Biology and Conservation of the European Sturgeon Acipenser Sturio L

Biology and Conservation of the European Sturgeon Acipenser sturio L. 1758 . Patrick Williot l Eric Rochard l Nathalie Desse-Berset l Frank Kirschbaum l Jo¨rn Gessner Editors

Biology and Conservation of the European Sturgeon Acipenser sturio L. 1758

The Reunion of the European and Atlantic Sturgeons Editors Dr. Patrick Williot Dr. Eric Rochard Sturgeon Consultant Cemagref Rue du Pas de Madame 4 Estuarine Ecosystems and Diadromous 33980 Audenge Fish Research Unit France Avenue de Verdun 50 [email protected] 33612 Cestas cedex France Dr. Nathalie Desse-Berset [email protected] Universite´ de Nice-Sophia Antipolis CNRS-CEPAM-UMR 6130, SJA3 Prof. Frank Kirschbaum Avenue des Diables Bleus 24 Humboldt-Universita¨t Berlin 06357 Nice cedex 4 Abt. Biologie und O¨ kologie der France Fische [email protected] Philippstr. 13 [email protected] 10115 Berlin Haus 16 Dr. Jo¨rn Gessner Germany Leibniz-Institut fu¨r [email protected] Gewa¨ssero¨kologie und Binnenﬁscherei Mu¨ggelseedamm 310 12587 Berlin Germany [email protected]

ISBN 978-3-642-20610-8 e-ISBN 978-3-642-20611-5 DOI 10.1007/978-3-642-20611-5 Springer Heidelberg Dordrecht London New York

Library of Congress Control Number: 2011935343

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Springer is part of Springer Science+Business Media (www.springer.com) Dedication to Dr. Etienne Magnin

Dr. E´ tienne Magnin (1922À1990) was one of the few contemporary researchers to take an interest in European sturgeon before it became an endangered species. His doctoral thesis and subsequent work on the biology of the species – and, more specifically, of the population of the Gironde basin – on its taxonomic position relative to Atlantic sturgeon and on its status, which was already considered a cause for concern in the 1950s, laid much of the scientific groundwork for a body of knowledge that others continue to build on and that will, we hope, some day make possible the survival of this magnificent fish. Paradoxically, Dr. Magnin, though he came late to the field of biology, was a pioneer, and not only with respect to his research in France on European sturgeon. After settling in Canada, where he was first a consultant (1962), then a professor in the Department of Biology at the Universite´ de Montreál (1964À1987), he was also a pioneer in the development of knowledge about the aquatic ecosystems of the St. Lawrence River and the James Bay region during the early stages of construc- tion of the large-scale hydroelectric projects in this vast, little-known northern territory. His areas of interest were many. Both an ichthyologist and a limnologist, Dr. Magnin authored or co-authored, with the graduate students and researchers who worked in his laboratory and that of the Socie´te´ d’eńergie de la Baie James where he was scientific director for more than 7 years, 81 scientific papers: 48 on fish (including 16 on European and North American Acipenseridae), 17 on benthic organisms, six on zooplankton, five on phytoplankton and five on various other subjects (such as amphibians). His book on the freshwater ecology of the James Bay region (1977) still remains a major reference work. Dr. Magnin was also an outstanding teacher. Those, like us, who had the opportunity of taking his zoology and comparative anatomy courses were able to appreci- ate his rigorous approach, his clarity of thought, his attention to detail, his respect for students and his sense of humour. He was also a mentor, whose knowledge, commitment, enthusiasm and support enabled him to train three PhD students and 28 master’s students in a span of 20 years. Many of these went on to their own research careers in academe and public service, contributing, in turn, to improving

v vi Dedication to Dr. Etienne Magnin our understanding and protection of freshwater ecosystems, preparing a new gen- eration of biologists and developing a profession that plays an increasingly integrated role in the decision-making processes of today’s society.

Pierre Dumont Ministe`re des Ressources naturelles et de la Faune du Que´bec Peter Harper Universite´ de Montre´al Dedication to Dr. Juraj Holcˇik

RNDr. Juraj Holcˇ´ık, CSc. – eminent Slovak zoologist was born on October 18, 1934 in Trnava in Western Slovakia. He completed his B.Sc. (1958) and PhD (1966) studies at Charles University in Prague. Before being appointed Director of the newly established Institute of Zoology and Ecosozology (in 1995 re-named Institute of Zoology), Slovak Academy of Sciences in 1990, Dr. Holcˇ´ık was research officer and after 1972 the Senior Research Officer of the Institute of Fishery Research and Hydrobiology in Brati- slava. He also worked at the Regional Museum at Trnava and the Slovak National Museum in Bratislava as the Curator of Zoology and Curator of Lower Vertebrates respectively. The scope of his scientific activity was very diverse, and included taxonomy, zoogeography, ecology, population dynamics, limnology, and ecosozology, as well as production, introduction, acclimatization and conservation of fish gene pool (genofond). However, ichthyology was the main focus of his work. He was an internationally renowned expert on Palearctic Petromyzontidae, Salmonidae, and Cyprinidae (especially the subfamilies Acheilognathinae and Percidae). He described six new fish species, three subspecies and two new hybrid forms of fish fauna in Slovakia, Romania, Mongolia, the Balkans and Iran. Of these, Gymnoce- phalus baloni (Holcˇ´ık and Hensel 1974) is considered to be one of the last new vertebrate taxa to be described on Slovak territory. He contributed to the development of a methodology for studying ecological fish production in open waters. He contributed significantly to the fact that the Slovak–Hungarian section of the Danube river is one of the best studied sections not only of the Danube but also of large rivers overall. He also contributed to our understanding of the importance of floodplains and knowledge of the causal relationships between the density, ecological production and activity of fish populations on one hand, and main abiotic environmental factors on the other hand. Concerning the population dynamics of fish in man-made water reservoirs, he amended principles of stepwise evolution of ichthyofauna. He was an uncompromising advocate of the conservation of the

