The Great Lake Sturgeon [1 ed.] 9781609173661, 9781611860788

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Copyright © 2013. Michigan State University Press. All rights reserved. The Great Lake Sturgeon, Michigan State University Press, 2013. ProQuest Ebook Central,

Copyright © 2013. Michigan State University Press. All rights reserved.

THE GREAT LAKE STURGEON

The Great Lake Sturgeon, Michigan State University Press, 2013. ProQuest Ebook Central,

Copyright © 2013. Michigan State University Press. All rights reserved. The Great Lake Sturgeon, Michigan State University Press, 2013. ProQuest Ebook Central,

Copyright © 2013. Michigan State University Press. All rights reserved.

THE GREAT LAKE STURGEON

Edited by Nancy Auer and Dave Dempsey

Michigan State University Press

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East Lansing

Copyright © 2013 by Michigan State University

i The paper used in this publication meets the minimum requirements of ANSI/ NISO Z39.48-1992 (R 1997) (Permanence of Paper). Michigan State University Press

p East Lansing, Michigan 48823-5245 Printed and bound in the United States of America. 19 18 17 16 15 14 13

1 2 3 4 5 6 7 8 9 10

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LIBRARY OF CONGRESS CATALOGING-IN-PUBLICATION DATA

The Great Lake sturgeon / edited by Nancy Auer and Dave Dempsey. pages cm Includes bibliographical references. ISBN 978-1-61186-078-8 (pbk. : alk. paper)—ISBN 978-1-60917-366-1 (ebook) 1. Lake sturgeon. I. Auer, Nancy A. II. Dempsey, Dave, 1957– QL638.A25G74 2013 338.3'72742—dc23 2012028146 Cover and book design by Charlie Sharp, Sharp Des!gns, Lansing, MI Cover image of sturgeon eggs on rocks is used courtesy of Nancy Auer and image of adult sturgeon is used courtesy of Andrew Muir. Michigan State University Press is a member of the Green Press

G Initiative and is committed to developing and encouraging

ecologically responsible publishing practices. For more information about the Green Press Initiative and the use of recycled paper in book publishing, please visit www.greenpressinitiative.org.

Visit Michigan State University Press at www.msupress.org

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Contents

prefaces . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . vii

Sturgeon: The Great Lakes Buffalo | Dave Dempsey . . . . . . . . . . . . . . . . . . . . . . . . . 1 Form and Function in Lake Sturgeon | Nancy Auer . . . . . . . . . . . . . . . . . . . . . . . . . 9 N’me | Jimmie Mitchell . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 The Lake Sturgeon as Survivor and Integrative Indicator of Changes in Stressed Aquatic Systems in the Laurentian Basin | Henry A. Regier, Robert M. Hughes, and John E. Gannon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27 Habitat, Foods, and Feeding | Edward A. Baker and Nancy Auer. . . . . . . . . . . . . 59 Recognizing the Genetic Population Structure of Lake Sturgeon Stocks | Amy Welsh . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79

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Restoration and Renewal: A Sturgeon Tale | Lauri Kay Elbing . . . . . . . . . . . . . . . 93 The St. Lawrence River Lake Sturgeon: Management in Quebec, 1940s–2000s | Pierre Dumont and Yves Mailhot . . . . . . . . . . . . . . . . . . . . . . . . . . 101 Bringing Us Back to the River | Marty Holtgren. . . . . . . . . . . . . . . . . . . . . . . . . . . 133 Sturgeon for Tomorrow | Brenda Archambo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 147 The Relationship between Lake Sturgeon Life History and Potential Sensitivity to Sea Lamprey Predation | Holly Muir and Trent M. Sutton . . . . . 153 Future Management and Stewardship of Lake Sturgeon | Nancy Auer . . . . . . . 173

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about the authors . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 187

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PR EFAC ES

Prefaces

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It was September 2007. Dave and I were both at an evening reception in the Seaman Mineral Museum at Michigan Technological University. Dave was the guest of honor and had just addressed a public and academic audience on his latest book, which was about the Great Lakes. His passion for the Lakes quickly became obvious. During the reception, I was introduced as the sturgeon biologist at MTU to Dave and he commented that his next idea for a Great Lakes book was one covering the lake sturgeon. Now my father used to say that I had a rubber face and could hide no emotion, but that night I hoped I had done a reasonable job at hiding my surprise, for my dream was to publish a book on lake sturgeon. I wanted some way to broaden and improve public understanding of and “liking” for this magnificent species, which I had studied for over 20 years. This was a pivotal moment for me, as until then I had found little time to move beyond a dream; writing takes time. I recall standing for a moment looking at Dave and saying to myself: Speak up, or forever you will lose an opportunity. So I shared with him my similar desire to publish a book and gain some n

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ground on preserving this great fish. To my surprise he suggested we put our heads together, and that very night he sent me an email with his proposed outline; I still have that email and remain ever grateful Dave said yes to a joint effort. Our intention with this text is to provide the reader with some history, biology and ecology, and human perspective, as well as suggest some future management ideas. Chapters included in this volume span stories from individuals who are relatively new to appreciating this largest of freshwater fish to those who speak for people whose lives and culture have been intertwined with lake sturgeon for decades or longer. The range of views runs from those who regard the sturgeon as a commercial species to those who wish to carve out a place for the species where it can live out a more natural existence like the wolves and moose of Isle Royale. Lake sturgeons are endemic to North America, so we have included authors from both the United States and Canada. The chapters are meant to allow readers to discover the inner beauty and mystery of a truly magical fish, one not often encountered or observed, one that some are striving to protect in a sustainable future. Caring for our natural resources takes relationship and respect, which can only be built when we understand how organisms and ecosystems are inextricably linked. We hope you will join us in that journey as you read through these chapters and grow to experience this marvelous fish.

DAVE DEMPSEY

Despite being a native of the Great Lakes Basin and working in the field of Great Lakes policy for a quarter century, I rarely gave thought to the lake sturgeon until the last 5 to 10 years. Fisheries didn’t escape my attention—salmon, lake trout, muskie, perch, and other species were impossible to ignore in the realm of policy. These are the sport of the great freshwater fisheries, these are the moneymakers, and these are the source of social and political controversy. And beginning in 1988, with the discovery of zebra mussels, invasive aquatic species vexed all of us who worked on, or cared about Great Lakes fish. But where was the sturgeon—so large and ancient, yet so invisible to me? My inattention was tantamount to a marine mammal policymaker overlooking whales. This personal fact is humbling for me to contemplate—and another illustration of the reason the lake sturgeon has had to claw back from near extinction in the last century. Right in front of us but sometimes left out of our policies and our hearts,

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the sturgeon is what it has always so magnificently been—it is we who are changing, by learning to appreciate it. That does not mean we will all appreciate it in the same way. Some will study it scientifically, some will incorporate it into their aboriginal lifeways as has been done for centuries, some will simply care for it, while others will want to harvest them sustainably. The sturgeon may be an icon, but each of us will behold a different fish. Each of us will enjoy a different relationship to the sturgeon—and it to us. In learning much more about the sturgeon in recent years, I have come to admire its resilience. I have also come to admire the people who treat sturgeon as a species worth working to conserve and protect. My coeditor, Nancy Auer, tops the list. She combines impeccable scientific ability, approach, and credentials with a deep regard for sturgeon. This combination of strengths has enabled her to add irreplaceably to the science of the sturgeon and to add incalculably to the Great Lakes community of sturgeon stewards. I am fortunate to know her. Because of what I have learned about the sturgeon on my own and through the review of the insightful data and feeling expressed about the species in the chapters that comprise this book, I will never again exclude them from my thoughts, my regard, and my sense of their place in the Great Lakes. I hope this book will promote a similar response in its readers. Although the lake sturgeon can rightfully take its place as a biological indicator of the health of the Great Lakes ecosystem, it must also take its place in the passions we all bring to caring for the place we love and protect for future generations.

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DAVE DEMPSEY

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Sturgeon: The Great Lakes Buffalo

The book of natural resource exploitation in the United States and Canada since European settlement contains multiple illustrations of a ravenous human hunger for fish and wildlife species that extinguished, or nearly so, that which it desired. Even the most casual student of conservation has heard the legend of the passenger pigeon. It goes like this: in the first half of the 1800s, unimaginable numbers of birds darken the sky as they pass overhead. After 50 years of unchecked market hunting followed by futile conservation work, the last passenger pigeon dies in a Cincinnati zoo in 1916. The passenger pigeon has plenty of company in the annals of catastrophic fish and wildlife consumption in North America. The most prominent example in the American imagination is the American bison, popularly known as the buffalo. Millions of bison ranged over millions of acres of the continent in the early 1800s. Desired primarily for their hides and meat, bison fell at the hands of market hunters by the millions for several consecutive decades. By the 1880s, they were rarely sighted anywhere in their native habitat. Managed enclosures and zoos were the bison’s remaining living space. The species’ monuments were stacks of skulls and piles of bones. n

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Said Theodore Roosevelt in a work published before his presidency: “A merciless and terrible process of natural selection, in which the agents were rifle-bearing hunters, has left as the last survivors in a hopeless struggle for existence only the wariest of the bison and those gifted with the sharpest senses” (Roosevelt 1893, 28). The bison is a magnificent creature, capable of stunning speed, gifted with a noble head. Its association with the European settling of the American West lent it romance and won it a reprieve. Today the survival and recovery of the bison, even the sale of bison burgers in grocery stores and restaurants, comforts the public with its narrative of near-extinction and successful last-gasp conservation. Humans, the tale suggests, may not only rescue, but also even restore a part of nature’s plenty. Yet even in their fondness for the symbol of bison, Americans know little about them. Of 2,000 Americans who completed a Wildlife Conservation Society questionnaire in 2008, fewer than 10 percent knew how many bison remain in the United States, but 74 percent “believed that bison are extremely important living symbols of the American West.” The Society called the public “woefully out of touch with the species’ prospects for long-term survival” (Wildlife Conservation Society 2008). A hundred years ago, the bison was safely, if not abundantly preserved. But the ultimate fate of sturgeon in the Great Lakes was unknown. It lacked, as yet, a meaningful human constituency for protection. Victim of contempt and plunder in turn, the lake sturgeon was frighteningly vulnerable to extinction. Many regarded it as ugly. It was far from being an example of what is now known as charismatic megafauna—a classification embracing polar bears, grizzly bears, cougars, wolves, and great apes (Buckley 2009). But as we have learned, the lake sturgeon is every bit as appealing as these stars of the natural world in its own way, and just as much a barometer of health for the ecosystem it inhabits. It may be useful to trace the arc of human appreciation that begins in the late nineteenth century in the Great Lakes. Before that, the sturgeon of the Great Lakes was often considered a nuisance. Thought to consume spawn of valuable species, the sturgeon got in the way of catching whitefish and had few known commercial uses. Looking back decades later, the Michigan Department of Natural Resources argued with only slight hyperbole that “no single animal was ever subjected to such deliberate wanton destruction as the lake sturgeon. By the time it finally became recognized as a valuable fish, it had largely been destroyed as a troublesome nuisance” (Michigan Department of Natural Resources 1973, 51). Sturgeon incidentally caught in nets were destroyed. Sometimes they were stacked in rows, dried, and burned. Some were used as fuel for boat boilers. Others were served up as pig feed or used to fertilize soil.

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But the sturgeon, it turned out, could serve humans tangibly. A Canadian fisheries commissioner lamented the only gradually growing esteem of his countrymen for the lake sturgeon: “The Sturgeon is hardly appreciated at its true value in Ontario, the greatest proportion of the fish caught in Canadian waters being shipped to the States for sale. It is a fish nevertheless, of high economic importance, its flesh being of excellent nutritive quality and good though somewhat meaty flavor. The sounds or air-bladders furnish the best quality isinglass, and the roe the expensive delicacy ‘caviare,’ but these accessory products are not properly taken advantage of in the Province” (Wright 1892, 441). An entry from the journal of John W. Kerr, a Canadian fishery official housed in Toronto, shows excitement about sturgeon harvest was growing. February 15, 1882: Sturgeon fishing at Niagara (with hooks and lines) is the biggest thing in fishing that has happened yet. Great success. One sturgeon, dressed weight 65 lbs. Dressing: first the head is taken off, tail and fins cut off, guts removed, then fish is skinned. Two original fishermen had caught and sold more than $300 worth of sturgeon. “Buffalo

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Boys” taught Niagara fishermen how to catch sturgeon. Niagara is a money-making place.

Under this pressure, remarkable abundance swiftly passed from the scene. The smoking of sturgeon brought the species into commercial demand. The development of sturgeon caviar, the use of sturgeon hides for leather, increased use of sturgeon oil, and the production of carriage glass using gelatin derived from the swim bladders of sturgeon increased appreciation for the fish. By 1888, a state agency observed, “The once despised sturgeon has become one of the most valuable, commercially, of the many fish that are caught in the great lakes and deep rivers of this state. . . . Nearly every part of it is utilized in some way” (Michigan State Board of Fish Commissioners 1888). But the newly discovered value of the sturgeon only increased its slaughter. From an 1880 catch of 4.3 million pounds, sturgeon harvests fell in the next 20 years to 140,000 pounds (Michigan State Board of Fish Commissioners 1888). By the early part of the twentieth century, some lake populations of the sturgeon had collapsed. For example, officials closed the Lake Michigan sturgeon fishery in 1929 when the catch fell to 2,000 pounds, down from 3.8 million pounds 50 years earlier (U.S. Fish and Wildlife Service 2009). Reckless, unsustainable fishing was only the direct cause of sturgeon’s demise. The destruction of spawning habitat for navigation and the construction of hundreds of dams that prevented spawning runs also contributed significantly. In the days long before the sturgeon came to be valued as a part of Americana, it even played a small role in the American pastime. “In the lake regions and other

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sections of the country where sturgeon were plentiful, baseballs were commonly made of the eyes of that fish. The eye of a large sturgeon contains a ball nearly as large as a walnut. It is composed of a flexible substance and will rebound if thrown against a hard base. These eyeballs were bound with yarn and afterward covered with leather or cloth. They made a lively ball, but were more like the dead ball of the present than any ball in use at that time” (Morris 2006, 271). The sum of all destructive forces affecting the sturgeon seemed likely to consign it to memory. “With the drastic decrease in lake sturgeon abundance, most fisheries managers and ecologists believed in the early to mid 1800s that lake sturgeon would eventually disappear as a result of compounding negative pressures. However, lake sturgeon in the Great Lakes proved to be more resilient than previously assumed” (Léonard, Taylor, and Goddard 2004, 232). Indeed. The survival of the lake sturgeon was synchronous with a remarkable rebound of the Great Lakes. Beginning in the late 1960s, public investments in sewage treatment, enforcement of strict environmental laws affecting business, and dawning environmental awareness among millions of Canadians and Americans reflected in stewardship actions contributed to a remarkable, visible recovery of the Great Lakes. As algal blooms abated in the lower Great Lakes, introduced salmon replaced unwanted alewives as a dominant species, and bald eagles began reproducing in earnest, the notion of a robust natural world took popular root. “It’s amazing how resilient natural processes are once we allow them to work,” said my friend, Elizabeth Harris, then executive director of the East Michigan Environmental Action Council in Bloomfield Township, Michigan, in the 1990s. The last decade of the twentieth century offered hope for all life in the Great Lakes after a near ecological collapse at the midpoint of the same century. The subject of careful census and study since the 1960s also, the lake sturgeon assumed significance before the new millennium began as a bellwether or indicator species for the Great Lakes. Their reintroduction in historic spawning habitats made the sturgeon, as one reporter put it, “a mascot” for a river’s recovery (Moule 2008). Evading extinction by clinging successfully to their last viable spawning grounds, the sturgeon, at an estimated 1 percent of its historic numbers, was tougher than the careless despoilers of 100 years before might have thought. In their persistence they appealed to the human heart; and in the light of a new historical narrative they became beautiful. Sensing the mood, elected officials quickly congratulated their partners and themselves for helping rescue the species. “We are indeed so proud to be part of this international success story of recovery of lake sturgeon in our shared Great Lakes waters,” said Canadian member of Parliament Jeff Watson. “It is so heartening to

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see the amazing success of this sturgeon habitat restoration for the Detroit River International Wildlife Refuge,” noted Congressman John D. Dingell. “No one thought this degree of success was possible only 30 years ago” (Friends of the Detroit River 2009). This is well known. But can a species whose tentative comeback depends not only on expert fisheries biologists but also on the fluctuation in government spending to support their work sustain that recovery? Can the sturgeon of the Great Lakes depend on its human constituency for continuing affection and conservation? The signals are mixed. The first aquarium in the United States devoted to freshwater fish, the Great Lakes Aquarium in Duluth, Minnesota, opened in 2000 and struggled for years to keep its doors open. Its heavy debt load was one reason, but it also had difficulty attracting a tourist following. Fish of the Great Lakes disappointed some visitors. In 2002, the aquarium’s managers “discussed adding saltwater exhibits for more pizzazz, such as bringing in a shark to compare it with a sturgeon” (Associated Press 2002). When Toronto, Ontario, considered its own aquarium in 2005, skeptics said it was a dubious proposal because it would lack “charismatic attractions” like whales, dolphins, and other marine mammals. By 2009, the Duluth facility had an Amazon exhibit and a seahorse display. The Toronto aquarium has not been built. As much as most anglers, scientists, and nature lovers value the restoration of lake sturgeon for their intrinsic value, poaching is no more eradicated than the species itself. In spring 2009, in plain sight of observers on the Grand River in downtown Grand Rapids, Michigan, a lawbreaker reeled in a five-foot sturgeon and drove away with it in spite of strict regulations (Grand Rapids Press 2009). “Given its size, it would have to be a fairly old fish,” Michigan Department of Natural Resources conservation officer Dave Rodgers said. “So losing even one can have an impact on the sturgeon population.” The 36- to 40-pound fish was one of only 35 to 40 thought to have spawned in the river in the spring of 2009. Some of those noting the recovery appear ambivalent. “There are tons of sturgeon in western Lake Erie, especially around the mouth of the Detroit River,’ said one commercial fisherman in 2005, echoing the complaints of his predecessors 150 years earlier. “They’re always tearing the hell out of our nets. I betcha there’s a lot more sturgeon there than they can imagine” (Currie 2005). A counterweight: the enthusiasm of some sturgeon protectors could be contagious, especially with children. The manager of the Detroit River International Wildlife Refuge, John Hartig, called the sturgeon “a show-stopper” for kids. “When you take kids and show them a fish that is five or six feet long, they are blown away. It’s a living dinosaur. It’s been around that long. They ask, ‘How has that thing survived when so many other things have gone extinct?’” (Henry 2009).

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The lake sturgeon was also the reasonably charismatic star of a popular IMAX documentary, Mysteries of the Great Lakes, which audiences across the region enjoyed in 2008 and 2009. With the haunting steel guitar of Gordon Lightfoot’s Wreck of the Edmund Fitzgerald in the background and soaring vistas of vast Great Lakes waters as a visual prelude, the movie lovingly portrayed the sturgeon run in a Great Lakes tributary and its human admirers. Elementary school teachers working off the documentary were given materials to help them tell the students “about the water and history of the Great Lakes and some of the aspects that make it an important and unique resource for us all. The common theme throughout all parts of this resource is our Great Lakes friend, Sally Sturgeon. Sally is a lake sturgeon. Sally is over 120 years old. Given everything Sally has been through it is amazing that she has survived so long.” It is; and it is equally amazing that the sturgeon is able to use some of humanity’s past mistakes in the Great Lakes as bootstraps for its recovery. A 2004 Michigan Department of Natural Resources report explained a reproducing population of sturgeon in Lake St. Clair’s North Channel was assisted by an artificial spawning reef: “The coal cinders at the North Channel site are believed to have been deposited during the late 1800s when coal-burning vessels moored to load salt from a nearby factory and emptied their cinders into the river. The cinder substrate is now zebra mussel encrusted, and the three-dimensional structure of the cinders combined with the zebra mussel layer provide a complex system of interstitial spaces that appears to provide excellent protection for deposited eggs and fry” (Thomas and Haas 2004, 9). The combination of an unconsciously crafty and opportunistic natural world and human fancy may well perpetuate the sturgeon’s good fortune. But while the behavior of nature is largely unalterable, the future of human beings is unknowable. Will our fancy again turn against the fish? “The outlook for lake sturgeon recovery rangewide is guardedly optimistic,” a recent scientific paper observed, “thanks in part to renewed interest in the species, novel approaches to management, new opportunities to eliminate long-standing data gaps, and continued progress in habitat restoration. Recent emphasis on maintaining biodiversity has prompted several new management initiatives to ‘bring back the natives . . . ’ Our guarded optimism, however, is not intended as an ‘all clear’ regarding threats facing the species” (Peterson, Vecsei, and Jennings 2007, 72–73). The buffalo/bison may be safe, but the lake sturgeon is not yet. Its populations are tenuous, its reproductive biology a liability, and the affinity of the top beast in the Great Lakes food chain always subject to change. What seems clear is that given half a chance, the lake sturgeon will add to its ancient Great Lakes history.

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REFERENCES

Associated Press. 2002. Great Lakes aquarium plagued by money woes. September 9. Http:// www.greatlakesdirectory.org/mn/090902_great_lakes.htm. Buckley, C. 2009. New love for the endangered uglies? June 29. Http://www.esa.org/ esablog/?tag=charismatic-megafauna. Currie, P. 2005. Great Lakes legend makes a comeback. Toronto Star, February 22. Http://www. greatlakesdirectory.org/on/022205_great_lakes.htm. Friends of the Detroit River. 2009. The lake sturgeon have arrived! (news release) May 22. Http:// www.detroitriver.org/fdr-pdf/2009-SturgeonAnncmtFinal_22May09.pdf. Grand Rapids Press. 2009. State offers $1,000 reward to land Grand River angler who took five-foot sturgeon. June 3. Http://www.mlive.com/outdoors/index.ssf/2009/06/ state_offers_1000_reward_to_la.html. Henry, T. 2009. Freshwater species making comeback in Great Lakes region. Toledo Blade, October 26. Http://www.toledoblade.com/apps/pbcs.dll/article?AID=/20091026/ NEWS16/910260310. Kerr, J. W., and F. Kerr. 1864–1892 and 1895–1896. Notes about lake sturgeon in Lakes Ontario, Erie and Huron and associated rivers in the diaries of John W. and his son Fred Kerr, Canadian Federal Fishery officials headquartered in Toronto. Léonard, N. J., W. W. Taylor, and C. Goddard. 2004. Multijurisdictional management of lake sturgeon in the Great Lakes and St. Lawrence River. In: Sturgeons and paddlefish of North America. G. T. O. LeBreton, F. W. H. Beamish, and R. S. McKinley, eds. Kluwer Academic Publishers. Michigan Department of Natural Resources. 1973. Michigan Fisheries Centennial Report, 1873–1973. Michigan Department of Natural Resources, Management Report 6. Copyright © 2013. Michigan State University Press. All rights reserved.

Michigan State Board of Fish Commissioners. 1888. Ninth report of state fish commissioners. Morris, P. 2006. A game of inches: The stories behind the innovations that shaped baseball. Ivan R. Dee. Moule, J. 2008. There’s something fishy in the Genesee—again. City Paper, December 24. Http:// www.rochestercitynewspaper.com/news/articles/2008/12/Theres-something-fishy-in-the/. Peterson, D. L., P. Vecsei, and C. A. Jennings. 2007. Ecology and biology of the lake sturgeon: A synthesis of current knowledge of a threatened North American Acipenseridae. Reviews in Fish Biology and Fisheries 17:59–76. Roosevelt, T. 1893. The American bison. In: Hunting the grisly and other sketches. Putnam and Sons. Thomas, M. V., and R. C. Haas. 2004. Abundance, age structure, and spatial distribution of lake sturgeon Acipenser fulvescens in the St. Clair system. December. Http://www.dnr.state. mi.us/publications/pdfs/ifr/ifrlibra/research/reports/2076rr.pdf. U.S. Fish and Wildlife Service. 2009. Great Lakes Lake Sturgeon Web Site. Http://www.fws.gov/ midwest/sturgeon/biology.htm. Wildlife Conservation Society. 2008. Survey says: Let bison roam. November 18. Http://www.wcs. org/new-and-noteworthy/survey-says-let-bison-roam.aspx. Wright, R. R. 1892. Preliminary report on the fish and fisheries of Ontario. In: Commissioners’ report, Ontario Fish and Game Commission. Warwick & Sons. (Henry A. Regier deserves thanks for unearthing this report.)

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Form and Function in Lake Sturgeon

The lake sturgeon, Acipenser fulvescens, was first described in 1818 by a botanist from Turkey named Constantine S. Rafinesque. He encountered lake sturgeon during a survey of the flora and fauna of the Ohio River (Rafinesque 1820). The lake sturgeon is the only endemic sturgeon of the genus Acipenser found throughout the three closely related, freshwater drainage basins in North America, those of the Mississippi River, Great Lakes, and Hudson Bay (Ferguson and Duckworth 1997). Eight other species or subspecies of sturgeons in two genera are recognized in North America (table 1). The sturgeons are one of the oldest fishes on earth, bridging evolutionary time between the closely related and similar-looking sharks (fishes with full cartilaginous skeletons) and the early true bony fishes first represented by the gars and bowfins (figure 1). Sturgeons possess some bone in the form of plates, called scutes, on their body surface and head that have persisted in the geologic record (plate 1). Fossils of sturgeon scutes date to between 100 to 200 million years of age, placing these fish on earth during the age of dinosaurs, the late Cretaceous (Hilton and Grande 2006).

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Table 1. North American Sturgeon Species Ordered by Known Maximum Size

LOCATION IN NORTH AMERICA

COMMON NAME

SCIENTIFIC NAME

West Coast

White

Acipenser transmontanus

East Coast

Atlantic

Acipenser oxyrinchus

60

Midwest

Lake

Acipenser fulvescens

154

Gulf Coast

Gulf

Acipenser oxyrinchus Desotoi

West Coast

Green

Acipenser medirostris

> 45†

2000‡ / 79

Midwest Rivers

Pallid

Scaphirhynchus albus

40

1900 / 75

East Coast

Shortnose

Acipenser brevirostrum

67

1200 / 47

Midwest Rivers

Shovelnose

Scaphirhynchus platorynchus

27

1000 / 39

Alabama & Mississippi Alabama

Scaphirhynchus Suttkusi

MAX. AGE (YEARS)

> 80

30*

unknown

MAX. TOTAL LENGTH (MM/IN.)

6500 / 2560 4100 / 161 2800 / 110 2400 / 94*

762 / 30§

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Sources: Data from LeBreton, Beamish, and McKinley 2004 unless otherwise indicated. *http://www.flmnh.ufl.edu/fish/Gallery/Descript/gulfsturgeon/gulfsturgeon.html † http://www.krisweb.com/biblio/klamath_usfws_nakamotoetal_1995_sturgeon.pdf ‡ http://www.nmfs.noaa.gov/pr/pdfs/species/greensturgeon_detailed.pdf § http://library.fws.gov/Pubs4/alabama_sturgeon.pdf

Figure 1. Large adult lake sturgeon at spawning time, Houghton, Michigan 1990. (Photo by N. Auer.)

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Why Have Sturgeon Survived for So Long? The 27 species of sturgeons known to live in the Northern Hemisphere have survived due to a combination of several life history strategies. Most sturgeons fill an ecological niche by retaining, from their shark ancestors, several features that have persisted over time, withstood the power of natural selection and proven beneficial to survival (Auer 2004). The sturgeons have combined several life history strategies, which include the following:

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• Large body size and shape, large muscle segments and oil-rich connective tissue that assist buoyancy and energy-efficient swimming • Early life development of fast growth and sharp boney scutes that help deter predators • Maturation of reproductive organs later in life and intermittent spawning • Feeding on benthic organisms and organic material, which allow a more passive feeding strategy, requiring less physical energy than a typical predator • Occupying the light-limited, bottom waters of lakes and large rivers that protect them from predators and unusual fluctuations in water temperature • Living to a great age, which allows them, over short or long time periods, to persist through flood, drought, warming, or cooling events. Fishing pressure, barriers, and dams blocking spawning migration routes and persistent chemical contamination are human impacts that have and continue to reduce the future success of this unusual organism in its remaining habitats. Let’s take a closer look at some of these life strategies.

Body Plan Sturgeons represent the transition between fishes with only cartilage, like sharks and rays often called elasmobranchs, and the true bony fishes most often eaten or caught in sport fisheries today. Sturgeon keep a similar body design to that of the shark, but instead of retaining oil in a large liver, as do the sharks for buoyancy, these fish possess large muscle segments and a rich oil between these segments that was either boiled off or smoked off prior to consuming the flesh (Harkness and Dymond 1961). The sturgeons also possess an air bladder that produces a gelatinous substance called isinglass that was used in clarifying beer and wine and in jam and jelly preparation (Harkness and Dymond 1961). The oil, muscle, and air bladder provide the body

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Figure 2. A sturgeon lifting off the bottom of a lake or stream might resemble an airplane at takeoff, using downward pressure by the tail and lift generated by pectoral fins as it moves forward. (Drawing by N. Auer.)

structure and buoyancy needed to move the fish without a bony internal support system. The bony plates they do have on the head and along the sides of the body in five rows are believed to be for protection, as the scutes have razor-sharp edges when the fish are very young. The scutes become dull and rounded with age. Sturgeons spend most of their lives cruising just above the lake, ocean, or river bottom, and must journey upstream during spring to spawn. However, they have adapted to using the currents and their body composition (high oil content and large air bladder) to their advantage for movement. A sturgeon starting to swim from a stopped position looks just like an airplane at the time of takeoff (Wilga and Lauder 1999) (figure 2). The sturgeon tail is broader on the top than at the bottom (termed heterocercal), and a single tail beat will actually push the back end of the fish down. Combine this with a small push up from the unusually large pectoral fins, and the streamlined head lifts into the current just as a plane takes off into the wind. Small energy-efficient pushes allow the sturgeon to use very little energy to lift itself into

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movement. The shark and sturgeon body plans are some of the most efficient for cruising over bottom substrate. Modern fishes such as salmon and sunfish can regulate their depth position in the water by direct secretion of gases into the air bladder from the blood. Since sturgeons are more primitive, they lack this ability and must gulp air to keep their air bladder inflated. When they inhabit deep water for any period of time they begin to lose air from the bladder due to hydrostatic pressure (weight of the water column above the fish creates pressure as scuba divers experience) so they rise or even rush to the surface and can be seen rolling or jumping to obtain air (Watanabe et al. 2008). The sturgeons have also kept impressive sensory development to achieve their successful life history strategy. Since these fishes live in deep, dark, turbid waters, they are believed to be nearsighted, yet they are sensitive to light. When very young, most sturgeon are photonegative (move away from light), often seeking out dark crevices and holes during daylight hours. Even as they mature they seem to avoid strong light, which is advantageous as potential predators less easily see them. Some features of the sturgeon eye confirm its place in the evolutionary tree between sharks and bony fishes. Sturgeons possess a feature typical of many early fish such as sharks called the tapetum lucidum (Nicol 1969). This name refers to the special cells, filled with a crystalline material in a layer just behind the retina in the eye, that reflect light back into the eye and help concentrate light and improve vision at night or in murky waters. This layer is common in many animals but obvious in sharks, cats, and deer. Sturgeons also have a simple retina dominated by rods and with single cones, in contrast to what is seen in later bony fishes, which have a more complex retina with multiple cone photoreceptors (Rodriguez and Gisbert 2001, 2002; Sillman et al. 2005). Eyes with mostly rods are generally found in animals adapted for life in low-light conditions (Sillman et al. 2005). The arrangement of the pigment in the sturgeon cornea also helps reduce eyeshine upward out of the pupil, making the eye less conspicuous when viewed from overhead (Nicol 1969), where other predatory fishes may be cruising. Sturgeons also have well-developed smell and taste receptors located in their nostrils and on the surface of the barbels and bottom of the head near the mouths (plate 2b). As they cruise along the lake and river bottom, they use their barbels to gather sensory information on what is on or near the substrate surface (Harkness and Dymond 1961). Sturgeons are known to consume all manner of organic debris, typically eating worms, small clams, crayfish, and decomposing fishes on the lake bottom. Some studies have even found sturgeon stomachs to contain items such as onions, corn or wheat from grain elevator or railroad spills, and cigarettes (Harkness and Dymond 1961). Recent reports indicate that adult sturgeons may be one of

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the few organisms that consume the exotic invasive zebra mussel (U.S. Geological Survey 2008). Sturgeon are known to be curious, often coming to investigate ice-fishing decoys (Harkness and Dymond 1961). Whether they see or sense these objects through sound or vibration traveling through the water and sensed through their large air bladders has yet to be determined. Sturgeon are also known to be sensitive to electrical transmission, migration routes possibly influenced by overhead transmission lines (J. Hayes, SUNY ESF, pers. comm).

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Growth Sturgeon are typically thought to be slow-growing fishes, but during their first year of life grow quickly, increasing in length from about 25 mm (one inch) at hatch to 125 mm (five inches) at about five months of life. Young hatch from small, ⅛-inchdiameter eggs spawned in the springtime in the rapids of large river systems (plate 3). The eggs and sperm are released together by the adult male and female fish, and fertilized eggs roll and adhere to the undersides of clean rocks. Depending on water temperature, the eggs will hatch in about five to eight days. Once hatched, the little fish burrow into the river gravel where it is dark and for a few days slowly absorb the yolk in the yolk sac remaining from the egg (plate 4). Once the yolk is gone, the little fish must begin to find food and start moving from the spawning location by rising out of the gravel at dusk and drifting with the current for a few miles before settling again and feeding and hiding during the daytime. This drift can occur for great distances until the young find a rich organic region within the river or near the river mouth in which to feed. Some young stay in rivers to feed, while others appear to move out into connecting lake systems, usually systems with large productive deltas. Some characteristics of sturgeon early in life, such as what they feed upon and exactly where they spend time, are still being investigated. Lake sturgeons will utilize both river and lake environments once they grow beyond their first year of life. They are usually found over sand or sand and pea gravel substrates and seem to feed on organic matter and organisms drifting with the currents (Harkness 1923). They remain juveniles, much like humans, for almost 15 to 20 years before they reach sexual maturity and return to natal rivers to spawn.

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Reproduction All sturgeon species are believed to spawn in their natal river, locating it by imprinting to chemical cues during early life much like salmon and trout. It is amazing that a fish 15–20 years old can return to a river after such a long absence. This is one of the reasons it is important to keep river corridors unblocked and river environments clean and healthy for fishes that must find their way back to natal spawning areas. In northern Michigan, lake sturgeon males reach maturity at about age 15 or 130 cm TL, while females take a bit longer and might be 20 years old before they first spawn at about 140–150 cm TL (Auer 1999). This strategy is beneficial as it increases the probability that male and female siblings will likely not spawn together in the same years and thus limit genetic variability. Males are found to spawn about every other year, while females spawn once every five to nine years. That is because it takes time and energy for a fish to produce eggs. A 25-year-old, 50-pound female may contain 250,000 eggs (Harkness and Dymond 1961), the resources for which are slowly accumulated over the long period away from the spawning river. The loss of one female through accident or poaching, therefore, can drastically impact a small, remnant stock of sturgeon. Producing and spawning a single large batch of eggs over a long and intermittent time frame is beneficial as a reproductive strategy for a fish under natural circumstances. Migrating to and spawning in rivers for fishes exposes them to possible predation, usually means food is limited, and costs a great deal of energy. By spawning in a stream in which they were originally spawned, the need to expend energy searching for adequate habitat is reduced. The site where they were spawned most probably had adequate water flow, temperature, and oxygen for egg development. The adults become concentrated in the rapids areas to spawn, so mate finding is easy. They depart the river area quickly after spawning, which reduces any possible predation on eggs of their own species as well. However, this strategy of reproduction can be compromised when rivers are influenced by human and industrial discharges, water withdrawals, or barriers.

Feeding Sturgeon feed along lake and river substrate, detecting organic matter with their four barbels, which are positioned in front of the mouth (plates 2a and 2b). They have no teeth, so they actually suck organic material and organisms, along with muck and sand, into the mouth. The sand and muck is rejected out the gill openings, while

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Figure 3. View of spiral valve in sturgeon stomach, showing continuous movement of material through the stomach, increasing absorption area. In this rendering, the fish mouth would be at the top of the image, and the narrower passage at the bottom would be close to the anus. (Drawing by. N. Auer.)

the organic material is swallowed. Sturgeons have a unique stomach with what is known as a spiral valve (figure 3), which reduces the need for a long gut to slowly digest food, thus saving internal space for energy storage and reproductive organs. Food in the spiral valve gut circulates over a spiral path, similar to that of a spiral staircase; the gut itself is full of folds to increase surface area and absorption of nutrients. Sturgeons are passive, opportunistic feeders. They usually don’t search out and track down organisms, but rather situate themselves in rich environments (lake shorelines, river mouths, wetlands, etc.) and feed on what is encountered there. This strategy requires less physical energy than that of a more aggressive predator like a salmon or trout seeking forage fishes on which to feed, so energy can be conserved and placed into growth and reproductive products. Feeding on a diverse diet of various organic products is also a good strategy for survival. When we think of food webs or food chains we often think of the typical piscivore (fish eating) or herbivore (vegetation eating) predator/prey dependent cycle, such as that of the lake herring or the cisco feeding on zooplankton or lake trout feeding on rainbow smelt. However, if for some reason such as disease or overharvest, the forage item (small fish or vegetation) become less abundant, then the specialized predator will also decline in number. A diverse diet, low on the food web, allows the lake sturgeon to find some type of food throughout almost the entire year.

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Habitat Most sturgeons select for environments with less light intensity than that found in surface or clear, open water. These fish are often found in large turbid or tannin-dark lake and river systems. The lake sturgeon is found throughout the Great Lakes, Mississippi, and Hudson Bay watersheds, yet is rarely encountered by people simply because to save energy sturgeon cruise along the lake and river bottoms.

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Age Lake sturgeon are known to reach great size and age. Two “record” sturgeon have been reported from the Great Lakes, one from Lake Superior in 1922 weighing 310 pounds and over seven feet long (Scott and Crossman 1973) and one from Lake Michigan in 1943 of about the same size (Becker 1983). These large sturgeon are believed to be over 150 years old (Scott and Crossman 1973). A life history strategy often found in large-bodied organisms is the ability to live to great age. Sturgeon are similar to humans and some mammals in that they reach reproductive readiness over a time frame of many years. This can be a life history advantage over smaller-sized creatures for many reasons, and may explain why they have not changed much physically over the millennia. First, they can persist over time by remaining in deep cool water or large bodies of water, where their metabolism can slow and energy demands become low. This could occur during chaotic global events such as cooling, drought, flooding, and heating, which can result from climate change. If for some reason they cannot spawn because of conditions that are unfavorable for reaching the spawning site, they can live to return in future years to try again.

