Professional Documents
Culture Documents
net/publication/44689600
CITATIONS READS
1,666 14,592
1 author:
Mark Vellend
Université de Sherbrooke
175 PUBLICATIONS 20,040 CITATIONS
SEE PROFILE
Some of the authors of this publication are also working on these related projects:
All content following this page was uploaded by Mark Vellend on 15 April 2014.
Mark Vellend
Departments of Botany and Zoology, and Biodiversity Research Centre, University of British Columbia,
Vancouver, British Columbia, Canada, V6T 1Z4
e-mail: mvellend@interchange.ubc.ca
keywords
dispersal, drift, community ecology, population genetics, selection, speciation
abstract
Community ecology is often perceived as a “mess,” given the seemingly vast number of processes that
can underlie the many patterns of interest, and the apparent uniqueness of each study system.
However, at the most general level, patterns in the composition and diversity of species—the subject
matter of community ecology—are influenced by only four classes of process: selection, drift, speciation,
and dispersal. Selection represents deterministic fitness differences among species, drift represents
stochastic changes in species abundance, speciation creates new species, and dispersal is the movement
of organisms across space. All theoretical and conceptual models in community ecology can be
understood with respect to their emphasis on these four processes. Empirical evidence exists for all of
these processes and many of their interactions, with a predominance of studies on selection. Organizing
the material of community ecology according to this framework can clarify the essential similarities and
differences among the many conceptual and theoretical approaches to the discipline, and it can also
allow for the articulation of a very general theory of community dynamics: species are added to
communities via speciation and dispersal, and the relative abundances of these species are then shaped
by drift and selection, as well as ongoing dispersal, to drive community dynamics.
183
184 THE QUARTERLY REVIEW OF BIOLOGY Volume 85
the metacommunity concept (Holyoak et al. motivation for conceptually organizing the
2005), which is explicitly concerned with the material in community ecology, and pro-
role of dispersal among local communities in vide operational definitions of important
influencing community patterns at multiple terms. I then illustrate, with separate sec-
scales. The movement of organisms across tions on theory and data, how the subject
space can have a variety of important conse- matter of community ecology can be pre-
quences in communities. sented using the proposed organizational
For each of the latter three processes—spe- framework, describing the ways in which
ciation, drift, and dispersal—conceptual devel- selection, drift, speciation, and dispersal
opments were motivated by a perceived lack of influence communities. I then touch on
emphasis in the literature on the impor- some of the general patterns that commu-
tance of the process in question. Selection, nity ecologists have traditionally been in-
in the form of deterministic interactions terested in, and I discuss how pattern is
among species and between species and connected with process. Finally, I compare
their environments, was always recognized the framework presented here with other
as important. With the additions of specia- conceptual frameworks in community ecol-
tion, drift, and dispersal, we now have a ogy.
logically complete set of process categories
within which all other more specific pro- Community Ecology Is a Mess
cesses can be placed. I believe that organiz- Based largely on empirical results, Lawton
ing the overwhelming number of specific (1999) famously called community ecology
ecological theories for communities under “a mess,” and ascribed this mess to the inher-
this scheme can help achieve at least two ent contingency of ecological patterns on
important goals. First, the essential similar- the details of how the underlying processes
ities and differences between different eco- or rules act. “The rules are contingent in so
logical models can be clarified in fairly many ways . . . as to make the search for
straightforward terms, thereby making the patterns unworkable” (Lawton 1999:181).
full set of models easier to understand, One source of motivation for the present
apply, and teach to students. Second, we paper is that even theoretical community
can articulate a very general theory of com- ecology can be considered a mess for much
munity dynamics, which may on the sur- the same reason: each and every twist added
face sound obvious and too generalized to to theoretical models seems to matter, mak-
make any specific predictions, but may, ing an overarching treatment of the subject
nonetheless, serve the same critical func- very difficult. Consider the number of differ-
tion as foundational theory in population ent models that can be constructed from the
genetics. simple Lotka-Volterra formulation of inter-
Before proceeding, I should emphasize actions between two species by layering on
that I am not arguing that the parallels be- realistic complexities, one by one. First, there
tween processes or models in population ge- are at least three qualitatively distinct kinds
netics and community ecology are perfect. of interactions (competition, predation, mu-
For example, selection among individuals tualism). For each of these, we can have ei-
across species can be manifested in ways that ther an implicit accounting of basal re-
are rare or absent within species (e.g., tro- sources (as in the Lotka-Volterra model), or
phic or parasitic interactions), and specia- we can add an explicit accounting in one
tion is a far more complicated process than particular way. That gives six different mod-
mutation. The list could go on. Rather, my els so far. We can then add spatial heteroge-
argument is that we can define a similar set neity or not (⫻2), temporal heterogeneity or
of four logically distinct processes in commu- not (⫻2), stochasticity or not (⫻2), immigra-
nity ecology in order to provide a coherent tion or not (⫻2), at least three kinds of func-
conceptual framework for the discipline. tional relationships between species (e.g.,
The rest of this paper is structured as predator functional responses; ⫻3), age/size
follows. I first specify more precisely the structure or not (⫻2), a third species or not
186 THE QUARTERLY REVIEW OF BIOLOGY Volume 85
(⫻2), and three ways that the new species A Theory of What Is Possible
may interact with one of the existing species Amazingly, the foundation of theoretical
(⫻3 for the models with a third species). population genetics was built in the near
Having barely scratched the surface of po- absence of data on patterns of genetic vari-
tentially important factors, we have 2304 dif- ation in natural populations—the very sub-
ferent models. Many of them would likely ject matter of the discipline (Provine 1971).
yield the same predictions, but, after consol- Perhaps for this reason, at least in part, a
idation, I suspect there still might be hun- theoretical foundation was built to de-
dreds that differ in ecologically important ways. scribe a logically complete range of the
As Lawton (1999) put it, “the necessary contin- basic possible processes that could cause
gent theory looks unworkably complicated” (p. evolutionary change, rather than a theory
180). skewed towards an emphasis on those pro-
One important manifestation of this mess cesses that are actually important in na-
is that textbook treatments of community ture. The latter is an empirical rather than
ecology and their associated university cours- theoretical issue. In contrast, long before
es—that is, the vehicles by which the subject the existence of ecological theory, patterns
matter is taught to students— have a struc- in nature were well-known to any keen ob-
ture whose logic is not easy to discern. Sec- server. Allen and Hoekstra (1992) describe
tion or chapter topics typically fall loosely ecology as a discipline “whose material study
under one or more of the following head- is part of everyday encounters: birds, bees,
ings: community patterns, competition, pre-
trees, and rivers” (p. 1). They go on to argue,
dation (plus other enemy-resource interac-
albeit in a somewhat different context, that
tions), niches, food webs, and issues of space
“It is, however, a mistake to imagine that this
and time (e.g., Putman 1994; Morin 1999;
familiarity makes ecology an easy pursuit-
Ricklefs and Miller 1999). This is a confusing
. . .the very familiarity of ecological objects
list because it includes unlike entities—pat-
presents the difficulties” (Allen and Hoek-
terns, processes (competition, predation),
stra 1992:1). I argue that this everyday fa-
concepts (niches, food webs), objects of
miliarity with ecological patterns pushed
study (food webs), or a consideration that is
ecological theory down the path of empha-
always important to think about (space and
sizing particular viewpoints on the pro-
time) (Vellend and Orrock 2009). In con-
trast, books and courses in population genet- cesses that are actually most important in
ics (e.g., Hartl and Clark 1997) are based nature, rather than emphasizing a logically
upon a structure that is easier to follow, with complete set of possible processes that
a consistent focus on the four processes of must play at least some role in community
selection, drift, gene flow, and mutation, and dynamics. The emphasis in ecology, there-
how these processes either individually or fore, has been on pattern before process
jointly determine patterns of genetic varia- (Roughgarden 2009; Vellend and Orrock
tion. In my opinion, ecology textbooks and 2009). Using the structure of population ge-
courses are a fairly accurate reflection of the netics theory as a guide, with details altered
way in which practicing community ecolo- where necessary for communities, the follow-
gists have self-organized around particular ing presents an organizational scheme for
research topics or themes, but I am not con- community ecology, within which all specific
vinced that this is the best way to organize models and frameworks can be described.
