1 Samuel M. Scheiner and Michael R. Willig, eds. 1. A General ...

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LIST OF CONTRIBUTORS Brandon T. Bestelmeyer USDA Agricultural Research Service Jornada Experimental Range Ingrid C. Burke Department of Botany University of Wyoming Mary L. Cadenasso Department of Plant Sciences University of California–Davis Jonathan M. Chase Department of Biology Washington University Robert Colwell Department of Ecology and Evolutionary Biology University of Connecticut Gordon A. Fox Department of Biology University of South Florida Steven D. Gaines Department of Ecology, Evolution and Marine Biology University of California–Santa Barbara Alan Hastings Department of Environmental Science and Policy University of California–Davis Robert Holt Department of Biology University of Florida Alan K. Knapp Department of Biology Colorado State University William K. Lauenroth Department of Botany University of Wyoming Mathew A. Leibold Section of Integrative Biology University of Texas–Austin Scott J. Meiners Department of Biological Sciences Eastern Illinois University Jay Odenbaugh Department of Philosophy Lewis & Clark College Debra P. Peters USDA Agricultural Research Service Jornada Experimental Range Stewart Pickett Institute for Ecosystem Studies Dov Sax Department of Ecology and Evolutionary Biology Brown University Samuel M. Scheiner Arlington, VA Andrew Sih Department of Environmental Science and Policy University of California–Davis Michael R. Willig Center for Environmental Sciences and Engineering University of Connecticut 2
Chapter 1: A General Theory of Ecology Samuel M. Scheiner and Michael R. Willig “In the absence of agreed protocols and overarching theory, Ecology with its numerous subdisciplines, can sometimes resemble an amorphous, postmodern hotel or rabbit warren with separate entrances, corridors and rooms that safely accommodate the irreconcilable.” (Grime 2007) The development of theory in ecology is a lively and robust enterprise (Pickett et al. 2007). Despite claims to the contrary, the science of ecology has a long history of building theories that fruitfully guide research and deepen understanding. Our goal with this book is to reveal a selection of those theoretical structures. In doing so, our hope is that ecologists will better appreciate the theoretical frameworks within which they do research and will engage those theories in designing observational, experimental, and modeling components of their research. Many theories in ecology contain unspoken or even subconscious assumptions. By bringing such assumptions to the forefront, we can understand their consequences and discover new mechanisms or patterns, as well as new linkages among theories. Theory sometimes seems to be distant or disconnected from everyday practice in ecology. By the end of this book, the relevance of theory to understanding in ecology and its role in advancing science should become clear. In this chapter, we present a general theory of ecology that serves as the supporting framework – a conceptual infrastructure – for the constitutive theories that appear in subsequent chapters. Chapters 2 (Kolasa) and 3 (Odenbaugh) consider the role that theory has played in ecology from the perspectives of a practicing ecologist and of a philosopher of science. The eleven chapters that make up the heart of the book delve into the theoretical underpinnings of a broad range of ecological subdisciplines. Although those chapters span the disciplinary range of ecology, they are representative rather than comprehensive. We could not possibly synthesize the full richness of ecological theory in a single book without it becoming encyclopedic. We encourage others to continue the process of theory development in other venues, and to re- engage theoretical discourse with ecological research (e.g., Pickett et al. 2007). We do not claim novelty for the general theory of ecology put forward here. Quite the contrary, the elements of the general theory have existed for at least the past 50 years. Many of 3 3
Page 1: TABLE OF CONTENTS THE THEORY OF ECO
Page 5 and 6: Chapter 10). Without such boundarie
Page 7 and 8: profound. As the statistician Georg
Page 9 and 10: In general, the domain of a theory
Page 11 and 12: Physiological ecology is likely ano
Page 13 and 14: of a theory. A theory is constantly
Page 15 and 16: component of the process of natural
Page 17 and 18: Literature cited Arnold, W., T. Ruf
Page 19 and 20: Scheiner, S. M. submitted. Towards
Page 21 and 22: Table 1.2. Definitions of terms for
Page 23 and 24: Chapter 2: Theory Makes Ecology Evo
Page 25 and 26: captured the multifaceted complexit
Page 27 and 28: proposition is less formal, some mi
Page 29 and 30: diversity and the size of the area
Page 31 and 32: a systematic analysis of the struct
Page 33 and 34: circumstance related to theory that
Page 35 and 36: vegetation types, a simple combinat
Page 37 and 38: author) was impressive. Although fe
Page 39 and 40: separate theoretical problems that
Page 41 and 42: indirect effects and enemy-mediated
Page 43 and 44: effects of diversity on the stabili
Page 45 and 46: to any major extent. Although ecosy
Page 47 and 48: • A larger, integrating theory is
Page 49 and 50: Forbes, S. 1887. The lake as a micr
Page 51 and 52: MacArthur, R.H. and E.O. Wilson. 19
Page 53 and 54:
Strong, D. R. 1984. Exorcising the
Page 55 and 56:
Table 2.1. Major theoretical develo
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1969 Individual thermodynamic budge
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c e n e a ra p a p f r o e rd O 20
Page 61 and 62:
Figure 2.4. Number of papers found
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Chapter 3: A General, Unifying Theo
Page 65 and 66:
the number of predators and the gro
Page 67 and 68:
Robert MacArthur, E. O. Wilson, Egb
Page 69 and 70:
Suppose we have a phase space where
Page 71 and 72:
group. The state variables and para
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superset of the former’s domain;
Page 75 and 76:
Literature Cited Abrams, P. (1983)
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Figure 3.1. Density-dependent selec
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foraging for resources. Although pr
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foraging on the same prey, and on p
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environment). In a later section, I
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published a seminal book that summa
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always be taken when encountered. S
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analyses of optimal sampling (DeGro
Page 91 and 92:
previous models, dynamic state-depe
Page 93 and 94:
functions that are part of these mo
Page 95 and 96:
Foraging behavior often balances th
Page 97 and 98:
esources. In recent years, renewed
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foragers often show adaptive respon
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y species of conservation concern.
