Plant succession

Plant succession can be defined as the process of gradual replacement of one plant community by
another plant community which is of stable type. Succession is the natural, orderly change in plant
communities that occurs over time. If left undisturbed, an open field may become an "aging" forest in
150-300 years. The first plant community which develops in a bare area is known as pioneer community
and the last invading community is called the climax community. The plant communities that develop
during the succession are called the seral communities. However natural and anthropogenic forces (soil
conditions, climate, permafrost, topography, fire, flood, insects, and human activity) can disrupt the
forest, altering the pattern of succession.

Figure: Plant Succession

BASIC TYPES OF SUCCESSION:

1. Primary Succession
2. Secondary Succession
3. Autogenic Succession
4. Allogenic Succession
5. Progressive Succession
6. Retrossive Succession

1. Primary Succession:

The formation of ecosystem from bare rock, sand or clear glacial pool where previous life does not exist
is called primary succession. In this case, the ecosystem is formed from the start. So it is a long process.
It often requires thousands of years.
 Figure: Primary Plant Succession

2. Secondary Succession:

The formation of a new ecosystem after the disturbance of an existing ecosystem is called secondary
succession. Die disturbance may be in the form of, forced fire or an abandoned farm field. The previous
community leaves some mark in the form of improved soil and seeds Therefore, secondary succession
occurs more rapidly than primary succession.

Figure: Secondary Plant Succession

On the basis of factors responsible for environment changes successions are sometimes classified as,

1. Autogenic succession:
2. Allogenic succession:
1. Autogenic succession:

The succession in which organisms themselves bring change in the environment during succession is
called autogenic succession. The organisms cause change in the soil. These changes include
accumulation of organic matter in form of humus or litter alteration of soil nutrients and change in pH of
soil. The structure of the plants themselves can also change the community. For example, larger species
like trees produce shade on to the developing forest floor. It destroys the light-requiring species. Shade-
tolerant species-establish in the area.

2. Allogenic succession:

The succession in which external environmental factors cause change in the environment during
succession is called allogenic succession. Soil erosion, leaching or the deposition of silt can changes the
soil. Similarly, clays can alter the nutrient content and water relationships in the ecosystems. Animals
also play an important role in allogenic changes. They act as pollinators. seed dispersers and herbivores.
They can also increase nutrient content of the soil in certain.

On the basis of community environment it can be classified as-

1. Progressive succession
2. Retrogressive succession

1. Progressive succession :

Progressive succession is a succession where the community becomes complex and contains more
species and biomass over time.

2. Retrogressive succession :

Retrogressive succession is a succession where the community becomes simplistic and contains fewer
species and less biomass over time.

DIFFERENT TYPES OF HABITATS IN WHICH SUCCESSION TAKES PLACE

 Hydrosere: succession occurs where water is plenty, e.g. pond.
 Derosere: succession occurs on a dry soil or rock.
 Xerosere: succession occurs on dry habitat like dry desert or bare rock.
 Psammosre: succession which occurs on sand dunes.
 Halosere: succession which occurs on saline soil.
 Lithosere: Succession on newly exposed rock surface.
 Oxylosere: Succession on acidic soil.
 Hydrosere

Oxylosere Derosere

TYPES OF
HABITAT
S
Lithosere Xerosere

Halosere Psammosre

Figure: Type of habitat in which succession can take place

HYDROSERE OR HYDRARCH
It is succession occurring in the aquatic environment. Such a type of succession does not necessarily lead
the aquatic communities toward the development of land communities.

If the body of water is large and very deep or very strong wave action and other powerful physical forces
are at work, the succession results in a stable aquatic community in which any considerable further
change is hardly recognizable.

Succession is recognizable only if the colonization of plant communities takes place in artificial small and
shallow ponds, lakes, etc. where wave action speeds up the process by allowing the erosion of soil
towards edge regions. In this way, the filling process also speeds up quickly and consequently the body
of water disappears within few years time.

