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12th International Symposium on TARDIGRADA V.N. Gaia, Portugal, 23-26 July 2012 Guest Editors: L. Rebecchi, D.R. Nelson, R. Bertolani, P. Fontoura
Journal of Limnology
Memorie dell’Istituto Italiano di Idrobiologia
The Journal of Limnology publishes peer-reviewed original papers, review papers and notes about all aspects of limnology. The scope of the Journal of Limnology comprises the ecology, biology, microbiology, physics, and chemistry of freshwaters, including the impact of human activities, management and conservation. Coverage includes molecular-, organism-, community-, and ecosystem-level studies both applied and theoretical. Proceedings of workshops, specialized symposia and conferences, may also be accepted for publication. The Journal of Limnology is published in three issues per year, open access online and each article is available for downloading for free. A printed version is also available at the annual subscription rate of € 120,00. For further information on subscription terms, rates, methods of payment and shipping, etc., please contact the Publisher at [email protected]
Editor-in-Chief Roberto Bertoni, CNR-ISE Institute of Ecosystems Study, Verbania, Italy
Associate Editors Esteban Balseiro, Laboratorio de Limnología, Universidad Nacional del Comahue, Bariloche, Argentina Mariano Bresciani, CNR-IREA Milano, Italy Cristiana Callieri, CNR-ISE Verbania, Italy Laurence Carvalho, Scottish Freshwater Ecosystems Group, Centre for Ecology & Hydrology, Midlothian, UK Gianluca Corno, CNR-ISE Verbania, Italy Stefano Fenoglio, DISAV, Università del Piemonte Orientale Amedeo Avogadro, Alessandria, Italy Diego Fontaneto, CNR-ISE Verbania, Italy Franz Hoelker, Leibniz Institute of Freshwater Ecology and Inland Fisheries, Berlin, Germany Andrea Lami, CNR-ISE Verbania, Italy Bernard Laval, Department of Civil Engineering, University of British Columbia, Vancouver, Canada David M. Livingstone, EAWAG, Dübendorf, Switzerland Marina Manca, CNR-ISE Verbania, Italy Aldo Marchetto, CNR-ISE Verbania, Italy Giuseppe Morabito, CNR-ISE Verbania, Italy Michela Rogora, CNR-ISE Verbania, Italy John P. Smol, Paleoecological Environmental Assessment and Research Lab., Queen’s University, Kingston, Canada Dietmar Straile, Limnological institute, University of Konstanz, Germany Kay Van Damme, University of Ghent, Gent, Belgium
Editorial Board Miguel Alonso, Departament d’Ecologia, Universitat de Barcelona, Spain Jean Christophe Auguet, Centre National de la Recherche Scientifique, France Marco Bartoli, Dipartimento di Scienze Ambientali, Università di Parma, Italy Stefan Bertilsson, Department of Ecology & Genetics, Limnology, Uppsala University, Sweden Angela Boggero, CNR-ISE Verbania, Italy Reinaldo L. Bozelli, Departamento de Ecologia, Instituto de Biologia, Universidade Federal do Rio de Janeiro, Brazil George S. Bullerjahn, Department of Biological Sciences, Bowling Green State University, USA Lluis Camarero, Centre de Recerca d’Alta Muntanya, Vielha, Spain Marco Cantonati, MTSN Sezione Limnologia e Algologia, Trento, Italy Manuela Coci, CNR-ISE Verbania, Italy William R. DeMott, Indiana-Purdue University, Fort Wayne, USA Claudia Dziallas, Department of Stratified Lake, Leibniz-Institute of Freshwater Ecology and Inland Fisheries, Stechlin, Germany Walter Geller, Centre for Environmental Research - UFZ, Freiburg, Germany Claudia Giardino, CNR-IREA Milano, Italy Niels Jepsen, Section for Freshwater Fisheries Ecology, Technical University of Denmark, Silkeborg, Denmark Ora Johannsson, Dept. of Fisheries and Oceans, Great Lakes Laboratory for Fisheries and Aquatic Sciences, Hamilton, Canada Christian Kamenik, Geographisches Institut - Physische Geographie, Universität Bern, Switzerland Suzanne N. Levine, Rubenstein School of Environment and Natural Resources, University of Vermont, Burlington, USA Steven Loiselle, Dipartimento Farmaco Chimico Tecnologico, Università di Siena, Italy Gian Marco Luna, CNR - Istituto di Scienze Marine, Venezia, Italy
