Sturgeons and of Fish and Fisheries Series VOLUME 27

The Kluwer Fish and Fisheries Series begins with the new millennium. The volumes in this series will cover topics ranging from the biology of individual species or groups of fishes, to broader concepts in fisheries science, conservation and management. The Series is directed to professionals and researchers in fish biology, senior undergraduate and postgraduate students, and those concerned with commercial production or harvest of fishes.

It is difficult to know if this is the best of times or the worst of times for fish and fisheries. For example, many of the historically important marine fisheries are at or very near harvest capacity, according to the best scientific data and predictions. Many of the changes in commercial harvests fit a predictable, depressing pattern. We tend to produce simplified communities, harvested by ever more efficient technology, at increasing rates of exploitation. Some would suggest that nothing can stop the apparently inevitable destruction of all commercially harvested fishes. Fish seem to be increasingly degraded, and the deliberate or accidental introductions of exotic species threaten endemic native fishes. We always do things to the limit of our technology.

However, in contrast, we can point to a number of very favourable examples of current success and future promise in fish and fisheries. Our knowledge of the basic biology of fishes continues to expand. We disseminate that knowledge with ever increasing speed to libraries and personal computers around the world. Many fishes are increasingly recognized as fundamentally important subjects for basic research. Studies of the zebrafish, Brachydanio rerio, have produced a veritable explosion of fundamental scientific information at major research institutions around the world. Fishes as diverse as arctic charr, Salvelinus alpinus, and stickleback, Gasterosteus aculeatus, are providing insights and new understanding of the fundamental processes of natural selection and speciation. Science and technology give us a better understanding of the implications of long term climate change for fish populations. We continue to see fundamental breakthroughs in our understanding of development, genetics and evolution of fishes on almost a daily basis. Production of fishes through our increasingly sophisticated and efficient aquaculture rivals or exceeds the harvest of wild fishes in many places. Our knowledge and understanding continue to develop to the limits of our science and technology.

Science and the promise it holds for us to deal with our questions and concerns about fish and fisheries, is the basis for this Series. The future is certainly not what it used to be. This and forthcoming volumes in the Kluwer Fish and Fisheries Series will define the scientific basis for our future interactions with fishes. It is truly an exciting time.

Dr. David L. G. Noakes Series Editor, Fish and Fisheries Series Professor of Zoology, University of Guelph Guelph, Canada and Paddlefish of North America

Edited by

Greg T.O. LeBreton F. William H. Beamish and R. Scott McKinley

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Sturgeon and paddlefish are strange and wondrous . The cartilaginous skeletons, heterocercal tail, scaleless skin, and the presence of the notochord in adults distinguish members of the order from all other bony fish (Birstein 1993, Scott and Crossman 1998). Indeed, their large size alone makes them a curiosity for anyone witnessing these increasingly rare individuals. Beyond their very unique physical characteristics these fish also possess some unexpected behavioural actions. While conducting field research we have been privileged to capture lake of incredible size, perform biopsies, and return them to their river. We have always been struck by how remarkably placid and complacent these enormous creatures were. In the lab, juveniles demonstrate an uncanny similarity to puppies, swimming in summersaults, wagging their tails and watching with beady, reflective eyes at feeding times. Indeed, when fed a diet of floating pellets, these fish will learn to feed, while “belly-up”, from the water surface. This “unfishlike” behaviour often allows an attachment between researcher and subject that is unknown with other fish species. During the last 30 years there has been an increase in scientific interest in sturgeon and paddlefish in North America. Aquaculture and conservation of these species has spawned a large pool of scientific literature. Other research into the genetics, reproduction, morphology, and embryological development of these species has increased our understanding of these fish and provided us with a wealth of information. While there have been compilations and synthesis of literature on acipenserids published in the past these have included species from around the world (Birstein 1993, Birstein et al. 1997). The North American sturgeon and paddlefish present their own unique history, biology, ecology, culture, and conservation issues and for this reason we feel that it is important that this book, dedicated solely to those species, be published. This book, through the work and assistance of many authors has compiled what is currently known about the biology North American sturgeon and paddlefish. For 100 million years fish similar to modern day sturgeon and paddlefish lived in the waters that were to become North America. These evolved into two families, Acipenseridae and Polyodontidae. While the latter possess only one surviving species, the paddlefish, Polyodon spathula, nine species in Acipenseridae survive in two genera, and . These are currently recognized as , Acipenser brevirostrum; lake sturgeon, A. fulvescens; , A. medirostris; Atlantic sturgeon, A. oxyrinchus oxyrinchus, and their sub-species , A. oxyrinchus desotoi; , A. transmontanus; , Scaphirhynchus albus; shovelnose sturgeon, S. platorynchus; and the Alabama shovelnose sturgeon, S. suttkusi. These fish were an integral and important part of the aboriginal culture. Prior to European influences, humans were dependant on sturgeon for survival and they revered the species. Early settlers to North America viewed the sturgeon as a nuisance fish which damaged nets used to capture other, more valuable, species. Captured individuals were killed and discarded, used as fertilizer, or as an oily fuel in steamships (Houston 1987). Only when European descendants discovered the economic value of sturgeon flesh and were these fish almost completely eliminated from North America during a strikingly short period of time at the turn of the 19th century. v vi