Overview Sturgeons are unique species of fish, and in North America the lake sturgeon is the largest, most widely distributed freshwater fish in inland freshwaters. This species is a historic fish; it represents one of the steps in evolution between fully cartilaginous fish (i.e., sharks and rays) and the more familiar ray-finned or “bony” fish (i.e., bass and salmon). Sturgeons have retained the form and sensory adaptations of the sharks yet show the beginnings of bony fish by the presence of protective scutes. These special fish can live to a great age, can grow to a great size, and have

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persisted for thousands of years. In fact sturgeons join the ranks of such aquatic creatures as horseshoe crabs and sea turtles, thriving pretty much unchanged in form and function since the days when dinosaurs ruled the world. Human actions now put sturgeons at risk of survival by a combination of overfishing, barricading of corridors needed to reach spawning sites, and destruction of important spawning, feeding, and resting habitats.

REFERENCES

Auer, N. A. 1999. Population characteristics and movements of lake sturgeon in the Sturgeon River and Lake Superior. Journal of Great Lakes Research 25(2): 282–293. —. 2004. Conservation. In: Sturgeons and paddlefish of North America. Greg T. O. LeBreton, F. William H. Beamish, and R. Scott McKinley, eds. Kluwer Academic Publishers. Becker, G. C. 1983. Fishes of Wisconsin. University of Wisconsin Press. Ferguson, M. M., and G. A. Duckworth. 1997. The status and distribution of lake sturgeon, Acipenser fulvescens, in the Canadian provinces of Manitoba, Ontario, and Quebec: A genetic perspective. Environmental Biology of Fishes 48:299–309. Harkness, W. J. K. 1923. The rate of growth and the food of the lake sturgeon (Acipenser fulvescens LeSueur). Univeristy of Toronto Biological Studies Series 24. Publication of the Ontario Fisheries Research Lab 18.

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Harkness, W. J. K., and J. R. Dymond. 1961. The lake sturgeon. Ontario Department of Lands and Forests. Hilton, E. J., and L. Grande. 2006. Review of the fossil record of sturgeons, family Acipenseridae (actinopterygii: acipenseriformes), from North America. Journal of Paleontology 80:672–683. LeBreton, G. T. O., F. W. H. Beamish, and R. S. McKinley, eds. 2004. Sturgeons and paddlefish of North America. Kluwer Academic Publishers. Nicol, J. A. C. 1969. The tapetum lucidum of the sturgeon. Contributions in Marine Science 14:5–18. Rafinesque, C. S. 1820. Icthyologia ohiensis. Author. Rodriguez, A., and E. Gisbert. 2001. Morphogenesis of the eye of Siberian sturgeon. Journal of Fish Biology 59:1427–1429. —. 2002. Eye development and role of vision during Siberian sturgeon early ontogeny. Journal of Applied Ichthyology 18:280–285. Scott, W. B., and E. J. Crossman, 1973. Freshwater fishes of Canada. Bulletin 84. Fisheries Research Board of Canada. Sillman, A. J., A. K. Beach, D. D. Dahlin, and E. R. Loew. 2005. Photoreceptors and visual pigments in the retina of the fully anadromous green sturgeon (Acipenser medirostrus) and the potamodromous pallid sturgeon (Scaphirhynchus albus). Journal of Comparative Physiology A: Neuroethology, Sensory, Neural, and Behavioral Physiology 191:799–811. U.S. Geological Survey, Great Lakes Science Center. Zebra mussel. 2008. Http://www.glsc. usgs.gov/main.php?content=research_invasive_zebramussel&title=Invasive%20

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Invertebrates0&menu=research_invasive_invertebrates. Watanabe, Y., Q. Wei, D. Yang, X. Chen, H. Du, J. Yang, K. Sato, Y. Naito, and N. Miyazaki. 2008. Swimming behavior in relation to buoyancy in an open swimbladder fish, the Chinese sturgeon. Journal of Zoology 275:381–390.

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Wilga, C. D., and G. V. Lauder. 1999. Locomotion in sturgeon: Function of the pectoral fins. Journal of Experimental Biology 202:2413–2432.

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JIMMIE MITCHELL

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N’me

I have been asked in a good way to share the influences and perspectives driving modern tribal resource management. In honoring this request, I am proud to share with you a project that is a premiere expression of modern-day tribal sovereignty, the n’me restoration project of the Little River Band of Ottawa Indians. Before I attempt to describe this project in greater detail, let me first explain the inspiration utilized in the creation of our sturgeon management plan, an Indigenous belief structure known as Baamaadziwin, which translates into “living in a good and respectful way.” The Anishinaabek (the name in our language we refer to ourselves as) have a belief system that has been in existence since time immemorial and has been passed on in the oral tradition from generation to generation to our present times. When we seek a teacher in Baamaadziwin, it is our understanding that we will receive more than basic instructions in a “good and just way to live.” As these teachings are learned, we are also introduced to the guiding spirits of Baamaadziwin, spiritual guides who will assist us on our new path till the end of our days on earth. This connection between spirit-world and our own is not obtainable by researching books or visiting the World Wide Web. The introduction is made in the presence n

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of teacher with student, connecting them to the unseen powers surrounding us all, a procession of knowledge and faith that has occurred since the beginning of our existence. I will explain later how this influence transcends the Anishinaabek understanding of the world, and how so much of what we understand has been learned from the wisdom of the wilderness. The spirit that is connected to our belief system guides the Anishinaabek to our respective responsibilities, beyond that of being “good and just” people—to being servants, devoting ourselves to making a difference in all that has occurred and may still be occurring within our respective communities and environment. This community-minded service for some of us also includes restoring the balance of our shared natural environment and of all inhabitants who are dependent upon a robust ecosystem. As is told in the creation story of the Anishinaabek, Giizhemanido (The Creator) put into existence all of the life-forms we have come to know. Some 360 million years ago, during the creation of the first n’me, Giizhemanido also decided to create the very first Anishinaabek from the remains of the first animals that had lived and died. In Giizhemanido’s infinite wisdom, the positive attributes that he admired most in those first animals thusly became inherent in the Anishinaabek; and therefore our dodem, or clan, system was created and set into motion. The phrase n’dodem in the language of the Anishinaabek literally translates into English as “I have the heart/spirit of . . .” A sturgeon clan person, for instance, would declare his or her clan in this way: n’me n’dodem, or “I have the heart/spirit of a sturgeon.” This introduction is very significant in the cultural interactions of the Anishinaabek because as we venture along this journey, living connected to the harmonious ways of Baamaadziwin, whenever we connect with other Anishinaabek, it is respectful to explain who we are, where we come from, and, if known, which clan we represent. Following the cultural practices of the Anishinaabek, clan responsibilities are expressed in fundamental rules to abide by. For instance, clan people are instructed not to marry into their own clan. This was done as a means of maintaining genetic diversity throughout the various Anishinaabek communities. Clan people are also charged with various tasks and responsibilities within the communities; some possess the role of teachers, others protectors, hunters, and healers, and through the clan system a well-balanced community was ensured. Upon the arrival of the dominant society, its effects began to negatively tilt the delicate balance of our world, including that of the clan system. In the throes of genocide, clan leaders were targeted as a threat opposing the emerging strides across the landscape driving a new concept known as Manifest Destiny.

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In the wake of the losses occurring to the world we knew, loved, and depended upon, tribal leaders opposing these changes were dealt with severely, and most often with extreme prejudice. As the balance of the environment teetered out of sync, new concepts like “resource management” and “public trust doctrine” were coined to leverage the needs of the dominant society against those of the Anishinaabek, causing negative environmental consequences that reverberate into our current landscape. Likewise, with the loss of leadership, land, and naturally occurring sustenance, the once vibrant cultural kinship between earth, animal, and the Anishinaabek wilted. To explain this detriment in greater detail: our healing ways suffered, as our doctors were no longer assisted by their clan animal counterparts who used to confer their healing powers on them, a significant component to our faith. The naturally occurring medicines we had relied upon to heal illnesses since the beginning of time no longer grew on the degraded soil, parched by the absence of the forest canopy, torn off by clear-cutting and subsequent burning. New diseases inflicted upon the Anishinaabek killed scores of our people, forcing us to turn toward the medicine of the dominant society. The loss to the indigenous animal and human populations within the Great Lakes Basin since first contact is profound. The dimension of suffering we have known and continue to know is reflected in the degraded condition of our environment, yet we still consider her sacred, so much so that we refer to her as Wegemind Aki, or Mother Earth. As our tribal nations begin to grow strong again, we haven’t forgotten our primary role: To mend the circle of life that has fallen so far out of balance, we must rely upon our codependency with our clan species in conjunction with the roles that were bestowed upon us by Giizhemanido. During the treaty negotiations of 1836, our chiefs consciously argued for language that secured the usual privileges of occupancy the Anishinaabek relied upon, then as now. The treaty was negotiated to pave the way for our lands to become a state; the negotiations were brought forward to divert a war about to break out between the United States and the Odawa, Ojibwe, and Bodwewadomi nations, the Three Fires Confederacy. In their wisdom, our chiefs knew that if they could continue to occupy our homelands and follow the natural order that Giizhemanido had granted us in this paradise that we called home, we could continue to exercise our unique identity as Anishinaabek. Then, shortly after the treaties were signed, the unthinkable occurred: the army took our children away from us by force and sent them to Indian boarding schools across the country, where they were taught in the cruelest of ways to be ashamed of themselves and our society as backward and lacking intellectual capacity. We had

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been stripped of our connection to the lands, animals, and medicines. For the next 160 years, the Anishinaabek languished as our families clung to the slim faith that one day we would return to our rightful place as a proud and unique society. Despite the intentional disenfranchisement from everything the Anishinaabek knew, loved, and held sacred, we endured. Remaining from the tenets of a nation that once stood as the proud original caretakers of the lands, the beliefs we clung onto eventually fanned the embers of our once great nation into flame, a flame that has illuminated our path, calling us home to begin anew in our rightful place in Creation. In 2003, the treaty of 1836 was called into question in the Sixth District Federal Court, targeting specifically the language that preserved our usual privileges of occupancy until the land was required for settlement. The usual privileges of occupancy during the 1836 treaty included hunting, fishing, and gathering practices, but certainly more with the economic change taking place within the region. The Tribe had already before 2003 been relying on the language preserved by our chiefs in 1836 to hunt, fish and gather, but also in our efforts to restore sturgeon populations in the Big Manistee River and to initiate economic development to fund the endeavors of the Nation. After an exhausting and costly negotiation process that lasted nearly four years, in the fall of 2007, the Tribes were able to reaffirm the original language in the1836 treaty, securing the rights not only to hunt, fish, and gather, but also to conduct co-management initiatives, including the restoration, reclamation, and enhancement of species and habitat that are of cultural significance to the Tribe. These inherent rights serve not only the Tribe; their benefits are to be shared with people from all races of life. In keeping with our Seventh Generation teaching, we have to ensure that the next seven generations of the people yet to be born inherit the same benefits that we enjoy and utilize today. Despite the advancements in resource management, the continued progress of humans continues to create a detriment to all forms of naturally occurring life. For example, it is difficult for me, as an Indian person, to drive my vehicle and see the massive number of animals killed along the road, animals that do not possess the wherewithal to comprehend the physics involved in cars speeding toward them. In the teachings that I have received, I’ve been taught to ask for forgiveness for doing something wrong but also to take steps to make amends. The Tribe, through its sturgeon-rearing program, is striving to make amends for 170 years of destruction done against this noble species. To qualify this detriment in regards to sturgeon populations, we have to travel back to the time when the last of the great forests were cut in Michigan. After the last of the trees were felled, the steamships had no fuel left to burn in order to propel these

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vessels across the lakes. During the sturgeon runs of the time, nets were stretched across the rivers. The caught sturgeon were dried whole and stacked like cordwood and subsequently burned as a new source of fuel, at least until the population was decimated, in the same fashion as the great forest. A current harm to the remaining sturgeon population comes with the massive number of lake trout and salmon fry that are stocked in the rivers in the spring of each year, about the same time the sturgeon larvae begin their drift. At this time, tribal biologists enter the dark waters of the river around midnight and painstakingly collect the elusive larvae from the frigid waters of the Big Manistee. The larvae are then carefully transported to the Tribe’s patented streamside rearing facility, where they are fed, monitored, treated for disease, and raised until their protective plates are formed. The process takes approximately four months, when tribal biologists feel the sturgeon survival rate is ensured. The Tribe’s annual sturgeon release ceremony takes place each September along the shores of the same river the larvae were retrieved from, the source that we feel imprints them with the water of the Manistee. Miigwetch (thank you) for listening to our story of n’me, and our humble attempts to preserve this amazing being that is loved and revered by so many of you.

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R EG IER , HU G HES, A ND GA N N ON

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Lake Sturgeon in the Laurentian Basin

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HENRY A. REGIER, ROBERT M. HUGHES, AND JOHN E. GANNON

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The Lake Sturgeon as Survivor and Integrative Indicator of Changes in Stressed Aquatic Systems in the Laurentian Basin

Our Laurentian Basin is ancient geologically, and the lake sturgeon lineage in our basin is ancient biologically. During the last five centuries, much of the southerly half of this basin’s waters has been transformed—mostly by humans of European origin—from a vast, clean, cascading “riverine system” to clotted strings of confined and dirty reservoirs and lakes with deformed tributaries and “connecting channels,” that is, into what we may term a “reservoirine system” (see box 1). The living part of the ecosystem in most of the southerly waters has changed from intricately organized and elaborately choreographed systems of native, freshwater, mostly riverine taxa to a highly altered and relatively disordered state with unwanted alien taxa from shallow seas and brackish bays far away. But that longlasting trend may not be our destiny. As complex self-organizing systems, these waters and their sturgeon populations retain resilient propensities in spite of their old age and man-caused disabilities. There have been three kinds of early waves of entry of unwelcome alien creatures into our basin:

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Box 1. Reservoirine Alterations

In 1971 some of us of an older generation convened a Symposium on Salmonid Communities in Oligotrophic Lakes (Loftus and Regier 1972). We compared and contrasted the case histories of numerous lakes that were once dominated by salmonine associations to see whether we could infer some major effects of three cultural stresses: poor fishing practices, environmental pollution, and introduction of undesirable alien species. The lakes that we selected fell into three sets; lakes at the margin of the Laurentian Shield in North America including the Great Lakes; deep Fennoscandian lakes; and lakes of the European Alps. During our comparative consideration within and between these three sets of lakes we noted that researchers on the Alpine lakes added an additional important stress related to hydrographic and hydrological modifications that they called Verbauung in German. We found that this term, which often had a pejorative connotation, meant something like “human alterations that obstructed or debased natural physical processes purportedly for the immediate benefit of the human obstructors/debasers.” What we refer to here as “reservoirine alterations” as a set has features like those that follow such Verbauung. A key feature of “reservoirine alterations” is our culture’s primary focus on the physical mass of water in aquatic ecosystems. (The aquatic ecosystem reaches into soils and aquifers and perhaps into local weather patterns, etc.) Nested within this primary focus on the mass of water are secondary physical foci: water level and its fluctuations; water flow and its fluctuations; copious fluid in a depression in a landscape for dumping wastes some of which were carried downstream and out of sight by currents; concentration of pollutants such as acids that corrode engineering works; temperature of the water; and so on. In recent decades Sproule-Jones (e.g., 2002) has described the role of historic preemption by use and by law of features of our natural ecosystems. Such preemptive practices that permeate current realities in our Laurentian Basin can be traced back to ancient history in Europe and Asia. With each major

• The Europeans themselves with their effective invasion strategy that included explorers, soldiers, missionaries complemented by traders, and settlers strongly influenced by an exploitative-consumptive culture • The human diseases that Europeans unintentionally brought with them • The alien terrestrial and aquatic species, including domesticated livestock and

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technological initiative like this, prior residents of the affected lands, including Aboriginal peoples, likely were harmed deeply in ways for which they received poor compensation at best. Preemption of water as physical mass relates to old-fashioned “progressive” industrial interests with respect to our aquatic ecosystem. Here “industrial” includes water for agriculture, including irrigation and drainage; cleansing and cooling things; electrical services including potential energy and cooling liquid; and floating and passing commercial vessels. At a macro level, the U.S. Army Corps of Engineers, the U.S. Bureau of Reclamation, and Public Works Canada serve such interests and the preoccupations of these federal institutions dominate and suppress the interests that are not primarily and secondarily as sketched above. The “obstructive” practical engineering methods that follow from such a set of primary and secondary foci have led to a transformation of a pristine riverine ecosystem to a debased reservoirine ecosystem that has adapted to old and crude industrial practices. The leading initiatives of the older basinwide governance institutions (the binational International Joint Commission and Great Lakes Fishery Commission, the Great Lakes Commission with its connections to Canadian provinces) all explicitly or implicitly accept the preeminence of what we have termed primary and secondary focal interests above. So “reservoirization” is a meta-adaptation of our basin’s aquatic ecosystem to old-fashioned obstructive industrialism. The reservoirization syndrome includes damping floods and droughts of natural flow regimes that reset the dynamics of water bodies but are troublesome to human enterprises; destroying wetlands, riparian zones, complex channels, large woody debris, floodplains, and temporary streams and ponds—all key to natural ecosystem processes; disconnecting surface and ground water, leading to salt deposition in xeric areas and flash flooding in others; and severing critical migratory routes for aquatic species such as fish and mussels.

plants, which were deliberately or accidentally introduced as a result of various kinds of stocking and barrier removal Humans, mostly of European descent, used and abused our basin’s waters in countless ways. The historical sequence of initiation of various human activities with

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major adverse ecological effects took place approximately as follows: the trapping of beaver and removal of the impermanent dams, ponds and the wetlands that they created; physical restructuring of streams and shorelines through snagging, morepermanent damming, channeling, and hardening of banks; fishing exploitatively; loading of organic wastes and sewage; introduction of alien fish deliberately or accidentally; loading of plant nutrients; loading of toxic chemicals, then radionuclide physicals, then trace organo-contaminants and pharmaceuticals; introduction of a broad spectrum of alien species mostly via ships’ ballast waters; acid rain; and change in the climate. Each of these classes of stress, as well as any transformational synergism among stresses, has likely affected adversely the numerous lake sturgeon populations in these waters. Nevertheless some sturgeon populations did survive, in small numbers, and are now showing signs of recovering resiliently. We, the three authors, are of European descent ourselves. As “Europeans” we wish to lay no claim to being the worst ever of the world’s conquerors throughout history, noting the work by Diamond (1997) on this subject. But here in the Laurentian Basin, half a millennium ago, it was Europeans who entered as external invaders and eventually affected ecological and other aspects of the basin’s history deeply, if often in ways that would have been unintended had they given the matter some ethically informed thought. With our broad-brush approach in this heuristic chapter, we will use the term “European” without making an effort to strike a fair balance, from some distant and disinterested perspective, between goods and bads that Europeans brought with them. All of the phenomena mentioned above—except sea lamprey as an alien invader, acid rain, and climate change—are much more apparent in the southern half of our basin than in the upstream northern half (Danz et al. 2007). Aboriginal peoples, who have lived here since time immemorial but with greatly diminished numbers during recent centuries, tend sacred flames in some locales that are not as severely abused as is the rest of our basin. Through the previous half millennium of gross change in our basin, some ancient species like the lake sturgeon and common loon have survived, in decreased numbers. Five centuries from now, will these species still be here? Overall, the cumulative effects of degrading cultural stresses (see box 2) in this vast ecosystem may have peaked or plateaued about 1950, at least in southerly waters. Gradually the overall intensities of some abuses have been reduced, but legacy effects from the settlement era (Trautman 1977) remain 200 years later. Remediation of some old stresses may have offset additional new stresses since 1950. Perhaps the present century will bring with it major net recovery and rehabilitation, but it seems highly unlikely to us that Lake Erie, for example, will again come to resemble closely its ecosystemic state in the year 1500.

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With precautionary commitment to sustainability, the inevitable new stresses may not cripple desirable ecosystem features to the extent that occurred with classes of old stresses over previous centuries. But sooner or later, a very human technician may do something foolish that will cause a nuclear electricity-generating plant to malfunction and blast radionuclides far and wide. Another 500 years of economic and population growth do not bode well for some of the remaining ancient features of these ecosystems, given past trends (Naidoo and Adamowicz 2001; Clausen and York 2008), especially with respect to freshwater fish—and including lake sturgeon (Miller Reed and Czech 2005; Rose 2005; Leprieur et al. 2008). Those past trends in economic and population growth have occurred even longer along the Atlantic coast of North America and in Europe. And the European sturgeon Acipenser sturio has been extirpated from the Seine Basin (Oberdorff and Hughes 1992) and most of Europe, whereas Atlantic sturgeon A. oxyrinchus, shortnose sturgeon A. brevirostrum, and Alabama sturgeon Scaphirynchus suttkusi have been listed as endangered in the United States (Jelks et al. 2008). As a result of overexploitation, habitat fragmentation, and water pollution, the International Union for Conservation of Nature considers sturgeon the most threatened group of animals, with 85 percent of the species at risk of extinction (http://www.iucn.org/?4928). Our approach here may be termed “heuristic” and consistent, we suggest, with the “Prescription for Great Lakes Ecosystem Protection and Restoration” (Bails et al. 2005). We emphasize general concepts and inferences and do not mobilize in a quantitative and critically convincing way much of the empirical information in hundreds of references as might be expected in a state-of-the-art meta-analysis. Other chapters in this volume contain much relevant empirical information. We suggest that all the concepts and data combined critically may demonstrate that an appropriate measure of the ecological health of rehabilitating populations of the lake sturgeon species can serve as an integrative indicator of the state of ecological health of the aquatic part of our basin ecosystem. Who better to act as guardians of recovering sturgeon populations than their friends since time immemorial, the Aboriginal peoples?

Ancient Eco-History Ancestral fish that resemble our present lake sturgeon have thrived in our Laurentian Basin and nearby watersheds for millions of years. Aboriginal humans, treading lightly in the lands and floating smoothly on the waters, “settled” the basin ecosystem many millennia ago, to be followed some centuries ago by many Europeans, with heavy tread and erosive navigation, followed in turn by many less-abusive humans of

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Box 2. Terms for Cause-Effect Sequences in Complex Ecosystems

As we learn more about life in ecosystems, including microbial life involving vast numbers of microbes not yet identified specifically except through a few genes in their genomes, we now realize that full scientific understanding of any dynamic ecosystem is far beyond human capabilities. So the particular effects on an ecosystem of particular actions by humans cannot be fully inferred. The terms “cause” and “effect” may seem unambiguous, but, applied to the consequences of human actions on ecosystems, they cannot be made fully operational. Numerous sets of terms have been used by ecosystem researchers in reference to cause-and-effect sequences, for example, action and reaction; stimulus and response; stress and strain (in physical sciences); stress and stressed; stressor and stress (in biological and psychological sciences); pressure, state, and response; and drivers, pressures, state, impact, and response. Note that in the physical sciences “stress” usually refers to a cause. In biological and psychological sciences influenced by the 1960s works of Hans Selye, “stress” refers to an effect of some relevant cause or “stressor.” Years after Selye introduced his terminology he came to realize that it was logically inconsistent with conventional uses in the physical sciences, but by then Selye decided not to change his terminology to align it with that of the physical sciences. In our title for this chapter we have used the term “stressed” for an effect of a causal “stress,” that is, “stressed” implies a “strain.” We continue with this convention throughout the chapter. In diagnostic and forensic sciences a researcher may try to deduce or induce causes from observed effects. With ecosystems and other complex phenomena,

African and Asian origins only some decades ago. Each of these waves of immigrants came to interact with the Basin ecosystem in somewhat different ways. The most pervasive, prolonged and violent interactions between humans and other parts of these ecosystems came and stayed with the European invaders acting in what they thought were enlightened ways. The story of how the complex of lake sturgeon populations has fared through history in our basin’s waters is introduced in this chapter, starting billions of years ago. Here the waters of the “Laurentian Basin” are defined as those that now drain naturally into the string of large “lakes” (Superior, Huron-Michigan, Erie, Ontario) and large “rivers” (St. Mary’s, St. Clair, Detroit, Niagara, St. Lawrence, Ottawa, Champlain) that eventually flow into the sea northeast of Quebec City, with tidewater

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inferring reliably about causality is difficult for many reasons. For example, the dynamics linking causes and effects may not be quantitatively linear but instead may involve nonlinear state shifts that resemble qualitative rather than quantitative consequences. Also, an ecosystem may respond to a variety of stimuli in ways that seem quite similar to an inexpert observer. Thus “eutrophication” is commonly used as a catchall term for a set of somewhat similar consequences, that is, a general adaptive syndrome, to a variety of quite different human actions that affect an aquatic ecosystem. Teasing out the particular cause(s) of an “eutrophication event,” that is, from a general adaptive syndrome, with diagnostic or forensic scientific methods is difficult (Rapport et al. 1985). That said, the art and science of ecological assessments have improved greatly in recent years with increased availability of large, consistently collected databases and sampling from hundreds of sites at continental or subcontinental scales (e.g., Pont et al. 2006; Brazner et al. 2007; Paulsen et al. 2008). In those studies, spatially extensive stresses such as row crop agriculture, urban development, hydrological alterations, and mining were determined to limit biological indicators from algae to fish. Paulsen et al. (2008), using a relative risk assessment approach developed by Van Sickle and Paulsen (2008), reported that for the coterminous United States excess total phosphorus, excess total nitrogen, and excess fine streambed sediments most often limited macroinvertebrate assemblages in wadeable streams—at least among the limited number of stresses quantitatively examined. These same stresses likely limit biota in lakes as well.

as an approximate downstream boundary. Important structural features of these waters’ geological basin reflect ancient faulting in the underlying core or craton of our continent, which was first formed four or more billion years ago. Thus the wide “valley” that extends in an approximately straight line from Chicago northeastward past Quebec City and beyond parallels an ancient fault, to which other faults at right angles to it are joined in various locations. The faults, as deep geological features of our part of the craton, are still active, as demonstrated by occasional earthquakes of low intensity (Dineva, Eaton, and Mereu 2004). The faults have been complemented at the surface by effects of numerous continental glaciers that arose periodically to the north of our basin and gradually flowed southward through it, approximately at a right angle to the natural

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flow of these waters when the glaciers were not present. Lakes were carved out and deepened; and the land’s surface was scoured away. The scraped-off clay, sand, gravel, and rocks were deposited in massive humpy moraines at the southernmost edge of a particular glacier’s motion. Many earlier moraines were overrun and deformed by later glaciers that proceeded further southward. Glacial moraine complexes are of great importance to the aquatic ecosystems of the Laurentian Basin. Because of their porous structure, the moraines act like sponges that are filled during heavy rains of spring and fall and then drain continuously during the drier winter and summer months. Mining the moraines for sand and gravel diminishes their capacities to store and then deliver water into headwater streams, so flows of many streams during dry periods must now be less than was the case five centuries ago (e.g., Carlisle, Wolock, and Meador 2011). Perhaps some streams that once served as spawning habitat for sturgeon no longer do so now for this reason alone. Springs that originate in moraine aquifers have relatively constant temperatures over a year that are about 1 to 2°C above average annual air temperature at the moraine site; snow cover acts as an insulating blanket in winter to keep the annual mean groundwater temperature a bit above the annual mean air temperature. So moraine springs average about 1°C at the northern edge near Lake Nipigon to about 12°C at the southern edge south of Cleveland (Schlesinger and Regier 1982). The drainage of the insulated water from moraines keeps the streams relatively warm and flowing in winter and relatively cool and flowing in summer, to the advantage particularly of certain fish species and other aquatic taxa. From a limnological perspective, the waters from the southerly moraines echo some of the seasonal features of the waters from the northerly part of our basin, with its porous ancient rock. Some species, like brook trout, appear in streams farther south latitudinally than would be the case in the absence of such moraines to provide habitat of suitable temperature year round (see, e.g., Moerke and Lamberti 2006). During the past million years, these waters with their biota and especially the fish of the Laurentian Basin’s aquatic ecosystem have been pushed south repeatedly by advancing continental glaciers into the headwaters of more southerly basins and have shifted north again as the glaciers melted thousands of years later. The biota of this relatively open aquatic ecosystem, as a whole or in several parts, likely always maintained some self-organizational ecosystemic integrity as it/they advanced southward in front of the ice and returned northward as the ice melted. Such integrity likely included complex structures such as “holonic” nesting of smaller ecosystems within larger ecosystems through a number of self-organized levels. The term “holonic” refers to a more generalized notion of vertical and horizontal nesting with reciprocal interactions than is usually implied by the etymologically misleading

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term “hierarchic” or “priestly order” (Regier 2008). At any particular time, spatial boundaries, usually indistinct, of ecosystems at various levels of nesting were likely related more clearly to hydrostatic gradients and hydrorheic features than to solid hydrographic features. If photographs, say from an observation platform in space, could have been taken annually at midsummer over the past million years, the photos would likely show this sequence:

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• At times between ice ages as now, an aquatic riverine treelike network with numerous vertical and lateral bulges (i.e., “lakes”) of various sizes and with combined outflow northeastward to the Atlantic • Early in an ice age, a distorted aquatic system with its outflow to the northeast dammed by ice twisting and shifting southward with advancing glaciation that broke into a number of subbasins with outflows southward during that ice age • Late in an ice age, the hydrologically adapting aquatic system shifting northward and eventually settling into a configuration somewhat similar to that of the previous ice-free period when the ice dam in the northeast had melted The natural aquatic ecosystem of five centuries ago, before the invasion by Europeans, had been affected by numerous north-south oscillations, with its separation into different southerly watersheds or catchments followed by a reverse shift into one watershed, as at present. From such a geological perspective, one would expect that the biota of this aquatic ecosystem would have a strongly riverine character. The native fish of our basin demonstrate evolution and adaptation to a metariverine habitat. One of the key considerations is that few of our basin’s native fish species were thoroughly adapted to an offshore, pelagic lifestyle that included midwater spawning. Almost all spawned in rivers and tributaries or on reefs in the “lakes” across which currents flowed. The lake sturgeon was a part of such riverine wanderings for countless millennia. Note that the Great Lakes are often said to be “young” geologically. The present/ recent hydrological/hydrographical manifestation is less than 10,000 years old. But many biotic parts of these ecosystems—the biota self-integrated into flexible, adaptive, riverine ecosystems—are much older than that. This becomes apparent when considering the complex stock structures of the salmonid and percid families of fish, say. So ecologically the biotic part of our basin ecosystem was not “young” five centuries ago; it was predominantly old-growth riverine, which has been mistaken by some for youth. A complementary error has been common: the “artificial eutrophication” that followed massive loadings of plant nutrients a century ago was said to have triggered

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“premature aging” of the relevant aquatic ecosystem when the effect could perhaps more realistically be caricatured as induced obesity through forced feeding of the pelagic part of the ecosystem, or “pelagification” (Regier and Kay 1996; Kay and Regier 1999; Dobson, Regier, and Taylor 2002). In aquatic ecosystems like those of our basin, the normal self-organizational “succession” over centuries leads to dominance by the benthic association of species, in a process that may be termed “benthification.” Our Basin’s fish thrive in heterothermic environments and cannot control their body temperatures (cold-blooded) except by purposely selecting habitat of preferred temperature, if available. For such fish, the temperature of various parts of their aquatic habitats is a key determinant for the spatiotemporal organization of various features of life histories. Many of the basin’s fish species have rather narrow, and different, ranges of temperature that are optimal for spawning or for growth, say. Not far beyond an optimal temperature for growth lies a lethal upper temperature, which may be less that 8°C above the growth optimum (Regier et al. 1996). During ice ages, the presence of ice in some waters during summer likely permitted the salmonid family to thrive there. Members of this family now thrive in or near those parts of the basin’s waters that are coldest in summer, such as moraine streams to the south, deep lake waters saturated with dissolved oxygen throughout the basin, and unpolluted surface waters to the north. This family includes salmonines like char, trout, and salmon, thymallines like the grayling, and coregonines like the whitefish, cisco, lake herring, and chubs. Farther downstream from glaciers and moraines and in surface waters separated from hydrologically isolated deep cold waters, the water warms in summer to a temperature suboptimal for salmonids. Cool-water percids thrive in such waters, as well as other families, including in particular the acipenserid, the lake sturgeon. Farther downstream from the sources of cold moraine water in summer, and in shallow waters protected from coastal upwelling of cold bottom water and from long-shore currents of cool water, the warm-water family of centrarchid sunfish and black bass thrive together with other families. To emphasize, we expect that a spectrum of fish families—with cold, cool, and warm habitat preferences in summer—has existed in the basin ecosystem during ice-free ages and in the parts of the displaced ecosystem during ice ages for millions of years. During the past two centuries, the amount of habitat for the cold-adapted fish has shrunk through warming of many streams and reductions of oxygen concentrations in some deep waters. For example, the grayling was eliminated from Michigan following land clearing in the early 1900s (Richards 1976). Similarly, the warm habitat has shrunk in many southerly locales because of destruction of the shallow land-water ecotone. Cool-water habitat may not have been diminished as much as the other two types. Lake sturgeon are at home in the cool waters, in company with

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the walleye and yellow perch. These two percid species can cope behaviorally both in benthic-dominated and in pelagic-dominated ecosystems, but the sturgeon cannot thrive in midwaters because of its heavy bony features. According to the “Ontario Freshwater Fishes Life History Data Base” (http:// ecometric.ca/fishdb/main.htm), based to a major extent on the work of Scott and Crossman (1973), the preferred habitat of lake sturgeon is the bottom water of lakes and large rivers 5 to 10 meters deep, where it feeds on benthic invertebrates in clay, mud, sand, and gravel bottom. It spawns in rivers in spring with water temperatures between 9 and 18°C; its estimated temperature preference in summer is 15 to 17°C, though field data suggest that the sturgeon may actually prefer a range of temperatures a couple of degrees higher than that. With that range of temperature preferences, during summer the lake sturgeon leaves warm stream, river, and epilimnetic “lake” waters, at least in the more southerly parts of the basin, and moves downslope within lakes to cooler waters but not into the cold hypolimnetic waters. Retrospectively, this species would likely have found appropriate habitat during any stage of glaciation and deglaciation of our basin in the past million years. Numerical measures of many indicators of the state of health of our basin ecosystem are used now and more are being proposed (e.g., Niemi and Kelly 2007). Various fish species can be used as integrative indicators of ecosystem health with respect to those parts of the ecosystem that they inhabit. The lake trout as a representative salmonid has been used in this way for decades in our basin with respect to large cold waters (Regier 1992); to a lesser extent the walleye is used with respect to cool waters and smallmouth bass with warm waters. It may be timely to document the case for using lake sturgeon as another integrative indicator of ecosystem health for cool waters in summer; much of the relevant information is contained in other chapters of this volume. The implicit rationale that was used by a number of agencies to select the lake trout as an integrative indicator of ecosystem health (Regier 1992) can be used with lake sturgeon.

Riverine Features of the Near-Pristine Waters in the Year 1500 and Subsequent Changes Rivers normally wander across a relatively flat landscape over the ages, even in the absence of glaciers. Even during glacial periods, the water in the form of deep ice extending a thousand meters skyward flows, if only slowly, but it does flow, as do streams in and under the ice. In the preceding section, the whole water mass of our basin is described as having river-like features, but with massive vertical and horizontal bulges in places we call lakes or glaciers. Internal to these bulges,

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whether downward as in lakes or upward as in glaciers, there are currents with some river-like features. In our basin, some nonecological experts of decades ago routinely referred to large rivers like the St. Clair and Detroit as “connecting channels” between lakes. Implicitly this terminology depreciates the importance of these rivers with respect to the lakes that they “connect.” To us it makes more sense to emphasize the riverine features and flows of the lakes and treat the lakes as “connecting bulges” or large pools (with some riverine features) between the natural rivers. (This is also similar to the way some limnologists view the Amazon Basin and its lakes, slowly flowing swamp streams, and flooded forests.) The tributaries from headwaters to one of the large lakes are obviously riverine. Especially in its northerly half, our basin has thousands of small lakes that rest on top of our billions-of-years-old craton, the Laurentian Shield. It also has numerous medium-sized lakes such as Fox, Nipigon, St. Clair, Nipissing, Simcoe, Kawarthas, Finger Lakes, Oneida, Opeongo, Champlain, and so on. There are four large lakes: Superior, Huron-Michigan, Erie, and Ontario. Five centuries ago, all of these lakes, whether of small, medium or large size, manifested notable riverine features, some of which have now been debased into “reservoirine” features (see box 1). An important feature of natural waters in our part of the continent is that they are bordered by a land-water ecotone or riparian zone profusely vegetated in locales other than wave-swept, rocky shorelines (Minshall et al. 1985; Gregory et al. 1991; Sedell et al. 1991). Five centuries ago, woody plants were found bordering shores often in dense thickets; woody debris of all sizes was found in headwater streams and down the tributaries along the shores and on the bottoms of rivers and lakes of increasing size all the way downstream to the St. Lawrence River. From what we now know about woody debris in relatively natural waters such as the Amazon Basin of South America and relatively pristine watersheds elsewhere in North America (Gregory, Boyer, and Gurnell 2003; Sedell and Froggatt 1984; Triska 1984; Hughes, Rinne, and Calamusso 2005a; Benke and Cushing 2005), we infer that woody debris had major ecosystemic importance in the pristine waters of our basin. In the downstream half of our basin, much “wood” was already cut and cleared out of the terrestrial and aquatic parts two centuries ago. But some of the cut wood was then dumped into the waters as bark and sawdust that took decades to decompose anoxically; old sawdust and bark still remains in many inland lakes near old lumber mill sites (R. M. Hughes, personal communication). Some of the secondgrowth deciduous trees that followed thorough removal of the white pine forests then toppled into naturally eroding rivers to good ecological effect, but in recent decades these have been removed from popular canoeing rivers to facilitate passage. Natural

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freshwater ecosystems in forested regions like our basin need constant supplies of foliage from living woody plants and of dead wood (though not sawdust) to behave naturally, so our basin’s aquatic ecosystems have become progressively pauperized for this reason, as well as others. In their natural state, beaver in countless locales dammed smaller rivers. We infer that the beaver did not “engineer” their dams to withstand an intensity of natural flooding that occurred less frequently than about once in a decade or two, maybe. So these dams were generally topped by high flows and freshets and were breached repeatedly. Beaver preferred early-successional trees like aspen, which thrive on old beaver meadows, so their engineering design may have incorporated “planned obsolescence.” In any case, beaver apparently did not permanently separate parts of streams behind permanent dams. Starting a century before the European agricultural settlers came, the demand for beaver pelts by the fur traders led to the great reduction in beaver and consequently of intact beaver dams. In thousands of streams, of all sizes from headwater brooks to the St. Lawrence River, artificial dams (of earthen, timber, rock, concrete, and steel materials) were built for many purposes by the European invaders. Many of the first-generation dams, built with less expertise than were the beaver dams, were quickly washed out by spates that had been made more “flashy” and violent by clearing the woody plants and debris upstream, but a fifth-generation dam some decades later may have then persisted for many decades. A dam that was not breached or overtopped by annual floods became a permanent barrier to migrant fish spawners that may habitually have homed to spawning areas upstream of such a dam. In aggregate, this happened with respect to thousands of gorges, rapids, and reaches in streams and rivers. Smith (1995) summarized some numerical data on the historical changes that occurred as follows: “the forests of the Lake Ontario drainage were described as ‘unbroken’ in 1784. . . . The process of forest cutting progressed from east to west; by the 1890s, the drainage area of Lake Ontario in western New York was described . . . as ‘almost entirely deforested.’ Concurrent with the removal of forests was the drainage of swamps to create farmland and the construction of dams for water-powered mills. The construction of mills to process forest and farm products started in the late 1700s. By 1845, there were 7,406 sawmills run by water power in the state of New York . . . and a somewhat lesser number of grist mills, plaster mills, tanneries, and other water-powered industries on both the United States and Canadian sides of Lake Ontario.” (Note that the catchments for all these streams are not stated explicitly to have been parts of the Lake Ontario catchment.) Large permanent dams also led to nonnatural, artificial flow regimes. Some changes fostered emergence of “dead zones” of anoxic mud and stagnant bottom water upstream of dams and in offshore deep waters in lakes.

