UNIT 

10: ECOLOGICAL PRINCIPLES 

A. The Environment: Physical environment; biotic environment; biotic and abiotic

interactions.
 
Ecology  is  the  division  of  science  that  examines  the  Indirect factors: These include soil structure, soil
interactions  between  living  organisms  and  their  organisms, altitude, wind, slopes etc.
environments.  Ecology  term  was  Coined  by  Ernst 
Haeckel  from  the  Latin  words  Oikos  (home)  and  According to Oosting (1948), the environment is
Logos  (study  of).  Ecology  is  the  study  of  the  complex of variable factors or causes, which includes (i)
interrelationships  between  organisms  and  their  substances (soil, water), (ii) conditions (temperature,
environment  and interaction of the biotic and abiotic  light), (iii) forces (wind, gravity), (iv) organisms (plants,
components of the environment.   animals), and (v) time.
 
“Ecology  is  the  study  of  the  structure,  function,  Daubenmire (1959) classified the factors into seven
behavior,  and  interaction  of  the  natural  systems  coordinate headings as (i) Soil, (ii) Water, (iii)
that comprise the biosphere.”   Temperature, (iv) Light, (v) Atmosphere, :vi) Fire, and
  (vii) Biotic factor.
A  biological  species  includes  all  organisms  of  a 
group that reproduce and produce fertile offspring in  Most of the ecologists, however, prefer to recognise four
nature.  In  simplest  terms,  organisms  that  do  not  categories of ecological factors, which Life as follows
achieve  this  in  nature  are  different  species  or  kinds, 
because  gene  flow  among  them  is  not  possible  1. Climatic or aerial factors: These include
without  reproduction.  Most  species  are  divided  up  (1) Light.
into  local  populations  that  are  partially  separated  (2) Temperature of air (atmospheric temperature).
from  one  another  by  a  physical  (e.g.,  mountain)  or  (3) Rainfall (precipitation).
(4) Humidity of air.
chemical  (e.g.,  saline)  barrier.  Gene  flow  is  reduced 
(5) Atmosphere (gases and wind).
but  not  cut  off.  The  several  populations  of  different 
species  that  live  together  and  interact  form  a 
2. Topographic or physiographic factors:
community.  The  community  of  organisms  and  their 
These are the factors related with physical geography of
physical/chemical  environment  constitute  an 
earth, and include
ecosystem.  
(1) Altitude.
 
(2) Direction of mountain chains and valleys
According to Taylor/1936: “Ecology is science of all  (3) Steepness and exposure of slopes etc.
relations of all organisms to all their environment”. 
  3. Edaphic factors: these deals with the formation of
According  to  R.  Mishra:  “Ecology  is  interaction  of  soil, its physical and chemical properties and details of
forms, functions and factors”  related aspects.
 
4. Biotic factors: These are all kinds of interactions
ENVIRONMENT between different forms of life, i.e., plants, animals,
microorganisms etc.
Environment has different meanings for different people.
Most definitions include the physical, chemical and
The Real Environment-Interaction of Ecological
biological components that influence the life of an
Factors
organism. The term environment etymologically means
surroundings. Thus, environment is a complex of so
An important aspect of the effects of environment on the
many things (light, temperature, soil, water, etc.) which
life of an organism is the interaction of ecological
surround an organism. Any external force, substance or
factors. Actually under natural conditions, at the same
condition, which surrounds and affects the life of an
time, the life or organism is affected by the sum of all
organism in any way, becomes a factor of its
ecological factors-the environment and not by any
environment. These factors have been variously called as
individual factor. All the factors are interrelated.
environmental factors, ecological factors or simply
Variations in one may affect the other. For instance, an
factors and may be living (biotic) as well as non-living
increase in temperature would certainly bring out a
(abiotic). The sum of all these living and non-living
decrease in humidity values. Increased light intensity, in
factors makes the environment of an organism. The
addition to affecting the rate of photosynthesis, would
place, where an organism lives - habitat, indeed presents
affect the atmospheric humidity. Thus, at a given time
a particular set of environmental conditions - the
effect on the organism's life is an overall effect of these
environmental complex.
interrelated factors, and thus it becomes very difficult
indeed sometimes to say as which of the individual
Various ecological factors of this environmental
factors is actually responsible for the marked effect on
complex have been grouped and discussed in different
organism's life. Thus ecological factors in affecting an
ways. Sometimes all the factors are classified into two
organism's life under natural conditions, operate in
groups
conjunction and not in isolation, as we have an
interaction between various factors.
Direct factors: These are light, temperature, humidity of
air, soil air, soil water, soil nutrients etc.

INSTITUTE FOR ADVANCED STUDIES, JODHPUR  1
UNIT 10: ECOLOGICAL PRINCIPLES 
Thus, in considering the ecological factors separately for
distinguishing and measuring the effect of each, one
must remain aware that in nature these factors never act
alone. Organism is subjected to many influences at the
same time, and the effect of one factor usually modified
by the other factor(s). The 'real life' of the organism thus
involves the simultaneous and continuing impact of all
existing interacting ecological factors. Though efforts
are made to focus on various components of
environment separately (in isolation of each other), the
environment in fact is not even the sum of all its
components put together but it is the product of
interactions between all the components.
Since various ecological factors affect the life of an
organism in a holistic manner (i.e., all the interacting
factors operate in conjunction and not in isolation), it
becomes difficult to understand this mechanism of the
nature of influence of individual factor. To understand
this mechanism of environmental influence, it thus
becomes essential to study the effect of each factor
separately, and this is said to be an analytical approach.
We shall now discuss in detail each of the above said
ecological factors separately to understand how it might
influence the life and distribution of organisms. We
begin with climatic factors.
I. Climatic Factors:
Climate of any region is chiefly determinated by such
meteorological influences as relative humidity of air,
temperature, wind pressures and evaporation rates.
Climatic factors characterise, in general, a particular
region. These factors, concerned mainly with the aerial
environment of organisms, include light, temperature of
air, humidity of air, precipitation (rainfall) and gaseous
components of atmosphere including wind. We will
discuss each of them in detail.
Light Factor
Light is well known for its effects on such basic
physiological processes of plants as photosynthesis,
transpiration, seed germination, flowering etc., and thus
is of much concern in ecology, where it plays important
role in the species composition and development of
vegetation. Light intensity, reaching the earth's surface,
shows much spatial variations being influenced by some
such factors as (i) atmosphere, chiefly atmospheric
gases, nitrogen and oxygen, that absorb and disperse
small fractions of the shorter wavelengths, (ii) suspended
particles, solid particles dispersed in air, such as dust and
smoke, or in water, such as clay, silt, plankton, bog
colloids etc., have a screening effect, (iii) water layers,
(iv) layers of vegetation, chiefly in terrestrial habitats, as
forests, and (v) such topographic factors as direction and
slope of the land surfaces which cause marked variations
in intensity and daily duration of insolation. Radiant
energy of sun, reaching the earth is transformed to other
forms of energy, such as thermal, mechanical, electrical
etc., which bring about changes in several physical
processes in atmosphere as well as on earth. Light
intensities are closely related with atmospheric
temperature and relative humidity. Thus light affects
organisms directly as well as indirectly.
2 INSTITUTE FOR ADVANCED STUDIES, JODHPUR
The great deal of solar radiation is absorbed in the
atmosphere. The amount of light thus is greater at higher
altitudes than at sea level. This is due to thinner layers of
air at higher altitudes. On a cloudy day, light may be
reduced to 4% of the normal intensity. Although red and
blue parts of the spectrum are useful in photosynthesis,
in ecology relatively less importance is given to the
quality of light for its role in plant distribution.
The layers of vegetation bring about variations in light
intensities reaching at various heights of mountain. This
becomes evident, when we study light conditions in
forests, where a major proportion of light intensity is
absorbed by tree vegetation and the light reaching to the
lower part of the ground vegetation is considerably
reduced by 90-98% of that in the exposed areas. The
amount of light reaching the forest floor depends upon
the height of canopy, crown development of trees, age of
trees, and phenological characteristics of the constituent
species. Thus in a forest, the mature tallest tree receives
full insolation, under-shrubs receive subdued
illumination, and herbs and especially epigenous
cryptogarns grow in weaker light conditions, When
leaves of a tree are fully expanded, its canopy may
reduce light to less than 1 % of full solar radiation.
Salisbury (1916), as a result of his studies on light
conditions in forests, showed that light intensities govern
the development of ground flora of a forest. But it is
seen that light conditions of forests, with and without
ground flora, are not very much different, and thus it
seems that not only light, but other factors as
temperature, moisture, plant growth, mineral nutrition,
humus content, and root competition are also involved. It
is also seen that during shade phase i.e., when full leaves
are present on the trees, light conditions are poor, but in
nature sun-flecks and movement of leaf canopy of trees
play an important role and these compensate for the
reduced light, making it available to the ground flora.
Costing and Shirely (1945) too, on the basis of a series
of experiments, concluded that for each light intensity
growth of forest plants may be increased by reducing the
root competition. Generally, the cause of death of the
tree seedlings in forest stands is the increased
competition in roots for the nutrients rather than reduced
light intensities.
Layers of water have pronounced effects on light
intensities. In aquatic bodies, plants are found growing
in distinct zones, although such a zonation is not always
due to different light intensities at various depths.
Submerged plants receive weaker illumination than the
exposed plants, as some of the light is reflected at the
water surface and of the remainder, much is absorbed by
the upper layers of water. Light penetration in water
depends upon the turbidity, solute content, motion, and
planktonic growth of the water. The intensity of light
decreases successively with increasing depth. Itowever,
in their studies on distribution of plants in English lakes,
Pearsall (1920) and Misra (1938) could show that the
zonation in such water bodies is not solely due to
differences in light conditions at various depths, but the
physical and chemical characteristics of the rooting
medium i.e., edaphic factors also play an important role
in plants' distribution. Importance of mud characteristics
in plants' distribution has also been recognised for plants
of low-lying lands by Misra (1946).
 
