Monarch butterflies are among the most popular insect species in the world and are an icon for conservation groups and environmental education programs. Monarch caterpillars and adults are easily recognizable as welcome visitors to gardens in North America and beyond, and their spectacular migration in eastern North America (from breeding locations in Canada and the United States to overwintering sites in Mexico) has captured the imagination of the public. Monarch migration, behavior, and chemical ecology have been studied for decades. Yet many aspects of monarch biology have come to light in only the past few years. These aspects include questions regarding large-scale trends in monarch population sizes, monarch interactions with pathogens and insect predators, and monarch molecular genetics and large-scale evolution. A growing number of current research findings build on the observations of citizen scientists, who monitor monarch migration, reproduction, survival, and disease. Monarchs face new threats from humans as they navigate a changing landscape marked by deforestation, pesticides, genetically modified crops, and a changing climate, all of which place the future of monarchs and their amazing migration in peril. To meet the demand for a timely synthesis of monarch biology, conservation and outreach, Monarchs in a Changing World summarizes recent developments in scientific research, highlights challenges and responses to threats to monarch conservation, and showcases the many ways that monarchs are used in citizen science programs, outreach, and education. It examines issues pertaining to the eastern and western North American migratory populations, as well as to monarchs in South America, the Pacific and Caribbean Islands, and Europe. The target audience includes entomologists, population biologists, conservation policymakers, and K-12 teachers.
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First published 2015 by Cornell University Press
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Library of Congress CataloginginPublication Data
Monarchs in a changing world : biology and conservation of an iconic butterfly / edited by Karen S. Oberhauser, Kelly R. Nail, Sonia Altizer. pages cm Includes bibliographical references and index. ISBN 9780801453151 (cloth : alk. paper) 1. Monarch butterfly. 2. Monarch butterfly—Conservation—North America. I. Oberhauser, Karen Suzanne, editor. II. Nail, Kelly R., editor. III. Altizer, Sonia M., editor. IV. Oberhauser, Karen Suzanne. Monarchs and people. Container of (work): QL561.D3M6635 2015 595.78'9—dc23 2014034844
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Cover photograph: Three monarchs congregating in the early morning at the Cerro Pelon overwintering colony in Mexico (February 2013). Photograph by Kelly R. Nail.
C o n t e n t s
Prefaceix
Mode Programs or Cîtîzen Scîence, Educatîon, and Conservatîon: An Overvîew Karen S. Oberhauser1 1. Environmental Education and Monarchs: Reaching across Disciplines, Generations, and Nations Elisabeth YoungIsebrand, Karen S. Oberhauser, Kim Bailey, Sonya Charest, Brian Hayes, Elizabeth Howard, Jim Lovett, Susan Meyers, Erik Mollenhauer, Eneida B. MontesiñosPatino, Ann Ryan, Orley R. Taylor, and Rocío Treviño 5 2. Contributions to Monarch Biology and Conservation through Citizen Science: Seventy Years and Counting Karen S. Oberhauser, Leslie Ries, Sonia Altizer, Rebecca V. Batalden, Janet KudellEkstrum, Mark Garland, Elizabeth Howard, Sarina Jepsen, Jim Lovett, Mía Monroe, Gail Morris, Eduardo RendónSalinas, Richard G. RuBino, Ann Ryan, Orley R. Taylor, Rocío Treviño, Francis X. Villablanca, and Dick Walton 13 3. Monarch Habitat Conservation across North America: Past Progress and Future Needs Priya C. Shahani, Guadalupe del Río Pesado, Phil Schappert, and Eligio García Serrano 31
