Fundamentals of Ecology

UNIT 8 ECOLOGICAL SUCCESSION AND

CLIMAX

Structure
8.1 Introduction
8.2 Objectives
8.3 Causes and Trends of Succession
8.4 Different Kinds of Succession
8.5 General Process of Succession
8.5.1 Hydrosere
8.5.2 Lithosere (Xerarch)
8.6 Climax Community
8.6.1 Monoclimax Concept
8.6.2 Polyclimax Concept
8.7 Ecological Adaptations
8.7.1 Tolerance Ranges
8.7.2 The Liebig’s Law of Minimum
8.7.3 The Law of Tolerance
8.7.4 Ecological Adaptation in Hydrophytes
8.7.5 Ecological Adaptation in Mesophytes
8.7.6 Ecological Adaptations in Xerophytes
8.7.7 Ecological Adaptations in Halophytes
8.7.8 Ecological Adaptation in Animals
8.8 Let Us Sum Up
8.9 Key Words
8.10 References and Suggested Further Readings
8.11 Answers to Check Your Progress

8.1 INTRODUCTION
The gradual change in species composition and processes of communities over
time is known as ecological succession or community development.
Understanding the process, rates and pattern of ecological succession is important
for the management of ecosystems and for understanding vegetation potential
and dynamic changes in the landscapes. From the stand point of time, ecosystem
and community changes can be divided into two types:
i. Changes occurring over geological time scale (million years), and
ii. Those occurring over medium time scale, say in 1-1000 years.
Community changes occurring over geological time period are called Palaeo-
ecological changes. These changes are synthesized on the basis of fossil records,
e.g., leaves, twigs, cones, pollens and seeds. For example, fossil evidences indicate
that in Rajasthan desert of India, the vegetation during the tertiary period consisted
largely of trees species ascribed to humid environment. Later on, in response to
drier climate the desert plants prevailed in this area. In this unit, we would be
156
discussing the causes of succession, types of succession, processes of succession Ecological Succession and
Climax
and the ecological adaptation in hydrophytes, mesophytes, halophytes and animals.

8.2 OBJECTIVES
After studying this unit, you should be able to:
• explain the causes of succession;
• describe the types of succession;
• explain the processes of succession; and
• explain the ecological adaptation in hydrophytes, mesophytes, halophytes and
animals.

8.3 CAUSES AND TRENDS OF SUCCESSION

The causes of succession are as follows:
i. Initial/Initiating causes: These are climatic as well as biotic. The factors include
erosion and deposition, wind, fire, activities of organisms, etc. These causes
produce the bare areas or destroy the existing population in the area.
ii. Ecesis/Continuing causes: These are the processes such as migration, ecesis,
aggregation, competition, reaction, etc., which cause successive waves of
populations as a result of changes, chiefly in the edaphic features of the
area.
iii. Stabilizing causes: These cause the stabilization of the community. According
to Clements, climate of the area is the chief cause of stabilization, other
factors are of secondary value.

Trends in Succession
i. Change in species composition (i.e., kinds of plants change continuously
with succession, the number of species often increase).
ii. Change in variety or diversity (the diversity of species tends to increase
with succession).
iii. Progressive increase in biomass (the amount of both living and dead organic
matter).
iv. Shift in community metabolism (a decrease in community production and
corresponding increase in community respiration). In a young pond, P/
R>1(P=production, R=respiration), whereas, in a stable pond (heterotrophic
succession) P/R=1, P/R<1.

