LEARNING LANDSCAPE ECOLOGY
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1 LEARNING LANDSCAPE ECOLOGY
2 Springer New York Berlin Heidelberg Hong Kong London Milan Paris Tokyo
3 LEARNING LANDSCAPE ECOLOGY A PRACTICAL GUIDE TO CONCEPTS AND TECHNIQUES SARAH E. GERGEL National Center for Ecological Analysis and Synthesis MONICA G. TURNER University of Wisconsin, Madison Editors Springer
4 Sarah E. Gergel Department of Forest Sciences University of British Columbia Vancouver, British Columbia Canada V6T 1Z4 Monica G. Turner Department of Zoology University of Wisconsin Madison, WI USA Library of Congress Cataloging-in-Publication Data Learning landscape ecology : a practical guide to concepts and techniques / edited by Sarah E. Gergel, Monica G. Turner. p. cm. Includes bibliographical references and index. ISBN (sc) 1. Landscape ecology. I. Gergel, Sarah E. II. Turner, Monica Goigel. QL L35 L dc Printed on acid-free paper. ArcExplorer and the GIS by ESRI emblem are trademarks provided under license from Environmental Systems Research Institute, Inc Springer-Verlag New York, Inc. All rights reserved. This work may not be translated or copied in whole or in part without the written permission of the publisher (Springer-Verlag New York, Inc., 175 Fifth Avenue, New York, NY 10010, USA), except for brief excerpts in connection with reviews or scholarly analysis. Use in connection with any form of information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed is forbidden. The use of general descriptive names, trade names, trademarks, etc., in this publication, even if the former are not especially identified, is not to be taken as a sign that such names, as understood by the Trade Marks and Merchandise Marks Act, may accordingly be used freely by anyone. Production coordinated by WordCrafters Editorial Services, Inc., Sterling, VA, and managed by Steven Pisano; manufacturing supervised by Jacqui Ashri. Typeset by Matrix Publishing Services, Inc., York, PA. Printed and bound by Edwards Brothers, Inc., Ann Arbor, MI. Printed in the United States of America (Corrected Printing, 2003) ISBN SPIN Springer-Verlag New York Berlin Heidelberg A member of BertelsmannSpringer Science Business Media GmbH
5 Preface Landscape ecology continues to grow as an exciting, dynamic ecological discipline. With its broadscale emphasis and multidisciplinary approach, landscape ecology lends itself both to basic research and to applications in land management, land-use planning, wildlife management, ecosystem management, and conservation biology. Landscape ecology makes a unique contribution to the scientific community in its attention to ecological dynamics across a broad range of spatial and temporal scales, and as a result it has become increasingly important for students in the natural sciences to gain a basic understanding of the subject. Colleges and universities across the United States are incorporating courses in landscape ecology into their curricula. However, nearly every book on landscape ecology is a book to be read, lacking a hands-on approach. This text is intended to fill that void by providing a comprehensive collection of landscape ecology laboratory exercises. These teaching exercises stress the fundamental concepts of landscape ecology, rather than highly specialized, technical methods. While students will gain experience using a variety of tools commonly used in landscape ecology, we stress the conceptual understanding necessary to use these techniques appropriately. This book attempts to convey the myriad approaches used by landscape ecologists (as well as a multitude of approaches to teaching) and include group discussion, thought problems, fieldwork, data analysis, spatial data collection, exposure to Geographic Information Systems (GIS), simulation modeling, analysis of landscape metrics, spatial statistics, and written exercises. v
