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Suggested Citation:"Front Matter." National Academies of Sciences, Engineering, and Medicine. 2020. A Research Strategy to Examine the Taxonomy of the Red Wolf. Washington, DC: The National Academies Press. doi: 10.17226/25891.
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Suggested Citation:"Front Matter." National Academies of Sciences, Engineering, and Medicine. 2020. A Research Strategy to Examine the Taxonomy of the Red Wolf. Washington, DC: The National Academies Press. doi: 10.17226/25891.
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Suggested Citation:"Front Matter." National Academies of Sciences, Engineering, and Medicine. 2020. A Research Strategy to Examine the Taxonomy of the Red Wolf. Washington, DC: The National Academies Press. doi: 10.17226/25891.
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Suggested Citation:"Front Matter." National Academies of Sciences, Engineering, and Medicine. 2020. A Research Strategy to Examine the Taxonomy of the Red Wolf. Washington, DC: The National Academies Press. doi: 10.17226/25891.
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Suggested Citation:"Front Matter." National Academies of Sciences, Engineering, and Medicine. 2020. A Research Strategy to Examine the Taxonomy of the Red Wolf. Washington, DC: The National Academies Press. doi: 10.17226/25891.
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Suggested Citation:"Front Matter." National Academies of Sciences, Engineering, and Medicine. 2020. A Research Strategy to Examine the Taxonomy of the Red Wolf. Washington, DC: The National Academies Press. doi: 10.17226/25891.
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Suggested Citation:"Front Matter." National Academies of Sciences, Engineering, and Medicine. 2020. A Research Strategy to Examine the Taxonomy of the Red Wolf. Washington, DC: The National Academies Press. doi: 10.17226/25891.
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Page viii Cite
Suggested Citation:"Front Matter." National Academies of Sciences, Engineering, and Medicine. 2020. A Research Strategy to Examine the Taxonomy of the Red Wolf. Washington, DC: The National Academies Press. doi: 10.17226/25891.
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Suggested Citation:"Front Matter." National Academies of Sciences, Engineering, and Medicine. 2020. A Research Strategy to Examine the Taxonomy of the Red Wolf. Washington, DC: The National Academies Press. doi: 10.17226/25891.
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Suggested Citation:"Front Matter." National Academies of Sciences, Engineering, and Medicine. 2020. A Research Strategy to Examine the Taxonomy of the Red Wolf. Washington, DC: The National Academies Press. doi: 10.17226/25891.
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Suggested Citation:"Front Matter." National Academies of Sciences, Engineering, and Medicine. 2020. A Research Strategy to Examine the Taxonomy of the Red Wolf. Washington, DC: The National Academies Press. doi: 10.17226/25891.
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Suggested Citation:"Front Matter." National Academies of Sciences, Engineering, and Medicine. 2020. A Research Strategy to Examine the Taxonomy of the Red Wolf. Washington, DC: The National Academies Press. doi: 10.17226/25891.
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Page xiii Cite
Suggested Citation:"Front Matter." National Academies of Sciences, Engineering, and Medicine. 2020. A Research Strategy to Examine the Taxonomy of the Red Wolf. Washington, DC: The National Academies Press. doi: 10.17226/25891.
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Suggested Citation:"Front Matter." National Academies of Sciences, Engineering, and Medicine. 2020. A Research Strategy to Examine the Taxonomy of the Red Wolf. Washington, DC: The National Academies Press. doi: 10.17226/25891.
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Below is the uncorrected machine-read text of this chapter, intended to provide our own search engines and external engines with highly rich, chapter-representative searchable text of each book. Because it is UNCORRECTED material, please consider the following text as a useful but insufficient proxy for the authoritative book pages.

