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"Scale - the understanding of ecological phenomena through levels of biological organization across time and space - is one of most important concepts in ecology. It is often challenging for ecologists to find systems that lend themselves to study acrossscales; however, Sarracenia, a pitcher plant indigenous to the eastern United States, is unique because it can be studied at a hierarchy of scales: individuals, communities, and whole ecosystems. Ecologists Aaron Ellison and Nicolas Gotelli have studied Sarracenia for decades and, in this book, they synthesize their research and show how this system can inform the broad and challenging question of scaling in ecology. The authors' goal is to deepen the current understanding of major ecological processes, and how they operate across scales"--

A groundbreaking approach to scale and scaling in ecological theory and practice

Scale is one of the most important concepts in ecology, yet researchers often find it difficult to find ecological systems that lend themselves to its study. Scaling in Ecology with a Model System synthesizes nearly three decades of research on the ecology of Sarracenia purpurea—the northern pitcher plant—showing how this carnivorous plant and its associated food web of microbes and macrobes can inform the challenging question of scaling in ecology.

Drawing on a wealth of findings from their pioneering lab and field experiments, Aaron Ellison and Nicholas Gotelli reveal how the Sarracenia microecosystem has emerged as a model system for experimental ecology. Ellison and Gotelli examine Sarracenia at a hierarchy of spatial scales—individual pitchers within plants, plants within bogs, and bogs within landscapes—and demonstrate how pitcher plants can serve as replicate miniature ecosystems that can be studied in wetlands throughout the United States and Canada. They show how research on the Sarracenia microecosystem proceeds much more rapidly than studies of larger, more slowly changing ecosystems such as forests, grasslands, lakes, or streams, which are more difficult to replicate and experimentally manipulate.

Scaling in Ecology with a Model System offers new insights into ecophysiology and stoichiometry, demography, extinction risk and species distribution models, food webs and trophic dynamics, and tipping points and regime shifts.