vii viii Dedication to Dr. Juraj Holcˇik

Danube inland delta below Bratislava which he supported even as a Member of Parliament of the Czecho–Slovak Federative Republic. During his career he published, either alone or with co-authors, 170 original articles, over 200 popular science articles, and 25 books or book chapters. Some of them were published in several languages and multiple editions. For example, the book Holcˇ´ık J, Mihlik J and Maly´ J: Freshwater Fishes was published in four English editions as well as in seven additional editions in five different languages. Dr. Holcˇ´ık is coauthor of an ongoing book series entitled The Freshwater Fishes of Europe, for which he created the structure of individual book chapters and was the editor of the first part of the series (Vol. 1/I: Petromyzontes, 1986, Vol. 1/II: General introduction to Fishes, Acipenseriformes, 1989, AULA Verlag Wiesbaden) for which he also authored several chapters. It is noteworthy that in 1998 he published the first and still the only textbook of ichthyology in the former Czechoslovakia (Ichtyolo´gia, Vydavatelˇstvo Prı´roda, Bratislava). This significant scientific output is a demonstration of his creativity, extraordinary industriousness, and determination. In the 1960s J. Holcˇ´ık led a Czechoslovak limnological expedition to Cuba. In addition, he worked as a visiting professor at the University of Waterloo (Canada, 1986–1987; 1992), and as a technical advisor for FAO in Iran (1989–1991) and Ghana (1995–1996). He also supervised and graduated several promising scientists. Dr. Holcˇ´ık’s publications and his activity in international ichthyological organizations (which was negatively influenced by political discrimination from the 1950s to the 1980s) had a significant impact on his scientific reputation. He was frequently asked to chair congresses and symposia, became a member of editorial boards of five international scientific journals (Biologia, Folia Zoologica, Bios, Environmental Biology of Fishes, and Voprosy ichtiologii), and was an active board member in many, primarily ichthyological organizations. For his lifetime achieve- ments he was awarded the World Wide Fund for Nature Award for Conservation Merit in 1996 and the G.J. Mendel honour medal of the Czech Academy of Sciences in 1998. He passed away on May 16, 2010.

Kristina Holcˇik Preface

General Introduction, Context, Origin and Setting Up of the Project

The European sturgeon, Acipenser sturio L. 1758, used to be one of the most widespread sturgeon species (Magnin 1959). However, like most Eurasian sturgeon species, its status has now become critical (Williot et al. 2002), it has been extir- pated from the great majority of its natural biotopes, and survives only in the French Atlantic coast façade, the Garonne basin, with a population on the verge of extinction (Rochard et al. 1990). The species was exploited more or less intensively for decades and even centuries, as it was in most European countries bordering the Mediterranean Sea (Italy, France, and Spain), the Eastern Atlantic Ocean (Spain, France), the North Sea (the Netherlands and Germany), and the Baltic Sea (Classen 1944; Holcˇik 1989). The species can therefore be looked upon as an emblematic European species. Compared with terrestrial animals or plants, fish combine several handicaps: they are not visible, they move, sometimes far away from the administrative limits of human societies. Fisheries are under-managed, as can be seen from changes in fish population status, and fish conservation is almost ignored. The sturgeon is no exception. Moreover, the European sturgeon is further handicapped. It is an anadromous species which colonises alternately a great variety of biotopes in fresh, brackish, and marine waters which do not tick the administrative boxes. Other biological characteristics such as longevity, late puberty, and non-yearly oogenesis make them even more susceptible to exploitation in spite of their apparent robust- ness (Boreman 1997; Jager et al. 2008).

Much has been done in France in the last few decades in favour of the preservation– conservation–restoration of the species, especially by research bodies. However, despite great efforts and protection, the status of the species has deteriorated. The conservation programme of the European sturgeon in France was begun in the early 1970s. In 2007 for the ﬁrst time, a breakthrough was achieved with the ﬁrst