UNIT 10: ECOLOGICAL PRINCIPLES 
Generally at dawn, sunset, and in water, light intensities
are weaker. At equator daylight prevails for about 12
hours out of every 24, in both summer and winter.
Towards the poles, in summer day length becomes
longer than 12 hours.
Light in relation to plants
Directly or indirectly light affects the plants' life in the
following ways:
Chlorophyll production: Expect few, like seedlings of
conifers young fronds of ferns, some mosses and algae
which may become green even without light, most of the
plants need light for chlorophyll formation. Thus
existence of other life forms, which depend upon plants
for food is due to light.
Heating action: Exposure of parts of the plants to light
raises their temperature, which effect their related
processes.
Effect on transpiration rate: Indirectly light affects
transpiration rates through increase in temperature.
Transpiration rates correspondingly affect water
absorption also. Thus high light intensities are always
associated with dry habitats and high transpiration rates.
Stomatal movement: Opening and closing of stomata is
regulated by light. This is thus related to transpiration
and absorption.
Distribution of plants: Light conditions at poles are
different from other parts of earth. Thus total amount of
radiation received by the earth's surface differs with
latitude (distance from the equator). This may be one of
the reasons for differences in the vegetation at poles and
at other parts of the earth.
Overall vegetative development of plant parts:
Sometimes plants are classified ecologically on the basis
of their relative light requirements and the effects of
light on their overall vegetative development into:
Heliophytes: Growing best in full sunlight, and
Sciophytes: Which grow best at lower light intensities.
There are some heliophytes, which grow best in sun, but
can grow fairly well under shade. Such plants are called
facultative sciophytes. Similarly there are facultative
heliophytes which, although grow best at lower light
intensities, but can grow well in full sunlight. Some
plants, likely to be injured by bright light, develop some,
characteristics in themselves to minimize such effects.
For example, plants of sunny habitats have vertically-
oriented leaf blades. Some variations in the pigments
complex of leaves also take place. In moist climates,
vegetation is generally a complex series of superimposed
layers of tall trees, low trees, shrubs, herbs and mosses
etc., most of which such as maples, beech, spruces,. firs,
and hemlocks are obligate or facultative sciophytes. In
dry climates, species of the flora are mostly heliophytes,
which are mostly pines, soft maple, willows, tulip tree,
birches, larches and junipers.
In comparison to plants grown in the shade, those
developing in full sunlight, usually exhibit such
morphological and physiological features as : thicker
INSTITUTE FOR ADVANCED STUDIES, JODHPUR  3
stems with well developed conducting elements and
mechanical tissues, shorter internodes, more frequent
branching, smaller chloroplasts, well-developed palisade
with poorly developed spongy parenchyma, roots longer,
more in number and much branched with higher root
shoot ratio, less chlorophyll content, low photosynthetic
rate per unit surface, high respiration rate, low water
content (dry weight basis), higher concentration of salts
and sugar and high osmotic pressure, decrease in acidity
of cell sap, low potassium contents, early appearance of
flowers, and much more resistance to temperature injury,
drought and parasitic organisms.
Photoperiodism: Total length of the daily light period to
which plants are exposed - photoperiod has pronounced
effect on the vegetative growth as well as flowering of
the plants. On the basis of their photoperiod, the plants
may be
(i) short-day plants: that develop normally only when
the photoperiod is less than a critical (between 12-14
hours) maximum. Examples are Salvia splendens,
Datura stramonimum, Cannabis sativa, Andropogon
virginicus and Cosmos bipinnatus, and
(ii) long-day plants: those demanding a photoperiod in
excess of certain critical minimum. For example,
Brassica rapa, Nigella avroensis, Secale cereale,
Sonchus oleraceum Sorghum vulgare. Most of the plants
are short-day, which require a relatively long period of
uninterrupted darkness for flowering.
(iii) Some plants are indifferent to the above conditions,
and are called day neutral plants. For example, Cucumis
sativus, Gossypium hirsutum, Nicotiana tabacum,
Phaseolu lunatus, Poa annua and Solanum tuberosum.
Thus, light as photoperiod plays an important role in
local distribution of plants, by affecting stem elongation,
flowering, fruit development and other processes.
Light in relation to animals
So much has been said of the role of light in the life of
plants; what about animals? Light has far reaching
effects on animals also, by affecting their several types
of activities like pigmentation, reproduction,
development, growth, locomotion, migration etc., which
may be summarised as follows:
Metabolism: Light affects metabolic processes of
animals through its heating effect on tissues and by
ionization of the protoplasm. This all results into an
increase in the enzymatic activities, general metabolic
rate and degree of solubility of salts and minerals.
Animals receiving poor light in caves thus shows slow
rate of metabolism. Light also affects photo-oxidation
and respiration rates.
Reproduction: In animals like birds, light initiates the
breeding activities. In some birds gonads become active
with increased light intensity during summer. Thus
animals are long-day, short-day, and indifferent to day-
length. Some sheep, deer and goats are short-day
animals, which may be brought to sexual act by
decreasing the length of exposure to daylight. Spring
breeding birds and animals as freest, starlings, turkeys
and others are long-day animals. Ground squirrels,
UNIT 10: ECOLOGICAL PRINCIPLES 
guinea pigs etc., are indifferent towards the length of
exposure to daylight.
Development: In Salmon, larvae undergo normal
development only under sufficient light conditions, in
the absence of which there may take place rapid death of
larvae. In Mytilus, larvae in their earlier stages grow
larger in darkness than in light.
Eyes: The degree of eye development sometimes
depends upon the available light intensities. In animals,
living in caves as Proteus anguinus, and in the deep sea
fishes, the eyes are absent or rudimentary. In surface
dwelling animals as crustaceans and fishes, the ratio of
the eyes to head is called as normal. In the ocean, with
increasing depth, size of eyes in animals generally goes
on increasing.
Vision: Higher animals including man are able to see
various objects only in the presence of one or other form
of light. Some fishes, as Lepomis, also depend on
eyesight for location of their food.
Pigmentation: Most of the chemical changes that result
into the formation of pigments are initiated by light
energy. The process of pigmentation in animals is thus
influenced by light in a number of ways as
(i) in the development of skin colour: For instances, in
some animals living in caves, as amphibians and some
fishes, the pigments in skin are lacking due to absence of
light,
(ii) in protective colouration: Some animals develop
colouration in their skin which more or less resembles
with the background in which they live. Thus leaf insect
Phyllium, Australian fish, Phyllopteryx eques and
butterfly, Kallima paralecta etc., have colouration
exactly like that of their background, and this affords a
good means of protection from their enemies. Similarly
some birds, mammals and fish display differential
shading, with darker pigmentation on back and lighter
one below, that helps them in providing protection,
(iii) colour changes: Some insects, crustaceans, fishes,
amphibians, reptiles and cephalopods are able to change
their colour or patterns rapidly according to the
environmental conditions where they live. This helps in
thermoregulation and in affording protection from
enemies.
Locomotion: The speed to locomotion in some lower
animals is regulated by light This phenomenon is known
as photokinesis. There is evidence, that blind larvae of
mussel crab move faster if exposed to increased light
intensities. More or less similar responses in terms of
locomotion towards different light intensities are known
to occur in some aquatic invertebrates. In some animals,
light plays a role in the orientation of locomotion, and
this phenomenon of movement of animals in response to
light is called phototaxis, and such movements are
known as phototactic. Animals like Euglena, Rantara
etc. are positively phototactic, moving towards the light
source, whereas earthworms, slugs and some
zooplanktons like copepods etc. are negatively
phototactic. In some animals, like many tubicolous
worms and polyps of many coelenteratces, only a part of
their body shows movement in response to light and this
has been termed as phototropism.
4 INSTITUTE FOR ADVANCED STUDIES, JODHPUR
Photoperiodism: Animals are known to respond to the
length of the day, through the effect of total light period
on such processes as gonad activities, reproduction,
metamorphosis, migration etc. In certain catfishes longer
days induce early' maturity. Migration of eels, salmons
and birds is known to be affected by photoperiodism, as
some birds are seen migrating towards north during
summer when days are longer, and towards south during
winter when days are shorter. In some animals
development of fur and feathers during winter is shown
to develop in response to photoperiodism.
Daily responses of animal to light conditions are known
as circadian rhythms, whereas the annual rhythms
known as circannual rhythms. Circadian rhythms are
internally driven or endogenous, are affected little by
temperature changes, are sensitive to a great variety of
chemical inhibitors, and are innate, not learned from or
imprinted upon the organism by the environment. The
innate character of circadian rhythm is demonstrated by
several animals. When Drosophila are kept under
constant conditions from the larval stage on, they will
still emerge from pupae with a regular circadian rhythm.
Eggs of chicken and lizards kept under constant
condition produce animals that later show regular
circadian cycles. The circadian rhythms have been
observed in zooplanktons polychaete annelids, many
insects, most birds and some mammals. Planktons of sea
and lakes show diurnal changes in their vertical
distribution.
Solary day, lunar day, tidal rhythms, monthly and annual
rhythms are also of common occurrence among animals.
Endogenous annual cycles or circannual rhythms have
been known in ground squirrels, warblers and other
birds, some crayfishes and slugs.
Temperature Factor
The term 'temperature' here, besides the aerial
temperature of terrestrial organisms, also includes the
temperature of water for aquatic life. In fact the
atmosphere to which aquatic forms of life are exposed, is
the water medium. Temperature is well known for its
effects upon plants and animals. Temperature affects
plants and animals in various ways, which are as
follows:
Effects on metabolism: All metabolic processes are
influenced by temperature Since temperature regulates
the activity of enzymes, all chemical reactions in the
body of organism are controlled by temperature. It
affects the rates of transpiration, photosynthesis in plants
and respiration rates and other metabolic processes in
plants as well as animals. In plants it affects seed
germination also.
Effects on reproduction: Flowering in plants is affected
by temperature through thermoperiodism i.e., the
response of plant to rhythmic diurnal fluctuations in
temperature. Temperature is an important factor, in the
phenology of plants (i.e., study of periodical phenomena
of plants, as the time of flowering in relation to climate).
In animals, the maturation of gonads or sex cells and the
liberation of gametes take place at a particular
temperature, which varies from species to species.
Breeding in some animals remains unaffected by
 
