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Distribution and Abundance of Larvaceans in the Southern Ocean

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Distribution and Abundance of Larvaceans in the Southern Ocean Distribution and abundance of Larvaceans in the Southern Ocean By Margaret Caroline Murray LINDSAY BSc. (Env. Mang. & Eco) Hons. GCAS Submitted in fulfilment of the requirements for the Degree of Doctor of Philosophy University of Tasmania June 2012 _________________________ _______ _____ _ Declaration of originality______________________ This thesis contains no material which has been accepted for a degree or diploma by the University or any other institution, except by way of background information and duly acknowledged in the thesis, and to the best of my knowledge and belief no material previously published or written by another person except where due acknowledgement is made in the text of the thesis, nor does the thesis contain any material that infringes copyright. Margaret CM Lindsay June 2012 ___ ____________________________ ______ _____ Statement of authority of access___ _____________ This thesis may be made available for loan and limited copying in accordance with the Copyright Act 1968. Margaret CM Lindsay June 2012 _______________________________ _ ________ _ Statement regarding published work contained in the thesis___ __ _______ ___ The publishers of the papers comprising Chapter 5 hold the copyright for that content, and access to the material should be sought from the respective journals. The remaining non published content of the thesis may be available for loan and limited copying in accordance with the Copyright Act 1968. Margaret CM Lindsay June 2012 ii _______________________________ _ _________ Statement of co-authorship____ ___ ___________ The following people and institutions contributed to the publication of the work undertaken as part of this thesis: Chapter 5: Distribution and abundance of Larvaceans in the Southern Ocean between 30° and 80°E. Published in Deep-Sea Research II 57 (2010) 905–15 Margaret Lindsay (candidate and first author) (70 %) and Guy Williams (co- author) (30 %) of the Institute of Low Temperature Science, Hokkaido University, Sapporo, Japan. The author’s roles were the candidate, Margaret Lindsay, developed the idea, completed the papers formalisation and development, completed the sampling and data analysis and also completed the refinement and presentation. Guy Williams (co-author) contributed to the papers formalisation and development and assisted with refinement and presentation. We the undersigned agree with the above stated “proportion of work undertaken” for the above published peer-reviewed manuscripts contributing to this thesis. Prof Thomas Trull Craig Johnson Supervisor Head of School Institute for Marine and Antarctic Studies Institute for Marine and Antarctic Studies University of Tasmania University of Tasmania September 2011 iii ______________________________ _ ___________ Abstract___________________________ _________ Larvaceans (also known as appendicularians) are zooplankton that inhabit most oceans, coastal waters and estuaries, and are often found in abundances that are second only to copepods among the meso-zooplankton. They form a gelatinous “house” through which they circulate seawater to filter very small particles onto mucopolysaccharide mesh. This concentrates sub-micron and micron-sized protists to 100 -1000 times the ambient concentrations. The house is discarded when the mesh clogs which can occur several times per day. These contribute significantly to the large particles of marine detritus known as “marine snow”. In addition, larvaceans form tightly compacted, rapidly sinking faecal pellets that contribute to carbon “export” from surface waters. Thus larvaceans play a rather unusual role in marine microbial food-webs, by transferring matter across many orders of magnitude in organism size, and moving it into the ocean depths. Previously there was little known about larvaceans in the Southern Ocean. The aims of this study were to increase the knowledge of the ecological role of the larvaceans species: by first determining their distribution and abundance in the Sub-Antarctic Zone (SAZ), Southern Oceans Permanently Open Ocean Zone (POOZ) and the Sea Ice Zone (SIZ) and secondly, by determining the diet of Southern Ocean larvaceans. Larvaceans were collected during four Southern Ocean marine science voyages between 2006 and 2008 which surveyed the different zones through different seasons in the East Antarctic sector of the Southern Ocean. Larvaceans were collected using a number of sampling devices; a purpose-built ring net, Rectangular Mid-Water Trawl (RMT1), Working Party 2 (WP2) net, HYDRO- BIOS MultiNet, Visual Plankton Recorder (VPR) and the Continuous Plankton Recorder (CPR). Larvacean distributions were complex. A significant fraction of