Concept of community and basic terms
A population consists of organisms of a particular species and characteristics like natality, mortality, age structure, growth dynamics, and so on. But, when several populations share a common habitat and its resources; they interact among themselves and develop into a BIOTIC ‘community’. Thus a community is a larger unit since it includes more than one species’ population.
A biotic community has its own characteristics:
- Animal populations sharing a common habitat and interacting among themselves form an animal community, and Plant populations of an area form a plant community.
- But, the biotic community includes all populations of living organisms of a common habitat, ranging from a tract of forest to the whole forests, from a small pond to a lake and so on.
- It also exhibits certain distinguishable characteristics, such as – diversity, dominance, density, composition and stratification.
The biological potential of species’ population determines a tolerance range for environmental conditions. The range of environmental conditions which a taxon can tolerate is called the ecological amplitude.
The composition of a biotic community in any habitat is dependant upon the prevalence of environmental conditions in that habitat and the ecological amplitude of species’ populations.
Ecotone
Each community has its spatial limits. Sometimes the boundary between two communities may be very sharp (such as that between a forest and a lake) or gradual (as between a forest and grassland). This transitional zone or junction zone between two or more diverse communities is called ecotone.
The ecotone harbors a community (called an ecotonal community) including organisms of each of the overlapping communities, and in addition, organisms peculiar to it. Generally the number of species and their population densities are greater in ecotones than in the communities flanking it. The occurrence of increased diversity and density of organism at the ecotone is due to the edge effect of two distinct communities.
For example, if man settles in a forest, he reduces the forest to scattered small areas interspaced with grasslands, agricultural lands and other open habitats. If he settles in the plains, he plants trees around his habitation. Thus by his activities, man creates an ecotone – some of the original organisms of the forest and plains survive in the man-made forest edge ad many species of insects, birds and mammals often increase in number in these zones.
Community function
Community (in ecology) is a dynamic system, which changes over time through the functions of community dynamics and succession. Moreover, to add on to this, we have seen that communities are made up of populations which interact with each other in many ways and influence their development over-time.
Ecological succession involves an orderly process of community changes, which are directional and hence predictable. It involves a modification of the physical environment, culminating in the establishment of a stable community. A sequence of temporary communities replace one another in a given site, thus bringing about changes in the physical environment, which in-turn determines the pattern of succession. In other words, succession can be regarded as the changes which increase the community complexity over time.
The transitional series communities which develop in a given area are called ‘sere’ or ‘seral’ stages, while the final stable and mature community is called the climax. Ecological succession is therefore community controlled although the environment determines the successional pattern. Succession is of two types, a).Primary succession, and, b). Secondary succession.
Keystone Species: In many communities one or a few species appear to be very important in maintaining the community structure… these are called “Keystone”
Primary Succession (PS) begins on a sterile area (an area not previously occupied by a community), such as a newly exposed rock or sand dune where conditions for existence may not be favorable initially. PS involves the following stages,
- Nudation – this is the exposure of the new surface and is followed by the arrival of the propagates/propagules or seeds from a neighboring region through the process called migration.
- Germination phase – where the seeds and propagules depending on the environmental conditions germinate.
- Ecesis phase – this is where the seedlings establish themselves on the land’s surface.
- Full-scale colonization – this is the 4th phase.
Colonization by successive off-spring and new migrants help increase the population, through a process called aggregation. Plants of organism (autotrophic) which are the first to colonize and aggregate are called pioneers. PS refers to autotrophic succession – which begins in a predominantly in-organic environment, or, succession which takes place when bare, lifeless substrate becomes available for colonization (VERY slow process).
Secondary Succession (SS) refers to community development on sites previously occupied by well-developed communities (the site may be rich in nutrients and hold favorable survival conditions). SS can be regarded as re-colonization following disturbance (much faster than PS). For example, SSs that occur in cut-over forests and abandoned croplands – here, the environment is both in-organic and organic based. Another type of succession occurs on a fallen log or in an aquatic environment (ecosystem) loaded with sewerage. Here the environment is predominantly organic and heterotrophic organisms usually dominate. This type of succession is called heterotrophic succession.
