Table Of ContentOUP CORRECTED PROOF – FINAL, 08/06/19, SPi
Essentials of Landscape Ecology
OUP CORRECTED PROOF – FINAL, 08/06/19, SPi
OUP CORRECTED PROOF – FINAL, 08/06/19, SPi
Essentials of
Landscape Ecology
Kimberly A. With
1
OUP CORRECTED PROOF – FINAL, 08/06/19, SPi
1
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OUP CORRECTED PROOF – FINAL, 08/06/19, SPi
Preface
Writing a textbook is rather like being enrolled in a of the material. Some students require an introduction
multi-year course on the subject, in which the roles of to basic ecological concepts and principles, whereas
student and teacher are essentially interchangeable. others are looking for synthesis, critical review, and a
Since we are all really students at heart, my approach deeper treatment of the analytical, methodological, or
to writing this textbook has been from the standpoint modeling approaches they are hoping to implement in
of a student, starting with myself. I know what difficul- their own research.
ties I have faced in understanding certain topics and The principal challenge in writing this textbook, as
analytical approaches, and after teaching landscape in the classroom, has thus been to develop a curricu-
ecology for more than 20 years, I know what difficul- lum that serves the needs of a diverse audience. To that
ties many of my students encounter when first exposed end, I have tried to provide a mix of basic concepts,
to particular concepts or methods. This textbook is examples, and case studies, as well as a more advanced
thus for the student of landscape ecology, whatever treatment of certain topics. The examples and case
your academic standing, professional rank, or title. studies were selected to emphasize a range of organ-
Welcome to our shared course on landscape ecology. isms, systems, and geographic locations to the extent
The scientific domain encompassed by landscape possible, although this was often influenced by the
ecology is incredibly vast and wide-ranging. Landscape idiosyncratic nature of scientific research as well as by
ecology is built upon contributions from various fields my idiosyncratic research into that literature. I therefore
in the natural and social sciences, and in turn contrib- offer advance apologies if I appear to have overlooked
utes to both basic and applied problems in each of an important paper or study, perhaps even one of your
its constituent fields. No single textbook—including own contributions to the field. I fear this is inevitable,
this one—can possibly do it justice. My focus in this given the vastness of the literature, the sheer size of the
textbook is thus on the ecology of landscape ecology. task at hand, and the limited space available to address
I am an ecologist by training, and besides landscape each topic.
ecology, much of my research has been in the areas In terms of the book itself, I thank Sharon Collinge,
of behavioral, population, and community ecology. Henri Décamps, Kevin McGarigal, Jean Paul Metzger,
In addition, I have long been interested in conservation John Wiens, and Jingle Wu for their suggestions, sup-
and the applications of landscape ecology for conser- port, and enthusiasm for the initial book proposal. I am
vation biology. Thus, my perspective herein is inevitably also indebted to the following colleagues who took the
that of an ecologist, with an eye toward applications time to review and provide comment on various sec-
of landscape ecology for conservation and natural tions or chapters of the book (and sometimes, on more
resource management. than one): John Briggs, Thomas Crist, Lenore Fahrig,
Despite this—or maybe even because of this—it is Olivier François, Janet Franklin, Marie-Josée Fortin,
my hope that this book will be of interest to students Doug Goodin, Eric Gustafson, Colleen Hatfield, Nancy
and professionals in other areas of landscape ecology, McIntyre, Jean Paul Metzger, Emily Minor, Rick Ostfeld,
such as geographers and landscape planners, who Luciana Signorelli, Mark Ungerer, Helene Wagner,
might desire an ecological overview to complement Lisette Waits, Jingle Wu, and Patrick Zollner. I would
their training or design perspective. Judging from the also like to thank the graduate students who have par-
composition of my own landscape ecology course over ticipated in my landscape ecology course at Kansas
the years, I envision an audience that is likely to be quite State University, and who provided input on many
diverse regardless. In the past, this course has attracted of the initial chapter drafts and offered suggestions
students from all over campus and from fields as for improvement, including other examples or topics
diverse as ecology, environmental science, entomology, I had overlooked. Special thanks in this regard are
geography, agronomy, range sciences, forestry, conser- due to Rachel Pigg, Jay Guarani, Mark Herse, Sean
vation biology, fisheries, and wildlife management. Hitchman, Nate Cathcart, Reid Plumb, Emily Williams,
The academic background, training, and expectations and E. J. Raynor.
