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

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can straddle two or more general theories if some of its models ultimately address a central question of each general theory. One way to decide if a constitutive theory straddles two general theories is to consider the assumptions of those general theories. If the constitutive theory simply accepts all of the assumptions in a particular general theory and never questions or tests them, it likely is not a member of that general theory. A corollary of the previous statement is that any given model of necessity explores or tests one or more of the assumptions, fundamental principles or propositions of a theory. For example, a continuing issue in ecology concerns the identity of parameters that can be treated as constants and those that need to be treated as variables in a particular theory or model. If a parameter is treated as a constant, the average value of that parameter is assumed to be sufficient because either the variation has no effect or acts in a strictly additive fashion relative to the causative mechanisms under examination. In some instances, ecologists make assumptions without ever testing them. For example, it is reasonable to assume that we can average over quantum fluctuations (from the domain of physics) in ecological processes. On the other hand, the physiological variations that occur from minute to minute in a mammal so as to maintain body temperature (from the domain of the theory of organisms) (Scheiner submitted; Zamer and Scheiner in prep.) may matter for ecological processes and should not be averaged in some instances. For example, basal metabolic rates in large mammals can vary substantially between winter and summer. Failure to account for this variation can seriously overestimate winter energy expenditures and food intake requirements (Arnold et al. 2006). A subdomain can overlap two domains. For example, ecosystem science has some constituent theories that are part of ecology and some that are part of geosciences. Such overlaps can extend to the level of individual models. For example, foraging theory (Sih Chapter 4) contains some models that are ecological, others that are evolutionary, and others that are both. This sharing of subdomains shows that the boundaries of domains are not distinct and can be somewhat arbitrary. A domain as defined by a general theory, constitutive theory, or model should be a coherent entity. Some named areas are not domains, but collections of domains. For example, evolutionary ecology consists of a set of constituent theories, some of which are within the domain of the theory of ecology and others that are within the domain of the theory of evolution. 10 10
Physiological ecology is likely another such collection of domains, although at this time it is difficult to tell because a theory of physiology has not yet been articulated. THE FUNDAMENTAL PRINCIPLES OF ECOLOGY The general theory of ecology consists of eight fundamental principles (Table 1.3). The roots of these principles can be traced to the origins of ecology in the 19th century. They were in place and widely accepted by the 1950s, were recently codified as the components of a general theory (Scheiner and Willig 2008), and continue to evolve. In particular, we have added an eighth fundamental principle (number 3) so that the numbering of this set differs somewhat from our previous list. Heterogeneous distributions The first fundamental principle – the heterogeneous distribution of organisms – is a refinement of the domain of the theory of ecology. The heterogeneity of distributions is one of the most striking features of nature: all species have a heterogeneous distribution at some if not most spatial scales. Thus, this principle encompasses a basic object of interest, is its most important property, and serves to guide the rest of the theory. All of the other parts of the theory of ecology serve to either explain this central observation or to explore its consequences. Arguably, the origins of ecology as a discipline and the first ecological theories can be traced to its recognition (Forster 1778; von Humboldt 1808). This heterogeneous distribution is both caused by and a cause of other ecological patterns and processes. Environmental interactions The second fundamental principle – interactions of organisms – includes within it the vast majority of ecological processes responsible for heterogeneity in time and space. They include both intraspecific and interspecific interactions such as competition, predation, and mutualism, as well as feedbacks between biotic and abiotic components. Within this principle, particular interactions that are part of constituent theories act to elaborate the general theory (see later chapters). Many definitions of ecology are restatements of this principle (Scheiner and Willig 2008). 11 11
Page 1 and 2: TABLE OF CONTENTS THE THEORY OF ECO
Page 3 and 4: Chapter 1: A General Theory of Ecol
Page 5 and 6: Chapter 10). Without such boundarie
Page 7 and 8: profound. As the statistician Georg
Page 9: In general, the domain of a theory
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
Page 57 and 58: 1969 Individual thermodynamic budge
Page 59 and 60: 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
Page 63 and 64:
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
Page 73 and 74:
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
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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
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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
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Salo, P., E. Korpimaki, P.B. Banks,
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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
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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
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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
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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
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Botkin D.B. and Sobel M.J. 1975. St
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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
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Principles underlying the ecologica
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As suggested above, not all authors
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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
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Theory 2: The effects of disturbanc
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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
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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
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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
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Chapter 13: A Theory of Ecological
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Although the constitutive theory th
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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
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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
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scale spatial context of biogeograp
Page 395 and 396:
each for many compelling and longst
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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
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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
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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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