Professional Documents
Culture Documents
3
Nathan J. B. Kraft and David D. Ackerly
Contents
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68
A Brief History of the Development of Community Assembly Concepts . . . . . . . . . . . . . . . . . . . . . 68
Dispersal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71
Abiotic Filtering . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
Biotic Interactions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
Relationship Between Community Assembly and Coexistence Theory . . . . . . . . . . . . . . . . . . . . . . . 76
Phylogenetic Patterns . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77
Biogeography and the Build Up of Species Pools . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80
Scale Dependence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83
Future Directions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84
Pattern-to-Process Mapping . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84
Coexistence Theory and Community Assembly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85
Methods for Multitrophic Interactions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85
Abstract
• Communities are located within a larger species pool of potential colonists.
The study of community assembly considers the mechanisms by which local
communities are formed from the species pool.
• Dispersal from the species pool, abiotic tolerance of colonists, and biotic
interactions can all influence membership in local communities.
• Phenotypic similarities and differences of co-occurring species can be used
(within limits) to make inferences about the role of alternative processes
contributing to community assembly.
• In many plant groups, close relatives tend to share similar phenotypic traits.
Therefore, patterns of phylogenetic relatedness within a community can also
be used to make inferences about community assembly mechanisms.
• As the community and the species pool can be defined at a number of
different spatial and temporal scales, community assembly patterns often
show strong scale dependence. In some cases, a single process can produce
contrasting phenotypic patterns at different scales of analysis, while in other
cases different processes may have stronger influences on community assem-
bly at different scales.
• Species pools are shaped by dispersal of lineages among biogeographic
regions, in situ speciation within regions, and extinction. The characteristics
of the species pool often persist in local community patterns.
• Community assembly studies are often limited in the extent to which specific
mechanisms can be inferred from community pattern. Future work should
focus on improved models of competition and coexistence dynamics in
community assembly as well as methods for considering multitrophic
interactions.
Introduction
Community assembly is the study of the processes that shape the identity and
abundance of species within ecological communities. Central to most studies of
community assembly is the concept of a species pool that is larger in geographic
scope than the local community under study. The species pool contains potential
colonists of the community, and many studies in this area focus on developing an
understanding of the role of dispersal, responses to abiotic conditions, and biotic
interactions in shaping local assemblages. Thus, community assembly considers
both the ecological interactions that shape local communities and the evolutionary
and biogeographic processes that lead to variation in the diversity and composition
of species pools across the globe.
There are two persistent and central concepts in the study of community assembly.
The first is the “species pool,” defined as the suite of possible colonists for a
local site under study, and the second is the metaphor of a “filter” or a “sieve”
that represents abiotic or biotic barriers to successful establishment at a local
site. The two concepts can be traced back to two distinct sources: the study of
species assemblages on oceanic islands and the study of succession following
disturbance.
Perhaps the best-known precursor to community assembly theory is MacArthur
and Wilson’s seminal theory of island biogeography, which describes the fate of an
3 Assembly of Plant Communities 69
Fig. 1 Basic conceptual model of community assembly in terms of species functional traits
(phenotypes). Empirically, an ecologist can consider the local community in relation to the species
pool of potential colonists. Habitat filtering is often hypothesized to limit the range of traits that
can successfully survive and establish at a site, as well as sometimes shifting the mean value
relative to the species pool. Competition, in its earliest forms in community assembly theory, was
predicted to favor the coexistence of species that differed in resource use or requirements, reflected
here in their functional traits. In the example here, competitive exclusion leads to a local
community of species with trait values that are more dissimilar than species in the original species
pool. Additional community assembly processes are not shown. (After Woodward and Diament
(1991))
Dispersal
Dispersal refers to the movement of an individual organism during its lifetime, from
its place of birth to the location where it produces offspring. As plants are sessile
organisms, in most species, dispersal only occurs once during the life cycle at the
seed stage. Once a plant germinates, it occupies a single location for the rest of its
life. In addition to this mechanism, a small number of species are able to disperse via
72 N.J.B. Kraft and D.D. Ackerly
Abiotic Filtering
the seed bank (and testing for seed viability) may be required to definitively
conclude that a species is absent from a site. This is most relevant in communities
that exhibit substantial variation in abiotic conditions over time, as different species
may use the same habitat at different times of the year or in different years,
depending on year-to-year variation in weather, remaining dormant in the seed
bank at other times.