Monarcs as Herbîvores, Prey, and Hosts: An Overvîew Jacobus C. de Roode 43 4. Macroevolutionary Trends in the Defense of Milkweeds against Monarchs: Latex, Cardenolides, and Tolerance of Herbivory Anurag A. Agrawal, Jared G. Ali, Sergio Rasmann, and Mark Fishbein 47 5. Invertebrate Natural Enemies and StageSpecific Mortality Rates of Monarch Eggs and Larvae Alma De Anda and Karen S. Oberhauser 60 6. Lacewings, Wasps, and Flies—Oh My: Insect Enemies Take a Bite out of Monarchs Karen S. Oberhauser, Michael Anderson, Sophia Anderson, Wendy Caldwell, Alma De Anda, Mark Hunter, Matthew C. Kaiser, and Michelle J. Solensky 71 7. Monarchs and Their Debilitating Parasites: Immunity, Migration, and Medicinal Plant Use Sonia Altizer and Jacobus C. de Roode 83
Monarcs în a Cangîng Cîmate: An Overvîew Kelly R. Nail and Karen S. Oberhauser 95 8. What’s Too Hot and What’s Too Cold? Lethal and Sublethal Effects of Extreme Temperatures on Developing Monarchs Kelly R. Nail, Rebecca V. Batalden, and Karen S. Oberhauser 99
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viContents
9.Microclimatic Protection of Overwintering Monarchs Provided by Mexico’s HighElevation Oyamel Fir Forests: A Review Ernest H. Williams and Lincoln P. Brower 109 10. Effect of the 2010–2011 Drought on the Lipid Content of Monarchs Migrating through Texas to Overwintering Sites in Mexico Lincoln P. Brower, Linda S. Fink, Ridlon J. Kiphart, Victoria Pocius, Raúl R. Zubieta, and M. Isabel Ramírez 117 11. Estimating the Climate Signal in Monarch Population Decline: No Direct Evidence for an Impact of Climate Change? Myron P. Zalucki, Lincoln P. Brower, Stephen B. Malcolm, and Benjamin H. Slager 130
Conservîng Nort Amerîcan Monarc Butterlîes: An Overvîew Lincoln P. Brower and Linda S. Fink 143 12. Understanding and Conserving the Western North American Monarch Population Sarina Jepsen and Scott Hoffman Black 147 13. Threats to the Availability of Overwintering Habitat in the Monarch Butterfly Biosphere Reserve: Land Use and Climate Change M. Isabel Ramírez, Cuauhtémoc SáenzRomero, Gerald Rehfeldt, and Lidia SalasCanela 157 14. Monarch Butterflies and Agriculture John M. Pleasants 169 15. Fires and Fire Management in the Monarch Butterfly Biosphere Reserve Héctor MartínezTorres, Mariana CantúFernández, M. Isabel Ramírez, and Diego R. PérezSalicrup 179 16. Project Milkweed: A Strategy for Monarch Habitat Conservation Brianna Borders and Eric LeeMäder 190 17. Grassland and Roadside Management Practices Affect Milkweed Abundance and Opportunities for Monarch Recruitment Kristen A. Baum and Elisha K. Mueller 197
New Perspectîves on Monarc Mîgratîon, Evoutîon, and Popuatîon Bîoogy: An Overvîew Andrew K. Davis and Sonia Altizer 203 18. Tracking the Fall Migration of Eastern Monarchs with Journey North Roost Sightings: New Findings about the Pace of Fall Migration Elizabeth Howard and Andrew K. Davis 207 19. Potential Changes in Eastern North American Monarch Migration in Response to an Introduced Milkweed,Asclepias curassavicaRebecca V. Batalden and Karen S. Oberhauser 215 20. Migration and Host Plant Use by the Southern Monarch,Danaus erippus Stephen B. Malcolm and Benjamin H. Slager 225 21. Monarchs in the Mist: New Perspectives on Monarch Distribution in the Pacific Northwest Robert Michael Pyle 236
