Environmental Sustainability

BLOCK 2
FUNDAMENTALS OF ECOLOGY

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Fundamentals of Ecology
BLOCK 2 FUNDAMENTALS OF ECOLOGY

Ecology is the study of the environment in which an organism lives. Ecology is

the study of the interactions of organisms with their surrounding environment.
Every living species whether plant, animal or microbe along with their physical
and chemical environment could be part of ecological studies. The study of
ecology is of quite significance which can help in better understanding of
consequences of human interventions at various levels and can also suggest
suitable solutions for the unsustainable changes occurring in nature. The study
of ecology includes levels of organization in ecology, environmental factors and
different types of biotic relationships. Ecology is studied at various levels of
organization from species level to biome. The population is studied as a collective
group of organisms of the same species which occupy a particular space at a
particular time. The population parameters include density, natality, mortality,
age distribution, biotic potential, dispersion, and growth form. As regards the
community organisation, a group of populations of different species interact in a
given area. Various interspecific and intraspecific interactions occur among
organisms. The world’s major communities, called biomes are classified according
to the predominant vegetation and characterized by adaptations of organisms to
that particular environment.

Unit 5 “Ecology” discusses the concepts in ecology, its scope, levels of

organization in ecology, environmental factors and different types of biotic
relationships.

Unit 6 “Population Parameters and Regulation” discusses the parameters of

population estimation, demographic techniques, various approaches employed
for population regulation and evolutionary implications of natural regulation.

Unit 7 “Community Organisation and Interaction among Organisms” deals with

the processes that affect community organisation and also various interspecific
and intraspecific interactions among organisms.

Unit 8 “Ecological Succession and Climax” discusses the causes of succession,

types of succession, processes of succession and the ecological adaptation in
hydrophytes, mesophytes, halophytes and animals.

Unit 9 “Major Biomes of the World” delves on the major biomes of the world and
characteristic features of the aquatic and terrestrial biomes.

We hope that after studying this block, you will acquire an understanding of the
concept and scope of ecology, biotic relationships, population parameters and
community organization.

Wishing you success in this endeavour!

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 Ecology
UNIT 5 ECOLOGY

Structure
5.1 Introduction
5.2 Objectives
5.3 Basic Concepts in Ecology
5.4 Scope of Ecology
5.5 Levels of Organization in Ecology
5.6 Applied Ecology
5.7 Environmental Factors
5.7.1 Climatic Factors
5.7.2 Topographic Factors
5.7.3 Edaphic Factors
5.7.4 Biotic Factors
5.8 Biotic Relationships
5.9 Let Us Sum Up
5.10 Key Words
5.11 Suggested Further Reading/References
5.12 Answers to Check Your Progress

5.1 INTRODUCTION
Ecology word has its origin in two Greek words; ‘Oikos’= house and ‘Logos’= to
study. Ecology is the study of environment in which an organism lives. Reiter (1868)
has coined the term for the first time. Haeckel (1886) first used the word ‘Ecology’
and defined its meaning as relations of organisms to their surrounding outer world and
to organic and inorganic conditions of existence which now could be called as biotic
and abiotic factors to which organisms interact with. Among so many definitions of
ecology available, the most common could be considered is the study of interactions of
organisms with their surrounding environment. Every living species whether plant, animal
or microbe along with their physical and chemical environment could be part of
ecological studies.

Humans are affecting environment in myriad ways. In this regard, study of ecology is of
quite significance which can help in better understanding of consequences of human
interventions at various levels and can also suggest suitable solutions for the unsustainable
changes occurring in the nature. In this unit, we would be discussing the concepts in
ecology, its scope, levels of organization in ecology, environmental factors and different
types of biotic relationships.

5.2 OBJECTIVES
After studying this unit, you should be able to:
• explain the basic concepts of ecology;
• describe the various levels of organization in ecology;
• explain different environmental factors; and 91
Fundamentals of Ecology • describe different types of biotic relationships.

5.3 BASIC CONCEPTS IN ECOLOGY

According to Misra (1991), there are four basic concepts in ecology. These are
explained briefly:

1. Holism: This term was coined by J.C. Smuts in 1926 in the book ‘Holism
and Evolution’. W. Ophuls (1974) suggested that Holism is the base of
Ecology. The various units of ecology from lower to higher levels are as
follows: individual, population, community, ecosystem, biome, biosphere.
Holism suggests that each unit is built as a whole by interactions of lower-
level wholes into higher level wholes.

2. Ecosystem: This term was coined by A.G. Tansley (1935). Ecosystem forms
the structural and functional unit of nature where both biotic and abiotic
components interact and work in a complex.

3. Succession: It is a long-term process where an ecosystem is converted into

a higher biomass rich and trophically diverse stable ecosystem. This occurs
due to change in environment which influences development of organisms
and in turn, organisms also affect the environment by their activities. Due to
this dynamism in both environment and organisms, process of succession
starts and keeps going on till a stable community is developed. The final
stabilized stage is known as climax.

4. Conservation: It is the necessity for living in a sustainable manner. Various

factors of ecosystem interact with each other and regulate themselves in
order to keep the ecosystem healthy for long. Due to degrading environment,
pollution, climate change and some other problems, conservation strategies
are required for their effective solution and maintenance of a healthy planet.

5.4 SCOPE OF ECOLOGY

Ecology is a multi-disciplinary science focusing on diverse spatial and temporal
scales. Study of ecology could be done at various levels starting from nucleic
acids and proteins in biochemistry and molecular biology, cells in cellular biology,
organisms in botany, zoology, microbiology with details in morphology,
physiology, taxonomy, genetics, evolution, developmental biology, toxicology,
etc. and other sub-division levels such as phycology, mycology, bacteriology,
virology, ornithology, etc. Ecology could also be studied at the level of population,
community and ecosystem. Human ecology itself is a vast field of investigation
and so is “Biome Ecology”.

Ecology not only involves biotic components but also the abiotic components
such as various environmental factors, thus study of geography, geology,
meteorology, pedology. geochemistry and paleoscience are also under scope of ecology.
Study of an organism with respect to its environment such as changes in behavior or
changes at genetic level are part of ecology. An ecologist must also have knowledge
about mathematics, statistics, physics, chemistry and computational biology. Further,
applied ecology is studied along with theoretical ecology to solve various
environmental problems of an area. Thus, Ecology is rightly considered as a
92 “Holistic Science”.
 Ecology
5.5 LEVELS OF ORGANIZATION IN ECOLOGY
The ecology could be studied at various levels of organization.
Species
John Ray (1686) first time defined “Species” in his work ‘Historia Plantarum’.
According to Mayr (1982), a biological species is a group of reproducing natural
populations inhabiting a particular niche and incapable of effective breeding
with other such groups. Generally, organisms similar in morphological characters;
can breed inter se and produce fertile offspring are considered to belong to same
species. For example, plants of Azadirachta indica (Neem) could be fertilized
by pollen grains of Neem; Homo sapiens could breed with Homo sapiens and
produce fertile progeny, etc. Thus, they belong to rank of Species. But the
ecological species concept is mainly about ecological competition where members
of a species compete more with each other than with members of other species.

Population
It is a group of individuals of same species who inhabit a particular geographic
area. Each population has its own characteristics like natality, mortality, age
structure, growth dynamics etc. For example, population of a moth species in a
specific area, population of Neem plants in a given area, etc.

Community
It is a group of population of different species who are interacting in a given
area. For example, a grassland community will include all populations inhabiting
there such as plants, animals and microbes. Sometimes, in a narrow sense, terms
like algal community, bacterial communities are used. Biocoenosis term is also
used to denote community.

Ecosystem

In a specific area, all biotic communities interact with their physical environment
in such a manner that a self-supporting sustainable system is formed where there
is a defined flow of energy and material cycling in every trophic level. That
system is called as Ecosystem or Biogeocoenosis. Some examples of such self-
sufficient ecosystems are pond, lake, grassland, desert, forest etc. Thus, there are
two basic components of ecosystem; biotic and abiotic. Biotic components include
living species at various trophic levels of producer, consumer, decomposer which
are linked together in food chains and form complex food webs. Abiotic
components consist of various environmental factors including both climatic
and edaphic factors.

Biome
It is a large area where many ecosystems coexist. So, one biome may have many
units of ecosystems. The sharing of common climatic region by different
communities of plants, animals etc. is represented by a biome. As biotic
communities adapt in various manner to a particular climate, biomes are usually
defined by abiotic factors like climate, soil, vegetation, geology, etc. Thus, a
forest biome could be grouped into Tropical Rain Forest Biome, Tropical Dry
deciduous Forest Biome, Evergreen Forest biome, etc. Similarly, aquatic biomes
could be grouped into fresh water biomes of lakes, ponds, rivers, streams and
marine biomes of seas, oceans and estuaries. 93
Fundamentals of Ecology Biosphere
It is also known as “Ecosphere”. It includes entire living species of earth interacting
with its physical environment. This giant ecosystem is almost a self-sufficient largest
unit and known as “Biosphere”.
Autecology

In this, relation of an individual species is studied with its environment and details
about its external morphology, geographic distribution, life cycle, interaction with
various environmental factors is studied.
Synecology

It deals with group of organisms in form of population, community, biome or

ecosystem with their environment. Interaction among plants, animals, microbe
communities occur frequently, hence synecological approach is applied for
ecological studies.
Biosphere

Biome

Ecosystem

Communities

Population

Species

Cell

Genes
94 Fig. 5.1: Levels of organization in Ecology
 Ecology
5.6 APPLIED ECOLOGY
Theoretical ecology may give new insights for using ecology in solving basic
problems of management of natural resources, which is one of the important
applications of ecology. Both theoretical and applied ecology are linked with a
feedback loop and not by a unidirectional supply link. As humans have an important
role in modifying natural components, habitats, etc., it is therefore, necessary to
develop a framework to create balance between man and various natural
components by applying appropriate knowledge about ecology. Thus, appropriate
management strategies in fields of conservation biology, global change, ecosystem
restoration, protected area design and management, eco-toxicology, environmental
pollution, environmental policies, bio-monitoring and bio-indicators of
environmental quality and biodiversity and economics should be applied for
management of agro-ecosystem, forest, rangeland, wildlife (including game),
aquatic resources, landscape use, disturbance including fires and floods, biological
invasions, pests and weeds, epidemiology, environmental design, etc. Applied
ecology thus, relates various ecological theories and concepts in proper
management of environmental problems.

5.7 ENVIRONMENTAL FACTORS

The complex of things which surround an organism is known as environment.
There are various environmental or ecological factors which affect the life of an
organism. These include both living (biotic) and non-living (abiotic) factors.
Ecologists classify these factors in different ways. Mostly, these environmental
factors are categorized into four factors:
A) Climatic factors
B) Topographic factors
C) Edaphic factors
D) Biotic factors
These factors are discussed in brief in the following section:

5.7.1 Climatic Factors

Climatic factors mainly include light, temperature, rainfall, humidity and
atmospheric gases. These influence climate of any region in a combined manner
and make the climate specific for that region. We will discuss one by one.

Light
Light, one of the most important climatic factors is a source of energy for living
organisms. Sun is the biggest source of energy on earth and constantly gives heat
energy in form of solar radiations. Many of the physiological processes in plants
for example, photosynthesis, seed germination, flowering, etc. are affected by
light. Not only intensity of light but its wavelength also affects growth of plants.
Light also affects various physiological activities in animals for example,
reproduction, development, vision, metabolism, locomotion, etc.

There are some plants which grow best in direct sunlight and termed as Heliophytes
that is sun loving plants for example, Xanthium, Amaranthus, Salix, etc. There
are also some plants which grow best in low light intensity or shade loving plants 95
Fundamentals of Ecology for example, Acalypha, Fagus, Abies, etc. and known as Sciophytes.

Among the heliophytes, some plants can survive only in sunlight are known as
obligate heliophytes whereas some plants prefer sunlight for growth but can also
survive in shadow and known as facultative sciophytes. Similarly, among
sciophytes, some plants are obligate sciophytes which means they cannot tolerate
sunlight and can survive in shade only. Some plants are known as facultative
heliophytes which means they are basically sciophytes, but in the absence of shade,
they can also survive in light.

Temperature
It is another very important climatic factor affecting reproduction, metabolism,
growth and development of organisms. Reproductive ability of animals is affected
by temperature for example; production of more eggs at higher temperature than at
low temperature by some insects, breeding only during summers or maturation of
gonads at a particular temperature, etc. Metabolic activities are regulated by actions
of different enzymes and functions of enzymes are affected by a specific
temperature. Hence, various metabolic activities like respiration, photosynthesis,
seed germination, etc. are controlled by temperature factor.

Growth and development of plants and animals are also affected by change in
temperature. Exposure to very low temperature causes chilling injury in plants
of hot climate whereas extreme low temperature causes freezing injury in plants
of temperate climate. Some perennial plants show cold resistance by tolerating
low temperature. Extreme high temperature causes heat injury in plants by
affecting various metabolic processes.

Temperature also affects development of animals for example, dependence of

hatching time of eggs, increase in body size of oysters, corals, sea urchins, etc.
with an increase in temperature etc. In some animals, temperature has been shown
to affect sex ratio, for example, production of more males with increase in
temperature in Macrocyclops albida. Likewise, dark color body of warm climate
animals than light color races of same species living in cool climate shows effect
of temperature on body coloration and also known as Gloger rule.

Effect of temperature on morphology of animal body is known as Bergman’s

rule. For example, big size of birds and mammals in cold region than in warmer
areas whereas smaller body parts of mammals in cold regions than in warmer
regions is known as Allen’s rule. Narrow and acuminate wings of birds of colder
areas than broad wings of birds of warm regions are known as Rensch’s rule. In
some animals, phenotypic changes are also evident due to effect of temperature
for example, changes in eye size or number of legs of fruit fly Drosophila
melanogaster.

Besides, there is also temperature effect on geographical distribution of plants

and animals. Change in latitude and altitude affect temperature values of different
regions of earth. Variation in soil water, direction of mountain chains, steepness
of slope, etc. also affects temperature which affects distribution of various plant
species. For example, on mountains, temperature decreases with increase in
altitude which is evident through different zones of vegetation on mountains from base
upwards. World’s vegetation has been divided into four classes on the basis of
temperature as 1) Megatherms in equatorial and tropical regions having tropical
96 rain forests as dominant vegetation due to prevailing high temperature almost
throughout the year, 2) Mesotherms in tropical and subtropical regions having Ecology
tropical deciduous forests due to alternating high and low temperature, 3)
Microtherms in high altitude regions having mixed coniferous forests due to
prevailing low temperature 4) Hekistotherms in polar and alpine regions having
alpine vegetation due to presence of very low temperature at higher altitude.

Similarly, effect of temperature on distribution of animals has also been observed,

for example, coral reefs cannot grow in colder regions where temperature is
below 21°C; Speckled trout cannot live in streams where temperature is above
25°C.

Plants and animals also show various adaptations to tolerate extremes of

temperature for example, production of thick-coated spores, seeds, cysts, eggs,
etc., increase in osmotic concentration, hibernation, aestivation, etc.

Rainfall
Rainfall is the one of the major events of water cycle on earth. Precipitation
occurs in forms of drizzle, rain, snow, dew, frost, sleet and hail on earth, out of
which rainfall is the most important source of soil water. Amount of rainfall affects
distribution and type of vegetation for example, evergreen forests are found in
tropical regions where heavy rains occur throughout the year, sclerophyllous forests
are found where heavy rains occur during winter and low during summer season,
grasslands are observed where heavy rains occur during summers and low during
winter season and xerophytic vegetation is visible at places with scanty rainfall.

Humidity
Moisture present in the form of invisible vapor in the atmosphere is known as
humidity which is expressed as relative humidity of air. The amount of moisture
which air can hold at saturation at the existing temperature is known as relative
humidity and it is measured by Psychrometer or Hygrometer. There are many
factors which affect humidity for example, temperature, altitude, wind, solar
radiation, etc. Habitat also affect relative humidity like places near oceans have
same humidity levels throughout the year whereas daily variations are visible in
deserts. Various physiological processes like absorption of water, transpiration,
evaporation is affected by changes in atmospheric humidity levels. Humidity also
affects life cycle of some plant groups like orchids, lichens, mosses and events
such as germination of spores, etc.

Atmospheric gases
Wind
Wind is defined as moving air which affects lives of plants on mountains, plains
and coastal regions. It causes changes in various physical, anatomical and
physiological processes of plants. Mechanical damage and uprooting of plants
are one of the important physical effects of high velocity wind on plants. There
are natural windbreaks present in the dense forests which reduce the wind velocity
by 80% and thus save the vegetation from physical, anatomical and physiological
damage. Nowadays, special trees and shrubs are planted in the fields as windbreaks
at right angles to direction of wind. Deformation of shoots of trees growing in
coastal regions by strong winds is a well-known effect of wind on plants. Flattening
of herbaceous plants and grasses against the ground is also a type of wind injury
known as Lodging. Abrasion of buds and other plant parts by soil/ice particles
97
Fundamentals of Ecology carried through wind is another type of wind injury. Exposure of roots of plants
due to soil erosion is result of high velocity wind. Strong wind also affects
transpiration and evaporation processes of plants as due to imbalance in internal
water of cells, plants suffer from desiccation. Sometimes dwarfing is visible in
the plants growing in coastal, arctic or alpine timberline due to dehydration and
loss of turgidity as a result of drying winds.

