AVIAN ECOLOGY AND CONSERVATION

IN AN URBANIZING WORLD

AVIAN ECOLOGY AND CONSERVATION

IN AN URBANIZING WORLD

edited by

John M. Marzluff College of Forest Resources

University of Washington

Reed Bowman Archbold Biological Station

Roarke Donnelly College of Forest Resources

University of Washington

SPRINGER SCIENCE+BUSINESS MEDIA, LLC

Library of Congress Cataloging-in-Publication Data

Avian ecology and conservation in an urbanizing world / edited by John Marzluff, Reed Bowman, Roarke Donnelly,

p. cm. Includes bibliographical references (p. ). ISBN 978-1-4613-5600-4 ISBN 978-1-4615-1531-9 (eBook) DOI 10.1007/978-1-4615-1531-9 1. Birds—Ecology. 2. Urbanization—Environmental aspects. 3. Birds, Protection of. I.

Marzluff, John M . II. Bowman, Reed. III. Donnelly, Roarke.

QL698.95 .A84 2001 598.17—dc21

2001042818

Copyright © 2001 by Springer Science+Business Media New York Originally published by Kluwer Academic Publishers in 2001 Softcover reprint of the hardcover 1st edition 2001

A l l rights reserved. No part of this publication may be reproduced, stored in a retrieval system or transmitted in any form or by any means, mechanical, photo­copying, recording, or otherwise, without the prior written permission of the publisher, Springer Science+Business Media, L L C .

Printed on acid-free paper.

TABLE OF CONTENTS

Preface ix

SECTION 1. Introduction to the Study of Birds in Urban Environments

1. A historical perspective on urban bird research: trends, terms, andapproaches

John M. Marzluff, Reed Bowman, and Roarke Donnelly 1

2. Worldwide urbanization and its effects on birds

John M. Marzluff 19

3. Synanthropic birds of North America

Richard F. Johnston 49

4. Human perception and appreciation of birds: A motivation forwildlife conservation in urban environments of France

Philippe Clergeau, Gwenaelle Mennechez, Andre Sauvage, and AgnesLemoine 69

5. Quantifying the urban gradient: linking urban planning and ecology

Marina Alberti, Erik Botsford, and Alex Cohen 89

6. Urbanization, avian communities, and landscape ecology

James R. Miller, Jennifer M. Fraterrigo, N. Thompson Hobbs, David M.Theobald, and John A. Wiens 117

7. The importance of multi-scale analyses in avian habitat selectionstudies in urban environments

Mark Hostetler 139

SECTION 2. Processes Affecting Birds in Urban Environments

8. Urban birds: Population, community, and landscape approaches

Douglas Bolger 155

9. Interactions among non-native plants and birds

Sarah Hayden Reichard, Linda Chalker-Scott, andSolai Buchanan 179

VI

10. Urban sprawl and juniper encroachment effects on abundance ofwintering passerines in Oklahoma

Bryan R. Coppedge, David M. Engle, Samuel D. Fuhlendorf, Ronald E.Masters, and Mark S. Gregory 225

II. Nest predator abundance and urbanization

David G. Haskell, A. M. Knupp, and M. C. Schneider 243

12. Bird tolerance to human disturbance in urban parks of Madrid(Spain): Management implications

Esteban Fermindez-Juricic, Maria Dolores Jiminez, and Elena Lucas 259

13. Settlement of breeding European Starlings in urban areas:importance of lawns vs. anthropogenic wastes

Gwenaelle Mennechez and Philippe Clergeau 275

14. Variation in the timing of breeding between suburban and wildlandFlorida Scrub-Jays: Do physiologic measures reflect differentenvironments?