Page 103 and 104:
Charnov, E.L., G.H. Orians and K. H
Page 105 and 106:
Lima, S.L. 1998. Stress and decisio
Page 107 and 108:
Salo, P., E. Korpimaki, P.B. Banks,
Page 109 and 110:
Werner, E.E. and J.F. Gilliam 1984.
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12. Foraging behavior often balance
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113 Figure 4.2. A graphical present
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Chapter 5: Ecological Niche Theory
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ecologists, and evolutionary biolog
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simply intended to serve as a backd
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esource is in mass balance (sensu H
Page 123 and 124:
Proposition 2 For more than one spe
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non-equilibrial (cycling) dynamic i
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also likely as a result of disturba
Page 129 and 130:
Literature Cited Abrams, P. A. and
Page 131 and 132:
Gravel, D., C. D. Canham, M. Beaude
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Shurin, J. B., P. Amarasekare, J. M
Page 135 and 136:
Table 5.1. The background and propo
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Chapter 6: Single Species Populatio
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DOMAIN OF THE THEORIES The domain o
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As another example, one can focus o
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where adults lay eggs, which are su
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We note that both of these solution
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density dependence has long transie
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ecognizing these assumptions is an
Page 151 and 152:
Literature Cited Caswell, H. 2001,
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Chapter 7: Natural enemy-Victim Int
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among species in the kinds of resou
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2. Satiation in predators, and time
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eproduction. Because natural enemy-
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assume the natural enemy is a speci
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A series of fundamental constraints
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General (but not universal) dynamic
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in demography and physical biology.
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∂f N ∂fP ∂f N ∂f P > , (9)
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interactions. In addition to its im
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many examples and so is a reasonabl
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ecause goose defense strategies bec
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So a ratio-dependent model such as
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aggressive interference) due to cau
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dS = R − i( I, S) − mS + γ I d
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zero, the model becomes identical t
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alternative food resources than the
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theoretical extensions for understa
Page 189 and 190:
Literature cited Abrams, P.A. 1992.
Page 191 and 192:
Deng, B., S. Jessie, G. Ledder, A.
Page 193 and 194:
Hochberg, M.E., R. Gomulkiewicz, R.
Page 195 and 196:
McCauley, E., W.A. Nelson and R.M.
Page 197 and 198:
Schoener, T.W. 1986. Overview: Kind
Page 199 and 200:
Density Figure 7.1. An example of d
Page 201 and 202:
numerator), and so weakens the appa
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metacommunity theory is still in it
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1) Dispersal contributes to local c
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theory in its general form is to hi
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to disturbances and environmental c
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more realistic view would model dis
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diversity’ starts out high and de
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• Niche Theory (Chase Chapter 5)
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iogeographic contingencies that cha
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interacting organisms in metacommun
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Cottenie K. & De Meester L. (2005).
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Skellam J. (1951). Random dispersal
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Invasible local communities should
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Patch Heterogeneity Figure 8.2. The
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Figure 8.4 Experimental results of
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to each other, and the meaning of e
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adaptation, physiology, and plastic
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Proposition 2: Successional pattern
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particularities required through it
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immigration. Thus, although the spe
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and Shelford 1939), or of Holdridge
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egion. The length of the resource g
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succession states that 1) if sites
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transition probabilities from earli
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exploiting contested and low levels
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appear in those current communities
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from society. When the theory was i
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systems. That spatial and temporal
Page 257 and 258:
Botkin D.B. and Sobel M.J. 1975. St
Page 259 and 260:
Farrell, T. M. 1991. Models and mec
Page 261 and 262:
Keever C. 1979. Mechanisms of plant
Page 263 and 264:
Pacala S.W., Canham C.D. and Siland
Page 265 and 266:
Tilman D. 1991. Constraints and tra
Page 267 and 268:
Figure 9.2. An idealized filter mod
Page 269 and 270:
Figure 9.4. A model template for th
Page 271 and 272:
Patch Dynamics Metapopulation Dynam
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Table 9.1. Contrasts between classi
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and to state the permissible relati
Page 277 and 278:
Principles underlying the ecologica
Page 279 and 280:
As suggested above, not all authors
Page 281 and 282:
Channel Islands of California (Diam
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increased if dispersal limitations
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some are larger, some more fecund,
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the 19 th Century (Sax et al. 2007)
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nearly-flat, but increasing very sl
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question remains whether modificati
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maintained, despite turnover in spe
Page 295 and 296:
Literature cited Allen, A. P., J. F
Page 297 and 298:
Huey, R. B., G. W. Gilchrist, M. L.