Fire
Mostly fire is caused by anthropogenic activities particularly in forests. Besides,
mass loss of plant species; it indirectly affects plants in various ways. Due to fire,
changes in soil fertility, litter & humus contents, pH, micro-flora & fauna, nutrient
cycle, rainfall are visible affecting the suitable habitat conditions for growth of
plant species. Large scars on plant stems generated by fire injury act as entrances
for various insect and parasitic fungi. Fire-tolerant plant species outgrows due to
reduction in competition by loss of fire sensitive plant species. However, treatment
of fire, positively influences germination of seeds of some grasses and shrubs;
thus, necessary for growth and development of those plant species. Similarly, in
some plants such as Betula papyrifera, killing of older shoots by fire is required
for development of new shoots.
Plants growing in areas of frequent fires develop some adaptations for survival.
For example, presence of lignotubers below the ground surface in Eucalyptus
species growing in Europe, North America and Australia which may give rise to
new shoots, presence of fire-resistant leaves due to lack of resin compounds or
oil content to check incidence of surface fires, development of fire-resistant seeds
or fruits with a rapid growth rate and short life cycle to avoid fire injuries, etc.

5.7.2 Topographic Factors

Climate of any region is indirectly affected by topographical factors. Topography
means physical geography of any area which includes mountains, hillocks, valleys,
slopes and any other surface irregularity of earth. These factors may sometimes
create specific microclimates peculiar of any region. Height of mountains is an
important factor in determining climate because with increase in altitude, changes
in humidity, temperature, pressure, solar radiation, etc., is observed which further
lead to change in vegetation. Direction of mountains determines direction of wind
flow and also affects humidity and rainfall. Due to these, some mountain sides are
rich in vegetation and some sides are poor in vegetation.
Slope is another important topographical factor affecting climate of an area.
Steepness of mountain slope is responsible for difference in amount of solar
radiation received and changes in soil characteristics. On steep slopes, water
flows down rapidly and upper layers of soil also runoff with water; hence, plants
are unable to grow there. Instead of this, in the valleys, water retains and permanent
water table is near to soil surface and supports growth of vegetation. Exposure of
slope is one of the important factors affecting vegetation types. For example,
slope which is much exposed to sunlight and wind is helpful for growth of plants
whereas the unexposed side has very less vegetation.

5.7.3 Edaphic Factors

It includes quality and texture of soil which affects life of plants in many ways.
Soil is the uppermost layer of earth and formed due to weathering of rocks into
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small mineral particles and degradation of plants and animals into organic matter Ecology
humus. The branch of science in which aspects related to soil are studied is known
as Pedology and the process of soil formation is known as Pedogenesis. Rocks
are weathered due to physical factors such as wind, water, temperature, gravity;
chemical processes like hydrolysis, hydration, carbonation, oxidation, reduction,
etc., and biological factors such as effect of lichens, fungi and bacteria on rock
surface.

If a straight trench is dug into soil, different natured horizons (layers) of soil are
found superimposed on one another and form ‘Soil Profile’. These layers are
termed as ABC layers from top to bottom (Fig. 5.2).

Fig. 5.2: Different horizons in soll profile

Top most soil is part of ‘A’ horizon where organic matter is present in its various
disintegration stages. The outermost layer of A horizon is named as A00 which
consists of pieces of branches, dead leaves, fruits, etc. The next layer is termed
as A0 which consists of partly decomposed organic matter where decomposing
matter can’t be recognized and termed as Duff. Layer below this is known as A1
which is dark in color and composed of organic matter humus mixed with mineral
matter. Next layer known as A2 layer is lighter in color with little amount of
organic matter and more amount of large size mineral matter. After A horizon a
mineral rich horizon ‘B’ is present which could also be further sub-divided into
B1, B2 and B3. B horizon is coarse-textured having silica, aluminum, iron,
manganese rich clay organic compounds and darker in color. Horizons A1, A2 and
B are collectively termed as Solum. The next horizon is known as ‘C’ horizon,
made up of incompletely weathered rocks below which true bedrock is present.
Looking at this profile, it is clear that the top most soil is the most fertile and
important for growth and development of plants. There is difference in thickness
and structure of these horizons at different places which affects development of
plants growing in those soils.
99
Fundamentals of Ecology Soil is made up of four major components besides others. These are: mineral
matter, organic matter, soil water and soil air.

Mineral matter
Due to variable degradation of mineral rocks, particles of varying sizes are present
in soil. These particles are named by International Society of Soil Science as
given in Table 5.1:

Table 5.1: Classification of Soil particles based on size

S. No. Name of Particle Diameter of Particle (mm)
1. Coarse gravel More than 5.0
2. Fine gravel 2.0-5.0
3. Coarse sand 0.2-2.0
4. Fine sand 0.02-0.2
5. Silt 0.002-0.02
6. Clay Less than 0.002

Due to difference in ratios of these particles, texture of soil becomes different in a

specific area. According to their relative ratios, soil is said to be of four major types as
given in Table 5.2.

Table 5.2: Soil texture based on percentage of various soil particles

Type and proportion of Particles Soil texture
85% sand + 15% clay or silt or both Sandy soil
70% sand + 30% clay or silt or both Loamy sand soil
50% sand + 50% clay or silt or both Loam soil
10% sand + 90% silt Silt

Soil texture affects soil water, air and root penetration of plants and also influences
nutritional status of soil. For example, sandy soil has less water holding capacity
and nutrients, with large pore space whereas clayey soil is heavy textured with
less aeration and more water holding capacity leading to water logging of soil.
Plants grow best in loamy soil where sand, silt and clay particles are almost
equal in quantity and thus good in fertility as well as good for easy movement of
air and water, root penetration and water holding capacity.

Soil organic matter

Though present in fewer amounts in the soil, organic matter widely affects the
growth of plants. It is formed by degradation of dead parts of plants and animals
by microorganisms living in the soil. The matter in which all dead fallen plant
parts and animal debris are recognizable and not degraded is called ‘Litter’.
Organic matter lying below litter which is degraded and not recognizable in
their original format is called ‘Duff’. Matter present below Duff is dark brown
or black in color and known as ‘Humus’. The process of degradation of dead
parts of plants and animals and their conversion into humus is called as
humification which depends on climatic factors like temperature, humidity, aeration
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and nature of degrading plant or animal parts as well as nature of microbial flora Ecology
and fauna available in soil for degradation.
Muller in late 1800s used two terms ‘Mor’ and ‘Mull’ humus for two different
types of humus. In Mor humus, soil is degraded at a very slow rate and humus is
raw, hard and non-porous in nature. It is devoid of earthworms with very less
amount of mineral matter. On the other hand, Mull humus layer is amorphous,
porous, loose and microbiologically richer than Mor humus with a fast rate of
degradation. Mull is usually formed in forests having broad-leaved trees.
Humus is indicator of a healthy soil and highly beneficial for growth of plants. It
is rich in minerals and plant nutrients and increases water holding capacity of
soil.

Soil water
It is one of the important factors in developing variations among plant communities.
There are pore spaces present between soil particles. Rainfall is the chief source of soil
water which fills in pore spaces and known as capillary water. It is the only water
which is available to plants among other soil water. Total amount of water available in
soil is known as Holard; whereas the water which is available to plants is known as
Chresard and rest of the water unavailable to plants is known as Echard.
According to Briggs (1897), three major types of water are available in the soil.
Hygroscopic water is tightly held around soil particles by surface forces and
cannot be removed by plants, Capillary water as described above and third is
Gravitational water which moves downwards due to gravitational force and
reaches up to deep saturated ground water zone of earth. Other types of water is
Combined water present as hydrated oxides of aluminium, iron, etc. in soil and
water vapor present in vapor form between pore spaces of soil particles. Water
holding capacity of soil is a very important measure of its quality. Percentage of
water retained as capillary water in soil against gravitational force is known as
water holding capacity. Due to water logging, aeration reduces in soil leading to
less absorption of water by plants and therefore, excess of soil water is harmful
for growth of plants.
Soil air
Oxygen is necessary for respiration and water absorption by underground roots
of plants in soil. Due to poor aeration, levels of CO2 increase in the soil leading
to production of some toxic compounds like acetic acid, formic acid, oxalic acid,
hydrogen disulfide, etc., which has harmful effect on growth of plants.
Degradation by various soil microbes also depends on concentration of oxygen.
Soil air is also important for seed germination and humification process. There
are pore spaces present between mineral particles in the soil whose size and
number differ in different textured soils like fine-textured soils silt/clay contains
small and less pore spaces than coarse-textured sandy soil.

Soil temperature
It is another important factor in affecting various physical and chemical processes
undergoing in the soil for example, absorption of minerals and water by roots,
growth of underground parts of plant and seed germination. Maximum metabolism
is observed in plants between 20° to 30° C. At the temperature below 20° C,
water absorption reduces and plant remains dwarf.
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Fundamentals of Ecology Soil flora and fauna

Organisms present in soil are also important components of soil. Both flora and
fauna are present in soil. Flora consists of bacteria, fungi, algae, actinomycetes
and fauna could be made up of nematodes, earthworms, rotifers, protozoa, termites
and burrowing animal species. Some of these microorganisms decompose the
complex organic matter present in dead plant and animal parts into simple
compounds and make it available again to other organisms. In this way fertility of
soil is maintained for years. Earthworm is one of the best friends of soil and helps
in proper aeration and gaseous exchange. Due to incomplete degradation of organic
matter, sometimes toxic compounds are produced in soil which negatively affects
growth of plants.

Soil reactions

Depending on concentration of H+ and OH- ions in soil, it could be classified into

acidic, neutral or alkaline. Neutral (pH=7) or partially acidic (pH 5.5-6.9) soil is
considered appropriate for growth of most of the plants. Excessive acidic or
alkaline soil could be harmful for plants. Soil microbes are sensitive to pH of
soil and thus, it is also an important factor in soil health, further affecting plant
growth.

Limiting Factors

An organism must get all essential materials for proper growth and reproduction
efficacy. There are several factors and complex conditions on which presence
and success of an organism depends. Any factor or condition which exceeds the
limits of tolerance is said to be the limiting factor or condition. Liebig (1840)
gave the ‘Law of minimum’ which states that under steady-state conditions,
growth of species depends on the material which is available to the species in the
minimum quantity. That material acts as limiting factor for growth and success
of that species. Shelford (1913) gave the ‘Law of tolerance’ which suggests that
success or failure of a species depends on qualitative or quantitative deficiency
or excess of any of the factors which approaches the limits of tolerance for that
species. It has incorporated limiting effect of both maximum and minimum as
limiting factor. The combined concept of limiting factors is based on the idea of
minimum and concept of limits of tolerance. It suggests that growth of any species
depends on firstly, the quantity of material which is required though in minimal
amount and secondly on the limits of tolerance. For example, oxygen gas which
is abundant in nature is not limiting for terrestrial species but it is a limiting
factor for aquatic organisms where it is scarce and available in variable amounts.
If a species has wide limits of tolerance in moderate quantity, then that could not
be limiting for that. Therefore, careful observation is required to determine which
factors are operationally significant and what are their effects on individual,
population or community.

5.7.4 Biotic Factors

All living species including plant, animal and microbes are considered as Biotic
factors. In nature, there is no independent existence of any organism, as all
organisms whether plants or animals interact and affects each other either directly
or indirectly.

102
Check Your Progress 1 Ecology

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 scope of ecology.
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2. List down the environmental factors that affect the living organisms.
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3. Explain the role of climatic factors in the distribution of plants.
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4. Explain the influence of soil properties on soil organisms.
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5.8 BIOTIC RELATIONSHIPS

For many biological processes such as pollination, fertilization, nutrition, growth,
seed dispersal, etc., biotic interactions occur among organisms. These could be
divided as: Intraspecific relationship, where interactions occur within the same
biological species and Interspecific relationship, where interactions occur
between two different biological species. These interacting activities act as biotic
factors affecting the ecosystem in various ways. In this regard, human beings are
an important biotic factor. Biotic relationships could be studied under various
categories as follows: 103
Fundamentals of Ecology Symbiosis
The term given by H.A. De Bary (1879) literally means ‘living together’. The
relationships between two species living together could be divided into two broad
categories positive or negative. Later on Clarke (1954) suggested for using the term
symbiosis only for positive interactions where the association is beneficial for either
one of the associating partner.

Odum (1971) suggested using the term ‘Symbiosis’ in a broader sense and kept
nine type of interactions under symbiotic interactions. Relationships among living
organisms could be broadly categorized into three i.e., neutral, positive and negative
interactions. These are summarized in the Table 5.3 where + indicates positive; -
indicates negative and 0 indicates neutral interaction.

Table 5.3: Types of interactions occurring among two-species populations

Sl. Species Type of Nature of interaction
No. interaction
1 2
1. 0 0 Neutralism No effect on each other
2. + 0 Commensalism Species 1 is benefitted; 2 is
not affected
3. + + Proto-cooperation Both are benefitted; but association
is not obligatory
4. + + Mutualism Both are benefitted; but association
is obligatory
5. - - Competition: Both actively inhibits growth of
Direct interference each other
6. - - Competition: Both indirectly inhibits each other
Resource use in using common resource which
is in short supply
7. - 0 Amensalism Species 1 is inhibited and 2 is not
affected
8. + - Parasitism Species 1 is parasite; smaller than
host 2
9. + - Predation Species 1 is predator; larger than
prey 2

Neutralism
It is a type of association where both species are living together but do not affect
each other in any manner. Such insignificant or negligible interaction is termed
as neutralism. Though, practically such type of interaction is very rare to find in
the nature.

Positive interactions
If the association between two species results in positive effects, it is called a
positive interaction. These associations are beneficial for single or both the
participating species. Following associations are part of positive interactions:
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Mutualism Ecology

It is also known as obligate symbiosis where both populations are essential for
survival of each other and mutually beneficial. For example:
• Pollination by insects while deriving nectar from flowers as their food
• Presence of nitrogen fixing bacteria Rhizobium species in root nodules of
leguminous plants where bacteria obtain food from host plant and in turn
fix atmospheric nitrogen and make it available for the plants.
• Mycorrhizal associations between fungi and roots of higher plants are also example
of mutualism.
• Association between termites and protozoans living in their guts where protozoans
help in digesting wood for termites and obtain food and shelter in return.
• Cellulose digesting anaerobic bacteria such as Bacteroides succinogenes,
Ruminococcus flavofaciens, etc. in rumen of herbivorous mammals like
sheep, goats, cattle, etc. which digest cellulose, provide vitamins to ruminants
and thus grow rapidly.
• A green alga Chlorella vulgaris providing food and oxygen to Hydra while living
within its gastrodermal cells and getting shelter and carbon dioxide in exchange.

Commensalism
In this type of association one species (commensal) is definitely benefitted but the
other species (host) is not affected in any way. Thus, it is unidirectional positive interaction
but not an obligatory symbiotic relationship. There is no physiological dependency
between host and commensal. One can find many examples of commensalism in nature
for example, the process of nitrification where product of one microbe is substrate for
another one that is oxidation of ammonium to nitrite by Nitrosomonas and further
oxidation to nitrate by Nitrobacter bacteria. Similarly, microbes growing in rhizosphere
around roots of plant use them without affecting the host plant are good example of
commensalism.

Proto-cooperation
It could also be said as non-obligatory mutualism where the association is beneficial
for both the partners but it is not essential for survival of each other. For example,
association between hermit crab and a coelenterate sea anemone where crab helps in
movement of sea anemone which is attached to its shells to fresh feeding sites and in
turn sea anemone protects crab by its nematocysts from external enemies.

Antagonism
All the relationships where any one of the associating partners is getting harmed are
termed as antagonistic association. According to this, all negative interactions are part
of antagonism.

Antibiosis
In such interaction, a harmful substance for competing population is produced by one
population and none of them derive any benefit from such interaction. Antibiotic
compounds secreted by some bacteria are most common example of antibiosis.
Microbial world is much familiar with phenomenon of antibiosis. Some blue-
green algae like Microcystis, etc., bloom on pond’s surface and cause death of
fish and other animal species by secreting harmful toxins like hydroxylamine. If 105
Fundamentals of Ecology growth of any plant species is inhibited by chemicals secreted by other species, it
is called as allelopathy. For example, a noxious shrub Lantana camara secretes
toxic compounds like lantadenes and does not allow survival of other plant species
near its existence.

Competition
Competitive interactions are mutual interactions which might occur for space,
food, nutrients, light, water, waste materials, susceptibility to disease, etc. These
may be of two types i.e., intraspecific – the competition occurring between
individuals of same population and interspecific – the competition occurring
between populations of two different species. If a competition arises due to common
physical space or limiting nutrient, then the dominant competitor will outgrow the
other competitor. It is also known as Competitive Exclusion Principle given by
E.F. Gause in 1934.

Parasitism and Predation are part of exploitation activities where one species
harms other by using it either directly or indirectly for food, shelter or support.

Parasitism
Organisms which derive their food either by living on or in the body of other
organisms are called as parasites. A typical parasite doesn’t kill its host organism.
These could be ectoparasites or endoparasites. There are also some vascular
plants acting as partial or total parasites. Parasitic flowering plants could be
classified as follows:
a) Total stem parasite: A root and leafless plant, Cuscuta living on stem of
Acacia or Ziziphus species is totally dependent on host plant for its survival.
Similarly, Cassytha is total stem parasite on Azadirachta indica.
b) Partial stem parasite: Loranthus spp., Viscum album, Dendrophthae spp.
are examples of partial stem parasites.
c) Total root parasite: Striga living on roots of Lepidagathis spp., Rafflesia on
roots of Vitis spp., Orobanche, Epifagus, Balanophora on roots of higher
plants are examples of this association.
d) Partial root parasite: Santalum album growing on roots of Albizia spp.,
Thesium, etc. are examples of partial root parasite plants.

