CHAPTER 4

THE PLANT COMMUNITY

A. Objectives
After studying this unit, students are expected to:
1
2

Explain scope of plant community

Explain the attributes of physiognomy concept, species composition,

and spatial patterns.

Explain the concept of species richness, evenness, and diversity

B. Introduction
In each type of habitat, certain species group together as a
community. Fossil records indicate that some of these groups (or very
closely related precursors) have lived together for thousands and even
millions of years. During that time, it is possible that an intricate
balance has been fashioned. Community members share incoming solar
radiation, soil water, and nutrients to produce a constant biomass; they
recycle nutrients from the soil to living tissue and back again; and they
alternate with each other in time and space. Synecologists attempt to
determine what is involved in this balance between all the species of a
community and their environment.
First, how is the community to be measured and how are the
measurements summarized for maximum, useful information? Second,
why do some cornrnunities change more rapidly over time than others?
How accurately can we predict future changes and reconstruct the past?
Are there community traits that transcend traits of the individual species
making up the community? A synecological view asks such questions,
and the answers take us to a level of complexity beyond autecology. The
term community is a very general one that can be applied to vege tation
types of any size or longevity. It can, for example, be applied to one
43

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stratum of plants in a very local area, such as the herbs, woody

seedlings, and mosses on the floor of a streambank forest; or to a very
widespread, regional vegetation type; or to a transitory plot of
vegetation undergoing rapid change in the species that compose it; or to
very stable vegetation that has exhibited no significant change for
hundreds of years. An association is a particular type of community,
which has been described sufficiently and repeatedly in several locations
such that we can conclude that it has: (a) a relatively consistent floristic
composition, (b) a uniform physiognomy, and (c) a distribution that is
characteristic of a particular habitat. In terms of its basic importance to
plant ecology and to the classification of vegetation, the association has
been compared to the species of taxonomy Just as a species is an
abstract, somewhat artificial synthesis of many individual plants, so an
association is a synthesis of many local examples of vegetation called
stands.
An observant person can measure and describe vegetation in such a
way that it can be subdivided neatly into association s. How natural are
these units and how much can their existence be attributed to human bias
and the need to classify?

Some ecologists imply or state that

associations are real, closed entities whose component species are

interdependent. We think that this organismic view needs more evidence
before it can be completely accepted.
C. The Organisms View
One of the traits of an association, first accepted by the
International Botanical Congress of 1910, is a relatively consistent
floristic composition. Wherever a particular habitat repeats itself in a
given region, the same cluster of associated taxa is found. This does not
mean that every species repeats itself, nor even that the majority of taxa

45

are repeated. Some species are extremely widespread, with broad

tolerance ranges, and these can be found in many habitats and in many
associations. Other species may have narrower range limits, but
nevertheless a few individuals can be found beyond the normal limits,
and they are occasional members of many communities.
According to the procedures of one method of community
classification, after accidental and ubiquitous taxa are removed from
consideration, certain species will remain that show a large degree of
association with each other and with a particular habitat. Associations are
defined by the presence of such char acteristic or indicator species, and a
particular stand is placed in an association if it contains a significant
fraction of such species. If clusters of species do repeatedly associate
together, that is indirect evidence for either positive or neutral (not
negative) interactions between them. Such evidence favors a view that
communities are indeed integrated unitsthat the whole is somehow
greater than the sum of its parts, much like an organism is greater than
the sum of its cells, tissues, or organs. Some of the interactions
described, such as mycorrhizae, are also evidence that com munities are
units. Clements (1916, 1920) metaphorically equated associations with
organisms. The pattern of species distribution and abundance predicted
by this organismic view The species in an association have similar
distribution limits along the horizontal axis, and many of them rise to
maximum abundance at the same points (noda). The ecotones between
adjacent associations are narrow, with very little overlap of species
ranges, except for a few ubiquitous taxa found in many associations.