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Cropping away of smaller streams as such throughout larger catchment areas in the southern half of our Laurentian Basin has occurred progressively for centuries. Tile drainage and adaptation of storm sewers to conduct streams underground are common in urban and agricultural areas (Smith 1971). Examination of a historic series of detailed topographical maps shows that many streams of low order—first, second, even third—do not appear as surface streams in later maps (Steedman 1986). For more recent examples of this phenomenon in the conterminous United States, see Stoddard et al. (2005) and Carlisle, Wolock, and Meador (2011). Gottgens and Evans (2007, 87) state that 632 dams are or were located on Ohio tributaries to Lake Erie. Of these, “29% are considered to be ‘highly hazardous’ and an additional 30% are considered to be ‘significantly hazardous.’ . . . Over time, hydraulic structures such as dams and reservoirs gradually change from assets to liabilities that cannot be ignored.” Presumably the assets were perceived as such from a conventional economic perspective and not necessarily from an ecological perspective. Large swampy areas were drained over a period of two centuries, such as the vast wet forest that extended from around the southwestern end of Lake Erie, northward around Lake St. Clair, and further northward to Saginaw Bay. Long stretches of the shores of larger rivers and lakes were diked. Some stretches of shorelines, especially those adjacent to dredged and deepened mouths of these rivers where they joined the large lakes and rivers, were sculpted and then hardened with concrete and steel to create harbors. The draining, diking, dredging, and hardening separately and together lopped off large ecotone areas of the natural riverine ecosystem, especially highly productive lake estuaries and floodplains. A different sort of alteration has occurred where Lake Huron drains into the St. Clair River. In 1859, a 4–5 m deep rapids existed there. To facilitate commercial navigation, channels were cut and blasted through the bedrock in the 1920s (7 m), 1930s (8 m), and 1960s (9 m). The river is now continuously eroding the softer substrate and currently ranges from 10–21 m deep, resulting in a lowering of Lake Huron and Lake Michigan by 40 cm to date. This amounts to a vast volume of water when multiplied by the lakes’ areas, and results in receding shorelines. Other changes to facilitate shipping (Erie Canal, Welland Canal, Soo [Sault] Locks, St. Lawrence Seaway) have opened up the Great Lakes to innumerable invasive alien species brought here in ships from throughout the world. Another ecologically important riverine feature of our large “lakes” is that currents of water flowing through a lake or circulating in gyres within a lake scrub up against the shore and bottom (Rao and Schwab 2007). The Coriolis force, as related to the forward momentum of water that is flowing downslope because of gravity but deflected to the right in our Northern Hemisphere because of the rotation of the earth, is one reason for the curving currents in our rivers and riverine lakes. Winds,

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which are markedly changeable in direction in our basin, alter direction of flows and contribute to the strongly episodic features of the currents. Another reason relates to rocky barriers that deflect and channel forward motion of water. A shore where currents often flow or winds persistently blow across a long fetch, though not necessarily constantly, has riverine features with respect to bottom materials, rooted aquatics, woody debris, the benthos, and the fish assemblage. It is like a one-sided, episodic river that may be spatially continuous for a long stretch of coastline when a large gyre is rotating or in inshore waters in spring when a “thermal bar” is present (Rao and Schwab 2007). The water of medium and large lakes shows one or more gyres, and sometimes gyres within gyres, of varying shape and velocity depending on seasonal meteorological and hydrological conditions. These have some features of slowly rotating whirlpools in rivers. Many kinds of “developments” that European invaders created in our basin have caused changes in the current regimes in rivers and lakes, especially in the southerly downstream half of the basin. Perhaps one of the “strains” in the rheic system caused by these stressful developments was that the current patterns have become much coarser and more grossly episodic (see box 2). If so, such a change must have been disadvantageous to many species, including lake sturgeon. Lake sturgeon prefer bottom habitats with currents; in such places the bottom may be of stone cobble and hard clay. For example, the underwater delta and distributary currents of the relatively warm Niagara River water into the colder Lake Ontario waters were a favorite resting, feeding, and spawning locale for sturgeon with a thriving fishery in the late nineteenth century. Rao and Schwab (2007) provide a description of the currents in this locale that show what must have been important features of the sturgeon’s habitat preferences. The Niagara River delta was fouled with raw sewage from upstream cities, poisoned by pesticides partly as a result of massive spraying of Buffalo’s elms with DDT against the Dutch elm bark beetle in the 1950s, and contaminated by the chemical industries along the shores upstream. Similar physical and chemical alterations have been documented to affect lake sturgeon elsewhere. Elimination of lake sturgeon spawning habitats and innumerable low-head dams were associated with extirpation of lake sturgeon in the Red River of the North (Aadland et al. 2005). Freeman et al. (2005) described how migration barriers have resulted in the listing of lake sturgeon as threatened in the Alabama River system. Habitat fragmentation has decimated lake sturgeon numbers in the Wabash River (Gammon 2005) and the Wisconsin River (Lyons 2005). One of the key sets of dynamic macro-features in pristine aquatic ecosystems in our basin was the incidence of massive episodic rheic features including river floods, shoreline storms, surface seiches, and thermal bars. When and where these

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were relatively predictable and gradual—on annual, decadal, and century-long scales—they were part of the natural context for healthy adapted ecosystems. On balance, what invading European settlers did to the streams and adjacent lands predisposed them to more violent consequences of floods, storms, and seiches, and they became more erratic, “peaky,” and erosive than was desirable for the kinds of stream biota that had adapted to the previous less violent and more gradual episodic regimes. Such changes have been particularly obvious in rivers in urbanized, surface mined, and overcultivated catchment areas (Dodge 1989). Hydrological innocents among the European invaders tried to counter the bad consequences of damming, diking, channelizing, hardening of shores, and other changes on land by additional and more massive earthen, concrete, and steel confining structures. Such obstructions in turn led to an increase in frequency and intensity of particularly destructive environmental events further downstream. More steel and concrete was often added, and the waters even farther downstream adapted again with increased violence, unpredictability, flashiness, and frequency of the most extreme manifestations in this pathological feedback syndrome. By the year 1970, the steel-and-concrete syndrome interacting with the channelize-dam-drain-and-dike syndrome had helped to transform large parts of our natural riverine ecosystem to a type dominated increasingly by an artificial “reservoirine” syndrome in form and behavior. This artificiality has been most prominent the further south one looks in the basin. But efforts to reverse this history are under way throughout our basin, mostly at a local level with removal of small dams that now serve no useful economic purpose (Evans and Gottgens 2007). One ecological example of the effects of the transformation of our waters from riverine to reservoirine is the fish-related part of this deep ecosystemic transformation. The 200 or so fish species of the near-pristine state were generally adapted to benthic riverine features in these waters and to shallow waters of the riparian ecotone. Currently the dominant fish species are aliens preadapted to enriched pelagic lacustrine (or marine continental-shelf waters) that experience less difficulty in our “reservoirine waters” than do our native species. Again, other human abuses have contributed importantly to this transformation of the fish assemblages from riverine to “reservoirine” (see box 1).

Ecological Effects of Fishing on Lake Sturgeon In ecosystems like those in our Laurentian Basin, the ecological effects of conventional fishing in historic times need to be teased out from the effects of other human

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stresses on these ecosystems. In our Laurentian Basin, such analyses, building on current empirical studies, have been reported as far back as the 1850s, when George Perkins Marsh undertook an investigation for the Vermont government to discover why the brook trout fisheries were failing in its streams. More recently, a team effort used a “forensic ecosystem approach” to infer causes for the failure of walleye fisheries in western Lake Erie in the 1960s (Regier, Applegate, and Ryder 1969). This was a difficult task because—as we came to realize—some generalized symptoms or effects of stress were not specific to particular stresses. The complex of nonspecific, nondiagnostic symptoms together with some specific, diagnostic symptoms could be termed a “general adaptive syndrome, GAS, to ecosystemic stress” as analogous to the GAS that Hans Selye had inferred with respect to the physiology of stressed mammalian organisms (Rapport et al. 1985). That 1969 walleye study in turn led to a Symposium on Salmonid Communities in Oligotrophic Lakes (Loftus and Regier 1972) in which severe effects of major stresses were teased apart. Again, a GAS was inferred; it involved suppression of the dominant “riverine” salmonid complex of species and emergence of a “reservoirine” complex of alien species, though we did not use the terms “riverine” and “reservoirine” at that time to summarize key features of these two types. Particularly for lake sturgeon, everybody’s favorite colleagues of years ago, W. Bev Scott and Ed J. Crossman (1973), summarized how European settlers had fished sturgeon nonsustainably to lead to deep population reductions already in the nineteenth century. In an exhaustive comparative study of historical changes in the fish assemblages in three bays of our basin, Whillans (1979) included information about lake sturgeon. He critically examined accounts of events from early settlement by Europeans in the eighteenth century to 1976 in Toronto and Burlington Bays of Lake Ontario and Inner Long Point Bay of Lake Erie: • Records of lake sturgeon in Toronto Bay began in 1800 when it was not one of the more preferred species by those with refined tastes. Sturgeon used Toronto Bay only for brief periods in the spring, when they congregated in the estuary of the Don River. By the 1840s, sturgeon were reported to be declining at a time while fishing was not intense; deforestation, milling, and construction of dams likely stressed the sturgeon population. • Records of sturgeon in Burlington Bay began in 1855. Transformations in this fish assemblage were first noted between 1859 and 1877; for example, the sturgeon population declined during this period. Between 1878 and 1892, sturgeon were fished heavily and their spawning grounds may have been harmed by water pollution.

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• Records of sturgeon in Inner Long Point Bay began between 1880 and 1885. Between 1892 and 1910, numbers declined in association with increased fishing pressure; between 1924 and 1937 sturgeon numbers fluctuated. Though Whillans noted numerous adverse practices on these bay ecosystems besides nonsustainable fishing, he inferred that the main cause of lake sturgeon decline in all three bays was selective commercial exploitation, which was not intense in waters in and near these bays until the 1870s. Historically, under rapidly intensifying fisheries in the second half of the nineteenth century that were conducted in the traditional European way (preferentially removing the largest sturgeon available with relatively nonselective gear), the abundance of lake sturgeon in our basin waters apparently did not show evidence of greatly increased fluctuations in abundance from year to year. Their abundance, as reflected in primitive measures of catch per unit effort of fishing, decreased gradually from year to year rather than fluctuating wildly. Moderate fluctuations in catch from year to year may have been due more to differences in the weather that affected temporal and areal aspects of fish schooling and fisher access and success rather than to the overall availability of sturgeon. But some degree of fluctuation may sometimes have been caused by conventional exploitation (see Whillans 1979). In our basin, very large sturgeon were occasionally caught decades after the commercial sturgeon fishery was catching only few sturgeon that were mostly small. Perhaps some sturgeon preferred habitats that were not fished intensely, with a consequence that something like sturgeon refuges may have existed and persisted somewhere in our basin’s waters. Venturelli, Shuter, and Murphy (2009, 919) conducted a meta-analysis of timeseries data on the reproductive success of 25 species of exploited marine fishes as related to the size and age distributions of the spawning population. They cautiously inferred that “a population of older, larger individuals has a higher maximum reproductive rate than an equivalent population of younger, smaller individuals, and that this difference increases in the reproductive lifespan of the population. These findings (i) establish an empirical link between population age structure and reproductive rate that is consistent with strong effects of maternal quality on population dynamics and (ii) provide further evidence that extended age structure is essential to the sustainability of many exploited fish stocks.” The report by Venturelli, Shuter, and Murphy supports the informed judgment of various experts on fisheries in our Laurentian Basin for many decades past that ensuring the presence of numerous large spawners in a fish population should be part of a sustainable fisheries policy. Refuges from which fishing is excluded rigorously may be a particularly helpful approach concerning the conservation of older and

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larger fish such as lake sturgeon. The underwater delta of the Niagara River at its mouth in Lake Ontario could be included in such a refuge for lake sturgeon. If additional research supports the work of these authors with respect to marine fisheries and also provides evidence that such inferences are valid with respect to freshwater fisheries and lake sturgeon in particular in our Laurentian Basin, then the regulatory and ethical codes of fishers should be revised to reflect those inferences (Dobson, Regier, and Taylor 2002).

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Benthification, Pelagification, and Surficialization Back to an ecosystemic general adaptive syndrome, GAS, adapted from the physiological GAS of Hans Selye: in our basin’s waters a dominant feature of this GAS was “pelagification” in that the normally dominant benthic association was suppressed and a somewhat unnatural pelagic association was fostered (Kay and Regier 1999). Again, such “pelagification” has often been termed “eutrophication,” a potentially misleading term as applied in our basin. More recently there have been reviews of relevant information about the failure of fisheries in multistressed enclosed seas (Caddy and Regier 2002). Such an “ecosystemic forensic approach” may not yet have been applied to problems in the Gulf of St. Lawrence, downstream from our basin ecosystem, which has its own sturgeon species. Studies like those mentioned above have generally led to inferences that the strains or adverse effects of numerous (but perhaps not all) human-related stresses that are typical of conventional economic and cultural development tend to interact synergistically in aquatic ecosystems to augment the adverse effects of each and thus to trigger a GAS as an ecosystemic phase or state shift. Apparently the effects of those other cultural stresses have not interacted with the effects of conventional fishing stress to counteract the deleterious effects of the latter. Instead, the negative effects of various stresses may more frequently have interacted synergistically with those of conventional fishing to exacerbate further the harm done to the native fish by such fishing. The artificial pelagification of the more stressed bays and shallow subbasins in our lakes that reached peak intensities several decades ago has been followed in turn during the past two decades by a suppression of that artificial pelagic association through the combined “benthification” to a new kind of artificial benthic association (alien zebra and quagga mussels, round goby, etc.) plus “surficialization” to an unusual kind of surface association (blooms of toxic algae, floating mats of decaying algae, etc.). This latter surface-bottom combination, with little systemic organization in

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the intervening waters, has contributed in some way to the emergence of a different type of “dead zone” in some deep waters, now with the presence of toxic organisms both at the top and at the bottom of these waters. Intense research is currently under way to describe and explicate phenomena that seem strange, at least at the scale and tempo with which they are appearing. Rationally reliable scientific methods for conducting meta-analyses in multistressed ecosystems are evolving gradually (Zwiers and Hegerl 2008). New generations of scientists in our basin will presumably come to use them, perhaps rather noisily at first. Humility in the face of uncertainty, as urged by Jasanoff (2007), has been a common, if not universal, trait within the invisible college of our basin’s researchers. New concepts and methods have come to be adopted smoothly, on the whole. With respect to other human-generated stresses than fishing, the transformation from a riverine to a reservoirine type of ecosystem must have been severely detrimental on balance for sturgeon. In particular, dams on rivers prevented sturgeon from ascending them to their ancestral spawning grounds. Also, emergence of oxygen-depleted bottom waters of reservoirine ecosystems (see below) likely served to exclude sturgeon from some of their ancestral habitat in summer and perhaps in winter.

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Phosphorus, Nitrogen, Oxygen, and “Dead Zones” In the aquatic ecosystems favored by lake sturgeon, some common chemicals to which humans are relatively insensitive should be at concentrations of only relatively few parts per million of the watery environment. In particular, this statement relates to dissolved oxygen and dissolved forms of some strongly oxidized, as well as some strongly reduced, “hydrogenized,” but still biologically reactive compounds of phosphorus and nitrogen. The role of biologically reactive phosphorus compounds, which dissociate in water to yield phosphate ions, came into clear focus in the late 1960s in our basin. Phosphates (and nitrates, see below) are taken up during photosynthesis by plants with oxygen produced as a waste product; concurrently, and at times when photosynthesis is not occurring, oxygen is taken up in respiration by most of the organisms that are abundant in natural ecosystems of our basin. Natural ecosystems in pristine states in our basin have ways to limit the concentrations of particularly reactive phosphate ions, for example, by complexing them—and thus rendering them “refractory” or less reactive biologically—on organic particles that originate from the slow decomposition of woody debris, say. But such

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complexing can be overridden by reductions in dissolved oxygen due to bacterial respiration during decomposition of readily oxidized organic material such as sewage and dead plankton. For purposes of respiration when dissolved oxygen falls to low levels, some organisms can steal oxygen from the strongly oxidized phosphate ions and thus transform them into more soluble phosphite ions (which can subsequently be oxidized again into temporarily uncomplexed phosphate in well-oxygenated water and contribute to photosynthesis again, etc.). So a positive feedback loop can emerge, as at the bottom of deep basins within lakes and reservoirs, that can result in a large anoxic “dead zone” during the four or five summer months when the overlying waters are stratified by temperature that blocks access of atmospheric oxygen to the deeper zones. In our basin, deep anoxic waters are colder (at 4–10°C, say) than those preferred by lake sturgeon (15–17°C or a couple of degrees warmer). But under unusual storm conditions, say involving a strong atmospheric front passing rapidly over a lake, waves may be induced at the surface and in the thermocline that separates cold and warm waters (Rao and Schwab 2007). Such a wave or seiche may result in anoxic waters sloshing into shallower, usually cool-water areas to the disadvantage of normal organisms that live there. Anoxic waters displaced by such an internal wave or seiche may harm strongly benthic lake sturgeon. In Lake Erie, for example, the band of water along the bottom that falls in the temperature range of 15–17°C in summer, as preferred by lake sturgeon, is not wide; thus sturgeon may sometimes have to choose between anoxic colder waters and oxic warmer waters rather than waters in their preferred temperature range. Such a seiche that induces them to greater mobility than usual to escape cold anoxic water may make them more vulnerable to capture in stationary fishing gear. Trapped sturgeon may then be “drowned” by the anoxic water, as has occurred with other species. Of course, what is usually termed a “dead zone” is not devoid of life. Unless it is poisoned by some nonselective toxic chemical or biocide, such a zone teems with life. But these life-forms, some of which resemble our genetic ancestors of billions of years ago and some that live in our own gastrointestinal tracts, thrive in the absence of dissolved oxygen. A relevant report by P. J. Mulholland and 30 colleagues (2008) is titled “Stream Denitrification across Biomes and Its Response to Anthropogenic Nitrate Loading.” This team undertook a carefully designed experiment that involved loading isotope-labeled nitrate compounds into 72 streams in eight regions of the coterminous United States. The experimental design was based on many previous, less-comprehensive studies. The team distinguished how the denitrification process differed in streams of three size categories draining smallish catchment areas that were relatively natural or subjected to urban or agricultural regimes. The team also distinguished how

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those denitrification processes differed as influenced by different concentrations of the soluble nitrate ions in the nine experimental cells (three size categories by three kinds of predominant human activity). In their 2008 report, the authors did not focus on historic changes in any stream, say as it progressed from a cold, permanently free-flowing first- or second-order stream to an underground conduit or an intermittent stream following urban or agricultural development. Their information could perhaps be interpreted to imply that such trimming away of first- and second-order streams made those waterways less effective in denitrifying excessive loads of nitrates that washed in from the overly enriched lands of a catchment area. They conclude: Our findings underscore the management imperative of controlling nitrogen loading to streams and protecting or restoring stream ecosystems to maintain or enhance their nitrogen removal functions. Controlling loading to streams and stream nitrogen export is a proven solution to eutrophication and hypoxia problems in downstream inland and coastal waters. Our findings suggest caution before implementing policies (for example, reliance on intensive agriculture for biofuels production) that may yield massive land conversions and higher nitrogen loads to streams. Associated increases in streamwater [nitrate] concentration may reduce the efficacy of streams as nitrogen sinks [through denitrification], yielding synergistic increases in downstream transport to estuaries and

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coastal oceans. (Mulholland et al. 2008, 204)

If each of the separate narratives of phosphates and nitrates in our basin ecosystems is already complex, the combined narrative is even more so. For starters, if phosphates are relatively more abundant than nitrates, the rapid photosynthesis and growth of aquatic plankton may demand more nitrates than are being brought into such waters through inflows. Surface blooms of specialized algae may then occur spontaneously that have the ability to transform gaseous nitrogen from the atmosphere into oxygenated nitrogen to be used in the rapid photosynthesis. Again, this may lead to a vicious cycle, or positive feedback, with more living and then dead plant material, more decomposition, less dissolved oxygen, regeneration of dissolved phosphates, more fixing of atmospheric nitrogen by surface algae, eruption of toxic algae, and so on. Altogether such a complex syndrome is dominated by nonlinear processes, including feedback and phase or state shifts, for which no conventional quantitative methods provide reliable predictions. In Lake Erie a “dead zone” that appeared with increasing regularity and size some six or more decades ago was part of the ecosystem-wide pelagification transformation. It was largely remediated through various controls on phosphate loading; and the relatively rapid natural flushing of Lake Erie helped. But then about

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two decades ago a somewhat different kind of “dead zone” appeared in Lake Erie as well as strange ecosystemic functions in other areas of bottom sediment that were not within a massive “dead zone.” Zebra and quagga mussels seemed to be playing strong roles in the strange behavior of Lake Erie’s aquatic ecosystem, as concerns the dynamics of phosphates, nitrates, oxygen, and various biota. Poisonous substances were being generated in the benthos in unnaturally large amounts that killed fish and then killed the birds that ate the fish. Poisonous algal blooms appeared more frequently at the surface of these waters. The many researchers at work studying this issue have yet to come to a consensus on what is causing this unwanted complex phenomenon. Presumably the ecosystem pathologies caused by excessive loadings of phosphates and nitrates into our basin waters, including complex effects that follow as the ecosystems try to adapt to these loadings with their synergistic feedback and interactive loops, do not make life more pleasant for lake sturgeon and most other native species of our basin. Unlike some alien species, lake sturgeon do not thrive in severely modified, reservoirine ecosystems. Of equivalent importance to the phosphate-nitrogen-oxygen complex pathology sketched above is the issue related to chemicals that strongly affect the genetic, hormonal, immunological, and other information capabilities of many organisms, including fish, birds, and humans. Chemical concentrations with the phosphatenitrate-oxygen complex sketched above are commonly measured in parts per million. However, concentrations of hazardous contaminants may be dangerous at levels of parts per billion or trillion. Many pharmaceutical, cosmetic, and sanitizing substances find their way into our waters via the sewage system; adverse ecosystem effects from their use have been coming into focus. Because of the widespread reliance on chemical medication, many in the medical profession, including both clinicians and public health types, seem compromised on this issue. But some public health professionals are trying to come to terms with adverse effects of residual medications prescribed by clinical physicians. It seems unlikely that permanent presence of hundreds of such chemical contaminants in these waters makes life more pleasant for lake sturgeon. Recent studies seem to imply that when some organisms that thrive in waters nearly saturated with oxygen are subjected to waters with depleted but not zero levels of oxygen concentrations, they may exhibit symptoms like those triggered by some chemical contaminants, for example, endocrine disruption. If so, then an anoxic “dead zone” may develop a surrounding penumbra of hazardous waters with depleted levels of oxygen. Do fish like sturgeon sense such oxygen diminution, and do they have behavioral capabilities to evade such waters?

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Climate Change A meta-analysis of empirical studies relevant to climate change by Rosenzweig and colleagues (2008) is titled “Attributing Physical and Biological Impacts to Anthropogenic Climate Change.” The authors examined thousands of data series, each longer than a 20-year duration, from across the globe. Many of these series relate to phenology; for example, the timing of a species’ life-history events in spring. For North America, the authors found 94 percent of the 405 observed physical changes (that were statistically significant) to be consistent with what the authors would have expected with climate warming. Similarly, they found 88 percent of the 579 biological changes (that were statistically significant) to be consistent with their understanding of effects of climate warming. If there had as yet been no effects from any climate change, then the expected percentages would have been 50 percent in both cases. The authors apparently found no scientific reports, appropriate to their meta-analysis, concerning temperature-related changes apparent in extant data series from the aquatic ecosystems in our basin. Numerous researchers have used a variety of approaches to assess some likely effects of forecasted scenarios of climate change on fish in our basin (e.g., Magnuson et al. 1989; Everett et al. 1995). Ralph Pentland (personal communication) estimates that climate change has already lowered the level of the Great Lakes by 5 cm, with more lowering expected. In general, researchers have inferred that the “moderate” extent of climate change to be expected in our basin would, in itself, not likely create insurmountable difficulties for many of our native species. Perhaps more correctly: forecasted climate changes would not likely cause many serious difficulties in the absence of the adverse effects of other human-caused stresses such as transformation of these waters from a riverine to a reservoirine state; introductions of alien species preadapted to a warmer climate; emergence of anoxic and toxic bottom waters and toxic surface algal blooms; and so on. But it seems unlikely that any of the other human-induced changes would predispose these systems to respond favorably to climate change, at least with respect to native species like the sturgeon. There may well be some alien species already present that will thrive with warming due to climate change, and more (including human pathogens) that are preadapted to a warmer climate will likely invade, or be released into our basin.

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Concluding Comments Our basin’s fish and fisheries “hit the wall” ecologically and economically about half a century ago, in the 1950s. Numerous remediative and rehabilitative measures have been undertaken since then. The record concerning corrective measures with exploitative fishing and loading of phosphate nutrients shows important improvements. The record with channel modifications, nitrates, contaminants and alien species is at best ambiguous, but generally poor. Climate change caused by continued economic and population growth will likely interact badly with remaining features of the old stresses as well as with new stresses for which current remediative and rehabilitative programs are clearly inadequate and precautionary methods are still weak (Regier et al. 1989). The lake sturgeon has been important to human fishers for at least two thousand years, judging from work by archaeologists (as in Smith 2004) and historians (as in Thwaites 1899), both with respect to sturgeon in the Straits of Mackinac. A quick search of the Internet has yielded five bays, 13 rivers, and two creeks in our basin named after sturgeon, not to mention human settlements with that name. Somehow, several small sturgeon populations have survived the past two centuries of intense ecosystemic abuse in our basin. During recent centuries, lake sturgeon populations were suppressed and some were extirpated in our basin’s waters (Scott and Crossman 1973, and other authors). A thorough forensic scientific study to identify the specific cause(s) of an extirpation of a particular lake sturgeon population was never undertaken (though work by Whillans [1979] deserves honorable mention) and cannot now be done in a fully convincing way. In recent decades some populations that survived all the abuse but in low numbers have been protected and are slowly increasing in abundance. If a full recovery of a population may take several of their generations, each of which may be 10 years long with sturgeon, then we should not expect to encounter a rehabilitated sturgeon population in our basin for half a century or so, and then only if ecosystemic rehabilitation—now stalled–resumes energetically. Assuming that successful sturgeon reproduction eventually comes to be dominated by large females (Venturelli, Shuter, and Murphy 2009), as with other long-lived large species, then we may not see widespread rehabilitation for a century. Similar declines and extirpations of sturgeons have occurred throughout their ranges as a result of reproductive failure associated with fisheries exploitation, water pollution, and migration barriers. Many sturgeon species are threatened or endangered (Williams et al. 1989; Oberdorff and Hughes 1992; Otel 2007; Jelks et al. 2008) with little hope for recovery in ecosystems that have suffered severe

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degradation. Unlike small, short-lived, and locally endemic species, the sturgeons are large, long-lived, and wide-ranging species that require unfragmented large water bodies in good chemical, physical, and biological condition to thrive. Such species were reported by Hughes, Rinne, and Calamusso (2005b) to be most affected by the flow and channel modifications common to large rivers. A variety of organisms, including some fish species like lake trout and walleye, are being used as integrative indicators of ecosystem adaptations (often of an unwanted and thus pathological type) to various cultural stresses, acting separately and jointly. It is timely now to produce appropriate documentation to legitimize lake sturgeon as an integrative indicator of how our basin’s aquatic ecosystem is faring with respect to a preferred, healthy ecosystem. Aboriginal folk, as friends or kin of the sturgeon, could be recognized as guardians of these survivors of centuries of abuse by European invaders, where such Aboriginals still practice valued traditional ways.

NOTES

Gavin Christie, David Maraldo, Kevin Reid, and Brian Shuter advised helpfully.

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Robert Hughes’s work on this manuscript was partially supported by USEPA cooperative agreement CR831682-01 with Oregon State University. The contents of this chapter should not be assumed to have an endorsement from any agency with which any of the authors have or have had a formal connection. So no endorsement has been sought, or offered, by the Great Lakes Fishery Commission, the U.S. Environmental Protection Agency, or the International Joint Commission.

REFERENCES

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Evans, J. E., and J. H. Gottgens, eds. 2007. Dam removals and river channel changes. Journal of Great Lakes Research 33 (special issue 2): 87–193. Everett, J. T., A. Krovnin, D. Lluch Belda, E. Okemwa, H. A. Regier, and J.-P. Troadec. 1995. Fisheries. In: Climate change 1995: Impacts, adaptations and mitigation of climate change, scientific-technical analyses. Contributions of Working Group II to the second assessment report of the Intergovernmental Panel on Climate Change. R. T. Watson, M. C. Zinyowera, and R. H. Ross, eds. Cambridge University Press. Freeman, M. C., E. R. Irwin, N. M. Burkhead, B. J. Freeman, and H. L. Hart Jr. 2005. Status and conservation of the fish fauna of the Alabama River system. In: Historical changes in large river fish assemblages in the Americas. J. N. Rinne, R. M. Hughes, and B. Calamusso, eds. American Fisheries Society Symposium 45. American Fisheries Society. Gammon, J. R. 2005. Wabash River fishes from 1800 to 2000. In: Historical changes in large river fish assemblages in the Americas. J. N. Rinne, R. M. Hughes, and B. Calamusso, eds. American Fisheries Society Symposium 45. American Fisheries Society. Gottgens, J. F., and J. E. Evans. 2007. Dam removals and river channel changes in northern Ohio: Implications for Lake Erie sediment budgets and water quality. Journal of Great Lakes Research 33 (Supplement 2): 87–89. Gregory, S. V., K. L. Boyer, and A. M. Gurnell, eds. 2003. The ecology and management of wood in world rivers. American Fisheries Society Symposium 37. Gregory, S. V., F. J. Swanson, W. A. McKee, and K. W. Cummins. 1991. An ecosystem perspective on riparian zones. Bioscience 41:540–551. Hughes, R. M., J. N. Rinne, and B. Calamusso. 2005a. Introduction to historical changes in large river fish assemblages of the Americas. In: Historical changes in large river fish

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Lyons, J. 2005. Fish assemblage structure, composition and biotic integrity of the Wisconsin River. In: Historical Changes in Large River Fish Assemblages in the Americas. J. N. Rinne, R. M. Hughes, and B. Calamusso, eds. American Fisheries Society Symposium 45. American Fisheries Society. Magnuson, J. J., D. K. Hill, H. A. Regier, J. A. Holmes, J. D. Meisner, and B. J. Shuter. 1989. Potential responses of Great Lakes fishes and their habitat to global climate warming. In: The potential effects of global climate change on the United States, Appendix E, Aquatic Resources. J. B. Smith and D. A. Tirpak, eds. EPA-230–05–89–055. U.S. Environmental Protection Agency. Miller Reed, K., and B. Czech. 2005. Causes of fish endangerment in the U.S., or the structure of the American economy. Fisheries 30(7): 36–38. Minshall, G. W., K. W. Cummins, R. C. Petersen, C. E. Cushing, D. A. Burns, J. R. Sedell, and R. L. Vannote. 1985. Developments in stream ecosystem theory. Canadian Journal of Fisheries and Aquatic Science 42:1045–1055. Moerke, A. H., and G. A. Lamberti. 2006. Relationships between land use and stream ecosystems: A multistream assessment in southwestern Michigan. In: Landscape influences on stream habitats and biological assemblages. R. M. Hughes, L. Wang, and P. W. Seelbach, eds. American Fisheries Society Symposium 48. American Fisheries Society. Mulholland, P. J., A. M. Helton, G. C. Poole, and 28 others. 2008. Stream denitrification across biomes and its response to anthropogenic nitrate loading. Nature 452:202–205. Naidoo, R., and W. L. Adamowicz. 2001. Effects of economic prosperity on numbers of threatened species. Conservation Biology 15:1021–1029. Niemi, G. J., and J. R. Kelly, eds. 2007. Coastal indicators. Journal of Great Lakes Research 33 (special issue 3): 1–318.

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Regier, H. A., V. C. Applegate, and R. A. Ryder. 1969. Ecology and management of the walleye in western Lake Erie. Great Lakes Fishery Commission Technical Report 15. Regier, H. A., and J. J. Kay. 1996. An heuristic model of transformations of the aquatic ecosystems of the Great Lakes–St. Lawrence River Basin. Journal of Aquatic Ecosystem Health 5:3–21. Regier, H. A., P. Lin, K. K. Ing, and G. A. Wichert. 1996. Likely responses to climate change of fish associations in the Laurentian Great Lakes Basin: Concepts, methods and findings. Boreal Environment Research (Helsinki) 1:1–15. Regier, H. A., and K. H. Loftus. 1972. Effects of fisheries exploitation on salmonid communities in oligotrophic lakes. Journal of the Fisheries Research Board of Canada 29:959–968. Regier, H. A., R. L. Welcomme, R. J. Steedman, and H. F. Henderson. 1989. Rehabilitation of degraded river ecosystems. In: Proceedings of the International Large River Symposium (LARS). D. P. Dodge, ed. Special Publication of the Canadian Journal of Fisheries and Aquatic Sciences 106. Canada Department of Fisheries and Oceans. Richards, J. S. 1976. Changes in fish species composition in the Au Sable River, Michigan from the 1920s to 1972. Transactions of the American Fisheries Society 105:32–40. Rinne, J. N., R. M. Hughes, and B. Calamusso, eds. 2005. Historical Changes in Large River Fish Assemblages in the Americas. American Fisheries Society Symposium 45. American Fisheries Society. Rose, A. 2005. Economic growth as a threat to fish conservation in Canada. Fisheries 30(8): 36–38. Rosenzweig, C., D. Karoly, M. Vicarelli, P. Neofotis, Qigang Wu, G. Casassa, A. Menzel, T. L. Root, N. Estrella, B. Seguin, P. Tryjanowski, Chunzhen Liu, S. Rawlins, and A. Imeson.

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2008. Attributing physical and biological impacts to anthropogenic climate change. Nature 453:353–357. Schlesinger, D. A., and H. A. Regier. 1982. Climatic and morphoedaphic indices of fish yields from natural lakes. Transactions of the American Fisheries Society 111:141–150. Scott, W. B., and E. J. Crossman. 1973. Freshwater fishes of Canada. Fisheries Research Board of Canada Bulletin 184. Sedell, J. R., and J. L. Froggatt. 1984. Importance of streamside forests to large rivers: The isolation of the Willamette River, Oregon, USA, from its floodplain by snagging and streamside forest removal. Internationale Vereinigung für theoretische und angewandte Limnologie Verhandlungen 22:1828–1834. Sedell, J. R., R. J. Steedman, H. A. Regier, and S. V. Gregory. 1991. Restoration of human impacted land-water ecotones. In: Ecotones: The role of landscape boundaries in the management and restoration of changing environments. M. M. Holland, P. G. Risser, and R. J. Naiman, eds. Chapman and Hall. Smith, B. A. 2004. The gill net’s “native country”: The inland shore fishery in the northern Lake Michigan basin. In: An upper lakes archeological odyssey. W. A. Lovins and C. E. Cleland, eds. Wayne State University Press. Smith, P. W. 1971. Illinois streams: A classification based on their fishes and an analysis of factors responsible for disappearance of native species. Illinois Natural History Survey, Biological Notes 76. Smith, S. H. 1995. Early changes in the fish community of Lake Ontario. Great Lakes Fishery Commission Technical Report 60. Sproule-Jones, M. 2002. The restoration of the Great Lakes. University of British Columbia Press.

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Steedman, R. J. 1986. Historical streams of Toronto. Toronto Field Naturalist 382:14–18. Stoddard, J. L., D. V. Peck, S. G. Paulsen, J. Van Sickle, C. P. Hawkins, A. T. Herlihy, R. M. Hughes, P. R. Kaufmann, D. P. Larsen, G. Lomnicky, A. R. Olsen, S. A. Peterson, P. L. Ringold, and T. R. Whittier. 2005. An ecological assessment of western streams and rivers. EPA 620/R05/005, U.S. Environmental Protection Agency, Washington, DC. Thwaites, R.G., ed. 1899. The Jesuit relations and allied documents: Travels and explorations of the Jesuit missionaries in New France 1610–1791. Vol. 55, 1670–1672. Burrows Brothers. Http://puffin.creighton.edu/jesuit/relations/relations_55.html. Trautman, M. B. 1977. The Ohio country from 1750 to 1977: A naturalist’s view. Ohio Biological Survey Biological Notes No. 10, Ohio State University. Triska, F. J. 1984. Role of wood debris in modifying channel morphology and riparian areas of a large lowland river under pristine conditions: A case history. Internationale Vereinigung für theoretische und angewandte Limnologie Verhandlungen 22:1876–1892. Van Sickle, J., and S. G. Paulsen. 2008. Assessing the attributable risks, relative risks, and regional extents of aquatic stressors. Journal of the North American Benthological Society 27:920–931. Venturelli, P. A., B. J. Shuter, and C. A. Murphy. 2009. Evidence of harvest-induced maternal influences on the reproductive rates of fish populations. Proceedings of the Royal Society B 276:919–924. Whillans, T. H. 1979. Historic transformations of fish communities in three Great Lakes bays. Journal of Great Lakes Research 5(2): 195–215. Williams, J. E., J. E. Johnson, D. A. Hendrickson, S. Contreras-Balderas, J. D. Williams, M.