Hugh MacIsaac, Great Lakes Institute for Environmental Research, University of Windsor, Canada Paco Melià, Dipartimento di Elettronica e Informazione, Politecnico di Milano, Italy Beatriz E. Modenutti, Laboratorio de Limnología, Universidad Nacional del Comahu, Bariloche, Argentina Luigi Naselli-Flores, Dipartimento di Biologia Ambientale e Biodiversità, Università di Palermo, Italy Liisa Nevalainen, Department of Geosciences and Geography, University of Helsinki, Finland Roberta Piscia, CNR-ISE Verbania, Italy Giampaolo Rossetti, Dipartimento di Scienze Ambientali, Università di Parma, Italy Michaela Salcher, Limnological Station, Institute of Plant Biology, University of Zurich, Switzerland Nico Salmaso, IASMA - Centro Ricerca e Innovazione, Fondazione Mach-Istituto Agrario di S. Michele all’Adige, Italy Hugo Sarmento, Departament de Biologia Marina i Oceanografia, Institut de Ciències del Mar, Barcelona, Spain Ruben Sommaruga, Institute of Ecology, University of Innsbruck, Austria Mauro Tonolla, Istituto Cantonale Batteriosierologico, Bellinzona, Switzerland Pierluigi Viaroli, Dipartimento di Scienze Ambientali, Università di Parma, Italy Jaroslav Vrba, Dept. of Ecosystem Biology, Faculty of Science, University of South Bohemia, Ceske Budejovice, Czech Republic Brian Whitton, School of Biological and Biomedical Sciences, University of Durham, UK Ian J. Winfield, Centre for Ecology & Hydrology, Lancaster Environment Centre, Lancaster, UK Editorial Assistant Igor Cerutti, CNR-ISE Verbania, Italy Managing Editor Nadia Moscato, Pavia, Italy
Autorizzazione Cancelleria del Tribunale di Verbania N. 58 del 7 Ottobre 1959 Istituto per lo Studio degli Ecosistemi, Verbania Pallanza, Italy Cover Photo Otranto lake, Italy (courtesy of R. Bertoni) Online Publication by PAGEPress Publications via G. Belli 7 27100 Pavia, Italy Tel. +39.0382.1751762 Fax +39.0382.1750481
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Hutchinson GE, 1975. A treatise on limnology. 3. J. Wiley & Sons, New York: 660 pp. de Bernardi R, Giussani G, Lasso-Pedretti E, 1979. Food suitability and availability, demographic parameters and population growth in Daphnia obtusa Kurz under laboratory conditions. In: R. de Bernardi (Ed.), Proc. Symp. Biological and Mathematical aspects in population dynamics. Mem. Ist. ital. Idrobiol. Suppl. 37:233-242. Muyzer G, Brinkhoff T, Wawer C, 1998. Denaturing gradient gel electrophoresis (DGGE) in microbial ecology, p. 1–27. In: A.D.L. Akkermans, J. D. van Elsas and F. J. Bruijn (eds.), Molecular microbial ecology manual. Kluwer Academic Publishers. We recommend the use of a tool such as EndNote or Reference Manager for reference management and formatting. EndNote reference styles can be searched for here: http://www.endnote.com/support/enstyles.asp Reference Manager reference styles can be searched for here: http://www.refman.com/support/rmstyles.asp
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Impact factor 2012: 1.473
12th International Symposium on
TARDIGRADA V.N. Gaia, Portugal 23-26 July 2012
Proceedings Guest Editors Lorena Rebecchi (University of Modena and Reggio Emilia, Italy) Diane R. Nelson (East Tennessee State University, USA) Roberto Bertolani (University of Modena and Reggio Emilia, Italy) Paulo Fontoura (University of Porto, Portugal)
INTRODUCTION TO THE PROCEEDINGS The Twelfth International Symposium on Tardigrada has been a further, wonderful occasion to meet together and to exchange suggestions and opinions as a part of a common experience. Luckily, every symposium on tardigrades has always been followed by printed proceedings, and always of good quality. The symposium of Vila Nova de Gaia is not an exception. First of all, for your contributions. Twenty-three papers are included in these proceedings, published in the prestigious Journal of Limnology. Even though focused on tardigrades, they look diversified and very interesting. The topics of the proceedings include taxonomy and phylogeny at morphological (structure and ultrastructure) and molecular level, faunal studies, biogeography, stress tolerance, ecology (communities, life histories), methodologies and datasets. We must thank all the authors who honored the symposium by sharing the results of their research with talks and poster presentations. All of them represent the outcome of hard work and fascinating interests.
