ATLANTIC SALMON IN MAINE

Committee on Atlantic Salmon in Maine

Board on Environmental Studies and Toxicology

Ocean Studies Board

Division on Earth and Life Studies

NATIONAL RESEARCH COUNCIL OF THE NATIONAL ACADEMIES

THE NATIONAL ACADEMIES PRESS
Washington, D.C. www.nap.edu



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Atlantic Salmon in Maine ATLANTIC SALMON IN MAINE Committee on Atlantic Salmon in Maine Board on Environmental Studies and Toxicology Ocean Studies Board Division on Earth and Life Studies NATIONAL RESEARCH COUNCIL OF THE NATIONAL ACADEMIES THE NATIONAL ACADEMIES PRESS Washington, D.C. www.nap.edu

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Atlantic Salmon in Maine THE NATIONAL ACADEMIES PRESS 500 Fifth Street, NW Washington, D.C. 20001 NOTICE: The project that is the subject of this report was approved by the Governing Board of the National Research Council, whose members are drawn from the councils of the National Academy of Sciences, the National Academy of Engineering, and the Institute of Medicine. The members of the committee responsible for the report were chosen for their special competences and with regard for appropriate balance. This project was supported by Grant No. 01-0008 between the National Academy of Sciences and the National Fish and Wildlife Foundation. Any opinions, findings, conclusions, or recommendations expressed in this publication are those of the author(s) and do not necessarily reflect the view of the organizations or agencies that provided support for this project. International Standard Book Number 0-309-09135-7 (book) International Standard Book Number 0-309-52977-8 (PDF) Library of Congress Control Number 2004110692 Cover design: The National Academies Press. The image is of a hen salmon from the Penobscot River painted by Walter H. Rich and is reprinted with permission of the Museum of Science, Boston, from Kendall, W.C., The Fishes of New England: The Salmons, 1935. Additional copies of this report are available from: The National Academies Press 500 Fifth Street, NW Box 285 Washington, DC 20055 800-624-6242 202-334-3313 (in the Washington metropolitan area) http://www.nap.edu Copyright 2004 by the National Academy of Sciences. All rights reserved. Printed in the United States of America

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Atlantic Salmon in Maine THE NATIONAL ACADEMIES Advisers to the Nation on Science, Engineering and Medicine The National Academy of Sciences is a private, nonprofit, self-perpetuating society of distinguished scholars engaged in scientific and engineering research, dedicated to the furtherance of science and technology and to their use for the general welfare. Upon the authority of the charter granted to it by the Congress in 1863, the Academy has a mandate that requires it to advise the federal government on scientific and technical matters. Dr. Bruce M. Alberts is president of the National Academy of Sciences. The National Academy of Engineering was established in 1964, under the charter of the National Academy of Sciences, as a parallel organization of outstanding engineers. It is autonomous in its administration and in the selection of its members, sharing with the National Academy of Sciences the responsibility for advising the federal government. The National Academy of Engineering also sponsors engineering programs aimed at meeting national needs, encourages education and research, and recognizes the superior achievements of engineers. Dr. Wm. A. Wulf is president of the National Academy of Engineering. The Institute of Medicine was established in 1970 by the National Academy of Sciences to secure the services of eminent members of appropriate professions in the examination of policy matters pertaining to the health of the public. The Institute acts under the responsibility given to the National Academy of Sciences by its congressional charter to be an adviser to the federal government and, upon its own initiative, to identify issues of medical care, research, and education. Dr. Harvey V. Fineberg is president of the Institute of Medicine. The National Research Council was organized by the National Academy of Sciences in 1916 to associate the broad community of science and technology with the Academy’s purposes of furthering knowledge and advising the federal government. Functioning in accordance with general policies determined by the Academy, the Council has become the principal operating agency of both the National Academy of Sciences and the National Academy of Engineering in providing services to the government, the public, and the scientific and engineering communities. The Council is administered jointly by both Academies and the Institute of Medicine. Dr. Bruce M. Alberts and Dr. Wm. A. Wulf are chair and vice chair, respectively, of the National Research Council www.national-academies.org