Abiotic factors can also cause mortality or prevent successful reproduction at
any time during the life cycle from germination through reproductive maturity.
Species vary in requirements for light, nutrients, and water as well as in tolerance to
drought and temperature, and any of these factors can cause mortality at any stage.
An important consideration is that brief, extreme climatic events can have strong
impacts on species survival. For example, the average climatic conditions at a site
may be ideal for the growth and reproduction of a species, but a brief period of
extreme cold or heat or a short but severe drought that occurs infrequently can cause
significant mortality and effectively remove particular species from a site. For
example, a severe drought associated with an El Niño event in the 1980s is thought
to have had persistent and long-lasting impacts on the species composition, and
associated functional traits, of a tropical forest on Barro Colorado Island, Panama
(Feeley et al. 2011). Therefore, in considering the role of abiotic conditions in
filtering species from a site, it may be just as important to consider the variance or
the extremes of abiotic conditions as it is to consider the average values.
Practically speaking, it can be challenging to distinguish between habitat filter-
ing and dispersal limitation when a species is completely absent from a site.
Simple experiments can be helpful in testing for habitat filtering. On the most
basic level, these experiments involve transplanting individuals either as adults or
as seeds to the site and monitoring germination and/or survival. In situations where
these experiments are impractical, seed traps or detailed examination of the seed
bank can be useful in ruling out dispersal limitation as the cause of a species
absence.
An important consequence of abiotic filtering is that species composition typi-
cally changes along environmental gradients. For example, there is widespread
evidence that plant communities change in predictable ways along gradients of
light, water availability, soil fertility, elevation and latitude, among other factors.
These changes in species identity are also often reflected in changes in the func-
tional traits of species, such that average trait values across species in the commu-
nity can shift along a gradient. For example, woody plant leaf functional traits
change consistently across a gradient of soil water availability in coastal California
and across microtopographic gradients in the Ecuadorian Amazon (Kraft
et al. 2008; Cornwell and Ackerly 2009). Another frequently documented pattern
is that the breadth or variance of strategies seen at any point along the gradient is
often smaller than is seen across the gradient as a whole. The significance of these
observations – i.e., shifts in the mean of trait values and reduction in the range or
variance in trait values at points along a gradient – is typically documented using a
null model approach, comparing observed communities to hypothetical communi-
ties assembled at random from the regional species pool.
3 Assembly of Plant Communities 75
Biotic Interactions
trees of the same species should suffer more negative effects than seedlings growing
far from adults, as natural enemies can become concentrated near adult trees
(reviewed in Wright 2002). In this case, the study of plant distribution patterns within
communities can offer some insight into the role of natural enemies. However, in
other cases, we likely need improved conceptual models and approaches to effec-
tively incorporate natural enemies into community assembly studies.
Positive interactions between species can also have profound impacts on com-
munity assembly, allowing species to establish or persist at sites where they would
otherwise be unable to survive. Many of these associations are between plants and
other organisms. For example, associations with mycorrhizae and nitrogen-fixing
bacteria allow many plant species to gain access to essential nutrients more
effectively, and many species rely on insect or animal pollinators for reproduction.
The absence of these mutualist partners can effectively exclude plants from partic-
ular sites. Our understanding of these relationships in a community assembly
context is hampered by the same limitations as our understanding of natural
enemies – many studies typically focus just on plants, not on other groups within
a community. While it is possible to study some consequences of plant-pollinator
interactions primarily through the plant community (Sargent and Ackerly 2008),
new approaches will be needed to fully incorporate positive interactions that extend
beyond a single trophic or functional group into community assembly studies.
