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Suggested Citation:"Front Matter." National Academies of Sciences, Engineering, and Medicine. 2019. Evaluating the Taxonomic Status of the Mexican Gray Wolf and the Red Wolf. Washington, DC: The National Academies Press. doi: 10.17226/25351.
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Suggested Citation:"Front Matter." National Academies of Sciences, Engineering, and Medicine. 2019. Evaluating the Taxonomic Status of the Mexican Gray Wolf and the Red Wolf. Washington, DC: The National Academies Press. doi: 10.17226/25351.
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Suggested Citation:"Front Matter." National Academies of Sciences, Engineering, and Medicine. 2019. Evaluating the Taxonomic Status of the Mexican Gray Wolf and the Red Wolf. Washington, DC: The National Academies Press. doi: 10.17226/25351.
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Suggested Citation:"Front Matter." National Academies of Sciences, Engineering, and Medicine. 2019. Evaluating the Taxonomic Status of the Mexican Gray Wolf and the Red Wolf. Washington, DC: The National Academies Press. doi: 10.17226/25351.
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Suggested Citation:"Front Matter." National Academies of Sciences, Engineering, and Medicine. 2019. Evaluating the Taxonomic Status of the Mexican Gray Wolf and the Red Wolf. Washington, DC: The National Academies Press. doi: 10.17226/25351.
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Suggested Citation:"Front Matter." National Academies of Sciences, Engineering, and Medicine. 2019. Evaluating the Taxonomic Status of the Mexican Gray Wolf and the Red Wolf. Washington, DC: The National Academies Press. doi: 10.17226/25351.
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Suggested Citation:"Front Matter." National Academies of Sciences, Engineering, and Medicine. 2019. Evaluating the Taxonomic Status of the Mexican Gray Wolf and the Red Wolf. Washington, DC: The National Academies Press. doi: 10.17226/25351.
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Page viii Cite
Suggested Citation:"Front Matter." National Academies of Sciences, Engineering, and Medicine. 2019. Evaluating the Taxonomic Status of the Mexican Gray Wolf and the Red Wolf. Washington, DC: The National Academies Press. doi: 10.17226/25351.
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Suggested Citation:"Front Matter." National Academies of Sciences, Engineering, and Medicine. 2019. Evaluating the Taxonomic Status of the Mexican Gray Wolf and the Red Wolf. Washington, DC: The National Academies Press. doi: 10.17226/25351.
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Suggested Citation:"Front Matter." National Academies of Sciences, Engineering, and Medicine. 2019. Evaluating the Taxonomic Status of the Mexican Gray Wolf and the Red Wolf. Washington, DC: The National Academies Press. doi: 10.17226/25351.
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Suggested Citation:"Front Matter." National Academies of Sciences, Engineering, and Medicine. 2019. Evaluating the Taxonomic Status of the Mexican Gray Wolf and the Red Wolf. Washington, DC: The National Academies Press. doi: 10.17226/25351.
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Suggested Citation:"Front Matter." National Academies of Sciences, Engineering, and Medicine. 2019. Evaluating the Taxonomic Status of the Mexican Gray Wolf and the Red Wolf. Washington, DC: The National Academies Press. doi: 10.17226/25351.
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Page xiii Cite
Suggested Citation:"Front Matter." National Academies of Sciences, Engineering, and Medicine. 2019. Evaluating the Taxonomic Status of the Mexican Gray Wolf and the Red Wolf. Washington, DC: The National Academies Press. doi: 10.17226/25351.
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Suggested Citation:"Front Matter." National Academies of Sciences, Engineering, and Medicine. 2019. Evaluating the Taxonomic Status of the Mexican Gray Wolf and the Red Wolf. Washington, DC: The National Academies Press. doi: 10.17226/25351.
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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 Evaluating the Taxonomic Status of the Mexican Gray Wolf and the Red Wolf Committee on Assessing the Taxonomic Status of the Red Wolf and the Mexican Gray Wolf 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 Contract No. 140F0918C0005 between the National Academy of Sciences and the U.S. Fish and Wildlife Service. Any opinions, findings, conclusions, or recommendations ex- pressed 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/10.17226/25351 Additional copies of this publication are available for sale 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 2019 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. 2019. Evaluating the Taxonomic Status of the Mexican Gray Wolf and the Red Wolf. Washington, DC: The National Academies Press. doi: https://doi.org/10.17226/25351. Prepublication Copy

The National Academy of Sciences was established in 1863 by an Act of Congress, signed by Presi- dent 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 re- search. 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. C. D. Mote, Jr., 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 Med- icine to provide independent, objective analysis and advice to the nation and conduct other activ- ities 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. Prepublication Copy