The scientific papers are preceded by the wishes of the organisers and the guest editors and more importantly by three tributes dedicated to dear colleagues, friends for some or most of us. In 2012, within just a few months, we lost Clark Beasley, Frank Romano and Jeanne Renaud-Mornant, a high number of colleagues with respect to our small community. The tributes here included are an occasion for many of us to remember interesting discussions and happy moments, and for some of us a scientific collaboration. Certainly these tributes do not fill the great void that these friends have left and that was already tangible during our meeting in Portugal, where we remembered Clark and we sent our greetings to Frank.
Regarding these proceedings, we must thank the availability of the Journal of Limnology, a milestone in tardigradology. This journal published the proceedings of the first symposium in 1975 under the old name of Memorie dell’Istituto Italiano di Idrobiologia dott. Marco de Marchi and before and later, with the same name, it published monographs on tardigrades by Ramazzotti (1962, 1972) and their supplements, the monograph by Ramazzotti and Maucci (1983), and several other papers by Ramazzotti. Under the name Journal of Limnology, in 2007 we had the proceedings of the 10th International Symposium on Tardigrada, held in Catania (2006), for which we can again appreciate the quality of the journal. Now we are at the third experience, and we can only thank the editors of the journal, Gianluigi Giussani and Roberto Bertoni, both very willing and competent persons, with whom it is very easy to collaborate. A special thank must be given to the many colleagues, tardigradologists and others, who have collaborated on the revision of the numerous contributions that enrich the proceedings of the 12th International Symposium on Tardigrada. In most cases three reviewers have considered the contributions, an approach that never created problems but on the contrary has allowed improvement of the content due to an evaluation from different points of view. Lastly, we cannot forget Paul Bartels, Harry Meyer and Sandra McInnes who, together with each one of us, have been involved in the grammar (and not only) corrections of several papers. To them and to other colleagues who directly revised the text, we send our special and certainly not less important thanks. GUEST EDITORS Lorena Rebecchi, Diane R. Nelson, Roberto Bertolani, Paulo Fontoura
12th INTERNATIONAL SYMPOSIUM ON TARDIGRADA
Tardigrada Symposia take place every three years and are unique opportunities for tardigrade scientific researchers, students and other tardigrade enthusiasts from all over the world to get together and to promote scientific exchange and friendship. Tardigradologists are not organised as a formal scientific society. Nevertheless, since the first Symposium, held in Pallanza, Italy, 1974, as a tribute to Giuseppe Ramazzotti, ten other conferences have been regularly organised. After the 11th meeting held in 2009 at the University of Tübingen, Germany, we were honored with the responsibility to organise the next conference on behalf of the Faculty of Sciences of the University of Porto and the Parque Biológico de Gaia, Portugal. We would like to thank our fellow tardigradologists, especially the deans Diane Nelson, Roberto Bertolani, Hartmuth Greven and Reinhardt M. Kristensen, for their trust in our capability to organise the meeting. The 12th International Symposium on Tardigrada was held in Vila Nova de Gaia, Portugal, from 23-26 July, 2012. The meeting took place at the auditorium of the Parque Biológico de Gaia, a permanent centre of environmental education and nature conservancy. There were a record number of attendees, more than 80, coming from twenty countries from all over the world, presenting and discussing their latest findings on many diversified topics on tardigrade biology. The scientific program was a success. The meeting included 36 oral and 40 poster presentations of high quality, about marine and limnic species, contributing to the integrative knowledge of the biology of these