Fig. 7.1 illustrates the different types of vegetation at different depths in a pond; floating plants in the
central region; rooted hydrophytes in shallow region, amphibious plants in the marginal mud and trees
developing in dry habitat.
 Figure: Hydrosere

In a new and virgin pond hydrosere starts with the colonisation of phytoplankton and finally terminates
into a forest (the climax community). The process of aquatic succession completes in the following
stages:

1. Phytoplankton stage.
2. Submerged stage.
3. Floating stage.
4. Reed swamp stage.
5. Sedge-meadow stage.
6. Woodland stage.
7. Climax stage.

1. Phytoplankton stage:

In the initial stage of succession algal spores are brought in the body of water. The simple forms of life
like bacteria, algae and many other aquatic plants (phytoplankton) and animals (zooplankton) floating in
water are the pioneer colonizers. All these organisms add large amount of organic matter and nutrients
due to their various life activities and after their death, they settle at the bottom of pond to form a layer
of muck.

2. Submerged stage:

The phytoplankton stage is followed by submerged plant stage. When a loose layer of mud is formed on
the bottom of the pond, some rooted submerged hydrophytes begin to appear on the new substratum.
The submerged aquatic vegetation develops in the regions of ponds or lakes where water depth is about
10 feet or more. The pioneers are Elodia, Potamogeton, Myriophyllum, Ranunculus, Utricularia,
Ceratophyllum, Vallisnena, Chara, etc.
These plants form tangled mass and have marked effects upon the habitat. When these plants die their
remains are deposited at the bottom of the ponds or lakes. The eroded soil particles and other
transported materials are also deposited at the bottom. This gradually raises the bottom of the ponds
and lakes up. As this process of stratification progresses the body of water becomes more and more
shallow, consequently the habitat becomes less suited for the submerged vegetation but more favorable
for other plants.

3. Floating stage:

When the depth of water reaches about 4 to 8 feet, the submerged vegetation starts disappearing from
its original place and then the floating plants make their appearance gradually in that area. In the
beginning the submerged and floating plants grow intermingled but in the course of time the submerged
plants are replaced completely. The most tolerant species in the area are able to reproduce and
perpetuate. Their broad leaves floating on the water surface check the penetration of light to deeper
layer of water.

This may be one of the main causes responsible for the death of submerged plants. Due to continuous
interaction between plant communities and aquatic environment, the habitat becomes changed
chemically as well as physically. More water and air borne soil and dead remains of plants are deposited
at the bottom. Thus, the substratum rises up in vertical direction. Important floating plants that replace
the submerged vegetation are Nelumbum, Trapa, Pistia, Nymphaea. and Limnanthemum etc.

Figure : Hydrosere
4. Reed-swamp stages:

When the ponds and lakes become too shallow (water depth one to three feet) and the habitat is
changed so much that it becomes less suited to the floating plants some other plants which are well
adapted to new environment will then come in Under these conditions, the floating plants start
disappearing gradually and their places are occupied by amphibious plants which can live successfully in
aquatic as well as aerial environment Important examples are Bothrioclova, Typha, Phragmites (Reed),
etc.

The foliage leaves of such plants are exposed much above the surface of water and roots are generally
found either in mud or submerged in water. The foliage leaves form a cover over submerged and
floating plants and thus they cut off light from the plants underneath them. Under such conditions
neither submerged nor floating plants can survive. Further deposition of soil and plant debris at the
bottom reduces the depth of water and makes the habitat less suitable for the pre-existing plants.

When the bottom reaches very close to the water surface many secondary species, such as those of
Polygonum, Sagittaria, etc. make their appearance. Later, they also bring about such reactions by which
the habitat becomes less suitable for most of the existing species, and consequently new successional
step follows.

5. Sedge Marsh or Meadow stage:

The filling process finally results in a marshy soil which may be too dry for the plants of pre- existing
community. Now the plants well adapted to new habitat begin to appear in the pre- existing community
in mixed state. Important plants that are well suited to marshy habitat are the members of cyperaceae
and poaceae. The species of sedge (Carex) and rushes (Juncus), species of Themeda, Iris. Dichanthium,
Eriophorum, Cymbopogon, Campanula, Mentha, Caltha Gallium, Teucrium, Cicuta, etc. are the first
invaders of marshy area.