Stephan J. Schoech and Reed Bowman 289

SECTION 3. Bird Populations in Urban Environments

15. The ecology of Western Gulls in habitats varying in degree of urbaninfluence

Raymond Pierotti and Cynthia Annett 307

16. Causes and consequences of expanding American Crow populations

John M. Marzluff, Kevin J. McGowan, Roarke Donnelly, and Richard L.Knight 331

17. Demographic and behavioral comparisons of suburban and ruralAmerican Crows

Kevin McGowan 365

18. Nest success and the timing of nest failure of Florida Scrub-Jays insuburban and wildland habitats

Reed Bowman and Glen E. Woolfenden 383

19. Synurbanization of the Magpie in the Palearctic

Leszek Jerzak 403

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20. Macaw abundance in relation to human population density in thewestern Amazon basin

Daniel M. Brooks and Alfredo J. Begazo 427

21. Waterbird production in an urban center in Alaska

Michael R. North 439

SECTION 4. Bird Communities in Urban Environments

22. Creating a homogeneous avifauna

Robert B. Blair 459

23. Avian community characteristics of urban greenspaces in St. Louis,Missouri

Jeffrey M. Azerrad and Charles H. Nilon 487

24. The importance of the Chicago region and the "ChicagoWilderness" initiative for avian conservation

Jeffrey D. Brawn and Douglas F. Stotz 509

25. Do temporal trends in Christmas Bird Counts reflect the spatialtrends of urbanization in southwestern Ohio?

Nancy A. Crosby and Robert B. Blair 523

26. Survey techniques and habitat relationships of breeding birds inresidential areas of Toronto, Canada

Jean-Pierre L. Savard and J. Bruce Falls 543

SECTION 5. SYNTHESIS

27. Integrating avian ecology into emerging paradigms in urban ecology

Reed Bowman and John M. Marzluff.. 569

PREFACE

Humans dominate earth's ecosystems in a variety of ways. One of themost striking and persistent is our ability to change land-cover as we settle aregion. Settlement modifies water and sediment fluxes; reduces, andfragments natural habitats; introduces exotic organisms; severely altersenergy flow and nutrient cycles; and overstresses entire ecosystems.Despite these important effects, the ecological implications of settlementand resulting urbanization are just beginning to be understood. Much of ourunderstanding derives from studies of birds, yet the existing literature isscattered, mostly decades old, and rarely synthesized or standardized. Inaddition, a new paradigm for urban ecological research is emerging, onebased solidly on traditional ecological theory. Increasingly, studies of birdsin urban environments consider the mechanisms by which human changesaffect birds, they consider these effects at multiple levels of ecologicalorganization, including the community, population, and individuals, andthey consider these effects across a gradient of different levels ofurbanization. Therefore, we felt it timely to summarize existing studies,suggest standard approaches, and showcase ongoing studies of birds inurban environments.

We investigate how birds respond to human settlement in 27 chaptersauthored by leaders in this expanding field. Our objectives were to: 1)review what is known about birds in urbanizing settings, 2) suggestimportant ways to standardize our study of birds and urbanization, 3)investigate connections between urbanization and individual birds, thedemography of bird populations and the structure and composition of birdcommunities, 4) probe the evolutionary importance of urbanization to birds,5) understand how best to conserve birds in the face of increasingurbanization, 6) investigate the policy and management implications ofurban bird studies, and 7) suggest important areas where research is neededto better understand these topics. We introduce these objectives andsummarize the authors' progress at fulfilling them in an introductory andsummary chapter.

This volume stands alone as a summary of current research on birds insettled environments ranging from wildlands to exurban/rural to urban. Assuch, we feel it would be suitable for the ecologist, land manager, wildlifemanager, evolutionary ecologist, urban planner, landscape architect, andconservation biologist. This wide array of professionals will find ourinformation useful because we address the conservation and evolutionaryimplications of urban life from an ecological and planning perspective.Graduate students in these fields also will find the volume to be a useful

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Graduate students in these fields also will find the volume to be a usefulsummary and synthesis of current research, extant literature, andprescriptions for future work. All interested in human-driven land-coverchanges will benefit from a perusal of this book because we present highaltitude photographs of each study area. Simply looking at these imagesshows the variety of ways in which human settlement affects earth--from thetropics to the arctic and from intensive urban centers to small nativesettlements.

We organized this volume into five sections. Section I introduces thestudy of birds in urban environments. Section II investigates the importantanthropogenic processes that affect birds in urban environments. SectionsIII and IV provide case studies of bird populations and communities inurban settings. These are all new empirical works, often summarizingdecades of research, that clearly relate the processes discussed in Section IIto demographic and community-level processes. Section V is a singlesynthetic chapter summarizing the book and highlighting its implications forfuture studies.