of these students are all very different, which demands Of course, this book would never have seen the light
a varied and diversified approach in the presentation of day without a publisher. I very much appreciate
OUP CORRECTED PROOF – FINAL, 08/06/19, SPi
vi Preface
the care and oversight provided by Ian Sherman and McIntyre, for uncovering many of the images con-
Bethany Kershaw, senior and assistant commissioning tained within these chapters. The end result is a text-
editors, respectively, for biology at Oxford University book that is as attractive as I hope it is informative.
Press, and Ioan Marc Jones, senior production editor Last, but certainly not least, I thank my husband,
for academic texts at OUP, for managing the book’s Gray Woods, for his unremitting support and encour-
transition through to its final production phase. Paul agement throughout the many years it has taken me to
Beverley had the unenviable task of copy-editing the complete this project. I dedicate this book to our son,
entire text, but the book is better for his efforts. This Johnathan, who I fear no longer remembers a time
project started out under the auspices of Sinauer when his mother wasn’t working on this book, becom-
Associates (now an imprint of OUP), and I therefore ing a sort of bibliographic sibling. Wise beyond his
owe a large debt of gratitude to Andy Sinauer for the years, he had asked me at the start, “How do you know
opportunity and for his support (and patience) during what to write?” I don’t recall my answer then, but I
this long process; production editors, Kathaleen have one for him now: figuring out what to write is
Emerson and Stephanie Bonner, as well as art director easy when one has access to such great material; it’s
and production manager, Chris Small, for their guid- figuring out when to stop that is hard. For, as I now
ance, design aesthetic, and attention to detail; Jan realize, the book will never truly be finished, but hope-
Troutt, for her beautiful artwork; Johannah Walkowicz fully, this is a good and worthwhile start.
and Michele Bekta for their efforts behind the scenes in
obtaining permissions; copy e ditor Carol Wigg for Kimberly A. With
comments and suggestions on the first set of chapters; Division of Biology
and photo researchers, Mark Siddall and David Kansas State University
OUP CORRECTED PROOF – FINAL, 08/06/19, SPi
Contents
Chapter 1 An Introduction to Landscape Ecology: Foundations
and Core Concepts 1
Why Study Landscape Ecology? 2
Birth of a Discipline 4
Regional Perspectives on Landscape Ecology 5
European Perspective 5
North American Perspective 7
Globalization of Landscape Ecology 8
Core Concepts of Landscape Ecology 9
Organization of this Book 11
Chapter Summary Points 12
Discussion Questions 13
Chapter 2 Scaling Issues in Landscape Ecology 14
Why is Scale so Important in Ecology? 14
Uses (and Misuses) of Scale in Ecology 16
Ecological Scale 16
Relationship between Grain and Extent 16
Effect of Changing Grain and Extent 18
Choosing the ‘Right’ Scale of Study 21
Hierarchy Theory and Landscape Ecology 24
Hierarchical Organization of Biological Systems 27
Structure of a Hierarchical System 28
Landscape Scale or Landscape Level? 29
Implications of Hierarchy Theory for Landscape Ecology 32
Extrapolating Across Scales 33
Extrapolating Within Domains of Scale 34
Extrapolating Across Domains of Scale 34
Uncertainty, Predictability, and Ecological Forecasting 36
Future Directions 39
Chapter Summary Points 40
Discussion Questions 41
Chapter 3 Landscape Heterogeneity and Dynamics 42
Heterogeneity and Disturbance Dynamics as Core Concepts in Landscape Ecology 42
Emergence of Heterogeneity and Dynamical Concepts in Ecology 44
Heterogeneity at Broad Geographic Scales: Biogeography and Life Zones 44
Dynamics of Plant Community Assembly: Climax State or Independent Assembly? 45
Patch Dynamics: A Paradigm Shift in Ecology 47
