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Characteristics of Vesicomyid Clams and Their Environment at the Blake Ridge Cold Seep W&M ScholarWorks Dissertations, Theses, and Masters Projects Theses, Dissertations, & Master Projects 2005 Characteristics of Vesicomyid Clams and their Environment at the Blake Ridge Cold Seep Taylor Perrine Heyl College of William & Mary - Arts & Sciences Follow this and additional works at: https://scholarworks.wm.edu/etd Part of the Fresh Water Studies Commons, Marine Biology Commons, and the Oceanography Commons Recommended Citation Heyl, Taylor Perrine, "Characteristics of Vesicomyid Clams and their Environment at the Blake Ridge Cold Seep" (2005). Dissertations, Theses, and Masters Projects. Paper 1539626834. https://dx.doi.org/doi:10.21220/s2-vdv4-1352 This Thesis is brought to you for free and open access by the Theses, Dissertations, & Master Projects at W&M ScholarWorks. It has been accepted for inclusion in Dissertations, Theses, and Masters Projects by an authorized administrator of W&M ScholarWorks. For more information, please contact [email protected]. CHARACTERISTICS OF VESICOMYID CLAMS AND THEIR ENVIRONMENT AT THE BLAKE RIDGE COLD SEEP A Thesis Presented to The Faculty of the Department of Biology The College of William and Mary in Virginia In Partial Fulfillment Of the Requirements for the Degree of Master of Science by Taylor Perrine Heyl 2 0 0 4 APPROVAL SHEET This thesis is submitted in partial fulfillment of the requirements for the degree of Master of Science Taylor Perrine Heyl Approved by the Committee, December 2004 Dr CindyNbee Van Dover, Chair Professor, Biology Departmem, College of William and Dr © g e r Mann, Professor, Virginia Institute of Marine Science Dr Randy M. Chambers, Biology Department, College of William and Mary To my father, Captain Phillip J. Heyl, who taught me to accept challenges, and to mother, Karen B. Hunter, who helped me follow them through. TABLE OF CONTENTS Page ACKNOWLEDGEMENTS vi LIST OF TABLES vii LIST OF FIGURES viii ABSTRACT x INTRODUCTION 2 G e o l o g ic a l s e t t in g a n d g l o b a l distribution o f s e e p s 4 B l a k e R id g e se e p 6 Geological setting 6 Biological setting 10 R eproductive s t r a t e g ie s in seep b iv a l v e s 15 METHODS 19 S a m p l in g a n d in it ia l c o r e p r o c e s s in g 19 G e o c h e m ic a l a n d p h y s ic a l a n a l y s e s 2 0 B io l o g ic a l a n a l y s e s 21 S t a t is t ic a l a n a l y s e s 22 RESULTS 2 4 S e d im e n t characterization b y h a b it a t 2 4 G e n e r a l c l a m characteristics 32 C l a m reproductive characteristics 38 General reproductive anatomy and condition 38 iv Page Relationship between reproductive characteristics and the environment 46 DISCUSSION 47 B l a k e R id g e v e s ic o m y id c l a m h a b it a t s 47 Sediment porewater geochemistry 48 V e s ic o m y id c l a m reproductive characterization 52 L ife h is t o r y st r a t e g ie s 53 APPENDIX 57 LITERATURE CITED 58 VITA 76 v ACKNOWLEDGEMENTS I would like to thank my major advisor, Cindy Lee Van Dover, for the incredible opportunity to work in the deep sea, for the chance to see the Blake Ridge seep from Alvin’s viewports and for guidance through this project. I would also like to thank my committee members Randy Chambers and Roger Mann for support and encouragement, especially in the absence of my advisor. Thank you to the Van Dover lab, past and present for emotional and intellectual support, most especially to Rachel Horak for plotting endless Eh graphs at sea and for standing with me in the cold room onboard the R/V Atlantis, Kristen Schmidt for sorting juvenile clams, Megan Ward for electron microscopy, and to Lizzie Blake and Jessica Wallace for their optimism and sense of humor. Thank you to Steve Macko and Bill Gillhooly at the University of Virginia for their expertise in geochemistry and for porewater sulfide and sulfate analyses. Thanks must also go to Linda Meneghini and Heidi Romine from the Virginia Institute of Marine Science for help with grain size analyses, Ken Moore for allowing a deep-sea student into his seagrass lab, teaching me how to take redox potential measurements and loaning instrumentation for use at sea, and to Beth Waterson for patience with deep-sea organic carbon and nitrogen analyses. Thank you to George Gilchrist for help with statistical analyses. Thanks to Tom Mayes, and Kirk and John from the William & Mary Physics machine shop for construction of Alvin pushcores. This work would not have been possible without the Captain and crew of the R/V Atlantis and expedition Leader Pat Hickey. Thank