PREPUBLICATION COPY A Research Strategy to Examine the Taxonomy of the Red Wolf Committee on Assistance to the U.S. Fish and Wildlife Service on Taxonomic Studies of the Red Wolf: A Review of Applications to Carry Out Research and Development of a Research Strategy Board on Life Sciences Board on Agriculture and Natural Resources Division on Earth and Life Studies A Consensus Study Report of

THE NATIONAL ACADEMIES PRESS 500 Fifth Street, NW Washington, DC 20001 This activity was supported by contracts between the National Academy of Sciences and U.S. Fish and Wildlife Service, Contract No. 140F0918C0005. Any opinions, findings, conclusions, or recommendations expressed in this publication do not necessarily reflect the views of any organization or agency that provided support for the project. Digital Object Identifier: https://doi.org/10.17226/25891 Additional copies of this publication are available from the National Academies Press, 500 Fifth Street, NW, Keck 360, Washington, DC 20001; (800) 624-6242 or (202) 334-3313; http://www.nap.edu. Copyright 2020 by the National Academy of Sciences. All rights reserved. Printed in the United States of America Suggested citation: National Academies of Sciences, Engineering, and Medicine. 2020. A Research Strategy to Examine the Taxonomy of the Red Wolf. Washington, DC: The National Academies Press. https://doi.org/10.17226/25891. Prepublication Copy

The National Academy of Sciences was established in 1863 by an Act of Congress, signed by President Lincoln, as a private, nongovernmental institution to advise the nation on issues related to science and technology. Members are elected by their peers for outstanding contributions to research. Dr. Marcia McNutt is president. The National Academy of Engineering was established in 1964 under the charter of the National Academy of Sciences to bring the practices of engineering to advising the nation. Members are elected by their peers for extraordinary contributions to engineering. Dr. John L. Anderson is president. The National Academy of Medicine (formerly the Institute of Medicine) was established in 1970 under the charter of the National Academy of Sciences to advise the nation on medical and health issues. Members are elected by their peers for distinguished contributions to medicine and health. Dr. Victor J. Dzau is president. The three Academies work together as the National Academies of Sciences, Engineering, and Medicine to provide independent, objective analysis and advice to the nation and conduct other activities to solve complex problems and inform public policy decisions. The National Academies also encourage education and research, recognize outstanding contributions to knowledge, and increase public understanding in matters of science, engineering, and medicine. Learn more about the National Academies of Sciences, Engineering, and Medicine at www.nationalacademies.org.

Consensus Study Reports published by the National Academies of Sciences, Engineering, and Medicine document the evidence-based consensus on the study’s statement of task by an authoring committee of experts. Reports typically include findings, conclusions, and recommendations based on information gathered by the committee and the committee’s deliberations. Each report has been subjected to a rigorous and independent peer-review process and it represents the position of the National Academies on the statement of task. Proceedings published by the National Academies of Sciences, Engineering, and Medicine chronicle the presentations and discussions at a workshop, symposium, or other event convened by the National Academies. The statements and opinions contained in proceedings are those of the participants and are not endorsed by other participants, the planning committee, or the National Academies. For information about other products and activities of the National Academies, please visit www.nationalacademies.org/about/whatwedo.

COMMITTEE ON ASSISTANCE TO THE U.S. FISH AND WILDLIFE SERVICE ON TAXONOMIC STUDIES OF THE RED WOLF: A REVIEW OF APPLICATIONS TO CARRY OUT RESEARCH AND DEVELOPMENT OF A RESEARCH STRATEGY Members JOSEPH TRAVIS, Chair, Florida State University FRED W. ALLENDORF, University of Montana LILIANA CORTÉS ORTIZ, University of Michigan MELANIE CULVER, U.S. Geological Survey and University of Arizona DIANE P. GENEREUX, Broad Institute KELLEY HARRIS, University of Washington ELAINE A. OSTRANDER, National Institutes of Health P. DAVID POLLY, Indiana University Bloomington ANNE STONE, Arizona State University Staff KEEGAN SAWYER, Study Co-Director CAMILLA YANDOC ABLES, Study Co-Director JESSICA DE MOUY, Senior Program Assistant Consultant ROBERT POOL, Editor Sponsor U.S. FISH AND WILDLIFE SERVICE Prepublication Copy v