D. The Continuum View of Community

Gleason (1953) sampled forest vegetation along a north-south gradient in

46

the Midwest and concluded that changes in species abundance and presence
occurred so gradually that it was not practical to divide the vegetation into associations. Even within a relatively homogeneous region, there were subtle but
important differences in the vegetation from one plot to another.
The sole conclusion we can draw from all the foregoing considerations is
that the vegetation of an area is merely the resultant of two factors, the
fluctuating and fortuitous immigration of plants and an equally fluctuating and
variable environment. As a result, there is no inherent reason why any two
areas of the earth's surface should bear precisely the same vegetation, nor any
reason for adhering to our old ideas of the definiteness and distinctness of
plant associations.Again, experience has shown that it is impossible for
ecologists to agree on the scope of the plant association or on the method of
classifying plant communities. Furthermore, it seems that the vegetation of a
region is not capable of complete segregation into definite communities, but
that there is a considerable development of vegetational mixtures.
It is now apparent that the method used to sample vegetation will
determine whether associations appear as distinct units or as arbitrary segments
along a continuum. If ampling is subjective, searching for stands that show the
presence of differential species, then the discrete view will be supported, for
only similar stands will be sampled, and intermediate stands will be ignored. It
is possible that if only climax stands are sampled, many intermediate stands
will also be ignored. If sampling is done less subjectively, ecotonal stands will
be included, and all the stands will be seen to lie along a continuum. Exceptions
to the continuum view do exist, of course, where there are sudden
discontinuities in the environment, such as a change in soil parent material, a
change in elevation or slope aspect, the advance of fire, or the presence of
landslide nibble.

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E. Some Community Attributes: Physiognomy, Species Composition,

and Spatial Patterns
Physiognomy is a combination of the external appearance of
vegetation, its vertical structure (its architecture or biomass structure), and
the growth forms of its dominant taxa. Physiognomy is an emergent trait of
communities. It could not, for example, be accurately estimated just from a
list of all the taxa present within the community. Life form includes such
plant features as size, life-span, degree of woodiness, degree of
independence, general morphology, leaf traits, the location of perennating
buds, and phenology Vertical structure refers to the height and canopy
coverage of each layer within the community.
Canopy coverage can be expressed as the percentage of ground covered by
the canopy, when the edges of the canopy are mentally projected down to the
surface. It may also be expressed as leaf area index (LAI). All leaves are
collected that project into a column of space that lies above an area of
ground (say, 1 m 2 ), and their cumulative surface area is measured. 'then,
total leaf area, one surface only LAI unit ground area Many crops, such
as corn, have an LAI of about 4, meaning that for every square meter of
ground, 4 m 2 of leaves lie above it. The species composition of a
community is also extremely important, because communities are partly
defined on a floristic basis. Several communities may have similar
physiognomies yet differ in the identity of dominants or other spe cies.
The abundance, importance, or dominance of each species can be
expressed numerically, so that different communities can be compared on
the basis of species similarities and differences.
The relative spatial arrangement of species within a community is
another community trait. As described in earlier chapters, individuals
within a species or individuals of different species may be distributed at
random with respect to each other, clumped (positive or neutral

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interactions), or overdispersed (negative interactions).

One theory of community stability, long-lasting communities
exhibit more species interactions and more species than transient, semi
communities. It this is so, then the fliChe:-. of species in stable
communities must be narrower, with less overlap in niche boundaries,
than those of less stable communities. Some ecologists have tried to
measure niche breadth and overlap to test such hypotheses, but the data
are so sparse that generalities cannot yet be made.
Part of

species' niche may include its unique metabolism. For

example, flowering plants exhibit three different metabolic pathways

of photosynthesis, C3, C4, and CAM, each of which is best fitted to
different environmental conditions or to different species' phonologies.
There have been some recent attempts to include such metabolic traits
in desert and chaparral community descriptions .
Community descriptions based on physiognomy, life form, niche
overlap, and other functional traits (as opposed to taxonomic traits
such as the identity of species in the communities) are useful because
they permit comparison of widely disjunct stands that have little or no
floristic similarity. These comparisons often show a convergence of
vegetation types, given a similar macroenviron ment.
F. Species Richness, Evenness, and Diversity
Species richness is simply the number of species in some area within a
community. Each species is not likely however, to have the same number of
individuals. One species may be represented by 1000 plants, another by 200, and
a third by only a single plant. The distribution of individuals among the species is
called species evenness, or species equitability. Evenness is maximum when all
species have the same number of individuals. Species diversity is a combination
of richness and evenness; it is species richness weighted by species evenness, and