Incredible levels of overfishing and destruction, in combination with the late maturation and intermittent spawning of these species, resulted in all populations being greatly reduced in numbers and distribution. The story of sturgeon and paddlefish extermination is one of human gluttony and greed. To be sure it is one of the most telling ways in which humanity has failed in our stewardship of the planet. The majority of chapters in this book focus on the various aspects of the biology, conservation, and culture of sturgeon and paddlefish and have been compiled from the latest published research material. These sections are designed to be used as a reference for readership from universities, colleges, and government agencies needing the most current of knowledge. This book also contains two chapters outlining the historical aboriginal uses and the decline of sturgeon brought on by overfishing. These chapters will appeal to anyone interested in understanding how these once prolific species have become so endangered. Perhaps the most remarkable fact surrounding the acipenserids of North America is that, though greatly reduced, their populations have survived almost 150 years of unchecked slaughter. With modern technology and human concern sturgeon are being successfully cultured in great numbers and indeed some wild populations have begun to recover. While the history of the North American species is dark, it is not without hope. It is possible, through our advanced understanding of the biology and importance of these species, that we might prevent their complete extinction. Many individuals have worked very hard to assist with this compilation, and each should be commended for their efforts and patience in getting this work published. The chapter written by Inga Saffron titled “The Decline of the North American Species” was graciously submitted for this work and was derived from the book “” published in 2002 by Broadway Books. The illustrations included by Paul Vecsei and photos submitted by many individuals have greatly added to the calibre of the work. It is with great appreciation that the principle editors would like to thank those involved in making this book possible. In addition, the editors would like to acknowledge the efforts of Colleen Bohan, Kevin Butterworth in collating and proof reading the final edition; Dalyce Epp for editing and production; and David Noakes for editing.

The editors are deeply indebted to Nancy Léonard (Editorial Coordinator) for her significant contributions to the review of manuscripts and the overall organizational structure of the book.

G.T.O. LeBreton, F.W.H. Beamish, and R.S. McKinley [Editors]

References

Birstein, V.J., J. Waldman and W.E. Bemis. eds. 1997. Sturgeon Biodiversity and Conservation. Developments in Environmental Biology of Fishes 17. Kluwer Academic Publishers, Dordrecht, Holland. 425pg. Birstein, V.J. 1993. Sturgeon and : Threatened fishes in need of conservation. Conservation Biology 7:773-787. Houston, J.J. 1987. Status of the Lake Sturgeon, Acipenser fulvescens in Canada. Canadian Field Naturalist 101(2) 171-185. Scott, W. B. and Crossman, E. J. 1998. Freshwater fishes of Canada. Galt House Publications, Ltd. pg 77- 100. Series Editor’s Foreword