UNIT 10: ECOLOGICAL PRINCIPLES 
temperature throughout the year, whereas in some,
breeding occurs only in summer or in winter. Some
species may have two breeding seasons, one in spring,
other in fall In blow-fly Calliphora sericate, the number
of eggs laid per female, increases with increase in
temperature up to 32.5°C, but the number decreases with
further increase in temperature.
Thus temperature also affects fecundity of animals.
Fecundity is the reproductive capacity of animals. The
female of the insect, acridid Chrotogonus traychpterus
becomes sexually mature at 30°C and 35°C than at 25°C
and maximum eggs per female laid at 30°C. In
grasshoppers, Melanoplus sanguinipes and Camnula
pellucida produced 20-30 times as many eggs at 32°C
than those at 22°C.
Effect on growth and development: Both, extremely
low and high temperature have adverse effect on the
growth of plants. Low temperatures bring about such
cold injuries as desiccation, chilling injury and freezing
injury. In desiccation, tissues are dehydrated and injured
due to rapid transpiration and slow absorption during
winter. Chilling injury is the killing or injury of plants of
hot climate, when exposed to low temperatures for
sometime. In some plants of temperate climates, if
exposed to low temperatures, water is frozen into ice
crystals in the intercellular spaces causing injury to cells.
This is known as freezing injury. Some perennials can
tolerate extremely low temperatures and this ability is
called cold resistance. Extremely high temperatures
cause stunting and final death of plants, which is due to
adverse effects on a number of physiological processes
as respiration, transpiration, protein metabolism etc. This
is called heat injury.
Effect on sex-ratio: In some animals, as rotifers and
daphnids, sex-ratio is affected by temperature. Under
normal condition daphnids give parthenogenetic eggs
that develop into females, whereas with increase in
temperature they give sexual eggs which after
fertilisation develop either into males or females. In
Macrocyclops albida there is significant increase is
number of males with increase in temperature. Similarly
in plague fea. Xenopsylla cheopes, males out numbered
females on rats on days when the mean temperature
remains in between 21-25°C.
Effect on coloration:Some insects, birds and mammals
in warm humid climates bear darker pigment than the
races of same species present in cool and dry climates.
This phenomenon is known as Gloger rule. In the frog
Hyla and the homed toad Phrynosoma low temperatures
induce darkening.
Effect on morphology: Temperature also affect the
absolute size of an animal and the relative proportions of
various body parts - Bergman's rule. Birds and
mammals attain greater body size in cold regions than in
warm areas. But poikilotherms are smaller in cold
region. The tail, snout, ears and legs of mammals are
relatively shorter in colder part than in the warmer areas
Allen's rule: For example, there occurs difference in the
size of ears of arctic fox (Alopex lagopus), red fox
(Vulpes vulpes) and the desert fox (Megatotis zerda).
Gazella picticanda of Himalayas has shorter legs, ears
and tail than G .benetli found in the plains.
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The races of birds with relatively narrow and more
acuminate wings tends to occur in colder regions, while
those in warmer areas tend to be broader (Rensch's
rule).
Temperature also affects the morphology of some fishes
and is found to have some relationships with the number
of vertebrae Jordon's rule. .
Thus, through its effect on organisms in various ways
described above, temperature affects geographical
distribution of plants and animals. Together with
moisture, temperature determines the climatic condition
of a particular region.
Variations in temperature and its effects on
distribution of plants
Temperature shows pronounced temporal as well as
spatial variations. Especially in a country like India,
there are experienced marked seasonal extremes and
fluctuations in the temperature values. Soil temperature
is the result of heat gained by the absorption of solar heat
energy, and soil temperatures are greatly influenced by
the latitude (distance from the equator) of the particular
place. Obviously, the temperature values are maximum
at the equator, decreasing gradually towards the poles.
The division of earth's vegetation into different zones, as
equatorial, tropical, coniferous, alpine vegetations etc.,
rests on the marked variations in temperature at different
latitudes. Similarly, altitude (height above the sea level)
affects temperature values. Besides latitude and altitude,
colour and composition of surfaces, plant cover, water
content of soil, and such physiographic factors as the
steepness of slope, exposure of slope, and direction of
mountain chains etc., affect greatly the temperature
conditions. For instance, on mountains, we generally
experience a decrease in temperature with increasing
altitude, and thus vegetation at different altitudes is
different, showing distinct zonation. Generally, mean
temperature of air decreases about 10°F for every degree
of latitude north of equator, and for every 300O of
altitude. In nature, valleys and lowlands are sometimes
much cooler due to sinking in of the heavier cold than
the mountain tops.
Changes in both, the latitude and the altitude, show more
or less similar effect upon the type of major vegetations
of the world. The various zones of different kinds of
vegetation at increasing latitudes from equator towards
poles and at increasing altitudes on mountains (Fig. 1)
are more or less similar, showing that increase in latitude
and altitude brings about more or less similar influence
upon vegetation. Thus, we may find distinct latitudinal
as well as altitudinal zonations in the vegetation.
UNIT 10: ECOLOGICAL PRINCIPLES 
On mountains, temperature and rainfall are commonly
said to determine the type of vegetation at different
altitudes. But Purl, G.S. has shown that at different
altitudes, not only temperature and rainfall but edaphic
conditions (soil characteristics) are also important. On
mountains with deviations in elevation, in general,
southern and western slopes register higher temperatures
than do the northern or eastern slopes. The angle of the
slope in relation to sun governs the total amount of
radiation. These differences are probably due to different
solar radiation received, differences in the amount of
rainfall, snow fall, relative humidity and wind movement
on the two slopes. At higher elevations, generally
temperature decreases and rainfall increases, and these
certainly affect the development of soil and vegetation.
Due to low temperature and high rainfall the organic
matter content of the soil increases at higher altitudes,
with an increase in soil nitrogen and a decrease in its pH
values. Generally, xerophytes are more common at lower
altitudes and chamaephytes occur at higher altitudes.
In Himalayas, temperature variations are quite evident,
and there is a general zonation of vegetation (Fig.1) from
lower to higher altitudes. The successive zones of
vegetation from base upwards are tropical and
subtropical, temperate, and alpine. However, there are
practically no sharp boundaries between these
vegetational zones due to differences in topography, soil
and geology. Thus on Himalayas, western as well as
eastern, the effect of temperature together with altitude
and other factors becomes quite evident after studying
the vegetational zones at different altitudes.
Some persons, on the basis of temperature conditions
divide world's vegetation into various classes as (i)
megatherms - where high temperatures prevail
throughout the year and dominant vegetation is tropical
rain forest, (ii) mesotherms - with high temperature
alternating with low temperature, and dominant
vegetation is tropical deciduous forest, (iii) microtherms
- where low temperatures prevail and vegetation is of
mixed coniferous forests type, and (iv) hekistotherms-
with very low temperatures and alpine vegetation being
dominant.
Effect of temperature on distribution of animals
As discussed above, there are much variations in
temperature and these affect plants distribution in
various ways. It also affects animals. All animals are
broadly classified into two group:
Homoiothermic or endothermic animals (warm-
blooded)
These animals like birds and mammals are able to
maintain their body temperature at a constant level
irrespective of the environmental temperature.
Poikilothermic or ectothermic animals (cold-blooded)
These are animals as reptiles, fishes, amphibians in
which the body temperature fluctuates with changes in
the environmental temperature.
Thermoregulation in homoiotherms: Though several
factors may operate, the most important role is played by
their skin. Almost each of the various structures present
in the skin play some part in the temperature regulation.
6 INSTITUTE FOR ADVANCED STUDIES, JODHPUR
A homoiotherm has different response to cold or heat as
follows.
Response to cold
When a typical homoiotherm as a mammal is subjected
to severe cold, following adaptations help the animal
from excessive cooling.
ƒ The subcutaneous fat serves as insulator and
reduces heat loss from body.
ƒ The hair is raised and brought up into a moreor-less
vertical position by the contraction of the erector-pili
muscles. Thus air gets trapped in the spaces between
hairs. This air is warmed by the body and being a poor
conductor of heat it serves as an insulatory layer around
the animals. In birds, the same function is done by
feathers.
ƒ The superficial blood vessels in the skin constrict so
that blood is directed from the surface to the deeper
layers. This reduces loss of heat from the blood to
surrounding atmosphere.
ƒ Extra heat is produced by increase in the metabolic
rate particularly of the liver and muscles.
Homoiotherm besides above said structural adaptations
also have physiological capability of increasing their
heat production.
Response to heat
Heat production is cut down and heat loss encouraged as
follows:
ƒ Animals of hot climates have relatively little
subcutaneous fat. The fat deposits are localised. Thus
camel's fat is stored only in hump and in buffalo and
bison on top of the neck.
ƒ The hair is lowered by relaxation of the erectorpili
muscles so that it lies flat against the body surface. There
is no space between the hair and no air can be trapped
against the skin,
ƒ The superficial blood vessels are dilated so that
blood is brought up near the surface from which it can
lose heat to atmosphere.
ƒ Sweating or panting occurs. Sweating by sweat
glands of skin and evaporation of sweat from the body
surface cools the skin and the blood flowing through it
ƒ Metabolic rate falls in hot conditions so that less
heat is generated by the body.
Thus in mammals the hypothalmic centre functions as a
thermostat It is sensitive to temperature changes of blood
flowing through it and responds by sending nerve
impulses through efferent nerves to the appropriate
effectors. If the temperature of the blood is slightly
higher than it should be, the thermo-regulatory centre of
hypothalamus of brain detects this and sets into motion
process that collectively encourage heat loss. On the
other hand, if the temperature falls below the normal the
ce'htre initials processes that produce and conserve heat.
Thermoregulation in poikilotherms
Some poikilotherms avoid both heat and cold by
underlying dormant in period of environmental stress.
Many insects, some crustaceans, mites and snails, enter
diapause, a state of dormancy and arrested growth. Eggs,
embryonic larvae and pupae are involved. Some
 