the subject matter for facilitating synthetic
and integrated understanding by students definitions
and practitioners alike. As elaborated below, Table 1 provides operational definitions
selection, drift, speciation, and dispersal may of the key terms used in this paper. With
not be of equal importance in understand- respect to the definition of community,
ing ecological patterns, but they fully repre- there has been considerable debate in
sent the logically distinct categories of impor- ecology concerning the degree to which
tant processes in community ecology. ecological communities are sufficiently co-
June 2010 SYNTHESIS IN COMMUNITY ECOLOGY 187
TABLE 1
Definitions of terms
Term Definition
Community A group of organisms representing multiple species living in a specified place
and time
Community ecology The study of patterns in the diversity, abundance, and composition of species
in communities, and the processes underlying these patterns
Community dynamics Changes over time in the relative abundances of species in a specified area,
including extinctions and species additions via dispersal or speciation
Species composition For a given community, a state defined by the abundances of all species
Species relative abundance The proportion of all organisms in a given area that are of a given species;
equivalent to species frequency.
Species density The number of organisms of a given species per unit of space
Community size The total number of organisms in a community
Coexistence The indefinite persistence of a specified set of species in a specified area
Absolute fitness The quantity of offspring produced by an individual organism per unit of
time, including survival of the organism itself
Relative fitness The absolute fitness of a given organism divided by the mean absolute fitness
across all individuals in the community
Species fitness (absolute or relative) The mean fitness (absolute or relative) across all individuals of a given
species in the community; for absolute fitness, this is equivalent to the
species per capita population growth rate.
Selection A deterministic fitness difference between individuals of different species
Drift Random changes in species relative abundances
Neutrality A state in which all individual organisms share identical demographic
properties
Speciation The creation of new species
Dispersal The movement of organisms across space
herent entities to be considered appropri- species diversity are considered part of commu-
ate objects of study (reviewed in Ricklefs nity ecology here).
2008). The definition of community used
here—that is, a group of organisms repre- The Four Processes of Community
senting multiple species living in a speci- Ecology: Theory
fied place and time— bypasses this issue by selection
recognizing that properties of communi-
ties are of central interest in ecology, re- Use of the term “selection” to describe
gardless of their coherence and integrity. deterministic fitness differences among in-
dividuals of different species (Table 1) re-
This definition of community also implic-
quires some explanation, as it is not yet
itly embraces all scales of space and time.
commonplace in ecology (but see Loreau
Studying communities in 1m2 plots or across and Hector 2001; Norberg et al. 2001;
entire continents requires different methods, Shipley et al. 2006; Bell 2008). Although
and the relative importance of different pro- the term is used in biology most often with
cesses likely varies across scales, but we are of- respect to evolutionary dynamics within
ten interested in understanding the same species, the definition of selection in no way
kinds of patterns (e.g., diversity, composition) restricts its application as such. Selection oc-
at these different scales. This represents an ex- curs when individuals in a population vary in
pansion of the purview of community ecology some respect, and when different variants
beyond its traditional focus on relatively small reproduce or replicate themselves at differ-
scales, without applying a new name to the ent rates (Darwin 1859; Bell 2008; Nowak
discipline (e.g., studies in “macroecology” of 2006). In its most generalized form, the con-
188 THE QUARTERLY REVIEW OF BIOLOGY Volume 85
cept of selection—and, more broadly, evolu- before doing so, it is first necessary to establish
tionary change— can be applied “based only the basic building blocks of community ecol-
on the assumption of a population of things ogy, with species as the fundamental category
that leave descendants and have measurable of accounting in the assessment of community-
phenotypes” (Rice 2008:4). level phenomena.
Applying the concept of selection to spe- In a community context, there are three
cies in a community rather than to alleles in relevant forms of selection: (1) constant,
a species’ population requires two changes (2) frequency- or density-dependent, and
to the frame of reference. First, rather than (3) spatially- or temporally-variable selec-
invoking selection at any level higher than tion. Constant selection is simple: if relative
that of the individual organism, we simply fitness is constant in space and time, inde-
define the “population” as containing indi- pendent of species’ densities but variable
viduals of multiple species; we call this pop- across species, the species with the highest
ulation a community. Second, the phenotype fitness will exclude all others (Figure 1A).
of interest, which may be under selection, The other forms of selection, however, are
is most often just the species identity. The more complicated.
species identity is a categorical phenotype, Frequency- or density-dependent selection is
assumed to have perfect heritability, ex- central to the vast majority of theoretical mod-
cept when speciation occurs, after which els with species interactions in community ecol-
new species identities are assigned (just as ogy. For simplicity, I will only use the term
mutation changes the identity of an allele). “density-dependent,” given that most ecologi-
In the same way that selection may favor cal models include densities rather than fre-
allele A over allele a within a species’ pop- quencies—a key distinction from the tradition
ulation, selection may favor species X over in population genetics (Lewontin 2004). If
species Y in a community. It is important to community size is constant, density and fre-
note that although the concept of selection quency are equal (as in Figure 1, for simplicity
in communities is easier to envision for of presentation). Density-dependent selection
species on the same trophic level than for occurs when individual fitness in a given
species on different trophic levels, the dif- species depends at least in part on the den-
ference is one of degree and not kind. For sity of that species, as well as the densities
example, a lynx and a hare are very differ- of other species. For two species, negative
ent organisms, but selection still favors density-dependent selection favors species
hares when lynx are declining, and it favors when they are at low density and is thus
lynx when hares are abundant (Krebs et al. capable of maintaining stable coexistence
2001). (Figure 1B), whereas positive frequency-
Rather than focusing only on species iden- dependent selection favors species at high
tities, it is also possible to define each species density and cannot maintain stable coexist-
by one or more traits (e.g., beak depth, leaf ence (Figure 1C). Selection can also de-
thickness) (McGill et al. 2006) and to then pend on species densities in more complex
apply tools from quantitative genetics at the ways, possibly allowing more than one stable
community level (e.g., Norberg et al. 2001; state at which coexistence can be main-
Shipley et al. 2006). This opens the door to tained (Figure 1D), or creating repeated
simultaneous consideration of selection both oscillations in interacting species’ abun-
within and among species. However, to sim- dances (Morin 1999). A major challenge in
plify the discussion and focus attention most ecology is presented by the nearly limitless
sharply at the community level, I hence- variety of configurations that the full set of
forth address selection in communities by intra- and interspecific density dependen-
assuming that individuals of a given species cies can take in a species-rich community.