fascinating creatures. A special note should also be devoted to the high number of young researchers that attended the conference. Their presence is a guarantee for the future of tardigradology. Nineteen presentations were candidates for the Young Scientist Awards instituted to recognize and stimulate the scientific work developed by young researchers on Tardigrada. After a very difficult decision due to the excellent level of the presentations, the Jury constituted by senior scientists honored the oral presentation of Eliana Andrea Beltrán-Pardo about the DNA repair system of tardigrades and the poster of Vladimir Gross in which the muscular organization of marine and limnoterrestrial species had been analysed. In addition to the promotion of scientific discussion and cooperation, symposia are events also providing contact with new cultures and moments of diversion. We have done our best to honor the traditional hospitality and authenticity of the Portuguese people, offering all the participants the opportunity to discover the charm, natural beauty and history of the cities of Gaia and Porto and to taste the wonderful Portuguese gastronomy. The Symposium included a visit to the port wine cellars, allowing the visitors to taste this worldwide famous nectar and get in touch with its history. A sightseeing tour at the seaside of Porto and Vila Nova de Gaia to admire the picturesque urban landscape and some stunning beaches, with a walk in the local nature reserve of the Douro estuary and park dunes of Aguda, ended by the conference dinner, was also included in the social program. Finally, an all-day boat cruise on the river Douro from Porto to Régua to admire the beautiful landscape combining lovely rural scenarios and vineyards shaped by human sweat was also held. The river Douro valley, rewarded in the 18th century by the Marquis of Pombal with the title of the first demarcated wine region in the world, is considered a World Heritage Site by UNESCO since 2001. During the conference sessions, an additional social program for accompanying persons was also offered. However, the success of the 12th International Symposium on Tardigrada would not be possible without the support of some public and private organizations. We want to thank the Municipality of Vila Nova de Gaia, the University of Porto, the Comissão de Coordenação e Desenvolvimento Regional do Norte (CCDR-N, responsible for the execution of the Program ON.2 O Novo Norte co-funded by the European Regional Development Fund - ERDF), the Eco-Ethology Research Unit, the Grupo Taper and the Árvore Digital. We also received the help of many people to whom we are greatful. In particular, the Organizing Committee wants to thank the colleagues Diane Nelson, Roberto Bertolani and Lorena Rebecchi for their indispensable scientific collaboration. Without their help we would not have been able to accomplish this initiative. A special acknowledgment is devoted to all the members of the staff of Parque Biológico for their enthusiastic and patient work and also to some students of Biology (University of Porto) for their voluntary collaboration. Last but not least, we are also indebted to all the participants in the Symposium. Their amiable nature and range of expertise were responsible for four days of an agreeable atmosphere of friendship and science. It was a great pleasure to host the 12th International Symposium on Tardigrada in Vila Nova de Gaia. We sincerely wish an even greater success for the next symposium, where we hope to meet the old and new tardigradologists. THE ORGANISING COMMITTEE Paulo Fontoura (Faculdade de Ciências da Universidade do Porto) Nuno Gomes Oliveira (Parque Biológico de Gaia) Alexandre Valente (Faculdade de Ciências da Universidade do Porto) Cristina Neves (Parque Biológico de Gaia)
12th INTERNATIONAL SYMPOSIUM ON TARDIGRADA July 23rd – 26th 2012, Vila Nova de Gaia (Portugal) The participants