As these plants grow most luxuriantly in the marshes, they modify the habitats in several ways. They
absorb and transpire a large quantity of water and also catch and accumulate plant debris and wind and
water borne soil particles. Consequently a dry habitat results which may be totally unfit for the growth
of normal hydrophytes. Gradually the mesophytes start appearing and after some time the sedge
vegetation is totally replaced by them.

6. Woodland stage:

In the beginning some shrubs and later medium sized trees form open vegetation or woodland. These
plants produce more shade and absorb and transpire large quantity of water. Thus, they render the
habitat drier. Shade loving herbs may also grow under the trees and shrubs. The prominent plants of
woodland community are species of Buteazon, Acacia, Cassia, Terminalia, Salix, Cephalanthus, etc.
7. Climax forest:

After a very long time the hydrosere may lead to the development of climax vegetation. As the level of
soil is raised much above the water level by progressive accumulation of humus and soil particles, the
habitat becomes more dry and certainly well aerated. In such a habitat, well adapted self-maintaining
and self-reproducing, nearly stable and uniform plant community consisting mostly of woody trees
develops in the form of mesophytic forest.

In the climax forest, all types of plants are met with. Herbs, shrubs, mosses and shade loving plants
represent their own communities. Trees are dominant and they have control over the entire vegetation.
Bacteria, fungi, and other micro-organisms are more frequently found in the climax vegetation. They
react upon the habitat and make the soil rich in the organic materials. At the climax stage, a complete
harmony develops between plant community and habitat. It is now clear that whole sere is a
continuously but gradually changing complex in which the changes are forced by biotic, topographic or
climatic factors. It is very slow process that cannot be observed in nature. It may require thousands of
years to reach the climax stage. One can however, observe the sequence of hydrosere as he moves in
the lake or pond from the deepest region towards the shallower margin.

Figure: Stage of Hydrosere & Xerosere

XEROSERE OR XERARCH
This is a type of succession originating on bare rock surfaces. The original substratum is deficient in
water and lacks any organic matter, having only minerals in disintegrated unweathered state. The
pioneers to colonise this primitive substratum are crustose type of lichens, and through a series of
successive seral stages the succession finally terminates into a forest which constitutes the climax
community.

Figure: Stage of Xerosere

Such as-

 Bare rocks
 Foliose and fruticose lichen stage.
 Moss stage.
 Herb stage.
 Shrub stage.
 Tree stage.
 Climax stage.
1. Lichen stage:

Due to great exposure to sun and extreme deficiency of water, the first pioneers on the bare rock area
are a few simple organisms. The most successful of such organisms are crustose lichens. These are able
to withstand extreme desiccation due to excessive dryness. During rainy season they absorb large
quantities of water and flourish rapidly.

Migration to distant rocks takes place either by spores or soredia by wind. The common species of
crustose lichens are Rhizocarpon, Rinodina, Lecanora etc. The lichen secretes carbonic acid causing rock
disintegration. Rock particles and dead organic matter of lichens accumulate to provide conditions
possible for the growth of higher forms of lichens.

As soon as little soil is formed by the activity of crustose lichens, higher forms of lichens such as foliose-
lichens appear. These include Dermatocarpon, Parmelia, Umbilicaria etc. These have large leafy thalli
which overlap the crustose-lichens and cause their gradual death and decay. In this way more and more
humus accumulates and gradually a thin layer of soil is formed which consists of rock particles, remains
of lichens, dust particles and moisture. Associated with the lichens a few mites make their appearance.
Along with them a few spiders also make their appearance in cracks and crevices of the rock.

2. Moss stage:

With the accumulation of dust and humus in small quantities the environment is altered enough to
allow the establishment of secondary communities in a rather definite sequence. Scattered patches of
mosses such as Tortula, Grimmia Byrum and Barbula etc. begin to invade the environment that had so
far been dominated by lichens. Later on, mosses like Funaria, Sphagnum and Polytrichum make their
appearance. Among the animals, mites become more varied; some small spiders and springtails as well
as tardigrades become associated with this secondary community.

3. Herbaceous stage:

As the mats of mosses become more extensive, more soil accumulates, much of the soil is blown in from
surrounding area during windy periods. More mineral material is added to the soil as acid leaches out
from the overlying vegetation and increases the depth of the mineral soil layer. Many annual weeds
develop which are, later on, followed by biennial and finally perennial grasses. Andropogon commonly
known as broom sedge becomes; dominant grass in many areas. With the influx of grasses, the fauna
(animals) also becomes varied. Nematodes and larval insects, collembola, ants and mites appear in the
gradually altered environment.