have the exact same phenotype (e.g., the The nature of density-dependent selec-
species identity). Relaxing this assumption tion between pairs of species depends on
forms the basis of a very active area of the qualitative ecological relationship be-
research (e.g., Hughes et al. 2008), but, tween them (e.g., competition, predation,
June 2010 SYNTHESIS IN COMMUNITY ECOLOGY 189
mutualism, and disease) and the quantita- 1999). From a given set of initial conditions,
tive form of this relationship. With more outcomes such as the exclusion of all but one
than two species, indirect interactions can species, the indefinite coexistence of all spe-
arise whereby fitness in one species de- cies, complex temporal fluctuations, or en-
pends on the density of a second species not tirely different equilibrium patterns depending
because of a direct interaction, but because on initial conditions are possible.
each of the two species interacts directly with a Selection, whether constant or density-
third (Strauss 1991). Even the nature of the dependent, may vary across space or time, with
direct interaction between two species can be potentially important consequences for com-
influenced by other species, amplifying even munity dynamics. Most importantly, the behav-
further the number of ways a community can ior of such models can deviate qualitatively
be configured. A massive edifice of theoretical from spatiotemporally invariant models when
research has addressed the community conse- the relative fitness of different species switches
quences of different forms of density- in different places or times, thereby allowing
dependent interactions among species (Morin for coexistence among species that would oth-
190 THE QUARTERLY REVIEW OF BIOLOGY Volume 85
erwise not be possible (Levene 1953; Ches- tion of how the species in a given area arose in
son 2000). More generally, species coexist- the first place, leaving such questions to the
ence always depends on trade-offs of some fields of biogeography and macroevolution
kind, with different species having fitness (Ricklefs 1987; Brown 1995). From the per-
advantages under different sets of condi- spective of understanding how species in-
tions, specified by some combination of teractions play out in homogeneous, small-
the abiotic environment and the densities scale localities, this is entirely defensible,
of the species themselves (Chesson 2000). because the origin of the local species pool
does not matter; what does matter is that
drift the species are present locally and possess
Because birth, death, and offspring pro- a given set of traits. But to compare com-
duction are inherently stochastic pro- munity patterns across different regions,
cesses, changes in any community with a and even across environmental gradients
finite number of individuals will also have a at quite local scales, it may be important to
stochastic component. This is ecological incorporate the biogeographic and macro-
drift. If individual-level demographic pa- evolutionary context in which the species
rameters are identical across all individuals pool originated (Ricklefs 1987; Ricklefs
in a closed community, drift is the only and Schluter 1993; Pärtel 2002). We can no
driver of community dynamics (i.e., there more afford to exclude speciation from com-
are no deterministic changes in abun- munity ecology than we can afford to exclude
dance; Figure 1E) and, eventually, all but mutation from population genetics, even if
one species will drift to extinction. The speciation is a far more complex process.
probability of each species reaching mono- I deliberately focus on speciation rather
dominance is equal to its initial frequency, than embracing extinction under this um-
and the rate at which this is achieved is brella as well, because, with an expanded spa-
negatively related to community size (Fig- tial and temporal scope of community ecology
ure 2). As such, declines in community size (see definitions section above), extinction is
(i.e., disturbance) may increase the impor- best considered as an outcome of selection and
tance of drift. Importantly, drift need not drift, rather than as a distinct process in and of
act alone to have an important impact on itself. When the last individual of a species dies,
community dynamics. The interaction of the species is extinct, and while the decline to
drift with speciation and dispersal (Hub- extinction may have many specific causes, they
bell 2001) will be described in subsequent must either be deterministic (selection) or sto-
sections; here, I address the interaction of chastic (drift). Even major geological events
drift and selection. (e.g., glaciation) are distinguished from more
If selection is relatively strong and the subtle environmental changes (e.g., slight acid-
community size is large, selection will over- ification of a lake) as agents of selection by
ride any effects of drift. But if selection is the rate, magnitude, and spatial scale of
relatively weak and the community size is change, rather than by a qualitatively distinct
small, drift can override the effects of selec- influence on communities. Such environ-
tion. Between these two extremes, selection mental changes may also alter the effects of
makes some community outcomes more drift via changes in community size.
likely than others, but it does not guarantee I focus here on some of the simplest ways
any particular outcome (Nowak 2006). For that speciation has been incorporated into the-
example, even with constant selection favor- oretical community models, as well as some
ing one of two species, there is some proba- empirically-motivated conceptual models. At
bility that the species with the higher fitness large spatial scales, such as entire conti-
will drift to extinction (Figure 2). nents, the rate of speciation can enter
mathematical models directly as a key de-
speciation terminant of community dynamics. For ex-
Most treatments of community ecology in- ample, Hubbell (2001) considered a neu-
herently exclude from their purview the ques- tral community of fixed size in which the
June 2010 SYNTHESIS IN COMMUNITY ECOLOGY 191
speciation rate is constant, with the rate of each species’ population must be smaller. Al-
extinction due to drift increasing with the ternatively, MacArthur (1969) posited that
number of species because, with more species, both speciation and extinction rates increase
192 THE QUARTERLY REVIEW OF BIOLOGY Volume 85
with the number of species, but with the ers, two kinds of models represent the ends of
increase decelerating for speciation and ac- a continuum. Mainland-island models assume
celerating for extinction, thus resulting in an one-way dispersal from a source community of
equilibrium number of species in the region. effectively infinite size (the mainland) to one
These models correspond to drift-speciation or more smaller, discrete local communi-
balance and selection-speciation balance, re- ties (the islands, or localities). These mod-
spectively, and, in both cases, a greater rate els assume that community dynamics in
of speciation leads to a larger species pool, small localities are sufficiently rapid rela-
all else being equal. In a model of multiple tive to those on the mainland, such that
local communities connected by dispersal the composition of the pool of dispersers is
along an environmental gradient, McPeek effectively constant. In contrast, island
(2007) found that the nature of the specia- models assume a network of small local
tion process influenced local species diver- communities linked by dispersal among
sity: greater ecological similarity between them, with no distinct mainland. Such net-
new and existing species extended times to works of local communities may be called
extinction, thereby elevating local species di- “metacommunities” (Holyoak et al. 2005).
versity at any given time. More conceptual Dispersal can interact with drift and specia-
(rather than mathematical) models address tion. In a mainland-island model with local
the consequences of variation in the rate at drift but no speciation or selection, dispersal
which species that are adapted to particular increases local species richness and causes local
conditions (e.g., regionally common or rare community composition to converge with that
abiotic conditions) are produced. The term of the mainland. For a given level of dispersal,
“species pool hypothesis” has been used to the number of new species introduced per unit
describe this type of conceptual model (Tay- of time will decrease as local species richness
lor et al. 1990). Through its effects on the increases, because fewer and fewer of the dis-
regional species pool, speciation then indi- persers will represent new species in the local-
rectly becomes a potentially important deter- ity. With fixed local community size, greater
minant of community dynamics and pat- species richness necessitates smaller popula-
terns, even at a local scale where the rate of tion size per species, so that the rate of species
speciation is negligible relative to other pro- extinction increases with species richness, at
cesses (e.g., Ricklefs and Schluter 1993; Pär- some point equaling the rate of species intro-
tel 2002). duction, and thus determining an equilibrium
number of species whose identities nonethe-
dispersal less change through time. This is the simplest
Dispersal involves the movement of or- form of the theory of island biogeography
ganisms across space, and, thus, its influ- (MacArthur and Wilson 1967). In an island
ence on community dynamics depends on model with local drift but no speciation or se-
the size and composition of the communi- lection, dispersal increases local diversity by
ties where the dispersers come from and of countering drift, and causes the similarity in
those in which they disperse to (Holyoak et composition among localities to increase
al. 2005). As such, the community conse- (Wright 1940). Without input via speciation
quences of dispersal can only be addressed in or dispersal from a separate metacommunity,
relation to the action and results of other pro- all but one species will ultimately drift to extinc-
cesses, selection and drift in particular. The tion. In a model with drift, speciation, and dis-
construction of theoretical community models persal but no selection, all community patterns
addressing the role of dispersal usually speci- are determined by the size of the local commu-
fies whether organisms are distributed contin- nities, the size of the entire metacommunity,
uously across space or in discrete patches. The and the rates of speciation and dispersal (Hub-
latter type of distribution will be adopted here bell 2001).