the net hauls (55%) contained no specimens, while others obtained high abundances (maximum iv 57.8 ind. m-3). The average abundances of larvaceans from the variety of nets used were 1.4 ± 5.4 ind. m-3, and for the CPR were 6.4 ± 29.7 ind. m-3. The surveys revealed that for the period of this study larvacean abundances varied between the Southern Ocean zones with the lowest abundances in the SAZ (0.6 ± 2.6 ind. m-3, maximum = 15.9 ind. m-3), highest in the POOZ (2.8 ± 10.6 ind. m-3, maximum = 57.8 ind. m-3) and 1.4 ± 4.8 ind. m-3 (maximum = 49.7 ind. m-3) in the SIZ. Possible controls on larvacean distributions were evaluated by comparing them to physical (latitude, longitude, water temperature, salinity, light and sea-ice) and biological (chlorophyll and total zooplankton) distributions. Significant correlations occurred with physical parameters of latitude, longitude, fluorescence, irradiance, water temperature and salinity and the biological distributions of other Southern Ocean zooplankton. The diet of the Southern Ocean larvacean was determined using scanning electron microscopy (SEM) and stomach dissections from the samples collected from the ring net during January to March 2006. Protists ranging from 3.5 µm – 240 µm were the main food items. By considering the alignment and sizes of the largest ingested protist, Corethron pennatum, the inferred feeding house mesh size was 5 - 82 µm. From this study Southern Ocean larvaceans were estimated to contribute ~10.5 million tonnes of (wet) biomass to the Southern Ocean. Comparing this to the ~250 million tonnes of copepods, which are considered to be a keystone taxon, emphasizes the need to consider the importance of larvaceans to Antarctic food webs. Key words___________________________________________ _____________ Larvaceans, Appendicularians, Southern Ocean, Sub-Antarctic Zone (SAZ), Permanent Open Ocean Zone (POOZ), Sea Ice Zone (SIZ), Diet, Biomass, Carbon contribution. v ______________________________ ____________ Acknowledgements___________________________ A huge thanks is required for my supervisors Professor Tom Trull, Dr Graham Hosie and Dr Simon Wright – this would not have been possible without your guidance. Thankyou. A high level of appreciation goes to the following organisations for the logistical support provided by the Australian Antarctic Division (AAD), Kingston; Institute for Marine and Antarctic Studies (IMAS) (previously the Institute of Antarctic and Southern Ocean Studies (IASOS)), Hobart; University of Tasmania (UTas), Hobart; and the Antarctic Climate and Ecosystems Cooperative Research Centre (ACE CRC), Hobart. The officers, crew and cadets on RSV Aurora Australis and TS Umitaka Maru, the Marine Science Support at AAD – especially Aaron Spurr, Tony Veness, Alan Poole Marine Science Voyage management and my fellow expeditioners, require special mention as without their support the successful voyages that made this a successful doctorate, would not have happened let alone been as enjoyable as they were. The officers, crew and cadets and CEAMARC researchers on the TS Umitaka Maru, especially Graham Hosie, Russ Hopcroft, Dhugal Lindsay, Andrea Walters, Jean Henri and Veronica Feuntes. This voyage was made possible thanks to the International Antarctic Institute and ACE CRC travel funding. This study would have been impossible without financial support from an Australian Postgraduate Award (UTas) and Antarctic Climate and Ecosystems Cooperative Research Centre (ACE CRC) Top-Up. The project was also supported in kind by the Australian Antarctic Division. Thanks also goes to other key academics and scientist for their support, including, Mark Curren, Patti Virtue, Guy Williams, Karen Westwood, Steve Candy, Simon Wotherspoon, Imojen Pierce, Russ Hopcroft, Brian Grifiths, Will Howard, Andrew Moy, Donna Roberts, David McLeod, Gerry Nash, Fiona Scott, Rob King and all those that have assisted in all the ways they could. Finally, though far from the least, my friends and relatives are thanked for remaining both friendly and related throughout the PhD journey. Especially those that were involved with insuring I remained sane enough to continue the PhD journey including my mum, Alice and my best friend and partner, Aaron, fellow students and work colleagues and all those that I have shared many adventures and cups of tea, coffee and chai with. The deepest thanks to each and every one of you. Margaret CM Lindsay June 2012 vi “Did you know the word plankton comes from the Greek for ‘born to wander’?” he said. I owned up to my ignorance. “I think that is why I love the little things. They’ve got the lifestyle I’d like to adopt: wandering the seas.” ‘Plankton, huh?’ “Sure! Give me the squidgy goey things every time. Whales get all the glory. I like the proletariat of the food chain, those amorphous gelatinous things that everybody ignores.
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