Rich Longcore ECOLOGY Advance praise for Ecological Consequences of Artificial Night Lighting E c Ecological Consequences “As a kid, I spent many a night under streetlamps looking for toads and bugs, or o l simply watching the bats. The two dozen experts who wrote this text still do. This o of isis aa definitive,definitive, readable,readable, comprehensivecomprehensive reviewreview ofof howhow artificialartificial nightnight lightinglighting affectsaffects g animals and plants. The reader learns about possible and definite effects of i animals and plants. The reader learns about possible and definite effects of c Artificial Night Lighting photopollution, illustrated with important examples of how to mitigate these effects a on species ranging from sea turtles to moths. Each section is introduced by a l delightful vignette that sends you rushing back to your own nighttime adventures, C be they chasing fireflies or grabbing frogs.” o n —JOHN M. MARZLUFF,, DenmanDenman ProfessorProfessor ofof SustainableSustainable ResourceResource Sciences,Sciences, s College of Forest Resources, University of Washington e q “This book is that rare phenomenon, one that provides us with a unique, relevant, and u seminal contribution to our knowledge, examining the physiological, behavioral, e n reproductive, community,community, and other ecological effectseffects of light pollution. It will c enhance our ability to mitigate this ominous envirenvironmentalonmental alteration thrthroughough mormoree e conscious and effective design of the built environment.” s TEPHEN ELLERT o —STEPHEN R. KELLERT,, TweedyTweedy OrdwayOrdway ProfessorProfessor ofof SocialSocial Ecology,Ecology, f YaleYale University, School of Forestry and Environmental Studies A r “A marmarvelous,velous, up-to-date summarsummaryy of the many negative influences of overabundant t i night lighting, or light pollution, on life of all kinds: mammals, birds, reptiles, fish, f i insects,insects, andand plants.”plants.” c i —ARRTHURTHUR UPGREN,, seniorsenior rresearchesearch scientist, YYaleale UniversityUniversity,, andand a l emeritus professorprofessor of astronomy, Wesleyan University N i g Catherine Rich holds degrees in psychology, geography, and law. Travis Longcore h isis researchresearch assistant professor of geography at the University of Southern California t and lectures regularly at University of California at Los Angeles. They co-founded L and run The Urban Wildlands Group, a Los Angeles–based conservation nonprofit. i g h Cover design by Brighid Willson t i Cover photo: “Wolf’s“Wolf’s Moon” by R. N. Cohen ISBN 1-55963-129-5 n g 90000 Edited by Washington • Covelo • London Catherine Rich • Travis Longcore www.islandpress.org All Island Press books are printed on recycled, All Island Press books are printed on recycled, 9 781559 631297 acid-freeacid-free paper.paper. About Island Press Island Press is the only nonprofit organization in the United States whose principal purpose is the publication of books on environmental issues and natural resource management. We provide solutions-oriented informa- tion to professionals, public officials, business and community leaders, and concerned citizens who are shaping responses to environmental problems. In 2006, Island Press celebrates its twenty-first anniversary as the leading provider of timely and practical books that take a multidiscipli- nary approach to critical environmental concerns. Our growing list of titles reflects our commitment to bringing the best of an expanding body of literature to the environmental community throughout North Amer- ica and the world. Support for Island Press is provided by the Agua Fund, The Geraldine R. Dodge Foundation, Doris Duke Charitable Foundation, The William and Flora Hewlett Foundation, Kendeda Sustainability Fund of the Tides Foundation, Forrest C. Lattner Foundation, The Henry Luce Founda- tion, The John D. and Catherine T. MacArthur Foundation, The Marisla Foundation, The Andrew W. Mellon Foundation, Gordon and Betty Moore Foundation, The Curtis and Edith Munson Foundation, Oak Foundation, The Overbrook Foundation, The David and Lucile Packard Foundation, The Winslow Foundation, and other generous donors. The opinions expressed in this book are those of the author(s) and do not necessarily reflect the views of these foundations. Ecological Consequences of Artificial Night Lighting Ecological Consequences of Artificial Night Lighting Edited by Catherine Rich • Travis Longcore / Washington • Covelo • London Copyright © 2006 Island Press All rights reserved under International and Pan-American Copyright Conventions. No part of this book may be reproduced in any form or by any means without permission in writing from the publisher: Island Press, 1718 Connecticut Avenue, N.W., Suite 