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Atlantic Salmon in Maine COMMITTEE ON ATLANTIC SALMON IN MAINE Members MICHAEL T. CLEGG (Chair), University of California, Riverside PAUL K. BARTEN, University of Massachusetts, Amherst IAN A. FLEMING, Oregon State University, Corvallis MART R. GROSS, University of Toronto, Toronto, Ontario LEWIS S. INCZE, University of Southern Maine, Portland ANNE R. KAPUSCINSKI, University of Minnesota, St. Paul PATRICK Y. O’BRIEN, ChevronTexaco Energy Technology Company, Richmond, CA BARBARA NEIS, Memorial University of Newfoundland, St. John’s, Newfoundland NILS RYMAN, Stockholm University, Stockholm, Sweden PETER E. SMOUSE, Rutgers University, New Brunswick, NJ JENNIFER L. SPECKER, University of Rhode Island, Narragansett ROBERT R. STICKNEY, Texas A & M University, College Station JON G. SUTINEN, University of Rhode Island, Kingston Staff DAVID POLICANSKY, Project Director SUSAN ROBERTS, Director, Ocean Studies Board RUTH E. CROSSGROVE, Senior Editor MIRSADA KARALIC-LONCAREVIC, Research Associate LEAH L. PROBST, Research Associate JENNIFER SAUNDERS, Research Associate BRYAN SHIPLEY, Research Associate JOHN BROWN, Senior Program Assistant SAMMY BARDLEY, Library Assistant Sponsor NATIONAL FISH AND WILDLIFE FOUNDATION

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Atlantic Salmon in Maine BOARD ON ENVIRONMENTAL STUDIES AND TOXICOLOGY Members JONATHAN M. SAMET (Chair), Johns Hopkins University, Baltimore, MD DAVID ALLEN, University of Texas, Austin THOMAS BURKE, Johns Hopkins University, Baltimore, MD JUDITH C. CHOW, Desert Research Institute, Reno, NV COSTEL D. DENSON, University of Delaware, Newark E. DONALD ELLIOTT, Willkie, Farr & Gallagher, LLP, Washington, DC CHRISTOPHER B. FIELD, Carnegie Institute of Washington, Stanford, CA WILLIAM H. GLAZE, Oregon Health and Science University, Beaverton SHERRI W. GOODMAN, Center for Naval Analyses, Alexandria, VA DANIEL S. GREENBAUM, Health Effects Institute, Cambridge, MA ROGENE HENDERSON, Lovelace Respiratory Research Institute, Albuquerque, NM CAROL HENRY, American Chemistry Council, Arlington, VA ROBERT HUGGETT, Michigan State University, East Lansing BARRY L. JOHNSON, Emory University, Atlanta, GA JAMES H. JOHNSON, Howard University, Washington, DC JUDITH L. MEYER, University of Georgia, Athens PATRICK Y. O’BRIEN, ChevronTexaco Energy Technology Company, Richmond, CA DOROTHY E. PATTON, International Life Sciences Institute, Washington, DC STEWARD T.A. PICKETT, Institute of Ecosystem Studies, Millbrook, NY ARMISTEAD G. RUSSELL, Georgia Institute of Technology, Atlanta LOUISE M. RYAN, Harvard University, Boston, MA KIRK SMITH, University of California, Berkeley LISA SPEER, Natural Resources Defense Council, New York, NY G. DAVID TILMAN, University of Minnesota, St. Paul CHRIS G. WHIPPLE, Environ Incorporated, Emeryville, CA LAUREN A. ZEISE, California Environmental Protection Agency, Oakland, CA Senior Staff JAMES J. REISA, Director DAVID J. POLICANSKY, Scholar

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Atlantic Salmon in Maine RAYMOND A. WASSEL, Senior Program Officer for Environmental Sciences and Engineering KULBIR BAKSHI, Senior Program Officer for Toxicology ROBERTA M. WEDGE, Senior Program Officer for Risk Analysis K. JOHN HOLMES, Senior Program Officer SUSAN N.J. MARTEL, Senior Program Officer SUZANNE VAN DRUNICK, Senior Program Officer EILEEN N. ABT, Senior Program Officer ELLEN K. MANTUS, Senior Program Officer RUTH E. CROSSGROVE, Senior Editor