BOARD ON LIFE SCIENCES Chair BARBARA A. SCHAAL, Washington University in St. Louis Members A. ALONSO AGUIRRE, George Mason University VALERIE H. BONHAM, Ropes & Gray LLP DOMINIQUE BROSSARD, University of Wisconsin–Madison NANCY D. CONNELL, Johns Hopkins Center for Health Security SEAN M. DECATUR, Kenyon College SCOTT V. EDWARDS (NAS), Harvard University GERALD L. EPSTEIN, National Defense University ROBERT J. FULL, University of California, Berkeley ROBERT NEWMAN, The Aspen Institute STEPHEN J. O’BRIEN (NAS), Nova Southeastern University LUCILA OHNO-MACHADO, University of California, San Diego CLAIRE POMEROY (NAM), Albert and Mary Lasker Foundation SUSAN RUNDELL SINGER, Rollins College DAVID R. WALT (NAE, NAM), Harvard Medical School PHYLLIS M. WISE, University of Colorado Staff KAVITA BERGER, Director FRAN SHARPLES, Advisor JO HUSBANDS, Scholar and Senior Project Director KATHERINE BOWMAN, Senior Program Officer ANDREA HODGSON, Senior Program Officer KEEGAN SAWYER, Senior Program Officer STEVEN MOSS, Program Officer AUDREY THÉVENON, Program Officer MATTHEW ANDERSON, Financial Business Partner JESSICA DE MOUY, Senior Program Assistant KOSSANA YOUNG, Senior Program Assistant vi Prepublication Copy

BOARD ON AGRICULTURE AND NATURAL RESOURCES Chair CHARLES W. RICE, Kansas State University, Manhattan Members ARISTOS ARISTIDOU (NAE), Cargill, Inc. SHANE C. BURGESS, University of Arizona, Tucson SUSAN CAPALBO, Oregon State University GAIL CZARNECKI-MAULDEN, Nestlé Purina PetCare BERNADETTE DUNHAM, The George Washington University GEBISA EJETA, Purdue University JAMES S. FAMIGLIETTI, University of Saskatchewan FRED GOULD (NAS), North Carolina State University JOHN HAMER, Data Collective DCVC DOUGLAS B. JACKSON-SMITH, The Ohio State University JAMES W. JONES (NAE), University of Florida ERMIAS KEBREAB, University of California, Davis STEPHEN S. KELLEY, North Carolina State University JAN E. LEACH, Colorado State University ROBIN LOUGEE, Independent Consultant JILL J. MCCLUSKEY, Washington State University KAREN I. PLAUT, Purdue University RICARDO SALVADOR, Union of Concerned Scientists V. ALARIC SAMPLE, George Mason University Staff ROBIN A. SCHOEN, Director CAMILLA YANDOC ABLES, Senior Program Officer KARA N. LANEY, Senior Program Officer JENNA BRISCOE, Research Associate SARAH KWON, Program Assistant Prepublication Copy vii

Preface This report draws from a large scientific literature, not only the literature on red wolves, but the general literature on species formation, hybridization and introgression, morphology, and tracing ancestry with population genomics. The answers to difficult, important questions usually lie at the confluence of many different intellectual streams, and understanding the red wolf is no exception to this rule. The committee’s work stands atop a broad foundation of biological research about the evidence necessary to understand the diversity of the various canid populations that may represent the red wolf lineage. The report is also based on the presentations offered to the committee by many scientific colleagues, the comments of those scientists who did not present to us but who generously answered specific questions addressed to them, and the thoughts of all those who contacted us through the study’s website. I thank the members of the committee for their dedication and commitment to every phase of this report, including critically assessing a very large literature and sifting that literature to develop a research strategy for the United States Fish and Wildlife Service. The committee devoted considerable effort to explaining very complicated genomic and statistical issues and making the logic of the research strategy clear. On behalf of the committee, I thank the co-Director study director, Keegan Sawyer, for her leadership, insights, and masterful manner of persuading us to work ever harder. On the committee’s behalf, I also thank study co-Director Camilla Yandoc Ables for her guidance and assistance with many aspects of our work, especially her ability to translate some of our rough-hewn ideas for figures and flow charts into excellent aids for presentation. All of us on the committee are grateful to Jessica De Mouy for her support in organizing and conducting our meetings and for her help with all of the technical aspects of the report’s preparation. On behalf of the committee, I thank all those who informed us about wolves and about the conceptual issues involved in tracing ancestry and discerning relationships among populations. Some of these scientists gave in-person presentations, others presented webinars, and still others responded to written queries. Appendix A is a list of these scientists and how they helped us. The committee’s knowledge of the critical issues was deepened and broadened by the contributions of so many people, and our thinking was constantly challenged by each increment in our knowledge. Everyone who spoke or wrote to us was unfailingly generous with their time and gracious with their patience. The committee’s gratitude for so much effort cannot be overstated. Joseph Travis, Chair Committee on Assistance to the U.S. Fish and Wildlife Service on Taxonomic Studies of the Red Wolf: A Review of Applications to Carry Out Research and Development of a Research Strategy Prepublication Copy ix

