A revision of Nereidinae (Polychaeta, Nereididae) Torkild Bakken Dr. scient. thesis Section of Natural History Norwegian University of Science and Technology Trondheim 2004 ISBN 82-471-6205-9 ISBN 82-471-6204-0 (electronic) Preface A copy of ”Identification of Annelida Polychaeta from northern European and adjacent Arctic waters” is a document of personal historic interest. Across the pages are scribbled short annotations beside several polychaete families, and two are circled, Sphaerodoridae and Nereididae. The annotations were made in the office of Torleif Holthe at the Directorate of Nature Management early in 1994, and marked the start of my cand scient education a few months later. I have much to thank Torleif for as my supervisor during my cand scient and dr scient education, he has taught me a lot about polychaetes especially relating to northern areas and a lot about general marine biology, the numerous long discussions we have had, often about completely different topics than polychaetes is much appreciated. He has always been helpful and supportive, and put in a lot of effort when it came to applications and job opportunities. I would like to thank Jon-Arne Sneli for passing along all that marine biology knowledge, not only as my supervisor during the cand scient period but also later. I have really appreciated the collaboration we have had, and still have. I am also grateful to Jarle Mork who has been my supervisor during this work, the quick replies of enquires from me have been valuable. This work would have been something else without the help and support of Robin Wilson. He taught me a lot more than I knew from before about nereidids, everything I know about DELTA, gave valuable insight into biogeography and provided lab space at Museum Victoria in Melbourne. The long discussions about specimens, taxa, methods, analyses, software, theories and projects have been very fruitful, as has those about Australian wine. I stayed two periods at Museum Victoria (MV), five month in 2001/2002 and seven month in 2002/2003 (climatic migration). I would like to thank staff, students and volunteers in the Marine Biology Section at MV for memorable moments and a lot of help. Thanks goes especially to Joanne Taylor for good discussions and valuable help in systematic questions, Gary Poore, Tim O’Hara, Mark Norman, Genefor Walker-Smith, Melissa Storey and Rachael King for sharing their knowledge and always being helpful, Tim also helped during fieldwork as did Jan Watson and David Staples, and Chris Rowley and Tania Bardsley for help in the lab. Janette Norman and Renae Pratt are thanked for their assistance, help and expertise with the molecular work. I would also like to thank Martin Gomon and Dianne Bray, and Les Christidis head of Sciences Department for making the opportunity for me to stay for a whole year in the department. During the time in Australia Chris Glasby were a great host when I visited Darwin, as was Pat Hutchings in Sydney. Pat along with Kate Attwood and Lynda Avery are thanked for a great field trip in northern New South Wales in February 2003. At last, but not least, I would like to thank Rosalind and Barry Poole to let me be a part of their family during my first stay in Melbourne, they taught me a lot about Australia and Australian culture (and heaps of other things). Librarians Else-Mari Leirvoll and Tore Moen and their colleagues at Gunnerusbiblioteket, NTNU have provided enormous help in finding all the strange, not to mention old, literature I have needed. This work would have been very different without their help and effort. After the 7th International Polychaete Conference I got the opportunity to participate in the Postconference Course in cladistics held at Sandgerdi, Iceland, for which I am grateful to Elín Sigvaldadottír, Fredrik Pleijel and Greg Rouse for organising and help during the course. I am also grateful to V.V. Khlebovich (St Petersburg) who provided translations of parts of his monograph on nereidids. Anja Schulze and Kristian Fauchald were very helpful and great hosts when I visited the Smithsonian Institution in December and January 2001/2002, as was Danny Eibye-Jacobsen when I visited Copenhagen in April and May 2002. I would also thank Arne Nygren for patiently answering a novice’s questions about genes and molecules. Staff at Section of Natural History and the former Department of Natural History (NTNU) are thanked for all help, it has been great to work here and I am especially grateful for the flexibility my position has been granted when it comes to travelling, time and that I was allowed to be undisturbed when I needed all the time I could get. I am especially grateful to Toril Loennechen Moen who has been a great discussion partner when no one else could answer