Predation
In contrast with parasitism, here for obtaining food, the predator catches and
kills its prey which is usually smaller than size of predator. Predators taking food
from animals are called as carnivores and predators taking food from plants are
called as herbivores. In a sense, human beings are also predators. Even some
fungi such as Zoophagus, Arthrobotrys, Dactylaria, Monacrosporium, etc. take
their food from insects and nematodes. There are some carnivorous plant species
which capture insects and small animals and digest them with proteolytic enzymes
and obtain their food. These are called as insectivorous plants for example Drosera,
Dionaea, Nepenthes, Utricularia, Sarracenia, etc. Some predatory bacteria are
also present in nature like Bdellovibrio, Vampirococcus, Daptobacter, etc. which
attack other bacteria.

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Check Your Progress 2 Ecology

Note: i) Use the space given below for your answers.

ii) Check your answers with those given at the end of the unit.
1. What is the difference between autecology and synecology?
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2. Define community.
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3. What is the difference between Mor and Mull Humus?
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4. What is Holard?
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5. Describe Mutualism and give one example.
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107
Fundamentals of Ecology 6. What is Allen’s rule?
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5.9 LET US SUM UP

Ecology is study of the environment in which organisms live. Abiotic and biotic
factors interact with each other and affect the ecology of a given area. Genes,
cells, species, population, community, biome and biosphere are different levels
of organization in ecology. It is a multi-disciplinary science where knowledge
about various disciplines is required for complete study of an area. Applied
Ecology is the appropriate utilization of different ecological theories and models
to solve various environmental problems and issues.

Among abiotic factors, there are climatic factors such as light, temperature, rainfall,
humidity, atmospheric gases, topographic factors like height, slope and direction of
mountains etc. and edaphic factors related to soil; its texture, temperature, aeration,
moisture content, mineral matter, organic matter etc. All these factors affect living beings
either directly or indirectly. Therefore, a thorough analysis is required to reach up to
conclusion about any environmental problem.

Biotic factors include various types of relationships occurring among organisms. These
could be mainly divided into two types of biotic relationships, i.e., intra-specific and
inter-specific. There exist positive and negative types of interactions among organisms
besides neutralism where no species affects each other. Among positive interactions,
there are mutualism, commensalism and proto-cooperation and among negative type
of interactions, there are competition, parasitism and predation kind of association.

5.10 KEY WORDS

Biome: A large geographical area with characteristic groups of organisms adapted to
that particular environment.
Community: A group of populations of different species that interact with each other.
Ecosystem: A community of organisms plus the abiotic (nonliving) factors that interact
with the community.

5.11 SUGGESTED FURTHER READING/

REFERENCES
Aldhebiani AY. 2018. Species concept and speciation. Saudi Journal of Biological
Sciences. 25:437-440.
Bertelsmeier C, Bonnaud E, Gregory S, Courchamp F. 2012. Applied Ecology.
In: Encyclopedia of Theoretical Ecology. Hastings A & Gross L (Eds.), University
of California Press, USA, pp. 52-60.

108
Odum EP. 1971. Fundamentals of Ecology. 3rd edition. W.B. Saunders company, Ecology
Philadelphia.
Sharma PD. 2007. Ecology and Environment. Tenth revised edition. Rastogi
Publications, Meerut.
Sharma PD. 2016. Microbiology. 3rd edition. Rastogi Publications, Meerut.
Singer FD. 2016. Ecology in Action. Cambridge University Press, United Kingdom.

5.12 ANSWERS TO CHECK YOUR PROGRESS

Check Your Progress 1
1. Refer section 5.4
2. The environmental factors are categorized into four factors:
A) Climatic factors
Light, temperature, rainfall, humidity, atmospheric gases are climatic factors.
B) Topographic factors
Height of mountains, direction of mountains, slope are few topographic factors.
C) Edaphic factors
Soil temperature, aeration, flora and fauna, soil reactions, soil organic matter
are few edaphic factors.
D) Biotic factors
All living species including plant, animal and microbes are considered as Biotic
factors.
3. Refer section 5.7
4. Refer section 5.7
Check Your Progress 2
1. Autecology is study of an individual species in relation to environment
whereas synecology is study of groups of organisms in relation to environment.
2. Community is a group of populations of different species within a defined
given area.
3. Mor humus lacks decomposing organisms. It is hard, non-porous with fewer
amounts of minerals whereas Mull humus is rich in decomposing organisms
and minerals. It is loose, porous layer where decomposition rate is higher.
4. The total amount of water available in soil is known as Holard.
5. Mutualism is a biotic relationship which is mutually beneficial for both the
partners and obligatory in nature. For example, Mycorrhizal structures
between roots of higher plants and fungi.
6. Effect of temperature on body parts as smaller parts in mammals living in
colder region than of living in warmer climate is known as Allen’s rule.

109
Fundamentals of Ecology
UNIT 6 POPULATION PARAMETERS AND
REGULATION

Structure
6.1 Introduction
6.2 Objectives
6.3 Definition
6.3.1 Density
6.3.2 Natality
6.3.3 Mortality
6.3.4 Population Dispersal
6.3.5 Age Distribution
6.3.6 Population Distribution
6.4 Population Growth
6.4.1 Factors Affecting Biotic Potential
6.4.2 Carrying Capacity
6.5 Population Regulation
6.5.1 Density Dependent Factors
6.5.2 Density Independent Factors
6.6 Genetic Diversity of the Population
6.7 Evolutionary Implications of Natural Regulation
6.8 Let Us Sum Up
6.9 Key Words
6.10 Suggested Further Reading/References
6.11 Answers to Check Your Progress

6.1 INTRODUCTION
The population is attributed with various characteristics that are unique
possessions of the group and are not characteristics of the individuals in the
group. In the present unit you will study the population as a collective group of
organisms of the same species which occupy a particular space at a particular
time. We will discuss here some of these properties like density, natality, mortality,
age distribution, biotic potential, dispersion, and growth form. It is also known
that populations possess genetic characteristics which are directly related to their
ecology, such as adaptiveness, reproductive fitness, persistence. The persistence
means the probability of leaving descendents over long periods of time. Also,
you will study the parameters of population estimation, demographic techniques,
various approaches employed for population regulation and evolutionary
implications of natural regulation.

6.2 OBJECTIVES
After reading this unit, you will be able to:
• define population and discuss the concepts of density, natality, mortality,
dispersal, population distribution and age distribution;
110
• explain that growth is the fundamental feature of the population which is Population Parameters and
Regulations
limited by the carrying capacity of our environment and population size is
regulated by various density dependent and independent factors; and
• appreciate the importance of genetic diversity for the future survival of mankind
and nature and how systems of natural regulations are affected by evolutionary
changes.

6.3 DEFINITION
Population can be defined as a group of organisms of same species occupying a
specific area at a particular time, such as all the deer or all the pine trees in a certain
wood land. We may even speak of human population of the world, of India or of
Delhi, simply by shifting the borders that enclose the group. Individual organisms that
interbreed are the ultimate constituents of the population and share a common gene
pool. Gene pool is the sum total of the genes of all the individuals in a population.
Populations may be further subdivided into demes or local population, which are the
smallest collective group or unit of interbreeding individuals.

A population has a number of group characteristics, that are statistical measures

unique to the population group and are not the characteristics of the individual.
These group characteristics are density, natality, mortality, immigration and
emigration, age distribution, dispersion, growth, regulation and genetic
composition. Density or size of population, however, is its basic characteristic
and is affected, by other group characteristics such as natality, mortality,
immigration and emigration. Any change in the density of the population implies
a change in one or more of these four characteristics. The relationship between
these factors is expressed as follows:

The study of the group characteristics or parameters of the population, their

changes over time and prediction of future changes is known as demography.
Techniques used for quantitative analysis of these statistics are known as
demographic techniques. In the following subsections we will discuss various
statistical parameters of population.

6.3.1 Density
Density is defined as number of individuals or population biomass per unit of
area or volume at any given time. Biomass refers to the total weight of all the organism
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Fundamentals of Ecology or of a specific group of organisms in a given area. Density is generally expressed
in terms of biomass when the size of individuals in a population is quite variable.
This measure of number or biomass of individuals per unit total area is called
crude density. However populations do not occupy all the space within the unit
area because whole of it is not a suitable habitat. Each organism occupies only
areas that can adequately meet its requirements, resulting in patchy distribution.
No matter how uniform a habitat may appear, it is not uniformly habitable,
sometimes because of even micro differences in light, moisture, temperature, etc.
Density, thus measured in terms of the amount of area as habitat or living space is
ecological density.

To cite an example we may talk of density of a goat population in a given area as

500 individuals per hectare, but goats might not utilise the entire area because of
various factors such as human habitation, lack of vegetation cover or lack of food.
Other examples can be of mango trees as 50 trees per acre, of diatoms as 5
million diatoms per cubic metre of water, of fishes as 200 pounds of fishes per
acre of water surface, of human population as hundreds of individuals per square
kilometres as it is in modern cities, etc. These examples indicate the range of
figures we have to study. Techniques to study density that work nicely with goats
cannot be applied to diatoms. The choice of technique is influenced by the size
and mobility of organism with respect to man. We will briefly discuss some of
these methods here.
1) Total Counts: The most direct way to find out how many organisms are
living in an area is to count them. It is possible with large or conspicuous
organisms or with those which aggregate into colonies. The best example of
this is human population census. Periodic head counts are made as we have
recently done it in India. India’s population stood at 843.93 million on March
1, 1991, with territorial birds, one can count all the singing males in the area.
Some animals, such as northern fur seal, may be counted when they are all
gathered in breeding colonies. Large plants on small areas can sometimes be
counted in total.
2) A quadrat is a sampling area of any shape. Although the word literally indicates
a four-sided figure, in ecology it has been used for all shapes of areas,
including circles.
Sampling Method: In this method a small proportion of the population is counted
A quadrat is a sampling and this sample is used to estimate the total. There are two ways of sampling
area of any shape. Al- which are given as follows:
though the word literally Use of quadrats involves counting or weighing organisms in several quadrats,
indicates a four-sided i.e. plots or transects of known size and number to estimate the average density.
figure, in ecology it has This average density is then extrapolated to the whole area. For example, if you
been used for all count 9 individuals of beetle species in a soil sample of 0.01 m2, you could
shapes of areas, includ- extrapolate it to 900 beetles per square meter of soil surface. In another example,
ing circles.
as shown in Fig. 6.1 there are 30 individuals (centipede) in 37 hexagonal quadrats.
The mean density is

30 individuals
= 0.811individual per quadrat
37 auadrats

Since each quadrat is 0.08 m2, estimated density is 10.1 individuals per square
metre. This sampling of centipedes was done by Lloyd in 1967 in central England.
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 Population Parameters and
Regulations

Fig. 6.1: Sampling of centipedes in 37 hexagonal quadrats. The estimated population

came out to be 10.1 individuals per square metre.

Quadrats have been extensively used in plant ecology and are most common method
for sampling plants. By doing the quadrat sampling for old trees and then for seedlings
in the same quadrat, one can determine if the populations were likely to change with
time. Foresters have deviced various ingenious quadrat sampling techniques for estimating
the forest trees.

Capture-recapture Method involves capture, marking, release and recapturing of

samples of a population. The proportion of marked individuals in the recaptured samples
helps in determining the population as follows:

Marked individuals in total population Marked individuals in the sample caught

=
Total population size Total caught in the sampe
A population is closed
For example, if 100 individuals of a population were marked and released and 10 out
if it is not changing in
of recaptured sample of 100 were found to be marked, then the population would be
size during the period
as follows:
of capture, marking
and recapturing. A
population is open if it
is changing in size
Therefore, total population (P) is of 1000 individuals. Capture-recapture method does during this period of
not work well when population is open, i.e. the density is undergoing a rapid change study.Real population
due to death and emigration. This method is used in animal ecology for the estimation are clearly open,
of the population of mobile and conspicuous animals like butterflies, snails, lizards, unless we sample
birds etc. them for a very brief
period.
Before proceeding further try the following check your progress to see whether you
have understood the concept of density.

Check Your Progress 1

Match the items in column I with those given in column II. Write your answer in the
space provided.
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Fundamentals of Ecology
Column I Column II
a) Ecological density [ ] i) Does not work with
populations whose density
change rapidly
b) Sampling method [ ] ii) Density per unit area as habitat
or living space
c) Capture-recapture methods [ ] iii) A small proportion of
population is counted and
used to estimate the total
d) Quadrats [ ] iv) Average density is measured
with the help of plots of same
size
So far you have read about one important parameter of population, i.e. density
which would be first one to get the attention while studying population. The
effect of population in any ecosystem depends both on type and number of
individuals. For example one crow in a 10 acres crop field might not affect the
yield, but 100 crows per 10 acres would certainly affect the yield. We will now
discuss briefly the other attribute of population which also affects density, i.e.
natality.

6.3.2 Natality
Natality is the ability of a population to increase. Natality rate is equivalent to
birth rate which means the production of new individuals by birth, hatching,
germination, or fission. Maximum production of new individuals under ideal
conditions of ecological and physiological factors is always theoretical and is
Natality is the ecological called maximum natality. It is constant for a population. However, the actual
concept which means increase in a population under a specific environmental condition is referred to
number of offspring as realized or ecological natality. This is not constant for a population and may
produced during a period vary with the size and composition of the population, i.e. the number of females
of time. Fertility is a in reproductive age at a particular time. It also varies with the physical
physiological notion that environmental conditions of the habitat a population is acquiring. For example,
indicates that the the realized natality rate for the human population may be only one birth per five
organism is, capable of years per female in the child bearing ages, whereas the maximum natality rate
breeding.
for humans is one birth per nine to eleven months per female in child bearing
ages.

Natality rate or birth rate is determined by dividing the number of individuals

born by unit time and is expressed as follows:
Symbol ∆(delta)
represents “the change
in” something, and by
writing it in front of the ∆ Nn = production of new individuals in a population
letter indicates that the
thing is changing. ∆t = unit time
Natality rate can also be determined as the number of new individuals per unit of
time per unit of population. This is called specific natality rate and can be
expressed as:

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 Population Parameters and
Regulations

N may represent the total population or only the reproductive part of the population,
i.e. females, for example, in higher organisms natality rate is per female. Natality rate is
zero or positive but never negative.

The measurement of natality or birth rate is highly dependent on the type of organism
being studied. Some species breed once a year, some breed several times a year and
others breed continuously. Some produce many seeds or eggs, and others few. For
example, a single oyster can produce 55 to 114 million eggs, whereas birds usually lay
between 1 and 20 eggs. Also the specific natality rate differs for individuals of different
age groups in the population. For example in a rabbit population for 1 to 2 year old
females the specific natality rate is average 4 young ones per year per female, while for
females of less than 1 year the rate is 1.5 on an average. Since natality is the concept
referring to the population and not to the isolated individuals, the average reproductive
capacity should be taken as the measure of natality, and not the capacity of the most
productive or least productive individuals.

6.3.3 Mortality
The death of an individual in a population is known as mortality. Mortality rate like
natality rate can be expressed as the number of individuals dying per unit time. Specific
mortality rate is expressed in terms of units of total population. Again like natality,
mortality can also be potential or ecological. Potential mortality also called minimum
mortality represents the death of an individual living under an ideal or non-limiting
environmental and physiological conditions. It is constant for a population. Ecological
mortality also called realized mortality is the loss of individuals in a population under
a given environmental and physiological conditions. It varies with populations and
environmental conditions. This means that under the best environmental conditions
individuals will die of old age determined by their physiological longevity. In most
populations in nature the average longevity is far less than the physiological inherent
longevity and so the realized mortality rate is much greater than the potential rate.

Only few organisms in nature attain their potential longevity. In most of them the life
span is shortened by predators, diseases and other hazards long before they reach
their old age. Measurement of mortality may be done directly or indirectly. Capture-
recapture method is a direct method about which we have discussed earlier in this unit.
One of the methods of indirect measure is that if one knows the abundance of successive
age groups in a population, one can estimate the mortality between these age groups.

What is really vital for the population is not which members die, but which member
survive? Consequently specific mortality rate of a population is expressed by survivorship
curve. To construct a survivorship curve, we start with a cohort of many individuals
newly added to the population and follow them to determine the age of death of each
member of cohort. The study is completed when the last individual dies. We then plot
number of survivors against ages. As shown in Fig. 6.2 survivorship curves may be of
three types.

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Fundamentals of Ecology

Fig. 6.2: Three main types of survivorship curves. In type I curve most of the mortality
occurs towards the end of the life. In type II curve age specific survival remains constant.
In type III curve mortality is very high during the young stages.

Life tables for human populations are used by National and regional planners and
Life Insurance companies to predict how much longer people of a given age are
likely to live. This is helpful in determining the price of insurance for people of
various ages.

In type I survivorship curve, most individuals of the cohort find environmental

conditions ideal and live out full physiological life and die as a result of old age.
However, a perfect type I curve never occurs because there is always some early
mortality. Most of the modern industrialized men approach a type I curve after
their first year of life. In first year the high infant mortality is from genetic or
developmental defects or birth accidents.

In type II curve which falls between types I and II the rate of mortality is constant
at all age groups, so that an individual’s chance of living another year is just as
good at one age as another. This curve is typical of several birds and of human
beings exposed to poor nutrition and hygiene.

In type III survivorship curve, most individuals die at an early age as eggs or
larvae, for example in many invertebrates, bony fishes plants and fungi. But those
few individuals that survive have a high life expectancy. Most survivorship curve
observed under natural conditions are intermediate between these & types:

Survivorship within a population can also be represented in the form of a life

table. A life table-is an age-specific distribution of a population that gives picture
of mortality and survival. An example of life table is given in Table 6.1.