Some salient points regarding SS
A plant community may be disturbed causing some plants to be destroyed, as from a fire or from human logging or cultivation. If the disturbance stops, the community will begin a SS, changes in the vegetation that will lead back to a climax community i.e. the progression of plant communities occurring on areas where there has been previous vegetation (destroyed by fire, farming, or other).
Since the soil is already in place, SS can take place five to ten times faster than PS.
It is important to remember that the abiotic factors (such as weather, humidity, and temperature) affect the nature of the plant community. Also the plant community affects these abiotic factors. Therefore, if a plant community is significantly disturbed, the loss of the vegetation may change the abiotic conditions. If this occurs and the habitat has changed, SS may lead to a different climax community.
What are the main ecological factors makes succession happen?
- Tolerance: only species which can tolerate full range of conditions survive early succession. Some pioneering species dominate because they have broader “tolerance ranges”.
- Facilitation- species present change the environment and make it more hospitable for others.
- Inhibition- species present change the environment and make it less suitable for themselves
An example of ecological succession (PS)
Consider the hydrarch succession or hydro-sere in which a pond and its community are converted into a land community. This could happen in the following stages:
In the initial stage – phytoplankton are the pioneer colonizers. They are consumed by zooplankton and fish. Gradually these organisms increase the content of dead organic matter in the pond. This is utilized by the bacteria and fungi and minerals are released after the decomposition.
The nutrient rich mud then supports the growth of rooted hydrophytes (Vallisneria, Gratophyllum etc) in the shallow water. Thereafter, the hydrophytes die and are decomposed by microorganisms thus releasing nutrients.
Besides, some dead organic matter lies in the mud, gradually reducing the margin of the pond, which is occupied by species whose leaves reach the water surface and roots remain in the mud (nelumbo, nucifera, Trapa spp, monocharia spp etc, grow in these conditions.
Gradually the water depth in the pond decreases due to evaporation and the decomposition of organic matter, and the concentration of nutrients increase.
Free floating plants like lemna, pistia etc increase in number because of the high nutrient availability. Gradually their dead parts fill up the pond bottom, which gets raised. The pond becomes (then) a swampy ecosystem. The reed species invade the pond and are gradually replaced by mesic communities as the water depth is reduced gradually. Thereafter, gradually land plants invade.
In association with the changes in water depth and vegetation the aquatic fauna also change and ultimately gets replaced by land animals.
Consider the case of natural water reservoirs that may be found in some plateaus and valleys. Hydrosere arising in these aquatic environments usually leads to the establishment of deciduous forests. A possible trend of succession in the aquatic environment can be summarized as follows.
Xerosere or xerarch secession
This begins on exposed parent rocks (lithosere) or drysand (psammosere). The pioneer plants are lichens, mosses, selagnella, which help in the soil formation by accelerating erosion. In course of time, grasses, animals and herbaceous vegetation grow on the soils deposited on rocks. Later the mixed woodland species (e.g. ficus etc) appears. In saline areas, the sere is called halosere.
An example of PS for a forest habitat that can happen on a barren rock
- Bare rock is first colonized by lichens and bacteria.
- Small amount of soil formed by the lichens is colonized by mosses, which do not have roots and require little soil, and ferns.
- As the seedless plants live and die, the soil continues to develop to the point that grasses can successfully grow and a grassland community forms.
- Over time, the soil level increases to the point that shrubs can grow in the grassland.
- The grassland is replaced by a shrub community.
- The shrub community may be gradually be replaced by a forest.
Lichens (pioneer species) à mosses and ferns à grasses à shrubs àtrees…
In this example, please note that each stage alters the habitat in such a way that it prepares the way for the next invasion of species. As succession proceeds, soil is formed and thickens – the result of decomposition.
When the changes in the composition of plants stop and the plant community remains generally the same for many years, the community is mature or at climax. A climax community is the relatively stable community at the end of succession.
Dynamic trends in succession
Some structural and functional changes which occur in the process of succession can be grouped/categorized as follows:
- Species composition – a change in the species composition occurs. The change is fast in the beginning and then is more gradual.