ix x Preface reproduction of cultured brood fish (Williot et al. 2009) which made it possible for the restoration programme to start again. This was the opportunity to draw up an extensive assessment of the long-term programme, to update available knowledge, describe the context, and make known some long-term concerns. The European sturgeon restoration–conservation programme has been a compli- cated programme, full of pitfalls. We realised that, among other things, besides its unusual scientific and technical aspects, conservation does not only have scientific relevance. Indeed, social, economic, political, financial, and administrative aspects are an integral part of any preservation–conservation programme. In the field of science, conservation issues are facing many difficulties. Research into biodiversity is favoured but not protection stricto sensu, and conservation actions are not considered with much enthusiasm in France. Thus, grants from this source are for the most part not accessible for concrete actions. In addition, in the absence of training courses in Conservation Biology, most people working in the field are undereducated, at least when they begin their careers. Thus, it is necessary to take time to explain, to discuss, to exchange and finally to get the consensus of all those involved, even though a specific decision might be not considered as the optimal one from scientific and technical points of view. Consen- sus is a key issue for going further. Many biological sciences are potentially concerned by fish management and conservation, sometimes at both fundamental and applied levels. Unfortunately, it is often difficult to mobilise specialists. One of the great challenges of conservation biology is that the situation of certain species is so critical that waiting for supposedly needed data would inexorably lead to the extinction of a species. The European sturgeon conservation programme was a good example of the difficulties that have been faced. Due to the high depletion status of the species, the decision was taken by a few people in a research institute to acquire experience and set up methods using another sturgeon species as a biological model, the Siberian sturgeon, Acipenser baerii Brandt 1869. Not only did the species play its role of model species almost to perfection, but also the further economic development achieved by farming the species enabled the leading French institute to obtain a new research facility. And this proved to be a key point for building the ex situ European sturgeon brood stock. Another fruitful conse- quence was that conservation and farming sturgeon activities enabled us to hold ACIPENSER, the first International Symposium on sturgeon (ISS1) in Bordeaux in 1989 (Williot (ed) 1991). The evaluation of the primary project submitted to Springer was rapid, positive and accompanied by two requests. The first was to include the restoration plan of the Atlantic sturgeon (Acipenser oxyrinchus) in the US, and the second to update the data regarding the status of the European sturgeon in the River Rioni in Georgia. No recent paper has been published on the status of the species in Georgia since the synthesis by Ninua (1976). At the time, it had been known for some years that the Atlantic sturgeon (A. oxyrinchus) inhabited the Baltic Sea (Ludwig et al. 2002; Tiedemann et al. 2007). Consequently, restoration actions for the species have started in both Poland and Germany. At almost the same time as our book project got underway, a French Preface xi archaeozoologist was publishing a scoop showing that the Atlantic sturgeon had inhabited the French Atlantic coasts for a long time (Desse-Berset 2009), from 5000 YBP to 300 YAD, which strongly suggested that the two species were sympatric in the area. We therefore seized the opportunity offered by the Springer incentive, and decided to broaden the Atlantic sturgeon issues in this book by introducing several actions in favour of the species, and looking for somebody able to provide the latest available data from the Georgian population. So this book is timely in taking into account the new situation of sturgeons in Western Europe, especially in France. It is worth noting that it was only recently in the 1960s that the two species were recognised as different (Magnin 1962, 1964; Magnin and Beaulieu 1963). And further genetic investigations have constantly pointed out the great proximity of the two species (e.g., Fontana et al. 2008). Clearly, such a project would have missed its objectives without the participa- tion of co-editors, including the unusual involvement of an archaeozoologist. This illustrates the need to open up conservation biology to new fields, and the need for cooperation, as mentioned above. All of them have brought very decisive inputs to the book, which would have been much less valuable otherwise. I would like to gratefully acknowledge Eric, Nathalie, Jo¨rn and Frank for their enthusiasm, kind- ness and efficiency in contributing to our work. The contents of the book were based on the following central themes: (a) pro- viding a book on conservation in practice, with the European sturgeon as an example, (b) updating knowledge in the field, (c) broadening the range of concerned disciplines, (d) highlighting aspects that are not strictly scientific, (e) assuming new issues from the presence of two sturgeon species, the European and the Atlantic sturgeons, and (f) showing internal coherence, illustrated by the numerous cross- references. The very great majority of those we contacted agreed to contribute. The contributors are greatly acknowledged, all the more so since many of them provided unpublished inputs, either results or analysis. The book is divided into six unequal parts. The first part deals with all the available biological data on species population. A very brief introduction provides the reader unfamiliar with sturgeon with some basic characteristics, with a special focus on the European sturgeon. In addition to cytogenetic matters, a large section is devoted to geographical extension, present and past, with different approaches, literature, archaeozoology, and palaeogenetics, to the history of populations and fisheries in diverse countries, especially those for which there was no recent synthesis, and to some specific biological traits. As the Atlantic sturgeon was “invited” to participate in the book, two chapters deal with comparisons of the two species in terms of morphology and osteometry of the bones and morphology for the juveniles. It is noteworthy that most of the physiological functions of the species were, and still are, to a great extent under-documented, e.g., reproduction, osmoregula- tion, endocrinology, nutrition, and haematology, with the exception of a preliminary investigation on the hydromineral balance (Magnin 1962), and on reproductive endocrinology (Davail-Cuisset et al. 2008). xii Preface

The second part is an illustrated interlude devoted to part of the iconography of the European sturgeon, i.e., mainly photos and stamps. The sources are limited to France for the photos. The third part focuses on the restoration–conservation actions that have been undertaken in France and later in Germany. Apart from the two chapters that draw an historic overview of European sturgeon-related management issues in France and Germany, the other chapters can be arranged under three topics dealing with: population, ex situ efforts, and efforts relying rather on non-scientific actions. The chapters reporting on population studies focus on methods and their applications, e.g., age determination, marking and tagging, monitoring of post-release sturgeon, and an attempt to model the future of these released fish. The ex situ actions cover sex determination and maturation staging, reproduction with two alternatives, wild and farmed brood fish because the management logics are different, larval rearing, brood stock building, its genetic variability, and sperm cryoconservation. The third topic in this part deals with the role of non-governmental stakeholders, awareness campaigns among marine professional fishermen, and a synthesis on French– German cooperation. The fourth part is focused entirely on Atlantic sturgeon management, illustrated by fisheries in Que´bec (Ca), conservation in the US, and restoration in the Baltic Sea. Some perspectives and prospective concerns are described in part five. They deal with the in situ life history, the potential effects of climate change, a population viability analysis, some preliminary investigation into cryobanking of somatic cells, and biological variability. Finally, part six contains some thoughts on the future of the species. As a preface to part 1, I am very pleased to dedicate the book to the memory of two renowned scientists because the present European sturgeon combined biology– restoration programme benefited from their previous studies and investment in the species. Dr Etienne Magnin started his career in France by publishing the first documented compendium on the biology of the European sturgeon and then went on to work in Que´bec, Canada where he also spent time working on sturgeon, especially (but not only) the Atlantic sturgeon. He was responsible for the first documented distinction between the two species, the European and Atlantic sturgeons. E. Magnin unfortunately missed our invitation to participate in ACIPEN- SER, the ISS1 in 1989. The second scientist is Dr Juraj Holcˇik, well known for many important works, but in particular for publishing The Freshwater Fishes of Europe, focusing on Petromyzontiformes and Acipenseriformes, with the last edition published in 1989. He personally wrote with colleagues the chapter that dealt with the European sturgeon, and continued to be involved in those issues. Due to health problems, he declined our invitation to contribute to the present book. Many thanks are due to Pierre Dumont and Peter Harper for the lines on E. Magnin. I am very grateful to Kristina Holcˇik for her kind offer of a text on Juraj’s activities.

