Complexity in Landscape Ecology

262 Pages·2020·9.205 MB·English

by  David G. Green, Nicholas I. Klomp, Glyn Rimmington, Suzanne Sadedin

Landscape Series
David G. Green
Nicholas I. Klomp
Glyn Rimmington
Suzanne Sadedin
Complexity
in Landscape
Ecology
Second Edition
Landscape Series
Volume 22
Series editors
Jiquan Chen, Department of Geography, Environment, and Spatial Sciences,
Michigan State University, East Lansing, USA
Janet Silbernagel, Department of Planning and Landscape Architecture, 
University of Wisconsin-Madison, Madison, USA
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David G. Green • Nicholas I. Klomp
Glyn Rimmington • Suzanne Sadedin
Complexity in Landscape
Ecology
Second Edition
David G. Green Nicholas I. Klomp
Faculty of Information Technology Central Queensland University
Monash University Rockhampton, QLD, Australia
Clayton, VIC, Australia
Suzanne Sadedin
Glyn Rimmington Fremont, CA, USA
Wichita State University
Wichita, KS, USA
ISSN 1572-7742 ISSN 1875-1210 (electronic)
Landscape Series
ISBN 978-3-030-46772-2 ISBN 978-3-030-46773-9 (eBook)
https://doi.org/10.1007/978-3-030-46773-9
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Preface to the Second Edition
We published the first edition of this book in 2006. However, the project really goes
back to the early 1990s, when we recognised the need for a book to introduce the
principles of complexity to ecologists. The changes since then have been enormous.
For example, when we published the first edition, complexity-based models in ecol-
ogy were rare; now they are mainstream. Likewise, monitoring, data analytics and
other new technologies are changing the way we do ecology. The increase in rele-
vant publications has been phenomenal.
There are some structural changes since the first edition. To make the subject
matter clearer, each chapter now begins with a title page, which includes a subtitle,
along with key words and a brief abstract of the main topics covered. We have reor-
ganised each chapter to highlight important and emerging concepts. Perhaps the
most obvious changes are the figures, which are essential for conveying concepts in
complexity. We have added 30 figures, updated others, and rendered all of them in
colour. We also plan to provide simulation models online of selected figures, so
readers can experiment with complexity themselves.
We have expanded every topic and added new ones not covered in the first edi-
tion. We have also added many recent references and contemporary examples.
Despite these additions, the second edition is still only an introduction to the field.
We do not claim to provide a comprehensive review of recent research. In the inter-
ests of keeping the book to a reasonable length, each topic and chapter is introduc-
tory in nature. For example, researchers have published work on literally hundreds
of metrics to quantify patterns of complexity in landscapes. To cover them all would
take an entire book in itself. Many of those metrics were developed for specialised
purposes that are beyond the scope of this book. We can give only a few examples
of metrics that we think best reflect the idea of complexity. The same limitation
applies to almost every topic we cover.
This second edition aims to be a useful introduction to complexity in ecology.
With this in mind, we have tried to make the book accessible to a wide, general audi-
ence. Most notably, we have tried to keep mathematics and other technical matters
in the background. This is challenging because research in the field has become
increasingly sophisticated and technical. Another challenge we faced was to update
v
vi Preface to the Second Edition
the case studies. The story of deforestation in the Amazon, for instance, is an on-
going tragedy that is continually documented. Similarly, alarming reports keep
appearing about the impact of human activity on biodiversity.
We are grateful to many people who contributed to the second edition. Dr
François Petitjean provided satellite images. Benjamin O’Leary tracked down many
references for the bibliography. Associate Professor Alan Dorin provided informa-
tion about foraging bees. Associate Professor David Dowe provided useful sugges-
tions about complexity measures. Dr Marc Cheong proofread the draft manuscript
and provided many useful suggestions. Ira (Maria) Djaja wrote programs for several
figures and the demonstrations that we will provide online. Dr Laura Trouille pro-
vided helpful information about the Zooniverse project.
Clayton, VIC, Australia David G. Green
Rockhampton, QLD, Australia Nicholas I. Klomp
Wichita, KS, USA Glyn Rimmington
Fremont, CA, USA Suzanne Sadedin
Contents
1 Complexity and Ecology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
1.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
1.2 What Is Complexity? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
1.2.1 Variety and Form . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
1.2.2 The Chicken and the Egg . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
1.3 What Makes Ecosystems Complex? . . . . . . . . . . . . . . . . . . . . . . . . 7
1.3.1 Measuring Diversity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
1.3.2 The Origins of Complexity . . . . . . . . . . . . . . . . . . . . . . . . . 9
1.4 Why Study Ecological Complexity? . . . . . . . . . . . . . . . . . . . . . . . . 12
1.5 The Complexity Paradigm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
1.5.1 Scientific Paradigms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
1.5.2 A New Ecology for a New Age? . . . . . . . . . . . . . . . . . . . . . 15
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
2 Seeing the Wood for the Trees: Emergent Order in Growth and
Behaviour . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
2.1 Plant Growth and Form . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
2.1.1 Factors Influencing Growth . . . . . . . . . . . . . . . . . . . . . . . . . 22
2.1.2 Branches and Leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
2.1.3 Overall Plant Form . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
2.1.4 Self-Organisation Versus Constrained Growth . . . . . . . . . . 33
2.2 Animal Behaviour . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
2.2.1 Searching for Food . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
2.2.2 Territory . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
2.2.3 Social Networks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
2.2.4 Animal Intelligence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
2.3 Multiagent Systems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
2.3.1 Turtle Geometry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
2.3.2 From Turtles to Agents. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
2.3.3 The Boids and the Bees . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