Page 299 and 300:
Simberloff, D., and E. O. Wilson. 1
Page 301 and 302:
Table 10.1: Propositions of the Equ
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Figure 10.2. The Equilibrium Theory
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NEE, net ecosystem exchange, is the
Page 307 and 308:
Application and advancement of thes
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activities, they control the elemen
Page 311 and 312:
Theory 2: The effects of disturbanc
Page 313 and 314:
fuels, forest products, water, and
Page 315 and 316:
Literature cited Aber, J., and J. M
Page 317 and 318:
Odum, H. T. 1983. Systems ecology.
Page 319 and 320:
Table 11.1. Relationship of Ecosyst
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Figure 11.1. Net ecosystem producti
Page 323 and 324:
Figure 11.3. There are 4 possible r
Page 325 and 326:
Ecosystem responses to global chang
Page 327 and 328:
Proposition 1. Interactions across
Page 329 and 330:
where dispersal to other patches be
Page 331 and 332:
environment is often altered to per
Page 333 and 334:
transport of propagules, toxins, an
Page 335 and 336:
develops into a hurricane (Gray 199
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ecognized and better appreciated as
Page 339 and 340:
multi-site analyses has been access
Page 341 and 342:
Literature Cited Albertson, F.W., a
Page 343 and 344:
Goldenberg, S.B., C.W. Landsea, A.M
Page 345 and 346:
Liu, J. and J. Diamond. 2005. China
Page 347 and 348:
Peters, D. P. C., R. A. Pielke, Sr,
Page 349 and 350:
The State of the Nation’s Ecosyst
Page 351 and 352:
Figure captions Figure 12.1. Two pr
Page 353 and 354:
Figure 12.1. 353
Page 355 and 356:
Figure 12.3 355
Page 357 and 358:
Chapter 13: A Theory of Ecological
Page 359 and 360:
Although the constitutive theory th
Page 361 and 362:
the world: we do not claim that all
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esource is limiting for all species
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areas of greatest resource or least
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about the long-term persistence of
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the species and type of environment
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from principle 7. Proposition 3 is
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that species-abundance curves are d
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level of resources, it may be usefu
Page 377 and 378:
confronted by such challenges, ecol
Page 379 and 380:
Literature Cited Abrams, P. A. 1988
Page 381 and 382:
Grime, J. P. 1973. Competitive excl
Page 383 and 384:
Oksanen, L., S. D. Fretwell, J. Arr
Page 385 and 386:
Waide, R. B., M. R. Willig, C. F. S
Page 387 and 388:
Table 13.2. Models of diversity gra
Page 389 and 390:
Figure 13.1. Gradient theory and sp
Page 391 and 392:
Chapter 14: Biogeographical Gradien
Page 393 and 394:
scale spatial context of biogeograp
Page 395 and 396:
each for many compelling and longst
Page 397 and 398:
heat source). Parameters were varie
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et al. (2001) (Proposition 9). In t
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anges are larger, given the same to
Page 403 and 404:
Models in this group have begun to
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exploration. With the empirical spe
Page 407 and 408:
example, nearly 70% of the 241 spec
Page 409 and 410:
that the latitudinal gradient in me
Page 411 and 412:
fourth issue is that the absolute s
Page 413 and 414:
Literature Cited Bell, G. 2000. The
Page 415 and 416:
Dobzhansky, T. 1950. Evolution in t
Page 417 and 418:
Kerr, J. T., M. Perring, and D. J.
Page 419 and 420:
Rahbek, C. and G. R. Graves. 2000.
Page 421 and 422:
Willig, M. R., D. M. Kaufmann, and
Page 423 and 424:
equires additional exposition to un
Page 425 and 426:
population, community) may conspire
Page 427 and 428:
evolve.” Using a historical persp
Page 429 and 430:
dynamics theory (Hastings Chapter 6
Page 431 and 432:
group of ecologists who were frustr
Page 433 and 434:
understanding the conditions that f
Page 435 and 436:
the end. But it is, perhaps, the en
Page 437 and 438:
Hagen, J. B. 1992, An Entangled Ban
Page 439 and 440:
Figure 15.1. The biological organiz
Page 441:
Figure 15.3. This conceptual model
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