for the sake of simplicity and clarity. Not surprisingly, the range of outcomes
With respect to the relative sizes of the when dispersal interacts with selection is vast.
source and recipient communities for dispers- The nature of selection among species in one
June 2010 SYNTHESIS IN COMMUNITY ECOLOGY 193
locality can take many different forms, and dis- tual frameworks in community ecology to
persal implies multiple localities, each of which their emphasis on selection, drift, specia-
may represent a unique selective environment. tion, and dispersal. Briefly, the idea of spe-
The consequences of dispersal depend on cies “niches” (Chase and Leibold 2003) is
these details. Also, in addition to the many essentially synonymous with selection, and
kinds of trade-offs in different models of pure many models of species interactions repre-
selection, dispersal ability itself may vary among sent different manifestations of selection.
species, possibly with a trade-off involving other Adding demographic stochasticity to selection-
aspects of fitness. Enumerating all possible based models represents a combination of
ways that dispersal can interact with selection drift and selection (e.g., Tilman 2004), as
and drift is well beyond the scope of the do models proposed under the rubric of a
present paper, but three generalized examples niche-neutral reconciliation (e.g., Shipley et
will provide us with a glimpse into how selec- al. 2006; Adler et al. 2007). The species-pool
tion and dispersal can interact. First, if selec- hypothesis (Taylor et al. 1990) and the
tion via competition or predation causes a spe- broader conceptual framework, based on
cies to go locally extinct, that species may the interaction between local and regional
nonetheless persist regionally, along with its processes (Ricklefs and Schluter 1993), rep-
competitor/predator, if it has a superior ability resent the interaction between speciation
to disperse to “open” sites where the supe- and selection and, to some degree, dispersal.
rior competitor/predator has gone ex- Classic island biogeography theory
tinct due to either an absence of prey or (MacArthur and Wilson 1967) represents a
for other reasons (Caswell 1978; Tilman balance between drift and dispersal, and
1994). Second, if selection favors different the full version of Hubbell’s (2001) neutral
species in different patches, dispersal can theory represents the combined influence
nonetheless maintain persistent local pop- of drift, dispersal, and speciation. Meta-
ulations of species, even in patches where community theory (Holyoak et al. 2005)
they are at a fitness disadvantage (Levene and the many specific models that fall into
1953). If species vary in their mean fitness this category, such as those involving colo-
across patches, then very high dispersal will nization-competition tradeoffs or “mass ef-
allow the species with the highest average fects,” emphasize dispersal first and fore-
fitness to exclude all others. Finally, in a most, and how dispersal interacts with
mainland-island context, dispersal to the selection and drift.
island determines the species pool in a way
closely analogous to the role of speciation The Four Processes of Community
on continents. Ecology: Data
A vast amount of empirical literature ad-
relating existing theories to the dresses the processes underlying the dy-
four processes namics of ecological communities. The
Four processes— or any four items— can purpose of this section is to illustrate the kinds
be considered singly or in combination in of evidence available from lab experiments-
15 different ways: each of the four alone, ,field experiments, and observations of nature
six pairwise combinations, four trios, and that speak to the importance of various forms
all four together. However, it is impossible of selection, drift, speciation, and dispersal in
to build a theoretical community model communities.
with only speciation and/or dispersal with-
out specifying the fate of new species or selection
dispersers, particularly with respect to se- Case studies of selection in ecological com-
lection or drift. Thus, the four processes munities number in the thousands, and most
can form the basis of theoretical models in communities documented in these studies ap-
12 different ways. pear to be characterized by unique combina-
Table 2 relates many of the influential tions of selective factors (Diamond and Case
and familiar theories, models, and concep- 1986; Putman 1994; Lawton 1999; Morin
194 THE QUARTERLY REVIEW OF BIOLOGY Volume 85
TABLE 2
Twelve combinations of selection, drift, speciation, and dispersal, and the ways in which existing ecological
theories relate to these combinations
Representative
Combination Selection Drift Speciation Dispersal Theories and models references
1 ⫻ Niche models of all kinds Tilman (1982); Chase
(e.g., resource & Leibold (2003)
competition, predator-
prey, food webs)
2 ⫻ Neutral theory I Hubbell (2001)
(demographic
stochasticity)
3 ⫻ ⫻ Niche-neutral models Tilman (2004); Adler
(any niche model with et al. (2007)
demographic
stochasticity)
4 ⫻ ⫻ Historical/regional MacArthur (1969);
ecology I (species pool Ricklefs (1987)
theory, diversity on
gradients, speciation-
selection balance)
5 ⫻ ⫻ Neutral model II (non- Hubbell (2001)
spatial)
6 ⫻ ⫻ Metacommunities - Holyoak et al. (2005)
deterministic (spatial
mass effects, spatial
food webs)
7 ⫻ ⫻ Neutral model III (island MacArthur & Wilson
biogeography) (1967); Hubbell
(2001)
8 ⫻ ⫻ ⫻ Historical/regional Ricklefs (1987)
ecology II
9 ⫻ ⫻ ⫻ Historical/regional Ricklefs (1987)
ecology III
10 ⫻ ⫻ ⫻ Neutral model IV Hubbell (2001)
(spatial)
11 ⫻ ⫻ ⫻ Metacommunities - Holyoak et al. (2005)
stochastic (colonization-
competition tradeoffs,
stochastic versions of
six)
12 ⫻ ⫻ ⫻ ⫻ The theory of ecological This paper
communities
1999). Important factors that underlie the in- types of limiting resources (e.g., renewable or
fluence of selection on community patterns non-renewable), and the presence and nature
include species’ responses to the abiotic envi- of indirect interactions among species (Put-
ronment, the disturbance regime, the types of man 1994; Morin 1999; Ricklefs and Miller
direct interactions between organisms (e.g., 1999). The following handful of examples
competition, predation, parasitism, herbivory, focuses mostly on competition and trophic
mutualism), the functional or behavioral re- interactions to illustrate the basic types of
sponses of organisms to different densities of selection (Figure 1) and the range of out-
interacting species, the degree of specialization comes of selection in local communities,
in interspecific interactions, the number and such as the exclusion of some species by oth-
June 2010 SYNTHESIS IN COMMUNITY ECOLOGY 195
ers, the stable coexistence of species, com- superior competitor for the resource most
plex fluctuations in abundance over time, or limiting to the other species (Si) and vice
alternation between different stable states. I versa (Cyclotella meneghiniana, and P), result-
focus on studies in which the outcome of ing in stable coexistence via negative density-
selective processes is measured as changes in dependent selection at intermediate Si/P ra-
the abundances or diversity of species, rather tios. Coexistence among grassland plant
than in the responses, such as individual species via the same mechanism has been
growth rates or body size, of focal species, found in field experiments as well (Tilman
which are often measured under the as- 1988). A trade-off between competitive abil-
sumption that they may have consequences ity and colonization ability can create nega-
at the community level (e.g., Van Zandt and tive density-dependent selection contribut-
Agrawal 2004). ing to the coexistence of protozoans and
Species can exclude each other; while rotifers in lab microcosms (Cadotte et al.
selection in any real situation is unlikely to 2006), although field evidence for this mech-
be constant across all species’ densities anism is more ambiguous (e.g., Levine and
(Figure 1A), one species may be at an ad- Rees 2002). Temporally variable selection via
vantage across the full range of possible den- environmental fluctuations can lead to sta-
sities. Lab experiments have demonstrated ble coexistence of diatoms under variable
competitive exclusion between species of temperatures in the lab (Descamps-Julien
paramecium (Gause 1934), phytoplankton and Gonzalez 2005), and also appears to be
(Tilman 1977), and flour beetles (Park a likely explanation for the coexistence and
1948), among many others, as well as the fluctuation of grassland plants in variable cli-
exclusion of prey species by a predator matic conditions in Kansas (Adler et al. 2006).