Audenge, France Patrick Williot Preface xiii

References

Boreman J (1997) Sensitivity of North American sturgeons and paddlefish to fishing mortality. Environ Biol Fish 48:399–405 Classen TEA (1944) Estudio bio-estadistico del esturion o sollo del Guadalquivir. Instituto Espanol de Oceanografia, Ministerio de Marina, Trabajos, N 19, 112 p + XVII planches Davail-Cuisset B, Lacomme S, Viaene E, Williot P, Lepage M, Gonthier P, Davail S, Rouault T (2008) Hormonal profile in adults of Atlantic European sturgeon, Acipenser sturio, adapted to hatchery in France. Cybium 32(2 suppl):169–170 Desse-Berset N (2009) First archaeozoological identification of Atlantic sturgeon (Acipenser oxyrinchus Mitchill 1815) in France. CR Palevol 8:717–724 Fontana F, Lanfredi M, Kirschbaum F, Garrido-Ramos MA, Robles F, Forlani A, Congiu L (2008) Comparison of karyotypes of Acipenser oxyrinchus and A. sturio by chromosome banding and fluorescent in situ hybridization. Genetica 1342:281–286 Holcˇik J, Kinzelbach R, Sokolov LI, Vassilev V P (1989) Acipenser sturio Linnaeus, 1758. In: Holcˇik J (ed) The freshwater fishes of Europe. Aula Verlag, Wiesbaden, pp 367–394 Jager HI, Rose KA, Vila-Gispert A (2008) Life history correlates and extinction risk of capital- breeding fish. In: Dufour S, Pre´vost E, Rochard E, Williot P (eds) Fish and diadromy in Europe (ecology, management, conservation). Hydrobiologia 602:15–25 Ludwig A, Debus L, Lieckfeldt D, Wirgin I, Benecke N, Jenneckens I, Williot P, Waldman JR, Pitra C (2002) When the American sea sturgeon swam east. Nature 419:447–448 Magnin E (1959) Re´partition actuelle des Acipense´ride´s. Rev Trav Inst Peˆches Marit 23(3): 277–285 Magnin E (1962) Recherches sur la syste´matique et la biologie des Acipense´ride´s Acipenser sturio L., Acipenser oxyrinchus Mitchill et Acipenser fulvescens Raf. Ann Station Centr Hydrobiol Appl 9:7–242 Magnin E, Beaulieu G (1963) Etude morphome´trique compareédel’Acipenser oxyrinchus Mitc- hill du Saint Laurent et de l’Acipenser sturio Linne´ de la Gironde. Le Naturaliste Canadien XC (1):5–38 Magnin E (1964) Validite´ d’une distinction spećifique entre les deux Acipense´ride´s: Acipenser sturio L. d’Europe et Acipenser oxyrinchus d’Ame´rique du Nord. Le Naturaliste Canadien XCI (1):5–20 Ninua NSh (1976) Atlantic sturgeon of the Rioni River. Editions Metsniereba, Tbilissi, pp 122 (in Russian) Rochard E, Castelnaud G, Lepage M (1990) Sturgeons (Pisces: Acipenseridae); threats and prospects. J Fish Biol 37A:123–132 Tiedemann R, Moll K, Paulus KB, Scheer M, Williot P, Bartel R, Gessner J, Kirschbaum F (2007) Atlantic sturgeons (Acipenser sturio, Acipenser oxyrinchus): American females successful in Europe. Naturwissenschaften 94:213–217. doi: 10.1007/s0014-006-0175-1 Williot P (ed) (1991) Acipenser. Actes du premier colloque international sur l’esturgeon. Cemagref, Antony, 518p Williot P, Arlati G, Chebanov M, Gulyas T, Kasimov R, Kirschbaum F, Patriche N, Pavlovskaya L, Poliakova L, Pourkazemi M, Kim Yu, Zhuang P, Zholdasova IM (2002) Status and management of Eurasian sturgeon: an overview. Int Rev Hydrobiol 87:483–506 Williot P, Rouault T, Pelard M, Mercier D, Jacobs L (2009) Artificial reproduction of captive endangered European Atlantic sturgeon, Acipenser sturio. Endang Species Res 6:251–257. doi: 10.3354/esr00174 . Acknowledgements