1 – F. Förster 2 – F. Bemm 3 – L. Grothman 4 – A. Grothman 5 – K. Oguchi 6 – G. Grothman 7 – B. Ramsay 8 – D. Horikawa 9 – R. Guidetti 10 – S. Gonçalves 11 – A. Jørgensen 12 – J. Eibye-Jacobsen 13 – M. Devetter 14 – K. Arakawa 15 – W.R. Miller 16 – J. Miller 17 – H.A. Meyer 18 – B. Luz 19 – Y. Kiosya 20 – M. Czerneková 21 – E. Santos 22 – V. Gross 23 – M. Bryndová 24 – C. Schulze 25 – M. Nottoli 26 – T. Tischer 27 – P. Bartels 28 – M. Rubal
29 – J. Cytan 30 – I. Poprawa 31 – P. Degma 32 – T. Kunieda 33 – S. Kawada-Suzuki with J. Suzuki 34 – B.V. Trygvadóttir 35 – J. Palhas 36 – A.I. Proença Pinto 37 – A. Cardoso 38 – D. Persson 39 – K.A. Halberg 40 - K. Zawierucha 41 – S. Quiroga 42 – C. Nilsson 43 – C. Rocha 44 – P. Fontoura 45 – R. Bertolani 46 – S. Ferrari 47 – V. Inshyna 48 – C. Gonçalves 49 – R. Londoño 50 – J. Nelson 51 – I. Jönsson 52 – T. Prasath 53 – D. Santos 54 – P. Veiga 55 – S. Tanaka 56 – S. Komine
57 – N. Guil 58 – A.M. Rocha 59 - T. Hashimoto 60 – S. Fujimoto 61 – B. Pappalardo 62 – S.J. McInnes 63 – Ł. Michalczyk 64 – N.G. Oliveira 65 – I. Šatkauskienė 66 – A.M. Jørgensen 67 – N. Møbjerg 68 – H. Greven 69 – L. Rebecchi 70 – O. Lisi 71 – G. Mayer 72 – E. Beltrán-Pardo 73 – J.G. Hinton 74 – S.M. Jørgensen 75 – R.M. Kristensen 76 – G.M. Kristensen 77 – T. Altiero 78 – D.R. Nelson 79 – A.C. Suzuki 80 – R.O. Schill 81 – N. Marley Not in the photo: A. Lyons C. Neves
Courtesy of Ralph O. Schill
TRIBUTE TO DR. CLARK W. BEASLEY
Dr. Clark W. Beasley passed away unexpectedly in July 2012. We received the notice of his death shortly before the 12th International Symposium on Tardigrada in Porto, Portugal, which he was planning to attend. A special tribute to Clark was presented at the beginning of the symposium by Dr. Paulo Fontoura. After receiving his PhD in 1968 from the University of Oklahoma, Clark joined the Biology Department at McMurry College in College Station, Texas, USA, in 1969, becoming the chair in 1973. He was awarded the
honour of Distinguished Professor in 1999. Even after his retirement from McMurry University in 2010, Clark continued his research on tardigrades and his interest in helping others with tardigrade taxonomy, especially those who were just beginning. With his first publication on tardigrades from Kansas in 1967, Clark began a life-long interest in tardigrades. After his first symposium participation in the 3rd International Symposium of Tardigrada at Johnson City, Tennessee, USA, in August 1980, he continued to be a regular at all the subsequent symposia, greeting all with his welcoming smile and hearty laughter. Assisted by others, Clark translated the Italian monograph Il Phylum Tardigrada (Ramazzotti and Maucci 1983) and published the English version, making it available worldwide. He also compiled a pdf bibliography of tardigrade publications through July 2010, which he shared with other tardigradologists. China held a special attraction for Clark, who made several field trips there, publishing articles on new records and new species from his extensive collections. Perhaps Clark’s greatest contribution came in his role as the editor for 77 tardigrade publications in Zootaxa from 2001-2010. Sandra McInnes summed it up well, Clark’s support, assistance and guidance to both early stage researchers and those with longer track records will be sadly missed. My friendship with Clark goes back 32 years and is filled with sharing many fond memories of tardigrade symposia. Diane R. Nelson
Courtesy of Jack Nelson
TRIBUTE TO DR. FRANK A. ROMANO III
Dr. Frank A. Romano III, retired chair of the Department of Biology, Jacksonville State University, Jacksonville, Alabama, USA, passed away on 17 August 2012 after an extended battle with esophageal cancer. Photos and videos of Frank and his students can be viewed on the JSU Biology web page, www.jsu.edu/biology With a PhD from Syracuse University, Frank joined the JSU faculty in 1989 and became chair in 2002. He retired in May 2012, due to his illness. Throughout his career, his professionalism and rapport with his students and