questions relating to systematics, molecules and software (questions are still unanswered but we are getting there). Mari-Ann Østensen were of great help in the molecular lab. Mona Ødegården, Anita Kaltenborn and Dag Altin have been fantastic when the deadline moved much faster than I did, they have inked and prepared most figures in Paper III and Paper IV. Anita and Dag are also thanked for all the fun with different projects, and so is also Jussi Evertsen who has from time to time diverted my time to nudibranchs and diving trips. Thanks also goes to my parents for being so supportive throughout the years although they have not got a clue what all this has been all about, and to Mona who has been so patient the last few months. Finally I would like to thank the following persons for access to collections, logistics during visits, and loan of material (institution acronyms are found in Paper III and Paper IV): Kate Attwood, Penny Berents and Pat Hutchings (AM, Sydney), Harry ten Hove (AZM, Amsterdam), Chris Glasby (MAGNT, Darwin), Fredrik Pleijel (NMHN, Paris), Leslie Harris (NHMLAC, Los Angeles), Stefan Lundberg and Sabine Stöhr (SMNH, Stockholm), Anja Schulze and Kristian Fauchald (USNM, Washington DC), Angelika Brandt and Brigitte Hilbig (ZMH, Hamburg), Jon Anders Kongsrud (ZMUB, Bergen), Danny Eibye-Jacobsen (ZMUC, Copenhagen), Cathrine Vollelv (ZMUO, Oslo), Wim Vader (ZMUT, Tromsø). Financial support was provided by NTNU through a position as Research Fellow, and through grants from Sparebanken Midt-Norge and Faculty of Natural Sciences and Technology (NTNU). The Norwegian Research Council (grants no 138717/419 and 151548/432) provided two scholarships that made it possible to stay 5+7 months at Museum Victoria, Melbourne, Australia. A visit to the Zoological Museum, Copenhagen (ZMUC) was made available through a grant from the European Commission’s programme ”Transnational Access to Major Research Infrastructures” to COBICE (Copenhagen Biosystematics Center). Torkild Bakken Trondheim, 9 October 2003 Contents Preface................................................................................................................. 2 Contents............................................................................................................... 4 List of papers ....................................................................................................... 5 Abstract................................................................................................................ 6 Introduction .......................................................................................................... 7 The family name......................................................................................................... 8 Family Nereididae − relationships and supra-family classification............................. 8 Classification and Phylogeny of Nereididae − an historic review .................................... 9 Approaches........................................................................................................ 14 Morphological characters ......................................................................................... 14 An attempt to include molecular studies .................................................................. 14 Aims and summary of papers ............................................................................ 16 Discussion.......................................................................................................... 18 Conclusions and prospects for further work............................................................. 19 References......................................................................................................... 20 4 List of papers I. Bakken, T. 2002. A new species of Neanthes (Polychaeta: Nereididae) from southern Australia. Memoirs of Museum Victoria, 59: 327-331. II. Wilson, R. S., Bakken, T. & Glasby, C. J. 2003. Nereididae (Polychaeta) - A DELTA database of genera, and Australian species. In R. S. Wilson, P. A. Hutchings and C. J. Glasby (Eds), Polychaetes: An Interactive Identification Guide (pp. Melbourne: CSIRO Publishing. III. Bakken, T. & Wilson, R.S. Phylogeny of nereidids (Polychaeta, Nereididae) with paragnaths. Submitted Zoologica Scripta. IV. Bakken, T. Revision of Pseudonereis (Polychaeta, Nereididae) based on type material. Manuscript. These are referred to in the text by the roman numerals. 5 Abstract Groups of taxa of the polychaete family Nereididae were studied with the aim of presenting a revision of the subfamily Nereidinae. Detailed description of characters starting with a selected group, Neanthes, was extended to include a wider range of species level taxa to include as much variation