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Patterns of Resource Allocation in Caribbean Coral Reef Sponges

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Patterns of Resource Allocation in Caribbean Coral Reef Sponges PATTERNS OF RESOURCE ALLOCATION IN CARIBBEAN CORAL REEF SPONGES Wai Leong A Thesis Submitted to the University of North Carolina Wilmington in Partial Fulfillment of the Requirements for the Degree of Master of Science Department of Biology and Marine Biology University of North Carolina Wilmington 2009 Approved by Advisory Committee ___Richard M. Dillaman__ ____Stuart R. Borrett_____ ____Joseph R. Pawlik____ Chair Accepted by ______________________ Dean, Graduate School TABLE OF CONTENTS ABSTRACT……………………………………………………………………………. ..iv ACKNOWLEDGEMENTS………………………………………………………………vii LIST OF TABLES……………………………………………………………………… .ix LIST OF FIGURES……………………………………………………………………. .. .x CHAPTER 1: FRAGMENTS VERSUS PROPAGULES: REPRODUCTIVE TRADE- OFFS FOR TWO CALLYSPONGIA SPP. FROM FLORIDA CORAL REEFS.................1 ABSTRACT......................................................................................................................2 INTRODUCTION ............................................................................................................3 MATERIALS AND METHODS......................................................................................5 RESULTS .........................................................................................................................8 DISCUSSION...................................................................................................................9 LITERATURE CITED ...................................................................................................12 CHAPTER 2: IS THERE A TRADE-OFF BETWEEN GROWTH AND DEFENSE AMONG CARIBBEAN CORAL REEF SPONGES? ......................................................18 ABSTRACT....................................................................................................................19 INTRODUCTION ..........................................................................................................20 MATERIALS AND METHODS....................................................................................22 RESULTS .......................................................................................................................23 DISCUSSION.................................................................................................................25 LITERATURE CITED ...................................................................................................30 CHAPTER 3: PATTERNS OF RESOURCE ALLOCATION IN CARIBBEAN SPONGES: IS THERE A TRADE-OFF BETWEEN REPRODUCTION AND DEFENSE? ........................................................................................................................39 ABSTRACT....................................................................................................................40 ii INTRODUCTION ..........................................................................................................41 MATERIALS AND METHODS....................................................................................44 RESULTS .......................................................................................................................45 DISCUSSION.................................................................................................................46 LITERATURE CITED ...................................................................................................52 iii ABSTRACT Trade-offs are a common theme in the ecological literature. Organisms allocate resources to physiological functions such as growth and reproduction. When resources are limiting, organisms must selectively allocate their resources, leading to resource allocation trade-offs. Among Caribbean reef sponges, some species produce secondary metabolites that deter predation. Yet, other species that do not produce any chemical defenses co-exist alongside their defended counterparts. Resource allocation trade-offs have been demonstrated to explain the co-existence of undefended and defended species in terrestrial plants. Species either deter predation by allocating resources to defense, or tolerate predation by allocating resources to growth and reproduction. Although previous work has provided some evidence for resource allocation trade- offs between chemical defense, growth and reproduction in Caribbean coral reef sponges, this is the first work measuring growth rates and reproductive output in sponge species for the purpose of comparing between the undefended and defended species most commonly found on Caribbean coral reefs. First, a resource allocation trade-off between growth and propagule production was studied in two undefended congeners with different growth forms. Callyspongia armigera is a