you to the pilots and technicians of Alvin, chief-scientist Carolyn Ruppel, and members of the Windows to the Deep science party for their assistance at sea. And finally, thank you to Peter Kingsley-Smith, who stood by me so long and so well. Note: Sulfide and sulfate porewater data reported herein make up a portion o f the Ph.D. thesis o f W. Gillhooly in Prof. S. Macko's lab at the University o f Virginia, and are used here with their permission as part of a collaboration between the Van Dover and Macko laboratories. T.P. Heyl assisted in all aspects o f the sulfide and sulfate chemical analyses, including collection o f samples, extraction o f porewater fluids, and wet chemistry analyses in the laboratory of Prof. Macko. Mineral sulfide data reported herein was analyzed by Randy Chambers at the College o f William and M ary’s KECK laboratory. Gillhooly, Macko and Chambers will be co-authors on any publication resulting from the work reported in this thesis. vi LIST OF TABLES Table Page 1. Environmental and reproductive variables in vesicomyid and 16 solemyid bivalves in chemosynthetic communities vii LIST OF FIGURES Figure Page 1. Site and sample locations A. Blake Ridge seep site 8 B. Location of pushcores 9 2. In situ photographs of the distribution of vesicomyid clams at Blake Ridge A. Clam bed showing distribution in small circular patches 12 B. Pushcore sampling in a shell bed 13 3. Photos of representative pushcores from 4 microhabitats at Blake Ridge 26 4. Redox potential, sulfide, and sulfate concentration (mM) profiles in representative pushcores from 4 microhabitats at Blake Ridge 27 5. A. Concentrations of porewater and mineral sulfides across all intervals within pushcores from 4 microhabitats at Blake Ridge 30 B. Sulfide retention at the 4 microhabitats at Blake Ridge 31 6. Grain size for sediments sampled from Blake Ridge 33 7. Percent organic carbon and total nitrogen in Blake Ridge sediments 34 8 . Size-frequency distribution of vesicomyid clam shell lengths 35 9. A. Relationship between number of juvenile and adult clams per pushcore 36 B. Relationship between clam density per pushcore and ZEfeSiocm 37 10. Relationship between clam shell length (mm) and EH 2Siocm 40 11. General anatomy of vesicomyid clams from Blake Ridge 41 12. Reproductive anatomy of vesicomyid clams from Blake Ridge A. Female gonad 42 B. Male gonad 43 13. Stages of reproductive condition in vesicomyid clams from Blake Ridge 44 14. Reproductive stages, mean and maximum oocyte diameters in vesicomyid clams from Blake Ridge vs. ZH 2SioCm 45 viii ABSTRACT Vesicomyid clams are one of the dominant megafaunal taxa found in deep-sea cold-seep communities. They rely on chemoautotrophic sulfide-oxidizing symbionts located in their gill tissues. Since their survival depends on a flux of sulfide, clam populations may be viewed as 'flux indicators' within chemosynthetic ecosystems. At the Blake Ridge methane hydrate seep, extensive beds of dead clams are interspersed with smaller (> 1 m diameter) circular patches of live clams or a combination of live clams and shell beds. The patchy distribution of live clams may reflect underlying heterogeneity in sulfide flux or in other environmental parameters that influence the health and reproductive condition of the clams. The research effort included sediment geochemical analyses from 4 distinct microhabitats (background sediments, clam beds, a combination of clam/shell beds, and shell beds) within the seep site. Porewater chemistry indicates sulfide concentrations are five times higher (10.3-14mM) in shell beds than in clam beds (0.1-3.1mM) and this could be the cause of massive clam mortality. Gametogenic characteristics of vesicomyid clams (collected from different microhabitats in July 2003) were investigated but there was no correlation between health of clams and geochemistry within or among microhabitats at the seep. This study suggests that Blake Ridge clams have a broad range of sulfide tolerance that limits local and population-wide extinction of the species but they have a critical sulfide tolerance limit above which mortalities due to toxicity occur. ix CHARACTERISTICS OF VESICOMYID CLAMS AND THEIR ENVIRONMENT AT THE BLAKE RIDGE COLD SEEP INTRODUCTION Cold seeps are found in environments where reduced sulfur and methane emerge from seafloor sediments without an appreciable temperature anomaly. These environments are among the most recently discovered marine habitats; the first seep was discovered just 20 yr ago in 1984, on the Florida Escarpment in the Gulf of Mexico (Pauli et al.
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