the field and in the lab. Sam Shiver, Jennifer Strykowski, and Natalie McCune all contributed many, many hours, braving cold, heat, and thunderstorms in the field and tedium in the lab. I would like to thank Holly Gray for drawing the first figure in Chapter Three. And lastly, but most importantly, I thank Matt Neilson for his tremendous support and fantastic home-cooked meals. TABLE OF CONTENTS List of Tables ............................................................................................................... iii List of Figures ................................................................................................................ v Abstract ........................................................................................................................vii Chapter One: A Synoptic Review of the Ecology of Rhizophytic Green Algae ................ 1 Rhizophytic Algae and Rhizoids .......................................................................... 1 The Importance of Rhizophytic Algae .................................................................. 3 Rhizophytic Algae, Succession, and Interaction with Seagrasses .......................... 4 Other Community Level Interactions ................................................................... 6 Organization of This Dissertation ....................................................................... 10 Literature Cited .................................................................................................. 11 Chapter Two: Rhizophytic Algal Communities of Shallow, Coastal Habitats in Florida: Components Above and Below the Sediment Surface ....................... 16 Introduction ....................................................................................................... 16 Methods ............................................................................................................. 19 Data analysis .......................................................................................... 23 Results .............................................................................................................. 25 Plant community composition ................................................................ 25 Aboveground thalli ................................................................................. 27 Sediments ............................................................................................... 28 Holdfasts ................................................................................................ 29 Above- and belowground comparisons ................................................... 33 Discussion ......................................................................................................... 35 Plant community composition ................................................................ 35 Above- and belowground components .................................................... 38 Ecological implications .......................................................................... 42 Conclusions ............................................................................................ 43 Literature Cited .................................................................................................. 44 Chapter Three: Regeneration and Recolonization of Rhizophytic Green Algae (Bryopsidales, Chlorophyta) by Vegetative Reproduction in Field Experiments ........................................................................................... 64 Introduction ....................................................................................................... 64 Methods ............................................................................................................. 70 Overview of study design and locations .................................................. 70 Regeneration from manipulated holdfasts ............................................... 71 i Holdfast size and regeneration ................................................................ 72 Recruitment of new plants ...................................................................... 73 Results ............................................................................................................... 75 Regeneration from manipulated holdfasts ............................................... 75 Holdfast size, regeneration, and handling control .................................... 76 Recruitment of new plants ...................................................................... 78 Discussion.......................................................................................................... 82 Summary ................................................................................................ 89 Literature Cited .................................................................................................. 90 Chapter Four: Secondary Metabolites Are Not Used by Rhizophytic Green Algae (Order Bryopsidales) to Inhibit Surface Fouling ................................... 112 Introduction ..................................................................................................... 112 Methods ........................................................................................................... 114 Collection and Handling of Seaweeds ................................................... 114 Surface Extractions ............................................................................... 114 Whole Plant Extractions ....................................................................... 115 Analysis of Extracts .............................................................................. 116 Surface Morphology ............................................................................. 116 Results ............................................................................................................ 117 Discussion........................................................................................................ 118 Literature Cited ................................................................................................ 120 Appendix A: Holdfast-Handling Control Experiment................................................... 128 Appendix B: Scanning Electron Micrograph of Caulerpa prolifera rhizoids ................ 130 ii LIST OF TABLES Table 2.1: The number of rhizophytic algal individuals sampled by date and location (total area = 1.375 m2 at each site) and the number of quadrats in which each species was found ................................................................. 49 Table 2.2: Mean (± SD) % cover of seagrass and number of quadrats with seagrass present ........................................................................................... 50 Table 2.3: Results of one-way analysis of similarity (ANOSIM) pairwise tests on abundance, ash-free dry weight (AFDW), and holdfast volume of rhizophytic algal species at all pairs of sites ................................................. 