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Habitat, Foods, and Feeding

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Habitat As the name implies, lake sturgeon are commonly found in lakes, and because of their biology they occupy large lakes. The Great Lakes lie at the center of the species native range (Harkness and Dymond 1961) and these lakes once supported a large concentration of lake sturgeon. Lake sturgeon were in all likelihood one of the most abundant large-bodied fish species in the Great Lakes prior to extensive settlement of the region, with estimates of abundance exceeding 16 million fish in all Great Lakes combined. The lake sturgeon commercial fishery that developed in the Great Lakes provides an indication of historic lake sturgeon abundance. According to statistics compiled by the Great Lakes Fishery Commission, during the peak of the Great Lakes commercial lake sturgeon fishery in the late 1800s, an average of over 4 million pounds of lake sturgeon was harvested annually. The maximum harvest occurred in 1885, when over 8.6 million pounds were harvested (Baldwin et al. 1979). The abundance of lake sturgeon in each of the Great Lakes was closely tied to the habitat and productivity of the individual lake. Lake sturgeon harvest was greatest in Lake Erie and was lowest in Lake Superior. Lake Erie has a large basin and is a relatively shallow, warm, and productive lake. There are also numerous large productive rivers n

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that feed into the lake. However, in Lake Superior abundance was much less than in the other lakes. Although it is the largest of the Great Lakes, Lake Superior is much deeper, colder, and less productive than the lower lakes. Estimated lake sturgeon abundance in 1885 was over 11 million fish in Lake Michigan but was only 870,000 in Lake Superior (Hay-Chmielewski and Whelan 1997). Much has changed in the Great Lakes since the late 1800s, including widespread alteration of habitat, and lake sturgeon abundance in the Great Lakes today is estimated to be less than 1 percent of the peak abundance of the 1800s (Hay-Chmielewski and Whelan 1997). The native range of lake sturgeon includes the Great Lakes and St. Lawrence River, the Hudson Bay drainage of Canada, and the Mississippi River upstream of northern Mississippi (Scott and Crossman 1973). Lake sturgeon were historically abundant and found in many of the large rivers and lakes in all of these major drainages. However, the status of lake sturgeon throughout its native range is similar to that of the Great Lakes; its abundance is greatly reduced from the peak before extensive settlement. The demise of lake sturgeon throughout the species range can be attributed to excessive harvest, but habitat changes have also played an important role in the decline. In the Great Lakes, these habitat changes are also likely preventing lake sturgeon populations from rebounding. Broadly defined, habitat includes the physical places occupied by an organism and can include the other organisms present with which a species interacts. In the case of lake sturgeon, the physical habitats occupied are varied and depend on season and the age of the fish. Most of a lake sturgeon’s life is spent in lake or large river habitat feeding and growing, so they are associated with habitats that produce the organisms on which lake sturgeon feed. Lake sturgeon prey on benthic animals or those that live directly on or in the bottom substrates of lakes and rivers. The bottom substrates in relatively shallow waters of lakes are the most productive for benthic animals. As a result, lake sturgeon spend most of their time in relatively shallow water in lakes, typically less than 30.5 m (100 ft) deep and most often less than 9 m (30 ft) deep in the Great Lakes (Harkness and Dymond 1961). Because fish are cold-blooded and their body temperature, and therefore metabolic processes, are dictated by the temperature of the surrounding water, most fish exhibit preferences for specific water temperatures that optimize their growth and development. The water temperature preferences of lake sturgeon are not as well understood as for other species. For example, several trout species are known to actively seek out cold water and largemouth bass actively seek warm water. Lake sturgeon water temperature preference for spawning is well known. They spawn when water temperatures reach about 12°C (54°F) and will continue until water temperature reaches 18°C (65°F) (Harkness and Dymond 1961; Kempinger 1988; Auer 1996a). However, little is known of water temperatures that adult lake sturgeon seek out

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during the extended period between spawning. Based on results from experimental hatchery rearing of lake sturgeon, it is likely that they seek relatively warm water. In an experimental hatchery setting, lake sturgeon growth was greatest when water temperatures were 15–18°C (60–64°F), and mortality was observed when water temperatures reached 22°C (72°F), suggesting this is near the upper lethal limit for lake sturgeon (Wehrly 1995; Diana, Webb, and Essington 2003). However, adult lake sturgeon congregations have been found in Lake St. Clair at a water temperature of 25.6°C (78°F) (M. Thomas unpublished). In streamside hatcheries that circulate river water, age-zero lake sturgeon have continued to feed at water temperatures as high as 28°C (82°F) for a few hours midday (E. Baker unpublished). Radio telemetry studies of juvenile (ages 1–5) lake sturgeon suggest that they move from deeper cool water during the daytime to shallow water at night and back to cooler deep water during the day (Holtgren and Auer 2004). Presumably the fish are feeding in the shallower water at night and resting in the cooler water during daytime, when shallow water habitats may become too warm, or move into deep water in response to intense light. Lake sturgeon roam shallow nearshore Great Lakes waters, feeding throughout the year, and are known to travel long distances in search of food. For example, Green Bay in Lake Michigan is shallow and productive. It is also known to have a relatively high concentration of lake sturgeon when compared to other areas in Lake Michigan. Recent research has demonstrated that the lake sturgeon found in Green Bay originated in rivers that empty directly into Green Bay (Menominee River, Peshtigo River, etc.) as well as rivers on the eastern shore of Lake Michigan, including the Manistee (Bott 2006). The productive waters of Green Bay appear to be an important feeding area for lake sturgeon from throughout the Lake Michigan basin, and sturgeon may migrate to Green Bay to feed and grow and then return to their natal rivers to reproduce. The same appears to be true of Saginaw Bay in Lake Huron, Lake St. Clair, and the bays at the western end of Lake Superior. Because lake sturgeon occupy relatively shallow nearshore waters of the Great Lakes and feed on the bottom, their migrations most likely follow the shoreline and do not cross the open waters of the lake. In large rivers where lake sturgeon may live their entire lives (e.g., Niagara River, St. Clair River), they also occupy the areas that are most productive for the benthic organisms they feed on and may travel long distances in search of food and suitable spawning habitat. The only time that lake sturgeon abandon lake and river feeding habitat is when, as mature adults, they respond to the urge to spawn. Spawning occurs in the spring, and adults return to the rivers of their origin to spawn. Because lake sturgeon roam great distances in search of food, the return trip to spawning rivers for some can be very long. For example, tag return information from Lake Superior indicates

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that lake sturgeon that spawn in the Sturgeon River in Michigan’s Upper Peninsula near Houghton travel as far east as Whitefish Bay near Sault Ste. Marie, Michigan, and as far west as Chequamegon Bay near Ashland, Wisconsin, while feeding in Lake Superior; distances in excess of 200 km (125 miles) (Auer 1999). Tag return information from Lakes Michigan and Huron has shown that lake sturgeon from Lake Winnebago, Wisconsin, that spawn in the Wolf River have traveled down the Fox River to Green Bay and continued on to Saginaw Bay near Saginaw, Michigan, in Lake Huron, a distance of over 600 miles (1,000 km). As winter gives way to spring and adult lake sturgeon shift their focus to spawning, they begin what can be a long migration up rivers to spawning habitats. As lake sturgeon ascend rivers, they depend on deep holes for rest and concealment (Threader, Pope, and Schaap 1998). Prior to extensive settlement of the Great Lakes region, most, if not all, large rivers were used by spawning lake sturgeon, and spawning migrations would take the fish dozens to over 100 miles (160 km) upstream to reach suitable habitat. Now, in the Lake Superior drainage, lake sturgeon ascend the Sturgeon River to spawn near the base of Prickett Dam, a distance of 43 miles (69 km) (Auer 1996a). In the Wolf River, Wisconsin lake sturgeon travel upstream 155 miles (250 km) from Lake Winnebago to reach their spawning habitats (Kempinger 1988). In many rivers in Canada lake sturgeon migrate upstream until they reach an impassable barrier such as a natural waterfall and spawn at the base of the falls (Harkness and Dymond 1961). Most river habitats used by spawning lake sturgeon can be best described as rapids characterized by high water velocity with clean substrates of large gravel, cobble, or even bedrock. An important component of spawning habitat is the presence of abundant interstitial spaces or gaps between the rocks where lake sturgeon eggs can settle and be concealed from predators. Water depth apparently has little influence on spawning habitat selection, as lake sturgeon have been observed spawning in the St. Clair and St. Lawrence rivers in water up to 60 feet (18 m) deep (LaHaye et al. 1992), while in smaller rivers like the Sturgeon River in Michigan’s Upper Peninsula and the Black and Manistee rivers in the northern Lower Peninsula of Michigan, lake sturgeon have been observed spawning in water only two to three feet (1 m) deep (Chiotti et al. 2008). Because depth does not appear to influence spawning site selection, the most important physical habitat cues for lake sturgeon spawning are apparently adequate water velocity and the presence of clean substrate. Reported water velocities at lake sturgeon spawning sites in rivers are in excess of 3 ft/s or 0.91 m/s (Threader, Pope, and Schaap 1998; Chiotti et al. 2008). There are also persistent, albeit unconfirmed, reports of lake sturgeon spawning in nearshore lake habitats. In Lake Michigan, spawning activity has been reported

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on a reef near the St. Joseph River and near Ludington, Michigan, along the eastern shore of the lake. In Burt Lake, a large lake in the northern Lower Peninsula of Michigan, lake sturgeon spawning activity has been reported along the eastern shore of the lake in relatively shallow water with rocky substrate. In instances where lake spawning has been reported, the habitat characteristics are similar and consist of an active wave zone along the eastern shoreline with gravel or cobble substrates. Whether spawning in lake habitats is actually occurring or successful is unknown. The presence of interstitial spaces in the spawning substrate is important for successful lake sturgeon spawning because they are broadcast spawners. Broadcast spawning fish do not construct nests for spawning, nor do they care for the eggs after spawning. Instead, lake sturgeon select a spot for spawning and simply release eggs and sperm into the current and the eggs are fertilized as they drift downstream with the current. Lake sturgeon eggs are negatively buoyant and sink to the substrate shortly after they are released by the female. When there are gaps in the substrate the eggs sink down into those spaces and adhere to the surfaces of the rocks, which protect them from large predators like suckers and crayfish and also shelter them from the main current. If spawning occurs where there are no gaps for the eggs to sink into, the eggs are more vulnerable to predators and can also be dislodged by the current and carried downstream to habitats that are unfavorable for incubation. Lake sturgeon eggs develop rapidly and hatching occurs in as few as five days after spawning (Kempinger 1988). At hatching, lake sturgeon are considered larvae because they are not fully developed and still have a large yolk sac that sustains them for another several days. They remain in the spaces in the substrate until the yolk sac is used up and they are ready to begin feeding. This stage lasts about seven days and ends when the larval lake sturgeon swim up out of the substrate and drift downstream to nursery habitat (Kempinger 1998; Auer and Baker 2002; Smith and King 2005). Little is known of the habitat requirements of lake sturgeon for the first few weeks after they hatch. They are less than 2.5 cm (1 in) long and darkly colored when they hatch and drift downstream at night, probably to avoid sight-feeding predators, and do not drift during daylight hours. Larval lake sturgeon drift has been recorded up to 64 km (40 miles) downstream from the spawning habitat (Auer and Baker 2002), but the daytime habitats used by larvae as they disperse downstream from the spawning site have not been documented. Eventually they stop drifting and begin feeding. By midsummer the fish are sandy-colored with dark mottling and blend in well with the sand and mixed sand/gravel substrates in the rivers where they feed and continue to grow. In the Sturgeon River of Michigan researchers found newly hatched young on gravel and mixed sand/gravel substrates but never on pure sand substrates (Holtgren and Auer 2004), while in the Peshtigo River, Wisconsin researchers studying young

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lake sturgeon habitat use always found newly hatched young on sand substrate (Benson et al. 2005). After the first summer and for the first few years of life, small juvenile lake sturgeon habitat use is somewhat more restricted than that of adults and larger juveniles. Many young remain in or near their river of origin, feeding and residing in and near the river mouth area until they are about five years old, at which time they begin to disperse away from the river mouth (Schram, Lindgren, and Evrard 1999). This tendency for lake sturgeon to remain in and near river mouths may partially explain why some populations are not rebounding. Many river mouths around the Great Lakes have been extensively altered by industrialization and shipping. The construction of piers and the channelization and dredging of river mouths to provide deep shipping channels have drastically altered river mouth habitats and eliminated productive channel-margin wetland habitats. These altered river mouth habitats may be unsuitable for young lake sturgeon survival. As previously mentioned, lake sturgeon habitat use in the Great Lakes is dictated by their feeding habits and the distribution of their prey. With their mouth positioned on the bottom surface of the head, lake sturgeon are adapted for feeding on benthic organisms, those that live directly on or in the substrate of lakes and rivers. The shallow waters with soft-bottom substrates of lakes and large rivers are the most productive habitats for the benthic organisms that these fish rely on for survival. In a study of habitat use in Black Lake, Michigan, lake sturgeon were commonly found over sand or muck, organic substrates. The most common benthic organisms in Black Lake were mayfly larvae, chironomid midge larvae, and crayfish, and these were most abundant in areas of sand and muck substrates (Hay-Chmielewski 1987).

Impacts of Habitat Change on Lake Sturgeon There are many factors that led to the decline of lake sturgeon in the late 1800s. Overharvest was probably the primary cause of the decline, but changes to Great Lakes habitat, particularly river habitat, also played a role in the demise of lake sturgeon (Houston 1987). For example, the entire Great Lakes region was extensively logged to produce lumber. This logging resulted in tremendous soil erosion as the vegetation that held soils in place was destroyed. As the region was logged, many of the rivers that empty into the Great Lakes and that lake sturgeon depended on for reproduction were used to carry logs to sawmills typically located near the mouths of rivers. Log drives caused additional erosion by digging into riverbanks and substrate that dramatically altered fish habitats. The erosion of sediments into the rivers filled in deep holes and reduced the amount

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Figures 1a and 1b. The clear-cut logging (above) and log drives (below) that occurred throughout the Great Lakes region resulted in tremendous erosion. The eroded sediments eventually ended up in rivers and caused marked changes to river habitats.

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of resting habitat that lake sturgeon relied on as they ascended rivers in the spring spawning run. Sediments also likely altered the spawning habitats in rivers by filling in the interstitial gaps that lake sturgeon eggs depended on for successful incubation. The widespread logging undoubtedly raised the temperatures of rivers because the canopy of trees that shaded river channels was removed and allowed more direct sunlight to hit the river surface and warm the water. The habitat change that has probably had the greatest long-term impact on lake sturgeon populations occurred following the peak of the logging era of 1870–1890. As settlement of the Great Lakes region expanded, many rivers that these fish used for spawning were dammed. Sturgeon species in general are noted for their long upriver migrations to find suitable spawning habitat (Auer 1996b). In the case of lake sturgeon these migrations may exceed 240 km (150 miles) (Kempinger 1988). The construction of dams, primarily hydropower dams that produce electricity, has eliminated access to habitat in most rivers where lake sturgeon previously spawned, blocking migrations and flooding the most suitable spawning habitat under the reservoirs created by the dams. In most cases the dams constructed on tributaries to the Great Lakes were built on the first high-gradient river reach upstream from the lake. These highgradient (defined by the drop in elevation over distance) river reaches are the most suitable for hydropower dam construction because they provide the highest hydraulic head and therefore are optimal sites for hydroelectricity generation. Hydropower dams were typically built at the base of high-gradient reaches, resulting in flooding of the river upstream of the dam. Unfortunately for lake sturgeon (and many other species), these high-gradient river reaches are the habitats needed for spawning. Dams in rivers throughout the Great Lakes region prevent access to the most critical habitat lake sturgeon need to complete their life cycle. The Menominee River forms the border between northeast Wisconsin and Michigan’s Upper Peninsula and is a good example of a river that once supported a large spawning run of lake sturgeon but has been severely impacted by dam construction. Prior to dam construction, lake sturgeon (as well as walleye, lake whitefish, lake trout, and other fish species) would migrate up the river in search of suitable spawning habitat. Lake sturgeon historically could migrate as far as Sturgeon Falls near Iron Mountain, Michigan, a distance of approximately 161 km (100 miles) (Thuemler 1985). However, the construction of six dams has fragmented the Menominee River system, and the first dam upstream from Lake Michigan is a mere 4 km (2.5 miles) from the lake. This dam prevents fish from Lake Michigan accessing and using more than 156 km (97 miles) of previously used river spawning and rearing habitat. Barriers to migration are common to most of the rivers historically used by spawning lake sturgeon across the Great Lakes (Holey et al. 2000). In Michigan alone, 90 percent of the largest rivers that feed the Great Lakes and that were historically used by

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Figure 2. Dam on Menominee River that prevents Lake Michigan lake sturgeon from migrating upstream. This dam is only 2.5 miles upstream from Lake Michigan and prevents access to more than 100 miles of previously utilized spawning and juvenile rearing habitat in the Menominee River. (Photo by D. Traynor, Michigan DNR.)

spawning lake sturgeon have dams that prevent use of historic spawning sites (HayChmielewski and Whelan 1997). It should not come as a surprise that lake sturgeon populations have not rebounded from the dramatic declines that occurred in the late 1800s because the fish do not have access to the vitally important habitats they need to reproduce. The detrimental impact of dam building and other habitat changes on lake sturgeon is perhaps best understood if we think of them as needing a large home range. Because lake sturgeon travel great distances in the open waters of the Great Lakes and up rivers for spawning, it should be rather obvious that the species needs large expanses of unimpeded habitat that could be considered their home range. However, few people seem to consider that large fish have these needs. All species of sturgeon travel over lake and river bottoms just as terrestrial organisms travel over field and forest. It is speculated that sturgeons move along habitual paths or corridors on lake bottoms and between lake systems such as exist in the Great

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Lakes. Some fish species, and sturgeon specifically, migrate to avoid unfavorable conditions, to increase feeding opportunities, and to improve chances of finding a mate for spawning (Northcote 1978; Tsyplakov 1978; McKeown 1984). When migrating to spawn, lake sturgeon are known to return to historic spawning grounds and natal spawning beds (Auer 1999; Lyons and Kempinger 1992). Such homing behavior in fish is thought to help develop population-specific adaptations to the habitat (Leggett 1977) and can over time create distinct stocks. For instance, in the Volga River, Russia, eggs of three species of sturgeon differ in membrane strength and resistance to rupture from water pressure (a factor of depth and velocity of water movement) (Nikolsky 1963). The largest sturgeon of the three, the beluga sturgeon, Huso huso, spawned furthest upstream and had the most resistant egg membranes, which would be beneficial in the higher water velocities encountered in upstream reaches. Such adaptations may improve survival of eggs in environments of differing harshness. There are 27 species of sturgeon known worldwide and these are categorized into four genera in the family Acipenseridae: Huso (2 sps.), Acipenser (19 sps.), Scaphirhynchus (3 sps.), and Pseudoscaphirhynchus (3 sps.) (Birstein 1993). Of all the sturgeon species known, only seven in the Acipenser and Scaphirhynchus genera are found in North America All sturgeons migrate in freshwater rivers to spawn, spawn in fast-flowing water, and are slow-growing, late-maturing fishes. All species take from 10 to 20 years to reach sexual maturity and most spawn intermittently. Three species of Acipenser are known to live their entire lives in freshwater. They include the Baikal sturgeon, A. baeri baicalensis (a subspecies of the Siberian sturgeon, A. baeri), which lives in Lake Baikal, Russia, and spawns in the tributaries of that system; the Yangtze sturgeon, A. dabryanus, from the lower Yangtze River, China; and A. fulvescens, the lake sturgeon known from three large drainage basins: the Great Lakes, Hudson Bay and the Mississippi River. The Yangtze sturgeon is considered very close to extinction (Doroshov and Binkowski 1985) particularly due to the construction of the Three Gorges dam (Wei et al. 1997). Of these three species, A. baeri and A. fulvescens occupy the coldest geographic ranges of any sturgeons, with A. baeri found throughout the major river systems of Siberia (Artyukhin 1995). In organisms other than fish, a relationship between body size and range is usually positive (Gaston 1990). There is a positive relationship between body size and range for several fish species (McAllister et al. 1986), and a similar relationship between body size as mass (kg) and migration distance for birds, mammals, walking insects, and some fish (salmon) (Peters 1983). Mean age at first spawning as reflected in body size and distance of upriver migration for Atlantic salmon also shows a positive correlation (Schaffer and Elson 1975). A similar relationship between body size and spawning migration distance is

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Figure 3. Relationship of average adult total length to river spawning migration distance for eight species of sturgeon. With lake sturgeon limited to 200 km, r 2 = 0.785; with lake sturgeon having maximum distance of 750 km (indicated by *), r 2 = 0.874 (adapted from Auer 1996b).

found in sturgeons (Auer 1996b). The body size as average total length of mature fish, and the maximum river spawning migration distance determined from values reported in the literature for historic or current spawning areas for sturgeons, show a positive relationship. There is no record of possible total migration distance for lake sturgeon; because dams or other river barriers have long impacted many populations, such a distance for this species remains unknown. Similar-size sturgeons migrate farther in rivers than historical data show lake sturgeon migrating. Comparing range and size data for all sturgeon species to those of an average adult lake sturgeon (about 145 cm TL, or 57 inches) it appears probable that this species could be capable of migrating distances of 1,000 to 1,800 km (620 miles). Current conditions to date show that lake sturgeon migrate shorter distances in rivers; relatively short segments of tributaries with natural barriers support lake sturgeon populations in the Great Lakes. In the Bad River, Wisconsin lake sturgeon migrate 32 km (20 miles) to spawn near natural falls and steep rapids (Shively and Kmiecik 1989). In the Grand River, Michigan lake sturgeon approach an impassable set of rapids 64 km (40 miles) upstream, while in the Sturgeon River of upper

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Michigan, lake sturgeon migrate 69 km (43 miles) to rapids about 1 km below a small hydroelectric facility (Auer 1996a). These data indicate that the lake sturgeon, which live exclusively in freshwater, appear to combine river and lake distances in their spawning migrations. This is shown by migrations where lake sturgeon travel 228 km (141 miles) to spawn in the Lake Winnebago, Wisconsin, system, and they move 200 km (125 miles) within the Wolf River and can add 27 km (17 miles) as they cross Lake Winnebago (Lyons and Kempinger 1992). Auer (1999) found telemetry tracked postspawning lake sturgeon to leave the Sturgeon River, Michigan spawning habitat, 69 km (or 43 miles) upstream from the lake, and move distances of 265 km (165 miles) from the river mouth into Lake Superior. Adult lake sturgeon movements in the St. Lawrence River in Canada showed sturgeon moving distances of 138 to 225 km (86 to 140 miles) (Dumont et al. 1987; Fortin et al. 1993). Combining river and lake migration distances increases the closeness of the match of size to migration distance that is seen in other species. Another reason open corridors for migration are so important for fishes is that spawning migrations are the time when hormones trigger the eggs and sperm to reach final development (McKeown 1984). Some species of female sturgeons, if prevented from reaching spawning areas because of barrier dams or manipulated flows, do not spawn and reabsorb eggs, or their spawned eggs show reduced survival (Artyukhin, Sukhoparova, and Fimukhina 1978; Veshchev and Novikova 1983, 1988). Scuba divers report sturgeon eggs, spawned below the Shawano Dam on the Wolf River in Wisconsin to be in masses as much as 15 cm (6 inches) thick (Kempinger 1988). Eggs in such masses are considerably more vulnerable to disease and predation. In the few remaining rivers where lake sturgeon can migrate onto natal spawning beds in natural rapids, with natural turbulent flows, their eggs adhere to the undersides of clean rock or fall into rock crevices, and thick masses of eggs are not observed. Years of research and monitoring have shown that lake sturgeon spawn in sections of rapids in the upper reaches of river systems and then return to feeding and wintering regions in lakes and river mouth reaches. Delta areas at the mouths of rivers are usually surrounded by rich organic sloughs or wetlands that are believed to provide food organisms and shelter from predators, necessary for the growth and survival of newly hatched and juvenile individuals. Identified sites for rehabilitation and restocking of lake sturgeon should consider allocation to a minimum of 250–300 km (155 to 185 miles) of unrestricted movement, and migratory distances of 750–1,000 km (465 to 620 miles) should not be considered unusual for this species. For lake sturgeon, protecting open tributary corridors between river and lake environments or providing fish passage must be given priority, combined with safe outmigration pathways and areas for feeding and wintering of all life stages.

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Other changes to Great Lakes habitat that may have affected lake sturgeon are associated with the logging era and industrial development. During the logging era, when sawmills were operating on many rivers, large quantities of sawdust were dumped directly into the rivers. This sawdust sank and in many places formed thick mats on the substrate of the river mouth and nearby Great Lakes bottomlands (Harkness and Dymond 1961). The mats of sawdust effectively eliminated large feeding areas vital to young sturgeon as they grew and developed in the lower reaches of rivers. Industrial pollution has also been widespread in the Great Lakes and contamination continues to plague some areas of the lakes. Lake sturgeon that inhabit areas of the lakes affected by industrial pollution have been shown to carry high body burdens of contaminants, including mercury, PCBs, dioxin, and others (Michigan DNRE unpublished data). The impacts of these contaminants on lake sturgeon are unknown, but reproductive impairment is possible. Exotic species have also changed habitat characteristics in the Great Lakes. Most notable are the zebra and quagga mussels. Zebra mussels are native to the European and Russian Black, Azov, and Caspian seas, and quagga mussels are native to the Dnieper River and Ponto-Caspian Sea. Both of these species were probably introduced to the Great Lakes in the ballast water of ocean freighters. Zebra and quagga mussels are filter feeders and have widely colonized most of the available habitats in the lower lakes; to date Lake Superior remains relatively unaffected. The exotic mussels have dramatically increased water clarity by filtering out nutrients, phytoplankton, and zooplankton from the water column. The long-term impact of these changes on lake sturgeon is unknown, but the establishment of zebra and quagga mussels is believed to be responsible for the near-disappearance of Diporeia from the lower lakes (Nalepa, Fanslow, and Lang 2009). Diporeia is a small shrimplike animal and an important link in the Great Lakes food chain. Although lake sturgeon have been known to eat zebra and quagga mussels, there is nothing to suggest that the establishment of the exotic mussels and subsequent ecosystem changes are a good thing for lake sturgeon or any other fishes of the Great Lakes.

Foods and Feeding As a general rule, as fish increase in size, the food items they consume also tend to increase in size. Therefore, as the largest fish found in the Great Lakes, lake sturgeon might be expected to eat the largest food items. However, lake sturgeon are an exception to the rule, and the foods they consume are among the smallest items available to them, such as insect larvae, small snails and clams, and crayfish. One of the unique aspects of lake sturgeon form is the position of the mouth.

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Most fish species found in the Great Lakes have a mouth positioned at or near the front of the head, in front of the eyes, and food is located by a combination of sight, sound, and smell. In contrast, the lake sturgeon mouth is located on the flat underside of the head directly beneath the eyes and is preceded by four sensory barbells. The mouth of the lake sturgeon is also unique in that it is protrusible, that is, can be extended out like a vacuum hose. Lake sturgeon locate food by swimming along the bottom of lakes and rivers and by dragging their sensitive barbels over the substrate. When the barbels encounter something edible, the mouth is quickly extended to the substrate and the prey is suctioned into the mouth. The prey can be detected by feel, taste, or the weak electric signals that are emitted by all living organisms. These weak electric signals are detected by sensory pits that are also located on the underside of the head (Boglione et al. 1999). The type of food eaten is directly determined by the lake sturgeon’s unique form and feeding habits, and their feeding habits change little as the fish grow throughout their lives. When newly hatched, they begin feeding on small zooplankton, and the smallest insect larvae available to them (Kempinger 1988). As the young lake sturgeon grow, the size of the insect larvae and zooplankton they consume also increases, but by the end of the first summer of life, when the young are 15–20 cm (6–8 inches) long, their diet is very similar to that of the adults that are 1.8 m (6 feet) long. As the young grow, they may add crayfish and clams to their diet when they become large enough to prey on them (Hay-Chmielewski 1987) but they also continue to eat small insect larvae and worms as they grow to very large size. As lake sturgeon feed, they invariably ingest some of the substrate (sand, small stones, etc.) along with their prey. However, these fish are able to work prey around in their mouth and begin the process of mechanically breaking down the ingested material. At the same time they are able to sort out most of the undesired materials like sand and gravel and expel them prior to swallowing. Lake sturgeon do not have teeth, but they do have strong cartilaginous plates in the mouth cavity that can be ground together to reduce prey size. In addition, these fish have a muscular gizzardlike stomach that can further break up ingested prey. By grinding food in the mouth and in the gizzard, lake sturgeon are able to crush small clams, snails, crayfish, and other hard foods to make them easier to digest. After the prey is mechanically broken down in the mouth and stomach, digestion and absorption of nutrients continues in the spiral valve intestine. The spiral valve intestine of lake sturgeon is similar to that found in sharks and is an adaptation that increases the surface area of the intestine and thus increases the absorption of nutrients as food is digested. In the spiral valve, a flap of tissue spirals down the length of the interior intestine wall much like a spiral staircase might follow

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the interior wall of a tower. This flap of tissue and the inner lining of the intestine absorb nutrients from digested food as the food passes down from the stomach. Lake sturgeon apparently do not use sight at all to find food but instead rely entirely on their sensitive barbels and sensory receptors alone. Lake sturgeon are classified as opportunistic predators that will eat anything they encounter. Because their barbels have sensory capabilities to help locate food, the fish avoid areas where aquatic plants are growing. Rooted aquatic plants probably interfere with the food search, and so lake sturgeon cruise areas of bottom that allow unobstructed foraging. Although there have been only a few rigorous scientific investigations of lake sturgeon diet, they all indicate that lake sturgeon readily eat what is available and in approximate proportion to its availability in the environment. For example, in Black Lake, Michigan, adult lake sturgeon diet in 1986 included large numbers of Hexagenia sp. mayfly nymphs and Chironomidae sp. midge larvae (Hay-Chmielewski 1987). These two taxa represented 77.3 percent of the stomach contents of fish sampled. At the same time that lake sturgeon diet was being analyzed, samples of the substrate in Black Lake indicated that these two organisms were the most abundant food available. Although mayfly and midge larvae were the most abundant food items in the environment and in the diet, crayfish were also apparently important food for Black Lake sturgeon. Crayfish, although only 2.3 percent of the stomach contents by number, represented 66 percent of the stomach contents by weight. Similar diets have been found for other lake sturgeon populations (Choudhury, Bruch, and Dick 1996; Beamish, Noakes, and Rossiter 1998; Werner and Hayes 2004). The ability of lake sturgeon to prey on a wide variety of food organisms has also been demonstrated by the few studies that have been conducted. As many as 42 different types of animals have been found in lake sturgeon stomachs from a single water body (Werner and Hayes 2004). Most of the prey eaten by lake sturgeon are invertebrates (insects, crustaceans, and worms), primarily mayfly and midge larvae, but the importance of particular prey varies by water body. In the St. Lawrence River, mayfly and midge larvae are eaten by lake sturgeon but the prey most frequently consumed is amphipods. The lake sturgeon diet may change to take advantage of prey that is seasonally abundant, as sturgeon have been known to feed on dead and dying gizzard shad during winter in Lake Winnebago, Wisconsin. Lake sturgeon have also been shown to adapt their feeding patterns to changes in prey availability. Zebra and quagga mussels are exotic mussels now widely established in the Great Lakes and are causing dramatic changes in ecosystem structure and function (Nalepa, Fanslow, and Lang 2009). Lake sturgeon appear to be taking advantage of this new food resource and are eating these exotic mussels. Although it may seem beneficial for lake sturgeon to be feeding on the newly established mussels, the ecosystem disruption that has been caused by exotic species may be

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causing increased mortality in lake sturgeon and other species. Botulism has been confirmed as the cause of bird deaths in some places in the Great Lakes (Brand et al. 1988) and is also suspected of causing occasional die-offs of lake sturgeon in some parts of the Great Lakes (Perez-Fuentetaja, Lee, and Clapsadl undated) . The exact mechanism of botulism poisoning is not known, but is believed to be linked to the expansion of zebra and quagga mussels as well as the establishment of round and tubenose gobies, exotic fishes that have also become widespread in the Great Lakes in recent years. Prior to scientific studies of the lake sturgeon diet there were a number of misconceptions about the role they played in the ecosystem. One of the early misconceptions was that lake sturgeon preyed on the eggs of more desirable fish species like walleye, trout, and salmon, and because of this, lake sturgeon were considered harmful to these species. However, in diet studies, the eggs of these and other game fishes have not been found in lake sturgeon stomachs. Furthermore, lake sturgeon are typically not found in the habitats that are used by walleye, trout, or salmon at the time these species are spawning, and thus it is unlikely that they would have the opportunity to prey on eggs of these species. Sturgeon are not considered a threat to such species and their reproductive success. Interestingly, the only known instances of lake sturgeon preying on relatively large quantities of fish eggs have occurred when they have consumed the eggs of their own species during spawning. In a few instances in the Sturgeon River, Michigan spawning males have been found to have lake sturgeon eggs in their feces when the fish have been captured for scientific study (Auer and Baker, personal observation). The same has been observed in the Black River, Michigan, and Wolf River, Wisconsin. Lake sturgeon feed on the same prey that other fish such as white suckers and redhorse consume. Therefore, there is at least the opportunity for competition for prey. However, there have not been studies that document harm to either lake sturgeon or other species from competition for food resources. Because these species are native to the Great Lakes they have coexisted for thousands of years and are able to occupy similar habitats and utilize similar food resources. During the course of feeding sturgeon do occasionally make mistakes. One of the most unusual items eaten by a sturgeon was reported from Germany and actually helped to solve a missing person’s case. In an account detailed by Harkness and Dymond (1961) a German newspaper reported a large sturgeon was caught in 1927 that had in its stomach an Iron Cross, a German military honor medal, with a soldier’s name on it. The soldier identified from the medal had been wounded and permanently disabled during fighting in World War I. The soldier had become despondent and had disappeared in 1920. Based on the discovery of the medal in the stomach of the sturgeon, the German police concluded that the soldier

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had committed suicide by jumping in the river. Other unusual items found in the stomachs of sturgeon included unopened sardine cans, a vanity case with a powder puff, cigarettes, coins, and fishhooks.

Conclusion The precarious status of lake sturgeon in the Great Lakes is a concern of many natural resource management agencies around the lakes, and management agencies are actively pursuing lake sturgeon reintroduction and rehabilitation across the watershed. It is recognized that the continued low abundance of the species is a symptom of historic overharvest as well as the drastic habitat changes that have taken place. If lake sturgeon management efforts lead to increased abundance and the lake sturgeon again becomes a prominent species in the Great Lakes, it will be an indication that the Great Lakes ecosystem has regained some of its historic health and function.

REFERENCES

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Artyukhin, E. N. 1995. On biogeography and relationships within the genus Acipenser. Sturgeon Quarterly 3(2): 6–7. Artyukhin, E. N., A. D. Sukhoparova, and L. G. Fimukhina. 1978. The gonads of the sturgeon, Acipenser guldenstadt, in the zone below the dam of the Volograd water engineering system. Journal of Ichthyology 18:912–923. Auer, Nancy A. 1996a. Response of spawning lake sturgeon to change in hydroelectric facility operation. Transactions of the American Fisheries Society 125(1): 66–77. —. 1996b. Importance of habitat and migration to sturgeons with emphasis on lake sturgeon. Canadian Journal of Fisheries and Aquatic Sciences 553(suppl. 1): 152–160. —. 1999. Population characteristics and movements of lake sturgeon in the Sturgeon River and Lake Superior. Journal of Great Lakes Research 25:282–293. Auer, N. A., and E. A. Baker. 2002. Duration and drift of larval lake sturgeon in the Sturgeon River, Michigan. Journal of Applied Ichthyology 18:557–564. Baldwin, N. S., R. W. Saalfeld, M. A. Ross, and J. J. Buettner. 1979. Commercial fish production in the Great Lakes 1867–1977. Technical Report No. 3, Great Lakes Fishery Commission. Benson, A. C., T. M. Sutton, R. F. Elliott, and T. G. Meronek. 2005. Seasonal movement patterns and habitat preferences of age-0 lake sturgeon in the lower Peshtigo River, Wisconsin. Transactions of the American Fisheries Society 134:1400–1409. Beamish, F. W. H., D. L. G. Noakes, and A. Rossiter. 1998. Feeding ecology of juvenile lake sturgeon, Acipenser fulvescens, in northern Ontario. Canadian Field-Naturalist 112:459–468. Birstein, V. J. 1993. Sturgeons and paddlefishes: Threatened fishes in need of conservation.

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Conservation Biology 7:773–787. Boglione, C., P. Bronzi, E. Cataldi, S. Serra, F. Gagliardi, and S. Cataudella. 1999. Aspects of early development in the Adriatic sturgeon Acipenser naccarii. Journal of Applied Ichthyology 15:207–213. Bott, K. 2006. Genetic analyses of dispersal, harvest mortality, and recruitment for remnant populations of lake sturgeon, Acipenser fulvescens, in open-water and riverine habitats of Lake Michigan. M.S. thesis, Michigan State University. Brand, C. J., S. M. Schmitt, R. M. Duncan, and T. M. Cooley. 1988. An outbreak of type E botulism among common loons (Gavia immer) in Michigan’s Upper Peninsula. Journal of Wildlife Diseases 24:471–476. Chiotti, J. A., J. M. Holtgren, N. A. Auer, and S. A. Ogren. 2008. Lake sturgeon spawning habitat in the Manistee River, Michigan. North American Journal of Fisheries Management 28:1009–1019. Choudhury, A., R. Bruch, and T. A. Dick. 1996. Helminths and food habits of lake sturgeon Acipenser fulvescens from the Lake Winnebago system, Wisconsin. American Midland Naturalist 135:274–282. Diana, J. S., P. W. Webb, and T. Essington. 2003. Growth and appetite of juvenile lake sturgeon Acipenser fulvescens. Michigan Department of Natural Resources, Fisheries Research Report 2063. Doroshov, S. I., and F. P. Binkowski. 1985. Epilogue: A perspective in sturgeon culture. In: North American sturgeons: Biology and aquaculture potential. F. P. Binkowski and S. I. Doroshov, eds. Dr. W. Junk Publishers.