We would like to thank all the colleagues at Cemagref who spent time and effort into this long-term programme, and more speciﬁcally Chantal Gardes for her decisive help with documentation and Philippe Camoin for his map drawing. I am also thankful to the EPBx research unit of Cemagref for the logistic help. Thanks are due to Antoine Pasqualini and Jean-Denys Strich (CEPAM-CNRS-Nice University) for their inputs in infography and photography, to Tamas Gulyas for his previous support in sperm cryoconservation and androgenesis investigations. This project would not exist without the efﬁcient and kind help of Verena Penning, Anette Lindqvist, and Athiappan Kumar from Springer.

xv . Contents

Part I Biology, History, Geographical Distribution and Status, from Past to Present

1 Brief Introduction to Sturgeon with a Special Focus on the European Sturgeon, Acipenser sturio L. 1758 ...... 3 P. Williot, E. Rochard, N. Desse-Berset, J. Gessner, and F. Kirschbaum

2 Cytogenetics as a Tool for an Exploration of A. sturio Status Within Sturgeons ...... 13 Francesco Fontana

3 Discrimination of Acipenser sturio, Acipenser oxyrinchus and Acipenser naccarii by Morphology of Bones and Osteometry ... 23 Nathalie Desse-Berset

4 Morphological Distinction Between Juvenile Stages of the European Sturgeon Acipenser sturio and the Atlantic Sturgeon Acipenser oxyrinchus ...... 53 Sven Wuertz, Stefan Reiser, Jo¨rn Gessner, and Frank Kirschbaum

5 Ontogeny of the European Sturgeon, Acipenser sturio ...... 65 Frank Kirschbaum and Patrick Williot

6 An Overview on Geographical Distribution from Past Descriptions ...... 81 Ge´raldine Lassalle, M. Be´guer, and E. Rochard

xvii xviii Contents

7 Ancient Sturgeon Populations in France Through Archaeozoological Remains, from Prehistoric Time Until the Eighteenth Century ...... 91 Nathalie Desse-Berset

8 Palaeogeographic Patterns of A. sturio ...... 117 Olivier Chassaing, Nathalie Desse-Berset, Marilyne Duffraisse, Gae¨l Pique`s, Catherine Ha¨nni, and Patrick Berrebi

9 Sturgeon in Iberia from Past to Present ...... 131 Arne Ludwig, Arturo Morales-Mun˜iz, and Eufrasia Rosello´-Izquierdo

10 Biological Cycles and Migrations of Acipenser sturio ...... 147 M.L. Acolas, G. Castelnaud, M. Lepage, and E. Rochard

11 Habitat, Movements and Feeding of Juvenile European Sturgeon (Acipenser sturio) in Gironde Estuary ...... 153 Laurent Brosse, Catherine Taverny, and Mario Lepage

12 Characteristics of the Reproductive Cycle of Wild Acipenser sturio ...... 165 Patrick Williot, Thierry Rouault, Re´mi Brun, and Jo¨rn Gessner

13 Sturgeon Fishing, Landings, and Caviar Production During the Twentieth Century in the Garonne Basin and the Coastal Sea ...... 177 Ge´rard Castelnaud

14 Historic Overview on the Status of the European Sturgeon (Acipenser sturio) and Its Fishery in the North Sea and Its Tributaries with a Focus on German Waters ...... 195 J. Gessner, S. Spratte, and F. Kirschbaum

15 History of the Sturgeon in the Baltic Sea and Lake Ladoga ...... 221 Ryszard Kolman, Andrzej Kapusta, and Jacek Morzuch

16 The Historical and Contemporary Status of the European Sturgeon, Acipenser sturio L., in Italy ...... 227 Paolo Bronzi, Giuseppe Castaldelli, Stefano Cataudella, and Remigio Rossi

17 European Sturgeon, Acipenser sturio in Georgia ...... 243 Ryszard Kolman Contents xix

18 Present Legal Status of the European Sturgeon Acipenser sturio ... 251 E. Rochard

Part II Iconography of the European Sturgeon

19 Iconography of the European Sturgeon in France ...... 259 Nathalie Desse-Berset and Patrick Williot

Part III Restoration: Conservation of Acipenser sturio, Scientiﬁc and Political Management

20 Historic Overview of the European Sturgeon Acipenser sturio in France: Surveys, Regulations, Reasons for the Decline, Conservation, and Analysis ...... 285 Patrick Williot and Ge´rard Castelnaud

21 Restoration of the European Sturgeon Acipenser sturio in Germany ...... 309 Frank Kirschbaum, Patrick Williot, Frank Fredrich, Ralph Tiedemann, and Jo¨rn Gessner

22 Fishery By-Catch of European Sturgeon in the Bay of Biscay ...... 335 M. Lepage and E. Rochard

23 Age Assessment in European Sturgeon ...... 343 Philippe Jatteau, E. Rochard, M. Lepage, and Christine Gazeau

24 Tagging European and Atlantic Sturgeons in Europe ...... 349 Philippe Jatteau, G. Castelnaud, E. Rochard, J. Gessner, and M. Lepage

25 Mass Marking in European Sturgeon ...... 357 Philippe Jatteau and Aude Lochet

26 Sex Determination and Staging of Gonads ...... 369 Patrick Williot

27 Reproduction of Wild Brood Fish from Garonne Basin ...... 383 Patrick Williot, Thierry Rouault, Re´mi Brun, Marcel Pelard, and Daniel Mercier

28 Preliminary Results on Larval Rearing the European Sturgeon, Acipenser sturio ...... 395 P. Williot, T. Rouault, R. Brun, M. Pelard, and D. Mercier xx Contents