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 Table 6.1: Life Table for the Total Population in a Developed Country Population Parameters and
Regulations
Age Interval Of 100,000 Borne Alive Average Remaining life span
Life span Number of Number Dying Mortality Average Number
Between living at the During age rate of Years of Life
two exact start of age Interval at the Beginning
ages interval (dx) of Age Interval
(in Years) (lx)
(x)
0-1 100,000 1,107 0.0110 73.6
1-5 98,893 269 0.0027 73.7
5-10 98,624 175 0.0017 71.7
10-15 98,449 181 0.0018 64.6
15-20 98,268 497 0.0050 59.7
20-25 98,771 673 0.0068 55.0
25-30 97,098 663 0.0068 50.4
30-35 96,435 725 0.0075 45.7
35-40 95,710 986 0.0103 41.0
40-45 94,724 1,483 0.0156 36.5
45-50 93,241 2,352 0.0252 32.1
50-55 90,889 3,483 0.0383 27.9
55-60 87,406 5,063 0.0579 23.9
60-65 82,343 7,281 0.0884 20.3
65-70 75,06 9,005 0.1196 17.9
70-75 66,057 12,214 0.1849 13.8
75-80 53,843 14,455 0.2684 10.4
80-85 39,388 14,467 0.3672 10.1
85 and over 24,921 24,921 1 9.2

To set up a life table, we must decide on age intervals to classify the population
data into groups. For example age interval for humans is generally 5 years, for
deer 1 year and for field mice 1 month.

Check Your Progress 2

Write (Y) for the correct statements and (N) for wrong statements.
1. Ecological natality is always constant for a population. [ ]
2. Natality rate measurement is dependent on the type of organism being studied.
[ ]
3. Potential mortality is the death of an individual under a given environmental
condition. [ ]
4. Very few organisms in natural populations attain their potential longevity.
[ ]

117
Fundamentals of Ecology 6.3.4 Population Dispersal
Population dispersal is the movement of individuals into or out of the population
or the population area. It occurs in three following ways in a population:
• emigration – one-way outward movement of individuals from an area.
• immigration – one-way inward movement of individuals into an area.
• migration – periodic departure and return of individuals to same area.
A population is inherently dynamic in nature since individuals are always leaving
or entering the populations. But such changes normally do not affect the size of a
population. It is because emigrations balance immigrations or because gains and
losses in terms of entry and exit of individuals are compensated by changes in
natality and mortality. However, mass dispersal may bring out rapid changes in
the population e.g., immigration may speed up population growth or in case of
extreme reduction may prevent extinction. Mass dispersal affects the balanced
population in other ways also e.g., the immigration of large number of blue gill
fishes into a pond already full of blue gill population may result in reduced growth
of the fish population and also result in smaller average size of fishes due to
environmental limitations. So even though the biomass density remains unchanged,
the size of fishes may be much reduced affecting the fishing process.
Dispersal is affected by the presence or absence of the barrier and vagility which
means inherent power of movement also called dispersal powers. Many plants
and lesser form of animals have greater dispersal power, although we all know
that birds and insects are much known for their ability to move around.
Migration is a special type of population dispersal process often involving the
mass movement of entire population. It occurs only in motile organisms and is
best developed in arthropods and vertebrates like certain fishes, birds and
mammals. Due to seasonal and diurnal migrations, the organisms are able to occupy
the regions which would be otherwise inhabitable during unfavourable conditions.
In most cases migration of population may occur for food, shelter or reproduction
and various ecological hazards such as temperature fluctuation, predation, etc.
Non-migratory populations in such unfavourable periods assume some form of
dormancy or undergo considerable reduction in number, for example, frogs
hibernate during winter season.
Apart from its influence on the size and density of a population, dispersal has few
advantages. It is the means by which new or depopulated areas are colonised. It
also contributes in gene flow and brings about gene exchange between populations
and hence the process of speciation.

6.3.5 Age Distribution

It is obvious that individuals in a population will be of different age groups.
Relative numbers of young and old individuals in a population will significantly
influence the behaviour of a population such as natality and mortality. The age
structure of any population can be classified into three categories, i.e. pre-
reproductive, reproductive and post-reproductive ages.
Mortality usually varies with age as chances of death are more in early pre-
reproductive and late post-reproductive periods. Likewise, natality is restricted
to the reproductive age of individuals in a population. The relative duration of
118
these ages, in proportion to life span is different in different organisms. For Population Parameters and
Regulations
example, in modern man the three ‘ages’ are relatively equal in length, about one
third of the life span. In comparison, the primitive man had much shorter post
reproductive period. Many animals and plants have quite long pre-reproductive
period. For example, a locust in its seventeen years of life has an extremely long
development history with adult life lasting only less than a year.

Functional age breakdowns can be useful to understand the importance of specific

environmental factors on the population as a whole. If we know both the time spent by
an organism in specific functional age groups and environmental factors that have
greatest effect on each age, we can gain a considerable insight into the
characteristics of that particular species. For example, this knowledge is essential
if we are trying to control an economically important crop and it is necessary to
know which stage in life cycle of the pest is most vulnerable to attack and by
which means.

The easiest and convenient way to picture age distribution in a population is to

arrange the data in the form of age pyramid. An age pyramid is vertical bar graph
in which the number or proportion of individuals in various age groups at any
given time is shown from youngest at the bottom of the graph to oldest at the top.
Fig. 6.3 shows the three types of hypothetical age pyramids which are as follows:
1) A pyramid with a broad base, indicating a high percentage of young individuals
in a population. This shows that population is expanding exponentially.
2) A triangular or bell shaped pyramid indicating moderate proportion of young
to old individuals. This is the characteristic of a stable population where
natality equals mortality.
3) An urn shaped figure indicating a low number of young individuals. This
pyramid is the characteristic of aging or declining population.

Fig. 6.3: Three types of age pyramid showing large (1) moderate (2) and small (3) number
of young individuals in a population. Vole population under laboratory conditions is
shown expanding (a), and stable (b) age pyramids. 119
Fundamentals of Ecology Also shown in the figure is the example of age pyramids of laboratory populations
of the vole (Microtus agrestis). Pyramid a show the expanding population with
an exponential expansion by increased number of young ones produced in unlimited
environment. Pyramid b shows the stable population where birth rates and death
rates are equal. Stable populations maintain the same age structure and may be
steadily increasing, decreasing or remain stationary. A stationary population is
the one in which number of individuals remain same over a period of time and
birth rate equals the death rate during the same period.

6.3.6 Population Distribution

Dispersion or distribution refers to the pattern of distribution of individuals of a
population. As shown in Fig. 6.4, individuals in a population may be distributed
in three broad patterns; a) uniform, b) random, and c) clumped. Uniform distribution
is more regular than random and may occur where competition between individuals
is severe or where there is antagonism which promotes even spacing. Random
distribution occurs where environment is very uniform and there is no tendency to
aggregate. This type of distribution is relatively rare in nature. Clumping of
individuals in groups is the most common pattern. In this case the groups could be
of same or of varying size.

Fig. 6.4: Distribution patterns of individuals in a population. Note that each rectangle
contains approximately the same number of individuals.

Again these groups could be randomly distributed, uniformly distributed or further

clumped with each other. All these ways of dispersions are found in nature. It is
obvious from the figure that if we examine small samples of dispersion from each
population, the results will be very different. For example, a sample from a
population with clumped distribution will give either too low or too high a density,
when the number in the sample is multiplied to obtain the total population. So we
can say that clumped populations require larger and more careful techniques for
study of populations than non-clumped ones.

Check Your Progress 3

Note: 1) Use the space given below for you answer
2) Check your answer with those given at the end of the unit.
1. State the ways of population dispersal.
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120
2. Differentiate between stationary and stable population. Population Parameters and
Regulations
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3. List the three patterns of population distribution.
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6.4 POPULATION GROWTH

By now you are aware that size of a population depends upon the balance between
natality and immigration through which individuals are added to a population
and mortality and emigration which remove them. If natality and immigration
exceed mortality and emigration, population will grow, if these are equal
population will remain same size, i.e., static and if death and emigration exceed
population will decrease.

Let us discuss the growth of population by taking few examples. The size of a
population of mice in a field seems to vary little from year to year despite the
fact that these organisms produce so many offspring that their population could
increase greatly from one year to the next. The size of such type of natural
populations is limited by environmental factors. Now consider another example
of Paramecium caudatum population studied by Russian ecologist G.F. Gause to
see how rapidly population could increase if nothing stopped their growth. Every
few hours a well nourished Paramecium divides to form two new individuals.
Gause set up tubes containing sufficient bacteria for food and introduced one
paramecium into each. If nothing checked the growth of paramecium, population
showed exponential growth, that is as time went on the number of individuals
added in each time period kept on increasing. When this type of increase in
population size is graphed on a linear axis, the exponential growth is plotted as a
curve that shows steep growth. When the population size is plotted on a logarithmic
axis, the exponential growth plots as a straight line (see Fig. 6.5 a,b).

Linear scale on the axis of a graph is taken when change in variables is not large
and can be accommodated on the axis. But if the change in variables is very large
viz. upto 100 or more, it is advisable to plot it on a logarithmic scale by taking the
log of the variables.
121
Fundamentals of Ecology

Fig. 6.5: Exponential growth of a population expressed oh a linear scale (a) and
logarithmic scale (b).

This type of exponential growth can be expressed in the form of following equation

dN
= γm N
dt

dN
Where N is the number of individuals in a population, is increase (or change)
dt
in number of individuals per unit time and gm is the maximum rate of population
growth per individual and is known as innate capacity, for increase or biotic
potential of the population. This is achieved when environment does not impose
any limitations, i.e. food and space are superabundant and there is no interference
from the members of other species.

Biotic potential also called reproductive potential is difficult to measure because

optimum conditions for growth almost never occur except under artificial
laboratory conditions. What we see in nature is the outcome of interactions
between a population’s biotic potential and various environmental factors that
restrict the actual growth. The actual rate of population increase is represented
by r which is the difference between birth rate and death rate per individual per
unit of time. In most natural populations g varies in response to interactions
between a population and its environment. Exponential growth occurs in nature
when a population has abundant supply of resources e.g., population explosion
occurs when bacteria invade the intestinal tract of an animal or when decomposers
invade a freshly dead animal or plant. However, exponential growth does not
necessarily mean that the population is growing at its biotic potential.

6.4.1 Factors Affecting Biotic Potential

Biotic potential differs from one species to another e.g., bacterial populations can
grow faster than population of oak trees. The rate of reproduction of any individual
can be increased in any or all of three following ways:
• by producing a large number of offspring each time, it reproduces,
• by having a long reproductive life, and
122 • by reproducing as early in life as possible.
Of these three factors, the last one is most important. Let us take an example to Population Parameters and
Regulations
understand this. A bacterium neither lives for a long time nor produces many offspring
each time it reproduces. Its reproductive potential is higher than that of a dog because
most bacteria can reproduce within an hour after being formed by cell division,
whereas a dog is not able to reproduce until it is at least 6 months old. So we can
say that shorter the generation time of a species, the higher its reproductive potential.
In case of organisms with equal generation time, the number of offspring produced
determines which has higher potential for population growth.

Thus the population of a plant that produces 100 seeds a year can potentially
grow faster than the population of a plant that produces 10 seeds a year. However,
with longer pre-reproductive period, number of offspring produced does not affect
much the biotic potential of a population.

6.4.2 Carrying Capacity

No population can grow exponentially for long. Gause found that his Paramecium
populations eventually stopped growing after reaching a certain level. So the
level beyond which no major increase can occur represents the saturation level
or carrying capacity which is represented by letter K. It is the number of individuals
of a particular species that a particular environment can support indefinitely.

Accordingly such type of population growth can be explained by following logistic

equation and the curve plotted is called logistic curve (see Fig. 6.6).

where N = population number

rm = innate capacity for increase

K = Carrying Capacity

The term indicates how much of the resources are still available to
populations. When N is much less than K, the term becomes approximately 1 and
the equation becomes = gm N (equation for exponential growth). As N almost
becomes equal to K, the term is almost zero and i.e. the growth rate also becomes
zero.
Another type of population growth curve called J-shaped curve is obtained when
the density of organisms increases rapidly and then stops abruptly as environmental
resistance or other limits become effective more or less suddenly. Carrying capacity
is determined by various factors including predation, competition and climatic
conditions. All factors which limit a population growth are collectively known as
the environmental resistance to population growth. Since such factors are many
and varied, it is clear that the carrying capacity of any area for a population may
vary over a period of time. In the next section we will discuss regulation of
population size, but before that try the following check your progress.

123
Fundamentals of Ecology

Fig. 6.6: S-shaped curve obtained as a result of growth of Paramecium population in a

culture medium with constant supply of bacteria as food each day. Population grew
exponentially at first but eventually reached zero as its numbers (N) approached the
carrying capacity (K).

Check Your Progress 4

Strike off the incorrect word in the following statements.

1. The size of natural populations is/is not limited by environmental factors.
2. Biotic potential is the innate capacity for decrease/increase of a population.
3. The length of the reproductive life of an individual affects/does not affect the
biotic potential of a population.
4. Carrying capacity is/is not the saturation level beyond which a population
does not grow.

6.5 POPULATION REGULATION

The number of individuals in a natural population varies with time. If the size of a
population declines too drastically due to some reason, it may become extinct, but
may later be re-established by immigration from other populations. On the other hand,
increase in size of a population is not infinite since the carrying capacity of the environment
always imposes a restriction upon it. In spite of such fluctuations, however, an outstanding
feature of most large populations is that their average size changes relatively little over
the years and certainly less than is expected from their biotic potentials. This indicates
that population sizes are regulated in such a way that small population grow fast, larger
populations grow more slowly and still larger populations decline. Let us see what
brings about such ecological homeostasis. In low-diversity, physically stressed
ecosystems or in those subjected to irregular or unpredictable external
perturbations, populations tend to be regulated by physical components such as
weather, water, chemical limiting factors, pollution, etc. In high diversity
ecosystems, or in those which are not physically stressed, populations tend to be
biologically controlled. In all ecosystems there is a strong tendency for all
populations to evolve through natural selection towards self-regulation such as
failure of reproduction and self-inflicted mortality. Even though this is difficult to
achieve under external stress. It is because over-population is not in best interests
124
of any population. Thus it can be said that limitation of number in any population Population Parameters and
Regulations
is brought about by the action and interaction of two basic regulatory processes
namely density dependent and density independent factors. We will discuss these
processes in the following subsections.

6.5.1 Density Dependent Factors

The density dependent factors are intrinsic or biotic factors and they depend on
interactions between individuals within same population or populations of different
species. Density dependent factors may stabilise the population at the level
determined by carrying capacity of the environment. The important density
dependent factors are reproductivity, emigration, competition for resources,
predation, parasites, and diseases. The contribution of these factors may vary
from species to species. You have already read how in large populations the
number of offspring produced are less, thus self-regulating the population size.
You have also read how emigration of individuals from a population reduces its
density. Competition can be between individuals of same species called
interspecific competition and between individuals of different species called
interspecific competition. Generally, members of the same species need same
resources and are bound to compete for them. In some bird species males and
females have different beak lengths enabling them to feed on different insect
prey.

Let us take an example, where seeds of white clover, Trifolium repens, were
planted at three different densities. Half of the plants at each density were watered
throughout the experiment but other half were watered only for first 18 days.
After seven weeks, the densities of the surviving seedlings were measured. As
shown in Fig. 6.7 among the seedlings that were watered regularly, mortality
was low regardless of density. However, the proportion of seedlings killed among
the deprived ones was three times greater in the high density plots than in the
intermediate density plots.

Fig. 6.7: Effect of seed density on survival of white clover seedlings subjected to water
shortage. Solid bars show that mortality was same at all population densities when water
was available. Unshaded bar represent the mortality of seedlings not watered after 18
days. 125
Fundamentals of Ecology Interspecific competition occurs when ecological niches of the species overlap.
Predation and diseases are a part of this type of competition and are partly density
dependent factors. It is because a disease-causing organism is more likely to
encounter a host, or a predator a prey, when there are more hosts or prey per unit
area. In dense animal populations individuals have decreased health and vigour
which make them more susceptible for predators and diseases.

6.5.2 Density Independent Factors

Density independent factors are the extrinsic factors which tend to regulate the
density of a population in ways that are not correlated with its density.
Environmental factors such as bad weather and scarcity of space, pollution etc.
are some factors. A hurricane, a severe winter, or a drought may kill most of the
individuals in a population irrespective of its density. In a bad weather only some
individuals may be able to shelter from it; if the number of shelters is limited.
Thus only a fraction of a large population will be protected. However, we cannot
pinpoint one or two factors and say that they determine the size of a particular
population. Often the sizes of natural populations are affected by many different
factors whose interactions can be complex.

Check Your Progress 5

Fill in the blanks in the following sentences with appropriate words from the text.
1. Ecological homeostasis is …………………… of population sizes.
2. Density dependent factors are …………………… factors which affect the
…………………… of population.
3. Competition for resources between individuals of different species is called
……………………
4. Environmental factors …………………… which affect the size of
population.

6.6 GENETIC DIVERSITY OF THE POPULATION

You will be aware that reduction in species and genetic diversity in historical
times has produced short-time benefits in agriculture and forestry. This can be
evidenced by the propagation of specialized, high yielding varieties over large
areas of the world’s crop and forest land. To call attention to the overall threat posed
by loss of diversity, biologists are organizing gene resources conservation programmes.

The biological diversity of animals, plants, and microorganisms is of fundamental

importance to human survival. The term “gene resources” may be defined as the
genetic diversity that is very essential for meeting the society’s needs in perpetuity.
This diversity is expressed in the differences between species as well as in the
variation among individuals that comprise a species. Gene resources include wild
and domestic species having many species of no commercial value. Every year
gene resources are utilized to provide crores of rupees worth of new and familiar
products e.g., food, clothing, shelter, pharmaceuticals, energy and hundreds of
industrial products. You will be aware that a wide range of species and their
products are required for medical and other research. Agricultural, forestry and
related industries are dependent, whenever needed on appropriate diversity as
for example resistance to plant diseases. It is this diversity that sets the limits to
126
which both wild and domestic species can successfully adopt to changes involving: Population Parameters and
Regulations
1) weather, insects and disease, 2) technology, 3) demand and, 4) human
preferences. Most of the biological diversity is still found in natural ecosystems
whose survival is dependent, in large part, on the diversity within them.