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Dumont, P., R. Fortin, G. Desjardins, and M. Bernard. 1987. Biology and exploitation of lake sturgeon in the Quebec waters of the Saint-Laurent River. In: Proceedings of a workshop on the lake sturgeon (Acipenser fulvescens). C. H. Olver, ed. Ontario Fisheries Technical Report Series No. 23. Fortin, R., J. R. Mongeau, G. Desjardins, and P. Dumont. 1993. Movements and biological statistics of lake sturgeon (Acipenser fulvescens) populations from the St. Lawrence and Ottawa River system. Canadian Journal of Zoology 71:638–650. Gaston, K. J. 1990. Patterns in the geographical ranges of species. Biological Reviews 65:105–129. Harkness, W. J. K., and J. R. Dymond. 1961. The lake sturgeon. Ontario Department of Lands and Forests. Hay-Chmielewski, E. M. 1987. Habitat preferences and movement patterns of the lake sturgeon (Acipenser fulvescens) in Black Lake Michigan. Michigan Department of Natural Resources, Fisheries Research Report 1949. Hay-Chmielewski, L., and G. Whelan. 1997. Lake sturgeon rehabilitation strategy. Michigan Department of Natural Resources Fisheries Division Special Report No. 18. Holey, M. E., E. A. Baker, T. F. Thuemler, and R. F. Elliott. 2000. Research and assessment needs to restore lake sturgeon in the Great Lakes. Great Lakes Fishery Trust, Workshop Results. Holtgren, J. M., and N. A. Auer. 2004. Movement and habitat of juvenile lake sturgeon (Acipenser fulvescens) in the Sturgeon River / Portage Lake system, Michigan. Journal of Freshwater Ecology 19:419–432. Houston, J. J. 1987. Status of the lake sturgeon, Acipenser fulvescens, in Canada. Canadian FieldNaturalist 101:171–185. Kempinger, J. J. 1988. Spawning and early life history of lake sturgeon in the Lake Winnebago

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system, Wisconsin. In: Eleventh annual larval fish conference. R. D. Hoyt, ed. American Fisheries Society Symposium 5. American Fisheries Society. LaHaye, M., A. Branchaud, M. Gendron, R. Verdon, and R. Fortin. 1992. Reproduction, early life history, and characteristics of spawning grounds of the lake sturgeon (Acipenser fulvescens) in Des Prairies and L’Assomption rivers, near Montreal, Quebec. Canadian Journal of Zoology 70:1681–1689. Leggett, W. C. 1977. The ecology of fish migrations. Annual Review of Ecological Systems 8:285–308. Lyons, J., and J. J. Kempinger. 1992. Movements of adult lake sturgeon in the Lake Winnebago system. Wisconsin Department of Natural Resources Research Publication RS-156- 92. McAllister, D. E., S. P. Platania, F. W. Schueler, M. E. Baldwin, and D. S. Lee. 1986. Ichthyofaunal patterns on a geographic grid. In: The zoogeography of North American freshwater fishes. C. H. Hocutt and E. O. Wiley, eds. J. Wiley and Sons. McKeown, B. A. 1984. Fish migration. Timber Press. Nalepa, T. F., D. L. Fanslow, and G. A. Lang. 2009. Transformation of the offshore benthic community in Lake Michigan: Recent shift from the native amphipod Diporeia spp. to the invasive mussel Dreissena rostriformin bugensis. Freshwater Biology 54:466–479. Nikolsky, G. V. 1963. The ecology of fishes. Academic Press. Northcote, T. G. 1978. Migratory strategies and production of freshwater fishes. In: Ecology of freshwater fish production. S. Gerking, ed. John Wiley and Sons. Perez-Fuentetaja, A., T. Lee, and M. Clapsadl. Undated. Botulism E in Lake Erie: Ecology and lower food web transfer. Final Report to U.S. Fish and Wildlife Service, Grant Award 26462. Peters, R. H. 1983. The ecological implications of body size. Cambridge University Press.

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Schaffer, W. M., and P. F. Elson. 1975. The adaptive significance of variations in life history among local populations of Atlantic salmon in North America. Ecology 56(3): 577–590. Schram, S. T., J. Lindgren, and L. M. Evrard. 1999. Reintroduction of lake sturgeon in the St. Louis River, western Lake Superior. North American Journal of Fisheries Management 19:815–823. Scott, W. B., and E. J. Crossman. 1973. Freshwater fishes of Canada. Fisheries Research Board of Canada Bulletin 184, Ottawa, Canada. Shively, J. D., and N. Kmiecik. 1989. Inland fisheries enhancement activities within the ceded territory of Wisconsin during 1988. Administrative Report 89-1. Great Lakes Indian Fish and Wildlife Commission. Smith, K. M., and D. K. King. 2005. Dynamics and extent of larval lake sturgeon Acipenser fulvescens drift in the Upper Black River, Michigan. Journal of Applied Ichthyology 21:161–168. Threader, R. W., R. J. Pope, and P. R. H. Schaap. 1998. Development of a habitat suitability index model for lake sturgeon (Acipenser fulvescens). Ontario Hydro Report Number H-07015.01-0012. Thuemler, T. F. 1985. The lake sturgeon, Acipenser fulvescens, in the Menominee River, WisconsinMichigan. Environmental Biology of Fishes 14:73–78. Tsyplakov, E. P. 1978. Migrations and distribution of the sterlet, Acipenser ruthenus, in Kuybyshev reservoir. Journal of Ichthyology 18:905–912. Veshchev, P. V., and A. S. Novikova. 1983. Reproduction of the stellate sturgeon, Acipenser

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stellatus (Acipenseridae), under regulated flow conditions in the Volga River. Journal of Ichthyology 23(1): 42–50. —. 1988. Reproduction of sevryuga, Acipenser stellatus, in the lower Volga. Journal of Ichthyology 28(1): 39–47. Wehrly, K. W. 1995. The effect of water temperature on the growth of juvenile lake sturgeon Acipenser fulvescens. Michigan Department of Natural Resources, Fisheries Research Report 2004. Wei, Q., F. Ke, J. Zhang, P. Zhuang, J. Luo, R. Zhou, and W. Yang. 1997. Biology, fisheries, and conservation of sturgeon and paddlefish in China. Environmental Biology of Fishes 48:241–255.

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Werner, R. G., and J. Hayes. 2004. Contributing factors in habitat selection by lake sturgeon (Acipenser fulvescens). Final report submitted to Environmental Protection Agency Great Lakes National Program Office.

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Plate 1. Scutes on juvenile sturgeon are razor sharp and run along the back, sides, and bottom edge on each side of the fish. (N. Auer.)

Plate 2a. The mouth of a sturgeon has no teeth. (N. Auer.)

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Plate 2b. Sturgeon locate food using the four sensitive barbels located just in front of the mouth as they swim over the lake bottom. (N. Auer.)

Plate 3. Eggs of lake sturgeon, which adhere to clean rocks in rapidly flowing, oxygen-rich water. (N. Auer.)

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Plate 4. Young juveniles showing dark saddle markings on back and rich copper color. (N. Auer.)

Plate 5. Lake sturgeon spawning rapids in the Sturgeon River, Michigan. (Photo by K. Koval, Michigan DNR.)

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Plate 6. Juvenile lake sturgeon shown with characteristic mottling that helps the fish blend in against sandy and mixed sand/gravel substrates. (Photo by E. Baker, Michigan DNR.)

30

40

50

60

70

80

90

100

110

120

Plate 7. Mitochondrial DNA sequences from two lake sturgeon individuals. The individual on the top is from the Detroit River and the individual on the bottom is from the Lower Niagara River. Arrows indicate polymorphisms, or places in the sequence where the two individuals differ.

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125,000

274 bp

Dye Signal

100,000 141 bp 75,000

50,000

218 bp 226 bp

25,000

0 0

50

100

150

200 Size (nt)

250

300

350

400

Plate 8. Microsatellite data from a single lake sturgeon individual from the Namakan River, Ontario.

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3

1

4

2 5 2

Plate 9. Stock structure of lake sturgeon in the Great Lakes basin. Each cube represents a spawning population. The different colors represent the different lake basins: red, Lake Superior; green, Lake Michigan; blue, Lake Huron; light purple, Lake Ontario; dark purple, St. Lawrence River system; orange, Lake Champlain. Circled and numbered regions are described in more detail in the text.

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Plate 10 (opposite). Major habitats of the lake sturgeon populations in the Quebec part of the St. Lawrence River system. Water discharge averages 7,500 m3/s in front of Cornwall, 8,000 m3/s in front of Montreal, and 12,600 m3/s in front of Quebec city. The average discharge of the Ottawa River at Carillon dam, at the head of Lac des Deux Montagnes (158 km2) is 2,000 m3/s. Water depth is generally low ( 100 kg) like those reported by Vladykov (1955) or Harkness and Dymond (1961) are now very rare or completely absent in the lower St. Lawrence River, fish exceeding 40 kg and 1.8 m are still observed in the spawning grounds (Fortin, D’Amours, and Thibodeau 2002; Dumont et al. 2011) and in the commercial catch. In the past 25 years, maximum age and weight reported in the commercial harvest were 96 years and 90 kg (Dumont, Fortin, et al. 1987). In the 1910s and 1920s, Quebec sport fishing rules imposed a closure period in June, and it seems that this measure was also applied to commercial fishing. In 1950, this protection moved to the month of May, in 1956, to a mid-May to mid-June period, and in 1967, to the beginning of the ice-over period (generally in December) to the end of April. A bag limit of two sturgeons appeared for the first time in 1971, simultaneously with a return to the mid-May to mid-June closure period. This period was extended in 1984 (mid-April to mid-June) and the bag limit reduced to one fish in 1988. The current period extends from mid-June to the end of October, with a closure period between August 1 and September 14 for the commercial catch in order to reduce incidental mortality during the warmer season. Another factor contributing to the relative stability of the lake sturgeon fishery, at least for the second half of the twentieth century, is the early development, from 1941, and use for management purposes, of scientific knowledge of the species’ biology in the Quebec part of the St. Lawrence and the Ottawa rivers watersheds under the patronage of the Station biologique de Montréal. The first research works were realized in cooperation with commercial fishermen, some of them being qualified as excellent naturalists (figure 3). Within 10 to 20 years, different aspects have been investigated : morphology and systematics (Vladykov and Beaulieu 1946; 1951; Roussow 1955a; Vladykov and Greeley 1963; Magnin 1962), feeding (Fry et al. 1941; Vladykov and Gauthier 1941; Vladykov 1948), movements of juvenile fish (Roussow 1955b; Magnin and Beaulieu 1960), growth (Cuerrier and Roussow 1951; Roussow 1955b; Magnin 1962), sexual maturation, spawning periodicity, and gonad

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development (Cuerrier 1945; Dubreuil and Cuerrier 1950; Roussow 1957, Cuerrier 1966), and commercial fishing (Cuerrier, Fry, and Préfontaine 1946; Roussow 1955b; Cuerrier 1962). With the exception of some concerns about dams’ impacts on sturgeon movements between Lac Saint-François and Lac Saint-Louis (Roussow 1955b), and some details about spawning grounds location in the Saint-François (Cuerrier 1962), Batiscan and Chaudière rivers (Vladykov 1955), little attention was paid to habitat before the work of Mongeau, Leclerc, and Brisebois (1982) in the 1960s and 1970s on the natural restoration of the sturgeon population in Lac des Deux Montagnes following an anoxic period in the early 1950s. A second phase of development of scientific knowledge on lake sturgeon biology and exploitation was initiated in 1981, when an increase in demand for commercial fishing licenses led us to study various aspects of the biology, habitat, and dynamics of this population, to revise the fishery management plan, and to implement various measures of habitat protection and improvement. This work has been done in close collaboration with the Department of Biological Sciences of the Université du Québec à Montréal.

Lake Sturgeon Biology in the Lower St. Lawrence River System

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DISTRIBUTION AND STOCK COMPOSITION

Lake sturgeon occur all along the Quebec part of the St. Lawrence River, where the species now forms two populations. The first one, located in Lac Saint-François, has been gradually separated from the downstream and upstream groups by the construction of the Beauharnois–Les Cèdres (1912–1961) and the Moses-Saunders (1958) hydropower complexes (plate 10). Tagging studies in the 1940s (Roussow 1955b) indicated that lake sturgeon were then able to migrate along the St. Lawrence River, from the limits of the brackish waters up to at least Ash Island, near the outlet of Lake Ontario (figure 4). Downstream Lac Saint-François, over a 350 km stretch from Beauharnois Dam, at the head of Lac Saint-Louis, to the brackish waters downstream of Quebec city (plate 10), lake sturgeon likely form a homogeneous phenotypic and genotypic stock (Guénette, Rassart, and Fortin 1992; Guénette, Fortin, and Rassart 1993). In the freshwaters of the upper estuary, downstream of Lac Saint-Pierre, this population co-occurs with the Atlantic sturgeon (A. oxyrynchus). This lake sturgeon population was not found to be significantly different from the population of Lac des Deux Montagnes, an enlargement of the lower Ottawa River downstream from Carillon Dam. However, the presence of a particular genotype in Lac des Deux Montagnes, and its absence in the rest of the St. Lawrence, support the hypothesis that sturgeon

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Figure 3. Mr. Napoléon Lalumière, a well-known and curious Lac Saint-Louis lake sturgeon commercial fisherman, with a girlfriend circa 1940. Mr. Lalumière also contributed to the identification of a new species of catostomid, the copper redhorse (Moxostoma hubbsi), an endangered species exclusive to southwestern Quebec.

movements between the two water bodies are limited, as shown by the results of tagging studies (Fortin et al. 1993) and morphological comparisons (Guénette, Rassart, and Fortin 1992). Mitochondrial DNA variation also suggests that genetic heterogeneity seemed lower in the St. Lawrence River population than in the James Bay drainage population, likely because this southern population has been more significantly influenced by overfishing and man-made habitat changes over a long period of time (Guénette, Fortin, and Rassart 1993). Small highly fragmented lake sturgeon groups also occur upstream of Lac des Deux Montagnes along the Ottawa River, where, in the last 580 km, nine reaches were gradually separated by dams during the nineteenth and twentieth centuries (Haxton and Findlay 2008).

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Figure 4. Movements of lake sturgeon tagged in the 1940s in the lower St. Lawrence system before the completion of large hydropower facilities downstream (Beauharnois complex, 1912–1961) and upstream (Moses Saunders complex, 1958) Lac SaintFrançois. More than 2,000 lake sturgeon had been tagged during this period. (Reproduced from Roussow 1955b.)

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Recent assessment of lake sturgeon population genetic structure in the Great Lakes basin and Hudson Bay drainage (Welsh et al. 2008), using 27 spawning locations, indicated that, even if the majority of the 25 spawning populations within the system were genetically distinct from each other, the Lac Saint-François population (represented by the Grasse River sample) and the Lower St. Lawrence River population (downstream from Beauharnois Dam, represented by the des Prairies River sample) were grouped with neighboring populations in the upper St. Lawrence River and in Lake Champlain. Before the construction of two dams on the Richelieu River in the 1840s, at Chambly and St-Ours, there was no obstacle to fish passage between Lake Champlain and the St. Lawrence River (Dumont et al. 1997).

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MOVEMENTS

In the Quebec part of the St. Lawrence River, mark-recapture experiments on more than 12,000 lake sturgeon indicate that, with the exception of spawning migrations, which are extensive, movements are restricted (Roussow 1955b; Magnin and Beaulieu 1960; Dumont, Fortin, et al. 1987; Fortin et al. 1993). Most multiple recaptures (up to four) were done near the tagging sites. Globally, juvenile sturgeon were found sedentary, but some movements were observed on a small proportion of them (< 5 percent). Larger sturgeon (> 85 cm), tagged on the des Prairies and l’Assomption rivers spawning grounds, and on pre- and postspawning concentration sites, were recaptured throughout the St. Lawrence River, from Beauharnois to the upper estuary (Dumont, Fortin, et al. 1987; Fortin et al. 1993). In this system, lake sturgeon occur in large numbers in small localized sites, increasing their vulnerability to fishing gear and to any intervention on these local habitats (filling, dredging, toxic outflows, etc.). Some sturgeon seem to form very stable groups; for example, at least on three occasions, pairs of fish tagged simultaneously were recaptured together (Dumont, Fortin, et al. 1987). Larvae drift downstream from the main spawning ground (des Prairies River), and size and age distribution of juveniles in the experimental samples (mostly age two to eight), and of subadults in the commercial harvest samples, suggest that sturgeon are mainly produced in the major spawning grounds, in the upper part of the system. Their offspring would then quickly drift downstream to the upper estuary and later, slowly and gradually colonize the river along a downstream-upstream gradient. Most juvenile sturgeon concentrations are found in the lower part, between the Lac Saint-Pierre archipelago freshwaters and the estuarine brackish waters, near Orléans Island (figure 5). In the commercial catch of the upper estuary (Gentilly), males and females are smaller, lighter, and younger, and only a small proportion of these fishes

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Figure 5. Average size of juvenile lake sturgeon in the 1992–1999 experimental multimesh gillnet catch from Lac Saint-Louis to the upper estuary. (Adapted from Dumont et al. 2000.)

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are maturing (table 1), while upstream in Lac Saint-Louis, they are longer, heavier, and older, and almost half the females are maturing. Intermediate values are generally observed in Lac Saint-Pierre and its archipelago.

GROWTH

Lake sturgeon average back-calculated total lengths at age 5, 10, 15, 20, 25, 30, 35, and 40 years in the lower St. Lawrence River are respectively 530; 770; 977; 1,165; 1,251; 1,338; 1,391; and 1,496 mm (Fortin, Guénette, and Dumont 1992; see also Fortin et al. 1993). Average corresponding weights are 0.8, 2.8, 6.0, 10.7, 13.6, 16.9, 19.2, and 24.4 kg. Condition factor (a function of length and weight that ranges from 0 to 1) increases with size (from .51 to .83 for 500 to 1,500 mm fish) and, at equal size, is higher in the St. Lawrence River than in the Lac des Deux Montagnes (Fortin, Guénette, and Dumont 1992; Guénette, Rassart, and Fortin 1992). In the river, as observed in the whole distribution area, growth rate decreases along a latitudinal gradient (Fortin, Dumont, and Guénette 1996). Compared to other eastern populations (≤80°W), lake sturgeon grow relatively rapidly in the St. Lawrence River; however, growth is faster in Wisconsin inland waters and Lake of the Woods, in the western part of the distribution area (Fortin, Dumont, and Guénette 1996).

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FOOD AND FEEDING

Juvenile lake sturgeon are generalists and opportunistic benthic feeders. In the lower St. Lawrence River, their diet has been found to be highly diversified and composed of at least 75 taxa, of which more than 50 occurred in more than 5 percent of the samples. This probably reflects the high diversity of the benthic fauna of this system, which is also much more productive (~2400 invertebrates/m2 compared for example to 0.1), but the two largest cohorts, those in 1994 and 2002, were associated with high larval production. On the other hand, the smaller-than-average 2003 cohort was associated with the highest larval production measured during the period.

Figure 9d. During this period, larvae production appeared highly and negatively related to the commercial landings in the previous year. In the 1990s, larval production decreased as the previous year’s recorded landings increased, and it rose starting in the year 2000 as the previous year’s landings began to decrease following the gradual application of a progressively lower commercial annual harvest quota (from 200 t in 1999 to 80 t in 2002). These observations suggest that larvae production quickly and positively responded to the implement of a decreasing quota between 2000 and 2002 but also demonstrate that environmental factors can alter the makeup of a cohort at a later stage. (Adapted from Dumont et al. 2011 and Mailhot et al. 2011.)

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lower St. Lawrence River sturgeon population, its habitat and exploitation, it will be important to achieve the following protective measures: • Prevent additional fragmentation of this 350 km stretch of fluvial habitat • Maintain the application to the fishery of conservative restrictions, measures of control, law enforcement, and periodic monitoring • Preserve and improve the quality of the known spawning grounds, feeding habitats, and deep water refuges, whose quality and areas will be reduced along with the St. Lawrence River flow reduction predicted by the climate changes models • Intensify the efforts of reduction of water pollution in the Great Lakes–St. Lawrence River system • Continue to deepen our knowledge of the biology and habitat of this species.

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NOTE

This chapter is dedicated to Réjean Fortin, who died prematurely in 2001. Réjean was an excellent scientist and pedagogue, and a great colleague. His contribution to the knowledge of sturgeon biology will remain pertinent for future decades. We also want to underline the work of J-P. Cuerrier, G. Roussow, V. D. Vladykov, E. Magnin, and J.-R. Mongeau, which still continues to inspire our collective effort to develop a comprehensive approach to the conservation of the St. Lawrence River lake sturgeon population. Finally, we wish to acknowledge the following persons for their involvement over the last 25 years in this long-term objective : L. Aubry, R. Bacon, M. Bernard, P. Bilodeau, A. Blanchard, A. Branchaud, V. Boivin, D. Bourbeau, L. Bouthillier, J. Brisebois, P. Brodeur, Y. Chagnon, S. Clermont, P. Y. Collin, C. Côté, J. D’Amours, M. Damphousse, G. Desjardins, S. Desloges, D. Dolan, B. Dumas, R. Dumas, R. Faucher, D. Fournier, N. Fournier, S. Garceau, D. Goyette, S. Guénette, F. Guilbard, D. Hatin, M. Lafleur, M. La Haye, J. Leclerc, P. Leclerc, M. Léveillé, G. Massé, H. Massé, M. Mingelbier, J. Morin, P. Nilo, G. Ouellet, A. Paquet, Y. Poiré, J. Robitaille, G. Roy, M. Rousseau, S. Thibodeau, G. Trencia, F. Veillette, and R. Verdon. M. Courtemanche provided interesting archaeological information on lake sturgeon use by First Nations people.

REFERENCES

Beamish, F.W.H., D.L.G. Noakes, and A. Rossiter. 1998. Feeding ecology of juvenile Lake Sturgeon, Acipenser fulvescens, in northern Ontario. Canadian Field-Naturalist 112:459468. Boucher, P. 1664. Histoire véritable et naturelle des mœurs et productions du pays de la NouvelleFrance vulgairement dite le Canada. Société historique de Boucherville, Québec, 1964. Callaway, C. G. 1989. Indians of North America: The Abénaki. Chelsea House. Clermont, N., C. Chapdelaine, and J. Cinq-Mars. 2003. L’Ile aux Alumettes. L’Archaïque supérieur dans l’Outaouais. Paléo-Québec 30, Recherches Amérindiennes au Québec,

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Montréal. Courtemanche, M. 2003. Pratiques halieutiques à la station 4 de la Pointe-du-Buisson (BhF1-1) au Sylvicole Moyen Tardif (920–940 A.D.). M. Sc. thesis, Département d’Anthropologie, Université de Montréal, Québec. Cuerrier, J-P. 1945. Les stades de maturité chez l’esturgeon du lac Saint-Pierre. Office de Biologie, MS 2212, Ministère de la Chasse et des Pêcheries, Québec. —. 1962. Aperçu général sur l’inventaire biologique des poissons et des pêcheries de la région du lac Saint-Pierre. Naturaliste canadien 89:193–213. —. 1966. Observations sur l’esturgeon de lac Acipenser fulvescens Raf. dans la région du lac Saint-Pierre au cours de la période de frai. Naturaliste canadien 93:279–334. Cuerrier, J.-P., F. E. J. Fry, and G. Préfontaine. 1946. Liste préliminaire des poissons de la région de Montréal et du lac Saint-Pierre. Naturaliste canadien 73:17–32. Cuerrier, J.-P., and G. Roussow. 1951. Age and growth of lake sturgeon from Lake St. Francis, St. Lawrence River. Canadian Fish Culturist 10:17–29. D’Amours, J., S. Thibodeau, and R. Fortin. 2001. Comparison of lake sturgeon (Acipenser fulvescens), Stizostedion spp, Catostomus spp, Moxostoma spp., quillback (Carpiodes cyprinus) and mooneye (Hiodon tergisius) larval drift in des Prairies River, Québec. Canadian Journal of Zoology 79:1472–1489. Doyon, C., S. Boileau, R. Fortin, and P. A. Spear. 1998. Rapid HPLC analysis of retinoids and dehydroretinoids stored in fish liver: Comparison of two lake sturgeon populations. Journal of Fish Biology 53:973–976. Doyon, C., R. Fortin, and P. A. Spear. 1999. Retinoic acid hydroxylation and teratogenesis in lake sturgeon (Acipenser fulvescens) from the St. Lawrence River and Abitibi region, Québec. Canadian Journal of Fisheries and Aquatic Science 56: 1428–1436.

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Dubreuil, R., and J.-P. Cuerrier. 1950. Cycle de maturation des glandes génitales chez l’esturgeon de lac (Acipenser fulvescens, Raf.). Institut de Biologie générale et de Zoologie, Université de Montréal, Québec. Dumas, R., F. Trépanier, and M. Simoneau. 2003. Fish problems and partnership solutions: The lake sturgeon case in the L’Asomption watershed. American Fisheries Society 133rd Annual Meeting, Québec City, Canada, August 10–14. Dumont, P., F. Axelsen, H. Fournier, P. Lamoureux, Y. Mailhot, C. Pomerleau, and B. Portelance. 1987. Avis scientifique sur le statut de la population d’Esturgeon jaune dans le système du fleuve Saint-Laurent. Ministère du Loisir, de la Chasse et de la Pêche du Québec et ministère de l’Agriculture, des Pêcheries et de l’Alimentation du Québec. Comité scientifique conjoint. Avis scientifique 87/1. 21 p. Dumont, P., P. Bilodeau, and J. Leclerc. 2005. Portrait sommaire de la faune ichtyologique du Courant Sainte-Marie (fleuve Saint-Laurent). Travail réalisé pour le Comité du Bassin du Havre, Ministère des Ressources naturelles et de la Faune, Longueuil, Québec. Dumont, P., J. D’Amours, S. Thibodeau, N. Dubuc, R. Verdon, S. Garceau, P. Bilodeau, Y. Mailhot, and R. Fortin. 2011. Effects of the development of a newly created spawning ground in the des prairies River (Quebec, Canada) on the reproductive success of lake sturgeon Acipenser fulvescens. Journal of Applied Ichtyology 27:394-404. Dumont, P., R. Fortin, G. Desjardins, and M. Bernard. 1987. Biology and exploitation of lake sturgeon (Acipenser fulvescens) in the Québec waters of the Saint-Laurent River. In: Proceedings of a workshop on lake sturgeon (Acipenser fulvescens), Feb. 27–28 1986, Timmins. C. H. Olver, ed. Ontario Fisheries Technical Report. Series 23: 57–76.

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Dumont, P., P. Lamoureux, G. Laforce, M. La Haye, and N. Fournier. 1989. Influence de la dimension de l’hameçon sur la sélectivité et le rendement de la ligne dormante pour la capture de l’esturgeon jaune (Acipenser fulvescens). Québec, Ministère du Loisir, de la Chasse et de la Pêche et Ministère de l’Agriculture, des Pêcheries et de l’Alimentation, Avis scientifique 89/1. Dumont, P., J. Leclerc, J.-D. Allard, and S. Paradis. 1997. Libre passage des poissons au barrage de Saint-Ours, rivière Richelieu. Québec, ministère de l’Environnement et de la Faune, Direction régionale de la Montérégie et Direction des ressources matérielles et des immobilisations, et ministère du Patrimoine canadien (Parcs Canada). Dumont, P., J. Leclerc, Y. Mailhot, E. Rochard, C. Lemire, H. Massé, Hélène Gouin, Denis Bourbeau, and Daniel Dolan. 2000a. Suivi périodique de l’évolution du recrutement de l’esturgeon jaune en 1999. In: Compte rendu du cinquième atelier sur les pêches commerciales, Duchesnay, 18–20 janvier 2000. M. Bernard and C. Groleau, eds. Québec, Société de la faune et des parcs du Québec. Dumont, P., Y. Mailhot, R. Dumas, and P. Bilodeau. 2000b. Plan de gestion de l’esturgeon jaune du fleuve Saint-Laurent 2000–2002. Société de la faune et des parcs du Québec, Directions de l’aménagement de la faune du Centre-du-Québec, de Lanaudière, de la Montérégie et de Montréal . Fleury, C., and D. Desrochers. 2004. Validation de l’efficacité des passes à poisson au lieu historique national du Canal-de-Saint-Ours saison 2003. Rapport final préparé pour Parcs Canada par Milieu Inc, Laprairie, Québec.

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Fortin, R., J. D’Amours, and S. Thibodeau. 2002. Effets de l’aménagement d’un nouveau secteur de frayère sur l’utilisation du milieu en période de fraie et sur le succès de reproduction de l’esturgeon jaune (Acipenser fulvescens) à la frayère de la rivière des Prairies. Rapport synthèse 1995–1999. Pour l’Unité Hydraulique et Environnement, Hydro-Québec et la Société de la faune et des parcs du Québec, Direction de l’aménagement de la faune de Montréal, de Laval et de la Montérégie. Département des Sciences biologiques, Université du Québec à Montréal. Fortin, R., P. Dumont, and S. Guénette. 1996. Determinants of growth and body condition of lake sturgeon (Acipenser fulvescens). Canadian Journal of Fisheries and Aquatic Science 53:1150–1156. Fortin, R., S. Guénette, and P. Dumont. 1992. Biologie, modélisation et gestion des populations d’esturgeon jaune (Acipenser fulvescens) dans 14 réseaux de lacs et de rivières du Québec. Québec, Ministère du Loisir, de la Chasse et de la Pêche, Direction régionale de Montréal et Service de la faune aquatique, Montréal et Québec. Fortin, R., J.-R. Mongeau, G. Desjardins, and P. Dumont. 1993. Movements and biological statistics of lake sturgeon (Acipenser fulvescens) populations from the St. Lawrence and Ottawa River system, Québec. Canadian Journal of Zoology 71:638–650. Fry, F. E. J., G. Préfontaine, et al. 1941. Alimentation de quelques espèces de poisson du lac SaintLouis et du lac des Deux-Montagnes. Rapport de la Station biologique de Montréal, pour l’année 1941. Ministère de la Chasse et des Pêcheries, Québec, Fascicule II, Appendice X, 188–218. Garceau, S., and P. Bilodeau. 2004. La dérive larvaire de l’esturgeon jaune (Acipenser fulvescens) à la rivière des Prairies, aux printemps 2002 et 2003. Ministère des Ressources naturelles, de la Faune et des Parcs, Direction de l’aménagement de la faune de Montréal, de Laval et de la Montérégie, Longueuil, Rapport technique 16-21. GDG Conseil Inc. 2001. Réfection de la centrale de La Gabelle. Programme de surveillance et de

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Hatin, D., S. Lachance, and D. Fournier. 2007. Effect of dredged sediment deposition on use by Atlantic sturgeon and lake sturgeon at an open-water disposal site in the St. Lawrence estuarine transition zone. American Fisheries Society Symposium 56:235–255. Haxton, T. J. 2008. A synoptic review of the history and our knowledge of lake sturgeon in the Ottawa River. Ontario Ministry of Natural Resources, Southern Science and Information Technical Report SSI 126. Haxton, T. J., and C. S. Findlay. 2008. Variation in lake sturgeon abundance and growth among river reaches in a large regulated river. Canadian Journal of Fisheries and Aquatic Sciences 65:646–657. Joliff, T. W., and T. H. Eckert. 1971. Evaluation of present and potential sturgeon fisheries of the St. Lawrence River and adjacent waters. New York Department of Environmental Conservation, Cape Vincent Fisheries Station. Khoroshko, P. N., and A. D. Vlasenko. 1970. Artificial spawning grounds of sturgeon. Journal of Ichthyology 10:286–292. La Haye, M., A. Branchaud, M. Gendron, R. Verdon, and R. Fortin. 1992. Reproduction, early life history, and characteristics of the spawning grounds of the lake sturgeon (Acipenser fulvescens) in des Prairies and L’Assomption rivers, near Montreal, Québec. Canadian Journal of Zoology 70:1681–1689. La Haye, M., S. Clermont, and C. Lemire. 1996. Localisation d’une frayère à esturgeons jaunes dans le cours inférieur de la rivière St-François. Enviro-science inc pour l’Association des Pêcheurs commerciaux du lac Saint-Pierre, Nicolet, Québec. La Haye, M., S. Desloges, C. Côté, J. Deer, S. Philips Jr., B. Giroux, S. Clermont, and P. Dumont.

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2003. Location of lake sturgeon (Acipenser fulvescens) spawning grounds in the upper part of the Lachine rapids. Société de la faune et des parcs du Québec, Direction de l’aménagement de la faune de Montréal, de Laval et de la Montérégie, Longueuil, Technical Report 16-15E. La Haye, M., S. Desloges, C. Côté, A. Rice, S. Philips Jr., J. Deer, B. Giroux, K. de Clerk, and P. Dumont. 2004. Search for and characterization of lake sturgeon (Acipenser fulvescens) spawning grounds in the upstream portion of the Lachine Rapids, St. Lawrence River, in 2003. Ministère des Ressources naturelles, de la Faune et des Parcs, Direction de l’aménagement de la faune de Montréal, de Laval et de la Montérégie, Longueuil, Technical Report 16-20E. La Haye M., S. Guénette, and P. Dumont. 1990. Utilisation de la frayère de la rivière Ouareau par l’Esturgeon jaune suite à l’éboulis survenu en mars 1990. Québec, Ministère du Loisir, de la Chasse et de la Pêche, Direction régionale de Montréal. Rapport technique 06-07. Magnin, E. 1962. Recherches sur la systématique et la biologie des Acipenséridés Acipenser sturio L., Acipenser oxyrhynchus Mitchill et Acipenser fulvescens Raf. Thèse présentée à la faculté des sciences de l’Université de Paris. Imprimerie nationale. —. 1977. Croissance, régime alimentaire et fécondité des esturgeons Acipenser fulvescens Rafinesque du bassin hydrographique de la Grande rivière (Québec). Naturaliste canadien 104:419–427. Magnin, E., and G. Beaulieu. 1960. Déplacements des esturgeons Acipenser fulvescens et A. oxyrhyncus du fleuve Saint-Laurent d’après les données de marquage. Naturaliste canadien 87:237–252.

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Mailhot, Y., and P. Dumont. 1998. Avis scientifique: Révision du statut du stock d’esturgeon jaune du fleuve Saint-Laurent. In: Compte rendu du troisième atelier sur les pêches commerciales, Duchesnay, Ministère de l’Environnement et de la faune du Québec, Québec, 13–15 janvier 1998. M. Bernard and C. Groleau, eds. —. 1999. Mise à jour de l’état du stock d’esturgeon jaune du fleuve Saint-Laurent. In: Compte rendu du quatrième atelier sur les pêches commerciales, Duchesnay, Faune et Parcs Québec, Québec, 12–14 janvier 1999. M. Bernard and C. Groleau, eds. Mailhot, Y., P. Dumont, and N. Vachon. 2011. Management of the lake sturgeon Acipenser fulvescens population in the lower St. Lawrence River (Québec, Canada) from the 1910s to the present. Journal of Applied Ichtyology 27: 405–410. Mélançon, C. 1936. Les poissons de nos eaux. Librairie Granger. Mingelbier, M., P. Brodeur, and J. Morin. 2004. Impacts de la régularisation du débit des Grands Lacs et des changementss climatiques sur l’habitat du poisson du fleuve Saint-Laurent. Vecteur Environnement 37(6): 34–43. —. 2005a. Recommendations concerning fish and their habitats in the fluvial St. Lawrence and evaluation of the regulation criteria for the Lake Ontario–St. Lawrence River system. Report presented to the International Joint Commission. Ministère des Ressources naturelles et de la Faune du Québec, Direction de la recherche faunique, Québec. —. 2005b. Modélisation numérique 2D de l’habitat des poissons du Saint-Laurent fluvial pour évaluer l’impact des changements climatiques et de la régularisation. Le Naturaliste Canadien 129:96–102. Mongeau, J.-R., J. Leclerc, and J. Brisebois. 1982. La dynamique de la reconstitution des populations de l’esturgeon jaune, Acipenser fulvescens, du lac des Deux-Montagnes, province de Québec, de 1964 à 1979. Québec, Ministère du Loisir, de la Chasse et de la

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Pêche, Rapport technique 06-33. Montpetit, C. 1897. Les poissons d’eau douce du Canada. C.-O. Beauchemin & Fils. Morin, J., and M. Leclerc. 1998. From pristine to present state: Hydrology evolution of Lake SaintFrançois, St. Lawrence River. Canadian Journal of Civil Engineering 25: 864–879. Ndayibagira, A., M.-J. Cloutier, P. D. Anderson, and P. A. Spear. 1995. Effects of 3,3,’ 4.4’-tetrachlorobiphenyl on the dynamics of the vitamin A in brook trout (Salvelinus fontinalis) and intestinal retinoid concentration in lake sturgeon (Acipenser fulvescens). Canadian Journal of Fisheries and Aquatic Science 52:512–520. Nilo, P., P. Dumont, and R. Fortin. 1997. Climatic and hydrological determinants of year-class strength of St. Lawrence River lake sturgeon (Acipenser fulvescens). Canadian Journal of Fisheries and Aquatic Science 54:774–780. Nilo, P., S. Tremblay, A. Bolon, J. Dodson, P. Dumont, and R. Fortin. 2007. Feeding ecology of juvenile lake sturgeon Acipenser fulvescens in the St. Lawrence River system. Transactions of the American Fisheries Society 135:1044–1055. Paradis, S., and R. Malo. 2003. Efficiency of the Vianney-Legendre fish ladders at the Saint-Ours Canal National Historical Site, Richelieu River, Quebec. American Fisheries Society 133rd Annual Meeting, Québec City, Canada, August 10–14. Peake, S., F. W. H. Beamish, R. S. McKinley, D. A. Scruton, and C. Katapodis. 1997. Relating swimming performance of lake sturgeon, Acipenser fulvescens, to fishway design. Canadian Journal of Fisheries and Aquatic Sciences 54:1361–1366. Robitaille, J. A., Y. Vigneault, G. Shooner, C. Pomerleau, and Y. Mailhot. 1988. Modifications physiques de l’habitat du poisson dans le Saint-Laurent de 1945 à 1984 et effets sur les pêches commerciales. Canadian Technical Report of Fisheries and Aquatic Sciences 1808.