8.4 DIFFERENT KINDS OF SUCCESSIONS

1. Primary succession: If succession proceeds from a primary bare area or
primitive substratum which has not been changed physically by organisms,
it is called primary succession. The first group of plants establishing there
are known as pioneers. The series of development stages are called prisere.
For example, succession on a bare rock (lithosere).
2. Secondary succession: If succession starts in a secondary area previously
colonized, but has been cleared off, it is called secondary succession. The 157
Fundamentals of Ecology series of developmental stages are called subseres. In subseres, the substratum
contains already formed soil, humus and may contain seeds. The rate of change
is more rapid and the time required for the completion of sere is much shorter
than in primary succession. For example, succession in a forest area where
vegetation has been devastated by natural catastrophes.
3. Autotrophic succession: Succession characterized by early and continued
dominance of autotrophic organisms like green plants. It begins in a
predominantly inorganic environment and the energy flow is maintained
indefinitely. There is gradual increase in the organic matter content supported
by energy flow.
4. Heterotrophic succession: Succession characterized by an early dominance
of heterotrophs, i.e., fungi, bacteria and animals. It begins in a predominantly
organic environment, and there is a progressive decline in the energy content.
5. Induced succession: The climax community has low productivity as
compared to initial communities. In a climax community respiration almost
balances the production of organic matter. Therefore, very little is left for
man to harvest.
6. Allogenic succession: Allogenic succession is due to major environmental
changes beyond the control of the indigenous organisms. Dust bowls, winds,
dry periods change the pattern of vegetation. The habitat is changed by the
action of outside factors like change in climate, leaching of soil nutrients,
increase in salt concentration of the soil and deposition of salt or sand.
7. Autogenic succession: Succession resulting from the resident population
altering its own environment. For example, plants of a developmental state
produce changes in the habitat initially to favor their growth but the changes
go on beyond the optimum so that the habitat becomes unsuitable for them.
It paves the way for the growth of another type of plant community.
8. Retrogressive succession: At times a climax vegetation may deteriorate
and get replaced by a community of an earlier stage of succession due to
destructive effects of organisms. Sometimes the development of the disturbed
communities does not occur and the process of successive instead of
progressive becomes retrogressive, e.g., forest may change to shrubby or
grassland.
9. Deflected succession: A succession in which the vegetation does not pass
through the normal stages of development but either adds or replaces a
successional type.
10. Serule (Microsere): It refers to the miniature succession of microorganisms
like fungi, bacteria, actinomycetes, etc., that occurs within a microhabitat
like fallen logs of decaying wood, tree, bark, etc. Serule is heterotrophic in
nature and begins on substratum rich in organic matter.