vii
viii Contents
3 Complexity in Landscapes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
3.1 The Eye of the Beholder . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
3.1.1 Geographic Information Systems . . . . . . . . . . . . . . . . . . . . 48
3.1.2 The Game of Life . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
3.1.3 Cellular Automata Models of Landscapes . . . . . . . . . . . . . . 52
3.2 Sampling and Scale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54
3.3 Complexity in Spatial Processes . . . . . . . . . . . . . . . . . . . . . . . . . . . 56
3.4 Complexity in Spatial Patterns . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58
3.4.1 Fractal Dimensions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
3.4.2 Fractals in Nature . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60
3.4.3 Measuring Landscape Complexity . . . . . . . . . . . . . . . . . . . 61
3.5 Are Landscapes Connected? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64
3.5.1 Connectivity in a Grid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64
3.5.2 Why Is a Starfish Like an Atomic Bomb? . . . . . . . . . . . . . . 66
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
4 Oh, What a Tangled Web … Complex Networks in Ecology . . . . . . . 73
4.1 The Roots of Complexity Theory . . . . . . . . . . . . . . . . . . . . . . . . . . 74
4.2 The Network Model . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76
4.2.1 Interactions and Connectivity . . . . . . . . . . . . . . . . . . . . . . . 76
4.2.2 Networks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76
4.2.3 Networks Are Everywhere . . . . . . . . . . . . . . . . . . . . . . . . . . 78
4.2.4 The Connectivity Avalanche . . . . . . . . . . . . . . . . . . . . . . . . 79
4.2.5 Phase Changes and Criticality . . . . . . . . . . . . . . . . . . . . . . . 80
4.2.6 The Order of Things . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82
4.3 Self-Organisation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84
4.3.1 Emergent Properties . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84
4.3.2 Modules and Motifs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86
4.3.3 The Shape of Complexity . . . . . . . . . . . . . . . . . . . . . . . . . . 88
4.4 Networks of Networks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95
5 The Imbalance of Nature … Feedback and Stability
in Ecosystems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97
5.1 Feedback . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99
5.1.1 Negative Feedback Promotes Stability . . . . . . . . . . . . . . . . 99
5.1.2 Positive Feedback Promotes Self-Organization . . . . . . . . . . 100
5.2 The Big Get Bigger . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
5.3 Who Eats Whom? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 104
5.3.1 Equilibrium and Stability . . . . . . . . . . . . . . . . . . . . . . . . . . . 105
5.3.2 Transients and Attractors . . . . . . . . . . . . . . . . . . . . . . . . . . . 107
5.3.3 Sensitivity to Initial Conditions . . . . . . . . . . . . . . . . . . . . . . 108
5.3.4 The Onset of Chaos . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109
5.3.5 Fractals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 112
5.4 Is There a Balance of Nature? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 112
5.4.1 Succession . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113
Contents ix
5.4.2 Ecosystems in Balance? . . . . . . . . . . . . . . . . . . . . . . . . . . . . 114
5.4.3 Does a Balance Really Exist? . . . . . . . . . . . . . . . . . . . . . . . 115
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116
6 Populations in Landscapes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 119
6.1 One Population or Many? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 120
6.2 Spatial Distributions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 122
6.3 Patches, Edges and Zones . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 124
6.3.1 Salt of the Earth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 125
6.4 To See the World in a Grain of Pollen . . . . . . . . . . . . . . . . . . . . . . . 126
6.5 Galloping Trees? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 129
6.6 Phylogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 131
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 133
7 Living with the Neighbours: Competition and Stability in
Communities . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137
7.1 Invasions and Persistence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 138
7.2 Disturbance and Competition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 140
7.3 Ecological Communities . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 142
7.3.1 Do Ecological Communities Exist? . . . . . . . . . . . . . . . . . . . 144
7.4 Networks of Interactions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 145
7.4.1 Food Webs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 145
7.4.2 Networks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 147
7.4.3 The Paradox of Stability . . . . . . . . . . . . . . . . . . . . . . . . . . . 148
7.4.4 Stability and Food Webs . . . . . . . . . . . . . . . . . . . . . . . . . . . 149
7.4.5 Resilience . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 152
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 153
8 Adaptation in Landscapes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 157
8.1 Genes and Selection . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 158
8.1.1 Evolutionary Trade-Off . . . . . . . . . . . . . . . . . . . . . . . . . . . . 160
8.2 Genetics in Heterogeneous Landscapes . . . . . . . . . . . . . . . . . . . . . . 161
8.2.1 Adaptation on a Gradient . . . . . . . . . . . . . . . . . . . . . . . . . . . 163
8.2.2 Fragmentation and Drift . . . . . . . . . . . . . . . . . . . . . . . . . . . . 164
8.2.3 Friends and Relations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 167
8.3 Catastrophes, Criticality and Macroevolution . . . . . . . . . . . . . . . . . 169
8.3.1 Mass Extinctions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 169
8.3.2 Landscape Phases and the Origin of Species . . . . . . . . . . . . 171
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 174
9 Virtual Worlds: The Role of Simulation in Ecology . . . . . . . . . . . . . . 177
9.1 Virtual Experiments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 179
9.1.1 From Landscapes to Virtual Worlds . . . . . . . . . . . . . . . . . . . 180
9.1.2 The Need for Simulation . . . . . . . . . . . . . . . . . . . . . . . . . . . 181
9.1.3 A World Inside a Computer? . . . . . . . . . . . . . . . . . . . . . . . . 182
9.1.4 The Limits to Growth Model . . . . . . . . . . . . . . . . . . . . . . . . 183
9.1.5 Just So Stories? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 184