In Section I, the reader learns: how we have studied birds in urbanenvironments (Chp. 1), standard definitions of settlement (Chp. 1), howhumans have settled Earth (Chp. 2), how birds respond to settlement (Chps.2, 3), the degree to which North American birds are synanthropic (Chp. 3),how human perception and appreciation of birds is affected by urbanization(Chp. 4), how to accurately quantify urban settlement patterns so they canrigorously be related to birds and other organisms (Chp. 5), and how toconsider multiple scales in our study of birds (Chps. 6, 7,). These latterchapters emphasize the need to consider many scales in ecological study aswell as policy implementation. We need to understand how individualanimals and landscapes respond to urbanization and then use thisunderstanding to motivate individual landowners and increasingly complexgovernments to plan urbanization in ways compatible with biodiversity.

Armed with this background, we discuss important mechanisms thataffect urban birds in Section II. The importance of "top down" mechanismssuch as predation and competition are contrasted with more subtle, lessstudied "bottom up" mechanisms such as arthropod availability (Chp. 8).Changes in vegetation caused by settlement are reviewed in detail (Chp. 9)and specifically related to bird changes in the midwestern United States(Chp. 10). The role birds play in aiding the spread of invasive plants is alsodiscussed (Chp. 9). The well-studied influence of urbanization on avian nestpredators is discussed in a case study from the American South (Chp. 11)and related to fragmentation of the American West (Chp. 8). Humans

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vIsItmg parks disturb native birds, but susceptibility depends on parkvegetation, amount of visitation, and bird size (Chp. 12). These resultscome from Europe, but have broad applicability to the design of trails inurban parks. European Starlings are one of the world's most successfulurban birds. Part of their success stems from their ability to utilize lawnsand anthropogenic foods in urban and suburban settings (Chp. 13). Foodsprovided to birds in suburban settings produce complex physiologicalresponses that are now amenable to study. For example, suburban FloridaScrub-jays have elevated plasma protein levels prior to egg layingsuggesting that anthropogenic foods may provide essential nutrients that, intum, cue breeders to lay eggs (Chp. 14). This may allow suburban jays tobreed earlier than wildland jays, but does not guarantee them greaterfledgling success (see below). The general mechanisms we cover in SectionII (exotic vegetation, food availability, exotic competitors, predation, andhuman disturbance) apply worldwide and are the primary reasons why birdpopulations and communities change in response to urbanization.

We illustrate many of the mechanisms discussed in Section II withspecific examples of how they affect bird populations in Section III.Western Gulls decline in response to urbanization because of changes innesting habitat and available food (Chp. 15). This is striking because gullsare usually thought of as being strongly synanthropic (Chp. 3). But, long­term observations and detailed experiments on Western Gulls indicate thatrelying on human refuse for food reduces reproductive performance (Chp.15). On the other hand, corvids appear to more effectively use human foodsand their populations increase in response to urbanization. American Crows(Chps. 16 and 17) appear to be most successful in suburban and ruralenvironments, but this success may fuel dramatic increases in urban settingsas well. Florida scrub-jays (Chp. 18) also use anthropogenic foods, butdifferences in food and predators between wildland and suburban areasappear to have a complex and interactive effect on nest success. Europeanmagpies are increasing rapidly in suburban and urban areas of the Palearctic(Chp. 19) because their reproduction and survivorship is increased as theyutilize anthropogenic foods and flourish in response to predator reductions.Human settlement of the Amazon and Arctic is low at present, but chapters20 and 21 anticipate the potential effects on native birds should settlement inthese regions increase. Parrots are likely to decline in the Amazon inresponse to direct harvest, changes in fruit abundance, and habitatmodification. Reserves that protect these species will likely need to be farfrom even small human settlements. Waterbirds in the Arctic are likely tohave mixed responses to human settlement; many may benefit from man-

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made water bodes while others may decline in response to wetland drainingand human use of wetlands. At least in Alaska, current policies (CleanWater Act) can be used to conserve these sensitive areas.