8.5 GENERAL PROCESS OF SUCCESSION

1. Nudation: Development of a bare area without any form of life. The area
may develop due to several causes such as landslide, erosion, deposition or
other catastrophic agency. The causes of nudation may be:
158
 Topographic: Soil erosion, landslide, volcanic activity, earthquake, etc. Ecological Succession and
Climax
Climatic: Glaciers, dry period, hails and storm, frost, fire, etc.
Biotic: Destruction of forest due to industrialization, agricultural expansion,
urbanization, diseases due to fungi, bacteria, etc.
2. Invasion: Successful establishment of a species in a bare area. The species
actually reaches this new site from any other area. It involves following
successive stages
i. Migration (dispersal): The seeds, spores, or other propagules of the
species reach the bare area. The agencies which bring about dispersal
are wind, water, animal including man.
ii. Ecesis (establishment): After reaching the area, the process of
successful establishment of the species, as a result of adjustment with
the conditions prevailing there, is known as ecesis. This process is
dependent upon the climatic, edaphic and biotic factors. The success
of plant depends upon the climatic, edaphic and biotic factors. In plants,
after migration, seeds or propagules germinate, seedlings grow, and
adults start to reproduce. Only a few of them are capable of doing this
under primitive harsh conditions and thus most of them disappear. As a
result of ecesis, the individuals of species become established in the
area.
iii. Aggregation: As a result of reproduction, the individuals of the species
increase in number, and they come close to each other.
3. Competition and coaction: This phenomenon involve struggle for existence
between two or more individuals growing in an area, that makes successive
demands, that are similar in nature, on the soil. The struggle is usually
between two individuals of same kind that have similar demands such as
space, nutrients, water, light, etc. Competition can either be interspecific
(between two different species) or intraspecific (within the individuals of
same species). As a result of competition, the weak individuals are eliminated
and the stronger ones are retained. The dead bodies of the eliminated plants
and animals’ decay and form humus enriching the soil.
4. Reaction: This is the most important stage in succession. The mechanism
of the modification of the environment through the influence of living
organisms on it, is known as reaction. As a result of reactions, changes take
place in soil, water, light conditions, temperature, etc., of the environment.
For example, plants change the structure and texture of soil in course of
time by addition of humus into the soil. Due to all these, environment is
modified, becoming unsuitable for the existing community which sooner or
later is replaced by another community.
5. Climax community: The reactions of individuals collectively have a wider
effect on the environment. The local climate is actually changed if the climax
vegetation is forest. The reactions keep the vegetation in active state till the
climax community is formed. Although strictly speaking vegetation can never
be called as stable. Yet at maturity the community being mesic, further mesic
changes being less possible the climax community can be called as relatively
stable. Thus, we find the development of community living in harmony
within the environment which has been the result of its reactions.
159
Fundamentals of Ecology 8.5.1 Hydrosere
Plant successions which begin in ponds, lakes, marshes, or elsewhere in water,
are termed hydrarch and different stages are called as hydrosere. The water is
deep in the middle and becomes progressively shallow towards the bank.
1. Pioneer stage: This is characterized by a bottom barren of plant life. The
pioneers include phytoplankton. This consists of microscopic algae, bacteria,
diatoms and protozoa. This phytoplankton after death settles to the bottom.
The soils are very much reduced with a pH value of not more than 5.
2. Submerged stage: This stage is found where the water is less than 20 feet
deep. The plants are entirely submerged. Prominent submerged plants include
pond weeds (Potamogeton), hornwort (Ceratophyllum), eelgrass
(Vallisneria), water weed (Elodea), Hydrilla, bladderwort (Utricularia),
Chara and Ranunculus. These are all rooted plants. When these plants die,
their remains sink to the bottom where they become humus. The humus binds
the soft muddy soil. These plants also help in depositing soil particles at the
bottom. As a result of these reaction the water becomes shallow and the
habitat becomes unsuitable for submerged plants, which in turn are replaced
by floating plants.
3. Floating stage: This stage is present where the water is only 6-8 feet, deep.
This stage includes rooted plants with floating leaves like Nyphaea (water
lily), Nelumbium, Limnanthemum, Aponogeton, Monocharia, Trap and free-
floating plants like Pistia, Azolla, Lemna, Spirodella, Wolffia, Eichhornia,
etc. The water level by now becomes very much decreased, making the pond
shallower. By their death and decay humus is formed which results in the
higher concentration of salts and organic matter and ultimately the water
becomes unsuitable for these floating plants which are then replaced by reed
swamp plants.
4. Reed-swamp stage: This stage also called as amphibious stage occurs where
the water is 1-4 feet deep and includes the plants which are partly submerged,
with their roots at the bottom and their foliage raised above the surface of
water. The important plants consisting this stage include cattail (Typha),
bulrush (Scirpus), reed grass (Phragmites), arrow head (Sagittaria), Rumex
etc. These plants cut off the light from the floating plants and in this way
make the water still shallower by settling down the sedimentary materials
washed into the lake and by very rapid accumulations of humus. This changed
habitat becomes highly suitable for the growth of plants of next seral stage
i.e., marsh-meadow stage.
5. Marsh-meadow stage: This stage includes hydrophytes or water loving
plants. The substratum at this stage is hardly covered by 1-2 inches of water,
so to say the soil becomes marshy. This is now invaded by numerous species
of sedge. Juncus, carice (Carex), spike rush (Eleocharis), Polygonum, etc.
Many species of herbs like mint (Mentha), marsh marigold (Caltha), bell
flower (Campanula), etc., also occur intermixed with sedges. All these
hydrophytes react upon the habitat, raise the surface by binding water carried
and wind-borne soil, accumulate plant debris, and transpire enormous
quantities of water. This makes the soil more suitable for the mesophytes
and terrestrial plants. Under these circumstances, hydrophytes cannot live
long, they migrate inward giving room for grasses and woody plants.
160
6. Woodland stage: If the climate is dry then a grassland develops but under Ecological Succession and
Climax
moist climate a woodland is formed containing certain shrubs and small
tress. This stage is characterized by the plants that can tolerate water-logged
soil around their roots. Shrubby willow (Salix), dog woods (Cornus), button
bush (Cephalanthus), alder (Alnus), cotton wood (Populus), tree willows
etc., are the plant species of the woodland stage. These plants by their reaction
make the soil unsuitable for themselves and more suitable for shade enduring
herbs which grow the trees and shrubs.

7. Climax forest: This represents the final stage of hydrarch. It includes mixed
forest of alder (Alnus), willow (Salix), cottonwood (Populus), elm (Ulmus),
ash tree (Fraxinus), oak (Quercus), etc. After a few generations a pure forest
oaks or hickories may develop.