Good area and regional planning can compensate to some extent for the reduction
in local diversity which tends to accompany intensive agricultural, forestry and
urban development. If crop and forest monocultures as well as tract housing (rows
and rows of similar houses on small, un-covered plots), are interspersed with
more diverse natural or semi natural ecosystems, the ecosystem can be preserved
in perpetuity as park, nature centres and so on. If floodplains and other wetlands
together with steep slopes and ravines are left underdeveloped, not only will
there be a pleasing landscape full of recreation possibilities, but also a high level
of diversity will have been safeguarded.

Landscapes can be planned to preserve diversity and yet accommodate urban and
industrial development. Thus the diversity is necessary for the future survival of humans
and nature.

6.7 EVOLUTIONARY IMPLICATIONS OF

NATURAL REGULATION
You would like to know that how systems of natural regulation are affected by
evolutionary changes. You will be aware of the interaction involved in co-
evolution of predator-prey systems and herbivore-plant systems. In many of these
interactions, evolutionary changes operate very slowly and are difficult to be detected.
But recent studies have shown that evolutionary changes may occur very rapidly, so
that the evolutionary time scale approaches the ecological time scale. Thus natural
selection may impinge upon natural regulation.

Many changes in abundance can be attributed to changes in extrinsic factors such as

weather, disease or predation. But some changes in abundance are the result of changes
in the genetic properties of the organisms in a population. Such evolutionary changes
are produced by the genetic feedback mechanism. It is believed by the biologists
that natural population regulation has its foundation in the process of evolution.

We will illustrate the type of systematic changes by a simple model which could
be involved in the genetic feedback mechanism. Consider a two-species system
of one plant and one herbivore, and to make the model simple, let us focus on only
one gene on one chromosome in the plant. The hypothetical gene has a major
effect on 1) the ability of the plant to survive in its environment and 2) the
palatability of the plant to the herbivore. Two different alleles (A and a) occur at
the hypothetical gene locus, and the properties of the genotypes areas shown in
Table 6.2.

Table 6.2: Genotype of Plants

AA Aa aa

Ability of plant to survive Good Poor Very poor

Palatability to herbivores High Low Very low

127
Fundamentals of Ecology Thus plants of genotype AA are to survive very well but attract many herbivores
because they are desirable foods. Each plant genotype can support only a limited
number of herbivores before it is killed by overgrazing. Finally we assume that
the reproductive rate of fie herbivore will be affected by the genotype of plant on
which it lives, so that highly palatable plants are best for herbivore reproduction.
We can give here some examples of genetic changes of this type playing a role in
population regulation. For example, the Hessian fly population was reduced
drastically in Kansas after 1942 when resistant varieties of wheat were introduced.
The herbivore population of Hessian flies was significantly reduced by changing
the genetic makeup of the wheat plant. Another example is the myxomatosis-rabbit
interaction in Australia. The European rabbit was introduced into Australia in
1859 and increased to very high densities within 20 years. After World War II, an
attempt was made to reduce rabbit numbers by releasing a viral disease from
South America, myxomatosis. The myxoma virus was highly lethal to European
rabbits, killing over 99 per cent of infected individuals. Since myxomatosis was
introduced into Australia in 1951 evolution has been going on in both the virus
and the rabbit. The virus has become attenuated so that it kills fewer and fewer
rabbits and takes longer to cause death. Since mosquitoes are a major vector of
the disease, the exposure time before death is critical to viral spread.
Rabbits have also become more resistant to the virus. By challenging wild rabbits
with a constant laboratory virus source, we can detect that natural selection has
produced a growing resistance of rabbits to this introduced disease. Evolution in
the rabbit myxoma system has thus been towards an intermediate rate of increase.
This is explained by selection operating at the individual level for the rabbit but
at the level of the group for the myxoma virus. Group selection occurred because
less virulent viral colonies are favoured over more virulent viral colonies because
they take longer to kill the host rabbit.
Self-regulatory populations present yet another problem not covered by the genetic
feedback mechanism. How does a population evolve the machinery to be self-
regulatory? Self-regulation is clearly a desirable adaptation for any population
that has the potentiality of destroying its resources.

Check Your Progress 6

Which features listed in Column II below, correspond to statements given in Column

I. List them under suitable headings.
Column I Column II
a) Genetic Diversity i) The Myxoma virus was highly
lethal to Europeans rabbits,
killing over 99 per cent of
individuals infected.
b) Evolutionary implications of ii) European rabbit was introduced
natural selection into Australia in 1859 and
increased very high densities within
20 years.
iii) Landscapes can be planned to
preserve diversity and yet to
accommodate urban and
industrial development.
128
 Population Parameters and
6.8 LET US SUM UP Regulations

In this unit you have studied that:

• Population is a group of interbreeding individuals occupying a particular space at
I a particular time. Density, natality, mortality are some characteristics which are
attributed to the whole group and not to any individual.
• Density of any population is influenced by natality rate, mortality rate, immigration,
emigration and dispersal of individuals in the population. Life table tabulates the
statistics of mortality for various age groups in a population. Plotting of these data
gives a survivorship curves for various species.
• Populations have relative numbers of young and old individuals with different
distribution of pre-reproductive, reproductive and post-reproductive stages.
Age pyramids are the vertical graph of horizontal bars showing the number
or proportion of individuals in various age ranges with youngest at the bottom
and oldest at the top.
• Individuals in any population may be distributed in uniform, random or clumped
pattern. These three patterns of distribution are much dependent upon the various
resources such as food and space.
• Growth is the most fundamental feature that a population displays. Populations
have innate capacity to increase and can attain exponential growth. However,
there is a level beyond which no major increase occurs due to environmental
limitations representing the saturation level or carrying capacity.
• There is inherent tendency of all populations to increase their size. However, this
increase is not infinite and is regulated by various density dependent-and density
independent factors.
• Genetic diversity of a population can be evidenced by the propagation of
specialised, high yielding varieties over large areas of world’s crop and forest
land. Landscapes can be planned to preserve diversity and yet to
accommodate urban and industrial development. Genetic diversity is
necessary for the future survival of human race and nature as well.
• Recent studies have indicated that evolutionary changes may occur very
rapidly, so that the evolutionary time scale approaches the ecological time
scale and thereby natural selection may affect the natural regulation.
Check Your Progress 7
Note: i) use the space given below for your answer.
ii) Check your answer with those given at the end of the unit.
1) Discuss very briefly in the space given below the role of quadrats in measuring
the density of a population.
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129
Fundamentals of Ecology 2) Explain briefly about survivorship curves.
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3) Enlist the three types of hypothetical age pyramids.
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4) Explain briefly about carrying capacity.
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6.9 KEY WORDS

Carrying Capacity (K): The maximum population size that can be supported or
sustained by the environment.
Density-dependent Factor: A factor that reduces a population’s growth rate as
population density increases, by decreasing the birth rate and/or increasing the
death rate at high population density.
Density-independent Factor: A factor that influences a population’s growth rate
in the same manner regardless of population density.
K-selected Species: Species whose individuals compete effectively for resources
in predictable and stable environments, and whose populations tend to be at or
near their carrying capacity.

6.10 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
130 Publications, Meerut.
Singer, F.D. (2016). Ecology in Action. Cambridge University Press, United Population Parameters and
Regulations
Kingdom.
Singh, J.S., Singh, S.P. and Gupta, S.R. (2017). Ecology environmental science
and conservation. S. Chand Publishing house.

6.11 ANSWERS TO CHECK YOUR PROGRESS

Check Your Progress 1
a) ii, b) iii, c) i, d) iv
Check Your Progress 2
i) N ii) y iii) N iv) y
Check Your Progress 3
1. a) Emigration – One-way outward movement of individuals.
b) Immigration – One-way inward movement of individuals.
c) Migration - Periodic departure and return of individuals.
2. Stationary population remains the same size over the period of time. Stable
population may be steadily increasing decreasing or remain stationary.
3. Random distribution - occurs in very uniform environment.
Uniform distribution - occurs where competition between individuals is
severe.
Clumped distribution - individuals are distributed in group forms.

Check Your Progress 4

1) is, 2) increase, 3) affects, 4) is
Check Your Progress 5
1) regulation, 2) intrinsic, size, 3) interspecific competition, 4) density
independent
Check Your Progress 6
a) Genetic Diversity: iii)
b) Evolutionary implications: i), ii)
Check Your Progress 7
1) Quadrat is the sampling area of any shape in ecology. To measure the density of a
population the individuals are counted or weighed in various plots of known
size and number to estimate the average density. This average density is then
extrapolated to whole area.
2) Survivorship curves describe mortality in relation to age by plotting number
of survivors against the age groups. These curves are of three main types. In
type I most individuals live for a long time. In type III most individuals die at
an early stage. Type II is in between these two.
3) Age pyramids are of three types:
i) Pyramid with a broad base indicating a high percentage of young
individuals. 131
Fundamentals of Ecology ii) Triangular shaped pyramid showing the moderate proportion of young to
old individuals.
iii) Urn shaped pyramids indicate a low number of young ones.
4) Carrying capacity is the saturation level in the density of a population, beyond
which no substantial increase occurs in size and is represented by letter K.
The relationship of the number of individuals per unit time in a population
and its carrying capacity is shown by the following equation.

132
 Population Parameters and
UNIT 7 COMMUNITY ORGANISATION Regulations

AND INTERACTION AMONG

ORGANISMS

Structure
7.1 Introduction
7.2 Objectives
7.3 What is Community?
7.4 Community Gradients and Boundaries
7.5 Community Organisation
7.5.1 Habitat and Niche
7.5.2 Functional Roles and Guilds
7.5.3 Keystone Species
7.5.4 Dominant Species
7.5.5 Stability
7.6 Species Interaction
7.7 Competition
7.7.1 Competition in Laboratory Populations
7.7.2 Competition in Natural Populations
7.7.3 Results of Competition
7.7.4 Evolution of Competitive Ability
7.8 Let Us Sum Up
7.9 Key Words
7.10 Suggested Further Reading/References
7.11 Answers to Check Your Progress

7.1 INTRODUCTION
Although seemingly calm and quiet an ecosystem can be as busy as any large city
during rush hours – minus the noise. Let us consider a forest as an example. The soil
teems with bacteria, fungi, insects, mites, slugs, worms, spiders and scores of other
organisms that dig up the ground as they move and reproduce. Delicate seedlings
erupt through the surface, absorb nutrients recycled by the decomposers and eventually
grow into shrubs, trees, etc., that manufacture food for herbivores. Carnivores devour
the herbivores as well as other carnivores.

The organisms will vary from one ecosystem to another and the example here of the
forest illustrates how some organisms interact with each other. Some interactions benefit
one or both participants whereas, some harm one or both participants. In this unit, we
will first discuss the various approaches used to explain community organisation. The
following sections would deal with the interaction among organisms that make up the
community.

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Fundamentals of Ecology
7.2 OBJECTIVES
After reading this unit, you should be able to:
• describe the processes that affect community organisation and define niche, guild,
keystone and dominant species;
• define the various interspecific and intraspecific interactions giving examples of
each; and
• explain competitive exclusion principle and how potentially competing species
may coexist.

7.3 WHAT IS COMMUNITY?

In nature, different kinds of organisms occur in association with each other, sharing the
same habitat. Let us consider the example of a field to illustrate this point further. In a
field, different kinds of grasses, insects, worms, birds and mammals interact in various
ways. Grasses provide food for certain insects and mammals; insects provide food for
birds; and birds prey on small mammals and worms. The various kinds of organisms in
this field thus constitute a community, which is also known as field community. Similarly,
a forest, desert, pond, marsh and stream – are examples of natural communities. After
discussing briefly what a community is, we would now discuss six main features of a
community.

First, a community represents the biotic or living component of the ecosystem. If the
non-living (abiotic) factors, together with the living (biotic) entities are considered, then
we would be dealing with an ecosystem rather than a community.

Secondly, considering the functional aspect, communities are made up of organisms

with interlocking food chains and each species depends on many other species in a
community which are taxonomically unrelated. Try to recall a food web, this will help
you to understand this concept more effectively. Food web is a representation of the
food relationships between various types of species found in an ecosystem, and as you
know these biotic components constitute a community. While a species may not relate
to every other species directly in a community, nevertheless, they all are indirectly
interrelated. This situation can be compared with an engine in which various components
are interrelated and they together make the working of an engine possible.

134 Fig. 7.1: A deodara (Cedrus deodara) tree.

Thirdly, a community may be of any size. A temperate forest of deodar trees is an Community Organisation and
Interactions Amongs
example of a large community. (Fig. 7.1 shows a deodar tree). In contrast to this a Organisims
rotting log of wood harbouring many insects and worms represents a small community.
So, the size of a community may vary widely.

Fourthly, just like the concept of ecosystem, that it can be applied to any scale, that is,
the earth as a whole can be considered as a large ecosystem, on the other hand, a
bowl of water with various living organisms in it is an example of a small ecosystem.
Similarly, a forest is a community, so is a rotting log in that forest containing fungus,
insects like termites, and even mice. Similarly, a large number of microorganisms within
the gut of termite, that occurs in the rotting log of wood, also constitute a community.
This suggests that there is a community within a community, and the situation is just like
the toys shown in Fig. 7.2

Fig. 7.2: Wooden toys used as an example of community within a community.

Fifthly, some communities may be autotrophic, in the sense that they include
photosynthetic plants and obtain their energy from the sun. Other communities
such as those found in springs and caves are heterotrophic, as they depend upon
organic material such as detritus as a source of energy.

Sixthly, interrelated with the idea of community is that of the stand. In some
situations, these two terms mean different things; and in some other situations
these two terms mean the same thing, and are used interchangeably. In order to
avoid any confusion in the usage of these two terms, we shall illustrate these two
situations with the help of two examples, that are discussed below.

The first example is of a temperate forest consisting of deodars, pines and

rhododendrons (see Fig. 7.3). Have you noticed in the figure, that at places, the
plants of a particular species are forming groups, These groups are indicated in
the figure by the areas enclosed by the dotted lines. So, each of this area containing
plants of same species and almost of the same age is called a stand. You can also
observe form the figure that stands A, B and C of deodar trees are different from
each other as far as their number is concerned.
135
Fundamentals of Ecology

Fig. 7.3: Diagrammatic representation of a temperate forest.

The term stand is applied to a more or less uniform area of vegetation. While
studying a community, when we talk of a stand, it means we are talking about a
group of plants of a species in that community. After going through this example,
you would have seen that, here-community and stand refer to different things.

The second example is of a cultivated wheat field. The wheat plants in this field
constitute the wheat field community. Since, this field has a uniform vegetation,
and plants are of the same age, this can also be called as a stand of wheat. This
example shows one of the situations where the term stand and community mean
the same thing and can be used interchangeably.

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.
i) A biotic community is an assemblage if:
a) food webs
b) interacting populations
c) closely related species
d) ecosystems
(Choose the correct option)
ii) What are the salient features of a community?
......................................................................................................................
......................................................................................................................
......................................................................................................................
......................................................................................................................
......................................................................................................................
136
iii) Fill in the spaces with question mark, using appropriate word(s). Community Organisation and
Interactions Amongs
Organisims

7.4 COMMUNITY GRADIENTS AND

BOUNDARIES
It is often difficult or impossible to determine where one community ends and the
next begins. Many communities, in fact, grade continuously into each other with
no sharp boundaries. For example, if two forests, pine forest and spruce forest
are nearby, one cannot see the boundaries between them. But if one moves from
one end of the pine forest to the other end spruce forest, one can observe difference
in species composition between the two; yet one cannot demarcate a sharp boundary
between these two forest communities. There are, however, instances where sharp
boundaries between the communities are seen, especially where the physical
environment changes abruptly – for example, at the transition between aquatic
and terrestrial habitats between distinct soil types, or between north-facing and
south-facing slopes of a mountain.

Ecotone
The zone of vegetation separating two different types of communities is called
ecotone. It is also known as a transition zone. The border between forest and
grassland, the bank of a stream running through a meadow are examples of ecotones.
Ecotone is a region where the influence of two different patterns of environment
work together and hence the vegetation of ecotones is highly specialized. An
ecotone may be narrow or wide. For instance, the ecotones between adjacent
plots-one fenced and protected from grazing, and other without fence and openly
exposed to grazing; or between as ponds and an adjacent upland are quite sharp
and narrow whereas among any other types of communities ecotones are very
wide and community boundary differentiation is not easy.
137
Fundamentals of Ecology A general characteristic of ecotone is that it has sufficiently greater number of
species and the diversity of most of the species at times is higher than that in the
neighboring communities (also see Fig. 3.4). The phenomenon of increased variety
of plants at the boundary is called the edge effect and is essentially due to wider
range of suitable environmental conditions. The ecotone area contains organisms
from both of the adjoining communities and besides there are organisms which
are confined to the ecotone and can exploit the special conditions there.

Fig. 7.4: Ecotone – where two community types come together, such as a forest and field
species in the zone between them include both forest and field species and some
additional species that do not exist in either forest or field.

Some ecologists have introduced the continuum concept that means that there are
no distinct communities with well-defined boundaries but there is a gradual change
in space and time along a gradient which may be of moisture, temperature, soil
type, altitude or any combination thereof. There are no sharp borders or changes
in species composition in areas, according to the continuum concept.