6 vi PREFACE This book is divided into seven sections, which complement the companion textbook, Landscape Ecology in Theory and Practice: Pattern and Process, by M. G. Turner, R. H. Gardner, and R. V. O Neill. However, this textbook is also a useful stand-alone volume that can be used for teaching and learning. We also hope practicing landscape ecologists will find this book to be a useful reference. This book provides labs spanning a range of difficulty levels; thus, we have provided Suggestions for Instructors, which include difficulty ratings as well as several suggested lab sequences for undergraduate- and graduate-level courses. Many of the exercises also require computers. However, we intentionally designed the labs to be very user friendly in a PC environment. The enclosed CD-ROM includes files for many of the labs, including easily installable computer programs, simple DOS executable files, and data files accessible using commonly available word processing or spreadsheet programs. The CD also includes color images that are viewable using a web browser or Adobe Acrobat Reader software (available free on the web at Our intent is to make these teaching materials readily usable by colleges and universities without elaborate computing facilities. This volume focuses on computer-oriented labs more than field-oriented labs, even though field studies are a critical component of landscape ecology. There are several reasons for this emphasis. First, designing exercises that are transportable to any landscape is tricky because the patterns, relevant scales, and important biological processes differ among landscapes (however, two chapters involving fieldwork are included). Field-oriented labs, in general, may be best designed by the instructor to emphasize the important features and dynamics of a particular local area. Second, the quantitative techniques developed during the past decade or so in landscape are unfamiliar to many students, and thus this book fills an important void. Students have few opportunities to use simulation models and to apply the methods of spatial analyses. We hope this volume will help share the quantitative and modeling expertise of a few within an even broader scientific audience. However, the quantitative emphasis in this book is in no way intended to diminish the important role that fieldwork plays in landscape ecology. We strongly encourage students and instructors to embark on field studies in their local landscapes. Acknowledgments So many people were fundamental to this endeavor and deserve our praise and thanks. First, we d like to thank all the contributing authors for their creativity and enthusiasm in this endeavor, as well as their flexibility in allowing their chapters to be molded into part of the whole. Second, the following external reviewers deserve huge thanks for their critical assessment and enthusiasm: Tim Allen, Matthias Burgi, Jiquan Chen, Jonathan Chipman, Graeme Cumming, Don DeAngelis, Amy Downing, Mike DeMers, Curtis Flather, Marie-Josée Fourtin, Frank Golley, Steven Hamburg, Andy Hansen, Tom Hoctor, Lou Iverson, Tony Ives, Jeffrey Klopatek, David Lewis, Nancy
7 Preface vii Matthews, Kevin McGarigal, Nancy McIntyre, Todd Miller, Ron Moen, Kirk Moloney, Barry Noon, Bob O Neill, Volker Radeloff, Marguerite Remillard, Kurt Riitters, Tania Schoennagel, Fred Sklar, Pat Soranno (and her lab), Tom Spies (and his lab), Dean Urban, Steve Ventura, Karen Whitney, John Wiens, and the lab group of David Mladenoff. Rebecca Reed also deserves special thanks for her contributions during the beginning stages of the book. Equally important were the wide variety of students who tested these labs, helped find our mistakes, and offered suggestions for improvement. First and foremost, Sarah s graduate students in the fall 1998 landscape ecology lab course at University of Wisconsin, Madison, Jill Bukovac, Bruce Kahn, David Lewis, and Anna Pidgeon, tested 13 of the labs in one semester. Students in several years of Monica Turner and David Mladenoff s landscape ecology classes at University of Wisconsin, Madison, also deserve special thanks for their patience with the first drafts of many of these labs. We were continually impressed with their thorough evaluations and insightful, constructive suggestions. Instructors at other colleges and universities (and their students who are too numerous to mention here!) also deserve many thanks for testing labs, including Tara Reed s undergraduate biology class at Lawrence College and Joshua Greenberg s landscape ecology course at the University of Washington, Seattle, and