Fig 8.1: Various stages of Hydrosere

8.5.2 Lithosere (Xerarch)

Xerosere is the sequence of successional stage which occur on bare areas deficient
in water. Succession on a bare rock is as follows:
1. Pioneer stage (Crustose lichen stage): The rocky habitat is extremely xeric
and hostile. There is no water as the substratum does not absorb rain water.
There is no nutrient holding mechanism. When exposed to sun, the surface
temperature goes very high. In such a habitat, only the crustose lichens can
become pioneer colonies that have ability to bear high degree of desiccation
and temperature extremes. These lichens reach the bare rock through wind
borne soredia, lichen fragments and spores. The lichens produce carbonic
acid which has a corroding effect on rock matter:
161
Fundamentals of Ecology Co2 + H2o→H2Co3
Generally, species of Rhizocarpon, Rinodena, Lecidea and Lecanora
establish themselves on the bare rocks.
2. Foliose Lichen stage: Foliose lichens i.e., those attached to the substratum
at a single point or along a single margin appear as soon as a little soil has
accumulated on the non-weathered portion of rock and in depressions or
other slightly less exposed situations. They slowly replace the crustose form.
These expanding leaf-like thalli may completely over shadow the crustose
lichens causing the crustose species to die and decay. Above the foliaceous
invaders water has better chance to collect and to be absorbed. Evaporation
is greatly decreased. Wind and water borne lichen fragments and dust particle
lodge and humus is more rapidly accumulated because of its less rapid
oxidation. Acid produced by living and decaying plants are constantly eating
further into the rocks. Indeed, it is possible that change from crustose to
foliose lichen is a change of habitat.
After the crustose give away to foliose species such as Dermatocarpon,
Parmelia, Umbilicaria, a new type of invaders appears.
3. Moss stage: As soon as sufficient amount of soil has accumulated in the
minute crevices and depression xerophytic mosses begin to appear. These
are common species of Gerimmia, Polytrichum and Tortula. They may have
migrated long distances by wind-blown spores that are caught in minute
amount of soil and along foliose lichens and germinate there. Their rhizoids
compete with those of foliose lichens for water and nutrients. The erect stems
of mosses often exceeded the lichens in heights. The power of withstanding
desiccation is almost as marked along these pioneers as among the lichens.
These are the most exacting foliose species that may occur simultaneously or
indexed. The mosses may sometimes precede foliose lichens.
Soil rapidly accumulates among the erect stems as the plant die below and
continue to grow above and build up the substratum and constantly increase
their area. The depth of the soil under the cushion like mat is often one inch
or even more. The crustose lichens like Cladonia grow along with mosses.
The mosses form thick mats and play a significant role in building up thick
substratum of soil. Their continuous growth, death and decay for several
years builds up a good soil which is quite fit for the growth of herbaceous
flora.
4. Herbaceous Stage: The soil forming and soil holding reaction of mosses
are so pronounced that the seeds of some xerophytic herbs especially short-
lived annuals are soon able to germinate and grow to maturity. They grow
slowly and exhibit stunted growth because the soil is yet not very favourable
and lacks nutrients. Drought conditions also prevail. The roots of these xeric
herbs continue to grow and corrode the rocks. Their dead remains enrich the
soil further and more humus collects.
Depending upon the plants growing in surrounding communities the invading
herbs are Potentilla, Solidago and Saxifraga. Their growth makes the
conditions less dry. Bacteria, fungi and microfauna appear along with grasses.
Their death and decay further add to the soil layers.

162
5. Shrub stage: Woody shrubs like Rhus glabra, or Rubus and Sassafras invade Ecological Succession and
Climax
these areas. Their shade makes the growth of herbs impossible and thus they
disappear. The humidity increases and wind velocity is decreased. The
addition of organic matter to the soil increases water holding capacity of soil, its
texture and structure is changed so that the seeds of trees find suitable place for
growth.
6. Climax forest: The tress which make their appearance are dwarf sized,
xeric and grow separated apart. They are however followed by mesophytes
as the climate becomes more mesic. Quercus, Tilia are the trees which find
place in climax communities.