29 Post-release Monitoring Techniques ...... 407 M.L. Acolas, Charles Roqueplo, E. Rouleau, and E. Rochard

30 Modelling the Future of Stocked Fish ...... 417 E. Rochard and Patrick Lambert

31 Building a Brood Stock of Acipenser sturio in France ...... 425 Patrick Williot, Thierry Rouault, Re´mi Brun, Marcel Pelard, Daniel Mercier, Louis Jacobs, and Frank Kirschbaum

32 Reproduction of the Cultured Brood Fish ...... 439 Patrick Williot and Patrick Che`vre

33 Recent Progress in Larval Rearing of the European Sturgeon, Acipenser sturio ...... 449 Patrick Che`vre, Julien Saint-Sevin, Daniel Mercier, Louis Jacobs, and Patrick Williot

34 Genetic Variability of Cultured European Sturgeon Acipenser sturio ...... 455 Ralph Tiedemann, Anja Schneider, Patrick Williot, and Frank Kirschbaum

35 Sperm Cryopreservation in Sturgeon with a Special Focus on A. sturio ...... 465 A´ kos Horva´th, Patrick Che`vre, and Be´la Urba´nyi

36 How Non-governmental Stakeholders Have Contributed to the Conservation Programme in France ...... 477 Didier Moreau

37 Why, How and Results from an Awareness Campaign Within Marine Professional Fishermen for the Protection of Large Migratory Fish, the European Sturgeon Acipenser sturio .... 489 Nicolas Michelet

38 The French–German Cooperation: The Key Issue for the Success of the Preservation and Restoration of the European Sturgeon, Acipenser sturio, and Its Signiﬁcance for Other Sturgeon Issues ... 499 Patrick Williot and Frank Kirschbaum Contents xxi

Part IV Restoration: Conservation Programmes of Acipenser oxyrinchus

39 Conservation and Restoration of Acipenser oxyrinchus in the USA ...... 517 John Waldman

40 Atlantic Sturgeon (Acipenser oxyrinchus oxyrinchus) Fishery Management in the St. Lawrence Estuary, Que´bec, Canada ...... 527 Guy Verreault and Guy Trencia

41 Remediation of Atlantic Sturgeon Acipenser oxyrinchus in the Oder River: Background and First Results ...... 539 Jo¨rn Gessner, Gerd-Michael Arndt, Frank Fredrich, Arne Ludwig, Frank Kirschbaum, Ryszard Bartel, and Henning von Nordheim

42 The Past and Future of Sturgeons in Poland: The Genetic Approach ...... 561 Ana Stankovic´

43 Re-establishing the Atlantic Sturgeon (Acipenser oxyrinchus oxyrinchus Mitchill) in Poland ...... 573 Ryszard Kolman, Andrzej Kapusta, and Arkadiusz Duda

Part V Perspectives and Prospectives

44 Population Conservation Requires Improved Understanding of In Situ Life Histories ...... 585 M.L. Acolas, J. Gessner, and E. Rochard

45 Potential Effects of Forthcoming Climate Change and Biological Introductions on the Restoration of the European Sturgeon ...... 593 Ge´raldine Lassalle, M. Be´guer, and E. Rochard

46 Population Viability Analysis of the European Sturgeon (Acipenser sturio L.) from the Gironde Estuary System ...... 603 Ivan Jaric´, J. Knezˇevic´-Jaric´, G. Cvijanovic´, and Mirjana Lenhardt

47 One Alternative to Germ Cells Cryopreservation: Cryobanking of Somatic Cells in Sturgeon ...... 621 Catherine Labbe, Alexandra Depince, Pierre-Yves Le Bail, and Patrick Williot xxii Contents

48 Some Ex-Situ-Related Approaches for Assessing the Biological Variability of Acipenser sturio ...... 635 Patrick Williot

Part VI General Conclusions

49 What Is the Future for the European Sturgeon? ...... 649 Patrick Williot, Eric Rochard, Nathalie Desse-Berset, Jo¨rn Gessner, and Frank Kirschbaum

Index ...... 663 Contributors

M.L. Acolas Cemagref, Estuarine Ecosystems and Diadromous Fish Research Unit, 50 Avenue de Verdun, 33612 Cestas Cedex, France, [email protected]

Gerd-Michael Arndt Fisch und Umwelt Mecklenburg-Vorpommern e.V., Fischerweg 408, 18069 Rostock, Germany

Ryszard Bartel Instytut Rybactwa S´ro´dla˛dowego, Zakład Ryb We˛drownych, ul. Syno´w Pułku 37, 80-298 Gdan´sk, Poland

M. Be´guer Cemagref, Estuarine Ecosystems and Diadromous Fish Research Unit, 50 Avenue de Verdun, 33612 Cestas Cedex, France, [email protected]

Patrick Berrebi Institut des Sciences de l’Evolution, UMR 5554 CNRS/UM2/ IRD, Universite´ Montpellier 2, cc065, Place Bataillon, 34095 Montpellier Cedex 05, France, [email protected]

Paolo Bronzi World Sturgeon Conservation Society, via Piave, 8, 20854 Vedano al Lambro, Italy, [email protected]

Laurent Brosse Aqua-Logiq, 527 rue Zac Petite Camargue, 34400 Lunel, France, [email protected]

Re´mi Brun Cemagref, Estuarine Ecosystems and Diadromous Fish Research Unit, 50 Avenue de Verdun, 33612 Cestas Cedex, France

Giuseppe Castaldelli Department of Biology and Evolution, University of Ferrara, via L. Borsari, 46, 44121 Ferrara, Italy

xxiii xxiv Contributors

Ge´rard Castelnaud Cemagref, Estuarine Ecosystems and Diadromous Fish Research Unit, 50 Avenue de Verdun, 33612 Cestas Cedex, France, gerard.castel- [email protected]

