Zoogeographic and Richness Patterns in Southern Ocean Benthos
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Zoogeographic and richness patterns in Southern Ocean benthos A thesis submitted in accordance with the requirements of the Open University for the degree of Doctor of Philosophy By Huw James Griffiths (BSc hons) 31/03/2010 Director of studies: Dr D.K.A. Barnes Supervisors: Dr K. Linse & Prof L.S. Peck Sponsoring Establishment: British Antarctic Survey High Cross Madingley Road Cambridge CB3 0ET United Kingdom i ABSTRACT This thesis describes the large scale biogeographic patterns found in the Southern Ocean benthos. Using SOMBASE and SCAR-MarBIN, the two most comprehensive, georeferenced databases of Antarctic marine biodiversity ever compiled, a range of taxa were investigated but focusing on the Mollusca, Bryozoa and Pycnogonida. Over 8,000 species of marine invertebrates from over 5,000 sites constituting ~34,000 records were used in the analyses. The strong faunal links between the Antarctic and South America were confirmed but I found little evidence for a biogeographical relationship between the Antarctic or South America and New Zealand or Tasmania. Regional levels of Southern Ocean endemism proved the influence of the Antarctic Circumpolar Current upon the distribution of Southern Ocean benthos. My study shows the Southern Ocean as a ‘single functional unit’ with no evidence for an earlier proposed biogeographical split between East and West Antarctica. Some general rules on Antarctic benthic biogeography are viable, including species endemism rates of around 50% and a definite distinction between the sub-Antarctic islands influenced by either South America or by New Zealand. In the context of potential shifts in species distribution with climate change I investigated the current ranges of selected Southern Ocean taxa (Mollusca, Amphipoda, Ophiuroidea and Hexacorala), and looked for hotspots of coinciding northern and southern geographic range limits. Southern Patagonia, South Georgia and Kerguelen had the greatest range-limit hotspots. Monitoring range shifts in these key places and taxa will enable us to track the influence and effects of climate change on benthic species distributions in the Southern Ocean. ii ACKNOWLEDGEMENTS It was whilst bobbing about in the JCR somewhere on the Scotia Sea in 2006 that Dave Barnes offered to supervise me for a PhD, all I had to do was find a second supervisor. I was lucky enough to find two willing volunteers in the shape of Katrin Linse and Lloyd Peck. Three supervisors meant that at least one of them would be in the same country as me at any one time! Having spent the best part of ten years working for the British Antarctic Survey I probably have far too many people to thank than could ever fit onto a single page. However, very special thanks go to Andy Clarke who has been invaluable with his advice and help along the way, thanks to Hoc Ewing for making me smile and thanks to Cath Waller some important lessons. I would like to thank my various co-authors; Claudia Arango, Stefanie Kaiser, Sandra McInnes and Tomás Munilla. I would like to thank the Captain and crew of the RRS James Clark Ross and those who have made my trips to Antarctica so memorable including Claire Allen, Peter Enderlein, Jan Strugnell, Jenny Rock, Anna Hey, Rachel Malinowska, Hilary Blagbrough and Chester Sands. Thanks to Tom Jordan for being the perfect officemate. I have been lucky enough to be involved with the Census of Antarctic Marine Life and SCAR-MarBIN which have opened many doors for me and introduced me to some really good friends, including my Belgian–brother Bruno Danis and the ultra-organized Vicki Wadley. I would also like to thank all the researchers who published or contributed unpublished data used to create the database and all those people who helped me get it into the computer. Finally, thanks to my family and friends who have put up with me jetting off all over the world and especially to Steve for his patience. iii CONTENTS ABSTRACT ...................................................................................................................... II ACKNOWLEDGEMENTS ........................................................................................... III CONTENTS ..................................................................................................................... IV LIST OF FIGURES ........................................................................................................ IX GENERAL INTRODUCTION ........................................................................................ 1 Biogeography .............................................................................................................. 1 Assessing biodiversity and biogeography ................................................................... 2 Global biogeographic patterns .................................................................................... 3 The Southern Ocean .................................................................................................... 4 CHAPTER 1 - ANTARCTIC MARINE BIODIVERSITY – WHAT DO WE KNOW ABOUT THE DISTRIBUTION OF LIFE IN THE SOUTHERN OCEAN? ............................................................................................................................ 7 Introduction to the region ............................................................................................ 7 Biological structure of the Southern Ocean .............................................................. 15 Research and species discovery in Antarctica .......................................................... 17 Sources of data for analysis ....................................................................................... 19 Sampling intensity ..................................................................................................... 22 Which taxa have been adequately sampled and which have not? ............................. 23 Known and Unknown ............................................................................................... 29 Major threats to biodiversity ..................................................................................... 30 Potential and priorities for future discovery and research in region ......................... 36 iv CHAPTER 2 - DISTRIBUTION OF MACROBENTHIC TAXA ACROSS THE SCOTIA ARC, SOUTHERN OCEAN. ......................................................................... 39 INTRODUCTION.......................................................................................................... 39 METHODS .................................................................................................................. 41 Study area .................................................................................................................. 41 Sample collection ...................................................................................................... 42 RESULTS ................................................................................................................... 46 DISCUSSION .............................................................................................................. 59 Richness .................................................................................................................... 59 Abundance and biomass ............................................................................................ 62 CHAPTER 3 - QUANTIFYING ANTARCTIC MARINE BIODIVERSITY: THE SCAR-MARBIN DATA PORTAL. ..................................................................... 67 INTRODUCTION.......................................................................................................... 67 Antarctic biological diversity: what we know so far ................................................ 67 The creation of SCAR-MarBIN ................................................................................ 70 METHODS .................................................................................................................. 71 Selection and handling of data .................................................................................. 72 RESULTS ................................................................................................................... 73 Patterns within individual phyla ................................................................................ 77 DISCUSSION .............................................................................................................. 85 Geographical distribution of sample stations ............................................................ 85 The effect of sampling intensity ................................................................................ 88 Patterns for individual taxonomic or ecological groups ........................................... 91 Linking the pelagic and benthic realms .................................................................... 92 v FUTURE PERSPECTIVES .............................................................................................. 95 CHAPTER 4 - TOWARDS A GENERALIZED BIOGEOGRAPHY OF THE SOUTHERN OCEAN BENTHOS. ................................................................................ 99 INTRODUCTION.......................................................................................................... 99 MATERIALS AND METHODS .................................................................................... 102 Biogeographical regions and data ..........................................................................