Fig 8.2 Various stages of Lithosere

Check Your Progress 1

Note: i) Use the space given below for your answers.
ii) Check your answers with those given at the end of the unit.
1. Explain the causes of succession.
......................................................................................................................
......................................................................................................................
......................................................................................................................
2. State the differences between primary and secondary succession.
......................................................................................................................
......................................................................................................................
......................................................................................................................

163
Fundamentals of Ecology 3. Explain the stages in hydrosere.
......................................................................................................................
......................................................................................................................
......................................................................................................................
4. Describe the sequential stages in lithosere.
......................................................................................................................
......................................................................................................................
......................................................................................................................

8.6 CLIMAX COMMUNITY

The end product of succession after seral communities is the climax community,
and it is a relatively stable community. It is generally believed that once the climax
is attained the community does not change at all, but this is not exactly so, climax
communities may also be changed by aging, storm, disease and by other biotic
and abiotic factors.
Hanson and Churchill (1961) characterized a climax community as:
i. The climax community is in steady state regarding its productivity, structure
and population.
ii. There is a diversity, stability, and homogeneity of the species populations
within and between the stand of the same climax community.
iii. Each stand is self-maintaining and long persisting.
iv. Replacement and fluctuation changes are operative on a continuous basis
within the climax, while all environmental factors determine its composition
and population so that there is a mosaic of climax types corresponding to the
mosaic of habitats.

8.6.1 Monoclimax Concept

This concept was advanced by an American plant ecologist E.E. Clements in
1916. According to him, in a climatic region, only one true climax community is
possible which is mainly controlled by the climatic factors. Such a climax is,
therefore, climatic climax. It is not affected by soil or topography. Edaphically
controlled stable communities are exceptions, and are not true climax.
In order to account for a variety of more or less stable communities but different
from regional climatic climax, Clements proposed four other terms:
• Subclimax- succession gets arrested at a stage which persists for a long time in
response to physiological or edaphic factors, before being replaced by the climatic
climax;
• Disclimax- the vegetation replacing the true climax as a result of some persistent
biotic disturbance in the environment, e.g., the grasslands in the Gangetic Plains
which occur due to grazing in a deciduous forest climax;
• Preclimax- slightly drier localities in a given climatic climax area may have a self-
perpetuating community different from the climatic climax, e.g., some of the pine
164
 forests in the Himalaya; Ecological Succession and
Climax
• Post-climax- slightly “better moisture” areas in the climatic climax region
may support a different yet self-perpetuating community, e.g., Terminalia
arjuna community growing near river banks in a dry deciduous forest climax.

The monoclimax concept has been severely criticized on the ground that in the
concept regionally prevailing undisturbed vegetation occupying the largest part
of the land surface was regarded as real climax and other stabilized plant
communities in the same area were recognized as subclimaxes which only
theoretically could be replaced by the climax.

8.6.2 Polyclimax Concept

This concept was advocated by Whittaker (1953) and supported by Tansley.
According to this concept a climax reflects not only the climatic factors but also
other factors of the environmental complex, viz., edaphic, biotic, etc. The climax
can be of several other kinds which are different from the climatic climax of the
area. Climate is not the only factor that determines the climax in a large area.
Other factors like edaphic, topographic, and biotic factors are equally important
so that edaphic, topographic and biotic climaxes may occur in localized regions
within the same climatic zone.

Tansley (1920) recognized the existence of a number of climax communities,

forming a mosaic corresponding to the mosaic of habitats and suggested a
polyclimax theory. However, this view too identified a climatic climax community
that is the stable and most extensive in different habitats for an area. Tansley
recognized following climax types:
i. Climatic climax: Climax under normal conditions of climate, soil and
topography and no disturbance.
ii. Edaphic climax: Substrate peculiarities are well pronounced to produce
self-perpetuating vegetation, which is different from the climatic climax of
the area.
iii. Topographic climax: Changes in the topography enough to cause variant
microclimates, each giving rise to self-perpetuating vegetation.
iv. Fire climax: Recurrent burning of vegetation eliminates fire-sensitive
species and self-perpetuating vegetation develops.
v. Zootic climax: Self-perpetuating community in response to zoological
factors e.g., grazing gives rise to zootic climax of grassland.

Climatic, edaphic and topographic climaxes are primary climaxes, while fire
and zootic climaxes are secondary disclimax.