Toward a Non-Equilibrium View of Ecology 48
Homogeneity: The Frictionless Plane of Ecological Theory 49
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viii Contents
Patch-Based Theory in Ecology 50
Toward a Landscape-Mosaic View of Environmental Heterogeneity 52
How is Heterogeneity Defined? 54
Spatial Heterogeneity 54
Temporal Heterogeneity 55
How are Disturbances Defined? 56
The Disturbance Regime 56
The Intermediate Disturbance Hypothesis 58
Large Infrequent Disturbances 59
Implications of the Disturbance Regime for Landscape Dynamics 61
Formation and Evolution of Landscapes: Geomorphological Processes 63
Formation of Montane Landscapes 63
Formation of River Networks 68
Formation and Diversity of Soils 69
Landscape Dynamics: Abiotic Disturbances 73
Volcanic Eruptions 73
Landsliding 76
Flooding and the Natural Flow Regime 78
Windstorms, Hurricanes, and Storm Surges 85
Drought 89
Fire 94
Landscape Dynamics: Biotic Agents of Landscape Formation and Disturbance 101
Organisms as Geomorphic Agents 102
Organisms as Ecosystem Engineers 102
The Keystone Role of Species 105
Relative Impact of Species on Landscapes 107
Humans as the Primary Driver of Landscape Change 107
Stages of Anthropogenic Landscape Transformation 108
Types of Human Land Use 110
Land-Use Legacy Effects 120
Future Directions 122
Chapter Summary Points 123
Discussion Questions 125
Chapter 4 Landscape Pattern Analysis 127
On the Importance of Landscape Pattern Analysis in Landscape Ecology 127
Sources of Landscape Data 128
Historical Land Surveys 129
Remote Sensing 131
From Landscape Data to Landscape Data Analysis 153
Geographical Information Systems 156
Data Input Subsystem 157
Data Processing and Database Management Subsystem 158
Data Manipulation and Analysis Subsystem 159
Data Output Subsystem 165
Landscape Metrics 165
Measures of Landscape Composition 167
Measures of Landscape Configuration 168
Effects of Pattern and Scale on Landscape Metrics 178
Use and Misuse of Landscape Metrics 184
Spatial Analysis 186
A Primer to Spatial Statistics 186
Spatial Statistics 189
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Contents ix
Future Directions (and Some Caveats) 202
Chapter Summary Points 203
Discussion Questions 205
Chapter 5 Landscape Connectivity 206
What is Landscape Connectivity and Why is it Important? 206
Elements of Landscape Connectivity 208
Structural versus Functional Connectivity 209
Patch Connectivity versus Landscape Connectivity 210
Patch-Based Connectivity Measures 210
Which Patch-Connectivity Measure to Use and When? 212
From Patches to Landscapes 213
Methods for Assessing Landscape Connectivity 214
Neutral Landscape Models 214
Graph-Theoretic Approaches 221
Assessing Connectivity in Heterogeneous Landscapes 227
Assessing Connectivity in River Networks 232
Which Landscape Connectivity Approach to Use and When? 235
Beyond Landscape Connectivity 236
Should Landscape Connectivity be a Dependent or Independent
Variable? 236
Future Directions 237
Chapter Summary Points 237
Discussion Questions 238
Chapter 6 Landscape Effects on Individual Movement and Dispersal:
Behavioral Landscape Ecology 239
Why are Movement and Dispersal Important from a Landscape Ecological
Perspective? 240
Scales of Movement 241
Movement Responses to Hierarchical Patch Structure 242
Allometric Scaling of Movement 245
Movement Responses to Patch Structure 245
Movement out of Patches (Emigration) 246
Movement Between Patches 256
Movement into Patches (Immigration) 260
Analysis of Movement Pathways 264
Tracking Animal Movements 265
Scaling Issues in Tracking Animal Movements 268
Measuring Movement Pathways 269
Models of Animal Movement 272
Mathematical Models of Animal Movement 272
Spatially Explicit Models of Animal Movement 277
Space Use and Home-Range Analysis 280
Methods of Home-Range Estimation 280
Measuring Plant Dispersal 284
Future Directions 286
Chapter Summary Points 287
Discussion Questions 289
