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The Pennsylvania State University the Graduate School Department of Biology the ECOLOGY of SEEP COMMUNITIES in the GULF of MEXIC

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The Pennsylvania State University the Graduate School Department of Biology the ECOLOGY of SEEP COMMUNITIES in the GULF of MEXIC The Pennsylvania State University The Graduate School Department of Biology THE ECOLOGY OF SEEP COMMUNITIES IN THE GULF OF MEXICO: BIODIVERSITY AND ROLE OF LAMELLIBRACHIA LUYMESI A Thesis in Biology by Erik E. Cordes Copyright 2004 Erik E. Cordes Submitted in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy December 2004 The thesis of Erik E. Cordes was reviewed and approved* by the following: Chuck Fisher Professor of Biology Thesis Advisor Chair of Committee Katriona Shea Assistant Professor of Biology Peter Hudson Willaman Professor of Biology Michael Arthur Professor of Geosciences Doug Cavener Professor of Biology Head of the Department of Biology *Signatures are on file in the Graduate School iii ABSTRACT Cold seeps are common habitats along the continental margin in all the world’s oceans. In the Gulf of Mexico, they occur in the salt dome province of the upper Louisiana slope, and along the base of the continental rise from Florida to Texas. Some of the most common inhabitants of cold seeps are vestimentiferan tubeworms which are entirely reliant on internal sulfide-oxidizing chemoautotrophic symbionts for their nutrition. The most common vestimentiferan tubeworm of the upper Louisiana slope is Lamellibrachia luymesi. This, and other species of tubeworms, form aggregations of hundreds to thousands of individuals which harbor a diverse community. In this study, a total of 40 tubeworm aggregation and mussel bed samples containing at least 171 macrofaunal species were collected at seeps from 520 to 3300 m depth. The upper Louisiana slope communities progress through a predictable sequence of successional stages. The youngest aggregations contain high biomass communities dominated by endemic species, with biomass decreasing over time as the relative abundance of non- endemic fauna in upper trophic levels increases. This process is mainly driven by the abundance of hydrogen sulfide in the epibenthic layer. Models support the hypothesis that L. luymesi alters its environment by releasing the sulfate generated by its internal symbionts into deeper sediment layers. This alters the distribution of sulfide leading to declines in sulfide concentration among the tubeworm tubes. The combination of these lines of evidence support the assertion that L. luymesi is a significant ecosystem engineer at hydrocarbon seeps in the Gulf of Mexico. iv TABLE OF CONTENTS LIST OF FIGURES .....................................................................................................vi LIST OF TABLES.......................................................................................................viii ACKNOWLEDGEMENTS.........................................................................................ix Chapter 1 Introduction and A Review of Gulf of Mexico Cold Seep Ecology ..........1 Introduction...........................................................................................................1 Gulf of Mexico Cold Seep Ecology......................................................................5 Geological Setting ................................................................................................5 Symbiotic Fauna ...................................................................................................14 Tubeworms....................................................................................................15 Mussels..........................................................................................................18 Clams.............................................................................................................20 Iceworms .......................................................................................................22 Community Ecology.............................................................................................23 Upper Slope Communities ............................................................................24 Deep Seep Communities ...............................................................................32 Conclusions...........................................................................................................34 Chapter 2 High Hydrogen Sulfide Demand of Tubeworm Aggregations Modifies the Chemical Environment at Deep-Sea Hydrocarbon Seeps ..............................36 Abstract..........................................................................................................36 Introduction ...................................................................................................37 Methods .........................................................................................................39 Results ...........................................................................................................43 Discussion......................................................................................................50 Chapter 3 Modeling the Mutualistic Interactions Between Tubeworms and Microbial Consortia..............................................................................................58 Abstract..........................................................................................................58 Introduction ...................................................................................................58 Methods .........................................................................................................61 Population growth model .......................................................................62 Geochemical setting ...............................................................................67 Model implementation ...........................................................................73 Results and Discussion..................................................................................79 Chapter 4 Succession of Hydrocarbon Seep Communities Associated with the Long-Lived Foundation Species Lamellibrachia luymesi....................................88 v Abstract..........................................................................................................88 Introduction ...................................................................................................89 Methods .........................................................................................................92 Results ...........................................................................................................98 Discussion......................................................................................................107 Chapter 5 Impact of an Ecosystem Engineering Tubeworm on the Habitat Characteristics and Community Structure of Gulf of Mexico Hydrocarbon Seeps.....................................................................................................................119 Abstract..........................................................................................................119 Introduction ...................................................................................................120 Methods .........................................................................................................123 Results ...........................................................................................................128 Discussion......................................................................................................134 Chapter 6 Community Structure of Gulf of Mexico Cold Seeps: Do General Theories of Deep-sea Ecology Pertain to Chemosynthetic Ecosystems?.............141 Abstract..........................................................................................................141 Introduction ...................................................................................................142 Materials and Methods ..................................................................................146 Results ...........................................................................................................152 Discussion......................................................................................................159 Bibliography ................................................................................................................169 Appendix......................................................................................................................192 vi LIST OF FIGURES Figure 2-1: Population growth characteristics of Lamellibrachia luymesi: Mortality rate ........................................................................................................44 Figure 2-2: Population growth characteristics of Lamellibrachia luymesi: Recruitment rate of aggregation BH7...................................................................45 Figure 2-3: Model output for Lamellibrachia luymesi aggregation BH7....................46 Figure 2-4: Size-frequency of Lamellibrachia luymesi in aggregation BH7...............49 Figure 2-5: Hydrogen sulfide uptake rate for whole Lamellibrachia luymesi aggregations..........................................................................................................50 Figure 3-1: Model construction ...................................................................................62 Figure 3-2: Growth model for L. luymesi. ...................................................................65 Figure 3-3:Concentration profiles of sulfate, sulfide, and dissolved organic carbon (DOC) ...................................................................................................................69
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