22. Monarchs across the Atlantic Ocean: What’s Happening on the Other Shore? Juan FernándezHaeger, Diego Jordano, and Myron P. Zalucki 247 23. Unraveling the Mysteries of Monarch Migration and Global Dispersal through Molecular Genetic Techniques Amanda A. Pierce, Sonia Altizer, Nicola L. Chamberlain, Marcus R. Kronforst, and Jacobus C. de Roode 257 24. Connecting Eastern Monarch Population Dynamics across Their Migratory Cycle Leslie Ries, Douglas J. Taron, Eduardo RendónSalinas, and Karen S. Oberhauser
References283 Contributors313 Index317
Color plates are at the end of the e-book
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Contents
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P r e f a c e
Weer you are a monarc cîîzen scîenîs, an enomoogîs, a popuaîon bîoogîs, a conservaîon poîcy maker, a eacer, or jus îneresed în mon-arcs’ amazîng bîoogy and eîr împressîve annua mîgraory cyce, you are readîng îs book because monarcs ascînae you. Wî conrîbuîons rom dozens o îndîvîduas across e gobe,Monarcs în a Cangîng Word: Bîoogy and Conservatîon o an ïconîc Butterlye unîque and remark- îgîgs abe naura îsory o monarcs and eîr compex and muîaceed îneracîons wî peope. Fîve înernaîona conerences ave been ed on monarc bîoogy and conservaîon, our o wîc rîggered e creaîon o edîed voumes: e Sympo-sîum on e Bîoogy and Conservaîon o e Mon-arc BuerLy (Moreos, Mexîco, 1981), e Second ïnernaîona Conerence on e Monarc BuerLy (os Angees, Caîornîa, 1986; Macom and Zauckî 1993), e Nor Amerîcan Conerence on e Mon-arc BuerLy (Moreîa, Mîcoacán, 1997; o e a. 1999), e Monarc Popuaîon Dynamîcs Con-erence (awrence, Kansas, 2001; Oberauser and Soensky 2004), and e Monarc Bîoogy and Con-servaîon Meeîng (Mînneapoîs, Mînnesoa, 2012; îs voume). Wî ree exîsîng compendîa o monarc bîo-ogy and conservaîon, wy do we need anoer one? Here as been a verîabe exposîon o knowedge abou monarcs în e as decade. Monarcs con-înue o provîde a wîndow îno some o e mos ascînaîng quesîons acîng bîoogîss and e pub-îc, and we’ve compîed recen Indîngs a uîîze cuîng-edge geneîc oos and anayîca ecnîques,
as we as rîed-and-rue meods în aboraory and Ied bîoogy. Sînce pubîcaîon o e as voume în 2004, e Nor Amerîcan Monarc Conservaîon Pan was pubîsed (CEC 2008) and e Monarc Joîn Venure was ormed (Monarc Joîn Venure 2013). Hese efors ave broug even more aen-îon o monarc conservaîon, wî a concomîan încrease în oca, regîona, naîona, and înerna-îona conservaîon efors. He pas decade as aso broug an exposîon o îneres by cîîzens în coec-îng scîenîIc daa; ese cîîzen scîenîss înves ens o ousands o vouneer ours în monîorîng mon-arcs every year. A e same îme, monarc num-bers are decreasîng în response o envîronmena canges broug on by abîa oss and oer sress-ors; îke many oer organîsms, monarcs requîre our aenîon and care o survîve ese canges. Now îs a crucîa îme or monarcs, and îs book îs no ony a ceebraîon o eîr amazîng bîoogy and our ove o îs carîsmaîc însec, bu aso a ca o arms. He cuîng-edge scîenîIc deveopmens descrîbed în e oowîng capers buîd on a ong radîîon o researc. Monarcs ave eped answer unda-mena bîoogîca quesîons abou ow organîsms mîgrae, Ind and dîges ood, and cope wî a word în wîc naura enemîes aack em as eggs, ar-vae, pupae, and adus. Hey’ve aso been e ocus o sudîes a ave eped eucîdae ow genes are ransaed îno moecues a aow organîsms o uncîon approprîaey as ey age and ace a dîver-sîy o envîronmena condîîons. New aspecs o monarc bîoogy ave come o îg în e pas ew years, încudîng undersandîng o arge-scae rends