8.7 ECOLOGICAL ADAPTATIONS

Adaptations are structural and functional characteristics which organisms have
developed during the course of evolution enabling them to survive and reproduce
in a particular environment. The term adaptation has therefore an evolutionary
significance.
165
Fundamentals of Ecology 8.7.1 Tolerance Ranges
Under natural conditions, each organism possesses a range of tolerance to
variations in its physical and chemical environment. The organism responds to
variations in environmental conditions in terms of their growth, reproduction and
distribution. Any of the physical or chemical components of the environment that
may inhibit the growth of living organisms, through either its lack or excess, is
said to be limiting condition or limiting factor. For example, low temperature
limits plant growth at higher elevation, water availability limits plant growth in
deserts, and changing levels of salinity affect organisms in estuaries, whereas
low phosphorus availability limits the growth of phytoplankton in deep lakes.

8.7.2 The Liebig’s Law of Minimum

The Liebig’s law of minimum states that “growth of plants is dependent on the
amount of food stuff which is present to it in minimum quantity”. Limiting factors
most often affecting terrestrial organisms are moisture and temperature; those
affecting aquatic plants and animals are mainly oxygen and light. The law of
minimum and the law of limiting factors operate within steady state conditions,
when inflows balance outflows of energy and materials.

8.7.3 The Law of Tolerance

However, not only too little of something is a limiting factor, but also too much
may limit the growth and distribution of an organism. The concept of the effect of
maximum as well as minimum has been incorporated in to the law of tolerance by
Victor E. Shelford (1913). According to this law, for most environmental factors,
such as temperature, light and moisture have an ecological minimum (‘too little’)
and an ecological maximum (‘too much’). The range between these two conditions
represents limits of tolerance.

Fig. 8.3 The bell-shaped curve shows the response of an organism to a range of single
166 environmental variable.
The tolerance curve is a measure of organism’s fitness and survival. A bell shape Ecological Succession and
Climax
results when measures of survival or fitness are plotted against the environmental
gradient (fig. 8.3). Those environmental conditions under which an organism grows
and reproduce maximally comprise the optimum range. The organisms show wide
distribution due to wide range of tolerance for all factors and restricted distribution
if the tolerance range for one or more than one factor is narrow. At limiting levels,
reproduction of an organism is most critical, whereas factor interaction influences
the fitness of an organism.

8.7.4 Ecological Adaptation in Hydrophytes

The plants which remain permanently immersed in water are hydrophytes. They
may be completely submerged or partly submerged and show the presence of
aerenchyma (large air spaces) in the leaves and petioles. Aerenchyma tissue helps
to store oxygen produced during photosynthesis and permits its free diffusion to
other parts, including roots which may be in aerobic soil. These tissues also
impact buoyancy to the plants.
Characteristic adaptation features of hydrophytes:
• Sponginess of the tissue because of the presence of large air chambers or cavities
(lacunae).
• Roots may be entirely absent (e.g., free-floating Wolffia, submerged
Ceratophyllum or they are poorly developed, without root caps, root hairs
and branches. Thus, they do not require too many stomata or a thick cuticle to
regulate the water loss.
• Cuticle in leaves is either absent, as in submerged plants like Potamogeton,
or is poorly developed. Similarly, stomata are absent or are non-functional
in some submerged plants, they are present only on the upper leaf surface in
floating plants and mainly in emergent their presence can be seen on both
leaf surface. To cope up with low light availability, leaves may be finely
dissected and chloroplasts may be confined to outer cell layers.
• Tissue differentiation is poor, including those of vascular bundles, and mechanical
tissue, such as sclerenchyma are generally altogether absent.
• Leaves in submerged species are thin, linear and finely dissected. The finely
dissected leaf was thought to be a useful adaptation to high-speed currents
which may tear the entire leaves to pieces.
• Most of the hydrophytes are characterized by a rapid vegetative growth and have
the potential to reproduce by asexual means. They can simultaneously reproduce
by rhizomes, tubers and turions as well as seeds produced from asexual
reproduction. E.g., Potemogeton pectinatus.