There is nothing like a vintage automobile to attract public attention anywhere in North America - except sturgeons and paddlefish. And for many of the same reasons. They are striking in appearance, with an apparently ancient body plan, as Paul Vecsei’s illustrations so clearly illustrate. No matter how often we see a live sturgeon, or even the static image of one of these creatures we pause in whatever we are doing. Anyone can recognize a sturgeon or paddlefish. All sturgeons are relatively minor variations on what must be a successful morphological theme. They are the largest fishes we can encounter in freshwater. Leviathan. Monster. Huge, shadowy prehistoric figures rising slowly from the depths. I remember my first view of a live sturgeon in the wild. My colleague Bill Beamish and I carried out a multi-year study of lake sturgeon, Acipenser fulvescens, in northern Ontario. On our first day in the field we set large mesh gill nets. We lifted those nets early next morning and in our first net we had a sturgeon. Its distinctly shark-like body rose from the dark water of the Groundhog River and created a mixture of awe, enthusiasm and wonder that continues to this day. Sturgeons and paddlefish should not be living among us. Conventional wisdom says that they have all the attributes that doom them by human exploitation. They are slow- growing, long-lived, late-maturing, large sized and highly valued individually and as species. Their anadromous life cycles make them vulnerable to fishing harvest under the worst possible circumstances. Large mature adults are supremely vulnerable to capture during their spawning migration and nowadays the focus of such fishing is invariably on mature females. Females are killed at precisely the worst possible time, for the harvest of their maturing ovaries for caviar. Caviar, as Inga Saffron so aptly describes is the world’s most coveted delicacy. The history of caviar is inextricably intertwined with politics, economics and intrigue. The present of sturgeons and paddlefish cannot be understood without that perspective. She admirably provides that in this volume. The history of the utilization by these fishes by native North American peoples, and the present conservation issues and multijurisdictional management are major themes in this volume. Despite all that seems primeval and pessimistic, there is clearly reason for hope for North American sturgeons and paddlefish. Federal legislation is now in place in both the U.S.A. and Canada to identify and direct attention to the conservation and restoration of threatened species and those at risk. All sturgeons and paddlefishes of the world are probably at risk. The details might vary, but the general conclusions are the same for all species because of their common biological and ecological attributes. This volume shows the reason why there is hope, at least for North American species. Legislation and efforts to sensibly manage these species exist in North America and the social, cultural and political threats the species face elsewhere in the world are less serious here. The importance of multijurisdictional approaches to management is a key contribution of this volume. There are also contributions on the full range of scientific knowledge of North American sturgeons and paddlefish. Habitat and environmental requirements, ecomorphology, metabolism and swimming and their bioenergetics are reviewed in detail to place these fishes in their environmental context. Chapters on genetics, metabolism, respiration and embryology review those aspects of biology. Of course conservation and management are major contributions to this volume, as is the growing interest and involvement of intensive aquaculture. This volume is a landmark vii viii in our understanding and appreciation of these species. It lays the groundwork for continuing scientific studies of these fishes. Most importantly, it points to the way forward for the management and conservation of these magnificent animals.

Dr. David L. G. Noakes Series Editor, Fish and Fisheries Series Professor of Zoology, University of Guelph Guelph, Canada TABLE OF CONTENTS

Foreword ...... v

Series Editor's Foreword.……………………………………………………………...vii

Introduction: The Decline of the North American Species ...... 1 Inga Saffron

1. Native American Utilization of Sturgeon ...... 22 Tim E. Holzkamm and Leo G. Waisberg 1.1 Introduction...... 22 1.2 Areas Of North America Where Native Americans Harvested Sturgeon and Paddlefish ...... 22 1.3 Conclusion ...... 35 1.4 References...... 36

2. Distribution, Habitat, and Movements...... 40 Jennifer A. Wilson and R. Scott McKinley 2.1 Introduction...... 40 2.2 Distribution and Status...... 41 2.3 Habitat...... 53 2.4 Migratory Behaviour...... 62 2.5 Management ...... 67 2.6 References...... 69

3. Environmental Requirements, Preferences, and Tolerance Limits of North American Sturgeons...... 73 Joseph J. Cech Jr. and Serge I. Doroshov 3.1 Introduction...... 73 3.2 North American Species ...... 73 3.3 References...... 83

4. The Ecology and Functional Morphology of Feeding of North American Sturgeon and Paddlefish ...... 87 Michael J. Miller 4.1 Introduction...... 87 4.2 Feeding ecology of North American sturgeons and paddlefish ...... 88 4.3 References...... 100

5. Sturgeon Ecomorphology: A Descriptive Approach ...... 103 P. Vecsei and D. Peterson 5.1 Introduction...... 103 5.2 Evolution and Brief Overview of Family Acipenseridae...... 103

ix x

5.3 Form, Function, and Movement...... 104 5.4 Descriptive Morphology ...... 105 5.5 Ecomorphology...... 109 5.6 A New Ecological Guild: The Supra-Benthos Cruisers...... 109 5.7 Descriptive Morphology of North American Species...... 114 5.8 Intraspecific Morphological Variation...... 122 5.9 References...... 132