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there are formulae that permit the diversity of a community to be expressed in a

single index number.
It is important to emphasize that richness and diversity are quite different.
Although richness and diversity are often positively correlated, environmental
gradients do exist along which a decrease in richness is accompanied by an
increase in diversity (1 hirlbert 1971). Community A, with five species but
uneven numbers of individuals in each species, has a lower diversity than
community 13, with four species that have a very similar number of individuals
in each. Community A has a higher species richness, however. Biologically,
diversity is the measure of population heterogeneity of a community.
Many simplifications are made in every calculation of species diversity
(Peet 1974). In the equations, all individuals of any species are equal. This may
not he true, especially in regard to animals of different sex or of different
developmental stages, or to plants of different phenological stages (dormant,
full-leaf, flowering, juvenile, senescent adult), different sizes (seedling, sapling,
suppressed adult, overstory adult), or different ecotypes. For this reason,
diversity is often calculated for each stratum in a community rather than for the
entire community. It is also assumed that all species are equally different,
whether the difference is in morphology or niche breadth. This assumption is
generally made even when it is unlikely to be valid, simply because we have no
way to quantify species differences. Finally, diversity is sometimes expressed as
numbers of individuals, biomass, or productivity, as canopy cover, and
sometimes in other ways. Obviously plant diversity from community to
community can be compared only if the same units are used. Some units arc
more appropriate to plants, and some to animals; possibly there is no one unit
suitable for calculation of an ecosystem-wide diversity index.
A further assumption, and one that is made in any statistical procedure, is
that the sample of the community was large enough to represent the community
adequately. The data on numbers of species and the relative abundance of each

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are dependent on sample size. Given a large enough sample, there will be some
species with few individuals, some species with many, and many species with an
intermediate number. The data will form a bell-shaped curve on a log-normal
plot. A small sample, however, will probably not include the full spectrum of
rare species.
What Does Diversity Signify? A lot of attention has been paid to
species diversity in the ecological literature: first, in the development
and comparison of several formula; second, in searching for general
trends in diversity along environmental gradients; and third, in trying to
attribute functional explanations for diversity gradients and some
ecological value that diversity might convey about communities.
Despite all this attention, we think it best to treat an index of diversity
as simply one descriptive attribute of a community, on a par (for
example) with a list of all species found in the community, or an
estimate of tons of biomass above the ground, or leaf area index. Generally,
there is a gradient of increasing species diversity (and richness) from the
poles to the equator, and from high elevations to low elevations. These
gradients follow complex environmental gradients of increasing warmth,
among other factors. It has been stated that diversity increases as any
particular stress lessens, but this is not true for all stress gradients. An
aridity stress gradient may show the opposite trend, with greater diversity
in semiarid grassland and desert than in savanna, woodland, or forest.
Diversity has been equated' with productivity and stability, but some very
diverse semiarid grasslands and deserts are low in terms of productivity
and stability, and there are many other exceptions. Diver sity has been taken
as a reflection of the many interactions that supposedly characterize
complex communities. However, one model of community structure that
assumes no species interactions shows that species diversity "may be main tained in spite of, rather than because of, such interactions" .

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Maintenance of high diversity appears to require episodic, random

(stochastic)
homogeneous

disturbance.
communities

Very

stable,

exhibit

regionally

lower

species

extensive,

and

diversity

than

communities composed of a mosaic of patches disturbed at various times in

the past by wind throw, fire, disease, etc. Following disturbance, diversity
increases with time up to a point where dominance by a few, long-lived,
large-sized species reverses the trend, and diversity falls thereafter.
Communities differ in their utilization of certain essential nutrients
(that is, how much of each element is required for normal growth, or at
least how much is absorbed from the soil solution and translocated to
leaves and growing points). They also differ in the rate at which the
nutrients are returned to the soil in litter fall and the efficiency of the
plant-soil-plant cycle. Early successional commu nities, for example,
may require little soil nitrogen, accumulate very little of any nutrient in
their tissues, and return nutrients rapidly to the soil, but erosion
removes a large fraction of the returned nutrients because of their low
cover or seasonal absence. Climax communities may require greater
quantities of some nutrients, store great quantities of nutrients in wood,
and return only a small fraction to the soil in leaf litter, but prevent
erosive losses by shielding the soil with a permanent, closed canopy .
Thus, climax communities have fewer leaks in nutrient cycles and more
efficiently hold the nutrients in the plant-soil-plant cycle.
The low efficiency and net productivity of desert communities does not
mean that the component species are themselves inefficient or capable of only
very low photosynthetic rates. Net productivity is a community trait and to
some extent a climatic one, for it is affected by the leaf area index, temperatures
during the growing season, distribution of rainfall, soil moisture storage, and
the length of the growing season. If net productivity or efficiency is expressed
on a 12-month basis, it is clear that the tropical rain forest, with a leaf area