(Gause 1934; Huffaker 1958). Similarly, field Patterns of species composition are often
experiments have revealed competitive ex- closely correlated with environmental condi-
clusion—for example, between barnacle spe- tions, with spatially-variable selection almost
cies at particular tidal depths (Connell certainly playing an important role (Whittaker
1961), or plant species under particular re- 1975).
source conditions (Tilman 1982, 1988). In Trophic interactions among species can
many cases of competitive exclusion, the win- lead to coexistence with fluctuations over time
ner in competition depends on environmen- via complex forms of density-dependent selec-
tal conditions, thus establishing the possibil- tion. Predators and their prey can coexist over
ity (in lab experiments) or existence (in field the long term with regular cycles, both in lab
studies) of spatially-variable selection. Many microcosms, such as those with different spe-
species distribution patterns have been inter- cies of mites (Huffaker 1958) or rotifers and
preted as evidence of competitive exclusion algae (Fussman et al. 2000), as well as in
between functionally similar species (e.g., Di- field populations, such as snowshoe hares
amond 1975), although it is very difficult to and lynx in the boreal forest (Krebs et al.
confidently infer process from pattern in 2001). Density-dependent species interac-
such cases (Strong et al. 1984). Regardless tions can also lead to complex, chaotic dy-
of the strength of direct interspecific compe- namics with species persistence in aquatic
tition, past environmental change (e.g., laboratory food webs (Benincà et al. 2008).
glacial cycles) has acted as an agent of selec- Positive and negative density-dependent
tion among species, favoring some but caus- selection over different ranges of species’
ing others to decline, sometimes to the densities can lead to switches between mul-
point of extinction (McKinney 1997; Wil- tiple states with respect to community com-
liams et al. 2004). position. Changes in the initial abundances
Competing species often exist in stable com- of species in aquatic microcosms can lead to
binations via negative density-dependent selec- very different and seemingly stable final spe-
tion. With multiple limiting resources and two cies compositions (Drake 1991)—a result
phytoplankton species, Tilman (1977) found that has been found in a variety of lab and
that one species (Asterionella formosa) was a field experiments (Schröder et al. 2005). In
196 THE QUARTERLY REVIEW OF BIOLOGY Volume 85
where relatively low soil pH has predomi- tial source habitats can also influence species
nated, the opposite is true (Pärtel 2002). composition, with distant localities contain-
Even if different habitats have been equally ing a preponderance of good dispersers
available over time, current species richness (e.g., Kadmon and Pulliam 1993). With vari-
might be greatest in conditions under which ation in local species composition created by
a particular group of organisms initially dispersal, either related to locality isolation
evolved and, therefore, where diversity has or stochastic variation, density-dependent se-
had more time to accumulate via speciation. lection via species interactions can further
For example, Wiens et al. (2007) found sim- magnify local variation in species composi-
ilar rates of diversification at different eleva- tion, as in freshwater communities of zoo-
tions for a clade of tropical salamanders, but plankton and their insect or fish predators
they noted that mid-elevation habitats were (Shurin 2001).
colonized earliest in the evolution of the Dispersal can interact with selection or
clade, thus helping explain a mid-elevation drift to influence community patterns at
peak in species richness patterns. In sum- the regional scale as well as at the local
mary, although selection is a key determi- scale. In an experimental metacommunity
nant of compositional change along gradi- of protozoans and rotifers, local species
ents, speciation is likely a critical process richness was maximal at intermediate rates
contributing to many diversity-environment of dispersal (Cadotte 2006a). The shift
relationships (Ricklefs 2004). from low to moderate dispersal increases
dispersal the rate of addition of new species to local-
ities and allows competitively inferior spe-
Dispersal can have manifold consequences cies to find temporary refuges, whereas the
for community patterns at multiple spatial shift from moderate to high dispersal al-
scales. First, much like speciation, dispersal lows superior competitors to dominate
is a key contributor to the regional species across the metacommunity. In the same ex-
pool and, consequently, the various com- periment, compositional variability among
munity consequences that it entails (Rick-
localities was maintained to the greatest de-
lefs and Schluter 1993).
gree with low to moderate dispersal, thus
From the local habitat perspective, a
maximizing richness across the entire meta-
common empirical result is that increasing dis-
community (Cadotte 2006a). A meta-analysis
persal into the locality increases species diver-
sity. For areas undergoing primary succession, of similar experiments found that local diver-
such as Krakatau following its volcanic erup- sity was generally maximized at intermediate
tion, dispersal is required in order to establish dispersal rates in animal communities, but,
a community and increase diversity (Whittaker at the highest dispersal rates in plants, there
et al. 1989). In the field, experimental dispersal was either a negative effect on regional di-
via seed addition into established plant com- versity or no effect was observed at all (Ca-
munities often results in increased species dotte 2006b). For the pond amphibians
diversity (Turnbull et al. 2000), and the prox- mentioned above, species turnover was
imity of an island or habitat patch to potential strongly influenced by both connectivity and
sources of dispersers often correlates positively environmental factors, thus suggesting an
with local species diversity in a wide range of important interaction between dispersal and
organisms (e.g., MacArthur and Wilson 1967). selection (Werner et al. 2007a,b).
For amphibians in a network of ponds, for The body of research on the community
instance, increased connectivity positively influ- consequences of dispersal and its interac-
enced species turnover, suggesting that dis- tion with selection is still comparatively
persal can affect not only species composition small. As with selection, it seems likely that any
and diversity, but their temporal rates of theoretically plausible effect of dispersal on
change as well (Werner et al. 2007a). community dynamics will be found in some
Since species vary in their propensity for experimental or natural community, while at
dispersal, the proximity of a locality to poten- the same time, many hypotheses concerning
June 2010 SYNTHESIS IN COMMUNITY ECOLOGY 199
perspective, although metacommunity mod- does not matter whether selection is the
els fit comfortably within it. best term for deterministic fitness differ-
In addition to these three conceptual ences among species, but it is critical to
frameworks specifically focused on community recognize that different mechanisms un-
ecology, two highly influential theoretical derlying selection, such as competition or
frameworks that cut across sub-fields of biol- predation, share more in common with
ogy, but with some connections to community one another than either does with drift,
ecology, are worth mentioning: the metabolic dispersal, or speciation.
theory of ecology (Brown et al. 2004) and eco- Recognizing how different theoretical
logical stoichiometry (Sterner and Elser 2002). traditions in community ecology relate to
In simple terms, these two approaches explore one another based on fundamental, logi-
the consequences of considering organisms es- cally distinct categories of process can po-
sentially as physical entities that process energy tentially prevent students from concluding
optimally given their size (metabolic theory), from a Web of Science search that research
or that interact with their environment based on species sorting or metacommunities
largely on their chemical composition (stoichi- goes back no more than 15–20 years. We
ometry). In my opinion, these frameworks might also make more modest—and, I be-
are most powerful in aiding our under- lieve, realistic—assessments of the degree
standing of the functioning of individual to which popular areas of research truly
organisms or the fluxes of energy and represent new paradigms or, more likely,
chemicals in whole ecosystems. The contri- incremental advances on previous work.