It is also well understood that plants can have positive effects on each other.
These impacts most commonly involve an amelioration of environmental stress
or a reduction in herbivore pressure via associational defenses. For example, in
hot and dry environments, some species are known to function as “nurse plants”
by modifying the nearby microclimate enough to allow other species to be able to
establish. However, many positive interactions between plants are known to be
highly context dependent. For example, in one globally replicated experimental
study, plants growing at lower elevations on mountains were often found to
compete with one another, while species growing at higher elevations on the
same mountains (which is presumably a more stressful environment) were found
to have positive effects on one another (Callaway et al. 2002). These findings
highlight that most positive interactions (and perhaps many species interactions
in general) typically include both a positive and a negative component and that
the relative importance of these components for community assembly can shift as
abiotic conditions change. This also highlights a general but understudied chal-
lenge within the topic of community assembly – disentangling the interactions
among abiotic and biotic filters.
Phylogenetic Patterns
In a famous quote from the Origin of Species (1859), Darwin noted that
“As the species of the same genus usually have. . .much similarity in habits
and constitution, . . . the struggle will generally be more severe between them, if
78 N.J.B. Kraft and D.D. Ackerly
they come into competition with each other, than between the species of distinct
genera.” His observation reflected the general knowledge of any experienced
systematist or field naturalist that related species tend to be ecologically similar;
e.g., one would expect two grass-eating rabbit species to compete directly for the
same food sources, whereas a grain-eating mouse and a carnivorous fox are
utilizing quite different resources. In the first half of the twentieth century, exper-
imental studies of competition by Gause and the development of Lotka-Volterra
competition theory led to the development of the competitive exclusion principle
(Hardin 1960), discussed earlier, which posits that species competing for the same
resources could not coexist in a community. Putting these ideas together, ecologists
in the mid-twentieth century suggested that species of the same genus would not
live together in local communities, at least not as often as one might expect if
communities were assembled randomly from the available species in a regional
species pool. This prediction was supported in studies of animal communities on
islands, compared to the fauna of adjacent mainland regions. These studies provide
some of the earliest examples of null models in ecology, discussed above.
Starting in the 1960s, the study of phylogenetics was revolutionized by concep-
tual, computational, and empirical advances, most notably the breakthroughs in
molecular biology leading to the modern era of DNA sequencing. With high-
resolution, well-supported phylogenies available, new methods have been devel-
oped to reexamine classical questions in ecology and evolutionary biology.
The study of plant communities presented particular challenges, as the deeper
structure of the angiosperm phylogeny had never been well understood and molec-
ular data brought a number of surprises. The first breakthroughs came in the 1990s,
quickly leading to a broad community effort under the Angiosperm Phylogeny
Group and a rapidly growing consensus about major patterns in flowering plant
phylogeny. Plant ecologists moved quickly to utilize the newly available phylog-
enies to tackle large-scale problems in adaptive evolution, diversity, and commu-
nity assembly (Webb 2000). Molecular data also provide branch lengths that
quantify the degree of relatedness among species, and fossil calibrations can be
applied to estimate branch lengths in millions of years since species diverged from
their most recent common ancestor. The phylogenetic distance between two species
is defined as the distance from one species down the phylogeny to the common
ancestor and back up to the other species (in other words, two times the age of their
most recent common ancestor).
The phylogenetic structure of a community can be described in a number of
ways, using quantitative metrics based on the phylogenetic relationships for the
community, “pruned” from the larger phylogeny of all plants. As in the examples
discussed above, statistical analyses of phylogenetic community structure consider
a local community relative to a null model of communities assembled from a
regional species pool. Two simple measures of phylogenetic community structure
are the mean phylogenetic distance, defined as the average of the phylogenetic
distances between all pairs of taxa in a community, and the mean nearest neighbor
distance, defined as the average distance from each species to its closest relative.
