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Dissertation Torben Riehl A PHYLOGENETIC AppROACH TO THE CLASSIFICATION OF MACROSTYLID ISOPODS AND FAUNAL LINKAGES BETWEEN THE DEEP SEA AND SHALLOW-WATER ENVIRONMENTS Dissertation zur Erlangung des Doktorgrades im Department Biologie, der Fakultät für Mathematik, Informatik und Naturwissenschaften der Universität Hamburg vorgelegt von Torben Riehl aus Geesthacht Hamburg, 2014 Acknowledgements It is one of the enjoyments in the final period of writing up a thesis to look back over the past years and remember all the colleagues, friends and family who have helped and supported me along this road. It was long and tedious, but exceptionally enjoyable thanks to them! I would like to express my gratitude to everyone who has supported, encouraged and inspired me whether on expeditions, and excursions, in meetings, at coffee and lunch breaks, conferences, over a pint in a pub, or at the Kicker table, and thus made this period. More specifically, my thanks begin with my main supervisor, Angelika Brandt (University of Hamburg), for her support, and for the “long lead” that she used to guide me – no matter on which con- tinent or ocean I was. Thanks for introducing me to the world of science and inspiring me with the see- mingly endless energy you spawn and invest in order to achieve what you aim for. I would like to express my heartfelt gratitude especially for your trust in me as well as your patience. Furthermore, I am very grateful to Pedro Martínez Arbizu (Senckenberg, German Center for Marine Biodiversity Research – DZMB) for taking on the second supervision, for providing advice, support and constructive criticism, whenever needed. You gave me the great opportunities, for example to participate in the DIVA 3 cruise, and to study isopod DNA at the Smithsonian National Museum of Natural History, and I am much appreciative for this. Exceptional gratitude is owed to George D. F. (Buz) Wilson (Australian Museum) for great colla- boration and hospitality at the Australian Museum in Sydney. I undoubtedly have benefited hugely from his effort getting me Downunder to work with him, and his enthusiasm introducing me to numerous re- searchers and even more isopods, which has resulted in some great ideas, discussions and opportunities! You inspired me to always delve a bit deeper and the barbies at your place were brilliant. Thanks, mate! Special thanks is due to Stefanie Kaiser. For several years, she was a fantastic lab mate and has become a true friend. She participated significantly in organizing my sometimes confused ideas about this thesis. She has greatly assisted me in writing up and helped me focus. I am grateful this and for your thoughtful and detailed comments. This thesis was made possible with the support of many colleagues, friend, and family members and funding from the German National Academic Foundation (Studienstiftung des deutschen Volkes) for which I am most thankful. The CeDAMar taxonomic exchange program, Foundation of the University of Hamburg, the Geddes Fellowship of the Australian Museum, as well as the Studienstiftung helped me fund several inter-continental trips to participate in conferences or visit laboratories and natural-history collections. I appreciate the “pharmacological“ support by Marina Malyutina and Simon Bober that kept me alive during these last days. Thanks to Jörundur Svavarsson (University of Iceland), T. Chat Walther and Amy Driskell (Smithsonian) for generous hospitality, extraordinary samples, recommendations and discussions as well as friendliness and good mood. Without Nils Brenke, none of our joint expeditions would have been half as amusing. Without the gorgeous crews of the research vessels Polarstern, Meteor, and Sonne, I would not have accomplished any of my work. All the collection managers and peers that contributed to my thesis and papers are kindly acknowledged. During the last three years, I was affiliated with three laboratories: “Niedere Tiere II” at the Zoo- logical Museum Hamburg, the German Center for Marine Biodiversity Research (Senckenberg) and the Marine Invertebrates at the Australian Museum. All colleagues I have been involved with have been in- viting and supportive in allowing me to participate in lab activities