colleagues were unparalleled. He was selected for the 2010 Meritorious Teaching Award given by the Association of Southeastern Biologists. In honor of his retirement, the Dr. Frank Romano III Outdoor Classroom was dedicated at Jacksonville State University. Frank had a unique ability to interest students in pursuing research, especially sharing his fascination and enthusiasm for working with tardigrades. Many of his students earned their Master’s degrees at JSU studying the ecology of limno-terrestrial and marine tardigrades, and several presented their research at our international symposia. One of Frank’s students, Dr. Brent Nichols, who went on to earn his PhD at the University of South Florida under Dr. Jim Garey, played a major role in organizing the 9th International Symposium on Tardigrada in Tampa, Florida, USA, and served as one of the Guest Editors for the symposium volume published in Hydrobiologia. Those of you who were fortunate to know Frank from his attendance at our international symposia were blessed to have a friend and a colleague who shared his enthusiasm for life and tardigrades and his positive outlook, sense of humor, and pride in his Italian heritage. It has been 20 years since Frank asked me at an ASB meeting to help him and his students study tardigrades. I am privileged to have had such a long friendship with this dedicated man, who has meant so much to so many people. Diane R. Nelson
Courtesy of Jack Nelson
From Psammonalia, number 158, December 2012
TRIBUTE TO DR. JEANNE RENAUD-MORNANT
Jeanne Renaud-Mornant passed away on 18 September 2012 in Paris. Retired since 1989, she was Directeur de Recherche Honoraire of the National Centre of Scientific Research (CNRS) in France. Jeanne Renaud-Mornant was born in 1925 at Vellexon (Franche-Compté, Haute-Saône) near Besançon, France. After finishing the academic studies at the University of Bordeaux, she started her research activities at the Marine Station of Arcachon, France. The PhD thesis of Jeanne Renaud-Mornant, concluded in 1961, at Paris, under the supervision of Delamare-Debouteville, was an innovative document on the distribution of the interstitial micrometazoans in relation with the substrate. At that time, this document was considered a major event in meiofauna research. During her long and remarkable career, Jeanne Re-
naud-Mornant specialized in taxonomy and ecology. She participated in diversified marine scientific expeditions all around the world, with special emphasis in the Atlantic ocean (French coast, Angola and Namibian coasts, Antilles and Brazil) and in the Pacific ocean (French Polynesia). She studied the fauna of very different ecosystems (e.g. estuaries, sandy beaches, coral reefs, abyssal deeps). Jeanne devoted her attention to several meiofaunal groups, such as Gastrotricha, Annelida, Crustacea, Loricifera, but definitely, her main interest was the study of tardigrades. She published important papers on morphology and phylogeny of tardigrades and described an impressive number of new taxa: 56 new tardigrade species (more than 25% of all the marine species known today), 17 new genera and 5 new families or subfamilies. She also participated in three of the four international tardigrade symposia from 1974 (first) to 1985 (fourth). Her scientific and human qualities were widely recognized by her fellow researchers, allowing the development of numerous research projects in collaboration with a lot of international institutions (Universities, museums and marine stations), not only in Europe but also in North and South America. As a tribute to the work of Jeanne Renaud-Mornant, her name has been attributed to a large number of new taxa: several species of different phyla, and one genus and one family of tardigrades bear her name. The commitment of Jeanne to the formation of a new generation of marine biologists of diverse nationalities also deserves special mention. We have lost a remarkable scientist, undoubtedly one of the more important names in tardigradology, and we are grateful for her incomparable contributions to the development of science. The Guest Editors of the Proceedings of the 12th International Symposium on Tardigrada
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12th International Symposium on TARDIGRADA V.N. Gaia, Portugal, 23-26 July 2012 Guest Editors: L. Rebecchi, D.R. Nelson, R. Bertolani, P. Fontoura