Cheryl Schultz s graduate course in Landscape Ecology at The University of California, Santa Barbara. Of course, the many students enrolled in the courses taught by chapter authors deserve much thanks for their role in the development of these labs. Sharon Cowling, Lisa Dent, Dan Kashian, and Jack Williams, and all the post-docs at NCEAS who participated in the Eigenbeer challenge deserve special thanks for their attention to detail in the proofing stage of the book. Lastly, the artistic prowess of Michael Turner and Dirk Brandts in improving the figures in this book cannot be overstated. Finally, we would like to thank everyone in the Turner lab, especially Matthias Bürgi, Jeff Cardille, Mark Dixon, Dan Kashian, Tania Schoennagel, Mark Smith, and Dan Tinker, for many discussions, critiques, and help with endless aspects of this book. Sarah would also like to thank the recently retired Larry D. Harris for introducing her to the field of landscape ecology as an undergraduate. Sarah also thanks Jon Shurin for feeding her, enabling her to simultaneously finish this book and her Ph.D. We hope that this book will help aspiring landscape ecologists use the concepts and tools of landscape ecology to further our knowledge of how landscapes function and change, and more importantly, build and expand on the ideas presented here. Sarah E. Gergel Monica G. Turner Madison, Wisconsin
8 Suggestions for Instructors This book explores a variety of topics in landscape ecology, and as result, the difficulty levels of the labs also vary greatly. We hope this will provide a range of teaching tools appropriate for undergraduates, beginning graduate students, and advanced graduate students and researchers. We have assessed the difficulty of the labs using three categories: Undergraduate first introduction to landscape ecology, less emphasis on quantitative or computer-oriented labs, assumes a semester of ecology Graduate assumes at least one semester of statistics and a basic familiarity with a Windows PC and common word processing and spreadsheet software Advanced Graduate assumes experience in conducting research in landscape ecology, solid facility with PCs, and beginning familiarity with some aspects of modeling These categories assume a 3-hour class period. Thus, while a graduate lab might be used with undergraduates, extra time for completion should be allowed; when using an undergraduate lab with graduates, it will likely be completed in less than 3 hours. Many labs also include discussion questions and write-up sections to be completed out of class. Some authors have also created more detailed Instructor s Notes for several chapters. Please see our website for availability (as they are subject to revision as questions arise). We also strongly recommend that the instructor viii
9 Suggestions for Instructors ix gain familiarity with the labs before use in the classroom. These labs have been tested extensively; however, unforeseen computer glitches may arise when using the programs on a different computer, with a different operating system, and so forth. Check our website for corrections or to report problems. Software Requirements Chapters 1, 2, 6, 10, 11, 12, 13, 14, 17, 18 and part of 7 use only Excel (.xls) files, Adobe (.pdf) files, or no files at all, and as such are compatible with Macintosh or Windows platforms which have Excel and/or Adobe Acrobat Reader installed. Adobe Acrobat Reader is available for free from the web at: Several other programs require a PC running a recent version of Windows, or require a Windows emulator for use on a Macintosh platform: Markov, HarvestLite, Rule, Fragstats, ReserveDesign, Folio, ArcExplorer, and Bachmap. All programs are compatible with Windows XP except for Fragstats 2.0. Fragstats 2.0 and Windows XP The version of Fragstats included on the CD (FRAGSTATS 2.0) is not compatible with Windows XP; however, it does work with Windows 95/98/2000. Using Fragstats on Windows XP requires downloading the latest shareware version of Fragstats 3.0 from the web at: Be sure to visit our website before you begin teaching (or if you experience any difficulties while teaching) as it is frequently updated with corrections and helpful tips: Some Suggested Course Sequences Undergraduate Capstone Course in Landscape Ecology/Land Management/Conservation Biology Introduction to Geographic Information Systems (GIS) Simulating Changes in Landscape Pattern Interpreting Landscape Patterns from Organism-Based Perspectives Landscape Context Modeling Ecosystem Processes (Basic Version) Reserve Design Graduate Course in Landscape Ecology Scale and Hierarchy Theory Collecting Spatial Data at Broad Scales Creating Landscape Pattern Introduction to Markov Models