whilst pursuing my PhD studies. And thanks to you I never ran out of coffee. In this context, it is my pleasure to emphasize the contributions of Saskia Brix-Elsig, Karen Jeskulke, Martina Vortkamp, Sarah Schnurr, Sabine Holst, Antje Fischer, Sven Hoffmann, Karin Meißner, Jasmin Renz, Petra Wagner, and Kathrin Philipps Bussau in Hamburg, Anna Murray, Steven Keable, Shane Ahyong, Frank Köhler, Eunice Wong, and Francesco Criscione in Sydney. A warm thank you to my friends Arne, Philipp, Steffi and Steffi, Jörg and Julia, Maria, Tim, Gerald, Stefan and Benjamin for their invaluable friendship, patience with my enduring absence and help with difficult decisions. I would not have attempted this road if not for my parents, Monika and Peter, my granddad Otto, as well as my sister Annika who implanted within me a love of creative pursuits and scientific endeavor and technical understanding, which drove me into becoming a biologist. Last but not least, I owe my current sanity, health and happiness to the caring support of my partner, Jasmin. She was key witness to the buzzing brain and immense time and energy that I invested in to completing this thesis. Her love, humor and wisdom, have taught me to see the wider picture and what life really is about; taught me about confidence and being content. I don’t want to miss one day! For sharing passions with me, kindling dreams and encouraging me to keep going, I thank you. TABLE OF CONTENTS Summary . 11 Zusammenfassung. 13 . Chapter 1: General Introduction . 15 . The deep-sea environment . 16 The deep sea and the importance of alpha taxonomy and phylogeny . 17 Age and origin of the deep-sea fauna . 17 Isopod crustaceans and deep-sea colonizations . 18 Deep-sea asellote isopods . 18 Macrostylidae . 19 Origin and colonization of the continental shelves . 19 Aims . 20 Content . 20 References . 21 Chapter 2: New Macrostylidae Hansen, 1916 (Crustacea: Isopoda) from the Gay Head-Bermuda transect with special consideration of sexual dimorphism . 27 Abstract . 28 Introduction . 29 Material and Methods . 29 Implicit Attributes . 30 Systematics . 35 Macrostylis dorsaetosa n. sp. .35 Macrostylis papillata n. sp. 42 Discussion . .51 Sexual dimorphism and terminal male stages . .51 Ecological and evolutionary implications . .52 Implications for future systematic work . 54 Acknowledgements . 54 References . 55 Author contributions . 57 Chapter 3: Conquered from the Deep Sea? A New Deep-Sea Isopod Species from the Antarctic Shelf Shows Pattern of Recent Colonization . 59 Abstract . 60 Introduction . 61 Material and Methods . 62 Systematics . 69 Macrostylis roaldi n. sp. 70 Development . 83 Molecular Results . .83 Discussion . 84 Morphological Affinities . 84 Developmental and Reproductive Notes . 84 Distribution. 85 Genetic Structure. 86 Evidence for Shelf Refuges? . 87 Conclusions . 89 Acknowledgements . 89 References . 89 Author contributions . 94 Chapter 4: Southern Ocean Macrostylidae reviewed with a key to the species and new descriptions from Maud Rise . 97 Abstract . 98 Introduction . 99 Material and methods . .100 Systematics . .104 Macrostylis scotti n. sp. 108 Macrostylis matildae n. sp. 118 Macrostylis antennamagna Riehl & Brandt, 2010 . 131 Macrostylis cerrita Vey & Brix, 2009 . 131 Key to the Southern-Ocean species of Macrostylis . 132 Macrostylis gerdesi (Brandt, 2002) . 135 Macrostylis obscura (Brandt, 1992) nom. dub. 135 Macrostylis sarsi Brandt, 1992 nom. dub. 136 Macrostylis setulosa Mezhov, 1992 . 136 Macrostylis uniformis Riehl & Brandt, 2010 . 138 Macrostylis vinogradovae Mezhov, 1992 . 138 Molecular results . 140 Discussion . .140 Characters used to distinguish species in the key . 141 Development and sexual dimorphism . 142 Molecular phylogeny. 142 Acknowledgements . .143 References . .144 Author contributions . .147 Chapter 5: Urstylidae – a new family of abyssal isopods (Crustacea: Asellota) and its phylogenetic implications . 149 . Abstract . .150 Introduction . .151 Material and Methods . .151 Results . .154 Character Conceptualization . 154 Systematics. 163 Urstylidae Fam. Nov.. 163 Urstylis gen nov. 165 Urstylis zapiola gen. et sp. nov. 166 Urstylis solicopia sp. et gen. nov. 173 Urstylis thiotyntlus gen. et sp..
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