Using these measures, the net relatedness index (NRI) and nearest taxon index (NTI)
3 Assembly of Plant Communities 79
As we have discussed above, local communities are assembled from a regional pool
of available species. Thus, an understanding of the processes that shape the
diversity and the functional and phylogenetic composition of regional biota is
valuable for a deeper understanding of local communities. How big is the regional
species pool? This is a difficult, perhaps impossible, question to answer precisely as
the appropriate temporal and spatial scale defining the regional pool will depend on
the size of the communities and the dispersal biology and longevity of the organ-
isms under consideration. In many cases, the delineation of the regional pool for the
purposes of empirical studies is constrained practically by availability of data,
though the growth and refinement of regional and global biodiversity databases
(e.g., GBIF, the Global Biodiversity Information Facility) may eventually over-
come this obstacle.
The diversity of a regional flora and fauna reflects the interaction of three
fundamental processes: arrival of new lineages by dispersal from other regions,
speciation within the region, and local extinction of lineages. Dispersal can occur
across substantial distances. As described earlier, the colonization of oceanic
islands by seeds dispersed by wind or water currents or by animal agents provides
direct evidence of the potential for long-distance dispersal. Similar types of long-
distance dispersal events occur across and among continents as well, though they
are harder to detect. As climates shift and the combination of continental drift and
fluctuating sea levels have altered the connections between landmasses, dispersal
can also occur as a stepping-stone process with populations migrating along
corridors that provide favorable environments for at least short-term establishment
and subsequent reproduction. For example, in the Northern Hemisphere, there has
been extensive migration between North America and Europe across a North
Atlantic land bridge, during the Eocene and possibly into the Oligocene, while
more recently Asia and North America have been connected by the Bering land
bridge during periods of low sea level.
On evolutionary timescales, speciation is a key process increasing diversity
within biogeographic regions. During the speciation process, evolutionary shifts
in habitat affiliation and the climatic tolerances of a lineage tend to change slowly.
As a result, individual clades will tend to diversify and spread across major climatic
zones, and these biotic similarities then come to define distinctive biogeographic
regions around the world. Based on phylogenetic and fossil evidence, it is believed
that flowering plants originated in tropical regions and diversified extensively
during the Cretaceous. Around 55 million years ago during the Eocene, the world
was much warmer overall, and tropical forests extended to midlatitudes, far beyond
their current distribution. Cooling and drying trends since then have led to the
3 Assembly of Plant Communities 81
tolerances, so close relatives spread out across the landscape and occupy different
habitats. Improved knowledge of phylogenetic history, the fossil record, and
the climatic history of different regions of the world will continue to shed light
on these fundamental questions in the evolution of regional floras and their influ-
ence on the assembly of local communities.
Scale Dependence
Fig. 3 A single community assembly process (habitat filtering into specific microsites) can
produce contrasting phenotypic patterns depending on the scale of analysis. If a microsite is
compared to the species pool, either microsite will show low dispersion (phenotypic clustering).
However, if a large spatial scale is used to define the community that includes both microsites, this
new larger community sample across sites may reveal higher phenotypic dispersion than expected
when compared to the species pool
Future Directions
The study of community assembly theory has seen considerable development since it
was pioneered in the middle of the twentieth century. However, considerable chal-
lenges remain, and here we highlight three essential areas where additional work is
needed, including better methods to distinguish between multiple processes in pro-
ducing community patterns, a more complete incorporation of coexistence theory into
community assembly studies, and better consideration of multitrophic interactions.
Pattern-to-Process Mapping
Most community assembly analyses focus within a trophic level, and most plant-
focused studies do not explicitly consider other trophic levels except as those
interactions play out implicitly among plants (e.g., Sargent and Ackerly 2008).
However, given the ubiquity of trophic interactions in shaping community patterns,
community assembly will not be able fully consider the multitude of ecological
interactions shaping local communities until it is able to explicitly incorporate
trophic interactions into analyses. Given the rapid pace of development of network
theory in recent years, it may be that a robust solution to this issue will emerge from
an integration of these two approaches.
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