Species Diversity
It is one of the most important and basic characteristics of a community. There are
various ways of measuring species diversity, the simplest is to enumerate the number of
species present in a given area. This is relatively easy to achieve for plants, and large or
sedentary animals but it is generally difficult to enumerate the various insect species
accurately. For large areas such as forests, islands etc., it may take many years to
prepare a reasonable estimate of species numbers. Assessment of species diversity
on the basis of species list is often an involved exercise. An, unless an elaborate
exercise is undertaken there is a good possibility that a number of species may be
left out. Since so many species in a sample are likely to be rare, we should not
ignore this fact while measuring diversity. For example, compare two imaginary
138
samples of 100 individuals each with two species A and B. The details of number Community Organisation and
Interactions Amongs
of species of each sample are given in Table 7.1. Organisims

Table 7.1: Number of Species in Two Sampling Units

Sample Number of Species
A B
I 50 50

II 99 1

In sample I there are 50 individuals of A and 50 individuals of B, but in sample II

there are 99 A and 1 B. Is the diversity of these two samples really same? If we
choose to measure diversity as the numbers of species present in each sample
then the answer is yes, but most ecologists would consider the community with
50A and 50B to be more diverse than the one with 99A and 1B. Let us see how.
To determine species diversity, the most widely used index is Shannon’s Index of
Diversity (H’) and it is calculated as below:

i=s i=s
' '
H = - � pi loge pi H = - � pi loge pi
i=1 i=1

H’ = Index of species diversity

S= Number of species
p= Proportion of the total sample belonging to the ith species (in rank), which
is calculated by dividing the number of individuals in species i by the total
number of individuals in the sample.
e= base of natural logarithms (loge pi = 2.302 × log10 pi)
i=s
means that-add up the following expression, for values
(Remember that �
of i from i=1 to i=s).
i=1

The larger the value of H’, the greater the uncertainty about predicting the next
species to be encountered and so the greater the diversity. Let us now compare the
two samples I and II, each of 100 individuals, and see whether sample I has
higher index of diversity H’ or sample II?
Sample I:
H’ = - [(0.50 × loge 0.99) + (0.01 – loge 0.01)]
= - [2 (0.50 × -0.69)]
= - [(-0.01) + (-0.05)]
= 0.69
Sample II:
H’ = - [(0.99 × loge 0.99) + (0.01 – loge 0.01)]
= - [(0.99 × -0.01) + (0.01 × - 4.61)]
= - [(-0.01) + (-0.05)]
= 0.06
139
Fundamentals of Ecology Sample I has the higher index of diversity.

7.5 COMMUNITY ORGANISATION

You know that a community contains populations of many species. We shall now
The difference between
see how a community is organized in the ecosystem. Three processes – competition,
habitat and niche can be
made clear if we take a predation and symbiosis help to organize communities. For instance, competition among
simple example. To plants, herbivores and carnivores could control the diversity and abundance of species
become acquainted with in a community. Predation could organize the community along feeding lines while
a person we need to know symbiosis which includes important interactions like mutualism that link species could
his address, i.e., where he help to increase community organisation in a positive way. This would be made clear
would be found, i.e., as you study these interactions in later sections.
habitat. To really know
the person well we would Before we examine the various factors that influence the organisation of a community,
want to know about his we should first review the important concepts of habitat and niche.
occupation, interests,
associations and his role 7.5.1 Habitat and Niche
in community, this world
be his niche. The habitat of an organism is where it lives, you could say its mailing address. When
we describe an organism’s habitat it is best to be specific and describe in as much
detail as possible the exact location.

Although the word niche recalls to mind a small space, in ecology it means much
more. The niche of an organism refers to the role it plays in the ecosystem. It includes
tolerance of physical factors such as temperature, light, soil, moisture, pH and nutrient
requirements. It also includes biological aspects such as - how it acquires its food,
what season of the year it reproduces and how it interacts with other organisms in the
community. In short, the niche defines a particular species role in the community, and is
unique for each species. Table 7.1 lists some of the factors that should be taken into
account when describing the niche of a species.
Table 7.1: Aspects of Niche
Plants Animals
Season of year for growth Time of day for feeding and season
and reproduction for reproduction

Sunlight, water, soil, pH, Habitat and food requirements

temperature requirements

Relationship with other organisms Relationships with other organisms

Effect on abiotic environment Effect on abiotic environment

In a community two niche dimensions usually taken into consideration are –

niche width and niche overlap (Fig. 7.5). Niche width or niche size is the sum
total of the different resources exploited by the organism. Measurement of niche
usually involve the measure of some ecological variable such as food size or
habitat space. Niche width may be described as narrow or broad. A narrow width
indicates a specialized species while a broad width would indicate that the species
is a generalist and can use a wide range of resources. Niche overlap indicates that
two or more species use a portion of the available resources, such as food, or
space, simultaneously. You can see in the figure that some niche space is shared
140 and some is exclusive.
The concept of niche suggests that related species that differ with respect to niche Community Organisation and
Interactions Amongs
are able to coexist in a habitat or community because their niche difference Organisims
partitions the resources and thus competition is avoided (Resource partitioning).
This is true for both plants and animals and we shall discuss this in more detail in
this section.

Fig. 7.5: Niche parameter in a community.

7.5.2 Functional Roles and Guilds

To find out how a community is organized we can use several approaches. The
simplest approach would be to group the species present in the community
according to feeding habits. In this way we can get information about the food
web and then each trophic level into guilds. These guilds are groups of species
exploiting a common resource base in a similar fashion. For instance, all organisms
like monkeys, parrots or other birds that eat fruit in a forest can form a guild; or
ants, rodents, and birds that eat seeds in a desert habitat can form a single guild.
Guilds may serve to pinpoint the basic functional roles of the species and different
guilds interact amongst themselves and provide the organisation that we see in a
community. The guild is a convenient unit for studies of interactions between
species in a community and should help us to study its organisation better by
making it unnecessary to study each and every species as a separate entity.

7.5.3 Keystone Species

When the activities of a species determine community structure that species is
called keystone species. For example, consider the case of the starfish, Pisaster
ochraceous. When this starfish was removed from the rocky intertidal areas of
western North America, the mussel Mytilus californianus was able to occupy the
space and excluded other invertebrates and algae which require attachment sites.
However, under natural conditions, predation of mussels by starfish keeps their
population under control and does not allow it to become dominant. This permits
other species requiring attachment sites to survive in such habitats.

Another example of a keystone species could be of the African elephant. By their

feeding habits elephants destroy shrubs and small trees and push woodland habitats
towards open grassland. Large mature trees can be destroyed by elephants feeding
on their bark. As more grasses invade the woodland habitats, the frequency of
fires increases, which accelerates the conversion of woods to grassland. This
141
Fundamentals of Ecology works to the disadvantage of the elephant, because grass is not a sufficient diet for
elephants, and they begin to starve as woody species are eliminated. However,
other ungulates that graze the grasses are favoured by the elephants’ activities.
Thus, in this community elephants play an important role in shaping the community
organisation.
Keystone species may be relatively rare in natural communities or may not be
easily recognized. At present, few terrestrial communities are believed to be
organized by keystone species, but in aquatic communities, keystone species may
be common.

7.5.4 Dominant Species

These are recognized by their greater number or more biomass and in general, are
described separately for each trophic level. Dominance is related to the concept of
species diversity. This means that some of the measures of species diversity could be
considered as measures of dominance. we may thus define simple community
dominance index as follows:

Community dominance index = percentage of abundance contributed by the two

most abundant species.
y1 + y2
=
y
Where y1 = abundance of most abundant species
y2 = abundance of second most abundant species
y= total abundance for all species
Abundance may be measured by density, biomass or productivity. Dominance,
however, is not always closely related to diversity. Dominant species are usually
assumed to be competitive dominants. However, dominant species in some
communities may be a matter of chance. For example, of the many invertebrates,
species present in a rotting log community, many species may be dominant at
least once but none would be dominant in every fog. Thus, a species could be
dominant in one log while very rare in the adjacent log. It seems that the
determining factor is very much a question of who gets there first.
Available data suggests that dominance can be achieved in three ways; i) the
first species to reach a new resource like in a rotting log, may become dominant,
ii) a species may become dominant by specializing on one part of a resource set
that is widely distributed, and iii) a species may generalize, so that it could use a
wide variety of resources.
Dominance is an important component of community organization, although it
is still poorly understood. Dominant species may affect not only the organisation
of a community but also its stability. Therefore, let us now investigate community
stability.

7.5.5 Stability
It is a dynamic concept that refers to the ability of a system to absorb change and
return back from disturbance. Let us look at figure 7.6. It explains the concept of
stability in an ecosystem. The black ball represents the community on a surface
which represents environmental conditions. In (a) the community is stable as the
142
system will return back to point I after disturbance. In (b) the community is locally Community Organisation and
Interactions Amongs
stable but if perturbed beyond a limit it will move to other positions of relative Organisims
stability (II and III). In (c) large disturbances will cause extinction of some species
and recolonization by newer species.

Fig. 7.6: Stability concept in an ecosystem.

Thus stability is a dynamic concept but is equilibrium centered. There must be

one or more equilibrium limits or points at which the system remains when faced
with a disturbing force. Stability may be local or global. Local stability is the
tendency of the system to return to its original position from a small disturbance;
gaps in forests filling in with similar species of trees are examples of local stability.
Global stability is the tendency of a community to return to its original condition
from all possible disturbances. Eucalyptus forest’s return to its original condition
after an outbreak of fire represents global stability. Communities show resistance
and resilience. Resistance is a measure of the degree to which a system is changed
from an equilibrium state after disturbance. Resilience is the speed with which a
perturbed system returns to equilibrium. A rapid return is an evidence of high
resilience.

The simple and appealing notion that diversity of species causes stability is
incorrect. ln fact increasing complexity reduces stability in mathematical models
and therefore, if diversity causes stability as is often said for tropical communities,
it is not an automatic consequence of species interactions. Natural communities
are products of evolution where non-random combination of interacting species 143
Fundamentals of Ecology is produced in which diversity and stability are related. The stability of whole
communities has rarely been studied in detail in spite of the great number of
perturbations caused by man.

Aquatic communities have been disturbed by pollution of human origin and the
stability of aquatic systems under pollution stress is a critical focus of applied
ecology today. It is important that we acquire information on how much we can
perturb the community I before it changes to a less desirable configuration. At
present, it is done by trial-and-error method only.
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.
State whether the given statements are true or false. Justify your answer where
you think the statement is false.
1. A guild consists of taxonomically similar organisms sharing the same food
resource.
......................................................................................................................
......................................................................................................................
......................................................................................................................
......................................................................................................................
2. Basically, a niche is the functional role of an organism in a community.
......................................................................................................................
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......................................................................................................................
3. Habitat is synonymous with the niche of an organism.
......................................................................................................................
......................................................................................................................
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4. Keystone species commonly determine the structure of aquatic communities.
......................................................................................................................
......................................................................................................................
......................................................................................................................
5. Stability in a community means that the community faces no changes.
......................................................................................................................
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144
 Community Organisation and
7.6 SPECIES INTERACTION Interactions Amongs
Organisims
Now that we have learnt how communities are organized. Let us see how
populations of plants and animals interrelate within the community. Individuals in
a species population interact amongst themselves – intraspecific interactions as
well as with individuals of other species population – intraspecific interactions.
Some have minimal influence on one another while some such as parasites and
their hosts, predators and their prey, have very distinct and immediate relationships.
At an individual level these relationships can be harmful or beneficial; at a
population level they can reduce, stabilise or enhance the rate of population growth.

The effects of these interactions can be positive, negative or neutral (see Table
7.2). Neutral interactions (OO) have no effect on the growth of population.
Positive interactions (+ +) benefit both populations and if the relationship is
mutually detrimental then the interaction is negative (– –). When one species
maintains or provides a condition necessary for the welfare of the other but does
not affect its own well-being by doing so, the interactions is called commensalism
(O+). An example is an epiphytic plant growing on the trunk of a tree. The tree
provides support and the epiphyte gets its nourishment through its aerial roots.
The interaction (O–) in, which one species reduces or adversely affects the
population of another but remains unaffected itself is called amensalism. An
example may be the release of toxic substances by one organism that inhibits the
growth and survival of another. This is known as allelopathy. An example is
juglone a chemical substance released into the soil by black walnut tree which
suppresses the growth of other plants near it. Amensalism may be considered a
form of competition.
Table 7.2: Interspecific Relationships
Type of Interaction Species Nature of Interaction
1 2
Neutralism o o Neither population affects the other
Competition – – Direct inhibition of each species by the other
Predation + – Population 1 predator kills prey species 2
(including herbivory)
Parasitism + – Population 1 parasite lives on population 2
prey without killing it
Commensalism + o Population 1 commensal benefits population
2 is unaffected
Mutualism + + Interaction favourable to both
Amensalism o – Population 1 unaffected population 2 harmed

Ecologically more important than both amensalism and commensalism is the

relationship which benefits both population (+ +). Such interactions are termed
mutualism. For example, bacteria present in termites and stomachs of ruminant
animals help in the digestion of cellulose. The bacteria get a warm environment
and help their host to extract nutrients. The nitrogen fixing bacteria Rhizobium
found in the root nodules of leguminous plants are another common example of
mutualism.

145
Fundamentals of Ecology Negative interactions are competition (– –), which is detrimental to populations
of both species and predation and parasitism (– +), in which the population of
one species benefits at the expense of another. Parasitism involves one organism’s
feeding on another and the prey or ‘host’ is seldom killed outright. The host survives,
though its fitness is reduced and when it dies it is due to reduced resistance to
other infections. Familiar parasites are tapeworm, fleas, Plasmodium and
numerous other disease-causing microorganisms.

Ecologists have studied negative interactions involving competition and predation

much more than positive ones, because impacts of positive interactions are not
easily demonstrated. In this unit we will study interactions between two species
involving competition and predation.

7.7 COMPETITION
Competition occurs over resources. For plants light, nutrients, and water may be
important resources. Plants may compete for pollinators or for attachment sites. Water,
food and mates are possible resources for animals, and they may compete for space
such as nesting sites, wintering sites or places that are safe from predators. Thus, we
see that resources can be complex and diverse.
• There are two types of competitive interactions: Exploitative or scramble
competition occurs when a number of organisms of same or different species
that are in short supply.
• Interference or contest competition occurs when organisms seeking a resource
will harm one another in the process even if the resource is not in short supply.
When a shared resource is sufficient like oxygen in terrestrial environment or in the
aquatic habitat there is no competition for it amongst organisms. But most resources
are generally in short supply, therefore, organisms with niche overlap enter competition.
The greater the niche overlap, the more intense the competition (Fig.7.7). Because
members of the same species require many of the same resources intraspecific
competition is more intense than interspecific competition between members of
different species.

Fig. 7.7: Niche overlap and intensity of competition. Each graph compares food and habitat
requirements for two species A and B.
146
Simplified communities and laboratory experiments allow ecologists to single out and Community Organisation and
Interactions Amongs
study qualitatively the various interactions. Mathematical models have been used Organisims
extensively to build up hypothesis about what happens when two species live
together either sharing the same food or occupying the same space or preying on
one another. The best-known models were developed independently by two
mathematicians Lotka (1925) in U.S. and Volterra (1926) in Italy. Lotka-Volterra
equations apply to predator-prey situations and non-predatory situations involving
competition for food and space.

The Lotka-Volterra equations for competition between organisms based on the

logistic growth equation, one for each species, can be written in the following
form.
dN1 K1 - N1 - αN2
= r1 N1
dt K2

dN2 K2 - N2 - βN1
= r2 N1
dt K2

where N1, N2 = population of species 1 and 2

K1, K2 = Equilibrium population size for each species in the absence of
the other
t = time
r1, r2 = per capita rate of increase of species 1 and 2 respectively.
α is a constant representing the inhibitory effect of species 2 on species 1 and β is
the constant signifying the inhibition of species 2 by species 1. Lotka-Volterra
models assume that: 1) the environment does not change, 2) migrations are
unimportant, 3) coexistence requires stable equilibrium points, 4) competition. is
the only important biological interaction. Now if we put these two species N1 and
N2 together what might be the outcome of this competition?
1) both species coexist
2) species 1 becomes extinct or
3) species 2 becomes extinct
The theoretical Lotka-Volterra equations stimulated studies on competitions in the
laboratory where under controlled situations the outcome is easily determined.

7.7.1 Competition in Laboratory Populations

Although species with small niche differences are able to co-exist in a community,
those with identical niches cannot, even if only one shared resource is in short
supply. Competition becomes so intense that one species is finally eliminated.
Greater number of offsprings of the ‘winner’ species with better suited traits
gradually displace members of the less efficient species. This is known as the
competitive exclusion principle. It was first demonstrated in the laboratory by
G.F. Gause (1934) in mixed cultures of closely related paramecium species.
Although each population survived when grown individually, only one survived
when grown together with a fixed amount of food (Fig.7.8).

147
Fundamentals of Ecology

Fig. 7.8: Competition between two species of paramecium. When grown in pure culture,
P aurelia and P caudatum exhibit rapid growth. When grown in mixed cultures
P aurelia is the better competitor and P caudatum dies out. (After Gause 1934).

Competition between species does not always lead to expansion of one population
and restriction of another. Gause showed in another experiment that when two
different species of paramecia P. aurelia and P. bursaria occupy the same tube,
both survived because P. aurelia could feed on the yeast suspension in the upper
layers of the fluid whereas P. bursaria could feed on the yeast in the bottom
layers. This difference in the feeding behaviour between these species allowed
them to coexist. It was thus demonstrated that two or more similar species can
live together only if their niches differ.

