NATURE OF VEGETATION

There are 3 basic concepts of the nature of vegetation as it exists on a given piece of ground
(space) at a given time :-

1. The "Individualistic Concept" advanced by Gleason over 75 years ago.

2. The "Plant Community Concept" as advocated by Clements.

3. The "Continuun" or "Gradient" concept - developed over the last 40 years, particularly
by Curtis and Whittaker in the United States, but first recognized and advanced by Russian
ecologist Ramenskii over 70 years ago.

The concept that a research ecologist accepts will determine his objectives and approach to any
vegetation problem. This in turn will predispose him towards using particular methods and
techniques - which in turn leads him to collect particular kinds of data, with their own
limitations when it comes to the interpretation of results.

THE INDIVIDUALISTIC CONCEPT

This concept was advanced by ecologist Gleason over 75 years ago. He argued that no
"patches" of vegetation are exactly alike in the combination of species present or the
proportions (abundance) of these species. This is a fact that can easily be verified in the field
even today.

Gleason argued that the properties of vegetation depend completely on the properties of
individual plant species composing it (hence the term "individualistic"). He argued that the
vegetation found at a particular area or locality is the result of selection by an environment that
fluctuates and varies in both time and space; and that such selection will have acted on plant
species that arrived at that particular locality through a similarly fluctuating and unpredictable
process of immigration. Communities are not fixed or stable; they change over time due to
migration, dispersal, and environmental changes. As such, chance plays a major role in the
development of vegetation, according to this concept.

Today, some ecologists hold the view since no new ideas has emerged that would lead one to
reject this concept.

PLANT COMMUNITY CONCEPT

Although no two patches of vegetation are exactly alike, it is equally true that "patches" of
vegetation growing under similar environmental conditions, and under similar histories of both
environment and plant migration, will often be very alike in terms of species composition and
abundance. As such, it is possible to recognize clearly definable vegetation "types" that are
similar in species composition and in the abundance of these species.

This frequent similarity of vegetation "types" led to the idea of "plant communities", as
vegetation units that repeated themselves in space. It led some ecologists, such as Clement, to
consider such a community as being analogous to an ‘organism’, Clements likened succession
to the development of an organism, where each stage represents a developmental phase
contributing to the whole. (as advanced in the Clementian/Classical succession model with the
stages: nudation, migration,ecesis, competition, stabilisation climax community).

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Most ecologists today hold the view that such an analogy is untenable. A "patch" of vegetation
is often comprised of plant species that have very different evolutionary histories. Such a
"patch" of vegetation can not be traced along a distinct ancestral phylogenetic line, as one can
do with an organism. Hence, the "patch" of vegetation is fundamentally different from an
organism.

That notwithstanding, a patch of vegetation does show the development of certain properties
which are characteristic of the species assemblage as a whole, rather than of the individual
species that make up that patch of vegetation. These characteristics include :-

• Many of the component species may have been evolving in close proximity to each
other over a long duration of time, and each species can be presumed to be still evolving
under selection pressures that include the influence of surrounding species. Therefore,
the process of (natural) selection both affects and is a function of the vegetation as a
whole.

• Species may develop adaptations such that they are substantially dependent on others
for their niches - e.g. the phenomenon of mutualism (e.g. epiphytes).

• There may be marked structural organization within the vegetation as a whole - e.g.
vertical stratification - a characteristic that is not applicable to individual species.

• The behaviour of a species population growing in a mixture of species can differ

markedly from that of the same species population growing alone in a pure stand.

The plant community concept is central to vegetation classification. Any time an ecologist tries
to classify vegetation, he is in essence trying to distinguish different "vegetation types", and
has de-facto accepted the plant community concept. The identification of such vegetation types
has practical value in ecological research - instead of dealing with hundreds of individual
species, the ecologist can deal with a single vegetation entity (community), on which he can
generalize. This concept is also central to the Classical/Clementian succession model (note that
there are alternative models).

However, it must be noted that plant communities are not separated by sharp/distinct
boundaries. Rather, they merge or inter-grade into each other. Hence, any attempt to demarcate
a plant community will in essence involve placing arbitrary boundaries. For this reason, some
ecologists today question whether plant communities actually exist in nature, or whether they
are simply a creation of the human mind.

CONTINUUN (GRADIENT) CONCEPT

Under natural conditions, changes in vegetation tend to be gradual, with one vegetation type
merging or inter-grading into the next. Situations where one vegetation type is clearly
demarcated from another are not common (but they do occur, for example where an area has
an environmental history that is critically different from that of the area surrounding it - e.g. an
aquatic/terrestrial habitat interface for a lake)

The continuum concept emphasizes (and is based on) the way in which different vegetation
types inter-grade - or the way in which species and species assemblages are related along
environmental gradients.

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The methods and procedures developed to investigate vegetation from this viewpoint are
collectively referred to as Gradient Analysis. (c.f. methods and procedures developed to
investigate vegetation from the "plant community" viewpoint - classification procedures).

The continuum (gradient) concept was actually advanced over 70 years ago by Russian
ecologist L. G. Ramenskii, who has largely been ignored in the western literature. Over the
past 40 years, This concept has been independently advanced and developed over the last 40
years, especially by American ecologists Whittaker and Curtis.

Whittaker demonstrated how, under natural conditions, species and species assemblages tend
to show broadly overlapping population distributions along environmental gradients, mostly in
the form of binomial distribution. When such distribution is plotted, it tends to appear as
bell-shaped curves (see illustration below).

He demonstrated that this distribution was a reflection of different environmental tolerances of

the individual species.

From this viewpoint, vegetation is seen as a complex pattern of overlapping species

populations. This view has proved very useful in studying the ecology of species populations
and the relationships between species and the environment.