Stefano Cataudella Department of Biology, University of Tor Vergata, via della Ricerca Scientiﬁca, 00173 Rome, Italy

Olivier Chassaing Paleógeńe´tique et Evolution Molećulaire, Institut de Geńomi- que Fonctionnelle de Lyon, Universite´ de Lyon, Universite´ Lyon 1, CNRS, INRA, Ecole Normale Supe´rieure de Lyon, 46 alleé d’Italie, 69364 Lyon Cedex 07, France; Institut des Sciences de l’Evolution, UMR 5554 CNRS/UM2/IRD, Universite´ Mon- tpellier 2, cc065, Place Bataillon, 34095 Montpellier Cedex 05, France

Patrick Che`vre Cemagref, Estuarine Ecosystems and Diadromous Fish Research Unit, Moulin de la logerie, 33660 Saint-Seurin-sur-l’Isle, France; Cemagref, CREA de Saint-Seurin-sur-l’Isle, Moulin de la logerie, 33660 Saint-Seurin-sur-l’Isle, France, [email protected]

G. Cvijanovic´ Institute for Multidisciplinary Research, Kneza Visˇeslava 1, 11000 Belgrade, Serbia, [email protected]

Alexandra Depince INRA, UR1037 SCRIBE, Cryopreservation and Regenera- tion, F-35000, Rennes, France

Nathalie Desse-Berset Universite´ de Nice-Sophia Antipolis – CNRS-CEPAM- UMR 6130, SJA3, 24 avenue des Diables Bleus, 06357 NICE Ce´dex 4, France, [email protected]; [email protected]

Arkadiusz Duda Department of Ichthyology, Inland Fisheries Institute, Oczapowskiego 10, 10-719 Olsztyn-Kortowo, Poland

Marilyne Duffraisse Paleógeńe´tique et Evolution Molećulaire, Institut de Geńomique Fonctionnelle de Lyon, Universite´ de Lyon, Universite´ Lyon 1, CNRS, INRA, Ecole Normale Supe´rieure de Lyon, 46 alleé d’Italie, 69364 Lyon Cedex 07, France

Francesco Fontana Department of Biology and Evolution, University of Ferrara, Via L. Borsari 46, 44100 Ferrara, Italy, [email protected]

Frank Fredrich Leibniz-Institut fu¨r Gewa¨ssero¨kologie und Binnenﬁscherei, Mu¨ggelseedamm 310, 12587 Berlin, Germany

Christine Gazeau Cemagref, Estuarine Ecosystems and Diadromous Fish Re- search Unit, 33612 Cestas Cedex, France Contributors xxv

Jo¨rn Gessner Leibniz-Institute of Freshwater Ecology and Inland Fisheries, Mu¨ggelseedamm 310, 12587 Berlin, Germany, [email protected]

Catherine Ha¨nni Paleógeńe´tique et Evolution Molećulaire, Institut de Geńomi- que Fonctionnelle de Lyon, Universite´ de Lyon, Universite´ Lyon 1, CNRS, INRA, Ecole Normale Supe´rieure de Lyon, 46 alleé d’Italie, 69364 Lyon Cedex 07, France

A´ kos Horva´th Department of Aquaculture, Szent Istva´n University, 2100 Go¨do¨llo˝,Pa´ter K. u. 1., Hungary, [email protected]

Louis Jacobs Cemagref, Estuarine Ecosystems and Diadromous Fish Research Unit, Moulin de Logerie, 33660 Saint-Seurin-sur-l’Isle, France

J. Knezˇevic´-Jaric´ Ecological Society “Endemit”, Oracˇka 42, 11080 Belgrade, Serbia, [email protected]

Ivan Jaric´i Institute for Multidisciplinary Research, Kneza Visˇeslava 1, 11000 Belgrade, Serbia, [email protected]

Philippe Jatteau Cemagref, Estuarine Ecosystems and Diadromous Fish Research Unit, 33612 Cestas Cedex, France, [email protected]

Andrzej Kapusta Department of Ichthyology, Inland Fisheries Institute, Oczapowskiego 10, 10-719 Olsztyn-Kortowo, Poland

Frank Kirschbaum Faculty of Agriculture and Horticulture, Biology and Ecology of Fishes, Humboldt University, Philippstr. 13, Haus 16, 10115 Berlin, Germany, [email protected]; [email protected]

Ryszard Kolman Department of Ichthyology, Inland Fisheries Institute, Oczapowskiego 10, 10-719 Olsztyn-Kortowo, Poland, kolrys@inﬁsh.com.pl

Catherine Labbe INRA, UR1037 SCRIBE, Cryopreservation and Regeneration, F-35000, Rennes, France, [email protected]

Patrick Lambert Cemagref, Estuarine Ecosystems and Diadromous Fish Re- search Unit, 50 Avenue de Verdun, 33612 Cestas Cedex, France

Ge´raldine Lassalle Cemagref, Estuarine Ecosystems and Diadromous Fish Research Unit, 50 avenue de Verdun, 33612 Cestas Cedex, France; UMR 6250 LIENSs, University of La Rochelle – CNRS, Institut du Littoral et de l’Environnement, 2 rue Olympe de Gouges, 17000 La Rochelle, France, [email protected] xxvi Contributors

Pierre-Yves Le Bail INRA, UR1037 SCRIBE, Cryopreservation and Regeneration, F-35000, Rennes, France

