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Patterns and Predictions Across Scales A STANDING STOCKS AND FAUNAL ZONATION OF DEEP-SEA BENTHOS: PATTERNS AND PREDICTIONS ACROSS SCALES A Dissertation by CHIH-LIN WEI Submitted to the Office of Graduate Studies of Texas A&M University in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY May 2011 Major Subject: Oceanography Standing Stocks and Faunal Zonation of Deep-Sea Benthos: Patterns and Predictions across Scales Copyright 2011 Chih-Lin Wei STANDING STOCKS AND FAUNAL ZONATION OF DEEP-SEA BENTHOS: PATTERNS AND PREDICTIONS ACROSS SCALES A Dissertation by CHIH-LIN WEI Submitted to the Office of Graduate Studies of Texas A&M University in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY Approved by: Chair of Committee, Gilbert T. Rowe Committee Members, Jay R. Rooker Antonietta Quigg Daniel C. O. Thornton Head of Department, Piers Chapman May 2011 Major Subject: Oceanography iii ABSTRACT Standing Stocks and Faunal Zonation of Deep-Sea Benthos: Patterns and Predictions across Scales. (May 2011) Chih-Lin Wei, B.S., National Chung-Hsing University (Taiwan); M.S., Texas A&M University Chair of Advisory Committee: Dr. Gilbert T. Rowe The deep ocean (> 200-m depth) covers more than 65% of earth’s surface and is known as the largest active carbon sink of the planet. Photosynthesis fixes inorganic carbon into organic rich-compounds to fuel the biological production in the upper ocean. A small portion of the photosynthetic carbon eventually sinks to the seafloor to support diverse deep-sea life. In this dissertation, the phytoplankton production and export flux of particulate organic carbon (POC) to the seafloor were linked to standing stocks and compositional changes of the deep-sea soft bottom assemblages. The pattern and processes of energy transfer from the surface ocean to the deep sea was examined by modeling the global benthic bacteria, meiofauna, macrofauna, and megafauna biomass from remotely sensed ocean color images and the seafloor relief. The analysis was then scaled down to the macrofauna of the Gulf of Mexico (GoM) to examine the global pattern on regional oceanic features with contrasting productivity regimes. These results suggested a universal decline of benthic standing stocks down the continental margins that is caused by an exponential decrease of export POC flux with depth. A revisit of iv historical epibenthic invertebrate sampling in the North Atlantic showed that both individual species and multi-species assemblages occurred in narrow depth bands that hugged the topography from the upper continental slope out to the Hatteras Abyssal Plain. The continuum compositional change suggested that the continuous decline of benthic food supply with depth was the potential driving force for the pattern of bathymetric faunal zonation. A broad, systematic survey across multiple depth transects in the northern GoM suggested that macrofauna zonation is not only taking place across isobaths, but also form the northeast to the northwest GoM due to a horizontal productivity gradient created by the nutrient-laden Mississippi River. Analyses of long- term demersal fish data from 1964 to 2002 in the northern GoM showed no evidence of large-scale faunal change across different sampling times. Base on the pooled data, a shift in rate of fish species replacement may be caused by complex biological interactions or changes in environmental heterogeneity along depth or productivity gradients. v ACKNOWLEDGEMENTS I would like to thank my committee chair, Dr. Gilbert T. Rowe, and committee members, Dr. Jay R. Rooker, Dr. Antonietta Quigg, and Dr. Dan C. O. Thornton, for their guidance, encouragement, and support throughout the course of my dissertation research. Their office doors were always open for me while searching for advice and invaluable discussions. I am grateful to Dr. Fain Hubbard, our formal lab manager and graduate student mentor, who oversaw our day-to-day macrofauna sorting activity. He passed away shortly after completion of the DGoMB project, but his influence has imprinted on the many student researcher s of the deep GoM benthos. Thanks go to Archie Ammons, Clif Nunnally, Lindsey Loughry, Min Chen, Matt Ziegler, Xiaojia Chen, and Yuning Wang for the wonderful time working together at sea and sample sorting in the lab. To Amélie Scheltema, Fain Hubbard, George Wilson, Iorgu Petrescu, John Foster, Mary Wicksten, Min Chen, Roe Davenport, Yuning Wang, and Yousria Soliman for macrofauna identification, and to John McEachran for the deepwater fish identification. I would like to thank my co-authors of publications and manuscripts generated from this dissertation including Gilbert Rowe, Elva Escobar-Briones, Antje Boetius, Thomas Soltwedel, Julian Caley, Yousria Soliman, Falk Huettmann, Fangyuan Qu, Zishan Yu, Roland Pitcher, Richard