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Rousseaux, C. G., A. Branchaud, and P. A. Spear. 1995. Evaluation of liver histopathology and erod activity in St. Lawrence lake sturgeon (Acipenser fulvescens) in comparison with a reference population. Environmental Toxicology and Chemistry 14:843–849. Roussow, G. 1955a. Quelques observations sur les variations de forme et de couleur chez les esturgeons de la province de Québec. Annales de l’Acfas 21:79–85. —. 1955b. Les esturgeons du fleuve Saint-Laurent en comparaison avec les autres espèces d’Acipenséridés. Office de Biologie, Ministère de la Chasse et des Pêcheries, Province de Québec, Montréal. —. 1957. Some considerations concerning sturgeon spawning periodicity. Journal of Fisheries Research Board of Canada 14:553–572. Scott, W. B., and E. J. Crossman. 1973. Freshwater fishes of Canada. Fisheries Research Board of Canada. Bulletin 184. Thiem J.D., T. R. Binder, J. W. Dawson, P. Dumont, D. Hatin, C. Katopodis, D. Z. Zhu, and S. J. Cooke. 2011. Behaviour and passage success of upriver-migrating lake sturgeon Acipenser fulvescens in a vertical slot fishway on the Richelieu River, Quebec, Canada. Endangered Species Research 15:1–11. Trencia, G., and P.-Y. Collin. 2006. Rapport d’aménagement d’une frayère pour le poisson à la rivière Chaudière. Ministère des Ressources naturelles et de la Faune du Québec, Direction de l’Aménagement de la Faune Chaudière-Appalaches, Lévis. Veillette, F. 2007. Étude de différents indicateurs biologiques chez l’esturgeon jaune (Acipenser fulvescens) du Québec. M. Sc. Thesis. Université du Québec à Montréal. Vladykov, V. D. 1948. Rapport du biologiste du département des pêcheries. 2. Nourriture de

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l’esturgeon. In: Rapport général du Ministre de la Chasse et des Pêcheries de la Province de Québec concernant les activités du département des pêcheries pour l’exercice financier 1947–48, Québec. —. 1955. Fishes of Quebec. Sturgeons. Album 5. Department of Fisheries, Quebec. Vladykov, V. D., and G. Beaulieu. 1946. Études sur l’esturgeon (Acipenser) de la province de Québec. I: Distinction entre deux espèces d’esturgeons par le nombre de boucliers osseux et de branchiospines. Naturaliste canadien 73:143–204. —. 1951. Études sur l’esturgeon (Acipenser) de la province de Québec. II: Variations du nombre de branchiospines sur le premier arc branchial. Naturaliste canadien 78:129–154. Vladykov, V. D., and C. Gauthier. 1941. Remarques sur le régime alimentaire de l’esturgeon (A. fulvescens) dans le lac Saint-Louis. Rapport de la Station biologique de Montréal, pour l’année 1941. Ministère de la Chasse et des Pêcheries, Québec, Fascicule III, Appendice IX: 384–387. Vladykov, V. D., and J. R. Greely. 1963. Order Acipenseroidei. In: Fishes of the Western North Atlantic, no. 1, part 3. Soft-rayed bony fishes. H. B. Bigelow, ed. Memoir Sears Foundation of Marine Research, Yale University.

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Welsh, A., T. Hill, H. Quinlan, C. Robinson, and B. May. 2008. Genetic assessment of lake sturgeon population structure in the Laurentian Great Lakes. North American Journal of Fisheries Management 28:572–591.

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MARTY HOLTGREN

Bringing Us Back to the River

The annual nmé (lake sturgeon) return and its celebration by our Peoples assure the renewal and continuation of human and all other life.

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—Little River Band of Ottawa Indians Nmé Stewardship Plan

The history of nmé in the Great Lakes is a story of harmony, tragedy, and an opportunity for redemption. The tragedy has been well documented by historians, academics, and writers chronicling a time when sturgeon were an abundant member of the lakes and their later spiral toward near extinction (Auer 1999; Tody 1974; Schoolcraft 1970; Harkness and Dymond 1961). I have sat through many lectures on sturgeon where I was presented with the same exhausting information about how their habitat was destroyed, how they were overharvested, and why the current outlook is so bleak. I believe it is time for a different story for the sturgeon, one of harmony and commitment toward their recovery, where the story comes from many voices. One such source is from Native American people who have lived in harmony with sturgeon for millennia. Their relationship with the sturgeon may be characterized by conservation approaches, stewardship, and religious beliefs (LRBOI 2008; Rettig, Berkes, and Pinkerton 1989). Native American oral history provides us information about a time when the sturgeon and human communities were in balance within the Great Lakes ecosystem. This history, and the worldview that comes with it, can provide a road map to restoring this species’ place in the Great Lakes—an opportunity for redeeming the tragedy that has been so well documented. n

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This opportunity for the Little River Band of Ottawa Indians (LRBOI) came in the form of a cargo trailer on the banks of the Big Manistee River. When I tell people that I am a fisheries biologist for the LRBOI and doing sturgeon restoration, I usually get a blank look followed by puzzlement and then a set of common questions. “Why would a tribe have a natural resources department and be involved with managing fish?” “Would a tribal management approach be different from that of the other agencies?” Within this chapter I hope to make apparent the answer to these questions by detailing the road map to redemption I spoke of earlier. I will use a case study from the LRBOI to demonstrate a unique tribal approach to restoration and stewardship. The answer to the first question is embedded in a deep history of cultural, social, and political elements. Simply put, the tribes in the Great Lakes manage natural resources to protect cultural sovereignty and to meet the generational and unique needs of tribal members. Cultural sovereignty is the process of tribes making decisions internally that protect traditions and customs (Coffey and Tsosie 2001), and this is evident throughout the Great Lakes, as the tribes are managing watersheds, reservation natural resources, and species that are of great consequence to them. The tribes have a need to manage the fishery in addition to other management institutions because tribal needs and worldviews are often very different from those of the general population (Berkes 2009; Mattes and Kmiecik 2006; Kimmerer 2000; Salmon 2000; Notske 1995; Busiahn 1989). The sturgeon population in the Big Manistee River, Michigan, is an example where for over 100 years the population was overlooked, where decisions and concerns for resource managers were often how many exotic trout should be stocked or what the fisherman’s opinion may be about that year’s harvest. Out of necessity and a cultural responsibility, the tribes began to work on restoring the sturgeon because it was unacceptable to them to lose a species that is revered and belongs in the watershed (LRBOI 2008). So why different management strategies for different agencies? The basis of state fisheries management relies on ownership of the fisheries resource. This is known as the common property principle, where the entire populace owns the fishery and the state government has the right and responsibility of being the trustee (Henquinet and Dobson 2006; Nielsen 1999). Within this framework, the state has the difficult task of maintaining open access to the fishery while ensuring the protection, sustainability, and productivity of the resource. Over the past century many of the species that are important to the tribes did not receive priority under the auspices of the public trust. Examples are abundant; in the state of Michigan millions of dollars have been spent on introducing nonnative species with little funding being spent on endemic species of significance to the tribes, including lake sturgeon, sucker, and the extirpated Arctic

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grayling. Therefore, the tribes have a need to keep these species present regardless of what the current state management perspective or funding priorities may be. The LRBOI has a past that recognizes the importance of sturgeon and a future that is focused on preserving it (LRBOI 2008; McClurken 2009). The Big Manistee River defines the tribe’s reservation, which is one of the few rivers on the eastern shoreline of Lake Michigan supporting a sturgeon population known to have a small group of spawners. Historically, tribal people would gather on the banks of the river each year for the lake sturgeon, sucker, and Arctic grayling spawning runs. Jay Sam, tribal cultural preservation director, says this about this historic event: “The grandfather fish (sturgeon), and its relatives the undermouth fish (sucker), they would sacrifice themselves during the sucker moon so the people would have food until the other crops were available.” The sturgeon is a clan spirit (LRBOI 2008). The historical importance of these clan spirits within tribal communities is evidenced on the pages of documents from the 1800s, where Tribal people would “sign their names” not in English but with symbols and their Aniishinabek names that represented their family lineage and clans (LRBOI 2008; GLIFWC 2007). On these documents is the distinct image of the sturgeon. These clan spirits are often the focus of tribal natural resources departments: the LRBOI, Menominee Indian Tribe of Wisconsin (Runstrom et al. 2002), and White Earth Nation with sturgeon, the Little Traverse Bay Bands of Odawa Indians with wolves, and the Grand Traverse Band of Ottawa and Chippewa Indians with martin are just a few examples. In 1836 a treaty was signed where five tribes ceded about one-third of what is now known as the state of Michigan. The cession guaranteed that the tribes would be granted the “Usual privileges of occupancy” to hunt, fish, and gather from the land and waters. I cannot help but wonder how the tribal chiefs at the time of the treaty signing viewed the Great Lakes. Could they envision a time when the clan spirits and the species so vital to their communities would be gone? The list is exhaustive. The Arctic grayling and woodland caribou have gone extinct, and the populations of wolf, martin, and moose are only a fraction of what they once were. This is where the tribes have continued to play a large role in management by participating in natural resource management decision making, developing tribal stewardship plans, conducting biological assessments and restoration projects.

New Beginnings In 2001 the LRBOI began to quantify fish populations within its reservation waters. Past research had shown that the sturgeon population was very small, and no true evidence of recent natural reproduction was available. The tribe hired fisheries

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professionals, purchased equipment, and began collecting information on the lake sturgeon. From the very beginning the tribal community was excited about the sturgeon program and at the prospect of seeing them restored. The excitement produced what was called a “Cultural Context Group” made up of tribal members and biologists that would develop goals and objectives for their sturgeon program and ultimately a stewardship plan that would guide the tribal natural resources department in their lake sturgeon restoration. This group had representatives from many different sections of the tribal community; men and women, elders and youth, artisans and pipe carriers. Rather than just having biologists determine restoration strategies, this group would provide a tribal voice in the management direction. This “voice” was an amalgamation of cultural, biological, political, and social elements, all being important and often indistinguishable from each other. The biologists of the group recognized early on that an exciting part of developing a sturgeon plan was using sound biological principles to meet objectives that were not necessarily or exclusively biological. For instance, the first goal of the stewardship plan was “Restore the harmony and connectivity between nmé and the Anishinaabek and bring them both back to the river.” Bringing the sturgeon back to the river was an obvious biological element; however, restoring harmony and connectivity between sturgeon and people was steeped in the cultural and social realm. Within these Cultural Context Group meetings, we observed that the depth of the relationship between the sturgeon and the tribe would bring a unique management perspective. Each “meeting” began with a ceremony, and the conversation was held over a feast, including wild rice soup and fry bread. After almost two years the stories, the friendly dialogue, and the vision shared at these meetings would produce a plan that would guide the LRBOI’s sturgeon work for the next seven generations. The four goals of the plan were these: • Restore the harmony and connectivity between nmé and the Anishinaabek and bring them both back to the river • Restore the nmé and reclaim the environment on which it depends for future generations of nmé and Anishinaabek in perpetuity • Emphasize strategies that promote natural reproduction and a healthy watershed • Protect tribal sovereignty and treaty rights

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Medicine and Drift Nets One of the most challenging species to rehabilitate in the Great Lakes may be nmé, largely due to a life history that is extreme when compared to other freshwater species. Probably the most daunting realization for me as a nmé biologist is that the management actions that we implement today will not be fully realized within my career because nmé mature so slowly and may spawn for the first time only after 10–20 years of life (Auer 1996). The first challenge in implementing a restoration plan was to fully understand the status of the sturgeon population. One of the best ways to determine the status is to capture and count the newly hatched larvae. A larval drift survey may be one of the more challenging and demanding surveys that biologists conduct within the Great Lakes Basin, and “larval drift” therefore is a phrase that is banned from the LRBOI natural resources department once the surveys are completed in the spring because of the exhaustion and stress that these surveys cause (this is only a partial joke!). Why the disdain for drift surveys? Unfortunately for fisheries staff, nmé larvae (fry) can most effectively be captured at night. After nmé hatch they remain buried in the gravel for a few days; once they have absorbed most of their yolk sac they drift downriver starting just after sunset through the early morning hours. To capture nmé larvae fisheries staff need to become nocturnal, face the rain, snow, and ice, hike through the woods, and wade into chest-deep, swift-moving rivers with headlamps to set anchors and check drift nets. This is done every night for many consecutive days. When we started conducting these surveys in 2002, no lake sturgeon larvae had been captured in any Lake Michigan tributary including the Big Manistee River even after two years of drift surveys had been conducted by a university. This was alarming because it indicated that either no nmé were successfully reproducing or only a few were surviving. Either way the prospects appeared bad. We decided to move our drift study site closer to the expected spawning area than the past researchers to increase the probability of capturing nmé. Three LRBOI staff, Mark Bowen, Stephanie Ogren, and I, began heading down to a desolate location on the bank of the Big Manistee River at night to attempt capture of young nmé. We set out four drift nets into the fast-flowing water, collected our catch every hour, and meticulously went through the samples. For the first few nights we caught almost everything in the river but sturgeon—thousands of sucker larvae, aquatic insects, lamprey amocetes, salmon, and trout fry. We were frustrated and disheartened until one evening Mark Bowen told us that we needed to follow a teaching that had been passed on to him. Both Stephanie

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and I watched as he pulled out his medicine pouch, put tobacco into the water, and prayed that the grandfather fish would allow itself to be captured. I mention this because it is an example of trying to achieve the first goal within the plan, bringing the sturgeon and the people back to the river (restoring the fish/human relationship). This also exemplifies the integration of culture and biology—the tobacco floating upon the river right next to drift nets. That night I remember anticipating our first net pull to see if we would find a nmé. As I poured my sample slowly into a white tray I scanned for signs of a swimming nmé with my flashlight. As I looked at dozens of frantically swimming sucker larvae I saw something different, a grayish fish with a blunt head that was swimming entirely differently. For a moment I didn’t know what species of fish it was. I had seen hundreds of sturgeon larvae before on the Sturgeon River (Barage County, Michigan) with my graduate advisor, Nancy Auer, but this time I was the one responsible for identifying the sturgeon, and this fish was smaller than any one I had observed before. I didn’t realize it at the time but I was holding my breath, and my two partners, both aware of this, had stopped going through their samples and were intently watching me. When I looked up at them with a grin on my face, they both knew we had a fish, a sign that there were still reproducing sturgeon in the Big Manistee River. Mark leaned over our table, over the little sturgeon, and gave me a hug. After the fish was measured Mark carefully took the fish in his hand and released it back to the river. The connection between people and sturgeon had begun. This little sturgeon, and the few others we captured that year, demonstrated there was a small amount of natural reproduction taking place and also gave us an idea that would eventually be applied across the Lake Michigan Basin for restoring sturgeon.

Bringing Back the Sturgeon Tribes are not often recognized for the large amount of fishery research and restoration they do. Each year tribes in the Great Lakes conduct thousands of hours of biological assessments and numerous habitat restoration projects and bring millions of dollars into fishery improvements, such as fish stocking, improved road-stream crossings, bank stabilizations, and dam removals (USDOI 2007; Snyder et al. in preparation). This work benefits not only the tribal members but each and every person who enjoys the natural resources of the area. The LRBOI began dedicating hundreds of hours to sturgeon research after capturing the first sturgeon larvae. We captured eggs that in turn identified spawning locations, we found juvenile fish that demonstrated survival from larvae, and we began to determine what habitat

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conditions were present and needed for spawning lake sturgeon. With this information, we could begin considering management approaches for restoring the nmé. Since 2002 a dedicated group of scientists has gathered every two years to discuss lake sturgeon rehabilitation strategies. One of the critical topics discussed at the first meeting was the unique genetic arrangements and structure that was observed in remnant populations of nmé across the Great Lakes. By taking a tissue sample from a nmé (a small piece of a fin) researchers were able to assign quite accurately the river in which that particular fish had originated (Welsh et al. 2010; Welsh et al. 2008; DeHann et al. 2006). Genetic structuring indicated that sturgeon were philopatric, meaning that the spawning fish returned faithfully to the same river that they themselves originated (DeHaan et al. 2006). This imprinting of fish to a particular area for spawning is exhibited in other species as well, notably salmon, and changes the way we should look at management of these fish. This meant something very important to those at the meeting and for those making management decisions for nmé; this genetic structuring (and imprinting) needed to be maintained because of important evolutionary traits that could be unique to each of the populations. The LRBOI started to develop a strategy for increasing sturgeon abundance in the Big Manistee River. One strategy historically used in sturgeon restoration was rearing fish in an off-site hatchery often far away from the river where the fish would be stocked. However, during the LRBOI Cultural Context Meetings we had discussed this strategy, and many of the participants were not supportive of the idea based on cultural values. It was clearly communicated that the sturgeon was a grandfather fish and part of the Big Manistee River watershed at all stages of life for a reason. The risk of altering their behavior by being reared in water from an off-site hatchery was not acceptable. The stewardship plan was clearly guiding us to keep the fish in the river at all times: “The Creator put the nmé in the Big Manistee River. The nmé and the rivers they use are part of our sense of place. The Creator put us here where the nmé return. We are obliged to remain and protect this place.”(LRBOI 2008). The group believed that maximizing the chances of imprinting was essential, not only for the Big Manistee River sturgeon but also for the small surrounding populations where straying of Big Manistee River fish could adversely impact those populations as well. As Don Stone, a tribal elder, put it, “These fish were here when our ancestors were here,” and we needed to make sure that they would come back as they always had. The biologists needed to determine a strategy that would accommodate the cultural and biological perspectives related to sturgeon restoration. The second question I posed above, “Would a tribal management approach be different from that of the other agencies?” has a simple answer—yes. There are often different management goals based on the unique culture of the tribes that in turn create different strategies toward management.

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I find it interesting that at the same time sturgeon managers from across the Great Lakes were questioning traditional fishery management techniques for restoring sturgeon, tribal people were coming up with similar conclusions based on culture and biology. It demonstrated to me the great conservation potential that may be gained by including multiple perspectives and developing a shared knowledge where the outcome (which will be described below) will often be much richer and innovative than if only one perspective was included (Natcher, Davis, and Hickey 2005). By combining these perspectives there is much more than sturgeon restoration being accomplished. Jay Sam and Art deBres describe this philosophy as, “We are not introducing, we are rehabilitating . . . we are assisting and saving our mother, grandfather and cousin.” (LRBOI 2008).

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A New Approach to Sturgeon Management: Streamside Rearing In 2003 the LRBOI decided to design and operate a streamside rearing facility to rear lake sturgeon (Holtgren et al. 2007). The larvae that were captured during drift surveys would be protected for a few months inside the rearing facility and released back into the river (plate 12). Even though streamside rearing had been used successfully for salmon, trout, and walleye (Dupuis and Dominy 1994; Steward 1996), it had never been applied to sturgeon. The Big Manistee River system provided its own unique challenges for keeping the streamside-rearing facility running effectively. The challenge for us was to create a system that pulled the water from the river without having the rearing tanks inundated with the silt and sand that the river carried, especially during high flow events. The system also had be cost-effective, incorporate genetic conservation, and address the concerns of imprinting and spawning site fidelity (Holtgren et al. 2007). The streamside rearing facility was built inside a cargo trailer for mobility (plate 13). It is pulled by truck to a forested area along the banks of the Big Manistee River each spring and put into storage each fall. From the outside it appears to be indistinguishable from one you would see on the road except for the large tribal logo and black lettering that says, Nmé Kooginaawsawin Koh-ge-now-sa-win, which means “The sturgeon home, where children are raised” in Anishinaabemowin (the language of the Ottawa). However, the inside is complete with water quality monitors, alarms, and safety systems. The water is pumped into the facility through 100 meters of underground piping and enters a set of mechanical sediment filters, which remove a portion of the silt and sand load. The water is then forced into a large reservoir above the trailer and gravity-fed through the sidewall into the tanks and immediately drains back into the river. Because of the remote location of the

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rearing facility, a safety system was necessary in case of a malfunction. We have a notification system that activates a phone when the water decreases to a low flow rate or stops and practically calls everyone in the department until someone checks to see what the problem may be. There have been nights when someone receives a call at two or three in the morning and our department phone tree lights up. No one rests until the fish are safe. Our larval collection procedure and the streamside rearing approach were appealing for many reasons. First, by capturing larvae for rearing we were allowing the spawning sturgeon to continue their natural process. Instead of capturing the prespawn fish and “stripping” their eggs and sperm, the sturgeon were selecting how they would mate and therefore continue the unique genetic structuring found within the population. Also, we decided to only collect 10 percent or less of the drifting larvae to ensure that if something did happen to the fish we were rearing, an adequate number of wild fish could provide that year’s production. In a strict sense of the word, we were not necessarily even “stocking” fish but augmenting the population by removing fish already within the population and simply increasing their prospects of survival. When sturgeon are larvae they are quite vulnerable to many sources of predation. Many of these predators are relatively new to the watershed (intentional and unintentional releases) and are species the sturgeon has not necessarily developed defense strategies against. By rearing collected larvae we could also increase the chances of imprinting by keeping the fish in their own river water throughout their early life.

Bringing Both Back to the River The week leading up to the release of streamside-reared fish is hectic, and we are busy being scientists. Each fish is marked with a small internal tag so if it is later captured it can be identified and we can evaluate if the program is meeting its goals and objectives. Tissue samples are collected, including a small snip of the tail fin, to better understand the genetic makeup of the population and to ensure that what we release is representative of that in the wild. We attach small radio transmitters to a portion of the fish to monitor how the reared fish behave compared to their wild counterparts. After those activities, the week is no longer about science but about people and fish coming together next to the banks of the river. Marcella Leusby described this concept by offering, “When putting the sturgeon back in the river, I felt it was one of the most meaningful acts the LRBOI had done. It was very emotional.” (LRBOI 2008). The nmé release is a celebration, a celebration with drumming and dancing,

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Figure 1. Drummers playing at the nmé release ceremony.

Figure 2. Young people releasing young sturgeon for future generations.

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Figure 3. Jimmie Mitchell singing a song while nmé are released.

Figure 4. Mark Bowen with a transmittered nmé about to be released.

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recognizing invaluable partners and finally commencing with a pipe ceremony. After the pipe, the nmé are taken from the raceways, carried overland in five-gallon buckets, and then released one by one, by hand, into the river. After the fish are released, a crowd of a couple hundred people stand silently for a few moments and begin to talk about the nmé and about how they will be ready to celebrate again next year. The gathering of people is a mixture of the watershed community encompassing both tribal and nontribal people, where the differences among cultures are celebrated and the similarities are apparent. Bringing nmé and people back to the river is now a goal of the watershed community. Yes, the tribe largely funds the program (with support from the U.S. Fish and Wildlife Service) and provides the expertise, but the nontribal community provides tremendous support. The day when we release nmé into the river is just as much about human ecology, how humans fit into the natural and social environments. It is at this type of celebration that I understand we are once again learning humans are part of the ecosystem, not an organism somehow separated. By putting our hands in the water and having the young nmé slowly swim away, we are seeing ourselves in a different way. After rearing sturgeon for over seven years, the stewardship plan of the LRBOI is beginning to be realized. Each year adds new memories and stories to the rich relationship between humans and nmé. The beginning of nmé rearing begins with a simple act, someone taking their hand and gently guiding the newly captured nmé larvae into a holding tank to be transported to the streamside rearing facility. The end of nmé rearing also begins with a hand gently guiding the larger nmé (around 250 mm TL [10 inches]) back into the Big Manistee River.

Final Thoughts I stated at the beginning of this chapter that a goal of the stewardship plan was to promote tribal sovereignty and treaty rights. Tribal sovereignty allows tribes to conduct projects like nmé restoration in the Big Manistee River, to bring people back to the river, to appreciate where we come from, and ultimately to understand each other. In this example, sovereignty is not some scary prospect where there is misunderstanding, confusion, and even anger between tribal and nontribal people. This is about harmony.

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REFERENCES

Auer, N. A. 1996. Importance of habitat and migration to sturgeons with emphasis on lake sturgeon. Canadian Journal of Fisheries and Aquatic Sciences 53(S1):152–160. —. 1999. Lake sturgeon: A unique and imperiled species in the Great Lakes. In: Great Lakes Fisheries Policy and Management: A Binational Perspective. W. W. Taylor and C. P. Ferreri, eds. Michigan State University Press. Berkes, F. 2009. Evolution of co-management: Role of knowledge generation, bridging organizations and social learning. Journal of Environmental Management 90:1692–1702. Busiahn, T. R. 1989. The development of state/tribal co-management of Wisconsin fisheries. In: Cooperative Management of local fisheries: New directions for improved management and community development. E. Pinkerton, ed. University of British Columbia Press. Coffey, W. and R. Tsosie. 2001. Rethinking the Tribal Sovereignty Doctrine: Cultural sovereignty and the collective future of Indian nations. Stanford Law & Policy Review 12:191–202. DeHaan, P. W., S. T. Libants, R. F. Elliott, and K. T. Scribner. 2006. Genetic population structure of remnant lake sturgeon populations in the upper Great Lakes Basin. Transaction of the American Fisheries Society 135:1478–1492. Dupuis, T., and L. Dominy. 1994. Introduction to satellite rearing: Installation and operation manual. Atlantic Salmon Federation. Great Lakes Indian Fish and Wildlife Commission (GLIFWC). 2007. A guide to understanding Ojibwe treaty rights. Harkness, W. J. K., and J. R. Dymond. 1961. The lake sturgeon. Ontario Department of Lands and Forests, Fish and Wildlife Branch.

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Henquinet, J. W. and T. Dobson. 2006. The public trust doctrine and sustainable ecosystems: A Great Lakes fisheries case study. New York University Environmental Law Journal 14(2): 322–373. Holtgren, J. M., S. A. Ogren, A. J. Paquet, and S. Fajfer. 2007. Design of a portable streamside rearing facility for lake sturgeon. North American Journal of Aquaculture 69:317–323. Kimmerer, R. N. 2000. Native knowledge for native ecosystems. Journal of Forestry 98:4–9. LRBOI (Little River Band of Ottawa Indians). 2008. Nmé (Lake Sturgeon) stewardship plan for the Big Manistee River and 1836 reservation. Natural Resources Department, Special Report 1. Mattes, W. P., and N. Kmiecik. 2006. A discussion of cooperative management arrangements within the Ojibwa ceded territories. In: Partnerships for a common purpose: Cooperative fisheries research and management. A. N. Read and T. W. Hartley, eds. American Fisheries Society Symposium 52. McClurken, L. M. 2009. Our people, our journey: The Little River Band of Ottawa Indians. Michigan State University Press. Natcher, D. C., S. Davis, and C. G. Hickey. 2005. Co-management: managing relationships, not resources. Human Organization 64(3): 240–250. Nielsen, L. A. 1999. History of inland fisheries management in North America. In: Inland fisheries management in North America. 2nd ed. C. C. Kohler and W. A. Hubert, eds. American Fisheries Society. Notske, C. 1995. A new perspective in aboriginal natural resource management: Co-management. Geoforum 26(2):187–209.

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Rettig, B. R., F. Berkes, and E. Pinkerton. 1989. The future of fisheries co-management: A multidisiplinary assessment. In: Cooperative management of local fisheries: New directions for improved management and community development. E. Pinkerton, ed. University of British Columbia Press. Runstrom, A., R. M. Bruch, D. Reiter, and D. Cox. 2002. Lake sturgeon (Acipenser fulvescens) on the Menominee Indian Reservation: An effort toward co-management and population restoration. Journal of Applied Ichthyology 18:481–485. Salmon, E. 2000. Kincentric ecology: Indigenous perceptions of the human-nature relationship. Ecological Applications 10:1327–1332. Schoolcraft, H. R. 1970. Narrative journals of travels through the northwestern regions of the U.S. extending from Detroit through the great chain of American lakes to the sources of the Mississippi in the year 1820. M. L. Williams, ed. Arno Press and the New York Times. Steward, C. R. 1996. Monitoring and evaluation plan for the Nez Perce Tribal Hatchery. U.S. Department of Energy, Bonneville Power Administration, Report 36809-2. Tody, W. H. 1974. Whitefish, sturgeon, and the early Michigan commercial fishery. In: Michigan Department of Natural Resources. Michigan fisheries centennial report, 1873–1973. Michigan Department of Natural Resources, Management Report 6. USDOI (U.S. Department of the Interior). 2007. Fishery status update in the Wisconsin treaty ceded waters. 4th ed. Bureau of Indian Affairs. Welsh, A., R. Elliott, K. Scribner, H. Quinlan, E. Baker, B. Eggold, J. M. Holtgren, C. Krueger, and B. May. 2010. Genetic guidelines for the stocking of lake sturgeon (Acipenser fulvescens) in the Great Lakes Basin. Great Lakes Fishery Commission Special Publication. 2010-01.

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Welsh, A., T. Hill, H. Quinlan, C. Robinson, and B. May. 2008. Genetic assessment of lake sturgeon population structure in the Laurentian Great Lakes. North American Journal of Fisheries Management 28:572–591.

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Sturgeon for Tomorrow

One of the most common questions I’ve been asked over the years is, “How did you become so involved with the lake sturgeon?” I grew up logging hundreds of hours in an ice shanty on Black, Burt, and Mullett lakes in northern Michigan’s Cheboygan County. My late father and brother Dock McCall and James McCall taught me to fish from an early age. I am so blessed they took me into the wilderness and upon our great waters, and for letting me roam free and discover earth’s natural treasures, beauty, and peace. I have never lost the wanderlust for life they and my late mother have instilled in me. In the mid-1990s, I traveled to southern Michigan to spend time with my grandfather Roy Naugle. Grandpa farmed his whole life. He worked hard from daylight till dark eleven months out of the year. His passion was to spend the other month ice fishing on Black, Burt, and Mullett lakes. During my visit, I knew Grandpa was dying. We cried, we laughed, and we talked about sturgeon. Grandpa held my hand, looked me in the eye, and said, “You do what you need to do to keep the sport alive.” I was with my grandfather the first time I saw a sturgeon. I was about six years

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old, and I will never forget that moment. I knew then that there was something extraordinary about this creature. Grandpa and I were sitting in a fish shanty on Burt Lake one February morning in the late 1960s. Suddenly a commotion came from a shanty nearby. We threw the door open to see what was going on. There lay a huge sturgeon on the ice, bigger than I had ever seen. A few dozen anglers from surrounding shanties began to gather around the sturgeon. We trudged through the snow to see the fish close up. The spirit in the air was powerful. I remember looking into the eye of the sturgeon. It amazed me. The diamond shape of the pupil reminded me of pictures I had seen of dinosaurs. That moment and the excitement of that day on the ice with Grandpa have stayed with me ever since. And so have the traditions associated with the sturgeon in our region. Onaway, a quaint community five miles south of Black Lake, is called the Sturgeon Capitol of Michigan. There the Black Lake Sturgeon Shivaree debuted in 1963. The Shivaree is a family fun weekend ice-carnival on Black Lake celebrating sturgeon. A large tent is set up on the ice. It is heated and a central meeting place for everyone to gather, socialize, and register for events. Numerous events for all ages include snowmobile races, ice skating, cross-country skiing, adult and children’s games, food and outfitter vendors, fishing contests, and of course, plenty of beer and a sense of place and community. A sturgeon king or sturgeon queen is crowned for harvesting the largest sturgeon. Area businesses thrive during Shivaree weekend and during ice fishing. Getting out and enjoying the great outdoors is a perfect opportunity for families to gather and break the monotony of cabin fever. Black Lake has long been recognized by local residents and conservation groups for its natural resources and, is, we believe, a key aquatic biodiversity site of the Great Lakes ecoregion. In addition to large kettle lakes, large forested areas, and an expansive network of streams and wetlands, the Black Lake watershed is home to a variety of threatened and endangered aquatic species, including not just the iconic lake sturgeon but also the Michigan monkeyflower and the Hungerford’s crawling water beetle. Several wetlands also provide important nesting habitat for rare birds such as the bald eagle, the common loon, and the great blue heron. Every spring for decades, locals have paddled the Upper Black River to see lake sturgeon spawning in all their glory. Families camp and picnic at several of the known spawning sites, where you can see dozens of the majestic sturgeon spawning in a short span of the river. By the 1990s, it became clear there were poachers on the river, especially at night. Poachers illegally harvested the sturgeon in the spring spawning run, selling

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the roe and smoked fillets on the black market. Poaching was reducing spawning stocks and hindering natural reproduction. This practice has a long history. Beginning in the middle to late 1800s, North Americans became aware of the value of sturgeon. Europeans considered caviar a delicacy and so the demand for sturgeon exploded. In addition to caviar, sturgeon were harvested for a number of purposes: sturgeon meat was delicious, especially smoked; the skin was tanned for leather; and the swim bladders were used for isinglass, a high-quality gelatin used for pottery cement, waterproofing, and clarifying wine and beer. Poaching is and was a family tradition. Some family members who poached them in the past are still bent on taking them, primarily for caviar and their meat. Poachers created an underground network whereby the roe would be bootlegged to Chicago and then to New York, where it was sold on the black market. Today, the Convention on International Trade of Endangered Species (CITES) monitors the import and export of sturgeon, while the U.S. Fish and Wildlife Service is the watchdog for international trade of threatened and endangered animals including sturgeon. Legal harvest of sturgeon is a family tradition, too. In our area there are two, three, and in some instances four generations of avid sturgeon anglers. Ice fishing on area lakes is deeply entrenched in the local culture. Sturgeon is a trophy fish, and fishing for them is indeed a hunt of a lifetime. Fish decoy carvers and spear craftsmen handcraft one-of-a-kind spears and award-winning decoys. Many ice anglers became deeply concerned about the future of their sport of spear fishing through the ice. I was committed and passionate about working to protect this tradition and to ensure a self-sustaining sturgeon population. In 1995 I learned the Michigan Department of Natural Resources (DNR) was reviewing statewide sturgeon regulations. In 1997, the DNR conducted a lake survey to assess the lake sturgeon population in Black Lake. The estimated adult spawning population was 550, and the entire population was thought to be about 1,300. DNR recommended closing the sturgeon harvest completely in all three lakes. We believed poaching was largely responsible for the sturgeon’s decline and that the more recent survey methodology was not standardized with the survey of 1975. It was believed poachers would take many more sturgeon illegally than were taken in legal harvest. It was commonplace. Everyone did it. Fathers of today’s poachers had done it. Their grandfathers, uncles, and cousins had done it. But at several public meetings in 1996–1997 conducted to receive community input on the proposed regulation changes, the state’s position was that the sturgeon population was not reproducing because of fishing pressure. We conducted a statewide outreach campaign, and with the help of local anglers

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and businesses, raised enough money to purchase several larval drift nets to loan to the DNR to assess if there was or was not in fact natural recruitment in the river. Studies have determined there is natural reproduction occurring in the river. The outreach campaign informed ice anglers of the proposed sturgeon regulation changes. Included in the outreach materials were goals and recommendations from the State of Michigan Lake Sturgeon Rehabilitation Strategy (LSRS). For water bodies with sturgeon populations of 500 or more breeding adults (Black Lake’s estimates were 550 breeding adults), there could be a 3 percent harvest for an expanding population, or 6 percent harvest for a sustaining population. Black Lake fell within these parameters. Mullett and Burt Lakes had limited data on sturgeon population assessments. Thirty days into the outreach campaign, we received over 1,300 petition signatures from anglers supporting the goals and recommendations of the LSRS. The overarching goal of the campaign was to determine if there was enough interest in lake sturgeon rehabilitation, and the tenacity to save the sturgeon and save the sport that had become deeply entrenched in the local culture. Indeed there was, and is today! Regardless of the regulation changes, poaching needed to be reduced. Here we had a valuable, rare, living fossil, and not enough was being done to protect it. Local DNR officials, who knew for years there was a huge poaching problem, had set up sting operations that were largely ineffective. So members of the community and I began tossing out ideas. In 1999, the Black Lake Chapter of Sturgeon for Tomorrow (SFT) incorporated as a 501(c3) nonprofit organization. The mission: To assist fisheries managers in the rehabilitation of lake sturgeon. That year, we coordinated the first annual Sturgeon Guarding Program, modeled after the successful program at Wisconsin’s Lake Winnebago. We began recruiting volunteers to stand watch and camp along the Black River during the spawning run to protect the sturgeon from would-be poachers. Boy Scout troops, Vietnam veterans, volunteer off-duty National Guardsmen, sportsmen and women, retirees, and people from throughout the Great Lakes Basin and Canada volunteered to patrol the area around the clock all year round. Today, about 350 volunteers contribute 3,800 hours along the river each year as a deterrent to poachers. The sturgeon migration is an amazing spectacle. We can identify sturgeon that have been tagged before when they come up the river. Although the literature says this return should happen every two to four years, we see some males returning to the river every year, and they are contributing to reproduction. We have also seen some previously tagged females. Some of these are just under seven feet in length. The annual spawning run is a wonder of nature at its finest. It is spectacular to see the sturgeon in all its glory. You look down from the cliff, look up and see eagles,

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and hear the rippling of the water. When sturgeon are spawning, you can hear them thrash and literally feel the ground shake. Since so many people were out on the river during spawning, it made sense to conduct research, compile data, and learn more about the reproductive capacity and early life history of the spawning population. Little was known about the spawning population of the Black Lake sturgeon. In 2000, Central Michigan University took on the project. In recent years, Michigan State University and the state Department of Natural Resources have directed the research. Researchers net and tag the adults to collect population data, and small fin samples are analyzed to determine genetic diversity. Larval sampling has quantified natural reproduction. The naturally produced larvae are collected, transferred, and reared at the streamside-rearing facility, then released after three months. Eggs and milt are also cultured to maximize production. There have been over 41,000 sturgeon fingerlings released into Black, Burt, and Mullett lakes since 2000. The Tower-Kleber Limited Partnership owns the dam and property on which the hatchery operates. The DNR leases the facility. With the leadership of Huron Pines and the hands of dozens of SFT volunteers, habitat improvements and streamside interpretive signage have been implemented to preserve some of the last known spawning habitat in northern Michigan. Guided eco-tours to see the spawning sturgeon and hatchery tours have proven to be highly popular. Meanwhile, SFT and collaborators are developing interpretive programming, including a visitors pavilion near the hatchery to expand outreach and education. Our annual banquet, golf scramble, memberships, and contributions primarily fund these initiatives, and grants have funded university research. Today, Sturgeon for Tomorrow has a vision of delisting the sturgeon from threatened status in the state and creating a world-class fishery. Sturgeon populations must be closely assessed and managed. States, tribes, universities, and nongovernmental organizations should collaborate to reestablish healthy, sustainable populations. Management plans will be developed with input from all stakeholders and then implemented. Since 2010, the LSRS has been under revision. Rehabilitation efforts, including biological research and strict regulations with stiff penalties for violators, should ensure there will be sturgeon populations for future generations to enjoy. But while sturgeon are being aided by these actions, people must remember that since sturgeon are such slow-growing, long-lived fish, it will be many years before we see populations restored to the levels resembling anything experienced in the past. Through our outreach programs, we will continue to educate, engage, and mobilize diverse constituencies to treasure the majestic lake sturgeon and our natural world and not take them for granted. We must all work together to keep our great lakes—great.

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In my view, conservation embraces values that spring from an early and profound childhood experience in nature. Sense of place deepens. The common thread of my childhood is that I spent a lot of time outdoors because that’s where the things that interest me live. It’s where I developed many of my core values. Understanding the threats to the places I love and where I’d gained self-assurance made me a conservationist and convinced me that connecting people with nature, especially children, is one of the major tasks to winning this great war, and standing up to crimes against nature. We must learn how best to live sustainably and in harmony with nature, and to pass this rich heritage onto future generations. The sturgeon is the oldest and largest fish in the Great Lakes, the elder statesman of Michigan fish species. It is an ancient fish. Sturgeon are our ancestors, a living fossil. Their sheer existence and wonderful display every spring in the river, and in the winter the ice-fishing heritage, have become deeply entrenched in our culture. Like the sturgeon, we are inextricably an element of the ecosystem.