300, Washington, DC 20009. ISLAND PRESS is a trademark of The Center for Resource Economics. No copyright claim is made in the work of Robert N. Fisher, an employee of the federal government. Library of Congress Cataloging-in-Publication Data Ecological consequences of artificial night lighting / edited by Catherine Rich and Travis Longcore p. cm. Includes bibliographical references. ISBN 1–55963–128–7 (cloth : alk. paper) — ISBN 1–55963–129–5 (pbk. : alk. paper) 1. Exterior lighting—Environmental aspects. I. Rich, Catherine. II. Longcore, Travis. QH545.E98E26 2005 577.27'2—dc22 2005020202 British Cataloguing-in-Publication Data available. Book design: Brighid Willson Printed on recycled, acid-free paper Manufactured in the United States of America 10 9 8 7 6 5 4 3 2 1 For our parents, and in memory of Cherryl Wilson Night is certainly more novel and less profane than day. —HENRY DAVID THOREAU (1817–1862), “Night and Moonlight,” 1863 Contents Preface xiii 1. Introduction 1 Catherine Rich and Travis Longcore PART I. MAMMALS Night, Venezuela • Alexander von Humboldt 16 2. Effects of Artificial Night Lighting on Terrestrial Mammals 19 Paul Beier 3. Bats and Their Insect Prey at Streetlights 43 Jens Rydell PART II. BIRDS Night, Maine Woods • Bernd Heinrich 62 4. Effects of Artificial Night Lighting on Migrating Birds 67 Sidney A. Gauthreaux Jr. and Carroll G. Belser xi xii Contents 5. Influences of Artificial Light on Marine Birds 94 William A. Montevecchi 6. Road Lighting and Grassland Birds: Local Influence of Road Lighting on a Black-Tailed Godwit Population 114 Johannes G. de Molenaar, Maria E. Sanders, and Dick A. Jonkers PART III. REPTILES AND AMPHIBIANS Night, Tortuguero • David Ehrenfeld 138 7. Protecting Sea Turtles from Artificial Night Lighting at Florida’s Oceanic Beaches 141 Michael Salmon 8. Night Lights and Reptiles: Observed and Potential Effects 169 Gad Perry and Robert N. Fisher 9. Observed and Potential Effects of Artificial Night Lighting on Anuran Amphibians 192 Bryant W. Buchanan 10. Influence of Artificial Illumination on the Nocturnal Behavior and Physiology of Salamanders 221 Sharon E. Wise and Bryant W. Buchanan PART IV. FISHES Night, Atlantic • Carl Safina 253 11. Artificial Night Lighting and Fishes 257 Barbara Nightingale, Travis Longcore, and Charles A. Simenstad Contents xiii PART V. INVERTEBRATES Night, Tropics • Philip J. DeVries 278 12. Artificial Night Lighting and Insects: Attraction of Insects to Streetlamps in a Rural Setting in Germany 281 Gerhard Eisenbeis 13. Effects of Artificial Night Lighting on Moths 305 Kenneth D. Frank 14. Stray Light, Fireflies, and Fireflyers 345 James E. Lloyd 15. Artificial Light at Night in Freshwater Habitats and Its Potential Ecological Effects 365 Marianne V. Moore, Susan J. Kohler, and Melani S. Cheers PART VI. PLANTS Night, Massachusetts • Henry David Thoreau 386 16. Physiology of Plant Responses to Artificial Lighting 389 Winslow R. Briggs 17. Synthesis 413 Travis Longcore and Catherine Rich About the Contributors 431 Index 435 Preface For people who care about the natural world, the unrelentingly destruc- tive activities of humanity are all too familiar. Most of these are painfully obvious to anyone watching the transformation of long-appreciated land- scapes. We know death by bulldozer when we see it. Seeing may be the key. We are for the most part diurnal, and visual, and as we consider the transformation of the planet, mostly we envision changes that we are able to see by day. It is an inherent bias in our collec- tive thinking. But as we light up the night, with ever more powerful lights, is it not obvious, giving the matter some thought, that we are wreaking havoc on creatures with physiologies far more delicate than ours, inter- fering with the lifeways of suites of organisms that have evolved over the millennia with a dependable pattern of light and dark? All this happens at times when most of us are sleeping. As the night is ever more brightly lit, at least we can close the shutters. But what of the animals and plants? I imagine myself in the position of other creatures, unable to control or escape their environments, which we have controlled for them—whales subject to sonar far beyond any sound their ears were meant to accommo- date, birds migrating thousands of miles, thrown off course by lights with no limits. We are ultimately deprived of their beauty as they die off by the millions, from our careless expansion into all realms of the Earth. My own love for night started simply as an aesthetic appreciation, and over the last twenty-five years or so I have been every year more pro- foundly saddened by the glare foisted on us by brighter and brighter lights. Long ago, as an undergraduate student, I loved to walk the neighborhoods