Mirjana Lenhardt Institute for Biological Research, Despota Stefana 142, 11000 Belgrade, Serbia

Mario Lepage Cemagref, Estuarine Ecosystems and Diadromous Fish Research Unit, 50, Avenue de Verdun, 33612 Cestas Cedex, France, mario. [email protected]

Aude Lochet Cemagref, Estuarine Ecosystems and Diadromous Fish Research Unit, 33612 Cestas Cedex, France

Arne Ludwig Department of Evolutionary Genetics, Institute for Zoo and Wildlife Research, Alfred-Kowalke-Str. 17, 10315 Berlin, Germany, [email protected]

Daniel Mercier Cemagref, Estuarine Ecosystems and Diadromous Fish Research Unit, Moulin de logerie, 33660 Saint-Seurin-sur-l’Isle, France

Nicolas Michelet French National Committee for Marine Fisheries and Sea Farming (CNPMEM), 134 Avenue de Malakoff, 75116 Paris, France, nmichelet@ comite-peches.fr

Arturo Morales-Mun˜iz Laboratorio Arqueozoologia, Universidad Auto´noma de Madrid, Darwin, 2, 28049 Madrid, Spain

Didier Moreau WWF-France (consultant), 34 rue des Souche`res, 26110 Nyons, France, [email protected]

Jacek Morzuch Department of Ichthyology, Inland Fisheries Institute, Oczapows- kiego 10, 10-719 Olsztyn-Kortowo, Poland

Marcel Pelard Cemagref, Estuarine Ecosystems and Diadromous Fish Research Unit, Moulin de logerie, 33660 Saint-Seurin-sur-l’Isle, France

Gae¨l Pique`s Arche´ologie des socie´te´sMe´diterrane´ennes, UMR 5140 CNRS, 390 Avenue de Pe´rols, 34970 Lattes, France

Stefan Reiser Institute for Hydrobiology and Fisheries Science, University of Hamburg, Olbersweg 24, 22767 Hamburg, Germany

Eric Rochard Cemagref, Estuarine Ecosystems and Diadromous Fish Research Unit, 50 Avenue de Verdun, 33612 Cestas Cedex, France, [email protected] Contributors xxvii

Charles Roqueplo Cemagref, Estuarine Ecosystems and Diadromous Fish Research Unit, 50 Avenue de Verdun, 33612 Cestas Cedex, France

Eufrasia Rosello´-Izquierdo Laboratorio Arqueozoologia, Universidad Auto´noma de Madrid, Darwin, 2, 28049 Madrid, Spain

Remiggio Rossi Department of Biology and Evolution, University of Ferrara, via L. Borsari, 46, 44121 Ferrara, Italy

Thierry Rouault Cemagref, Estuarine Ecosystems and Diadromous Fish Research Unit, Moulin de logerie, 33660 Saint-Seurin-sur-l’Isle, France

E. Rouleau Cemagref, Estuarine Ecosystems and Diadromous Fish Research Unit, 50 Avenue de Verdun, 33612 Cestas Cedex, France

Julien Saint-Sevin Cemagref, Estuarine Ecosystems and Diadromous Fish Re- search Unit, Moulin de Logerie, 33660 Saint-Seurin-sur-l’Isle, France

Anja Schneider Unit of Evolutionary Biology/Systematic Zoology, University of Potsdam, Karl-Liebknecht-Straße 24-25 (Haus 26), 14476 Potsdam, Germany

S. Spratte Landesamt fu¨r Landwirtschaft, Umwelt und la¨ndliche Ra¨ume des Landes Schleswig-Holstein (LLUR), Abt. 3 Fischerei, Dezernat Binnenﬁscherei und Aquakultur, Hamburger Chaussee 25, 24220 Flintbek, Germany

Ana Stankovic´ Department of Biology, Institute of Genetics and Biotechnology, University of Warsaw and Institute of Biochemistry and Biophysics, Polish Academy of Sciences, Pawin´skiego 5A, 02-106 Warsaw, Poland, [email protected]

Catherine Taverny Cemagref, Estuarine Ecosystems and Diadromous Fish Re- search Unit, 50 Avenue de Verdun, 33612 Cestas Cedex, France

Ralph Tiedemann Unit of Evolutionary Biology/Systematic Zoology, Institute of Biochemistry and Biology, University of Potsdam, Karl-Liebknecht-Straße 24-25 (Haus 26), 14476 Potsdam, Germany, [email protected]

Guy Trencia Ministe`re des Ressources Naturelles et de la Faune, 8400 Sous-le-Vent, Charny, QC, Canada, G6X 3S9

Be´la Urba´nyi Department of Aquaculture, Szent Istva´n University, 2100 Go¨do¨llo˝, Pa´ter K. u. 1., Hungary

Guy Verreault Ministe`re des Ressources Naturelles et de la Faune, 186 rue Fraser, Rivie`re-du-Loup, QC, Canada, G5R 1C8, [email protected] xxviii Contributors

Henning von Nordheim Fachgebiet Meeres- und Ku¨stennaturschutz, Bundesamt fu¨r Naturschutz, Außenstelle Insel Vilm, 18581 Putbus, Germany

John Waldman Biology Department, Queens College, 65–30 Kissena Boulevard, Flushing, New York, NY 11367, USA, [email protected]

Patrick Williot Sturgeon Consultant, 4 Rue du pas de madame, 33980 Audenge, France, [email protected]

Sven Wuertz Leibniz-Institute of Freshwater Ecology and Inland Fisheries, Mu¨ggelseedamm 310, 12587 Berlin, Germany, [email protected]