- Species diversity – some plant species which were present in the initial stages may not be found in an advanced stage of succession. However, in the climax stage – there may be more kinds of autotrophs and heterotrophs than in the earlier seral stages.
- Density and biomass of organisms – there is usually a marked increase in the number (density) of organism. The number may decline in the latter stages, but in the climax stage the biomass structure remains very high. The total biomass gradually increases and reaches a maximum in the climax stage.
- Heterotroph population – the number of species usually goes on increasing as the food chain relationships become more complex in the climax stage.
- Chlorophyll – green pigments go on increasing during the early phases of PS. The ratio of yellow/green pigments remain around 2 in the early stages and increases to 3 to 5 in the climax stage. Pigment diversity also increases.
Functional Changes during succession
There is a progressive increase in the amount of living biomass and dead organic matter. There is an increase in grass as well as net primary production in the initial and seral stages. Thus there is more biomass accumulation, gradually reaching a huge biomass structure in the climax stage.
The food-chain relationships become more complex as succession proceeds. Researchers have established that the stability of the climax community is associated with high species diversity, large accumulation of the living bio-mass and complex food chain relationships. Figure below summarizes the changes in biomass and primary production that can occur through the passage of succession.
The community respiration increases but the P/R ratio (‘production to respiration’ ratio) remains more than 1 in the seral stages. The huge living biomass respires a lot in the climax stage and the P/R ratio equals 1 (P:R = 1). Thus in the early stages P>R, and in the climax P = R. Refer to figure below.
The complexity of the climax community increases number of ecological niches and the routes of flow through the system. These attributes make the climax community more stable. Furthermore, it has also been established that productivity increases from poles to the tropics, and hence it may mean that a climax tropical forest is more stable than a temperate climate deciduous forests (and, so on).
The climax concept in ecological succession
Many theories (to date) have been put forward by ecologists to explain the climax concept such as,
- The mono climax theory,
- The poly climax theory, and
- The Information theory.
1. The mono climax theory
Clements (1916), was of the view that in a given climate, the successional stages (seral stages) will ultimately end up as climatic climax vegetation. If this to be true, succession is a progressive phenomenon. The emphasis is on the fact that only one type of climax vegetation develops. This is called the mono climax theory. But some communities like savannahs/prairies with grass land climax (in Canada, USA, Africa etc) also prairies, shrubs and forest patches in low areas in the mesic belt as stable vegetation.
Ecologists supporting the mono climax theory argue that this vegetation is post climax. But others feel that the post climax concept is confusing. It has also been observed that different types of stable vegetation occur within the same climatic belt. Yet other plant ecologists consider this vegetation pre-climax. Pre-climax vegetation is that which has not reached the climax stage due to the prevalence of some adverse climatic, topographic and edephic environmental conditions. Due to human activites, such as cutting, fire, grazing and so on, the successional process may be checked and the community may not reach the climax stage. However, some sort of stable vegetation may develop under such circumstances. Such vegetation is called dis-climax or post climax.
2. Poly climax theory
Looking a the occurrence of several types of climax vegetation (in Europe), Braun-Blanquet (1932) proposed the polyclimax theory, which states that there may be climatic climax, edaphic climax and biotic climax depending upon the situations in which the climax vegetation has developed.
Therefore, succession may not always be progressive. In course of succession, a community with longer life forms like forests may be degraded and patches of grass land may occur within them (retrogressive succession)
Note: the two above said theories explain the climax concept from a structural view point. However, looking at the problem/case of succession from an energistics point of view is also possible (since energy drives all functions in a community or in an ecosystem). This has led to the development of the ‘information theory’ – to explain the concept of the climax community.
3. Information theory
In this the community is considered a thermodynamic unit. It receives energy from the sun and concerts it into chemical energy from the sun and converts it into chemical energy, performs its activities and dissipates heat energy. In the seral stages, the dissipation energy is usually less than the input energy – therefore there is more of net production, and the community grows. In the early stages, the species diversity is low and the food chain relationships simple; but in the course of scission the total information in the community increases (i.e. the food chain relationships become more complex and the possible interactions between individuals, species and materials increase).