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The Relationship between Lake Sturgeon Life History and Potential Sensitivity to Sea Lamprey Predation

The lake sturgeon Acipenser fulvescens, a species native to the Laurentian Great Lakes, has a unique life history. Like other sturgeons, lake sturgeon are a slow-growing, long-lived species with delayed maturation; first spawning for males typically occurs between ages 12 and 15, while females become mature between ages 18 and 27. In addition, lake sturgeon spawn intermittently, with females spawning only once every four to nine years and males spawning every one to three years (Roussow 1957; Scott and Crossman 1973; Fortin, Dumont, and Guénette 1996; Bruch 1999; Bruch, Dick, and Choudhury 2001). Although these life-history traits are advantageous for buffering against extreme environmental conditions, they increase susceptibility to human-induced mortality and the negative effects of aquatic invasive species (Hay-Chmielewski and Whelan 1997; Auer 2004). Despite supplemental stocking, efforts to improve water quality, and permitted harvest reductions, lake sturgeon populations have been slow to recover from their imperiled state throughout the Great Lakes basin (Welsh et al. 2008). This slow recovery is not surprising, considering the great age-to-maturity characteristic of lake sturgeon. However, another potential factor that may be currently impeding lake sturgeon rehabilitation is aquatic invasive species. For example, the invasive parasitic n

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sea lamprey Petromyzon marinus preys on and kills lake sturgeon in confined tank experiments (Patrick, Sutton, and Swink 2009). The objective of this chapter is to review the effects of aquatic invasive species on Great Lakes lake sturgeon populations, with an emphasis on the potential effects of sea lamprey predation on both the decline and slow rate of recovery of lake sturgeon.

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Aquatic Invasive Species in the Great Lakes The introduction and establishment of undesirable, nonnative plant and animal species is one of the greatest threats to the future of the Great Lakes and to species and community conservation worldwide (Finster 2007; Thresher 2008). The Great Lakes have been subject to invasion by nonnative species since European settlement (Mills et al. 1993). Development of the basin, including timber harvest, agriculture, hydropower development, and canal construction, as well as invasion of marine organisms, fish stocking, and overharvest, have resulted in rapid changes in what was once a simple, slowly evolving ecosystem (Fetterolf 1980). At least 182 nonindigenous species have been introduced into the Great Lakes since 1840, with over 40 percent of these invaders occurring after the opening of the St. Lawrence Seaway in 1959 (Ricciardi 2006). The Great Lakes have the highest rate of nonnative species invasions and introductions recorded in any freshwater ecosystem. Since 1960, the invasion rate is estimated to be 1.8 species per year, equivalent to one new invader being discovered every 28 weeks. Though not all nonnative species jeopardize the Great Lakes ecosystem (some, such as the Pacific salmon, have been intentionally introduced to support commercial or recreational fisheries), those that are considered to be injurious are capable of inflicting significant damage to the environment and the economy (Finster 2007). The degree to which native fish and their habitats are affected by injurious aquatic invasive species depends on the life-history traits of invading and of native species, as well as the ability of the ecosystem to withstand change (Ricciardi 2006). To date, little is known about the injurious effects of aquatic invasive species on lake sturgeon.

The Voracious Sea Lamprey Of all the aquatic invasive species thought to impact lake sturgeon abundance, the sea lamprey poses the most serious threat. The sea lamprey is similar in appearance to the American eel Anguilla rostrata, but lacking a jaw and paired fins, and having seven pairs of gill pouches instead of the usual gill structure of bony fishes. The

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life cycle of the sea lamprey involves distinctively different larval and adult feeding phases. Whereas larval lampreys, called ammocoetes, feed primarily on organic detritus, adults in their predacious stage feed on the blood of other fishes (their hosts) (Lowe, Beamish, and Potter 1973; Sutton and Bowen 1994). After spending an extended larval phase (3 to 10 or more years) buried in the sediment of streams, sea lamprey larvae metamorphose into parasites, developing eyes and teeth, and enter the lakes to feed. The period of metamorphosis between these two feeding phases is called “transformation,” and lampreys at this stage are referred to as “transformers,” or macropthalmia because of their large eyes (Applegate 1950). Each summer and fall, one age group of parasitic sea lamprey is actively feeding. The next spring, this group migrates back to tributary streams to spawn and die. When feeding on a fish, sea lamprey attach to their host with their suction-cup-like mouth, called an oral disc, which is armed with concentric rows of sharp teeth. In the center of their mouth is a sharply toothed tongue, which is used to rasp holes in the flesh of their prey to feed on its blood and tissue. Sea lampreys are also equipped with a buccal-gland system that secretes anticoagulant, called lamphredin, to thin the blood and tissues of its host. Because the average sea lamprey grows from an approximately 10 g transformer to a 150 g adult in only 18 months, one can imagine how much blood must be consumed and, in turn, how many fish a lamprey must kill, during its parasitic life stage. A single sea lamprey can kill an estimated 18 kg of fish or more during its life (Farmer 1980).

Origins of the Sea Lamprey in the Great Lakes The sea lamprey is native to the Atlantic Ocean and; although its origin in the Great Lakes, Lake Champlain, and the large lakes in New York has been long debated, landlocked freshwater populations also exist in these lakes (e.g., Bryan et al. 2005). Most proponents support one of three zoogeographic scenarios explaining the possible origins of freshwater sea lamprey populations. Freshwater populations in Lake Ontario and Lake Champlain may have arisen from (1) natural migrations from marine sources in the St. Lawrence River or the Atlantic coastal drainages; (2) as a result of historical bio-geographic events such as periods of marine submergence (i.e., the incursion of the Champlain Sea [11,500–13,000 years ago]); or (3) by recent invasion from the Hudson River through canals built for transportation purposes in the nineteenth and twentieth centuries (i.e., Erie and Champlain canals) (Bryan et al. 2005). Proponents of the theory that sea lamprey entered Lake Ontario by way of natural migration maintain that this species is indigenous to Lake Ontario and

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its tributaries (Hubbs and Lagler 1947; Bigelow and Schroeder 1948; Rostlund 1952; Lawrie 1970; Scott and Crossman 1973; Bailey and Smith 1981; Smith 1985; Daniels 2001; Waldman et al. 2004). These supporters hold that the discontinuous distribution between the freshwater populations in the New York Finger Lakes and the Hudson River population is evidence that sea lamprey are native to Lake Ontario and its tributaries (Mills et al. 1993). However, DeKay (1842) found sea lamprey as far upstream in the Hudson River as Albany, New York (Mills et al. 1993). Modeling results of Bryan et al. (2005) also support the hypothesis that sea lamprey migrated to the freshwater lakes via the St. Lawrence River. For example, the presence of a rare genetic composition (allele) in landlocked Lake Ontario and Lake Champlain populations is evidence that populations in these lakes have been separated from the putative North American progenitor populations on the Atlantic Coast for a considerable period of time. Another argument supporting the hypothesis that lampreys are native to Lake Ontario is that the presumed poor water quality conditions of nineteenth-century navigation canals, in combination with the impediment of numerous locks, make canals an unlikely vector for lamprey dispersal (Daniels 2001). The conclusions regarding endemnicity of the Lake Ontario population may be overstated (Eshenroder 2009). Because observations of sea lamprey in Lake Ontario were not reported until the 1830s, a second belief is that the sea lamprey is not native to Lake Ontario, but rather invaded the lake in historical times after the construction and opening of the Erie Canal in the early 1800s (Aron and Smith 1971; Emery 1985; Mandrak and Crossman 1992; Mills et al. 1993; Smith 1995). Most support for the invasion-by-canal hypothesis lies in the disbelief that sea lampreys could have been historically present, but gone unobserved in Lake Ontario until after the construction of the Erie Canal (Eshenroder 2009). Even if the sea lamprey is indeed native to Lake Ontario, Niagara Falls served as an insurmountable natural barrier that effectively prohibited the migration of the parasite into the remaining Great Lakes. It is well established that the construction of the Welland Ship Canal in 1829 provided the necessary thoroughfare for the sea lamprey to gain access to Lake Erie, and eventually, the upper Great Lakes. The sea lamprey now inhabits all five of the Great Lakes, where it is considered by U.S. and Canadian management agencies to be an invasive species. Even if sea lampreys are native to Lake Ontario, they have proven to be capable of exhibiting characteristics of invasive species when predator-prey dynamics are disturbed.

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The Effects of Sea Lamprey Predation on Fish Communities Sea lamprey were first reported in Lake Erie in 1921, Lake St. Clair in 1934, Lake Michigan in 1936, and Lake Huron in 1937 (Dymond 1922; Trautman 1949; Smith and Tibbles 1980). The rapids and Soo Locks at the lower end of Lake Superior apparently hindered their invasion; the first confirmed record in Lake Superior was a parasite near Isle Royale in 1946 (Applegate 1950). Sea lampreys and lake trout Salvelinus namaycush coexisted in Lake Ontario for a long time. Annual production of lake trout in Lake Ontario did not appear to decline abruptly as it did in the upper Great Lakes; annual production from 1867 to 1940 averaged 164.7 MT and declined to only 14.5 MT in the years 1941–1967 (Smith 1971). Reports of parasitic-phase lampreys in Lake Erie were sporadic for the first decade following their initial appearance. The slow rate of establishment in Lake Erie is generally attributed to the limited number of spawning streams with suitable habitat, poor water conditions, and a limited forage base of desirable prey species until more recent times (Lawrie 1970; Pearce et al. 1980). The establishment of sea lamprey in the upper three Great Lakes (Huron, Michigan, Superior), however, was followed by an abrupt and continual decline in commercial lake trout production that was intensified by elevated fishing mortality (Lawrie 1970). The sea lamprey invasion of Lake Erie and the upper Great Lakes was also followed by extensive changes in the fish communities (Schneider et al. 1996). Prior to sea lamprey invasion, competition, especially in offshore areas, was quite low (Smith 1971). During this time, there were only two apex (i.e., top) predators in the deep waters of the lakes, lake trout and burbot Lota lota. All Great Lakes fishes are susceptible to attack by a sea lamprey, but some species are favored over others (Farmer and Beamish 1973). Host habitat, body size, and scale covering influence susceptibility. Sea lampreys tend to occupy cool water and are believed to actively select the largest available host. Thus, large deepwater species, including lake trout, burbot, whitefish, and ciscoes Coregonus spp. were the first to decline in the upper Great Lakes (Smith 1972). The combined effects of sea lamprey parasitism, overexploitation, and other variables such as habitat degradation, led to the ultimate loss of lake trout from lakes Ontario, Erie, Huron, and Michigan, leaving behind only a few populations in Lake Superior, and small remnant populations in Lake Huron (Krueger and Ebener 2004). Although sea lamprey predation was well documented on the aforementioned Great Lakes fishes, it is probable that lake sturgeon were also preyed upon, particularly when abundance of preferred hosts was low. Reports by regional biologists of lake sturgeon with sea lamprey marks and wounds support this hypothesis. The

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question that remains is whether these attachment events ever lead to mortality in lake sturgeon.

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Controlling Sea Lamprey Populations Because the Great Lakes intersect the political boundary between Canada and the United States, sea lamprey management is a complex endeavor that falls under the jurisdiction of two nations, eight states, one province, and several tribal and First Nation groups. To facilitate coordinated, binational fisheries management, the federal governments of Canada and the United States negotiated and ratified the Convention on Great Lakes Fisheries in 1954 and 1955 respectively. The Great Lakes Fishery Commission was formed in 1956 pursuant to the agreement (www. glfc.org). Since its inception, the goals of the commission have been to control sea lamprey populations in the Great Lakes basin; to coordinate, communicate, and conduct research; and to improve the coordination of fisheries management agencies. The commission works with Fisheries and Oceans Canada, the U.S. Fish and Wildlife Service, the U.S. Army Corps of Engineers, the U.S. Geological Survey, and universities throughout the Great Lakes basin to achieve its sea lamprey control and management goals. The control program operates under an integrated pest management approach and consists of four main components: assessment, barriers, trapping, and lampricides. The control program has historically relied heavily on chemical methods of control, primarily 3-trifluoromethyl-4-nitrophenol (TFM), costing more than $18 million annually. However, the development of alternative control methods was a keystone of the commission’s vision for the past decade and the resulting effort has set the stage for deployment of one or more new alternate control methods in this decade. The sea lamprey control program, in association with stocking, has permitted the successful rebuilding of lake trout populations in Lake Superior and has reduced the impact on other native fishes in all of the Laurentian Great Lakes.

Sea Lamprey and Lake Sturgeon Little is known about the extent or the effects of sea lamprey predation on lake sturgeon in the wild. The unique morphological traits of lake sturgeon make it probable that the effects of an attack vary greatly compared to effects on common teleost fishes. First, lake sturgeon are armed with rows of hard, bony plates called scutes (a defense mechanism evolved to provide protection from predation in

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general), which intuitively seem difficult to penetrate. Scutes are much sharper and more plentiful on smaller, juvenile sturgeon and become more rounded and less prevalent as the fish matures. The sharp scutes offer more protection from predation to young sturgeon when their blood volume is less and their chance of dying due to a sea lamprey attack is increased. Sea lamprey are thought to be a size-selective predator, choosing the largest available host on which to feed (Farmer 1980; Cochran 1985; Swink 1990). Increased body size means increased blood volume; however, increased body size does not necessarily allow for better survival following a sea lamprey attack. Collection of lake trout carcasses in Lake Ontario from 1982 to 1985 (Bergstedt and Schneider 1988) confirmed that sea lamprey predominantly kill large lake trout when the trout are abundant, because of the increased frequency of multiple attachments by lamprey for large hosts (Swink 1990). Scott and Crossman (1973) recounted an observation of a 70 kg lake sturgeon caught in the Bay of Quinte in October 1969 that, when captured, had 15 large sea lampreys attached to its body and bore scars from several previous attacks. The sturgeon was reported to have been sluggish and in poor body condition, probably due to the loss of blood from attacks. Recent accounts of sea lamprey attacks on lake sturgeon are comparatively rare, in part because abundance of other host species, such as lake trout, has rebounded and abundance of sturgeon is currently low. However, field observations do indicate that lake sturgeon still occasionally serve as a host for parasitic sea lampreys. For example, the rate of sea lamprey wounding on subadult and adult lake sturgeon in the St. Marys River, Michigan-Ontario, from 2000 to 2002 was estimated to be 23 percent (T. Sutton, unpublished data). Sea lamprey predation likely continues to be a threat to the future restoration of lake sturgeon. Results from modeling simulations conducted by Sutton et al. (2003) gave reason to believe that sea lamprey predation could affect lake sturgeon rehabilitation more than initially believed. In response to concerns for lake sturgeon in the Sturgeon River, Michigan, the sea lamprey control program established a protocol in 1989 to protect larval lake sturgeon against mortality caused by lampricide application. Under this protocol, the maximum TFM treatment was reduced to 1.3 times the minimum lethal concentration (MLC). Allowable treatment levels were further reduced in 1998 after results of laboratory toxicity experiments showed larval and age-0 lake sturgeon less than 100 mm in length were particularly sensitive to TFM, the primary chemical used for sea lamprey control in the Great Lakes (Johnson, Weisser, and Bills 1999; Boogaard, Bills, and Johnson 2003). Boogaard, Bills, and Johnson (2003) suggested that the prohibition of treatment for at least 90 days after lake sturgeon spawning would allow sturgeon to reach this length. As a result, allowable treatment levels were not to exceed 1.0 times the MLC for TFM or 1.2 times the MLC for TFM/2% niclosamide and application could not

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take place until August 1. This protocol is referred to as the “No-Observable-Effect Concentration” (NOEC) protocol or the “sturgeon” protocol. However, modeling results from Sutton et al. (2003) suggested that this protocol may be counterproductive. Less effective lampricide treatments at lower concentrations may result in more predatory lampreys and, consequently, greater levels of parasitism-induced mortality of adult lake sturgeon and other host species. Throughout the 50-year history of sea lamprey control, only minimal numbers of dead age-0 lake sturgeon have been observed during treatments of streams with lampricides. More specifically, during 1,800 U.S. stream treatments from 1959 to 2000, only 10 dead age-0 lake sturgeon were observed in over 1,600 assessment collections (Adair and Young 2004). Similarly, only 13 dead age-0 sturgeon have been observed during the history of treatments of Canadian streams, with the last observation occurring in 1997 (M. Steeves, Department of Fisheries and Oceans–Sea Lamprey Control Centre, personal communication). A field study was conducted by T. Pratt, M. Steeves, and L. O’Connor (Department of Fisheries and Oceans–Sea Lamprey Control Centre) in 2008 to assess survival of age-0 lake sturgeon subjected to regular and sturgeon protocol TFM treatments (M. Steeves, DFO Sault Ste. Marie, personal communication). Age-0 lake sturgeon (mean length 78.7 mm; range 57–101 mm; hatchery and native origin) were caged at three sites: the upper Mississagi River, the lower Mississagi River, and the Spanish River (M. Steeves, DFO Sault Ste. Marie, personal communication). The upper portion of the Mississagi River was treated using the sturgeon protocol (1.2 times MLC), whereas the lower portion was treated using a boost to the normal application strategy (1.5 times MLC). Sturgeon were also caged in the Spanish River, which did not undergo treatment during the study period, to control for the effects of stress caused by the caging on survival. Overall, observed lake sturgeon mortality was extremely low (3 percent) and was equally spread among the three treatments. Sturgeon that did die were all notably smaller in size than the rest of the study fish and presumably died from starvation. Results of this study suggest that lake sturgeon mortality related to lampricide treatment is low. These results were surprising given that laboratory toxicity tests and on-site bio-assays have demonstrated that early life stages of lake sturgeon are sensitive to lampricide treatments (Johnson, Weisser, and Bills 1999; M. Boogard, U.S. Geological Survey, unpublished data). It is important to note that lampricide toxicity varies in relation to environmental factors, including stream pH and alkalinity, and may differ at other sites enough to warrant caution (Le Maire 1961; Dawson, Cumming, and Gilderhuis 1975; Marking and Olson 1975). As a result of these preliminary findings, the Great Lakes Fishery Commission has funded two additional years of field research at 12 additional rivers to further investigate the lethality of lampricide treatments on lake sturgeon in the field

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(M. Steeves, DFO Sault Ste. Marie, personal communication) and managers should err on the side of caution in the meantime. A trade-off exists between protecting larval lake sturgeon from lampricide toxicity and protecting juvenile and adult lake sturgeon from sea lamprey parasitism. The Sutton et al. (2003) model suggested that parasitism-induced mortality by sea lampreys could more negatively influence abundance of subadult and adult lake sturgeon, recruitment to age 1, and reproductive potential than lampricide-related mortality on early life stages. If this were true, the survival of subadult and adult lake sturgeon is more crucial than recruitment of early life stages for long-term population persistence. Obviously, this is only a model and models are based on unknowns, so further study is still necessary to investigate the validity of this prediction. The parasitism-related mortality estimates used in the sublethal model were based on the results of laboratory experiments that used lake trout as the host. As described above, the different life-history traits between lake trout and lake sturgeon likely mean that lethality would differ between these species. To address this uncertainty, a laboratory-based study was conducted by Patrick, Sutton, and Swink (2009). The goal of this study was to estimate the lethality of a single sea lamprey attack on four size-classes of lake sturgeon. A series of 55 experimental trials were conducted whereby one sea lamprey and one lake sturgeon were placed into a tank together after being weighed and measured. Sea lamprey were allowed to feed on lake sturgeon until detachment or mortality of the host or parasite, and each sea lamprey was only permitted one attachment per host. Blood samples were also collected from the lake sturgeon before and after each trial to determine changes in blood chemistry after a lamprey attack. Changes in lake sturgeon growth and blood chemistry were also monitored; the experimental design is described in Patrick, Sutton, and Swink (2009). Twenty-six percent (n = 14) of the lake sturgeon attacked by a sea lamprey (n = 55) died. The study showed that even a single sea lamprey attack can significantly affect the survival of lake sturgeon, particularly those less than 650 mm in fork length, probably because of their lower host-to-lamprey weight ratio (Patrick, Sutton, and Swink 2009). Changes in lake sturgeon hematology after a sea lamprey attack indicated that direct mortalities resulted from the onset of acute anemia, or the rapid loss of blood. Hosts that lose an amount of blood equivalent to their blood volume (i.e., fish that “bleed out”) generally die within two days, whereas hosts that lose 10 percent or less daily can replace those losses and survive (Farmer, Beamish, and Lett 1977). As a result, host survival depends on percentage of blood loss relative to total blood volume (Edsall and Swink 2001). Of fish that do not die directly from blood loss, many suffer from decreases in body condition and may develop fungal infections due to damage to the skin and can die indirectly from an attack.

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The 26 percent sturgeon mortality observed in the Patrick, Sutton, and Swink (2009) study was lower than that reported for other species, such as lake trout or burbot, in containment studies with similar study designs (Swink and Hanson 1989; Swink and Fredricks 2000; Bergstedt, Schneider, and O’Gorman 2001; and Swink 2003). For instance, mortality of lake trout, rainbow trout Oncorhynchus mykiss, and burbot attacked by a sea lamprey ranged from 40% to 65% (Swink and Hanson 1989; Swink 1990, 1993; Swink and Fredricks 2000). However, the Sutton et al. (2003) simulation model used an estimated range of mortality of 0–22 percent. The results from the laboratory study show that sea lamprey-induced mortality of lake sturgeon may be greater than 22 percent, especially for smaller ( 600 mm in total length) are; however, capable of using zebra mussels as a food source. Because zebra mussels supplement the benthic (i.e., bottom dwelling) food web at the expense of the pelagic (i.e., open water) web, a net gain may occur for sturgeon populations, but this increase in food depends on the successful growth of juveniles to a size category that can use zebra mussels or on behavioral shifts to use the macroinvertebrates inhabiting zebra mussel colonies (McCabe et al. 2006). Managers should consider this change in benthic community when developing lake sturgeon rehabilitation strategies.

Evaluating Effects of Aquatic Invasive Species on Lake Sturgeon The effects of parasitic-phase sea lamprey on Great Lakes fishes are monitored in several ways. One way is by recording sea lamprey wounds on host fishes to determine the extent of parasitism. Sea lamprey wounding data, such as the wound size and the number of each type and stage of wound, are often collected ancillary to other stock assessment information for a variety of fishes in the Great Lakes. Wounding rates, such as the number of observed wounds per 100 fish or the number of observed wounds per unit observation effort, have been shown to be directly proportionate to sea lamprey attack rates (Eshenroder and Koonce 1984).

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A dichotomous key for use in the classification of sea lamprey wounds on lake sturgeon was developed as part of the Patrick, Sutton, and Swink 2009 study (Patrick, Sutton, and Swink 2007). This key was developed based on the classification system used for lake trout for over 25 years (King 1980). The key was revised by Ebener, King Jr., and Edsall (2006) to include images of other Great Lakes species, such as lake whitefish Coregonus clupeaformis, cisco Coregonus artedii, walleye Sander vitreus, Chinook salmon Oncorhynchus tshawytscha, and white sucker Catostomus commersonii. A unique classification system for lake sturgeon was needed because this species differs morphologically from other Great Lakes species, especially in that lake sturgeon lack scales, which makes a difference in healing rates. For example, laboratory observations showed that Type-A wounds (those that create a pit in the skin of the fish) in lake sturgeon often do not penetrate the musculature because of their thick skin and scutes, and result in a more rapid healing rate (Patrick, Sutton, and Swink 2007). The use of the key in the field should provide increased data collection consistency for sea lamprey wounds on lake sturgeon and help to determine changes in rates of predation on lake sturgeon in the wild. More research is needed on methods to prevent the spread of exotics, and yet preserve migration corridors for native fishes (Auer 2004). The overarching goal of the Great Lakes Fishery Commission’s barrier and trapping research theme area is to “transition from using barriers to deny spawning-phase sea lampreys access to spawning habitat, to using barriers to block and selectively trap sea lampreys, and ultimately, to the development and deployment of barriers that are transparent to non-target fishes and of novel, barrier-free traps effective enough for control purposes” (McLaughlin et al. 2007). Much progress has been made to date, with many barriers today having built-in traps to remove sea lampreys and downstream pools to assist the passage of jumping fish. Maximizing sea lamprey control, while minimizing effects on nontarget native species, is a challenging balance. For example, many Lake Superior stakeholders are anxious to remove or alter the existing hydroelectric dam at the mouth of the Black Sturgeon River in Canada to enhance walleye spawning. Dam removal would have the added benefit of reestablishing connectivity between the Great Lakes and their tributaries for remnant Lake Sturgeon populations. However, this dam limits production to the lower 17 km of the 100 km river in a waterbody that could be an enormous source of sea lamprey to Lake Superior, so management options need to be very carefully considered Limiting the area of production reduces the amount of stream that needs to be chemically treated. Clearly, dam removal would increase the monetary requirements of the sea lamprey control program, as lampricide treatment frequency and magnitude would need to be increased. It is also likely that dam removal would

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negatively affect the ecology of Lake Superior because more parasitic phase sea lamprey would enter the population and prey on the fishes. Yet maintaining the dam presents challenges for migrating fishes. The Great Lakes Fishery Commission sponsored a two-day workshop in Turners Falls, Massachusetts, in January 2009, focused on addressing the unique passage issues associated with walleye and lake sturgeon. The potential exists for the Black Sturgeon Dam to become a valuable test case for evaluating fish passage issues.

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Conclusions Mitigating the effects of aquatic invasive species is an important aspect of lake sturgeon rehabilitation. Currently, the effects of sea lamprey predation on lake sturgeon survival are based on laboratory studies and modeling results. Mortality rates in the wild may differ owing to a variety of factors, including increased stress in a laboratory setting (e.g., fish handling) which may lead to an increase in secondary infection, and the presence of preferred alternative hosts (i.e., salmonines and coregonines) in the wild. In the Great Lakes, wounds on lake sturgeon are rarely recorded in comparison to wounds on other host species. The possibility exists, however, that wounds are not frequently observed because sea lamprey attacks on lake sturgeon result in mortality. If a lake sturgeon were to die from a sea lamprey attack (or multiple sea lamprey attacks), it would be unnoticeable. It is also equally possible that sea lamprey predation on lake sturgeon is not an issue for lake sturgeon, either because it doesn’t occur frequently enough in the wild for it to be a problem or, if it does occur, it rarely results in mortality. In any case, laboratory studies should be validated in the field to provide a better understanding of the effects of sea lamprey predation on lake sturgeon. The collection of wounding data using the newly developed dichotomous key for sea lamprey wounds on lake sturgeon will help address these uncertainties. Although sea lamprey may not be feeding extensively on lake sturgeon today because alternative preferred host species are abundant in the Great Lakes, sea lamprey predation may have contributed to mortality in the past when lake trout abundance was low. Sea lamprey parasitism should be considered a potential threat to lake sturgeon survival, especially in the context of modified sea lamprey control treatment protocols that strive to reduce use of lampricides. Further investigation is also needed to determine the effects of other aquatic invasive species on lake sturgeon rehabilitation. The importance of considering the effects of aquatic invasive species in lake sturgeon rehabilitation plans will become increasingly important if the rapid rate of new species introduction into the Great Lakes continues.

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REFERENCES

Adair, R. A., and R. J. Young. 2004. Standard operating procedures for application of lampricides in the Great Lakes Fishery Commission integrated management of sea lamprey (Petromyzon marinus) control program. USFWS Sea Lamprey Control, Marquette, Michigan, Special report 92-001.4. Applegate, V. C. 1950. Natural history of the sea lamprey (Petromyzon marinus) in Michigan. U.S. Fish and Wildlife Service Special Scientific Report—Fisheries 55. Aron, W. I., and S. H. Smith. 1971. Ship canals and aquatic ecosystems. Science 174:13–20. Auer, N. A. 2004. Conservation. In: Sturgeons and paddlefish of North America. G. T. O. LeBreton, F. W. H. Beamish, and R. S. McKinley, eds. Kluwer Academic Publishers. Bailey, R. M., and G. R. Smith. 1981. Origin and geography of the fish fauna of the Laurentian Great Lakes basin. Canadian Journal of Fisheries and Aquatic Sciences 38:1539–1561. Benson, A. C., T. M. Sutton, R. F. Elliott, and T. G. Meronek. 2006. Biological attributes of age-0 lake sturgeon in the lower Peshtigo River, Wisconsin. Journal of Applied Ichthyology 22:103–108. Bergstedt, R. A., and C. P. Scheinder. 1988. Assessment of sea lamprey (Petromyzon marinus) predation by recovery of dead lake trout (Salvelinus namaycush) from Lake Ontario, 1982–85. Canadian Journal of Fisheries and Aquatic Sciences 45:1406–1410. Bergstedt, R. A., C. P. Schneider, and R. O’Gorman. 2001. Lethality of sea lamprey attacks on lake trout in relation to location on the body surface. Transactions of the American Fisheries Society 130:336–340.

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Bigelow, H. B., and W. C. Schroeder. 1948. Cyclostomes. In: Fishes of the Western North Atlantic. Sears Foundation for Marine Research Memoir 1. Boogaard, M. A., T. D. Bills, and D. A. Johnson. 2003. Acute toxicity of TFM and TFM/ Niclosamide mixture to selected species of fish, including lake sturgeon (Acipenser fulvescens) and mudpuppies (Necturus maculosus), in laboratory and field exposures. Journal of Great Lakes Research 29 (Suppl. 1): 529–541. Bruch, R. M. 1999. Management of lake sturgeon on the Winnebago System—long-term impacts of harvest and regulations on population structure. Journal of Applied Ichthyology 15:142–152. Bruch, R. M., T. A. Dick, and A. Choudhury. 2001. A field guide for the identification of stages of gonad development in lake sturgeon, Acipenser fulvescens Rafinesque, with notes on lake sturgeon reproductive biology and management implications. Publication of Wisconsin Department of Natural Resources. Oshkosh and Sturgeon for Tomorrow. Bryan, M. B., D. Zalinski, B. Filcek, S. Libants, W. Li, and K. T. Scribner. 2005. Patterns of invasion and colonization of the sea lamprey (Petromyzon marinus) in North America as revealed by microsatellite genotypes. Molecular Ecology 14:3757–3773. Chotkowski, M., and E. Marsden. 1999. Round goby and mottled sculpin predation on lake trout eggs and larvae: Field predictions from laboratory experiments. Journal of Great Lakes Research 25:26–35. Christie, W. J., and D. P. Kolenosky. 1980. Parasitic phase of the sea lamprey (Petromyzon marinus) in Lake Ontario. Canadian Journal of Fisheries and Aquatic Sciences 37:2021– 2038. Cochran, P. A. 1985. Size-selective attack by parasitic lampreys: Consideration of alternate null

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hypotheses. Oecologia 67:137–141. Daniels, R. A. 2001. Untested assumptions: The role of canals in the dispersal of sea lamprey, alewife, and other fishes in the eastern United States. Environmental Biology of Fishes 60:309–329. Dawson, V. K., K. B. Cumming, and P. A. Gilderhuis. 1975. Laboratory efficacy of 3-trifluoromethyl-4 nitrophenol (TFM) as a lampricide. U.S. Fish and Wildlife Service, Investigations in Fish Control 63. Dymond, J. R. 1922. A provisional list of the fishes of Lake Erie. University of Toronto Studies. Publications of the Ontario Fisheries Research Laboratory 57-73. Ebener, M. P., E. L. King, Jr., T. A. Edsall. 2006. Application of a dichotomous key to the classification of sea lamprey marks on Great Lakes fish. Great Lakes Fishery Commission Miscellaneous Publlication 2006-02. Http:www.glfc.org/pubs/pub.htm#misc. Edsall, C. C., and W. D. Swink. 2001. Effects of nonlethal sea lamprey attack on the blood chemistry of lake trout. Journal of Aquatic Animal Health 13:51–55. Emery, L. 1985. Review of fish species introduced into the Great Lakes, 1819–1874. Great Lakes Fishery Commission Technical Report No. 45. Great Lakes Fishery Commission. Eshenroder, R. L. 2009. Comment: Mitochondrial DNA analysis indicates sea lampreys are indigenous to Lake Ontario. Transactions of the American Fisheries Society 138:1178–1189. Eshenroder, R. L., and J. F. Koonce. 1984. Recommendations for standardizing the reporting of sea lamprey wounding data: A report from the Ad Hoc Committee. Great Lakes Fishery Commission. Special Publication 84-1. Farmer, G. J. 1980. Biology and physiology of feeding in adult lamprey. Canadian Journal of Fisheries and Aquatic Sciences 37:1751–1761.

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Farmer, G. J., and F. W. H. Beamish. 1973. Sea lamprey (Petromyzon marinus) predation on freshwater teleosts. Journal of Fisheries Research Board of Canada 30:601–605. Farmer, G. J., F. W. H. Beamish, and P. F. Lett. 1977. Influence of water temperature on the growth rate of the landlocked sea lamprey (Petromyzon marinus) and the associated rate of host mortality. Journal of the Fisheries Research Board of Canada 34:1373–1378. Fetterolf, C. M., Jr. 1980. Why a Great Lakes Fishery Commission and why a Sea Lamprey International Symposium. Canadian Journal of Fisheries and Aquatic Sciences 37:1588– 1593. Finster, J. L. 2007. Investigating injurious species introductions as environmental crimes. M.S. thesis, Department of Fisheries and Wildlife, Michigan State University. Fortin, R., P. Dumont, and S. Guénette, 1996. Determinants of growth and body condition of lake sturgeon (Acipenser fulvescens). Canadian Journal of Fisheries and Aquatic Sciences 53:1150–1156. Harkness, W. J. K., and J. R. Dymond. 1961. The lake sturgeon: The history of its fishery and problems of conservation. Ontario Department of Lands and Forests, Fish and Wildlife Branch, Maple. Hay-Chmielewski, E. M., and G. E. Whelan. 1997. State of Michigan lake sturgeon rehabilitation strategy. Michigan Department of Natural Resources, Fisheries Special Report 18. Herbert, P. D. N., B. W. Muncaster, and G. L. Mackie. 1989. Ecological and genetic studies on Dreissena polymorpha (Pallas): A new mollusk in the Great Lakes. Canadian Journal of Fisheries and Aquatic Sciences 46:1587–1591. Hubbs, C. L., and K. F. Lagler. 1947. Fishes of the Great Lakes region. Cranbrook Institute of

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Science Bulletin 26. Johnson, D. A., J. W. Weisser, and T. D. Bills. 1999. Sensitivity of lake sturgeon (Acipenser fulvescens) to the lampricide 3-trifluoromethyl-4-nitrophenol (TFM) in field and laboratory exposures. Great Lakes Fishery Commission Technical Report 62. Jude, D. J. 2001. Round and tubenose gobies: 10 years with the latest Great Lakes phantom menace. Dreissena! (National Aquatic Species Clearinghouse, SUNY 11(4): 1–9, 12–14. King, E. L., Jr. 1980. Classification of sea lamprey (Petromyzon marinus) attack wounds on Great Lakes lake trout (Salvelinus namaycush). Canadian Journal of Fisheries and Aquatic Sciences 37:1989–2006. Krueger, C. C., and M. Ebener. 2004. Rehabilitation of lake trout in the Great Lakes: Past lessons and future challenges. In: Boreal shield watersheds: Lake trout ecosystems in a changing environment. J. Gunn, R. J. Steedman, and R. A. Ryder, eds. CRC Press. Lawrie, A. H. 1970. The sea lamprey in the Great Lakes. Transactions of the American Fisheries Society 99:766–775. Le Maire, E. H. 1961. Experiments to determine the effect of pH on the biological activity of two chemicals toxic to ammocoetes. Fisheries Research Board of Canada, Biological Report Series No. 690. Lowe, D. R., F. W. H. Beamish, and I. C. Potter. 1973. Changes in proximate body composition of the landlocked sea lamprey Petromyzon marinus (L.) during larval life and metamorphosis. Journal of Fish Biology 5:673–682. Lowe, E. A., J. E. Marsden, and W. Bouffard. 2006. Movement of sea lamprey in the Lake Champlain basin. Journal of Great Lakes Research 32:776–787. Mandrak, N. E., and E. J. Crossman. 1992. Postglacial dispersal of freshwater fishes into Ontario. Canadian Journal of Zoology 70:2247–2259.

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Marking, L. L., and L. E. Olson. 1975. Toxicity of the lampricide 3-trifluoromethyl-4-nitrophenol (TFM) to non-target fish in static tests. U.S. Fish and Wildlife Service, Investigations in Fish Control No. 60. McCabe, D. J., M. A. Beekey, A. Mazloff, and J. E. Marsden. 2006. Negative effect of zebra mussels on foraging and habitat use by lake sturgeon (Acipenser fulvescens). Aquatic Conservation: Marine and Freshwater Ecosystems 16:493–500. McLaughlin, R. L., A. Hallett, T. C. Pratt, L. M. O’Connor, and D. G. McDonald. 2007. Research to guide the use of barriers, traps, and fishways to control sea lamprey. Journal of Great Lakes Research 33 (special issue 2): 7–19. Mills, E. L., J. H. Leach, J. T. Carlton, and C. L. Secor. 1993. Exotic species in the Great Lakes: A history of biotic crises and anthropogenic introductions. Journal of Great Lakes Research 19:1–54. Mills, E. L., G. Rosenburg, A. P. Spidle, M. Ludyanskiy, Y. M. Pligin, and B. May. 1996. A review of the biology and ecology of the quagga mussel (Dreissena bugensis), a second species of freshwater Dreissenid introduced to North America. American Zoologist 36:271–286. Nichols, S. J., G. Kennedy, E. Crawford, J. Allen, J. French III, G. Black, M. Blouin, J. Hickey, S. Chernyák, R. Haas, and M. Thomas. 2003. Assessment of Lake Sturgeon (Acipenser fulvescens) spawning efforts in the lower St. Clair River, Michigan. Journal of Great Lakes Research 29(3): 383–391. Patrick, H. K., T. M. Sutton, and W. D. Swink. 2007. Application of a dichotomous key to the classification of sea lamprey marks on lake sturgeon Acipenser fulvescens. Great Lakes Fishery Commission Miscellaneous Publlication 2007-02. Http:www.glfc.org/pubs/pub.

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htm#misc. —. 2009. Lethality of sea lamprey parasitism on lake sturgeon. Transactions of the American Fisheries Society 138:1065–1075. Pearce, W. A., R. A. Bream, S. M. Dustin, and J. J. Tibbles. 1980. Sea lamprey (Petromyzon marinus) in the Lower Great Lakes. Canadian Journal of Fisheries and Aquatic Sciences 37:1802–1810. Ricciardi, A. 2006. Patterns of invasion in the Laurentian Great Lakes in relation to changes in vector activity. Diversity and Distributions 12:425–433. Rostlund, E. 1952. Freshwater fish and fishing in native North America. University of California Publications in Geography 9:1–314. Roussow, G. 1957. Some considerations concerning sturgeon spawning periodicity. Journal of the Fishery Research Board of Canada 14:553–572. Schneider, C. P., R. W. Owens, R. A. Bergstedt, and R. O’Gorman. 1996. Predation by sea lamprey (Petromyzon marinus) on lake trout (Salvelinus namycush) in southern Lake Ontario, 1982–1992. Canadian Journal of Fisheries and Aquatic Sciences 53:1921–1932. Scott, W. B., and E. J. Crossman. 1973. The freshwater fishes of Canada. Bulletin of the Fisheries Research Board of Canada 184. Smith, B. R. 1971. Sea lampreys in the Great Lakes of North America. In: The biology of lampreys, vol. 1. M. W. Hardisty and I. C. Potter, eds. Academic Press. —. 1972. Factors of ecological succession in oligotrophic fish communities of the Laurentian Great Lakes. Journal of the Fisheries Research Board of Canada 29:717–730. Smith, C. L. 1985. Inland fishes of New York State. Department of Environmental Conservation.