UNIT 10: ECOLOGICAL PRINCIPLES 
poikilotherms become dormant in temperature extreme
conditions. Amphibians and turtles bury them in the
pond mud, snakes and lizards seek burrows.
A few exceptional poikilotherms, especially insects,
certain amphibians and reptiles, exercise some degree of
thermoregulation by physiological or behavioural
mechanism. For example, hawk-moths can raise the
temperature of their flight muscle to 32°-36°C by
vibrating the wings before take off and gregarious butter
fly larvae may raise their temperature 1.5-2°C by
clustering together. Locust and grasshoppers may
increase temperature l0oC by basking sideways in the
sun. Ants move their larvae to warm or cool places.
When temperature drops, lizards bask in the sun to
achieve more temperature. Once this temperature is
attained, they will divide their time between sun and
shade to maintain it The frogs and reptiles lower their
body temperature slightly by evaporating cooling
through the skin or via the respiratory tract by panting.
Temperature affects the distribution of animals also, as
different species have different ranges of their
temperature requirements. For instance, coral reefs
require a minimum temperature of 21°C for their
suitable growth, hence are completely absent in colder
regions, where prevailing temperatures are below 21°C.
Speckled trout, Salvelinus fontinalis, prefers 14°C to
19°C, and thus cannot tolerate a temperature above
25°C, being thus completely absent from streams where
the temperature exceeds 24°C, for some time.
Distribution of intertidal bamacles is also affected by
temperature. For example, an intertidal bamacle, Pectin
groenlandicus, is hardly known to occur in areas with
O°C temperature, whereas another one, Balanus
balanoides, occurs at a temperature below 7.2°C.
Precipitation (Rainfall)
Precipitation is the chief source of soil water. The water
available to plants and animals from soil comes as a
result of rainfall. There occurs an interchange of water
between the earth's surface and the atmosphere, forming
the water or hydrologic cycle. The two important events
which are involved in this cycle are precipitation and
evapotranspiration.
Infect, water during rainfall is directly of little or no use
to the organisms. It is used only after reaching to the
soil. Precipitation is the result of gravitational pull on the
vapour in atmosphere. Precipitation occurs in various
forms. These are, drizzle, rain, snow, dew and frost, sleet
and hail. Drizzle involves minute drops appearing as to
float in air. Rain is the drops of liquid water, which are
larger than drizzle and also heavier. Snow is the
moisture as solid state, and dew and frost are formed due
to condensation of moisture directly on the surfaces of
objects, plants, animals, soil, etc. Sleet is in the form of
small grains or pellets of ice, whereas hail consists of
balls or lumps of ice. Snow is injurious to plants,
breaking tender branches, flowers and fruits. Hail and
sleet also cause similar damage. Some sedges grow in
snow patches.
Of all the above forms of precipitation, the rain is the
most important. It is the source of soil water and also
INSTITUTE FOR ADVANCED STUDIES, JODHPUR  7
affects humidity of atmosphere. Rains in India are
caused by monsoons. About 45% of the water available
during annual precipitation flows into rivers, 20%
percolates into the ground and the remaining 35% is lost
by evaporation.
The amount of annual rainfall greatly influence the
vegetation as well as animal population of a particular
region. Rainfall distribution in different seasons of the
year also may further be important in regional
distribution of vegetation. On high mountains, the
regions are correlated with the distribution of rainfall.
And, there are distinguished three regions: (i) lower
region with scanty rainfall, (ii) middle region or the
cloudy region with lot of rain, and (iii) upper dry region,
above the clouds covered with snow and ice.
Annual rainfall determines the type of vegetation in any
region. For example, in tropical areas with heavy rainfall
throughout the year, main vegetation are evergreen
forests. In countries with heavy rainfall during winter
and low during summer, there are present scierophyllous
forests. The plants are shrubs stunted in height, with
leathery, thick, evergreen leaves. The areas with heavy
rainfall during summer and low during winter are
characterized by the presence of grasslands. The regions
where rainfall is scanty, are seen with deserts and
xerophytic vegetation.
In India, due to differences in the amount of annual
rainfall and its distribution in different seasons of the
year, we find that vegetation types in different parts of
the country are much different from each other.
Similarly, with changes in vegetation we also find
different types of animals in different geographic
regions.
Humidity of Air
It is one of the different forms of water in nature.
Atmospheric moisture in the form of invisible vapour is
known as humidity. The humidity of air is expressed in
terms of relative humidity values. It is "the amount of
moisture in air as percentage of the amount which the air
can hold at saturation at the existing temperature".
'Humidity is greatly influenced by intensity of solar
radiation, temperature, altitude, wind, exposure, cover
and water status of soil. High temperature increases the
capacity of the air to retain moisture and cause lower
relative humidity. Low temperatures cause higher
relative humidity by decreasing the capacity of air for
moisture. Relative humidity is measured by the
instrument called psychrometer or by paper strip
hygrometer or a thenno-hydrograph.
Daily variations in relative humidity values depend upon
the type of habitat conditions. In plains and deserts, it
may show variations during day, whereas in oceanic
islands there is little variation, being same throughout
the year.
Of the other factors of aerial environment, humidity
plays an important role in life of plants and animals.
Processes as transpiration, absorption of water etc., are
much influenced by atmospheric humidity. The
saturation deficit, temperature and wind velocity modify
evapo-transpiration rates. Saturation deficit (as mm Hg)
 UNIT 10: ECOLOGICAL PRINCIPLES 