All three concepts can be justified, and the concept that is best suited to a particular task or
research problem is a matter of judgement by the researcher.

WHY SURVEY AND ANALYZE VEGETATION?

There are basically two reasons: -

1. Description and Mapping Purposes

This usually involves large areas, where the aim is description and classification of community
types found in the area. Normally, such an analysis does not concern itself with the interactions
between the vegetation and the environment.

2. The second reason may be termed "Ecological Purposes"

Here, the aim of the analysis is to investigate species to species and species to environment
relationships. One should keep in mind the fact that a fundamental objective of ecological
enquiry is to determine the factors, both biotic and abiotic, which control the occurrence,
abundance, and distribution of plant species.

Essentially, different methods will be suitable for the two approaches. In the first instance
(large scale), communities, or recurring vegetation units, can be recognized. In the second
instance, where more detailed work has to be carried out in much smaller areas, then the
"continuum" nature of vegetation has to be recognized. Here, classification methods may no
longer be appropriate, since classification is not the beast way of dealing with continuous
change. Classification may be useful here only to the extent that it will facilitate the
classification of the vegetation being studied. The more appropriate techniques in this case
would be gradient analysis and ordination procedures, since they specifically deal with change
in vegetation along gradients.

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PROCEDURES TO FOLLOW IN VEGETATION ANALYSIS
There are 4 basic steps that an ecologist must consider in any vegetation analysis:-

1. ENTITATION. Which is recognition of the vegetation (plant community), or part of

the vegetation that is to be studied.
2. TYPE, SIZE, AND SHAPE OF SAMPLE. The researcher must decide on the type,
size, and shape of the samples to be used in studying the selected vegetation. Since it is
impossible to consider all the individual plants that make up the vegetation (100%
census), then representative samples of the vegetation have to be analyzed.
3. SAMPLING DESIGN. The researcher has to decide on how the samples will be placed
within the vegetation to be studied. This has very important implications for subsequent
analyses of the data obtained from the samples.
4. ATTRIBUTES TO MEASURE/RECORD. The researcher must decide what is to be
measured or recorded once the location of the samples has been established.

Since fieldwork is both time consuming and expensive, a researcher will still need to pay
careful attention to all the above 4 steps. However, it must be emphasized that actual collection
of data (sampling) is not the ultimate aim of vegetation research; rather, it provides the raw
material necessary for proper documentation of the study. The importance of vegetation
sampling is that all subsequent treatment of the data, and the conclusions one may draw, depend
on the researcher having paid careful attention to the above 4 steps, especially the initial
selection of the sampling design and the characteristics of the sample.

These 4 steps are reviewed in more detail below:

1. ENTITATION
Entitation refers to recognition of the vegetation (plant community) that is to be studied. This
step is for most part subjective.

It is a process of identifying and describing discrete vegetation units (entities). Essentially, it

is a general inventory of an area's plant communities conducted by means of aerial photograph
interpretation and field reconnaissance (involves describing the plant type, name, plant height,
topography, dominant plant type…etc). The survey results in the identification of ecologically
distinct plant communities based on species and structure.

2. TYPE, SHAPE, AND SIZE OF SAMPLE

Study Area and Sampling Plot Sizes: The general site size should be large enough to capture
a representative sample of the area to be described. The subplot sizes within the site should be
large enough to include significant numbers of individuals, but small enough so that plants can
be identified without duplication or omission of individuals.

Sampling plot size may change depending on species measured.

Cain and Castro (1959) suggest the following sampling sizes:

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In general, there are two types of sampling that a research ecologist can use: -
i. Quadrats (Sampling Plots)
ii. Plotless samples

A transect, which in it's simplest form consists of a straight line running across the vegetation
to be sampled, can be used in combination with either of these two types of samples.

i. QUADRATS
A quadrat (subplots of a standard size) or sampling plot) can be used for most plant
communities.
A quadrat delimits an area in which vegetation cover can be estimated, plants counted, or
species listed. The vegetation contained within the quadrat is referred to as a stand.
Quadrats should be located randomly within a study site. Large quadrats with many plants may
require two or more people to obtain an accurate census, while one person may be sufficient
for smaller plots or those with sparse vegetation.

A quadrat is typically a square frame constructed of plastic or pvc pipe, metal rod, or wood that
is placed directly on top of the vegetation. Quadrats are also commonly called "plots."

Quadrats do not have to be square but their area must be known. Other quadrat shapes
commonly include circles and rectangles. The ideal shape of a quadrant depends on the specific
purpose, but typically, a square is considered ideal for sampling and analyzing distribution in
uniform grids. Squares minimize the complexity of edge effects compared to circles and
irregularly shaped rectangles (Squares have straight edges, making it easier to standardize the
inclusion or exclusion of partially present objects). In general, the difference obtained when
using quadrats of different shapes is very small, and may not be important.

Square quadrats can be any size. Common sizes include: 25 by 25 cm, 50 by 50 cm, 1 by 1 m
and similar sizes in feet. In general, a quadrat size of up to 1m2 is suitable for the study of
herbaceous vegetation, while a quadrat size of 20m by 20m is commonly used in the study of
tropical forest ecosystems.

Quadrats are required for estimating several vegetation attributes including:

Density - for counting the number of objects within the unit area of the quadrat.
Biomass - achieved by "clipping" all the material of a given type (e.g., grass, shrub or forb) or
species within a quadrat.
Cover - often accomplished by estimating the area of a quadrat that is covered by a plant's
canopy.
Frequency - the proportion of quadrats in which a species occurs is called frequency, thus
quadrats are required to estimate plant frequency.

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Once the quadrat is placed on the ground, begin listing all the plants that are covering the space
in the quadrat including any grasses, shrubs and trees.

ii. PLOTLESS SAMPLES

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