butions of these frameworks to community Placing ecological ideas in their full histor-
ecology, such as predictions concerning the ical context can curtail wheel reinvention
effect of temperature on species diversity (met- and thus help to advance and expand eco-
abolic theory) or of plant-herbivore interac- logical understanding in the long term
tions (stoichiometry), fall comfortably within (Graham and Dayton 2002).
the present framework, usually as mechanisms My hope is that the present framework will
underlying selection. be useful to practicing community ecologists
as a way to place their research in a process-
implications based context. I also think that this concep-
The first goal of this paper was to orga- tual framework can potentially be of great
nize the material of community ecology in use in teaching and communicating the sub-
a logically consistent way in order to clarify ject matter of community ecology to a
the similarities and differences among var- broader audience. As argued in the intro-
ious conceptual constructs in the disci- duction, the traditional presentation of com-
pline. One motivation was the common munity ecology can be confusing because
criticism that ecologists tend to repeatedly the common threads among topics such as
reinvent the wheel: we claim ideas as new food webs, competitive coexistence, and is-
that are only subtly distinct, or not distinct land biogeography are quite difficult to dis-
at all, from ideas put forth long ago (Law- cern. The essential similarities and differ-
ton 1991; Graham and Dayton 2002; Be- ences among these theoretical traditions can
lovsky et al. 2004). There are likely many be seen quite clearly in the present frame-
reasons for this, but one important reason, work (Table 2). The core subject matter in
at least in community ecology, may be the community ecology need not change, but I
lack of a coherent framework within which believe there can be great benefit to shifting
particular perspectives or theories can be the emphasis away from an organizational
described and related. As such, a plethora structure based on the useful lines of inquiry
of terms, each of which sounds new and carved out by researchers, to one based on
different, is often used to communicate the fundamental processes that underlie
much the same thing—such as niche pro- community dynamics and patterns.
cesses, species interactions, or species sort- The second goal of this paper was to artic-
ing all being used to describe selection. It ulate a general theory of community ecol-
202 THE QUARTERLY REVIEW OF BIOLOGY Volume 85
ogy. Such a theory might seem so general- important rule of heredity is decidedly facile:
ized as to be of little use, but the utility of the elephants give rise to elephants and daffodils
Modern Synthesis in evolutionary biology— to daffodils. However, on its own, the Mod-
warts and all (Pigliucci 2007)—suggests oth- ern Synthesis makes no predictions about
erwise. In essence, the Modern Synthesis can exactly how processes will interact to deter-
be summarized as positing that genetic vari- mine evolutionary change in any particular
ation is created in populations via mutation situation; rather, it simply establishes the fun-
and immigration, and is then shaped by drift damental set of processes that may be at
and natural selection to drive evolutionary work.
change (Kutschera and Niklas 2004). The We can likewise articulate a very general
fact that the all-important mechanism of he- theory of community ecology: species are
redity was essentially unknown until the re- added to communities via speciation and
discovery of Mendel made the construction dispersal, and the relative abundances of
of the Modern Synthesis a profound scien- these species are then shaped by drift and
tific achievement in a way that cannot be selection, as well as ongoing dispersal, to
matched in community ecology, where the drive community dynamics (Figure 4). The
June 2010 SYNTHESIS IN COMMUNITY ECOLOGY 203
precise way in which these processes inter- to make general predictions about how par-
act to determine community dynamics var- ticular processes have shaped real ecologi-
ies tremendously from case to case, just as cal communities. If the goal is to make
the processes that determine evolutionary general statements about the fundamental
change interact in ways that vary tremen- processes that can underlie community dy-
dously in each case. Stating a general theory namics and the possible ways in which
of community ecology in this way echoes the these can interact, then community ecol-
perspective of Ricklefs and Schluter (1993), ogy appears to be in excellent shape.
and I believe that recognizing this perspec-
tive as the community ecology counterpart acknowledgments
to the evolutionary Modern Synthesis high- Valuable feedback and insights on the topic of this
lights an important sense in which commu- manuscript or on the manuscript itself were provided by
nity ecology already has a general theoretical J. Goheen, C. Harley, R. Holt, J. Levine, J. Losos, J.
framework that is every bit as robust as that Orrock, R. Ricklefs, J. Roughgarden, J. Shurin, D. Sriv-
of population genetics. The oft-cited recalci- astava, M. Whitlock, A. Agrawal, J. Wiens, the UBC Com-
trance of community ecology to generally munity Ecology class of 2008, and two anonymous re-
applicable theory (e.g., Lawton 1999) seems viewers. This work was supported by the Natural
like a fair assessment if the goal is to be able Sciences and Engineering Research Council, Canada.
REFERENCES
Aarssen L. W., Schamp B. S. 2002. Predicting distri- Cadotte M. W. 2006b. Dispersal and species diversity: a
butions of species richness and species size in re- meta-analysis. American Naturalist 167(6):913–924.
gional floras: applying the species pool hypothesis Cadotte M. W., Mai D. V., Jantz S., Collins M. D., Keele
to the habitat templet model. Perspectives in Plant M., Drake J. A. 2006. On testing the competition-
Ecology, Evolution and Systematics 5:3–12. colonization tradeoff in a multispecies assemblage.
Adler P. B., HilleRisLambers J., Kyriakidis P. C., Guan American Naturalist 168(5):704 –709.
Q., Levine J. M. 2006. Climate variability has a Caswell H. 1978. Predator mediated coexistence: a
stabilizing effect on coexistence of prairie grasses. nonequilibrium model. American Naturalist 112:
Proceedings of the National Academy of Sciences USA 127–154.
103(34):12793–12798. Chase J. M., Leibold M. A. 2003. Ecological Niches:
Adler P. B., HilleRisLambers J., Levine J. M. 2007. A Linking Classical and Contemporary Approaches. Chi-
niche for neutrality. Ecology Letters 10:95–104. cago (IL): University of Chicago Press.
Allen T. F. H., Hoekstra T. W. 1992. Toward a Unified Chesson P. 1978. Predator-prey theory and variability.
Annual Review of Ecology and Systematics 9:323–347.
Ecology. New York: Columbia University Press.
Chesson P. 2000. Mechanisms of maintenance of spe-
Bell G. 2008. Selection: The Mechanism of Evolution. Ox-
cies diversity. Annual Review of Ecology and System-
ford (UK): Oxford University Press.
atics 31:343–366.
Belovsky G. E., Botkin D. B., Crowl T. A., Cummins
Chesson P. L., Warner R. R. 1981. Environmental vari-
K. W., Franklin J. F., Hunter M. L., Jr., Joern A.,
ability promotes species coexistence in lottery com-
Lindenmayer D. B., MacMahon J. A., Margules C. R.,
petitive systems. American Naturalist 117:923–943.
Scott J. M. 2004. Ten suggestions to strengthen the
Connell J. H. 1961. The influence of interspecific compe-
science of ecology. BioScience 54(4):345–351. tition and other factors on the distribution of the bar-
Benincà E., Huisman J., Heerkloss R., Jöhnk K. D., nacle Chthamalus stellatus. Ecology 42:710–723.