7.7.2 Competition in Natural Populations

Now let us see if these laboratory results apply to populations in nature. There is
a wide range of opinion on the importance of the competitive exclusion principle.
Although this principle has been repeatedly demonstrated in laboratory
experiments, it is not the rule in nature or rather it is not easy to see in natural
communities.

Let us first examine situations where competitive exclusion would not be expected
to occur. These situations are:
1) When the critical resource is in abundance. For example, six species of the
leafhopper Erythoneura can live on the same tree and feed on the same leaves.
Not only are their habitats and food source same but their life cycle phases
are similar. Apparently competitive exclusion is avoided because of an
abundance of resources.
2) When environmental conditions are unstable and change frequently. There is
just not enough time for one species to replace the other during the short
148
 period ‘ when resources become limited. For example, in oceans and temperate Community Organisation and
Interactions Amongs
lakes, changes sometimes occur so suddenly that there is no time for one species Organisims
of phytoplankton to increase so much in numbers so as to exclude the other,
despite intense competitions for limited nutrients. Because steady changes
take place during primary and secondary succession, competitive exclusion
has not been seen in communities undergoing succession.
Another reason why the process of competitive exclusion may go unnoticed in
nature is that is takes time for one species to exclude another. If researchers are
unable to observe the community continuously then they may miss the process
entirely. There is an interesting example to illustrate this. Goats were introduced
on the island of Abingdon in the Galapagos Archipelago in 1957. The goats browsed
on the same low-growing grass as the native tortoise, as well as other leaves and
higher stems. In the absence of any predators, the goats multiplied rapidly and the
low growing food that the tortoise required got exhausted. By the time the research
team revisited in 1962 all the tortoise were gone. Here competitive exclusion had
caused the extinction of the Abingdon tortoise.

7.7.3 The Results of Competition

Competitive exclusion is not the only outcome of competition. Sometimes a shared
resource can be partitioned in such a way that potential competitors use different
portions of the resource. Let us take a hypothetical case of 4 species of birds that live
in a similar habitat. They would feed on different position in the canopy and thus avoid
competition. In addition to partitioning of space, a shared resource may be exploited
at different times.

Fig. 7.9: Closely related Phalacrocorax species get their living in different ways and so
do not compete.

149
Fundamentals of Ecology These are example of resource partitioning. A study involving closely related
species of birds to test Gauses’ hypothesis was done on the cormorant
(Phalacrocorax carbo) and the shag (P. aristotelis), see Fig. 7.9. These species
occur in similar habitats and appear to have a wide niche overlap. They are both
cliff nesters and eat fish. It was shown that coromorant nests chiefly on flat broad
cliff edges and feeds chiefly in shallow estuaries and harbours; the shag nests on
narrow cliff edges and feeds mainly out at sea. Thus, because of these differences,
competition is minimized.

A similar partitioning of resources exists among plants. Species of plants cultivated

together exploit different soil depths (Fig. 7.10): Some have shallow fibrous roots
that draw water from the soil top. Another species may have sparsely branched
taproot that extends to an intermediate depth. Yet another may have a taproot that
is moderately branched in the upper layers but develops primarily below the
rooting zone of the other species.

In addition to spatial partitioning a shared-resource may be exploited at different

times. This is known as temporal partitioning. An example can be seen in
grasslands where a species of buttercup Ranunculus grows only in early spring
before competing perennial grasses begin to grow.

Fig. 7.10: Resource partitioning of the soil by a group of desert plants. Root system
morphology is species specific. Species (A) are shallow surface rooters, able
to take up moisture quickly during occasional rains. (B) have more spreading
roots at intermediate depths. Plants such as (C) hove deep tap roots.

Another alternative to competitive exclusion is character displacement where

intense competition affects evolution, leading to displacement or change in a
characteristic rather than extinction of a species. An example is seen in the forests
of Europe, where six species of a small bird called titmice (Parus) coexist because
each species has a slightly different beak size. These differences in beak size
prevent any two species from seeking the same food in the same feeding area. For
example, titmice with longer beak catches larger insects. Biologists have found
out that all the six species evolved from an identical ancestor and that variations
in beak size are the result of character displacement.

150
7.7.4 Evolution of Competitive Ability Community Organisation and
Interactions Amongs
Organisims
It would be obvious from the examples discussed in earlier subsections that if
two species are competing for a resource that is not abundant, then it would be In animals, aggressive
advantageous to both species to evolve differences – structural, physiological behaviour leads to
and behavioural – that would reduce competition. However, it is not always establishment of
possible to evolve such mutually beneficial changes because the species may dominance hierarchies
have other possible competitors. or rank order. The
dominance is
So, the only way to survive is to evolve competitive ability or ‘stay and fight’. determined by actual
This concept is not very easy to define. But in short it means that any mechanism fighting or a ritual
that prevents a competitor from gaining access to a limited resource will improve contest in which one
frightens the other
competitive ability. Territorial behaviour in birds is a good example. In animals,
away. The dominant
the evolution of a broad array of aggressive behaviour has been critical in
animal eats and mates
substituting ability in combat for ability in utilising resource in competition. On first, and drives the
the basis of this we can recognize an idealized evolutionary gradient. other away from his
territory.
Low density
leads to Colonisation and growth

High density
leads to Resource competition

High density
leads to Interference mechanisms that
prevent resource competition.

Check Your Progress 3

Note: i) Use the space given below for your answers.

ii) Check your answers with those given at the end of the unit.

1. According to the principle of competitive exclusion, species cannot remain

the same community if they have the same a) taxonomy, b) habitat, c) food
requirement, d) niche.
2. a) When members of the same or different species exploit common
resources that in short supply …………………. competition occurs.
b) ………………….. competition occurs when both individuals have
access to resources but one individual use more of it or more efficiently
than the other.
3. Species can avoid competition in the community if they …………………..
resources. Whereas, intense competition often leads to …………………..
……………….. rather than extinction of the species.

7.8 LET US SUM UP

In this unit, you have learnt some aspects of nature and structure of community. So
far you have learnt that:
• Communities are made up of population of organisms, occupying and
interacting in a given area. They constitute the biotic component of ecosystems.
151
Fundamentals of Ecology • Communities have several group characteristics which are not exhibited by
either its individuals or populations.
• The size of community may vary. Just like an ecosystem, a bigger community
too may be sub-divided into smaller communities.
• Based on the source of energy, a community may be autotrophic or
heterotrophic.
• Rarely, can different communities be sharply delimited, because they blend
together to form a continuum along some environmental gradient. Sometimes,
because of severe environmental disturbance(s), sharp boundaries between
the communities can be seen.
• The area where two communities blend is an ecotone. This zone has a high
species richness. It not only supports the species of the adjoining two
communities, but also a few species found exclusively in this zone.
• Communities can be organized by competition, predation and symbiosis among
organism working within the physical environment. Two important concepts in
community ecology are habitat and niche. Habitat is where the organism lives and
niche is its functional role in the community.
• Species in a community are organized into food webs and further into groups or
guilds that exploit a common resource base. Keystone species are those
species that can determine the community structure and can be recognized by
removal experiments. Dominant species are those that have maximum
abundance or biomass in a community and may affect the stability of the
community.
• Stability is the ability of a system to come back to the original condition after
a disturbance. The ecological generalization that diversity causes stability is
not supported by field or laboratory studies.
• There are two opposing views on community organisation. The equilibrium
hypothesis suggests that natural communities are stable and the nonequilibrium
hypothesis suggests communities are never stable and always recovering
from disturbances. High species richness is maintained in communities that
face moderate disturbances.
• The organisms of a community interact in a variety of ways that may be classified
as negative, positive or neutral interactions. Competition occurs over resources.
Theoretical models indicate that in case of competition between similar species,
one species may be displaced or both may reach a stable equilibrium. Possibility
of displacement gave rise to the competitive exclusion principle which states that
complete competitors cannot coexists. Species can coexist in the same community
only if they have slightly different niches. Intensity of competition increases with
niche overlap.
• The results of competition are resource partitioning and character
displacement. Organisms evolve competitive ability by becoming more
efficient resource users and by developing interference mechanisms that keep
competing species from using the same resources.

152
Check Your Progress 4 Community Organisation and
Interactions Amongs
Note: i) Use the space given below for your answers. Organisims

ii) Check your answers with those given at the end of the unit.
1. Comment on the following statement”
“Community is an association of interacting populations.”
......................................................................................................................
......................................................................................................................
......................................................................................................................
2. Fill in the blank spaces with appropriate words:
a) The area where two adjacent communities blend is an ………………….
b) A …………………………. Refers to an area of vegetation having
species of same kind and age.
c) In situations where there is a wide range of environmental conditions at
the junction of two communities ………………….. is seen.
3. Fill in the blanks with suitable community interactions.
both unaffected ……(a)
both harmed ……(b)
ORGANISM INTERACTIONS
one harmed ……(c)
one benefits ……(d)
one benefits ……(e)
one unaffected ……(f)
both benefit ……(g)
4. A certain island has two closely species of birds, one of which has a slightly
larger beak than the other. Interpret this finding with respect to competitive
exclusion principle and ecological niche, defining both terms.
......................................................................................................................
......................................................................................................................
......................................................................................................................
......................................................................................................................

5. Why would you recommend that animals such as wild dogs be not killed
even though they sometimes prey on farm animals?
......................................................................................................................
......................................................................................................................
......................................................................................................................
......................................................................................................................
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Fundamentals of Ecology
7.9 KEY WORDS
Ecotone: The zone of vegetation separating two different types of communities is
called ecotone. It is also known as a transition zone.
Niche: The set of environmental conditions and patterns of resource availability, in
which a species can survive, grow, and reproduce.

7.10 SUGGESTED FURTHER READING/

REFERENCES
Odum EP. 1971. Fundamentals of Ecology. 3rd edition. W.B. Saunders company,
Philadelphia.
Sharma PD. 2007. Ecology and Environment. Tenth revised edition. Rastogi
Publications, Meerut.
Singer FD. 2016. Ecology in Action. Cambridge University Press, United Kingdom.

7.11 ANSWERS TO CHECK YOUR PROGRESS

Check Your Progress 1
1. b
2.

3. a) represents biotic components of an ecosystem

b) consists of organisms with interlocking food chains
c) may vary in size
d) concept can be applied to any scale
e) may be autotrophic or heterotrophic
154 f) may consist of stand(s)
Check Your Progress 2 Community Organisation and
Interactions Amongs
1. False, a guild consists of groups of species that may not be taxonomically Organisims
related but have similar or comparable feeding roles in a community.
2. True.
3. False. Habitat describes the place where an organism is to be found. Niche,
however, includes the physical space the organism occupies as well as its
functional role in the community.
4. True.
5. Stability in a community means its ability to return to the original condition
after disturbances. It does not mean that stable communities never face any
fluctuations.
Check Your Progress 3
1. d
2. (a) scramble. (b) contest.
3. partition, character displacement.
Check Your Progress 4
1. Hint: Community is an assemblage of populations of various kinds of
organisms in a prescribed area. Community is the biotic component of an
ecosystem. And in an ecosystem, the various kinds of organisms are related
in terms of food and they form a food web. So, the populations interact with
each other directly or indirectly not any for food, but also influence the
existence of other populations.
2. a) ecotone
b) stand
c) edge effect
3. a) neutralism
b) competition
c) predation
d) parasitism
e) herbivory
f) commensalism
g) mutualism
4. Competitive exclusion principle states that no two species can occupy the
same niche. If closely related species are to survive, they must show some
difference, however, small in their niches. Ecological niche refers to the role
that the organism plays in the ecosystem. The related species of birds coexist
because a part of their niches i.e., beak size, differ so they cannot exploit the
same food resource.
5. In nature predator-prey populations often show oscillations. Predators help
control prey population. If all the wild dogs are removed then their natural
prey species like rodents, rabbits etc. will increase tremendously and cause
damage to the food crops.
155
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:
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 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.
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2. State the differences between primary and secondary succession.
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163
Fundamentals of Ecology 3. Explain the stages in hydrosere.
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4. Describe the sequential stages in lithosere.
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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.
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2. Explain the ecological adaptation in hydrophytes.
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3. Describe the characteristic adaptation features of mesophytes.
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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
Fundamentals of Ecology
UNIT 9 MAJOR BIOMES OF THE WORLD

Structure
9.1 Introduction
9.2 Objectives
9.3 Major Biomes of World
9.4 Classification of Biomes
9.4.1 Aquatic Biomes
9.4.1.1 Freshwater Biomes
9.4.1.2 Marine Biomes
9.4.2 Terrestrial Biomes
9.4.2.1 Forest Biomes
9.4.2.2 Grassland Biomes
9.4.2.3 Woodlands and Shrublands
9.4.2.4 Taiga
9.4.2.5 Tundra
9.4.2.6 Desert
9.5 Let Us Sum Up
9.6 Key Words
9.7 Suggested Further Reading/References
9.8 Answers to Check Your Progress

9.1 INTRODUCTION
Biomes are large geographical areas. Biomes are defined as “the world’s major
communities, classified according to the predominant vegetation and characterized
by adaptations of organisms to that particular environment”. Biomes are also
defined by factors such as plant structures (such as trees, shrubs, and grasses),
leaf types (such as broadleaf and needleleaf), plant spacing (forest, woodland,
savanna), and climate. In this unit, we would be discussing the major biomes of
world and characteristic features of the aquatic and terrestrial biomes.

9.2 OBJECTIVES
After studying this unit, you should be able to:
• explain the major biomes of world;
• classify the biomes; and
• explain the aquatic and terrestrial biomes.

9.3 MAJOR BIOMES OF WORLD

Biomes are basically large geographical areas. Within a biome, there are many
interrelated ecosystems. Ecosystems can be classified into biomes because
climate, along with other factors, determines the plant growth form best suited
to an area, and because plants with particular growth forms are restricted to
particular climates. These principles establish the close relationship between
172
climate and vegetation. Different physical conditions characterize each biome Major Biomes of the World
and its inhabitants are adapted to live under those conditions. Biomes are defined
as “the world’s major communities, classified according to the predominant
vegetation and characterized by adaptations of organisms to that particular
environment”. A biome (from the Greek “bios” life), also called bioclimatic
landscape or biotic areas is a particular part of the planet that shares the climate,
flora and fauna. Biomes are defined by factors such as plant structures (such as
trees, shrubs, and grasses), leaf types (such as broadleaf and needleleaf), plant
spacing (forest, woodland, savanna), and climate.

Many different environmental factors play a role in determining which biomes

are found where. Two key parameters are available moisture and temperature.
These biomes generally differ in their latitude, weather and relative humidity,
amount of sunlight, and topography. Each biome has a unique set of plants and
animals that thrive in its climate. A biome is different from an ecosystem. An
ecosystem is the interaction of living and non-living things in an environment. A
biome is a specific geographic area notable for the species living there. A biome
can be made up of many ecosystems. Similar biomes can occur in widely divergent
places as long as the environmental conditions are appropriate for their
development.

9.4 CLASSIFICATION OF BIOMES

Not all scientists classify biomes in the same way. Some use broad classifications
and count as few as five biomes. These are forest, grassland, freshwater, marine,
desert, and tundra.

Other scientists use more precise classifications and list dozens of different biomes.
For example, they consider different kinds of forests to be different biomes. Many
classification schemes for biomes exist. Biomes are fundamentally classified into a)
Terrestrial (land) biomes and b) Aquatic biomes (including freshwater biomes

Fig. 9.1: Whittaker’s Classification of Biomes (Source: Whittaker, 1975) 173

Fundamentals of Ecology and marine biomes). Robert H. Whittaker defined biomes first by their vegetation
type, and then devised a simple climate diagram. (Fig. 9.1) He based his
classification scheme on two abiotic factors namely precipitation and temperature.
Heinrich Walter classification scheme (Fig. 9.2) considers the seasonality of
temperature and precipitation. In this scheme, nine major biome types with
characteristic climate traits and vegetation types are defined. The biome boundaries
correlate to the conditions of cold stress and moisture. In fact, the moisture and
cold stress are determinants of plant form. So, the vegetation defines the region.
However, extreme conditions like flooding can create different kinds of
communities within the same biome.

Fig. 9.2: Heinrich Walter classification scheme (Source: Walter and Box, 1976)

9.4.1 Aquatic Biomes

The aquatic biome can be broken down into two basic regions namely freshwater
biomes and marine biomes. Freshwater biome consists of the lakes, ponds, rivers,
streams, and wetlands. The marine regions include oceans, coral reefs and
estuaries. The marine biome consists of the oceans, seas, coral reefs and estuaries.
These are habitats with high saline conditions. In these habitats we find
mangroves, salt marshes, and mudflats. Marine habitats consist of five zones -
intertidal, neritic, oceanic pelagic, abyssal and benthic zones. Freshwater biomes
are aquatic habitats with low saline conditions. The freshwater habitat is classified
into lotic (moving water bodies) and lentic (standing water bodies). Lotic
form includes streams and rivers. Lentic forms include bogs, lakes, ponds, and
inland wetlands. Freshwater habitats are also influenced by land of the surrounding
areas, the pattern and speed of water flow and the climatic condition.

9.4.1.1 Freshwater Biomes

Freshwater biomes have a salt content (or salinity) of less than 0.05%. On the
174 other hand, marine biomes have a salinity of 3.5% or greater. Freshwater biomes
are classified according to the movement of its water. Biomes in which the water Major Biomes of the World
is not flowing are classified as lentic freshwater biomes while those that contain
freely flowing water are known as lotic freshwater biomes Examples of lentic
freshwater biomes are lakes and pond. Examples of lotic freshwater biomes are
rivers and streams.