is the difference between the pressure of water vapour in

the atmosphere at a given time and the maximum vapour
pressure that it could contain at the same temperature. A
decrease or increase in the saturation deficit causes a fall
or a rise in evaporation rate respectively. Similarly
temperature variations also influence evaporation. Dry
winds decrease the amount of air moisture by removing
moist air about plants and mixing it with dry air which
decreases the humidity and thus increases transpiration.

Thus humidity affects the life of plants in various ways.

Effects of moist air on plants are more or less similar to
those of reduced light intensity. Some plants as orchids,
lichens, mosses etc., make direct use of atmospheric
moisture. In fungi and other microbes, it plays an
important role in germination of spores and subsequent
stages in life cycle.

Atmosphere-Gases and Wind

The term 'atmosphere' here includes only the thick

gaseous mantle surrounding the earth. As we will see
later, the soil too has its own atmosphere. There are pore
spaces in between soil particles as well as between
aggregates, and such spaces make the atmosphere of
soil. The present discussion on atmosphere as an Troposphere: The basal part of atmosphere which is
ecological factor would include the effects of principal about upto 20 km above the earth's surface (on poles
gaseous constituents present in atmosphere, wind and only Up to 8 km) is known as troposphere. For
fire on the organisms. organisms this is the most important zone of the
atmosphere. The important events, such as cloud
Atmospheric gases formation, lightning, thundering, thunder storm
formation etc., all take place in troposphere. Air
Up to the height of about 300 km above the earth's temperature in this zone gradually decreases with height
surface, there is present some sort of a thick gaseous at the rate of about 6.5°C per km. Towards the upper
mantle. With increasing height above sea level, the layers of troposphere, the temperature might decrease
atmospheric pressure gradually decreases. In upto -60°C, and this limit of troposphere, which
atmosphere, about 95% of the total air is present up to gradually merges into the next zone, is known as
the height of about 20 km above earth's surface and the tropopause.
remaining 5% in the rest about 280 lan. height. In the
gaseous mantle, there is found a mixture of different Stratosphere: Next to troposphere, the second zone of
gases in different proportions. Of these various gases, about 30 km height, is called stratosphere, where the
nitrogen, oxygen and carbon dioxide are the major temperature values show an increase up to 90°C. This
components. Relative proportions of different gases, by increase in temperature is due to ozone fonnation under
volume, are shown in Table 1. the influence of ultra-violet component of sunlight.
Upper layers of stratosphere form stratopause.
Per cent Per cent
Gases Gases Mesosphere: About 40 km in height, the zone, next to
(By volume) (By volume)
stratosphere, is called mesosphere. In this zone,
Nitrogen 78.0841 Crypton 0.00011 temperature shows again a decrease up to -80°C. Upper
Oxygen 20.9486 Xenon 0.00009 layers of this zone are mesopause.
Argon 0.9340 Hydrogen 0.00006
Carbon dioxide 0.0318 Methane 0.0002 Ionosphere: The rest above the mesosphere, upto the
Neon 0.00182 Nitrous oxide 0.00005 height of above 300 km above earth's surface is
Helium 0.00052 Ozone 0.000004 ionosphere. Most of the gaseous components which
become ionised under the influence of radiant energy,
Besides gases, some other constituents are water remain as ions. Due to their much less density, they are
vapours, industrial gases, dust and smoke particles in rarely present in molecular form. Thus radiant energy
suspended state, micro-organisms, pollen grains, fungal brings about considerable changes in the atmosphere.
spores etc. Cycling of gases as carbon dioxide, nitrogen Water vapour in the atmosphere affects the temperature
etc, between the atmosphere, soil and organism is the and light intensities of atmosphere.
regular feature of nature.
Wind Factor
The detailed structure of atmosphere as given by
Sutcliffe (1966) is shown in Figure 2. Atmosphere can Air in motion is called wind. It is an important
be roughly divided into four zones: ecological factor of the atmosphere, as it affects plants'
life mainly on flat plains, along seacoasts, and at high
altitudes in mountains. Wind is directly involved in