10 x SUGGESTIONS FOR INSTRUCTORS Understanding Landscape Metrics I Neutral Landscape Models Landscape Disturbance: Location, Pattern, and Dynamics Individual-Based Modeling: The Bachman s Sparrow Modeling Ecosystem Processes Feedbacks between Organisms and Ecosystem Processes Prioritizing Reserves for Acquisition Advanced Graduate Course or Spatial Modeling Introduction to Markov Models Simulating Changes in Landscape Pattern Understanding Landscape Metrics II: Effects of Changes in Scale Scale Detection Using Semivariograms and Autocorrelograms Alternative Stable States Landscape Connectivity and Metapopulation Dynamics Modeling Ecosystem Processes (Advanced Version)
11 Overview of Chapters Chapter Title Author Difficulty Level Related Chapters in Landscape Ecology in Theory and Practice: Pattern and Process, by M. G. Turner, R. H. Gardner, and R. V. O Neill 1 Scale and Hierarchy Theory O Neill Undergraduate; short discussion 2 lab 2 Collecting Spatial Data at Broad Scales Gergel et al. Graduate; requires significant 1, 5 out-of-class work 3 Introduction to Geographic Information Greenberg et al. Undergraduate Systems (GIS) 4 Introduction to Markov Models Urban and Wallin Graduate 3 5 Simulating Changes in Landscape Pattern Gustafson Undergraduate or Graduate 3, 4 6 Creating Landscape Pattern Delcourt Includes different versions for Undergraduate or Graduate; Understanding Landscape Metrics I is pre-requisite for Advanced version 4 7 Understanding Landscape Metrics I Cardille and Graduate 5 Turner 8 Understanding Landscape Metrics II: Effects of Greenberg et al. Graduate; essential prerequisites: 5 Changes in Scale Understanding Landscape Metrics I, Introduction to Geographic Information Systems (Continued)
12 Overview of Chapters (Continued) 9 Neutral Landscape Models Gardner and Advanced Graduate 6 Walters 10 Scale Detection Using Semi-variograms and Palmer Advanced Graduate 5 Auto-correlograms 11 Landscape Disturbance: Location, Pattern Turner et al. Graduate 7 and Dynamics 12 Multiple Stable States Peterson Advanced Graduate; strongly 7 recommended prerequisite: Introduction to Markov Models 13 Interpreting Landscape Patterns from Pearson Undergraduate 8 Organism-based Perspectives 14 Landscape Context Pearson Undergraduate 8 15 Landscape Connectivity and Metapopulation With Advanced Graduate; 8 Dynamics recommended prerequisite: Neutral Landscape Models; Section II is short discussion lab 16 Individual-Based Modeling: The Bachman's Dunning et al. Undergraduate or Graduate 8 Sparrow 17 Feedbacks Between Organisms and Ecosystem Wallace and Graduate 9 Processes Gray 18 Modeling Ecosystem Processes Gergel and Reed Includes different versions for Graduate or Undergraduate 9 19 Reserve Design Temple and Cary Undergraduate Prioritizing Reserves for Acquisition Urban Graduate 10
13 Contents PREFACE v SUGGESTIONS FOR INSTRUCTORS CONTRIBUTORS xvii viii SECTION 1 INTRODUCTION AND CONCEPTS OF SCALE CHAPTER 1 SCALE AND HIERARCHY THEORY 3 Robert V. O Neill and Mark A. Smith CHAPTER 2 COLLECTING SPATIAL DATA AT BROAD SCALES 9 Sarah E. Gergel, Monica G. Turner, and David J. Mladenoff CHAPTER 3 INTRODUCTION TO GEOGRAPHIC INFORMATION SYSTEMS (GIS) 17 Joshua D. Greenberg, Miles G. Logsdon, and Jerry F. Franklin xiii
14 xiv SECTION 2 MODELS AND CAUSES OF LANDSCAPE PATTERN CONTENTS CHAPTER 4 CHAPTER 5 INTRODUCTION TO MARKOV MODELS 35 Dean L. Urban and David O. Wallin SIMULATING CHANGES IN LANDSCAPE PATTERN 49 Eric J. Gustafson CHAPTER 6 CREATING LANDSCAPE PATTERN 62 Hazel R. Delcourt SECTION 3 QUANTIFYING LANDSCAPE PATTERN CHAPTER 7 CHAPTER 8 UNDERSTANDING LANDSCAPE METRICS I 85 Jeffrey A. Cardille and Monica G. Turner UNDERSTANDING LANDSCAPE METRICS II: EFFECTS OF CHANGES IN SCALE 101 Joshua D. Greenberg, Sarah E. Gergel, and Monica G. Turner CHAPTER 9 NEUTRAL LANDSCAPE MODELS 112 Robert H. Gardner and Steven Walters CHAPTER 10 SCALE DETECTION USING SEMIVARIOGRAMS AND AUTOCORRELOGRAMS 129 Michael W. Palmer SECTION 4 DISTURBANCE DYNAMICS CHAPTER 11 LANDSCAPE DISTURBANCE: LOCATION, PATTERN, AND DYNAMICS 147 Monica G. Turner, Daniel B. Tinker, Sarah E. Gergel, and F. Stuart Chapin III