6. Embryology...... 134 Jessica A. Bolker 6.1 Introduction...... 134 6.2 History of developmental research in sturgeons ...... 134 6.3 Scope of modern research...... 135 6.4 Developmental stages and timing ...... 136 6.5 Major stages of embryonic development ...... 136 6.6 Developmental bottlenecks and abnormalities...... 142 6.7 Formation of the body axis, cns and heart ...... 143 6.8 Conclusion ...... 144 6.9 References...... 144

7. Swimming and Respiration ...... 147 Stephan J. Peake 7.1 Introduction...... 147 7.2 Behaviour, performance, and respiration in forced swimming tests...... 147 7.3 Morphological constraints on swimming activity and respiration ...... 155 7.4 Environmental constraints on swimming activity and respiration ...... 159 7.5 Management implications...... 163 7.6 References...... 164

8. Sturgeon and Paddlefish Metabolism ...... 167 Thomas D. Singer and James S. Ballantyne 8.1 Introduction...... 167 8.2 Digestion...... 167 8.3 Oxidative Metabolism...... 169 8.4 Carbohydrate Metabolism...... 171 8.5 Lipid Metabolism...... 173 8.6 Ketone Body Metabolism ...... 177 8.7 Nitrogen Metabolism ...... 177 8.8 Hormonal Control of Metabolism...... 178 8.9 Toxicant Metabolism ...... 180 8.10 Environmental Effects on Metabolism ...... 181 8.11 Evolutionary Considerations...... 187 8.12 References...... 188

9. Growth, Bioenergetics and Age...... 195 G.T.O. LeBreton and F.W. (Bill) H. Beamish 9.1 Introduction...... 195 9.2 Growth and Environmental Factors ...... 200 xi

9.3 Growth and Biological Factors ...... 205 9.4 Age, Aging, and the Use of Hard Tissues in Ecological Investigations...... 210 9.5 Conclusion ...... 213 9.6 References...... 214

10. Genetics of North American Acipenseriformes...... 217 Michael R. Robinson and Moira M. Ferguson 10.1 Introduction...... 217 10.2 Genomic Organization...... 217 10.3 Genome Evolution ...... 220 10.4 Transmission Genetics...... 221 10.5 Phylogenetics...... 222 10.6 Species Relationships...... 223 10.7 Population Genetics ...... 224 10.8 Conclusions and Future Prospects ...... 226 10.9 References...... 227

11. Multijurisdictional Management of Lake Sturgeon in the and St. Lawrence River ...... 231 Nancy J. Léonard, William W. Taylor, and Christopher Goddard 11.1 Introduction...... 231 11.2 Lake sturgeon in the Great Lakes-St. Lawrence River ...... 232 11.3 Successes of cooperative management for lake sturgeon ...... 236 11.4 Current management of lake sturgeon ...... 238 11.5 The potential for multijurisdictional management of lake sturgeon ...... 240 11.6 International perspective on lake sturgeon...... 241 11.7 Future Outlook...... 245 11.8 References...... 247

12. Conservation...... 252 Nancy A. Auer 12.1 Introduction...... 252 12.2 Status of Acipenseriformes species...... 252 12.3 Reasons for the Initial Decline of the Species in North America...... 254 12.4 Persistent Threats to Recovery of North American Species ...... 254 12.5 Recent Advances in Recovery of Acipenseriformes in North America...... 261 12.6 The Future for Sturgeon and Paddlefish in North America – Walking a Tightrope...... 268 12.7 References...... 271

13. Aquaculture ...... 277 Joel P. Van Eenennaam, Frank A. Chapman, and Peter L. Jarvis 13.1 Introduction...... 277 13.2 White Sturgeon Culture ...... 278 13.3 Wild Broodstock...... 278 13.4 Growout Systems...... 281 13.5 Feed and Feeding Methods ...... 282 13.6 Water Quality...... 284 xii

13.7 Domestic Broodstock Development ...... 284 13.8 Commercial Production ...... 292 13.9 Paddlefish Culture...... 294 13.10 Culture of Other Sturgeon Species...... 297 13.11 Challenges...... 300 13.12 Summary...... 302 13.13 References...... 303

Conclusion: Ensuring The Future Of Sturgeon ...... 312 Kevin G. Butterworth and Nancy J. Léonard References ...... 314

Identification Key to Sturgeons and Paddlefish of North America ...... 315 Martin Hochleithner and Paul Vecsei References ...... 318

Glossary...... 319