Consensus Study Reports published by the National Academies of Sciences, Engineering, and Med- icine 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 Na- tional Academies. The statements and opinions contained in proceedings are those of the partici- pants and are not endorsed by other participants, the planning committee, or the National Acade- mies. For information about other products and activities of the National Academies, please visit www. nationalacademies.org/about/whatwedo. Prepublication Copy

COMMITTEE ON ASSESSING THE TAXONOMIC STATUS OF THE RED WOLF AND THE MEXICAN GRAY WOLF Chair JOSEPH TRAVIS, Florida State University Members FRED W. ALLENDORF, University of Montana DIANE K. BOYD, Montana Department of Fish, Wildlife, and Parks LILIANA CORTÉS ORTIZ, University of Michigan LORI S. EGGERT, University of Missouri DIANE GENEREUX, Broad Institute, Massachusetts Institute of Technology, Harvard University MICHAEL LYNCH (NAS), Arizona State University JESÚS E. MALDONADO, Smithsonian Conservation Biology Institute, National Zoological Park RASMUS NIELSEN, University of California, Berkeley Staff KEEGAN SAWYER, Study Director, Board on Life Sciences CAMILLA YANDOC ABLES, Senior Program Officer, Board on Agriculture and Natural Resources JENNA BRISCOE, Research Assistant, Board on Agriculture and Natural Resources ROBERT POOL, Editor Prepublication Copy v

BOARD ON LIFE SCIENCES Chair JAMES P. COLLINS, Arizona State University Members A. ALONSO AGUIRRE, George Mason University ENRIQUETA C. BOND (NAS), Burroughs Wellcome Fund DOMINIQUE BROSSARD, University of Wisconsin–Madison ROGER D. CONE (NAS, NAM), University of Michigan NANCY D. CONNELL, Johns Hopkins Center for Health Security SEAN M. DECATUR, Kenyon College JOSEPH R. ECKER (NAS), Howard Hughes Medical Institute SCOTT V. EDWARDS (NAS), Harvard University GERALD L. EPSTEIN, National Defense University ROBERT J. FULL, University of California, Berkeley ELIZABETH HEITMAN, University of Texas Southwestern Medical Center MARY E. MAXON, Lawrence Berkeley National Lab ROBERT NEWMAN, Independent Consultant STEPHEN J. O’BRIEN (NAS), Nova Southeastern University CLAIRE POMEROY (NAM), The Albert and Mary Lasker Foundation MARY E. POWER (NAS), University of California, Berkeley SUSAN RUNDELL SINGER, Rollins College LANA SKIRBOLL, Sanofi DAVID R. WALT (NAE, NAM), Harvard Medical School Staff FRAN SHARPLES, Director JO HUSBANDS, Senior Scholar KATHERINE BOWMAN, Senior Program Officer KEEGAN SAWYER, Senior Program Officer ANDREA HODGSON, Program Officer AUDREY THEVENON, Program Officer 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 SHANE C. BURGESS, University of Arizona, Tucson SUSAN CAPALBO, Oregon State University, Corvallis GAIL CZARNECKI-MAULDEN, Nestlé Purina PetCare, St. Louis, MO GEBISA EJETA, Purdue University, West Lafayette, IN JAMES S. FAMIGLIETTI, California Institute of Technology, Pasadena FRED GOULD (NAS), North Carolina State University, Raleigh DOUGLAS B. JACKSON-SMITH, The Ohio State University, Wooster JAMES W. JONES (NAE), University of Florida, Gainesville STEPHEN S. KELLEY, North Carolina State University, Raleigh JAN E. LEACH, Colorado State University, Fort Collins JILL J. MCCLUSKEY, Washington State University, Richland KAREN I. PLAUT, Purdue University, West Lafayette, IN JIM E. RIVIERE (NAM), Kansas State University, Manhattan Staff ROBIN A. SCHOEN, Director CAMILLA YANDOC ABLES, Senior Program Officer KARA N. LANEY, Senior Program Officer JENNA BRISCOE, Research Assistant Prepublication Copy vii