8.7.5 Ecological Adaptation in Mesophytes

Mesophytes are land plants that grow under normal conditions. Mesophytes grow in
environments where it is neither too dry nor too moist. They grow in areas where the
atmospheric temperature and the relative humidity are also normal. Examples of
mesophytes are wheat, pea, tomato, mango, guava, etc. The roots of these plants are
tightly embedded in the soil and the leaves are large and broad while the stem is erect.
Mesophytes make up the largest ecological group of terrestrial plants. Morphologically
167
Fundamentals of Ecology these plants do not special modifications. They have broad, flat and green leaves.
They have extensive fibrous root system to absorb water. The characteristic
adaptation features of mesophytes:
• The leaves of mesophytes are large and rarely small, broad, thin and are in a
variety of shapes. The leaves are oriented horizontally without any waxy
coating or hairs.
• In the aerial plant parts, the cuticle is well developed.
• The epidermis of the plant is very well-developed, the epidermis is without any
hair or waxy coating and the cells are without chloroplasts.
• The stems of mesophytes are aerial and are branched freely.
• Stomata are present generally on both surfaces of the leaves.
• Guard cells the opening and closing of the stomata which is usually a frequent
movement.
• The mesophyll of the leaves is differentiated into palisade and spongy parenchyma
with many intercellular spaces.
• The vascular tissues and the mechanical tissues are well-developed and well-
differentiated.
• The osmotic pressure of the cells is low and if less water provided, the plants will
quickly droop.
• During the summer midday temporary wilting is quite common in these plants.
• The mesophytes may also shed their leaves during the dry months.
• The evergreen plants show xeromorphic characters to overcome the drought
periods.

8.7.6 Ecological Adaptations in Xerophytes

Desert plants are adapted to cope with dry conditions of soil and high temperature.
The plants evading the dry conditions are known as ephemerals. With the onset of
rainfall, these plants quickly pass-through various growth phases, germination,
flowering and seed dispersal in only a few days. The plants become dominant
again with the advent of dry conditions. Photosynthesis and respiration are inhibited
by high temperature; photosynthesis is more sensitive to high temperature than
respiration. Temperature compensation point is defined as the temperature at which
CO2 release during respiration equals to that of CO2 fixed during photosynthesis.
Plants produce large number of low molecular mass heat shock proteins
(Chaperons) that function to facilitate protein folding and membrane fluidity during
high temperature stress episodes. The characteristic features of xerophytes:
• Leaves are small and compact, with high ratio of volume to surface. They
have thick blades, small sunken and dense stomata on lower surface, dense
covering of hair (pubescence), a thick cuticle, heavily lignified epidermal
cells, strongly developed palisade mesophyll and less inter-cellular spaces.
• In many types of dryland shrubs, leaves are replaced by thorns, which almost
have no transpiration. Hair have been considered to decrease transpiration,
but there is also evidence to indicate that transpiration was reduced with the
removal of hair. It is possible that hair of xerophytic plants serves other
purpose, such as prevention of insect attack and cooling of leaf by increasing
168 total radiating surface.
• Plant shows adaptations in terms of water storage in roots (e.g., Asparagus) Ecological Succession and
Climax
and stems (succulents), deep penetration of roots so that deep water-soil is
made available.
Check Your Progress 2
Note: i) Use the space given below for your answers.
ii) Check your answers with those given at the end of the unit.
1. Explain the polyclimax concept.
......................................................................................................................
......................................................................................................................
......................................................................................................................
......................................................................................................................
......................................................................................................................
......................................................................................................................
2. Explain the ecological adaptation in hydrophytes.
......................................................................................................................
......................................................................................................................
......................................................................................................................
......................................................................................................................
......................................................................................................................
......................................................................................................................
3. Describe the characteristic adaptation features of mesophytes.
......................................................................................................................
......................................................................................................................
......................................................................................................................
......................................................................................................................
......................................................................................................................