The demographic responses of birds to urbanization detailed in SectionIII eventually lead to restructuring of avian communities. Approaches andintroductions to such community-level responses were investigated earlier(Chps. 2, 6, 7, 8). In Section IV a series of empirical studies of aviancommunities in urbanizing areas is presented. Urban bird communitiesworldwide are characterized by ecologically similar species. Often, theyeven host identical species. This homogenization of avifaunas is exploredby contrasting western and mid-western sites in the United States (Chp. 22),but its implication for worldwide avian diversity is sobering. Many recentinvestigators of bird communities relate changes in land-cover, land-use,and resulting landscape patterns to community assembly. New empiricalwork on avian communities in St. Louis (Missouri), Chicago (Illinois),Cincinnati (Ohio), and Toronto are detailed in Chapters 23-26. Thesestudies quantify the large diversity of birds living in some of the mosturbanized regions of North America. They suggest that despite extensiveland-cover change in these areas, many native birds persist and rely on theseareas for breeding (Chps. 23, 26), during migration (Chp. 24), and overwinter (Chp. 25). Challenges abound to conserve diverse, native birdcommunities in highly urban areas, but these studies suggest that a variety ofpolicy instruments (e.g., greenspace reserves and private-publicpartnerships) are feasible and effective. The extensive group working toconserve native areas in the Chicago region is exemplary and is being usedas a model for international efforts (Chp. 24).

We conclude the book in a final section with a single chaptersummarizing the main points made by contributors and reflecting on thetype of research that is still needed to increase our ability to provide forbirds in the face of global urbanization. Current research suggests that theeffect of urbanization on birds is immense, yet our understanding of theseeffects is rudimentary. We identify three broad approaches that arenecessary to improve this understanding: (I) long-term research that utilizesan experimental and observational approach, (2) research that explores theinfluence of multiple urban spatial patterns, and (3) research that examinesurbanization effects on individuals, populations and communities acrossgradients of variable urbanization intensity. Many effects of urbanizationare obvious (loss and degradation of native habitat, introduction of exotics,changes in predator communities). However, it is important to rememberthe less obvious and often indirect effects of urbanization (climate change,

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human disturbance, ecosystem disruption, physiological stress, toxins, foodsupplementation) and determine how these affect birds. We end bysuggesting that effective conservation of birds in urban settings will requireinvolvement of all relevant stakeholders and interdisciplinary approaches.Our hope is that future urban avifaunas will retain a large and self-sustainingcomponent of that which existed before settlement. The thoughts presentedin this volume should help us attain that goal.

We could not have completed this book without the patience and supportof many. Each chapter was peer reviewed by two of our colleagues. Wethank David Aborn, Curtis Adkisson, Marina Alberti, Keith Bildstein, DavidBird, Robert Blair, Carl Bock, Jeff Bradley, Jeff Brawn, Dan Brooks, JoannaBurger, John Russell Butler, Carolee Caffrey, Dave Craig, Robert Curry,Richard DeGraaf, John Faaborg, Fred Gehlbach, Jon Greenlaw, JohnHadidian, Harlo Hadow, Lori Henning, Mark Hostetler, Richard Johnston,Jukka Jokimaki, Matthias Leu, Dave Manuwal, Don Mcivor, James Miller,Michael Molton, Ronald Mumme, Charles Nilon, Michael North, PeterPaton, Raymond Pierotti, Thane Pratt, Marco Restani, James Rodgers,Steven Rottenborn, Tom Scott, Dawn Sherry, Navjot Sodhi, Harrison B.Tordoff, Ian Warkington and Dave Willard for providing these reviews.John Marzluff and Roarke Donnelly were supported by The NationalScience Foundation (DEB 9875041), The Washington Department of Fishand Wildlife, and The University of Washington, particularly its RachaelWood's Graduate Program and the Tool's for Transformation Program.Reed Bowman was supported by Archbold Biological Station, the FloridaFish and Wildlife Conservation Commission, the United States Fish andWildlife Service, and the National Science Foundation (IDN-0077469).

John Marzluff, Seattle, WAReed Bowman, Lake Placid, FLRoarke Donnelly, Seattle, WA