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Smith, S. H. 1995. Early changes in the fish community of Lake Ontario, Great Lakes Fishery Commission Technical Report 60. Smith, B. R., and J. J. Tibbles. 1980. Sea lamprey (Petromyzon marinus) in Lakes Huron, Michigan, and Superior: History of invasion and control, 1936–78. Canadian Journal of Fisheries and Aquatic Sciences 37:1780–1801. Sutton, T. M., and S. H. Bowen. 1994. Significance of organic detritus in the diet of larval lampreys in the Great Lakes Basin. Canadian Journal of Fisheries and Aquatic Sciences 51:2380–2387. Sutton, T. M., B. L. Johnson, T. D. Bills, and C. S. Kolar. 2003. Effects of mortality sources on population viability of lake sturgeon: A stage-structured model approach. Great Lakes Fishery Commission project completion report. Swink, W. D. 1990. Effect of lake trout size on survival after a single sea lamprey attack. Transactions of the American Fisheries Society 119:996–1002. —. 1993. Effect of water temperature on sea lamprey growth and lake trout survival. Transactions of the American Fisheries Society 122:1161–1166. —. 2003. Host selection and lethality of attacks by sea lampreys (Petromyzon marinus) in laboratory studies. Journal of Great Lakes Research 29(Suppl. 1): 307–319. Swink, W. D., and K. T. Fredricks. 2000. Mortality of burbot from sea lamprey attack and initial analyses of burbot blood. In: Burbot biology, ecology, and management. V. L. Paragamian and D. L. Willis, eds. American Fisheries Society. Swink, W. D., and L. H. Hanson. 1989. Survival of rainbow trout and lake trout after sea lamprey attack. North American Journal of Fisheries Management 9:35–40.

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Thresher, R. E. 2008. Autocidal technology for the control of invasive fish. Fisheries 33(3): 114–121. Trautman, M. B. 1949. The invasion, present status, and life history of the sea lamprey in the waters of the Great Lakes, especially the Ohio waters of Lake Erie. The Ohio State University. The Franz Theodore Stone Laboratory. Waldman, J. R., C. Grunwald, N. K. Roy, and I. I. Wirgin. 2004. Mitochondrial DNA analysis indicates sea lampreys are indigenous to Lake Ontario. Transactions of the American Fisheries Society 133:950–960.

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Welsh, A., T. Hill, H. Quinlan, C. Robinson, and B. May. 2008. Genetic assessment of lake sturgeon population structure in the Laurentian Great Lakes. North American Journal of Fisheries Management 28:572–591.

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Future Management and Stewardship of Lake Sturgeon

Aldo Leopold is considered to have been one of the earliest conservation activists in the United States and is well known for his best-selling A Sand County Almanac, which calls for all humans to live with a “land ethic.” Doing so requires that we consider ourselves and all other organisms as valuable and integral partners within an ecosystem community (Leopold 1966). Because he worked for the U.S. Forest Service and wrote of a “land ethic,” we often think of Leopold’s focus mostly in terms of terrestrial systems. What few know is that some of his first employment obligations concerned aquatic systems and fish (Leopold 1918). His ethic included the aquatic systems within management areas. His initial position after graduating from Yale was with the U.S. Forest Service in Arizona and New Mexico, where he produced a Game and Fish Handbook (1915) calling on foresters to help maintain and support rare wildlife and not just popular game species. He championed the strategy that native fish species should be protected and preferred when considering stocking waters in national forests (Leopold 1918). Leopold’s “land ethic” resonated with environmentalists, contributing to the growth of preservation and management programs for parks, reserves, and forests

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over the last 100 years, yet a similar focus and concern for our aquatic ecosystems was much slower to develop. All of us can see more and more of the world’s natural wild lands being developed for housing, industry, and agriculture as our human population grows, but we are less inclined to realize the extent to which our freshwater resources and the organisms that live in those ecosystems are becoming depleted, polluted, or overexploited, unless there is some drastic outcome of our actions. Examples include the oil and debris spills in the Cuyahoga River in Ohio, which often caught fire in the 1950s and 1960s (Scott 2009) and the large die-offs of the invasive fish—the alewife—in the Great Lakes during the late 1960s (Anon. 1967). The Clean Water Act of 1972 sprang from the deplorable conditions of many lakes and rivers in the 1960s and the need to regulate discharges from industry and municipal sources. The result has been an improvement in water clarity and condition, especially for drinking, but attention seemed to remain focused on water as a resource and not as an integral part of an ecosystem with associated organisms. As human populations grow around the world, the demand for clean freshwater also grows. Since 2000, much has been written about safety and availability of freshwater resources, and slowly emphasis has been placed on freshwater and marine natural resources. Only within the last 10 to 15 years has the idea of a water ethic been championed (Postel 1997; National Catholic Rural Life Conference 2003), and in March 2008 a conference was held in Santa Clara, California, titled Common Grounds, Common Water: Toward A Water Ethic (proceedings found at http:// www.internationalwaterlaw.org/bibliography/Ethics/). As humans slowly connect the importance of water with that of life, that of both their own and plants and animals, we become more aware of what Leopold was suggesting almost 100 years ago—we need to protect all life-forms native to each ecosystem. These organisms have distinct roles within ecosystems that allow them to function and support life best. In the Great Lakes Basin ecosystem, invasive and introduced species such as the zebra and quagga mussels, sea lamprey, and round goby are creating ecological, economic, and industrial havoc, clogging water intakes, and disrupting fish food webs. Sustainability has become an important term as population growth, demand for resources, climate change, and continued pollution make us all aware of the delicate balance of all of our natural resources. A resource is sustainable if the manner in which we live on earth today allows us to meet our present need of that resource without destroying the ability of future generations to also have sufficient supply and experience of that resource (Garcia and Grainger 1997). In the past, we’ve often dealt with individual species in small regions but now realize the need to adapt a large ecosystem sustainability strategy.

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In the chapters that have preceded, we have learned that the native and unique lake sturgeon has begun to rebound in some regions of its historic range, especially in some portions of the Great Lakes. Few self-sustaining and unrestricted (by dams or habitat loss) groups of lake sturgeon remain, but these groups are rebuilding when conditions allow. As lake sturgeon populations begin to recover through current management practices, all of us need to decide where and how lake sturgeon will be managed and protected. New approaches to sturgeon management can have wider implications for our current approaches to fish management, which include maximum sustainable yields for commercial or sport fisheries or total protection for rare or small fishes. Fisheries management needs to more intentionally include management for sustaining the role of a species within its ecosystem, as Leopold suggested a century ago (Leopold 1918) as well as manage for sustaining the full potential of each species to adapt and evolve, that is, to reduce typical harvest practices of targeting the largest individuals of any population. Scientists are learning the effects of years of harvesting the largest individuals within a stock—we are actually reducing the overall attainable size of the species (Conover and Munch 2002). Lake sturgeon are the largest fishes in the Great Lakes, and if a fishery continues for those individuals that are the largest throughout the entire ecosystem, we will eventually reduce the potential size of these special fishes. Size matters. The controversy over fishing regulations worldwide that support the catch of the largest individuals must change, or we change the fish itself. Imposing a selective force on any population exposed to fishing will, over time, select for smaller fish, and large trophy fish may become a thing of the past (Conover and Munch 2002; Jorgensen et al. 2007). There are new strategies and scenarios in which better management practices can be applied.

So Why Restore or Protect Lake Sturgeon? Recently a young woman asked me to explain the benefit of rebuilding lake sturgeon populations. I found it an odd question, as it was couched in the typical thinking that all resources must be seen as a benefit to us. Even 100 years after Aldo Leopold suggested humans need to live harmoniously within our ecosystems and watersheds, underappreciation of the role of humans as part of our environment persists. There are many stories and descriptions of species now extinct, often as a direct result of human activity. Examples include the passenger pigeon in 1914 (Schorger 1955), Caribbean monk seal in 1952 (Roach 2009), and the Great Lakes deepwater cisco in the 1960s (U.S. Fish and Wildlife Service 2009). Once a species is lost, it cannot be returned. We will never know if any extirpated species held the key to a

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cancer cure or was a potentially more nutritious food source. But we can be sure it contributed in some way to the ecosystem balance.

What Needs to Be Considered for the Future Management of Lake Sturgeon?

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Basic habitat: As discussed in earlier chapters, lake sturgeon need open corridors to reach historic spawning habitat, and those places need good, clean natural freshwater flow to allow eggs to hatch and young to drift downstream to foodrich habitat. Ecosystem role: Lake sturgeon are benthic feeders; they keep lakes and rivers clean by consuming dead and dying organisms and by feeding on other organisms that help digest and decompose organic debris, thereby helping to maintain ecosystem stability. Human expectations: Humans have always needed to harvest plants and animals for food. Early Native Americans harvested sturgeon for food, often smoking or drying the flesh. Some Native Americans today still retain sturgeon clan names, and the fish is culturally important to tribes in the Great Lakes region. Early Europeans who settled North America harvested sturgeon using spears, and in Wisconsin and Michigan small spear fisheries are still allowed. This is because spear fishermen bring economic support to local communities that have a historic record of such activities. Some management agencies also benefit economically by these fisheries, as they supply additional research revenue through the sale of fishing stamps or licenses. In other areas where fishing for sturgeon long ago collapsed, some remnant stocks of fish exist and with some protection may produce a rebound in abundance if habitat is protected and corridors of migration remain open. As long as there is economic stability, humans may begin to acknowledge and appreciate natural resources not so much for their benefits to us as for their place in the ecosystem. This should create a growing populace satisfied simply to know that an organism as rare and as historic as the lake sturgeon exists and is afforded some protection. As we become educated in the importance of maintaining each organism and allowing at least some members of each organism to attain the size and age they might have without severe human depredations, fewer will want to kill or harvest them with abandon. New approaches to satisfying man’s desire to harmonize with the natural world, often for hunting and fishing, could include encouragement to take photos and releasing fish after capture, which has driven some management agencies to

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consider catch-and-release options for some popular fishes. On the U.S. West Coast large sport catch-and-release fisheries are growing for white sturgeon. That option is increasing in Michigan waters of Lake St. Clair and Lake Erie as well (M. Thomas, personal communication). But no matter what approach or strategy managers take for sturgeon, caution is needed. There is evidence that unrestricted catch-and-release programs can produce such fishing pressure that a single sturgeon can be caught many times in one season, and such use of energy (fighting hook capture) may deplete reserves of fat needed to sustain spawning and migration (McKeown 1984). There are also many new, unexpected problems in protecting lake sturgeon populations that may arise: invasive species, poaching, land use changes or sales, dam relicensing or deconstruction, economic imperatives to open lands for mining or other uses, global climate change, and warming waters, bringing increased incidents of disease and parasite spread (Alben et al. 2006).

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A Best Practice for Managing This Unique Species? Historically, fish populations were managed to produce MSY—maximum sustainable yields, again with the aim of retaining sustainable populations. This was done by collecting data on populations thought to have geometric growth (a constant amount of increase) and then harvesting some (usually adults or large individuals) at a rate at which they will be replaced by young fish growing into the adult population, based on life-history data. This type of calculation has not been beneficial to fish stocks because it often overlooked environmental instability (impacts of weather, disease, or predators that can wipe out young fishes), density dependence (when some fish become crowded into one area, their reproductive effort can decrease), and human impacts (lack of complete information on life history, industrial discharges, dams, actual catch). Computer modeling and the ability to handle large data sets have improved some predictions, but uncertainty remains, as ecosystems and organisms are ever changing. In the last 20 years fishery personnel have been examining additional ways to enhance fish management, especially in the ocean, where many fish stocks have been severely overfished. A relatively new approach is called adaptive management, which has two basic components: to monitor stocks closely and respond quickly. Management plans include constant measuring of important indicators and current status of populations with a planned response, which allows managers to change outcomes (Hilborn and Sibert 1988). Such efforts as closing a fishery when catch quotas are reached or suspending a fishery for a year (or more) if reproductive effort was lost

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can improve outcomes. Providing opportunities to view fish either in aquariums or at viewing stations can also inform people as to the plight of many fish species and encourage their support of management decisions. The most recent suggestion to improve management efforts is to move to the ecosystem scale with “nested” governance (Garcia and Grainger 1997). Fishes move between state, country, and oceanic boundaries and their value in trade and economics also changes. Cultural perspectives, local and international law, and general oversight vary, complicating the development of such goals. Walters (2007) suggests large-scale management can work with improvements in three areas. There needs to be an increase in resources for expanded monitoring at such large ecosystem scales, policymakers need to accept uncertainty, and finally, we need fishery managers with vision and a willingness to oversee such projects.

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Marine Protected Areas In October 1972, the U.S. Marine Sanctuaries Program was developed because there was a need to protect spawning grounds and nursery areas of valued marine species. This initiative was later renamed the National Marine Sanctuaries Act, the purpose of which is to conserve, protect, and enhance the biodiversity, ecological integrity, and cultural legacy of marine protected areas. The official federal definition of an MPA is “any area of the marine environment that has been reserved by federal, state, tribal, territorial, or local laws or regulations to provide lasting protection for part or all of the natural and cultural resources therein” (Executive Order 13158, May 2000). In practice, MPAs are defined areas where natural or cultural resources are given greater protection than the surrounding waters. In the United States, MPAs span a range of habitats including the open ocean, coastal areas, intertidal zones, estuaries, and the Great Lakes. They also vary widely in purpose, legal authorities, agencies, management approaches, level of protection, and restrictions on human uses (http://mpa.gov/all_about_mpa/basics.html). Most MPAs are located within the ocean/marine environment, but there are some in the Great Lakes region that focus on preserving historic shipwrecks. There is a call to develop MPAs for some important Great Lakes fish and aquatic resources. In the U.S. waters of the Great Lakes, as of April 2009, three MPAs had been identified: Isle Royale National Park, Huron Islands National Wildlife Refuge, and Thunder Bay National Marine Sanctuary and underwater preserve (www.mpa. gov). In Canada, one region within the Great Lakes, Georgian Bay, was designated as a unesco Biosphere Reserve in 1990 for the purpose of maintaining a balanced relationship between man and the environment. More recently Canada established

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the Lake Superior National Marine Conservation Area from Thunder Cape to Bottle Point along the Thunder Bay, Ontario, coastline, encompassing one-eighth of Lake Superior to protect and conserve representative examples of Canada’s Great Lakes (Parks Canada 2007).

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How Are U.S. MPAs Classified? Because there are many reasons a site may have importance to different “user” groups, there is a five-level system of classification for MPAs and activities that can be conducted within them (Wahle and Uravitch 2006). All have a focus on conservation yet can vary in the level, constancy, permanence, and ecological scale of protection. The focus of any MPA can be centered on natural or cultural heritage conservation (see figure 1, adapted from http://www.michigan.gov/deq/0,1607,7– 135–3313_3677_3701–14531—,00.html). This may or may not also include a focus on sustainable production—areas established to support extraction of some living renewable resource (fish, plant, etc.). Within these focus areas there can be a variety of levels of protection of resources depending on “use.” The protection can range from permanent to temporary, seasonal or year-round, and encompass an entire ecosystem or a single resource within the ecosystem (Wahle and Uravitch 2006). There is currently some discussion as to the effectiveness of using MPAs as tools to conserve and rehabilitate fisheries (Hilborn et al. 2004; Willis et al. 2003). There is evidence of improvements to fish populations when important stocks are protected within no-harvest areas, and these benefits have been well studied in oceanic environments. Depending on classification and size of the reserve or protected area, some fish and other marine resource populations under study doubled in density, and the average weight of individuals increased three times, with these results seen in as little as three years. Species diversity increased in protected areas by 20 percent, and systems with either several small connected areas or a single large reserve area showed similar improvements (Halpern 2003; Halpern and Warner 2002). Concern about the success of some reserves can be influenced by the life history of organisms slated for protection as harvestable size adults (large individuals) and pelagic (open water floating) young can regularly move outside the boundaries. Fishing pressure outside reserves can become so intense that improvements in size and abundance seen within the reserve are lost (Hilborn et al. 2004). Walters (2007) believes adaptive fishery management has failed in many protected areas through lack of leadership and oversight of such large projects, lack of willingness to experiment, and lack of funding for careful monitoring programs. Besides needing managers with vision and oversight, new fishery management tactics must involve

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Keewenaw Marquette

Whitefish Point

Alger DeTour Passage Grand Traverse Bay

Straits of Mackinac

Manitou Passage

Thunder Bay

Thumb Area

Sanilac Shores

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Southwest Michigan Figure 1. Michigan Marine Protection Areas

all stakeholders, encompass and model entire ecosystems, and explicitly address the issue of uncertainty (Walters 2007).

A Proposal for Fishery Managers in the Great Lakes with Regard to Lake Sturgeon We know that lake sturgeon have inhabited and continue to inhabit most regions of the Great Lakes, although in many locations their populations are more limited today than they were historically because of barrier dams and spawning habitat loss. Often, fish management plans have been developed on a lake-by-lake basis or state-by-state basis for species currently deemed sensitive, including sturgeon (Auer 2003; Hoff

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2003; Newman, DuBois, and Halpern 2003; Hay-Chmielewski and Whelan 1997). Rarely have Great Lakes fish been considered for complementary management strategies throughout their range or within an entire ecosystem. We propose that such a view and strategy be considered and developed for lake sturgeon within the Great Lakes. Using adaptive strategies, shared by many stakeholders, lake sturgeon management could exemplify new and optimal approaches to ensuring a place for this unique and important species within the entire ecosystem. Some populations are now so reduced in number that total fishery closures for indeterminate time periods alone may still not result in restoration. Other populations are being slowly and carefully managed to assure rebuilding sustainable stocks, while other populations are utilized for spear or hook-and-line fisheries. Some populations have rebounded so well that the public has begun to catch and release large individuals for the sport of it. Comprehensive management oversight is needed to ensure all stakeholder voices are considered so the lake sturgeon will forever remain at self-sustaining levels throughout the Great Lakes.

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MPAs and Managing for Lake Sturgeon When evaluating potential sites for designation as marine protected areas, managers must consider several criteria. Although evaluations should include social and economic criteria, for this chapter we will limit discussion to several ecological criteria suggested for use in evaluations (Roberts et al. 2003). Roberts et al., gathered as a working group on the Science of Marine Reserves funded by the National Science Foundation, suggest that proposed MPAs should cover many biogeographic regions and habitats, not just a few, thereby providing for all life stages of plants or animals of concern. Sites need to be protected from major human threat, must be of a significant size (or encompass several small sites with corridor connections), and should be focused on sensitive species or species that man exploits. Sites should also retain important ecological services for humans. With these criteria in mind, we propose that fishery managers consider an MPA as one part of lake sturgeon management in the Great Lakes basin. Lake sturgeon exhibit long-range movements, so effective management will occur only when Canadian and U.S. managers work together. In considering a total ecosystem approach to management and criteria for MPAs, managers must identify other important and sensitive species within the ecosystem that can benefit from designated reserve sites, as suggested in criteria for MPA establishment (Roberts et al. 2003). Using the classification system described earlier (Wahle and Uravitch 2006), we also need to identify within that ecosystem different use approaches such as total preserves,

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managed historic spear/hook-and-line fisheries, subsistence fisheries and cultural take, and proposed catch-and-release fisheries. Within the Great Lakes ecosystem, only one nonharvested, self-sustaining population of lake sturgeon remains. That population is found in Lake Superior (Auer 1999). All other populations have, until a very recent closure of the fishery in Ontario in 2008 (cosewic 2000; Vélez-Espino and Koops 2008), been subject to harvest—for example, stocks in Lake St. Clair Michigan/Ontario area, and those in the St. Lawrence Seaway. The Lake Superior stock is located in close proximity to one of the few remaining self-sustaining coaster brook trout populations and also an important walleye stock (Hoff 2003). All of these fish are culturally important to local tribes as well as to others for sport and commercial value. They are also key historic members of the Lake Superior ecosystem. Designing a MPA for this group of fish in south central Lake Superior would include the needed criteria of a broad biogeographic area, incorporation of protection for several sensitive species, and ecosystem services, as suggested by Roberts et al. (2003). The design and classification of subunits within a proposed MPA in Lake Superior for three critical fish species will require collaboration between state (Wisconsin and Michigan), tribal, and possibly Canadian agencies. Within the MPA, sections could be set aside as no-fishing zones or otherwise limit human activity. Areas covered by these protected sections could be different for each species or more inclusive of all species of special concern. These areas would allow at least one stock or population of each species to grow to large size and become monitored sections that contribute to our scientific knowledge on species life history. An MPA in the Great Lakes region would also provide stock for reintroductions, or simply be a base that “seeds” other regions. It can be managed as a large experiment in freshwater sustainable fisheries, proposed earlier as the future for sustainable fisheries. These strategies are being employed in the oceans and can be adapted to freshwater systems and species. Such a proposal does not limit or restrict establishment of other strategies in other locations within a state or other Great Lake region where current fishing or cultural use may be well established.

Proposal for a Lake Sturgeon / Coaster Brook Trout / Walleye Marine Protected Area Underwater preserves in the state of Michigan were established in 1980 to provide opportunities for sport diving and to protect historic shipwreck sites (Graf 2009). However, states have also designated underwater preserves to support the conservation of fish and wildlife, such as coral reefs, fishes, and monk seals. Hawaii declared

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a 100,000-square-mile underwater preserve in June 2006 (Shogren 2006). Such preserves, protecting natural resources, are most often associated with marine systems, usually establishing an area free of heavy harvest pressure. This allows stocks of fish, shellfish, and other organisms to recover and perpetually build populations, which eventually begin to “seed” and restore fisheries in adjacent areas. Underwater preserves for natural resource protection in the Great Lakes region have not been developed even though human development, exotic species, continued harvest, and barriers to migration corridors have prevented many native aquatic species from rebounding from the overharvest and development in the late 1800s (Auer 1999). Some of the fish species historically abundant and sought either in commercial or sport fisheries, and now absent or low in abundance in the lower Great Lakes, include lean lake trout, ciscoes, coaster brook trout, and lake sturgeon. Lake Superior remains a stronghold for all of these species, and the proposed United States site contains the only self-sustaining, nonharvested population of lake sturgeon in the Great Lakes Basin. Populations of lake sturgeon in the U.S. waters of the Great Lakes remain either threatened or endangered in all states from continued harvest or reduced habitat availability from dam construction for mills and hydropower (Auer 2004). Although there are areas of free-ranging lake sturgeon (St. Lawrence River, Lake St. Clair, and Detroit River), these populations remain subject to harvest, and spawning sites in river systems have been compromised in most cases. Also, lake sturgeon prefer rivers with a large river mouth delta that provides habitat for production of food organisms. Few unimpacted systems remain today. Only in Lake Superior and only in the Sturgeon River / Keweenaw Bay system do lake sturgeon move freely to spawn, and then return to the larger lake to feed and rest without threat of harvest or loss of historic habitat. This unique, self-sustaining population should be protected so that one population within the Great Lakes system remains as a case study for other regions and also as a source of material to aid in recovery of the species throughout the rest of the basin. The coaster brook trout, once abundant in Lake Superior, is now restricted to only a few river sites. One is the Salmon-Trout river in Marquette County, Michigan (Newman, DuBois, and Halpern 2003). Coasters moving out of this river need protection from fishing pressure and should be included in a reserve. A population of walleye in the Huron Bay watershed has been selected by managers for rehabilitation (Hoff 2003). The MPA must be of sufficient size to ensure protection for all life stages of these species. Combining agency efforts for an MPA for Lake Superior would be a groundbreaking approach to sustainable fisheries management in freshwater ecosystems.

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Graf, T. 2009. Michigan underwater preserves system. Http://www.michigan.gov/deq/0,1607,7135-3313_3677_3701-14531-,00.html. Halpern, B. S. 2003. The impact of marine reserves: Do reserves work and does reserve size matter? Ecological Applications 13(1): S117–S137. Halpern, B. S., and R. R. Warner. 2002. Marine reserves have rapid and lasting effects. Ecology Letters 5:361–366. Hay-Chmielewski, E. M., and G. E. Whelan. 1997. Lake sturgeon rehabilitation strategy. Michigan Department of Natural Resources, Fisheries Division, Special Report No. 18. Hilborn, R., and J. Sibert. 1988. Adaptive management of developing fisheries. Marine Policy 12:112–121. Hilborn R., K. Stokes, J. J. Maguire, T. Smith, L. W. Botsford, M. Mangel, J. Orensanz, A. Parma, J. Rice, J. Bell, K. L. Cochrane, S. Garcia, S. J. Hall, G. P. Kirkwood, K. Sainsbury, G. Stefansson, and C. Walters. 2004. When can marine reserves improve fisheries management? Ocean and Coastal Management 47:197–205. Hoff, M. H., ed. 2003. A rehabilitation plan for walleye populations and habitats in Lake Superior. Great Lakes Fishery Commission Miscellaneous Publication 2003-01. Jorgensen, C., and 16 others. Managing evolving fish stocks. Science 318:1247–1248. Kaufman, L., J. B. C. Jackson, E. Sala, P. Chislom, E. D. Gomez, C. Peterson, R. V. Salm, and G. Llewellyn. 2004. Restoring and maintaining marine ecosystem function. In: Defying ocean’s end: An agenda for action. L. K. Glover and S. A. Earle, eds. Island Press.

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Shogren, E. 2006. Vast Hawaii sea area now a national monument . June 15. Http://www.npr.org/ templates/story/story.php?storyId=5488173. Vélez-Espino, L. A., and M. A. Koops. 2008. Recovery potential assessment for lake sturgeon (Acipenser fulvescens) in Canadian designatable units. Fisheries and Oceans, Burlington, Canada Document 2008/007. U.S. Fish and Wildlife Service. 2009. Extinct species. Http://www.fws.gov/midwest/Endangered/ lists/extinct.html. Wahle, C. M., and J. A. Uravitch 2006. A functional classification system for marine protected areas in the United States. www.mpa.gov. Walters, C. J. 2007. Is adaptive management helping to solve fisheries problems? Ambio 36(4): 304–307. Willis, T.J., R.B. Millar, R.C. Babcock and N. Tolimieri. 2003. Burdens of evidence and the benefits of marine reserves: putting Decartes before des horse? Environmental Conservation 30(2): 97–103.

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ABOUT THE AUTHORS

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About the Authors

Brenda Archambo is the founder and president of the Black Lake, Michigan, Chapter of Sturgeon for Tomorrow. Her volunteer leadership established partnerships among the Michigan Department of Natural Resources, stakeholders, and universities to implement research, habitat, conservation, and outreach programs to better understand lake sturgeon reproductive ecology and early life history in order to manage self-sustaining populations of Michigan’s lake sturgeon. Archambo is the recipient of the Michigan Public Service Commission Innovative Spirit Volunteer Service Award , the American Red Cross Everyday Heroes Award , the Michigan United Conservation Clubs Special Conservation Award, and the Huron Pines O. B. Eustis Outstanding Individual Conservationist Award. She is also a graduate of the Straits Area Leadership Forum and Great Lakes Fisheries Leadership Institute. Archambo is also an outreach consultant with the National Wildlife Federation and president of the Cheboygan County Economic Development Corporation. Nancy Auer is an Associate Professor in the Department of Biological Sciences and is also the Department Graduate Program Director at Michigan Technological University. She received her BS, MS, and PhD degrees from the University of n

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Minnesota Duluth, The University of Michigan, and Michigan Technological University, respectively. Nancy began her journey with lake sturgeon in 1987 using a small nongame wildlife grant from the Michigan DNR to study these fish. She fell in love with them when visiting a laboratory rearing young sturgeon at the University of Wisconsin while working on an identification guide to larval fishes of the Great Lakes. Auer has an active research program in the areas of large lake research and restoration of native fish species. She has published in a variety of well-known journals and has authored several book chapters. She enjoys living in the north country, is an avid birder, and has a home on Lake Superior with her husband Martin and two Welsh Corgies, Peanut and Prairie.

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Edward A. Baker is a Michigan native and was an avid angler while growing up in Grand Haven. However, like most people living in the Great Lakes region, he had no idea what a lake sturgeon was. That changed when he entered college and began studying fisheries science. Baker has been a research biologist with the Michigan DNR working in Marquette after finishing graduate school in 1995 and has been studying lake sturgeon since that time. He has studied lake sturgeon distribution and status in Michigan, early life history, population assessment, restoration, streamside rearing, and population demographics. He has authored or co-authored numerous scientific papers on lake sturgeon and serves on a number of interagency committees and working groups. He has also been actively involved in the drafting of various management plans for lake sturgeon in the Great Lakes region. Dave Dempsey has been an environmental professional since 1982 and is the author or co-author of six previous books, including On the Brink: The Great Lakes in the 21st Century, which won a Michigan Notable Book Award. He was environmental advisor to Michigan Governor James Blanchard from 1983 to 1989 and served on the Great Lakes Fishery Commission from 1994 to 2001. He has a master’s degree in natural resources policy and law from Michigan State University. He lives in Rosemount, Minnesota. Pierre Dumont first studied biology at the Université de Montréal (1973) and received his MS (1977) and PhD degrees (1996) from Université du Québec à Montréal. He started serving as a fishery biologist in the beginning of the 1970s and was involved in the impact studies of the James Bay hydropower development. He has worked for the Québec government since 1978—mainly in the St. Lawrence River lowlands, the most urbanized part of the province. He is involved in scientific studies on the status and management of lake sturgeon, yellow perch, and American eel, on the long term monitoring of fish communities in the St. Lawrence River,

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on fish passage and fish habitat improvement, on invasive fish species, and on the recovery of the cooper redhorse, a rare and endangered species endemic to south western Québec. Dumont regularly collaborates with specialists of other national and international organizations of the Great Lakes on water level regulation, fish stocks status and management, and fish passage. He has also been involved in the restoration program of the European sturgeon since 1998, when he had the chance to work at the Cemagref (Bordeaux, France) for one year.

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Lauri Kay Elbing attended both Michigan State University and the University of Michigan, studying liberal arts and natural resource policy, respectively, and her work in public policy, political and issue campaigns spans more than twenty years. Perhaps most notable is her work for Congressman John D. Dingell, leading the establishment of the Detroit River International Wildlife Refuge and her work for the National Wildlife Federation developing a high donor advocacy program. She joined the government relations team of The Nature Conservancy in Michigan in March of 2010. John E. Gannon was born in Detroit, Michigan, in 1942 to a family of automobile factory workers. His career direction was highly influenced by learning to love the out-of-doors while spending summers at his grandparents’ cabin on Mullett Lake just south of the Straits of Mackinac in northern Lower Michigan. The lake is part of the inland water route that supports the only inland population of lake sturgeon in Michigan. Gannon has been fascinated with the “big fish” since childhood. He received his BS degree in biology at Wayne State University, MS in fisheries at the University of Michigan, and PhD in zoology (concentration in limnology) at the University of Wisconsin. Gannon worked on the Great Lakes throughout his over four-decades-long career. Following twelve years in academic research and teaching in Michigan and New York, he held various positions in U.S. federal government research and management with the U.S. Geological Survey Great Lakes Science Center (and its predecessor agencies) in Ann Arbor, Michigan, and the International Joint Commission in Windsor, Ontario. Gannon has broad interests in eutrophication, toxic substances, invasive species, habitat protection and restoration, and environmental education and communications. He focused most of his professional career at the interface between research, resource management, and policy. He also enjoyed teaching summers at the University of Michigan Biological Station and Ohio State University’s Stone Biological Laboratory. Gannon retired in 2010 and is currently Scientist Emeritus at the USGS Great Lakes Science Center. He also is engaged in various citizen science activities in gardening, birding, and watershed quality.

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Marty Holtgren has loved lake sturgeon since he was a youngster when he saw one at the Shedd Aquarium in Chicago. While growing up he lived on the St. Joe River in Niles, Michigan, which once held lake sturgeon but does no longer. He received his bachelor’s degree from Bethel College with coursework from Taylor University and the AuSable Institute. He searched to find a graduate position where he could enhance understanding of lake sturgeon to ultimately improve their condition. This led him to Michigan Technological University for a master’s degree studying lake sturgeon under the guidance of Dr. Nancy Auer, whom he has worked closely with for the past fifteen years. Currently, he works as the senior fisheries biologist for the Little River Band of Ottawa Indians where he merges science with tribal cultural values, leading to the first streamside rearing facility for lake sturgeon. Holtgren is a PhD candidate at Michigan Technological University. He lives in Manistee, Michigan, with his family (who have spent almost as much time with sturgeon as he has). Robert M. Hughes was born in 1945 in north central Michigan to working class parents. For the first eighteen years of his life, hunting, fishing, trapping, and a large organic garden were major sources of food for the family. In his early teens, he observed Michigan Department of Natural Resources fish biologists surveying the lake along which he lived and learned that a person could earn a living by studying fish. Hughes obtained multidisciplinary BA and MS degrees at the University of Michigan and a PhD in fisheries at Oregon State University. He conducted research for the U.S. Environmental Protection Agency for thirty-two years as an on-site contractor. He works part-time for Amnis Opes Institute focusing on biological assessments of streams, lakes, and rivers across large geographic extents in the United States, Europe, Brazil, China, and India. Hughes is President-elect of the American Fisheries Society and a member of Oregon’s Independent Multidisciplinary Science Team, which reviews state actions for rehabilitating salmon and watersheds. Yves Mailhot was born in Sainte-Anne-de-la-Pérade, Québec—a small village along the St. Lawrence River famous for its winter time spawning run and sport fishing festival of Atlantic tomcod. When he was five years old his father took him fishing, which probably contributed to the choice of his future career. He studied biology at the Université du Québec à Trois-Rivières and completed graduate studies in applied ecology at the Université Scientifique et Médicale de Grenoble, in France. Mailhot served as a fishery biologist in the St. Lawrence River for thirty-three years, working mainly for Québec’s Ministry of Natural Resources and Wildlife. His lifetime objective was to characterize as precisely as possible what is never seen—the status of the populations of the principal species of fish—in order to contribute to their sustainable exploitation. He developed a unique expertise about the sport

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and commercial fisheries in the whole St. Lawrence River and collaborated with neighbor colleagues from Ontario and Maritime provinces to national fisheries issues. Mailhot contributed to many scientific committees and to the production of the fishery management plans of the majority of the St. Lawrence River freshwater fish species, like lake sturgeon, yellow perch, American eel, Atlantic tomcod, walleye, sauger, northern pike and striped bass. With his colleague, friend, and co-author Pierre Dumont, Yves has contributed to scientific study and management of the St. Lawrence River lake sturgeon since 1985. They are also particularly proud for creating together a standardized scientific monitoring program of the St. Lawrence River fish communities—the Réseau de suivi ichtyologique (RSI), designed to describe the changes and trends over time of the fish populations and their habitats—in order to help future generations of biologists to manage the resources and evaluate the impact of climate change. Jimmie Mitchell is a Native American treaty rights activist, environmentalist, and Tribal citizen with the Little River Band of Ottawa Indians. Mitchell has actively fought to preserve Tribal hunting, fishing, and gathering rights secured within the 1836 Treaty of Washington. He has been an advocate of Tribal co-management related to existing natural resources and also in the development of programs focusing on the restoration, reclamation, and enhancement of degraded ecosystems and extirpated species significant to the cultural needs of his People. Today, Mitchell is known as a cultural practitioner for his Tribe, serves his Tribe as Natural Resources Director (since 2006), is an active board member on the Chippewa Ottawa Resource Authority (since 2003), and is a representative on the Bureau of Indian Affairs, Midwest Region’s Tribal Interior Budget Council. Holly Muir is a Science Communications Liaison for the U.S. Geological Survey Great Lakes Science Center in Ann Arbor, Michigan. Her Master’s project involved assessing host size selection and lethality of sea lamprey on lake sturgeon in a laboratory setting under the advisement of Dr. Trent Sutton at Purdue University. Prior to attending Purdue, she worked with lake sturgeon on Lake of the Woods and Rainy River as an Ontario Ministry of Natural Resources summer STEP student, gillnetting sturgeon for the Minnesota Department of Natural Resources and the Rainy River First Nations Lake Sturgeon Recovery Project in 2004. Henry A. Regier was born in 1930 in the bush of Northern Alberta in a log house built by his immigrant father the previous year from local timber and lumber. A precautionary version of sustainable practice in communitarian living came with that kind of childhood. He was educated at Queen’s, Toronto, and Cornell Universities,

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implicitly in an ecogenic mindscape, as he notes in retrospect. He served as a director of the Institute for Environmental Studies (1989–1994) and Professor in the former Department of Zoology (1966–1995) at the University of Toronto. As an empirical scientist, policy administrator, and practical activist, he has worked collaboratively at all level of governance from local to global with respect to the human population, fisheries, commercialization, chemical pollution, and climate change. The ecosystem approach to the Laurentian Great Lakes Basin that he shares with colleagues integrates all of those issues in ecogenic ways. Among other honors, he is recipient of the Centenary Medal of the Royal Society of Canada, the Conservation Award of the Federation of Ontario Naturalists, the American Fisheries Society Award of Excellence, and the Lifetime Achievement Award from the International Association of Great Lakes Research.

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Trent M. Sutton is a Professor of Fisheries Biology and the Chair of the Undergraduate Fisheries Program at University of Alaska Fairbanks (UAF), where he has been a faculty member since June 2007. Prior to this he was a faculty member at Lake Superior State University and Purdue University. Since 2001, he has conducted research on lake sturgeon in the Great Lakes, focusing on various aspects of their distribution and movement patterns, habitat use, and life history. Most notably, his research focused on the impacts of various biotic and abiotic stressors (e.g., predation, parasitism, discharge events) on the survival and recruitment of various life stages within the context of long-term population viability. Amy Welsh has been working on lake sturgeon genetics since 2001. She received a BS from the University of Maryland. While working on a master’s of forensic science degree from the George Washington University, she discovered the use of genetics in identifying sturgeon species represented in batches of caviar. She then went on to receive her PhD from the University of California–Davis studying lake sturgeon genetic population structure. She was an Assistant Professor at SUNY–Oswego, continuing genetic studies in the Great Lakes. Currently, Welsh is an Assistant Professor at West Virginia University researching fish and wildlife genetics. She lives in Morgantown, West Virginia, with her husband and two daughters (who have quickly become sturgeon enthusiasts).

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