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

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Variation of individuals The third principle – the variation of individuals – is the result of processes that derive from the theory of organisms (Scheiner submitted; Zamer and Scheiner in prep.). Ecological theories make predictions about the characteristics or aggregate properties of species, individuals or traits. The majority of ecological theories make predictions about species or collections of species (e.g., species richness of communities; see Chapters 8-10, 13, 14). Some theories, such as population ecology and behavioral ecology, concern themselves with predictions about individuals or collections of individuals (e.g., numbers of individuals in a population; see Chapters 4-8). Some theories make predictions about the properties of individuals or species (e.g., body size distributions; see Chapters 4, 5, 8, 10, 13, 14). Finally, some theories make predictions about the aggregate properties of individuals or species (e.g., ecosystem standing biomass; see Chapter 11). Groups of species or individuals share the property that the members of those groups differ in their characteristics, even though many theories and models assume invariance. For example, one of the most common hidden assumption in models of species richness is that all individuals within a species are identical (e.g., Fox et al. Chapter 13). Such assumptions may be reasonable for the purposes of simplifying models. Violations of this assumption may not substantially change predictions. However, in some cases relaxing this assumption has led to substantial changes in predictions. For example, when the chances of survival are allowed to vary among individuals within a population, treating all individuals as equal turns out to substantially mis- estimate the risk of local extinction from demographic stochasticity; depending on the model used for reproduction, treating all individuals as identical can over- or underestimate that risk (Kendall and Fox 2003). Contingency The fourth fundamental principle – contingency – has grown in importance in ecological theory and now appears in a wide variety of constituent theories and models. By contingency we mean the combined effects of two processes – randomness and sensitivity to initial conditions. Contingency is an important cause of the heterogeneous distribution of organisms, both at very large and very small extents of time and space (e.g., a seed lands in one spot and not another; a particular species arises on a particular continent). This principle exemplifies the dynamic nature 12 12
of a theory. A theory is constantly evolving, although substantive change typically occurs over decades. One hallmark of that dynamic is the emergence of new principles, such as occurred with this principle during the 1960s to 1980s. Heterogeneity of environmental conditions The fifth fundamental principle – environmental heterogeneity – is a consequence of processes from the theories of earth and space sciences when the environmental factors are abiotic, as well as the consequences of the second principle when those factors are biotic. For example, seasonal variation in temperature is the result of orbital properties of the Earth, whereas a variety of geophysical processes create heterogeneity in environmental stressors like salt (e.g., wave action near shores) or heavy metals (e.g., geologic processes that create differences in bedrocks). This principle is part of many constituent theories and contains a broad class of underlying mechanisms for the heterogeneous distribution of organisms, as seen in many of the constitutive theories presented in this book. As with the second principle, particular mechanisms pertain to particular constituent theories. Finite and heterogeneous resources The sixth principle – finite and heterogeneous resources – is again a consequence of processes from the theories of earth and space sciences or the second principle. Although variation in resources is similar to variation in environmental conditions, a fundamental distinction is the finite, and thus limiting, nature of these resources. Unlike an environmental condition, a resource is subject to competition. For example, seasonal variation in light and temperature are caused by the same orbital mechanisms, but light is subject to competition (e.g., one plant shades another) whereas temperature is a condition and not subject to competition. This distinction in the nature of environmental factors with regard to competitive processes can result in different ecological outcomes. For example, β diversity in plant communities is high in warm deserts and low in arctic tundra because diversity in warm deserts is controlled by water, a limiting resources, while diversity in arctic tundra is controlled by temperature, an environmental condition (Scheiner and Rey-Benayas 1994). Whether a particular environmental factor is a condition or a resource can be context dependent. For example, water is sometimes a resource subject to competition (e.g., plants in a desert) and sometimes a condition (e.g., fish in the ocean). Some heavy metals (e.g., 13 13
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 and 10: In general, the domain of a theory
Page 11: Physiological ecology is likely ano
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
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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;
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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
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functions that are part of these mo
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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.
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Charnov, E.L., G.H. Orians and K. H
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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
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Literature Cited Abrams, P. A. and
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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
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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
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Literature cited Abrams, P.A. 1992.
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Deng, B., S. Jessie, G. Ledder, A.
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Hochberg, M.E., R. Gomulkiewicz, R.
Page 195 and 196:
McCauley, E., W.A. Nelson and R.M.
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Schoener, T.W. 1986. Overview: Kind
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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
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Keever C. 1979. Mechanisms of plant
Page 263 and 264:
Pacala S.W., Canham C.D. and Siland
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Tilman D. 1991. Constraints and tra
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Figure 9.2. An idealized filter mod
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Figure 9.4. A model template for th
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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
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Literature cited Allen, A. P., J. F
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Huey, R. B., G. W. Gilchrist, M. L.
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Simberloff, D., and E. O. Wilson. 1
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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
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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
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Odum, H. T. 1983. Systems ecology.
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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
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Ecosystem responses to global chang
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Proposition 1. Interactions across
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where dispersal to other patches be
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environment is often altered to per
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transport of propagules, toxins, an
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develops into a hurricane (Gray 199
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ecognized and better appreciated as
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multi-site analyses has been access
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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,
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The State of the Nation’s Ecosyst
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Figure captions Figure 12.1. Two pr
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Figure 12.1. 353
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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
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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
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Figure 13.1. Gradient theory and sp
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Chapter 14: Biogeographical Gradien
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scale spatial context of biogeograp
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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
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example, nearly 70% of the 241 spec
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that the latitudinal gradient in me
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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
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evolve.” Using a historical persp
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dynamics theory (Hastings Chapter 6
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group of ecologists who were frustr
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understanding the conditions that f
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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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