Acknowledgment of Reviewers This Consensus Study Report was reviewed in draft form by individuals chosen for their diverse perspective and technical expertise. The purpose of this independent review is to provide candid and critical comments that will assist the National Academies of Sciences, Engineering, and Medicine in making each published report as sound as possible and to ensure that it meets the institutional standards for quality, 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 thank the following individuals for their review of this report: Melissa Hawkins, Smithsonian Museum of Natural History Jennifer Leonard, Estación Biológica de Doñana - CSIC Jonathan Losos (NAS), Washington University in St. Louis Roland Kays, North Carolina Museum of Natural Sciences Julie Meachen, Des Moines University Michael Schwartz, U.S. Forest Service Aaron Shafer, Trent University Lisette Waits, University of Idaho Jeff Wall, University of California, San Francisco Although the reviewers listed above provided many constructive comments and suggestions, they were not asked to endorse the conclusions or recommendations of this report. The review of this report was overseen by Dr. Barbara Schaal (NAS) of Washington University in St. Louis and Dr. James A. Estes (NAS) of the University of California, Santa Cruz. They were responsible for making certain that an independent examination of this report was carried out in accordance with the standards of the National Academies and that all review comments were carefully considered. Responsibility for the final content rests entirely with the authoring committee and the National Academies. Prepublication Copy xi

Contents SUMMARY ................................................................................................................................................. 1 1 INTRODUCTION ............................................................................................................................ 10 Purpose of the Study, 11 Why Do More Research on the Red Wolf?, 11 Committee’s Approach to Its Charge, 18 Organization of the Report, 19 References, 19 2 MORPHOLOGY, BEHAVIOR, AND ECOLOGY ...................................................................... 21 Morphology, 21 Behavior and Ecology, 28 Conclusions and Recommendations, 32 References, 32 3 COLLECTING GENOMIC DATA FROM HISTORICAL AND ANCIENT CANIDS........... 38 Sample Selection, 38 Partial Versus Whole Genome Data Collection, 39 Ancient DNA Analyses, 41 Data Filtering, 43 Data Analysis and Interpretation, 44 Recent Research on Ancient North American Canids, 47 Conclusions and Recommendations, 47 References, 48 4 GENOMIC ANALYSES OF EXTANT CANIDS ......................................................................... 53 Collection of Genomic Data, 54 Sampling, 55 Data Analyses and Interpretation, 57 Conclusions and Recommendations, 63 References, 63 5 SYNTHESIS OF RESEARCH STRATEGY ................................................................................. 66 Components of the Research Strategy, 66 Guiding Principles for the Research Strategy, 68 Overall Recommendations, 69 Reference, 70 GLOSSARY .............................................................................................................................................. 71 APPENDIXES A OPEN SESSION AND WEBINAR AGENDAS ............................................................................ 77 Prepublication Copy xiii

Contents B RED WOLF (CANIS RUFUS) MUSEUM SPECIMENS ............................................................. 79 C SUMMARY OF MUSEUM HOLDINGS OF KEY NORTH AMERICAN CANIDS ............... 95 D DNA SAMPLES FROM RED WOLVES ...................................................................................... 99 E WHOLE GENOME RESOURCES .............................................................................................. 100 F BIOGRAPHICAL SKETCHES OF COMMITTEE MEMBERS ............................................. 104 G FULL STATEMENT OF TASK ................................................................................................... 107 xiv Prepublication Copy

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A Research Strategy to Examine the Taxonomy of the Red Wolf provides independent guidance about taxonomic research on the red wolf, Canis rufus. Building from the 2019 report Evaluating the Taxonomic Status of the Mexican Gray Wolf and the Red Wolf, this report reviews and ranks research applications to determine the taxonomy of wild canid populations in southern Louisiana and other relevant locations. The report then develops a research strategy to examine the evolutionary relationships between ancient red wolves, the extant managed red wolf populations, and the unidentified canid populations.

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