9.4.1.2 Marine Biomes

Marine regions cover about three-fourths of the Earth’s surface. It includes oceans,
coral reefs, and estuaries. The largest of all the ecosystems, oceans are very large
bodies of water that dominate the Earth’s surface. Like ponds and lakes, the
ocean regions are separated into separate zones: intertidal, pelagic, abyssal, and
benthic. All four zones have a great diversity of species. It is said that the ocean
contains the richest diversity of species even though it contains fewer species
than there are on land. Estuaries are bodies of water where salt and freshwater
mix. Apart from having considerable scientific interest because of their unusual
properties, they are very important to humans. Estuaries frequently make good
ports. They tend to be very productive and have large fish and shellfish yields.
Coral reefs are widely distributed in warm shallow waters. They can be found as
barriers along continents (e.g., the Great Barrier Reef off Australia), fringing
islands, and atolls. Naturally, the dominant organisms in coral reefs are corals.
Corals consist of both algae (zooxanthellae) and tissues of animal polyp. Since
reef waters tend to be nutritionally poor, corals obtain nutrients through the algae
via photosynthesis and also by extending tentacles to obtain plankton from the
water. Besides corals, the fauna include several species of microorganisms,
invertebrates, fishes, sea urchins, octopuses, and sea stars.

9.4.2 Terrestrial Biomes

9.4.2.1 Forest Biomes

i) Tropical Rain Forest

Tropical forests are distributed in areas of the world near the equator and between
the Tropic of Capricorn and the Tropic of Cancer. Regions which receive heavy
rainfall all year round, with no distinct dry season, are referred to as tropical
rainforests. One of the major characteristics of tropical forests is their distinct
seasonality: winter is absent, and only two seasons are present (rainy and dry).
They are characterized by very hot temperatures and high average annual rainfall
(greater than 1500 mm per annum). They form habitat for a wide variety of species.
There are extensive tropical rain forests in South America, Africa, and Southeast
Asia. Soil in rainforest is highly acidic, the type of clay particles present in tropical
rainforest soil has a poor ability to trap nutrients and stop them from washing
away, the dead organic matter decompose quickly leading to low nutrient content
of the soil. Canopy in tropical forests is multilayered and continuous, allowing
little light penetration. Vegetation is dominated by a continuous canopy of tall
evergreen trees rising to 30–40 m. Occasional emergent trees rise above the
canopy to heights of 55 m or so. Climbing lianas, or woody vines, and epiphytes,
plants that grow on the branches of other plants and are not rooted in soil are
prominent in the forest canopy itself.
Rainforest can be further classified on the basis of seasonal distribution of rainfall:
Evergreen rainforest: with no dry season; Seasonal rainforest: short dry period
in a very wet tropical region; Semi-evergreen forest: longer dry season (the 175
Fundamentals of Ecology upper tree storey consists of deciduous trees, while the lower storey is still
evergreen); Moist/dry deciduous forest (monsoon): the length of the dry season
increases further as rainfall decreases (all trees are deciduous). Broad-leaved
evergreen trees; ferns; large woody vines and climbing plants; orchids and
bromeliads are commonly found. Dominant wildlife includes herbivores such
as sloths, tapirs, and capybaras; predators such as jaguars; anteaters; monkeys;
birds such as toucans, parrots, and parakeets; insects such as butterflies,
ants, and beetles; piranhas and other freshwater fishes; reptiles such as frogs,
Caymans, boa constrictors, and anacondas.

ii) Temperate Evergreen Forest

These are characterized with mild winters, heavy winter rains, and summer
fog create conditions that support extremely tall evergreen forests. Forests in
the northern hemisphere are dominated by oaks, laurels and camphor, while
species of southern beech and eucalypts dominate the south. Soil is fertile,
enriched with decaying litter. Canopy is moderately dense and allows light
to penetrate, resulting in well-developed and richly diversified understory
vegetation and stratification of animals. temperate forests have just two layers
of vegetation. The tallest trees have their foliage generally about 15-30 m
above ground and a layer of shrubs and smaller trees underneath, at
approximately 5-10 m. This is why the soil receives more light than in tropical
forests and the undergrowth is luxuriant: ferns, mosses and lichens, especially
in very rainy areas. Temperate-mixed evergreen forests represent a broad
transitional zone between temperate deciduous forests to the south and taiga
to the north.

iii) Temperate Deciduous Forest

Mild climates (warm summers and cool winters) and plentiful rains promote
the growth of deciduous (hardwood) forests in Eurasia, the northeastern United
States, and eastern Canada. It has four distinct seasons: winter, spring, summer
and fall. Winters are cold and summers are warm. The average annual
temperature in a deciduous forest is 50° F. The average rainfall is 30 to 60
inches a year. Soil tends to be slightly acidic and moderately leached, and
are brown in color owing to abundant organic matter. Warmer and drier parts
of the temperate seasonal forest biome, especially where soils are sandy and
nutrient poor, tend to develop needle-leaved forests dominated by pines.
Conifers like spruce, fir and pine trees can also be found mixed in with the
hardwood trees in this biome. Higher species diversity than coniferous forest
deer, wolf, bear, foxes, many migratory birds, squirrels, rabbits, etc. are
commonly found. Some species hibernate in winter.

9.4.2.2 Grassland Biomes

i) Temperate Grassland
Temperate grasslands are characterized as having grasses as the dominant
vegetation. Trees and large shrubs are absent. rainfall ranges between 30
and 85 cm per year, Summer temperatures can reach over 38 °C, while in
the winter are cold they can sometimes reaching below -40 °C. Precipitation
is infrequent, so organic detritus does not decompose rapidly, and the soils
are rich in organic matter Extensive grasslands are also found in central Asia,
where they are called steppes. Steppes are dry areas of grassland with hot
176 summers and cold winters. In North America, grassland is called a prairie.
 In South America, it is called a pampas. And in Australia, grassland is called Major Biomes of the World
an outback. Prairies are grasslands with tall grasses while steppes are
grasslands with short grasses.

ii) Tropical Grassland/Savanna Biomes

Savannas are found in parts of the tropics where there is not enough rainfall
throughout the year to create a rainforest. Savanna is grassland with scattered
individual trees. On a global scale, the savanna biome is transitional between
tropical rain forest and desert. They are always found in climates with
relatively hot, wet summers and warm, dry winters where the annual rainfall
is typically 90–150 cm per year, but the driest three or four months bring
less than 5 cm each. Different savannas support different grasses depending
on the amount of rainfall received and soil conditions. The soil of the savanna
is porous, with rapid drainage of water. Nutrients in soil are present mainly
due to a thin layer of humus. Fire and grazing undoubtedly play important
roles in maintaining the character of the savanna biome. The woody trees
are very limited and usually do not get very tall. Grasslands usually support
large numbers of both small and large wildlife predators such as lions,
leopards, cheetahs, hyenas, and jackals; aardvarks; herbivores such as
elephants, giraffes, antelopes, and zebras; baboons; birds such as eagles,
ostriches, weaver birds, and storks; insects such as termites

9.4.2.3 Woodlands and Shrublands

This biome actually goes by several names. It is found surrounding large parts of
the Mediterranean Sea and is thus sometimes called the Mediterranean Biome,
others call it a Woodland Biome, while others call it a Shrub or Chaparral Biome.
Temperate woodlands and shrublands are located in Western coastal regions
between 30° and 40° North and South latitude. Around the Mediterranean Sea,
southern parts of Australia, and Mexico. The soil type is quite fertile, rich in
minerals and other nutrients. The most dominate plant life includes aromatic
herbs such as: sage, rosemary, and thyme, as well as shrubs and other grasses.
Some prominent wildlife are coyotes, wild goats, mule deer, and the
Mediterranean gecko. During the dry season lightning started wildfires wreak
havoc on the shrublands. Because of this some plants have adapted to have fire
resistant capabilities, by holding on the more moisture in their leaves. Some
plants only release these seeds during fires, the heat cracks them open and they
begin to grow. The shrublands are also known as Chaparral. It is from the Spanish
word “chapa” or scrub oak

9.4.2.4 Taiga

Stretching in a broad belt centered at about 50°N in North America and about
60°N in Europe and Asia lies the boreal forest biome, often called taiga. Long,
severe winters (up to six months with mean temperatures below freezing) and
short summers (50 to 100 frost-free days) are characteristic, as is a wide range of
temperatures between the lows of winter and highs of summer. The winters are
extremely harsh and can last as long as six months. Typically, the soil freezes
during the winter. This makes it difficult for animals to stay year-round.
Precipitation ranges between 25 and 100 centimeters (10 to 40 inches) per year.
Needle leaf coniferous trees are the dominant plants of the taiga biome. Coniferous
forests are mainly found in broad circumpolar belt across the northern hemisphere
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Fundamentals of Ecology and on mountain ranges where low temperatures limit the growing season to a
few months each year - thus too unfavorable for most hardwoods. Spruce, fir,
pine, larch or tamarack, alder, birch, and aspen are common. Throughout the
southern half of Canada, parts of northern Europe, and much of Russia, there is an
evergreen coniferous forest.

9.4.2.5 Tundra

Tundra is the coldest biome on Earth. The word tundra comes from a Finnish
word for treeless land. There are two types of tundra—Arctic tundra, found in a
band around the Arctic Ocean, and alpine tundra, found high in mid-latitude
mountains. Arctic tundra has a growing season of only 50 to 60 days. The average
winter temperature is –34 °C. Summer temperatures rarely exceed 12 °C. As a
result of these cold temperatures, the ground is permanently frozen from 25
centimeters to about 100 centimeters below the surface. This frozen ground is
called permafrost. There is a thin layer of soil above the permafrost that does
thaw in summertime, but it is not deep enough to support the growth of trees.
Lichens, mosses, grasses, and a few woody shrubs are the most common plants in
the Arctic tundra.

Floral biodiversity includes low shrubs, sedges, reindeer mosses, liverworts,

and grasses about 400 varieties of flowers, crustose and foliose lichen. Plants are
short and group together to resist the cold temperature. They can carry out
photosynthesis at low temperatures and low light intensities. The snow protects
them during the winter. The growing seasons are short, and most plants reproduce
by budding and division rather than sexually by flowering. The fauna in the arctic
is also diverse herbivores that include lemmings, voles, caribou, arctic hares and
squirrels’ Carnivorous mammals: arctic foxes, wolves, and polar bears. Migratory
birds: ravens, snow buntings, falcons, loons, ravens, sandpipers, terns, and snow
birds, and various species of gulls. Insects like mosquitoes, flies, moths,
grasshoppers, blackflies and arctic bumble bees. Fishes like cod, flatfish, salmon,
and trout. Animals are adapted to handle long cold winters and to breed and raise
young quickly in the summer. Animals such as mammals and birds also have
additional insulation such as fat. Many animals hibernate during the winter because
food is not abundant. Another alternative is to migrate south in the winter, like
birds do. Reptiles and amphibians are few or absent because of the extremely
cold temperature. Because of constant immigration and emigration, the population
continually oscillates.

9.4.2.6 Desert

A desert is a climate region that averages less than 35 centimeters of rainfall per
year. Most deserts are found between the latitudes of 30° N and 30° S. Because of
the lack of cloud cover, deserts receive more than twice as much incoming solar
radiation as humid regions. They also emit almost twice as much radiation at
night. As a result, deserts have large variations in daily high and low temperatures.
Soils are course-textured, shallow, rocky or gravely with good drainage and have
no subsurface water. They are coarse because there is less chemical weathering.
The finer dust and sand particles are blown elsewhere, leaving heavier pieces
behind. Soil tends to be well drained, with very little or absent organic matter.
However, they are rich in nitrogen and other minerals. There are 4 types of deserts
commonly found:
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i) Hot and Dry Desert Major Biomes of the World

The seasons are generally warm throughout the year and very hot in the summer.
The winters usually bring little rainfall. Many mean annual temperatures range
from 20-25° C. The extreme maximum ranges from 43.5-49° C. Hot and dry
deserts are warm throughout the fall and spring seasons and very hot during
the summer. The winters usually have very little if any rainfall. Hot and dry
desert vegetation is very rare. Plants are almost all ground-hugging shrubs
and short woody trees. All of the leaves store nutrients. Some examples of
these kinds of plant are Turpentine Bush, Prickly Pears, and Brittle Bush. For
all of these plants to survive they have to have adaptations. Some of the
adaptations in this case are the ability to store water for long periods of time
and the ability to stand the hot weather. Animals include small nocturnal
carnivores. The dominant animals are burrowers. The animals stay inactive
in protected hideaways during the hot day and come out to forage at dusk,
dawn or at night, when the desert is cooler.

ii) Semi-arid Desert

The summers are moderately long and dry, and like hot deserts, the winters
normally bring low concentrations of rainfall. Summer temperatures usually
average between 21-27° C. It normally does not go above 38° C and evening
temperatures are cool, at around 10° C. Cool nights help both plants and
animals by reducing moisture loss from transpiration, sweating and breathing.
Furthermore, condensation of dew caused by night cooling may equal or
exceed the rainfall received by some deserts. As in the hot desert, rainfall is
often very low and/or concentrated. The average rainfall ranges from 2-4 cm
annually.

iii) Coastal Desert

These deserts occur in moderately cool to warm areas such as the Nearctic
and Neotropical realm. A good example is the Atacama of Chile. The cool
winters of coastal deserts are followed by moderately long, warm summers.
The average summer temperature ranges from 13-24° C; winter temperatures
are 5° C or below. The maximum annual temperature is about 35° C and the
minimum is about -4° C. In Chile, the temperature ranges from -2 to 5° C in
July and 21-25° C in January. The average rainfall measures 8-13 cm in
many areas. The maximum annual precipitation over a long period of years
has been 37 cm with a minimum of 5 cm.

iv) Cold desert

These deserts are characterized by cold winters with snowfall and high
overall rainfall throughout the winter and occasionally over the summer. They
occur in the Antarctic, Greenland and the Nearctic realm. They have short,
moist, and moderately warm summers with fairly long, cold winters. The
mean winter temperature is between -2 to 4° C and the mean summer
temperature is between 21-26° C. Cold Deserts have quite a bit of snow
during winter. The summer and the beginning of the spring are barely warm
enough for a few lichens, grasses and mosses to grow. Most of the
precipitation comes in the form of snow or fog. Annual precipitation is less
than 250mm.More than 99% of the land area is covered by ice. Plant growth
is limited as there are few areas not covered by ice. There are no trees and
shrubs present at all. The only vegetation able to survive are liverworts,
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Fundamentals of Ecology lichens and mosses, few amphibians, reptiles or mammals are native to cold
deserts, but humans have introduced some animals, such as rats and mice.
Native fauna includes spiders, earthworms, beetles and the Arctic fox.

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 aquatic biomes.
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2. What are the characteristic features of tropical rain forest?
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3. Explain the characteristic features of temperate forests.
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4. List the characteristic features of temperate grassland biomes.
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5. Explain the characteristic features of desert biomes.
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 Major Biomes of the World
9.5 LET US SUM UP
Biomes are the world’s major communities, classified according to the predominant
vegetation and characterized by adaptations of organisms to that particular
environment. Different environmental factors play a role in determining which
biomes are found where. Biomes generally differ in their latitude, weather and
relative humidity, amount of sunlight, and topography. Each biome has a unique
set of plants and animals that thrive in its climate. Similar biomes can occur in
widely divergent places as long as the environmental conditions are appropriate
for their development. In this unit, we discussed the major biomes of world and
characteristic features of the aquatic and terrestrial biomes.

9.6 KEY WORDS

Biome: A large geographical area with characteristic groups of organisms adapted
to that particular environment.
Taiga (boreal Forest): Terrestrial biome characterized by cold temperature,
moderate precipitation, and continuous forest.
Temperate Forest: Terrestrial biome characterized by moderate temperature,
substantial precipitation, and a moderately diverse assemblage of plant and animal
species.
Temperate Grassland: Biome in interior Eurasia and North America, with cold winters,
warm summers, and much grass.
Temperate Shrubland: Biome at 30–40° latitude on the southwest side of large
landmasses, with mild, wet winters, hot dry summers, occasional fires, and a
diverse community of drought-adapted shrubby plants.
Tropical Rain Forest: Terrestrial biome characterized by constant high
temperature and rainfall in a broadleaf evergreen forest with extraordinary species
diversity.

9.7 SUGGESTED FURTHER READING/

REFERENCES
Campbell, N.A. 1996. Biology, Fourth Edition. The Benjamin/Cummings
Publishing
Company, Inc. Menlo Park, CA.
http://en.wikipedia.org/wiki/Biome
http://www.blueplanetbiomes.org/world_biomes.htm
http://www.ucmp.berkeley.edu/glossary/gloss5/biome/
Jeffree, E. P., and C. E. Jeffree. 1994. Temperature and biogeographical
distributions of species. Functional Ecology, 8:640–650.
Odum, E.P. 1971. Fundamentals of Ecology. 3rdEd. W.B. Saunders Company
USA

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Fundamentals of Ecology Odum, E.P. and Barrett, G.W. 2001. Fundamentals of Ecology. 5thed. Thomson.
Brooks/Cole. 10.
Olson, D. M., et al. 2001. Terrestrial ecoregions of the world: A new map of life
on earth. BioScience 51(11):933–938.
Perry, D.A. 1994. Forest Ecosystems. The Johns Hopkins University
Press.Baltimore.
Singer FD. 2016. Ecology in Action. Cambridge University Press, United
Kingdom.
Walter H. and Box E. 1976, Global classification of natural terrestrial ecosystems,
32 (2), 75–81
Whittaker R. H. 1975, Communities and Ecosystems, 2nd ed., Macmillan, New
York

9.8 ANSWERS TO CHECK YOUR PROGRESS

Check Your Progress 1
1. Refer section 9.4.1
2. Refer section 9.4.2
3. Refer section 9.4.2
4. Refer section 9.4.2.2
5. Refer section 9.4.2.6

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