8 INSTITUTE FOR ADVANCED STUDIES, JODHPUR 
UNIT 10: ECOLOGICAL PRINCIPLES 
transpiration, in causing several types of mechanical
damage, and in dissemination of pollen, seeds and fruits.
It also modifies the water relations and light conditions
of a particular area. The velocity of wind is affected by
such factors as geographic situation, topography and
vegetation masses, and position with respect to
seashores. Effects of winds are much pronounced in
plants along the sea coasts and at high altitudes on
mountains. Air moves from a region of high to that of
low pressure. The pressure differences are mainly due to
differential heating of atmosphere. The equatorial
regions receive more heat than north or south regions,
thus low pressure occurs at lower latitude. The air
generally moves from the poles towards equator.
Wind brings about a number of physical, anatomical and
physiological effects on plants. These Physical effects
are as follows (i) Breakage and uprooting (ii)
Deformation: Strong winds from a constant direction
sometimes cause permanent alteration in the form and
position of the shoots. (iii) Lodging (iv)Abrasion (v)
Compression wood and (vi) Desiccation.
Fire Factor
Except those fires caused by lightning or volcanic
activity, which are rare and confined only to certain
specific areas, most of the fires very common in most of
the regions are of biological origin. These are mostly
man-caused, and sometimes, chiefly in forest, develop
due to mutual friction between tree (bamboos etc.)
surfaces.
Fires are generally classified as (i) ground fires - which
develop in such conditions where organic matter
accumulates richly as heaps and they catch fire which
generally smoulder for longer periods. These fires are
flameless and subterranean and kill almost all plants
rooted in the burning material except some woody
species, (ii) surface fires which sweep over the ground
surface rapidly and their flames consume the litter, living
herbaceous vegetation and shrubs, and also scorching the
tree bases if come in contact; (iii) crown fires - which
are most destructive, burning the forest canopy, surface
debris and bringing about vegetational destruction at
larger scale. These develop in dense woody vegetations.
Temperatures that prevail when fire is in progress may
reach up to 1300°F.
Effects of fire
In addition to its direct lethal (killing) effects on plants,
fire also affects them indirectly in the following ways :
ƒ Some trees, as a result of injury by fire, have large
scars on their stems. Such scars may serve as suitable
avenues of entry of parasitic fungi and insects.
ƒ Fire brings about marked alteration of such
environmental factors as light, rainfall, nutrient cycles,
fertility of soil, litter and humus contents of soil, pH, and
soil fauna. Sometimes the habitat needs thousands of
years in the return of normal conditions. The nature of
such alterations depends mainly on the nature of fires.
ƒ Fire plays an important role in the removal of
competition for surviving species. Such species that
fortunately survive fires (fire-tolerants) generally
increase in abundance at the expense of those killed by
fire (fire-sensitive) as a result of considerable reduction
INSTITUTE FOR ADVANCED STUDIES, JODHPUR  9
in competition and possibly due to alteration in other
conditions.
ƒ Some plants as Populus tremuloides and Epilobium
angustifolium are stimulated to growth by fires. A
number of such grasses as Aristida stricta, Cynodon
dactylon, Paspalum notatum etc. are stimulated by fire
to produce large quantities of seeds. In some grasses and
legumes the seeds would germinate only after these get
fire treatment. Their seeds have much degree of
tolerance to temperatures due to fires.
ƒ Common examples studied are species of Themeda,
Heteropogon, Andropogon, Rheus, Tephrosia and Stipa.
ƒ Some fungi, mainly some ascomycetes, grow in
soils of burnt areas. Such fungi are known as
pyrophilous, as for example Pyronema conjluens.
Adaptations to fire
In areas, where fires are very common, the plants have
developed some features that favour their survival under
such conditions. These adaptive features are as follows:
ƒ Some shrubs belonging to the genera Rheus,
Ceanothus, Aractostaphylos produce seeds in large
numbers and whose seed-coats are very hard. The
breakage of hard seed-coats depends on fIre, after which
seedlings appear in large numbers. It is experimentally
proved that their seeds need a temperature of about 80°C
for maximum germination.
ƒ Some woody plants that develop fIre-resistant seeds
or fruits have unique property of completing their life
cycles within a short period by virtue of their rapid rate
of growth and development. Within this short period,
there never accumulates organic debris and thereby
chances of surface fIres reduced. Seedlings of Pinus
palustris grow at rapid rate, and in P. muricata cone
development starts when plants are only one meter tall.
ƒ In some plants leaves are fire-resistant due to poor
contents of such compounds as resin or oil, and thus may
check surface fires. Similarly, Quercus macrocarpa,
Larix occidentalis, Pinus palustris, and P. ponderosa
have thick bark which often escape fire injury.
ƒ Some plants as Pinus rigida and species of
Eucalyptus have adventitious or latent axillary buds
which may develop into new branches. Similarly, Betula
papyrifera, and Vaccinium spp. may develop new shoots
after fire kills the older ones.
ƒ In chaparral vegetation of Europe, North America
and Australia, many shrubs, as species, of Eucalyptus
and Aractostaphylos have evolved special turnip-shaped
swellings of their axes remaining entirely below the
ground surface, which have latent buds. These structures
are known as lignotubers which may produce new
shoots.
ƒ A number of conifers, as Pinus contorta var.
latifolia, P. clausa, P. leiophylla, Picea mariana produce
serotinous cones i.e., cones remaining on trees with
viable seeds for several years sometimes, up to 75 years.
Fires cause the prompt opening of these cones.
III. Topographic (physiographic) Factors
It has just been pointed out that climate of any area is
chiefly determined by the interactions of solar radiation,