4. Shrub stage:

Further modification of the environment provides conditions for the germination and growth of shrubs
and perennial wood plants such as Acacia, Prosopis, Capparis, Zizyphus etc. With the approach of
shrubs, the animals also become vivid and numerous, and join hands with the vegetation in altering the
environment
5. Climax forest:

With the establishment of shrubs, more and more soil is formed and environment becomes increasingly
humid. This favours the growth of woody trees. In the beginning, trees show stunted growth and are
sparsely placed. Finally a climax forest community is established. The climax community is the last
aggregation in the successional series. If the climax condition does change and no catastrophic event
alters the area, the community maintains indefinitely.
 Climax Community
Theory of Climax Community:

The succession of community structure attains a stable and final equilibrium state called Climax
Community suggested by Shimwell (1971) and Clements (1936). This stable community is in accordance
with the predicting biotic and physically environment and is self- perpetuating. Birth rate, death rate and
growth process induce structural changes in climax community. The following are the four theories of
climax community:

1.Mono-climax theory

In 1936, Clements suggested the mono-climax theory of succession according to which all communities
of different regions to reach climax community. He believed that climate is a determining functional
factor of climax and vegetation is solely influenced by the climatic conditions of the community. This
climate based theory termed various stages of succession process as sub-climax, dis-, post- and pre-
climax with stabilized climax as last stage of community. Other than climate controlled community,
exceptional cases of edaphic (soil), topographic and biotic factors influencing communities are also
discussed under this theory.

2. Poly-climax:

Poly-climax theory In 1939, Tansley proposed Poly-climax theory which was supported by Daubenmire
(1966). According to this theory in an area the climax community may have varied type of vegetation
and these are called as climaxes. These climaxes are influenced not only by the climate rather by the
activities of animals, soil nutrients and moisture etc. Unlike mono-climax theory, climate is one factor
among all other influencing factors regulating the stability and structure of climax community and allows
multiple climaxes and thus called as poly-climax theory. In 1994, Kreb suggested that it is the time
dependent relative stability measurements which differentiate between the two theories. The factors
influencing climax varies with geological time scale and ecological time scale such as climate. Thus
different vegetation develops to reach different climax]leading to a continuous succession. Therefore,
the mono-climax theory and poly-climax theory mainly differ on understanding the functional factor
which is responsible for establishing the steady state elimax community.

3. Climax-pattern theory:

In 1953, Whittaker proposed climax pattern hypothesis. The climax pattern theory of succession holds
that the existing ecological community is affected by the entire pattern of environmental factors. The
environmental factors influencing the community can be climate site, activity of animals (biotic), soil,
species genetic structure, plant and animal species, dispersal, wind and fire etc. This theory emphasizes
on the total environmental conditions regulating the pattern of population constituting the climax
community. There is no single functional factor influencing the climax community and thus, there is no
mono- or poly- climax. Unlike mono-climax hypothesis resulting into one climatic climax in an area and
several climaxes in poly-climax theory, the climate pattern theory allow continuous inseparable climax
types under the influence of gradually changing gradient of environmental conditions.

4. Climax as Vegetation:

In 1954, Egler hypothesized in a broad sense that the whole pattern of vegetation represents the
climaxes. He favours the vegetation study of an area as by the careful observation of the vegetation we
can predict the community conditions. Thus from these entire hypothesis we can conclude that the
ultimate goal of succession is to achieve a stable form of community structure called climax. The climax
community can be disturbed and incompletely stable. The vegetation of an area is totally controlled by
the existing climatic conditions but the broad climatic zones leads to many climax populations under the
influence of animal activities, topographic and soil compositions.

Figure: Flow chart depicting different theories of Climax Community

Communities are regularly changing or dynamic entities. The development of the community initiates on
arrival of the propagules or their parent organisms to the area of study. The first developmental stage of
the community is called as the pioneering community which is made up of those few successfully
survived and established organisms or their propagules.