branching sponge, whereas Callyspongia vaginalis occurs as a collection of tubes. C. armigera had higher growth rates (0.36 ± 0.31 vs. 0.08 ± 0.11 % initial mass per day), higher number of attachment points (2.31 ± 1.47 vs. 1.03 ± 0.18), and lower propagule production (0.04 ± 0.22 vs. 0.53 ± 1.08 % area of reproductive propagules) compared to C. vaginalis . Branching sponges can disperse by fragmentation, and therefore would allocate fewer resources to propagule production. Results demonstrated a resource allocation trade-off between growth and iv reproduction that is linked with morphology. To reduce complications from morphology- linked resource allocation trade-offs, the remaining resource allocation trade-offs were examined using only branching sponges. Growth rates of undefended sponge species ( Callyspongia armigera, Iotrochota birotulata, and Niphates erecta ) and defended species ( Amphimedon compressa, Aplysina fulva , Aplysina cauliformis and Ptilocaulis walpersi ) were measured using predation exclusion experiments. Growth was greater for undefended than defended sponges (0.89 ± 0.01 vs. 0.77 ± 0.01 % g final g -1 initial day -1). Winter growth was diminished in both undefended and defended sponges compared to summer growth, but less so in undefended sponges than defended sponges (significant season x defense interaction: ANOVA, F = 10.01, df = 1, 1150, p = 0.002). Both comparative growth rates and seasonal patterns of growth support a resource allocation trade-off between growth and chemical defense among Caribbean coral reef sponges. Reproductive output was quantified for six of the above branching species (excluding Ptilocaulis walpersi ), and in the tube sponge Callyspongia vaginalis . Monthly samples were collected for a year, from which histological sections were made for the quantification of reproductive propagules. Propagule production was highly variable among the sponge species. On average, undefended and defended species had the same reproductive output, and a resource allocation trade-off was not found between reproduction and chemical defense. A simultaneous trade-off between growth and propagule formation, such as the one between Callyspongia armigera and C. vaginalis , could be confounding the pattern between propagule formation and chemical defense. Finally, resource allocations to defense, growth and reproduction were consolidated to v form a conceptual model of how resource allocation has influenced the evolution of sponge communities on Caribbean coral reefs. vi ACKNOWLEDGEMENTS None of this work would have been possible without the guidance, feedback and support from my advisor Dr. Joseph Pawlik. My graduate committee, Dr. Richard Dillaman and Dr. Stuart Borrett, provided many useful insights and suggestions. Mark Gay taught me everything I know about histological procedures, and Dr. James Blum was there to untangle my bouts of statistical confusion. I am indebted to my lab mates for being my family away from home, and for all their support and advice. Thanks especially to the new Dr. Timothy Henkel for always being there to answer my questions about life, courses, procedures and statistics. Steven McMurray, Tse-Lynn Loh, David Hines and Michael Echevarria were always happy to bounce ideas off with me. I spent many long hours with the denizens of the Dillaman lab - Carolina Priester, Anne Leaser, Ana Jimenez and Kristen Hardy, who were always a source of encouragement and support. The office ladies, Tracie, Debby, Eleanor, Carol and Lori saved me from being hopelessly entangled in bureaucracy. There are many other folks I’ve met along the way, too many to list by name, who have tirelessly answered my questions and provided technical support. These would include the crew and participants on the R/V Seward Johnson, the folks at NURC, other faculty and students in the department, and other people who work on sponges that I’ve written to for advice and suggestions. Dr. Henry Feddern did an excellent job of sample collection on my behalf. My family has always been a silent but ceaseless fount of love and support for me, and has given me the strength to be who I am. vii This research was funded by grants to Joseph R Pawlik from the National Undersea Research Program at UNCW (NOAA NA96RU-0260) and from the National Science Foundation, Biological Oceanography Program (OCE-0095724, 055468). viii LIST OF TABLES Table Page 1. Details of growth experiments run at North Dry Rocks in Key Largo, Florida. Species: ACO= Amphimedon compressa , ACA= Aplysina cauliformis , AF= Aplysina fulva , CA= Callyspongia armigera , IB= Iotrochota birotulata , NE= Niphates erecta , PW= Ptilocaulis walpersi ; bold species are defended ...............34 2. Table 2: ANOVA results for factors affecting growth of sponges in Key Largo, Florida. Significant factors are marked by an asterisk................................................34
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