Preface This report is based on the intellectual contributions of a host of scientists, from those who wrote the initial descriptions of these animals to those who have published the latest genomic analyses. The commit- tee examined research published between now and almost a century ago, proving that good science is time- less. Beyond the written literature, the report is 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, thoroughly discussing the evidence in that lit- erature, and cheerfully writing draft after draft to make the report clear and accessible. On behalf of the committee, I thank our 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 Camilla Yandoc Ables for her assistance with many aspects of our work, especially her ability to provide us with the resources we needed to do a thorough review of the large literature on this subject. All of us on the committee are grateful to Jenna Briscoe for her support in organizing and conducting our meetings and to the National Academies of Sciences, Engineering, and Medicine librarians, Jorge Mendoza-Torres and Rebecca Morgan, without whose skills the committee might well have been lost. On behalf of the committee, I thank all those who informed us about wolves, from in-person presen- tations to webinars to comments sent via the study website. The Appendix includes their names and how they helped us. The committee’s knowledge of the critical issues was deepened and broadened by the con- tributions of so many people, and our thinking was constantly challenged by each increment in our knowledge. All who spoke or wrote to us were generous with their thoughts, patient in fielding our ques- tions, and unfailingly gracious in their comments. The committee’s gratitude for so much effort cannot be overstated. Joseph Travis Chair, Committee on Assessing the Taxonomic Status of the Red Wolf and the Mexican Gray Wolf Prepublication Copy ix

Acknowledgment of Reviewers This Consensus Study Report was reviewed in draft form by individuals chosen for their diverse per- spectives 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: Eric Gese, USDA-APHIS Frank Hailer, Cardiff University Jody Hey, Temple University Anna Linderholm, Texas A&M University James Mallet, Harvard University Daniel Rubenstein, Princeton University Elizabeth Thompson (NAS), University of Washington (retired) Lisette Waits, University of Idaho Robert Wayne, University of California, Los Angeles Although the reviewers listed above provided many constructive comments and suggestions, they were not asked to endorse the conclusions or recommendations of this report nor did they see the final draft before its release. The review of this report was overseen by Dr. James A. Estes (NAS), University of California, Santa Cruz (emeritus), and Dr. Barbara A. Schaal (NAS), Washington University in St. Louis. 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 ..................................................................................................................................... 6 Purpose of the Study, 6 What Is a Wolf?, 8 Evolution and Taxonomy, 8 The Key Issues in Recognizing Species and Subspecies, 11 Organization of the Report, 12 References, 12 2 GUIDING PRINCIPLES FOR IDENTIFYING SPECIES AND SUBSPECIES .............................. 14 What Is a “Species”? What Is a “Subspecies”?, 14 Understanding “Species” in Light of Hybridization, 17 Establishing Guidelines for Determining Taxonomic Status, 21 A Framework for Establishing Taxonomic Designations, 21 References, 22 3 USING GENES AND GENOMES TO IDENTIFY SPECIES AND SUBSPECIES ......................... 25 Individual Nuclear Genes, 26 mtDNA, 27 Genomes, 28 References, 33 4 IS THE MEXICAN GRAY WOLF A VALID SUBSPECIES? ........................................................... 35 A Brief Taxonomic History of the Mexican Gray Wolf, 35 Is There Evidence for Distinctiveness of Mexican Gray Wolf Populations from Other North American Canis Populations?, 37 Is There Evidence for Continuity between the Historic Mexican Gray Wolf Lineage and the Present Managed Populations?, 40 Synthesis of Findings, 41 References, 41 5 IS THE RED WOLF A VALID TAXONOMIC SPECIES?................................................................ 45 A Brief Taxonomic History of the Red Wolf, 45 Is There Evidence that the Historical Population of Red Wolves Was a Distinct Lineage?, 45 Is There Evidence for Distinctiveness of Contemporary Red Wolves Populations from Gray Wolves and Coyotes?, 50 Is There Evidence for Continuity between the Historic Red Wolf Population and the Present Managed Populations?, 56 Synthesis of Findings, 60 References, 61 GLOSSARY ...................................................................................................................................................... 65 Prepublication Copy xiii

Contents APPENDIXES A OPEN SESSION MEETING AGENDAS .............................................................................................. 70 B LIST OF WEBINARS AND SOLICITED EXPERT INPUT .............................................................. 73 C BIOGRAPHICAL SKETCHES OF COMMITTEE MEMBERS AND PROJECT STAFF ......................................................................................................................... 74 xiv Prepublication Copy

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Scientists strive to develop clear rules for naming and grouping living organisms. But taxonomy, the scientific study of biological classification and evolution, is often highly debated. Members of a species, the fundamental unit of taxonomy and evolution, share a common evolutionary history and a common evolutionary path to the future. Yet, it can be difficult to determine whether the evolutionary history or future of a population is sufficiently distinct to designate it as a unique species.

A species is not a fixed entity – the relationship among the members of the same species is only a snapshot of a moment in time. Different populations of the same species can be in different stages in the process of species formation or dissolution. In some cases hybridization and introgression can create enormous challenges in interpreting data on genetic distinctions between groups. Hybridization is far more common in the evolutionary history of many species than previously recognized. As a result, the precise taxonomic status of an organism may be highly debated. This is the current case with the Mexican gray wolf (Canis lupus baileyi) and the red wolf (Canis rufus), and this report assesses the taxonomic status for each.

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