atmospheric temperature, relative humidity and annual
rainfall. Each of these climatic factors, besides latitude
and altitude, is greatly influenced by the topography of
the area. We have seen how light, temperature, moisture
UNIT 10: ECOLOGICAL PRINCIPLES 
--etc., may vary at different height on mountains
(altitude) as well as at different distances from the
equator (latitudes).
Topographic factors are concerned with the physical
geography of the earth in the area. Earth's surface is not
similar in different regions and may show several
irregularities. Such topographic factors influence
vegetation by producing variations in the climate of
geographical region. Such variations in climate due to
these factors give rise characteristic local or even
microclimates, or may even modify soil conditions.
Thus, the effect of topographic factors is mainly indirect
primarily through their effects on climatic factors.
The chief topographic factors have been discussed here.
Height of mountain chains
Mountains, hills, hillocks, valleys etc., are formed due to
irregularities in the surface of earth. Effect of different
heights (altitude) can be better seen on mountains. With
an increase in altitude above sea level, there are changes
in values of temperature, pressure, wind velocity,
humidity, intensity of solar radiation. Due to these
changes, vegetation at different altitudes differ much.
We have seen, how the two, eastern and western
Himalayas differ in respect of the vegetation types at
different altitudes. (Fig.1).
Direction of mountains and valleys
Mountains steer or deflect winds into different directions
and capture moisture from wind on certain sides.
Consequently water vapour may accumulate only in
some preferred directions as clouds which come down as
rain. This may be the reason, why on certain sides and at
a particular height one may see luxuriant forests,
whereas on other sides and at lower levels the vegetation
is scanty, in number as well as variety and show xeric
tendencies. Thus outer Himalayas show frequent rains
with luxuriant vegetation, while the middle and inner
Himalayas are dry with poor vegetation. In fact moisture
in the wind is condensed and deposited at the outer
Himalayas before it reaches the. middle and inner ones.
Similarly, southern side of Kulu valley (in Kashmir) is
moist with rich vegetation, whereas at lower level into
Lahul valley there is very poor vegetation.
High mountains not only act as climatic barriers to air
masses between neighboring areas or zones but also
cause local air circulation, giving rise to mountain
winds. The most common winds in such areas are slope
winds which blow down-hill during night and up-hill
during the day. These winds are frequent over glaciers in
Antarctica.
Relative humidity depends upon the direction of the air;
whether it comes from north or south, as well as on the
habitat i.e., from a sea, desert or forest. The heating of
earth's surface is directly proportional to the angle at
which solar rays arrive on the ground. Thus mountains
affect the climate through rainfall and other factors.
10 INSTITUTE FOR ADVANCED STUDIES, JODHPUR
Steepness of slope
Slope is the characteristic feature of mountains.
Steepness of a slope affects the amount of solar radiation
received during the day, and soil characteristics
primarily through its effect on rate of water flow. In the
northern hemispheres, at higher altitudes, steepness of
slope increases the exposure of the surface to sun on the
southern slopes, whereas the northern slopes remain
cooler. This is due to the fact that the steep southern
slopes receive the rays of the mid-day sun almost at right
angles, whereas the northern slopes receive only oblique
rays during morning and evening hours and sometimes
none at all except for a short period during summer. This
difference in solar radiation and consequent temperature
values bring about changes in the vegetation on the two
slopes. Thus, northern slopes (protected from sun)
support virgin forests with hygrophilous ground
vegetation whereas the southern slopes (heated by sun)
have only a xerophytic vegetation. These are due to local
climatic differences on two slopes.
Speed of water flow is also determined by the steepness
of slope. It affects soil characteristics. Most water on the
sloping side flows down rapidly and little is allowed to
reach the soil. Thus loss of water as run off is more with
increase in steepness of the slope. Thus two sides (one
sloppy and other plain) of mountain, inspite with similar
annual rainfall may bear different vegetation. On a steep
slope, plants are unable to establish, even if there is
sufficient rainfall.
Soil erosion is also related with the steepness of the
slope. Even with heavy rainfall on a steep slope, active
soil erosion and denudation due to run off result into
disappearance of the plants from the area. Eroded hills,
steep slopes and rock faces may produce a special
habitat where only a particular group of plants can adapt
themselves. Gravel, sand or clay eroded by water, wind
etc., from rock surfaces, are generally brought down by
rivers and streams and these become then deposited as
silt at their mouths, providing new habitats (like salt
marshes) which may support different kinds of plants.
Exposure of slope
Exposure of slope to sun and wind affects very much the
kind of plants growing there. Generally, the slope
exposed to the sun and wind supports vegetation which
may be entirely different from that which is less or not at
all, exposed to sun and/or wind. This is the reason, why
green houses and hot beds, in temperate climates are
always built in a way so as to face the sun or on southern
slopes, which receive much heat from the sun.
IV. Biotic Factors
Under natural situations, organisms live together
influencing each other's life directly or indirectly. Such
vital processes as growth, nutrition and reproduction
depend very much upon the interactions between the
individuals of same species (intraspecific interaction), or
between those of different species (interspecific
interaction).
Pollination, seeds and fruits dispersal, grazing,
parasitism, symbiosis are the common examples of such
interactions. As a whole, such interactions become
 