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Atlantic Salmon in Maine OCEAN STUDIES BOARD Members NANCY RABALAIS (Chair), Louisiana Universities Marine Consortium, Chauvin LEE G. ANDERSON, University of Delaware, Newark WHITLOW AU, University of Hawaii, Manoa ARTHUR BAGGEROER, Massachusetts Institute of Technology, Cambridge RICHARD B. DERISO, Inter-American Tropical Tuna Commission, La Jolla, CA ROBERT B. DITTON, Texas A&M University, College Station EARL DOYLE, Shell Oil (retired), Sugar Land, TX ROBERT DUCE, Texas A&M University, College Station PAUL G. GAFFNEY, II, Monmouth University, Long Branch, NJ WAYNE R. GEYER, Woods Hole Oceanographic Institution, Woods Hole, MA STANLEY R. HART, Woods Hole Oceanographic Institution, Woods Hole, MA RALPH S. LEWIS, Connecticut Geological Survey (retired), Hartford WILLIAM F. MARCUSON, III, U.S. Army Corp of Engineers (retired), Vicksburg, MS JULIAN MCCREARY, JR., University of Hawaii, Honolulu JACQUELINE MICHEL, Research Planning, Inc., Columbia, SC JOAN OLTMAN-SHAY, Northwest Research Associates, Inc., Bellevue, WA ROBERT T. PAINE, University of Washington, Seattle SHIRLEY A. POMPONI, Harbor Branch Oceanographic Institution, Fort Pierce, FL FRED N. SPIESS, Scripps Institution of Oceanography, La Jolla, CA DANIEL SUMAN, Rosenstiel School of Marine and Atmospheric Science, University of Miami, FL Staff SUSAN ROBERTS, Director JENNIFER MERRILL, Senior Program Officer DAN WALKER, Senior Program Officer JOANNE BINTZ, Program Officer ALAN B. SIELEN, Visiting Scholar ANDREAS SOHRE, Financial Associate SHIREL SMITH, Administrative Coordinator

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Atlantic Salmon in Maine JODI BACHIM, Senior Program Assistant NANCY CAPUTO, Senior Program Assistant SARAH CAPOTE, Program Assistant

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Atlantic Salmon in Maine OTHER REPORTS OF THE BOARD ON ENVIRONMENTAL STUDIES AND TOXICOLOGY Air Quality Management in the United States (2004) Endangered and Threatened Fishes in the Klamath River Basin: Causes of Decline and Strategies for Recovery (2004) Cumulative Environmental Effects of Alaska North Slope Oil and Gas Development (2003) Estimating the Public Health Benefits of Proposed Air Pollution Regulations (2002) Biosolids Applied to Land: Advancing Standards and Practices (2002) The Airliner Cabin Environment and Health of Passengers and Crew (2002) Arsenic in Drinking Water: 2001 Update (2001) Evaluating Vehicle Emissions Inspection and Maintenance Programs (2001) Compensating for Wetland Losses Under the Clean Water Act (2001) A Risk-Management Strategy for PCB-Contaminated Sediments (2001) Acute Exposure Guideline Levels for Selected Airborne Chemicals (4 volumes, 2000–2004) Toxicological Effects of Methylmercury (2000) Strengthening Science at the U.S. Environmental Protection Agency (2000) Scientific Frontiers in Developmental Toxicology and Risk Assessment (2000) Ecological Indicators for the Nation (2000) Waste Incineration and Public Health (1999) Hormonally Active Agents in the Environment (1999) Research Priorities for Airborne Particulate Matter (4 volumes, 1998–2004) Arsenic in Drinking Water (1999) The National Research Council’s Committee on Toxicology: The First 50 Years (1997) Carcinogens and Anticarcinogens in the Human Diet (1996) Upstream: Salmon and Society in the Pacific Northwest (1996) Science and the Endangered Species Act (1995) Wetlands: Characteristics and Boundaries (1995) Biologic Markers (5 volumes, 1989–1995) Review of EPA’s Environmental Monitoring and Assessment Program (3 volumes, 1994–1995) Science and Judgment in Risk Assessment (1994) Pesticides in the Diets of Infants and Children (1993) Dolphins and the Tuna Industry (1992)