Branco P., Van Nes E. H., Scheffer M., Ellner S. P. Cooper G. J. 2003. The Science of the Struggle for Existence:
2008. Chaos in a long-term experiment with a On the Foundations of Ecology. Cambridge (UK): Cam-
plankton community. Nature 451(7180):822– 825. bridge University Press.
Brown J. H. 1995. Macroecology. Chicago (IL): Chicago Cottenie K. 2005. Integrating environmental and spa-
University Press. tial processes in ecological community dynamics.
Brown J. H., Gillooly J. F., Allen A. P., Savage V. M., Ecology Letters 8:1175–1182.
West G. B. 2004. Toward a metabolic theory of Darwin C. 1859. On the Origin of Species by Means of
ecology. Ecology 85(7):1771–1789. Natural Selection, or the Preservation of Favoured Races
Cadotte M. W. 2006a. Metacommunity influences on in the Struggle for Life. London (UK): John Murray.
community richness at multiple spatial scales: a Dent C. L., Cumming G. S., Carpenter S. R. 2002.
microcosm experiment. Ecology 87(4):1008 –1016. Multiple states in river and lake ecosystems. Philo-
204 THE QUARTERLY REVIEW OF BIOLOGY Volume 85
sophical Transactions of the Royal Society of London, John R., Dalling J. W., Harms K. E., Yavitt J. B., Stal-
Series B 357(1421):635– 645. lard R. F., Mirabello M., Hubbell S. P., Valencia R.,
Descamps-Julien B., Gonzalez A. 2005. Stable coexist- Navarrete H., Vallejo M., Foster R. B. 2007. Soil
ence in a fluctuating environment: an experimen- nutrients influence spatial distributions of tropical
tal demonstration. Ecology 86(10):2815–2824. tree species. Proceedings of the National Academy of
Diamond J. M. 1975. Assembly of species communi- Sciences USA 104(2):864 – 869.
ties. Pages 342– 444 in Ecology and Evolution of Com- Kadmon R., Pulliam H. R. 1993. Island biogeography:
munities, edited by M. L. Cody and J. M. Diamond. effect of geographical isolation on species compo-
Cambridge (MA): Harvard University Press. sition. Ecology 74(4):977–981.
Diamond J. M., Case T. J. 1986. Community Ecology. Kingsland S. E. 1995. Modeling Nature: Episodes in the
New York: Harper and Row. History of Population Ecology. Second Edition. Chi-
Drake J. A. 1991. Community-assembly mechanics and cago (IL): University of Chicago Press.
the structure of an experimental species ensemble. Krebs C. J., Boonstra R., Boutin S., Sinclair A. R. E.
American Naturalist 137(1):1–26. 2001. What drives the 10-years cycle of snowshoe
Fauth J. E., Resetarits W. J., Jr., Wilbur H. M. 1990. hares? BioScience 51(1):25–35.
Interactions between larval salamanders: a case of Kutschera U., Niklas K. J. 2004. The modern theory of
competitive equality. Oikos 58:91–99. biological evolution: an expanded synthesis. Natur-
Fussmann G. F., Ellner S. P., Shertzer K. W., Hairston wissenschaften 91:255–276.
N. G., Jr. 2000. Crossing the Hopf bifurcation in a Lawton J. H. 1991. Warbling in different ways. Oikos
live predator-prey system. Science 290(5495):1358 – 60:273–274.
1360. Lawton J. H. 1999. Are there general laws in ecology?
Gause G. F. 1934. The Struggle for Existence. Baltimore Oikos 84(2):177–192.
(MD): Williams and Wilkins Company.
Lerner I. M., Dempster E. R. 1962. Indeterminism in
Gilpin M. E., Diamond J. M. 1976. Calculation of immigra-
interspecific competition. Proceedings of the National
tion and extinction curves from the species-area-
Academy of Sciences USA 48(5):821– 826.
distance relation. Proceedings of the National Academy of
Levene H. 1953. Genetic equilibrium when more
Sciences USA 73(11):4130–4314.
than one ecological niche is available. American
Goldberg D. E., Werner P. A. 1983. Equivalence of
Naturalist 87:331–333.
competitors in plant communities: a null hypoth-
Levine J. M., Rees M. 2002. Coexistence and relative
esis and a field experimental approach. American
abundance in annual plant communities: the roles
Journal of Botany 70:1098 –1104.
of competition and colonization. American Naturalist
Graham M. H., Dayton P. K. 2002. On the evolution of
160:452– 467.
ecological ideas: paradigms and scientific progress.
Lewontin R. 2004. The problems of population ge-
Ecology 83:1481–1489.
Grime J. P. 1973. Competitive exclusion in herba- netics. Pages 5–23 in Evolutionary Genetics: from Mol-
ceous vegetation. Nature 242:344 –347. ecules to Morphology, edited by R. S. Singh and C. B.
Hartl D. L., Clark A. G. 1997. Principles of Population Krimbas. Cambridge (UK): Cambridge University
Genetics. Third Edition. Sunderland (MA): Sinauer Press.
Associates. Loreau M., Hector A. 2001. Partitioning selection and
Holyoak M., Leibold M. A., Holt R. D. 2005. Metacom- complementarity in biodiversity experiments. Na-
munities: Spatial Dynamics and Ecological Communi- ture 412(6842):72–76.
ties. Chicago (IL): University of Chicago Press. Losos J. B., Schluter D. 2000. Analysis of an evolutionary
Hubbell S. P. 2001. The Unified Neutral Theory of Bioge- species-area relationship. Nature 408(6814):847–850.
ography and Biodiversity. Princeton (NJ): Princeton Lotka A. J. 1925. Elements of Physical Biology. Baltimore
University Press. (MD): Williams and Wilkins.
Hubbell S. P. 2005. Neutral theory in community MacArthur R. H. 1969. Patterns of communities in
ecology and the hypothesis of functional equiva- the tropics. Biological Journal of the Linnean Society
lence. Functional Ecology 19:166 –172. 1(1–2):19 –30.
Huffaker C. B. 1958. Experimental studies on preda- MacArthur R. H., Wilson E. O. 1967. The Theory of
tion: dispersion factors and predator-prey oscilla- Island Biogeography. Princeton (NJ): Princeton Uni-
tions. Hilgardia 27:343–383. versity Press.
Hughes A. R., Inouye B. D., Johnson M. T. J., Under- McGill B. J., Enquist B. J., Weiher E., Westoby M.
wood N., Vellend M. 2008. Ecological consequences 2006. Rebuilding community ecology from func-
of genetic diversity. Ecology Letters 11(6):609 – 623. tional traits. Trends in Ecology and Evolution 21(4):
Hu X.-S., He F., Hubbell S. P. 2006. Neutral theory in 178 –185.
macroecology and population genetics. Oikos 113(3): McKinney M. L. 1997. Extinction vulnerability and
548–556. selectivity: combining ecological and paleontolog-
June 2010 SYNTHESIS IN COMMUNITY ECOLOGY 205
ical views. Annual Review of Ecology and Systematics Ricklefs R. E., Miller G. L. 1999. Ecology. Fourth Edi-
28:495–516. tion. New York: W.H. Freeman.
McPeek M. A. 2007. The macroevolutionary conse- Ricklefs R. E., Qian H., White P. S. 2004. The region
quences of ecological differences among species. effect on mesoscale plant species richness between
Paleontology 50:111–129. eastern Asia and eastern North America. Ecography
McPeek M. A., Gomulkiewicz R. 2005. Assembling 27:129 –136.
and depleting species richness in metacommuni- Ricklefs R. E., Schluter D. 1993. Species Diversity in
ties: insights from ecology, population genetics Ecological Communities: Historical and Geographical
and macroevolution. Pages 355–373 in Metacom- Perspectives. Chicago (IL): University of Chicago
munities: Spatial Dynamics and Ecological Communi- Press.
ties, edited by M. Holyoak et al. Chicago (IL): Rosenzweig M. L. 1995. Species Diversity in Space and Time.