51 Table 2.4: Organic content (percent of sample weight ± SD) of whole aboveground rhizophytic algal thalli collected at each site ........................... 52 Table 2.5: Mean (± SD) of height (cm), holdfast depth (cm), and holdfast volume (ml) per rhizophytic individual for all algal species present at each site ........ 53 Table 2.6: Sediment organic and carbonate composition (mean percentage of sample dry weight ± SD, n = 3) for the four transect sampling sites in Florida ......................................................................................................... 54 Table 2.7: Organic and carbonate content (percent of sample weight ± SD) of rhizophytic algal holdfasts by site ................................................................. 55 Table 2.8: Pearson correlation of aboveground thallus wet weight (g) and holdfast volume (ml) and of thallus height (cm) with holdfast depth (cm) .................. 56 Table 2.9: Aboveground biomass (g/m2 AFDW) and belowground holdfast volume (ml/m2) data for dominant rhizophytic algal species by site .............. 57 Table 2.10: Comparison of rhizophytic algal thallus (and seagrass short shoot) densities by plant type and location .............................................................. 58 Table 2.11: Comparison of rhizophytic algal/seagrass community dry weight by plant type and location ................................................................................. 59 Table 2.12: Comparison of reported means of rhizophytic algal thallus height, holdfast depth, and holdfast volume by plant type and location .................... 60 iii Table 3.1: Summary of experiments including dates, locations, and essential features ........................................................................................................ 96 Table 3.2: Results of regeneration experiments showing the duration (days) each experiment ran, the number of holdfasts originally planted, the number recovered from undisturbed plots, the percentage of undisturbed holdfasts that regenerated new aboveground material, the mean (± SE) dry weight of new aboveground thallus material generated per holdfast, and the mean (± SE) height of the new material (top edge of the manipulated stipe stub to the uppermost part) for each species ................ 97 Table 3.3: Percent change and holdfast volume (mean ± SE) of tagged individuals from the start to the conclusion of the experiments/handling control by species and thallus manipulation................................................................... 98 Table 3.4: Pearson correlation coefficients (r) and p values from holdfast regeneration experiment on H. incrassata and P. capitatus at Crystal Beach in August 2009 .................................................................................. 99 Table 3.5: The number of individual rhizophytic algal colonizers and the number of plots colonized by species in the summer 2009 regeneration/ recolonization experiments ......................................................................... 100 Table 3.6: Mean (± SE) holdfast volume, height, and aboveground dry weight of individual thalli for each species of colonizer from the experiments at Sunset Beach (28-30 days) and Crystal Beach (22-24 days) in summer 2009 compared with mean values from the natural population in the same area at Sunset Beach sampled on 29 June 2007 .................................. 101 Table 3.7: Number (mean ± SE) of new rhizophytic algal thalli m-2 that recruited into cleared plots over 51 days (June-August 2008) at both sites in the natural recolonization experiment and in the regeneration/ recolonization experiments in June 2009 at Sunset Beach (28-30 days) and in August at Crystal Beach (22-24 days) .............................................. 102 Table 3.8: Braun-Blanquet scores of each species found in plot borders at Fred Howard Park and the number of individuals that recruited into the plots over 51 days....................................................................................... 103 Table 3.9: Braun-Blanquet scores of each species found in plot borders at Sunset Beach and the number of individuals that recruited into the plots over 51 days ....................................................................................................... 104 Table .4.1: Presence or absence of non-polar secondary metabolites in surface and whole tissue extracts as detected in 1HNMR spectra of four rhizophytic green algal species...................................................................................... 123 iv LIST OF FIGURES Figure 2.1: Transect sampling sites in Florida: Sunset Beach (28° 08.72' N, 82° 47.51' W), Mote Tropical Research Laboratory (24° 39.62' N, 81° 27.31' W), Bahia Honda (24° 39.37’ N, 81° 16.80’ W), and Little Duck Key (24° 40.83' N, 81° 13.78' W) ........................................................ 61 Figure 2.2: Aboveground rhizophytic algal biomass as total dry weight and ash- free dry weight by species composition and site/sampling date ..................... 62 Figure 2.3: Rhizophytic algal holdfast volume by species composition at each site/sampling date ......................................................................................... 63 Figure 3.1: Vegetative reproduction in the rhizophytic alga, Penicillus capitatus ......... 105 Figure 3.2A: Mean density of new aboveground thalli of each species that recruited into a single plot (0.04 m-2) of each treatment during over the 28-30 days of the June 2009 experiment at Sunset Beach ........................... 106 Figure 3.2B: Mean density of new aboveground thalli of each species that recruited into a single plot (0.04 m-2) of each treatment during over the 22-24 days of the August 2009 experiment at Crystal Beach ...................... 106 Figure 3.3: Mean (± SE) dry weight (g) of new aboveground thalli and holdfast volume (ml) that recruited into experimental plots during the 28-30 days of the June 2009 experiment at Sunset Beach and the 22-24 days of the August 2009 experiment at Crystal Beach ........................................ 107 Figure 3.4: Mean (± SD) density of new recruits per plot (0.04 m2) of each rhizophytic algal species given Braun-Blanquet (BB) score for that species in borders around plots in 51 day natural recolonization experiment at Fred Howard Park 2008........................................................ 108 Figure 3.5: Mean density of new recruits per plot (0.04 m2) of each species (noted on the x-axis) into those plots with the border Braun-Blanquet score for that species (also given on the x-axis) ................................................... 109 Figure 3.6: Mean length of Caulerpa cupressoides stolon inside a plot (0.04 m2) at Sunset Beach in 51 day natural recolonization experiment ...................... 110 v
Description: