Global Biogeographical Patterns of Plankton in Response to Climate Change
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Global biogeographical patterns of plankton in response to climate change By Ernesto Villarino A thesis submitted to the University of the Basque Country for the degree of DOCTOR OF PHILOSOPHY Thesis Directors Dr. Guillem Chust Dr. Xabier Irigoien Doctoral programme Marine Environment and Resources Department Plant Biology and Ecology University of the Basque Country Year 2016 (c)2017 ERNESTO VILLARINO PRADO The research carried out in this Dr. Philos thesis has been developed in AZTI and it has been financed by: Iñaki Goenaga – Fundacion Centros Tecnologicos (IG –ZTF) with a pre-doctoral training grant for the years 2012-2016 And it has been part of the following projects and co-founders: EURO-BASIN (European Union Basin-Scale Analysis, Synthesis and Integration), funded by the European Commission (Contract No. 264933). http://www.euro-basin.eu/ DEVOTES (DEVelopment Of innovative Tools for understanding marine biodiversity and assessing good Environmental Status) project, funded by the European Union under the 7th Framework Program. http://www.devotes- project.eu/ Malaspina 2010 Circumnavigation Expedition, funded by Consolider- Ingenio 2010,CSD2008-00077. http://www.expedicionmalaspina.es/Malaspina/Main.do Basque Government (Department Deputy of Agriculture, Fishing and Food Policy). A mi madre Please cite this document as: Villarino, E. 2016. Global biogeographical patterns of plankton in response to climate change. PhD Thesis. Department of Plant Biology and Ecology, University of the Basque Country, 223p. Cover illustration: Ion Zubiaurre Ros www.onyon.org. Photo by Sharif Mirshak, from "Plankton - Wonders of the Drifting World" by Christian Sardet http://planktonchronicles.org/en/the-project/ Acknowledgements It`s been fun to work with you Guillem! Despite you like soul and I love jazz, we rock it out! Thanks for helping me every time I sang out of tune. Said that, we played amazing material during these 4 years, both at winter and in summer! Xabier, what a pleasure to have you!! Thanks for the timely interactions that kept spirits high and for reminding me every full moon that it is the courage to continue what it counts for success. Honestly, having you both as directors made it easier to get things done, no matter the challenge we always tried!!! Thanks Javi Franco for that coffee in that bar. Leire Citores, for your time and help, at both rough and calm seas, I wish this only to be the start of many fruitful collaborations; you are a good one, talent all over! Aitor Larrañaga, for your boundless support in times of trouble. Thanks Ionan Marigomez, you are the boss (in a politically correct way).Tor Knutsen, from IMR Norway, for your advice “Keep it simple, just do it!” when we were off Greenland, by the Irminger Sea, at full-storm, approaching the North Pole… Johannes Paetch, Markus Kreus and Jian Su from ZMAW Hamburg, for being always by myside. Fernando Villate, thanks for sharing your passion for science. Oihana Solaun thanks for the last minute big help. Special appreciation to my AZTI colleagues and friends, who made my stage warm and pleasant, and helped me both morally and technically: Carlitos my brother, Maddito, Varona (el cabra), Evita, Neretxu, Corellix, Peter, Antoine, Sonia, Agur, Maite, Pınarbaşı, Basterre, N. Goikoetxea, Itsaso, Julien, Lucho, L.Ibaibarriaga, Bubu, Unaitxo, Lopez, Isa, Blanki, Pecoraro, Sarai, Iraide, Aizkorri, Mireia, Nagores, Ane, María, Curtin, Axelle, Marine, Fermin… También agradezco el apoyo del mejor osteópata de la noche, el gran Dani Irusta, brindemos por esas tantas olas que nos quedan por pillar, bro! A esos dos personajes de Elgoibar que tanto me quieren, gracias brothers. A ti, por esas noches de insomnio, por esos tantos rooibos, por esos paseos noctámbulos. Gracias por aguantarme durante el último tercio de esta tesis, tanto en la lluvia como al sol. Pero sobre todo agradezco a mi familia, a Patricia, al aita, y a la ama. La familia telerín. Por creer en mí y apoyarme incondicionalmente en todo momento para conseguir todas esas metas que me he propuesto. Por empujarme a explorar nuevos océanos. Ahora empieza el rock’n roll, let’s get the party started! Table of contents SUMMARY ....................................................................................................................................................... 1 RESUMEN ......................................................................................................................................................... 5 0. GENERAL INTRODUCTION ..................................................................................................................... 11 0.1 PLANKTON BIOGEOGRAPHY AND GENERAL CONCEPTS .............................................................................................. 11 0.1.1 Zooplankton as sentinels of climate change ....................................................................................... 13 0.1.2 Methods to study zooplankton distribution ........................................................................................ 14 0.2 SPECIES ECOLOGICAL NICHE............................................................................................................................... 16 0.2.1 Niche-theory ....................................................................................................................................... 16 0.2.2 Species Distribution Models ................................................................................................................ 19 0.3 COMMUNITY STRUCTURE AND MARINE BIODIVERSITY .............................................................................................. 24 0.3.1 Dispersal and connectivity .................................................................................................................. 24 0.3.2 Genetic flow ........................................................................................................................................ 27 0.3.3 Community assembly .......................................................................................................................... 28 0.3.4 Neutral theory of biodiversity ............................................................................................................. 31 0.4 CLIMATE CHANGE THREATS ................................................................................................................................ 33 0.4.1 Climate change impacts on marine organism and ecosystems .......................................................... 34 0.4.2 Climatic scenario for the XXI century .................................................................................................. 38 0.5 HYPOTHESIS AND OBJECTIVES ............................................................................................................................. 39 0.6 THESIS STRUCTURE .......................................................................................................................................... 42 0.7 LIST OF PUBLICATIONS AND ORAL COMMUNICATIONS .............................................................................................. 43 PART I: CLIMATE CHANGE AND HABITAT MODELLING OF PLANKTON ......................... 47 1. CHAPTER 1: “LONG-TERM TRENDS OF ZOOPLANKTON AND SEA WARMING ACROSS THE NORTH ATLANTIC AND MEDITERRANEAN SEA” .......................................... 49 1.1 INTRODUCTION ............................................................................................................................................... 51 1.2 METHODS ...................................................................................................................................................... 53 1.2.1 Study sites ........................................................................................................................................... 53 1.2.2 Sea temperature time-series .............................................................................................................. 55 1.2.3 Species by species analysis ................................................................................................................. 55 1.2.4 Community analysis ............................................................................................................................ 60 1.2.5 Correlations of species turnover with time and environmental predictors......................................... 61 1.3 RESULTS ........................................................................................................................................................ 62 1.3.1 Temperature trends ............................................................................................................................ 62 1.3.2 Species by species analysis ................................................................................................................. 63 1.3.3 Community analysis ............................................................................................................................ 63 1.4 DISCUSSION .................................................................................................................................................... 69 2. CHAPTER 2: “MODELLING THE FUTURE BIOGEOGRAPHY OF NORTH ATLANTIC ZOOPLANKTON COMMUNITIES IN RESPONSE TO CLIMATE CHANGE” .......................... 75 2.1 INTRODUCTION ..............................................................................................................................................