University of Chicago Press. Cambridge (UK): Cambridge University Press.
Mertz D. B., Cawthon D. A., Park T. 1976. An exper- Roughgarden J. 2009. Is there a general theory of
imental analysis of competitive indeterminacy in community ecology? Biology and Philosophy 24(4):
Tribolium. Proceedings of the National Academy of Sci- 521–529.
ences USA 73(4):1368 –1372. Scheffer M., Carpenter S., Foley J. A., Folke C., Walker
Morin P. J. 1999. Community Ecology. Oxford (UK) and B. 2001. Catastrophic shifts in ecosystems. Nature
Malden (MA): Blackwell. 413:591–596.
Norberg J., Swaney D. P., Dushoff J., Lin J., Casagrandi R., Schröder A., Persson L., De Roos A. M. 2005. Direct
Levin S. A. 2001. Phenotypic diversity and ecosystem experimental evidence for alternative stable states:
functioning in changing environments: a theoretical a review. Oikos 110(1):3–19.
framework. Proceedings of the National Academy of Sciences Shipley B., Vile D., Garnier E. 2006. From plant traits
USA 98(20):11376–11381. to plant communities: a statistical mechanistic ap-
Nowak M. A. 2006. Evolutionary Dynamics: Exploring the proach to biodiversity. Science 314(5800):812– 814.
Equations of Life. Cambridge (MA): Belknap Press Shurin J. B. 2001. Interactive effects of predation and
of Harvard University Press. dispersal on zooplankton communities. Ecology 82:
Palmer M. W. 1994. Variation in species richness: 3404 –3416.
towards a unification of hypotheses. Folia Geobo- Siepielski A. M., Hung K.-L., Bein E. E., McPeek M. A.
tanica et Phytotaxonomica 29:511–530. 2010. Experimental evidence for neutral commu-
Park T. 1948. Experimental studies of interspecies nity dynamics governing and insect assemblage.
competition. I. Competition between populations Ecology 91:847– 857.
of the flour beetles, Tribolium confusum Duval and Simberloff D. 2004. Community ecology: is it time to
Tribolium castaneum Herbst. Ecological Monographs move on? American Naturalist 163:787–799.
18(2):265–308. Srivastava D. S. 1999. Using local-regional richness
Pärtel M. 2002. Local plant diversity patterns and plots to test for species saturation: pitfalls and
evolutionary history at the regional scale. Ecology potentials. Journal of Animal Ecology 68:1–17.
83(9):2361–2366. Sterner R. W., Elser J. J., Vitousek P. 2002. Ecological
Pigliucci M. 2007. Do we need an extended evolution- Stoichiometry: The Biology of Elements from Molecules to the
ary synthesis? Evolution 61(2):2743–2749. Biosphere. Princeton (NJ): Princeton University Press.
Provine W. B. 1971. The Origins of Theoretical Population Strauss S. Y. 1991. Indirect effects in community ecol-
Genetics. Chicago (IL): University of Chicago Press. ogy: their definition, study and importance. Trends
Putman R. J. 1994. Community Ecology. New York: Chap- in Ecology and Evolution 6(7):206 –210.
man and Hall. Strong D. R., Simberloff D., Abele L. G., Thistle A. B.
Qian H., Ricklefs R. E. 2000. Large-scale processes 1984. Ecological Communities: Conceptual Issues and the
and the Asian bias in species diversity of temperate Evidence. Princeton (NJ): Princeton University Press.
plants. Nature 407(6801):180 –182. Taylor D. R., Aarssen L. W., Loehle C. 1990. On the
Rice S. H. 2008. A stochastic version of the Price relationship between r/K selection and environ-
equation reveals the interplay of deterministic and mental carrying capacity: a new habitat templet
stochastic processes in evolution. BMC Evolution- for plant life history strategies. Oikos 58:239 –250.
ary Biology 8:262. Tilman D. 1977. Resource competition between plank-
Ricklefs R. E. 1987. Community diversity: relative roles of ton algae: an experimental and theoretical ap-
local and regional processes. Science 235(4785):167– proach. Ecology 58(2):338 –348.
171. Tilman D. 1982. Resource Competition and Community
Ricklefs R. E. 2004. A comprehensive framework for Structure. Princeton (NJ): Princeton University Press.
global patterns in biodiversity. Ecology Letters 7:1–15. Tilman D. 1988. Plant Strategies and the Dynamics and
Ricklefs R. E. 2008. Disintegration of the ecological Structure of Plant Communities. Princeton (NJ):
community. American Naturalist 172:741–750. Princeton University Press.
206 THE QUARTERLY REVIEW OF BIOLOGY Volume 85
Tilman D. 1994. Competition and biodiversity in spa- nity: the role of local and regional factors. Oikos
tially structured habitats. Ecology 75(1):2–16. 116:1713–1725.
Tilman D. 2004. Niche tradeoffs, neutrality, and com- Whittaker R. H. 1975. Communities and Ecosystems. New
munity structure: a stochastic theory of resource York: MacMillan Company.
competition, invasion, and community assembly. Whittaker R. J., Bush M. B., Richards K. 1989. Plant
Proceedings of the National Academy of Sciences USA recolonization and vegetation succession on the
101(30):10854 –10861. Krakatau Islands, Indonesia. Ecological Monographs
Turnbull L. A., Crawley M., Rees M. 2000. Are plant 59(2):59 –123.
populations seed-limited? A review of seed sowing Whittaker R. J., Fernandez-Palacios J. M. 2007. Island
experiments. Oikos 88(2):225–238. Biogeography: Ecology, Evolution and Conservation.
Van Zandt P. A., Agrawal A. A. 2004. Specificity of induced Second Edition. Oxford (UK): Oxford University
plant responses to specialist herbivores of the common Press.
milkweed Asclepias syriaca. Oikos 104(2):401–409. Wiens J. J., Parra-Olea G., Garcı́a-Parı́s M., Wake D. B.
Vellend M., Geber M. A. 2005. Connections between 2007. Phylogenetic history underlies elevational
species diversity and genetic diversity. Ecology Let-
biodiversity patterns in tropical salamanders. Pro-
ters 8:767–781.
ceedings of the Royal Society of London, Series B 274:
Vellend M., Orrock J. L. 2009. Genetic and ecological
919 –928.
models of diversity: lessons across disciplines. Pages
Williams J. W., Shuman B. N., Webb T, III, Bartlein
439 – 461 in The Theory of Island Biogeography Revisited,
P. J., Leduc P. L. 2004. Late Quaternary vegetation
edited by J. B. Losos and R. E. Ricklefs. Princeton
dynamics in North America: scaling from taxa to
(NJ): Princeton University Press.
biomes. Ecological Monographs 74(2):309 –334.
Werner E. E., Skelly D. K., Relyea R. A., Yurewicz K. L.
2007b. Amphibian species richness across environ- Wright S. 1940. Breeding structure of populations in
mental gradients. Oikos 116(10):1697–1712. relation to speciation. American Naturalist 74:232–
Werner E. E., Yurewicz K. L., Skelly D. K., Relyea R. A. 248.
2007a. Turnover in an amphibian metacommu- Associate Editor: Anurag Agrawal