8.7.7 Ecological Adaptation in Halophytes

Halophytes are plants of saline environments, which are adapted to high
concentration of salt in soil or water. Tidal marshes and coastal dunes, mangroves
and saline soils support halophytes. In wet and marshy conditions of tropical
deltas and along ocean edges mangroves are found. Few species of mangroves
can excrete salt through salt glands on the leaf. Some plants can exclude salts
from the roots through pumping excess salt back to the soil for coping with condition
of high salt conditions and osmotic potential many mangroves have high level of
organic solutes such as proline, glycine betaine, amino acid mannitol and sorbitol.
Mangroves have developed adaptations e.g., pneumatophores, prop and stilt roots
and vivipary (seed germinate while on the tree) to cope with high concentration
of salt.
169
Fundamentals of Ecology In Avicennia, presence of pneumatophores helps to take up oxygen and transport it
to main roots. The pneumatophores are root extension that grow out of water and
up in the air. These roots have lenticels and spongy tissues that allows oxygen to
diffuse in them. The red mangroves (Rhizophora) seed germinates on the tree and
then drops into water. The wave transports the seed into shallow water so that
they can penetrate the mud. Prop and stilt roots in many species of mangrove give
support to the plant.

8.7.8 Ecological Adaptation in Animals

Carnivorous and herbivorous animals have adaptations to eat a certain kind of
food. Some animals have adaptations to avoid being eaten by the predators; others
have behavioral adaptations to attract a mate. The males of some animals
(particularly the plumage of the birds) have bright coloration, which gives
advantage in sexual selection and mate attraction. However, a majority of animal
adaptations to environmental variations and stress conditions are physiological
and behavioral. Camouflage in some animals is a common example of adaptation.
Some insects, reptiles and mammals have markings on their bodies which make it
difficult to distinguish them from shadows and branches or from other members of
the group. In very cold or dry environments, animals are not able to function
normally. Under such conditions, the species that are not capable of migration,
shift to a physiological dormant state. Hibernation is spending winter in dormant
condition, whereas aestivation is spending the dry-hot period in an inactive state.

Table 8.1: Behavioral strategies of adaptations in animals

Types Examples
Hibernation Northern ground squirrels
Aestivation Ground squirrels in south-west deserts
Cryptic appearance Leaf like grasshopper
Batesian mimicry The Vasp (Vespula vulgaris) and two
mimics, the wasp beetle (Cltytus arietis),
and the hoverfly ( Syrphus ribesii )
Mullerian mimicry Monarch butterfly and the mimic Queen
butterfly
Echolocation Horseshoe bat, dolphins
Auditory location system Barn owls
Electromagnetic sensing DogfishElectric ray

8.8 LET US SUM UP

The gradual change in species composition and processes of communities over
time is known as ecological succession or community development. The causes
of succession are initial/initiating causes, ecesis/continuing causes, and stabilizing
causes. We have discussed different kinds of successions which include primary
succession, secondary succession, autotrophic succession, heterotrophic
succession, induced succession, allogenic succession, autogenic succession,
retrogressive succession and deflected succession. In this unit, we have discussed
the processes of succession and the ecological adaptation in hydrophytes,
170 mesophytes, halophytes and animals.
 Ecological Succession and
8.9 KEY WORDS Climax

Invasive Species: A non-native species that is introduced into a new habitat, and
that often adversely affects numerous species in the new habitat.
Keystone Species: A species that has a much greater impact on the community
than would be expected by measuring its abundance or biomass.
Primary Succession: The changes that occur to a habitat following a severe disturbance
that kills virtually all organisms in that habitat.
Secondary Succession: The pattern of recolonization following a disturbance in which
only some of the organisms are killed within a habitat.

8.10 SUGGESTED FURTHER READING/

REFERENCES
Molles, M.C., and Molles, M.S. (2018). Ecology: Concepts and applications.
McGraw-Hill Education, United Kingdom.
Odum, E.P. and Barrett, G.W. (2005). Fundamentals of Ecology. 5th edition.
Thomson Brooks/Cole.
Sharma, P.D. (2007). Ecology and Environment. Tenth revised edition. Rastogi
Publications, Meerut.
Singer, F.D. (2016). Ecology in Action. Cambridge University Press, United
Kingdom.
Singh, J.S., Singh, S.P. and Gupta, S.R. (2017). Ecology environmental science
and conservation. S. Chand Publishing house.

8.11 ANSWERS TO CHECK YOUR PROGRESS

Check Your Progress 1
1. Refer section 8.3
2. Refer section 8.3
3. Refer section 8.5.1
4. Refer section 8.5.2
Check Your Progress 2
1. Refer section 8.6.2
2. Refer section 8.7.4
3. Refer section 8.7.5

171