15 Contents xv CHAPTER 12 ALTERNATIVE STABLE STATES 166 Garry D. Peterson SECTION 5 ORGANISM RESPONSE TO LANDSCAPE PATTERN CHAPTER 13 INTERPRETING LANDSCAPE PATTERNS FROM ORGANISM- BASED PERSPECTIVES 187 Scott M. Pearson CHAPTER 14 LANDSCAPE CONTEXT 199 Scott M. Pearson CHAPTER 15 CHAPTER 16 LANDSCAPE CONNECTIVITY AND METAPOPULATION DYNAMICS 208 Kimberly A. With INDIVIDUAL-BASED MODELING: THE BACHMAN S SPARROW 228 John B. Dunning Jr., David J. Stewart, and Jianguo Liu SECTION 6 ECOSYSTEM PROCESSES AT BROAD SCALES CHAPTER 17 FEEDBACKS BETWEEN ORGANISMS AND ECOSYSTEM PROCESSES 249 Linda L. Wallace and Steve T. Gray CHAPTER 18 MODELING ECOSYSTEM PROCESSES 266 Sarah E. Gergel and Tara Reed-Andersen SECTION 7 APPLIED LANDSCAPE ECOLOGY: INTEGRATING ACROSS THE LEVELS CHAPTER 19 RESERVE DESIGN 281 Stanley A. Temple and John R. Cary CHAPTER 20 PRIORITIZING RESERVES FOR ACQUISITION 293 Dean L. Urban INDEX 307
16 Contributors Jeffrey A. Cardille Department of Zoology University of Wisconsin, Madison Madison, WI John R. Cary Department of Wildlife Ecology University of Wisconsin, Madison Madison, WI F. S. (Terry) Chapin, III Institute of Arctic Biology University of Alaska, Fairbanks Fairbanks, AK Hazel R. Delcourt Department of Ecology and Evolutionary Biology University of Tennessee Knoxville, TN John B. Dunning, Jr. Department of Forestry and Natural Resources Purdue University West Lafayette, IN Jerry F. Franklin College of Forest Resources University of Washington Seattle, WA Robert H. Gardner Center for Environmental Science Appalachian Laboratory Frostburg, MD Sarah E. Gergel Department of Forest Sciences University of British Columbia Vancouver, British Columbia Canada V6T 1Z4 xvii
17 xviii CONTRIBUTORS Stephen T. Gray Department of Botany University of Wyoming Laramie, WY Joshua D. Greenberg Skagit County GIS/Mapping Mount Vernon, WA Eric J. Gustafson North Central Research Station USDA Forest Service Rhinelander, WI Jianguo (Jack) Liu Department of Fisheries and Wildlife Michigan State University East Lansing, MI Miles G. Logsdon School of Oceanography University of Washington Seattle, WA David J. Mladenoff Department of Forest Ecology and Management University of Wisconsin, Madison Madison, WI Robert V. O Neill, retired Environmental Sciences Division Oak Ridge National Laboratory Oak Ridge, TN eoneill@attglobal.net Michael W. Palmer Department of Botany Oklahoma State University Stillwater, OK carex@osuunx.ucc.okstate.edu Scott M. Pearson Biology Department Mars Hill College Mars Hill, NC spearson@mhc.edu Garry D. Peterson Department of Geography & McGill School of the Environment McGill University 805 Sherbrooke St. W. Montreal, Quebec Canada H3A 2K6 Tara Reed-Andersen Department of Natural and Applied Sciences University of Wisconsin, Green Bay Green Bay, WI reedandt@uwgb.edu Mark A. Smith Department of Wildlife Ecology and Zoology University of Wisconsin, Madison Madison, WI masmith7@students.wisc.edu David J. Stewart Department of Anthropology University of Georgia Athens, GA dstewart@julian.dac.uga.edu Stanley A. Temple Department of Wildlife Ecology University of Wisconsin, Madison Madison, WI satemple@facstaff.wisc.edu Daniel B. Tinker Department of Botany University of Wyoming Laramie, WY Tinker@uwyo.edu
18 Contributors Monica G. Turner Department of Zoology University of Wisconsin, Madison Madison, WI Dean L. Urban Nicholas School of the Environment Duke University Durham, NC Linda L. Wallace Department of Botany and Microbiology University of Oklahoma Norman, OK David O. Wallin Center for Environmental Science Huxley College of Environmental Studies Western Washington University Bellingham, WA Steven Walters U.S. Environmental Protection Agency Narragansett, RI Kimberly A. With Division of Biology Kansas State University Manhattan, KS xix
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