Preface xiii
Abbreviations xxi
1 Introduction: Why Scale?
1(10)
1.1 Time and Space
2(1)
1.2 Genes to Ecosystems
3(2)
1.3 Modeling: Metabolic Theory and Macroecology
5(1)
1.4 Mechanisms at Scales
6(1)
1.5 Organisms as Model Systems
7(1)
1.6 Summary
8(3)
Part I Ecophysiology, Nutrient Limitation, and Stoichiometry 11(40)
2 Context: Nutrient Limitation, the Evolution of Botanical Carnivory, and Environmental Change
13(7)
2.1 Background
14(4)
2.1.1 Nutrient Acquisition, Plant Traits, and the Evolution of Botanical Carnivory
14(1)
2.1.2 Anthropogenic Activities Alter Resource Availability and Fluxes
14(4)
2.2 Next Steps
18(2)
3 The Small World: Stoichiometry and Nutrient Limitation in Pitcher Plants and Other Phytotelmata
20(11)
3.1 Stoichiometric Manipulations of Sarracenia
21(5)
3.1.1 Effects of Soluble N from Atmospheric Sources
21(2)
3.1.2 Effects of Nutrient Inputs from Supplemental Prey
23(3)
3.1.3 Synthesis of Supplemental Feeding Experiments
26(1)
3.2 Nutrient Additions in Other Phytotelmata
26(3)
3.3 Summary
29(2)
4 Scaling Up: Stoichiometry, Traits, and the Place of Sarracenia in Global Spectra of Plant Traits
31(20)
4.1 Global Plant Trait Spectra
31(2)
4.1.1 Traits
32(1)
4.1.2 Trait Data
32(1)
4.2 Carnivorous Plants in Global Trait Spectra
33(15)
4.2.1 Nutrient Concentrations
33(4)
4.2.2 Nutrient Stoichiometry
37(1)
4.2.3 Stoichiometric Effects of Supplemental Prey on Carnivorous Plants
37(5)
4.2.4 Stoichiometric Effects of Adding Inorganic Nutrients to Carnivorous Plants
42(5)
4.2.5 Photosynthesis and Construction Costs
47(1)
4.3 Synthesis
48(3)
Part II Demography, Global Change, and Species Distribution Models 51(46)
5 Context: Demography, Global Change, and the Changing Distributions of Species
53(6)
5.1 Background
54(1)
5.2 SDMs, Demography, and Anthropogenic Drivers: Moving Beyond Temperature
54(3)
5.2.1 Weak Responses to Temperature
55(1)
5.2.2 Nutrient Enrichment as Another Global-Change Driver
56(1)
5.2.3 The Importance of Demographic Effects
57(1)
5.3 Next Steps
57(2)
6 The Small World: Demography of a Long-Lived Perennial Carnivorous Plant
59(23)
6.1 Demographic Models of Sarracenia purpurea
59(8)
6.1.1 A Deterministic, Stage-Based Demographic Model for Sarracenia purpurea
59(4)
6.1.2 Stochastic Stage-Based Models
63(4)
6.2 Experimental Demography
67(3)
6.3 Demography in a Changing World
70(10)
6.3.1 Forecasting Nitrogen Deposition
70(1)
6.3.2 Linking N-Deposition Rates to Stage-Transition Matrices
71(3)
6.3.3 Modeling Population Growth
74(3)
6.3.4 The Future Is Now: Nitrogen Deposition and Extinction Risk in 2020
77(3)
6.4 Summary
80(2)
7 Scaling Up: Incorporating Demography and Extinction Risk into Species Distribution Models
82(15)
7.1 Available Data
82(1)
7.1.1 Sarracenia purupurea Occurrence Data
82(1)
7.1.2 Environmental and Climatic Data
83(1)
7.2 Continental Scaling of Demographic Models
83(8)
7.2.1 Challenges and Simplifying Assumptions
83(3)
7.2.2 Including P Introduced Additional Complexity
86(2)
7.2.3 Continental Forecasts for S. purpurea Persistence
88(3)
7.3 Forecasting the Future Distribution of Sarracenia purpurea
91(2)
7.3.1 A MaxEnt Model for Sarracenia purpurea
91(1)
7.3.2 Comparison of Forecasts of Demographic and MaxEnt Models
91(2)
7.4 Additional Forecasting Scenarios, Past and Future
93(2)
7.5 Synthesis
95(2)
Part II Ecology of the Sarracenia Community 97(48)
8 Context: Community Ecology, Community Ecologies, and Communities of Ecologists
99(5)
8.1 Background
100(3)
8.1.1 What Is an Ecological Community?
100(1)
8.1.2 Substituting Space for Time, and Vice Versa
100(3)
8.1.3 The Importance of Networks
103(1)
8.2 Next Steps
103(1)
9 The Small World: Structure and Dynamics of Inquiline Food Webs in Sarracenia purpurea
104(20)
9.1 Composition and Structure of the Sarracenia purpurea Food Web
104(3)
9.1.1 The Inquilines
104(1)
9.1.2 Network Structure of the Sarracenia purpurea Food Web
105(2)
9.2 Co-occurrence Analysis of Sarracenia purpurea Inquilines
107(7)
9.2.1 Quantifying and Testing Inquiline Co-occurrence
107(7)
9.3 Succession of the Inquiline Food Web
114(3)
9.4 Dynamics of the Sarracenia purpurea Food Web
117(6)
9.4.1 Temporal Changes in Food-Web Structure
117(1)
9.4.2 A Model of Food-Web Temporal Dynamics
118(5)
9.5 Summary
123(1)
10 Scaling Up: The Generality of the Sarracenia Food Web and Its Value as a Model Experimental System
124(21)
10.1 The Sarracenia Food Web and Other Container Webs Are "Normal" Food Webs
125(1)
10.1.1 Food-Web Data
125(1)
10.1.2 Food-Web Structure
126(1)
10.2 Spatial Scaling of the Sarracenia purpurea Food Web
126(6)
10.3 The Sarracenia purpurea Food Web as a Model Experimental System for Understanding and Managing Food Webs
132(11)
10.3.1 Fishing Down the Sarracenia Food Web
135(1)
10.3.2 Is Wyeomyia smithii a Keystone Predator?
136(1)
10.3.3 Dynamic Food Webs in Dynamic Habitats
137(6)
10.4 Synthesis
143(2)
Part IV Tempests in Teapots 145(38)
11 Context: Tipping Points and Regime Shifts
147(6)
11.1 Background
148(3)
11.1.1 Examples of Regime Shifts and Alternative States
149(1)
11.1.2 Linking Empirical Data with Mathematical Models of Alternative States
150(1)
11.2 A Potential Need for Interventions
151(1)
11.3 Next Steps
151(2)
12 The Small World: Tipping Points and Regime Shifts in the Sarracenia Microecosystem
153(9)
12.1 State Changes in the Sarracenia Microecosystem
153(8)
12.1.1 Temporal Dynamics of Aerobic and Anaerobic Conditions in Sarracenia purpurea Pitchers
154(3)
12.1.2 An Alternative Approach
157(4)
12.2 Summary
161(1)
13 Scaling Up: Using *omics to Identify Ecosystem States and Transitions
162(13)
13.1 Protein Surveys of the Sarracenia Microecosystem
162(1)
13.2 Proteomics of Sarracenia Fed Supplemental Prey
163(3)
13.3 The Cybernetics and Information Content of the S. purpurea Proteome
166(2)
13.4 Early Warning Indicators, Hysteresis, and the Twisted Path of Funded Research
168(5)
13.4.1 Hysteresis, Environmental Tracking, and Anti-hysteresis in the Sarracenia Microecosystem
170(3)
13.5 Synthesis
173(2)
14 Conclusion: Whither Sarracenia?
175(8)
14.1 Resources, Nutrients, and Stoichiometry
176(1)
14.2 Demography and Species Distributions
177(1)
14.3 Food Webs and Other Networks
178(2)
14.4 Tipping Points, Regime Shifts, and Alternative States
180(3)
Appendices 183(72)
Appendix A: The Natural History of Sarracenia and Its Microecosystem
185(27)
Appendix B: The Basics of Resource Limitation
212(3)
Appendix C: Deterministic Stage-Based Models
215(3)
Appendix D: The Basics of Species Distribution Models
218(3)
Appendix E: A Brief History and Precis of Methods for Analyzing Ecological Communities
221(17)
Appendix F: On Tipping Points and Regime Shifts
238(11)
Appendix G: On Biodiversity, Ecosystem Function, and *omics
249(6)
Notes 255(4)
References 259(44)
Subject Index 303(6)
Taxonomic Index 309
Aaron M. Ellison is the Senior Research Fellow Emeritus in Ecology at Harvard University. Website unbalancedecologist.net Twitter @AMaxEll17 Nicholas J. Gotelli is the George H. Perkins Professor of Zoology at the University of Vermont. Website uvm.edu/~ngotelli/homepage.html They are the coauthors of A Primer of Ecological Statistics and A Field Guide to the Ants of New England.