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Atlantic Salmon in Maine Science and the National Parks (1992) Human Exposure Assessment for Airborne Pollutants (1991) Rethinking the Ozone Problem in Urban and Regional Air Pollution (1991) Decline of the Sea Turtles (1990) Copies of these reports may be ordered from the National Academies Press (800) 624-6242 or (202) 334-3313 www.nap.edu

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Atlantic Salmon in Maine exceptionally knowledgeable and cooperative committee. I learned a great deal from my colleagues, and I approach the end of this report with a sense of regret that the fellowship associated with our work has reached an end. The superb staff of the National Research Council also eased our task. Our study director, Dr. David Policansky, is himself an expert in fish biology, genetics, and conservation policy. Dr. Policansky contributed to the final report in numerous ways, and the report is much better because of his professional touch. Dr. Susan Roberts and Leah Probst contributed their expertise and skill to the report’s quality as well, and we are grateful to Jennifer Saunders, Bryan Shipley, Dominic Brose, and John Brown for their attention to the needs of the committee throughout its meetings and for their attention to detail in production of the report. We also thank Ruth Crossgrove and Mirsada Karalic-Loncarevic for their editorial and research efforts. Michael T. Clegg Chair, Committee on Atlantic Salmon in Maine

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Atlantic Salmon in Maine Acknowledgments This report has been enhanced by the contributions of many people. They are listed below and we are grateful to them all. This report has been reviewed in draft form by individuals chosen for their diverse perspectives and technical expertise, in accordance with procedures approved by the NRC’s Report Review Committee. The purpose of this independent review is to provide candid and critical comments that will assist the institution in making its published report as sound as possible and to ensure that the report meets institutional standards for objectivity, evidence, and responsiveness to the study charge. The review comments and draft manuscript remain confidential to protect the integrity of the deliberative process. We wish to thank the following individuals for their review of this report: Edward T. Baum, Atlantic Salmon Unlimited Barry Costa-Pierce, University of Rhode Island Perry R. Hagenstein, Institute for Forest Analysis, Planning, and Policy William K. Hershberger, National Center for Cool and Cold Water Aquaculture Irving Kornfield, University of Maine Kai N. Lee, Williams College Rosamond Lee Naylor, Stanford University Ralph Pisapia, Meridith, New Hampshire Jack Schmidt, Utah State University

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Atlantic Salmon in Maine Nancy Targett, University of Delaware James Wilson, University of Maine Although the reviewers listed above have provided many constructive comments and suggestions, they were not asked to endorse the conclusions or recommendations, nor did they see the final draft of the report before its release. The review of this report was overseen by John Dowling, Harvard University (review monitor), and John Burris, Beloit College (review coordinator). Appointed by the NRC, they were responsible for making certain that an independent examination of this report was carried out in accordance with institutional procedures and that all review comments were carefully considered. Responsibility for the final content of this report rests entirely with the authoring committee and the institution. We are grateful to Randall Peterman for his helpful advice on methods of risk assessment and risk analysis.

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Atlantic Salmon in Maine Presenters at Public Sessions John Banks, Penobscot Indian Nation Ed Baum, Atlantic Salmon Unlimited Ken Beland, Maine Atlantic Salmon Commission David Bell, Wild Blueberries Commission Sebastian Belle, Maine Aquaculture Association Russell W. Brown, NMFS Northeast Fisheries Science Center Elizabeth Butler, Pierce Atwood Mary Colligan, National Marine Fisheries Service David Courtemanch, Maine Department of Environmental Protection Laura Rose Day, Natural Resources Council of Maine Scott Dickerson, Coastal Mountains Land Trust Norman R. Dube, Maine Atlantic Salmon Commission Kevin Friedland, University of Massachusetts-National Oceanic and Atmospheric Administration John Gold, Texas A&M University Andy Goode, Atlantic Salmon Federation Terry Haines, University of Maine-U.S. Geological Survey Melissa Halsted, Kennebec Soil & Water Conservation District James Hawkes, NOAA-Fisheries, Maine Field Station The Honorable Angus King, Jr., Governor of Maine Thomas King, Craig Brook National Fish Hatchery, U.S. Fish and Wildlife Service Fred Kircheis, Maine Atlantic Salmon Commission John F. Kocik, NOAA