Haedrich, Mary Wicksten, Michael Rex, Jeffrey Baguley, Jyotsna Sharma, Roberto Danovaro, Ian MacDonald, Clifton Nunnally, Jody Deming, vi Paul Montagna, Mélanie Lévesque, Jan Marcin Weslawski, Maria Wlodarska- Kowalczuk, Baban Ingole, Brian Bett, David Billet, Andrew Yool, Bodil Bluhm, Katrin Iken, Bhavani Narayanaswamy, Amélie Scheltema, George Wilson, Iorgu Petrescu, John Foster, Min Chen, Roe Davenport, Yuning Wang, and Greg Boland. Thanks also go to Bob Carney, Michael Rex, Ron Etter, Paul Snelgrove, and anonymous reviewers for the comments and suggestions on my manuscripts and to Roland Pitcher, Peter Lawton, Nick Ellis, Stephen Smith, Lewis Incze, Michelle Greenlaw, Nicholas Wolff, Tom Shirley, and Paul Snelgrove for valuable discussions on data analysis. Thanks go to the men and women on board the R/V Eastwood, R/V Alaminos and R/V Gyre for their enthusiasm and devotion to explore deep ocean life. The North Atlantic epibenthic invertebrate data was derived from a dissertation at Duke University by Gilbert T. Rowe under the direction of Robert J. Menzies. This dissertation was supported by Minerals Management Service (MMS), now the Bureau of Ocean Energy Management, Regulation and Enforcement (BOEMRE). Thanks go to Erma Lee and Luke Mooney, the Oceanography Department (TAMU) and Marine Biology Department (TAMUG) for conference travel support. Additional funding thanks to Cross-Project Synthesis and Continental Margin Ecosystems (COMARGE) of Census Marine Life (CoML) that was supported by Alfred P. Sloan Foundation. Thanks to my friends, colleagues, department faculty and staff for making my time at the College Station and Galveston campuses an enjoyable memory. Finally, I would like to vii thank my parents, family, friends in Taiwan, and my wife, Shih-Ying for their encouragement, patience and love. viii TABLE OF CONTENTS Page ABSTRACT .......................................................................................................... iii ACKNOWLEDGEMENTS ................................................................................... v TABLE OF CONTENTS....................................................................................... viii LIST OF FIGURES ............................................................................................... x LIST OF TABLES................................................................................................. xiii CHAPTER I INTRODUCTION............................................................................. 1 II GLOBAL PATTERNS AND PREDICTIONS OF SEAFLOOR BIOMASS USING RANDOM FORESTS......................................... 5 2.1. Overview .............................................................................. 5 2.2. Introduction .......................................................................... 6 2.3. Materials and Methods .......................................................... 10 2.4. Results .................................................................................. 20 2.5. Discussion............................................................................. 37 III STANDING STOCKS AND BODY SIZE OF DEEP-SEA MACROFAUNA: A BASELINE PRIOR TO THE 2010 BP OIL SPILL IN THE NORTHERN GULF OF MEXICO ........................... 45 3.1. Overview .............................................................................. 45 3.2. Introduction .......................................................................... 46 3.3. Materials and Methods .......................................................... 52 3.4. Results .................................................................................. 60 3.5. Discussion............................................................................. 82 IV FAUNAL ZONATION OF LARGE EPIBENTHIC INVERTEBRATES OFF NORTH CAROLINA REVISITED........... 95 4.1. Overview .............................................................................. 95 4.2. Introduction .......................................................................... 95 ix CHAPTER Page 4.3. Data Analyses ....................................................................... 97 4.4. Results .................................................................................. 98 4.5. Characteristics of Each Depth Zone....................................... 105 4.6. Discussion............................................................................. 106 V BATHYMETRIC ZONATION OF DEEP-SEA MACROFAUNA IN RELATION TO EXPORT OF SURFACE PHYTOPLANKTON PRODUCTION ................................................................................. 107 5.1. Overview .............................................................................. 107 5.2. Introduction .........................................................................
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