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index of 10-11 and warm temperatures and adequate moisture all year, should
indeed have a greater net productivity and efficiency than a desert, with a leaf
area index of I or less and a growing season (based on water supply) of only a
few months. However, the photosynthetic rates of individual desert shrubs and
shrubs of mesic forests are very similar.
The allocation of net productivity to different organs also differs from
community to community. Grassland communities channel much of their energy
to below ground biomass, scrub communities less so, and forests even less so. If
we compare the distribution of carbon in a subalpine conifer forest, a temperate
broadleaf forest, and a tropical rain forest, there are significant differences.
Most carbon in the subalpine forest is in the form of humus atop or in the
surface soil. This is because the acidic, cold soil is not favorable to decomposers. Litter half-life is 10 or more years. Most carbon in the tropical rain
forest, in contrast, is locked up as inert wood. This means that leaf litter has a
major function in the tropical rain forest, for it represents the only mobile,
cycleable part of the nutrient bank. This is one reason why tropical soils cleared
of forest vegetation soon become infertile. The annual litter rain has been
stopped, and the limited soil reserves are soon depleted by crops or leached
from the soil. The litter decay rate is very rapid beneath the rain forest canopy;
litter half-life is a fraction of a year. Nutrient and productivity relationships of
plant communities.
G. Change Over Time
Plant communities that exhibit no directional change for several
centuries are considered to be in equilibrium with their environment, and
are

called

climax

communities.

Other

communities

may

exhibit

significant changes in such a time period. Some species decline in

abundance and may disappear from the site; invasive species may
increase in abundance; and the vegetation type itself may change, for

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example, from a meadow to a forest, or from a pine forest to a hardwood

forest. Such transient communities are called successional, or seral,
communities. It may be possible to recognize and describe an entire
sequence of successional communities that replace each other on one site,
finally culminating in a climax community. This sequence of communities
is called a succession, or sere.
Stability Communities differ in their response to disturbance or stress.
Intuitively, one can use terms such as stable and fragile to describe how
easily communities are perturbed by stress. More technically, however,
stability is a complex term that includes several distinctly different qualities.
One component of stability is resistance, which is the ability of a
community to remain unchanged during a period of stress. Resistance
appears to be characteristic of vegetation dominated by long-lived
perennials with moderately high species diversity and many paths of
interdependence (links or connections) between the component species.
Climax communities generally fit this definition. Such communities have
considerable inertia, changing slowly even in the face of regional climatic
change.
Resilience is a second component of stability; this is the ability of
a community to return to normal, or the rate at which this occurs,
following a period of stress or disturbance. Resilience appears to be
characteristic of vegetation dominated by short-lived, rapidly maturing
species of low diversity and with few interdependence links among
them. Early seral communities fit this defini tion. Climax communities
require considerable time to recover from destructive disturbance , thus
they have low resilience--but they have great resistance to less
destructive stress.
A third component is variance, which is the ability to exhibit patches
of variable abundance in some of the component species. Many climax

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communities appear homogeneous on a large scale, but at a very local

scale they show considerable variance. A fourth component is persistence,
which is the ability to remain relatively unchanged over time. Some
persistent communities are neither resistant nor resilient but owe their
continued existence to a protected, buffered environment. The redwood
forest along the northern California coast is a good example; it exists in a
very narrow, foggy, temperate belt.
Relatively small changes in global temperatures have created striking
changes in climate and vegetation. Most careful analyses suggest an average
global surface temperature difference between full glacial eras and the
present of only 4-6'C (Bryson 1974). This temperature difference, however,
has been correlated with changes in cloudiness, rainfall, length of the frostfree season, and other factors that have a large impact on vegetation.
H. Summary
The community concept is of general importance to synecology, just
as the ecotype concept is central to autecology and the species concept is
central to taxonomy. The precise nature of the community is ambiguous
because of the biases of individual ecologists and their sampling methods.
This does not make the concept useless, however; we must simply
appreciate its subjectivity. For the purposes of classification, stands can he
grouped into associations that have a fixed floristic composition,
physiognomy, and habitat range.
The discrete view of associations assumes that they are closed
systems,

with

interdependent

species

that

synchronously

peak

in

abundance and with narrow ecotones. The discrete view assumes that the
whole is greater than the sum of its parts. The individualistic view
assumes that associations are open systems, with independent species that
happen to associate together wherever their range limits and the chance

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arrival of propagules overlap; consequently, associations are at most

arbitrary units along a continuum. Communities differ in their demand for
essential nutrient elements, in the efficiency with which radiant energy is
converted into net productivity, in the fraction of the nutrient pool that is
stored, in the rate at which nutrients.
I. Evaluatioan
4 Explain scope of plant community!
5 Explain the attributes of physiognomy concept, species composition
6

and spatial patterns !

Explain the concept of species richness, evenness, and diversity!