In the climax community the input energy is more or less equal to the output energy (particularly) the dissipation energy), which makes for negligible NPP.
Time factor in ecological succession
Ecological succession is dependent on the time factor even though it is often taken for granted. For example, consider the foll cases; primary succession on sand dunes or recent lava follows takes about 1000 years.
Secondary succession on abandoned agricultural land in a tropical climate taken about 100 years; whereas for a moist temperate climate – it is 200 years
Secondary succession in grasslands (as per the observations/experiments in North America by ODUM in 1966) take about 50 to 60 years to reach a climax grass stage – this occurs in 4 distinguishable stages,
- 2 – 5 years of annual weed stage,
- 3 – 10 years of short-lived grass stage,
- 10 – 20 years of perennial grass stage, and,
- The climax grassland stage – reached in some 20-40 years.
Significance of ecological succession
The principles and trends of ecological succession indicate that seral stages are more productive, although comparatively less stable. The climax community is nature and stable with greater biological diversity, larger becomes structure and balanced energy flow and is able to buffer the physical environment. This community provides man with food, fuel, fodder, medicine and so on and is able to control the climate and keep a balance with regard to biogeochemical cycles. It is (as such), considered a multiple use system – since the P/R ratio in such a community is – there is not much NPP for harvesting. Since the fast growing human population needs a huge amount of food and other materials – man is forced to look for high NPP systems – which decreases that he needs early successional stages as a source of food.
Disturbances to ecosystems and the continuous cycle of succession
Disturbances (either by natural or anthropogenic means) are an integral part of some ecosystems, e.g. forest fire destructions are part of the dynamics of eucalyptus forestland in Australia. Any event that interrupts life in an ecosystem is (can be) defined as ‘disturbance’.
If there is no further manipulation of the system after the disturbance an ecosystem will slowly (gradually) develop and reach its climax stage. But, when there are intermittent shocks, the climax stage will not be reached and a lesser stable community dominating the system is also possibility to be considered. For example, think about the case-study on the grasslands in Batticaloa you have taken up for ENS 2133. Can you relate that example with SS? Please consider the diagram below as a ‘thought-provoker’.
Why is it important for us to understand the concepts behind succession?
As a budding environmental scientist, you need to know, that:
- Natural systems can maintain themselves, whereas disturbed systems cannot.
- Human influence on succession can be positive or negative – generally, humans knock out the climax community.
An interesting example of human interference with succession = producing a green, weed-free football ground (or lawn). Producing such a lawn means a constant battle against natural plant succession
- What you must do to maintain such a system
- plant grass seed – these are needed to complete against natural seed dispersal of many different species,
- mow – this limits the establishment of trees and other tall species,
- apply herbicides – to eliminate inter-specific competition (competition between different species) and to cut down on species diversity – must be done because broad-leaved species are often more adaptable and successful than grasses and can out-compete them,
- apply fertilizer – grass species use up soil nutrients quickly (particularly when clippings are bagged),
- irrigate – natural rainfall is often inadequate for lawn grass,
- apply fungicides and other chemicals to control disease and insect pests- must be done because an unnatural ecosystem (such as a lawn) is more prone to disruption.
- Thus, when humans interfere with plant communities – many problems that would normally be controlled in a natural systems occur at a blown-up level (this is same thing happens in farming).
- Without the continued interference of humans, the yard would have a natural sequence of succession over time.
Summary
We have seen the various aspects of ecological succession, and why it is an inevitable phenomenon in ecology. As student of environmental science, you should be able to correlate the theoretical elements behind this phenomenon to understand the natural environment, and to manage the environment sustainably. Knowing when and where ecological succession can be manipulated, gives you the power to design and improve ecosystems for the benefit of the human society (e.g. Coppock et al., experiment of 1993 in Africa). However, the ethical aspects of manipulating the natural environment should be given careful consideration before taking any drastic action to modify the natural composition(s).