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Atlantic Salmon in Maine Stephen Koenig, Project SHARE Irv Kornfield, University of Maine Donna Loring, Penobscot Nation Jerry Marancik, Craig Brook National Fish Hatchery, U.S. Fish and Wildlife Service Morten Moesswilde, Department of Conservation, Maine Forest Service Paul Nickerson, U.S. Fish and Wildlife Service Stephen Page, ContiSea Ducktrap River Fish Farm Lee Perry, Maine Department of Inland Fisheries and Wildlife Jeff Reardon, Trout Unlimited Timothy Sheehan, NOAA Alan Spear, Bangor Hydro-Electric Company Joan Trial, Maine Atlantic Salmon Commission Douglas Watts, Friends of Kennebec Salmon Mark Whiting, Maine Department of Environmental Protection Fred Whoriskey, Jr., Atlantic Salmon Federation

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Atlantic Salmon in Maine Site Visits The committee visited the following sites during its meetings and is grateful to the many people who provided information, shared their facilities, provided hospitality, and provided valuable information there. Coopers Mills Sheepscot Sites Craig Brook Hatchery Ducktrap Coalition program Ducktrap Timber Preserve Edwards Dam Head Tide/Sheepscot watershed Maine Atlantic Salmon, LLC facilities Pleasant River Weir Wild Blueberry Farms

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Atlantic Salmon in Maine Contents     SUMMARY   1 1   INTRODUCTION   16      Background,   16      The Listing of Salmon under the Endangered Species Act,   20      The Present Study and Report Organization,   21 2   SALMON LIFE HISTORY AND ECOLOGY   23      Introduction,   23      Salmon Life History,   30      Physiology,   38      Aquatic Environments,   42      Climate Change,   43 3   THREATS TO ATLANTIC SALMON IN MAINE   47      Introduction,   47      A History of Threats to Atlantic Salmon in Maine,   50      Natural Predation and Competition,   53      Aboriginal, Commercial, and Recreational Salmon Fisheries in Maine,   56      Forestry, Farming, and Freshwater Habitat Quality,   61      Dams,   70      Hazards of Chemical Contaminants in Rivers and Streams,   75      Hatcheries,   77

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Atlantic Salmon in Maine      Aquaculture,   81      Acidification of Streams and Related Problems,   90      Research and Monitoring,   92      Governance,   92 4   SETTING PRIORITIES FOR ACTION   108      Risk Assessment and Decision Analysis Basics,   108 5   ADDRESSING THE THREATS TO ATLANTIC SALMON IN MAINE   138      A Strategy for Conservation and Restoration,   138      Dams,   139      Hatcheries,   139      Aquaculture,   163      Fishing,   167      Mortality of Salmon in Estuaries and the Ocean,   169      Research and Monitoring,   169      Governance,   170      Additional Conservation Options with Multiple Environmental Benefits,   173      Costs of Options,   183 6   FINDINGS AND RECOMMENDATIONS   188      Findings,   188      Recommendations,   191     REFERENCES   195 Appendix A   Endangered Species Act Basics   227 Appendix B   Governance   233 Appendix C   Supportive Breeding, Effective Population Size, and Inbreeding   251 Appendix D   Supportive Breeding and Risks to Genetic Quality   258 Appendix E   Summary of Committee’s Interim Report   261 Appendix F   Stocking Numbers, 1871–1995   265 Appendix G   Biographical Sketches of the Committee’s Members   271

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Atlantic Salmon in Maine ATLANTIC SALMON IN MAINE

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