UNIT 10: ECOLOGICAL PRINCIPLES 
indeed very much complex. The effect of man upon
vegetation has been much pronounced. Under natural
conditions, we find interdependencies between animals
themselves, between plants themselves, as well as
between plants and animals. Such interactions are found
in various gradations, being for whole life or are casual
and temporary. Moreover, interdependency may exist
between species which are taxonomically widely
different, such as between trees and bacteria, or between
elephants and ants. The relationship between species
may be beneficial to both, harmful to both, or beneficial
or harmful to one and neutral for the other.
For different types of relationships among organisms,
various authors have used different terminology. There
is no general agreement on various terms. We shall now
consider the various kinds of relationships among
organisms and the different terms used for them.
Relationships among organisms
Most of the ecologists are in favour of the use of the
term symbiosis which literally means 'living together', in
its broader sense. The term 'symbiosis', with a similar
intention, was proposed by De Bary (1879). Thus, most
of the ecologists include all types of interactions
including parasitism under 'symbiosis'. Odum (1971)
too, agreeing with the use of the term in its broader
sense, divided all types of symbiotic relationships
between organisms into two groups, (i) positive and (ii)
negative interactions. But others, like Clarke (1954),
restrict the use of the term symbiosis only for such types
of interactions which are mutually beneficial, and where
one or both the species are benefited and neither is
harmed. The relations where at least one of the species is
harmed have been grouped under antagonism.
The various symbiotic phenomena between organisms
have been grouped and interpreted in various ways by
different authors. For instance, Haskell (1949) proposed
the idea of a 'stronger' and 'weaker' partner in the two
species involved during their association. On the basis of
the evident effects of such an association, there were
proposed a number of such terms, as predation,
allotrophy, symbiosis, amensalism, neutrality,
commensalism, synnecrosis, allolimy and parasitism.
McDougall (1918) grouped all, i.e., beneficial as well as
harmful effects of association under the single term
'symbiosis', and classified them into two major groups:
(i) disjunctive symbiosis - where associated organisms
are not in constant contact, and (ii) conjunctive
symbiosis - where dissimilar organisms live in contact
with each other. Each of the two has been further
subdivided into - social (no direct relationship) and
nutritive (with food relationship). The latter may be
antagonistic or reciprocal. Lianas and epiphytes have
been classified under 'social conjunctive symbiosis',
parasites under 'antagonistic nutritive conjunctive
symbiosis', and the lichens, mycorrhizae and nitrogen
INSTITUTE FOR ADVANCED STUDIES, JODHPUR  11
fixing organisms under 'reciprocal nutritive conjunctive
symbiosis'.
Haskell's elaborate classification of 'coactions' between
species is adopted by Burkholder (1952), who on the
basis of several combinations of 0 (no significant
interaction), + (growth, survival, or other population
attribute benefited), and - (population growth or other
attribute inhibited) between two species, the various
types of interactions have been grouped into nine types
of interactions. The possible combinations are 0 0, --, +
+, +0, -0, + -, three of which (++,--, and + -) have in turn
been commonly subdivided, and the whole scheme
resulted into nine types of possible interactions, which
have been explained in Table 1.
Odum (1971) used the term 'symbiosis' in its broader
sense and preferred to group all the above nine types of
symbiotic interactions into two major groups:
Positive interactions
Where populations help one another, the interaction
being either one-way or reciprocal. These include (i)
commensalism, (ii) protocooperation and (iii)
mutualism.
Negative interactions
Where members of one population may eat members of
the other population, compete for foods, excrete
harmful wastes, or otherwise interfere with the other
population. These have been sub-divided into (i)
competition, (ii) predation, (iii) parasitism, and (iv)
antibiosis.
In the present discussion by the author, the term
'symbiosis' has been used in its broader sense, and all
types of interrelationships, following Odum (1971), have
been grouped into above said two major groups.
However, the detailed scheme is somewhat different.
Before we proceed to discuss individual type of
interaction, it would be desirable to present first an
artificial key that would help understanding various
types of such interactions.
UNIT 10: ECOLOGICAL PRINCIPLES 
B1. Concepts of Habitat and Ecological Niche 
Home range: It’s an area where the individual travels to
conduct its normal activities, commonly larger that
defended territory. Theoretically it contains all resources
needed to survive and reproduce over time
Territory: Area of home range defended against con-
specifics. It contains resources that can be defended.
Size varies with resource abundance and population
density. Territorial behavior can restrict numbers of
animals in an area and therefore regulate populations
Habitat: The suite of resources and environmental
conditions that determine the presence, survival, and
reproduction of a population. It’s an organism’s address;
where an organism lives where food, water, cover, and
enough space is available. It actually represents set of
environmental conditions suitable for organism for its
successful growth.
1 Ecological Niche
a. The Concept of Grinell (Spatial or habitat niche) :
The idea of ecological niche has developed a great deal
since it was first defined by Grinell in 1904. According
to this author, niche corresponded to habitat. It concerns
with physical space occupied by an organism. Eg.
Distribution of seven species of millipedes in forest floor
of maple oak forest. As ecology became increasingly a
systemic science, scientists wished to give this simplistic
term a functional meaning, and it was abandoned for
other concepts.
b. The Concept of Odum (Tropic Niche ): The first
systemic definition was given by Odum in 1971. It is
concerned with tropic position of an orgaism. Eg.
Occurance of various species of birds in Galapagos
island.. On the analogy of a visiting card, Odum said that
the habitat of an individual represents an address, the
ecological niche a profession. A doctor does not
necessary live in his clinic, where patients find him
practising is skills. Under the term of profession, it is
necessary to consider the role of various species of the
biocenose, their adaptations, needs, tolerances, way of
life, in sum, their functions in the framework of the
ecosystem.
c. The Concept of Hutchinson: The most holistic
concept is that of Hutchinson, who combined the ideas
of habitats and functions. His idea can be illustrated by
an example cited by Blondel (1979) as described below.
a. Preference of Granivorous Birds vis-à-vis the Weight
of Grain Consumed
Imagine a forest ecosystem in which there are plants pro-
ducing seeds whose weight varies from 0.1 to 10 mg.
Imagine a bird pecking at grains of any weight. By
definition it is behaving like an r-strategist, or generalist.
In its digestive tract are grains of all sizes according to
the curve in Figure. It, in effect, consumes all sizes of
grains available to it.
For the small grains (2-3 mg, for example), the bird will
waste energy by trying to feed on them. Not every peck
will be successful, because the grains are very small.
12 INSTITUTE FOR ADVANCED STUDIES, JODHPUR
Nevertheless, "it expends energy in the form of ATP to
peck at them. It is the same for large grains (8-9 mg, for
example), which will often have to be broken up before
being ingested, which sometimes requires an expenditure
of energy higher than that recovered from the morsels
ingested.
What will happen ultimately is that other birds much
better adapted to picking up a given size of grain will be
more efficient and invade the ecosystem. A bird with a
fine beak will more accurately take the smaller grains,
one with a larger beak will take the large grains.
The r-strategist competing for use of the resources with
several very efficient K-strategists has no alternative but
to emigrate or to remain in the biotope in very small
numbers. .
2. Hutchinson's Multidimensional Model of
Ecological Niche: When the settlement is dominated by
r-strategists, there is a small number of species with a
large number of individuals. On their part, the K-
strategists can live together if each exploits a part of the
environmental resources. The specialist species do not
compete, since they complement each other. The jay, for
example, feeds mostly on acorns, and the woodpecker on
xylophagous larvae. The specialists also occupy all the
ecological niches. Ultimately, they limit losses and
waste.
Does this mean that one of the specialists we have
described in Blondel's example (fig) will be present in
all the ecosystems on earth in which grain of the ap-
propriate size (4-7 mg, say) is available? The answer is
no.
Each species shows preferences with respect to several
parameters. Our bird could also frequent dry zones and
accept only a rainfall between 400 and 800 mm. To go
further, it could prefer temperatures between 12 and
18°C. There would thus be three parameters to consider,
a small number in relation to all the demands and toler-
ances presented by each species taken separately, such as
hygrometry, insolation, wind speed, and competition, to
name only a few.
With the three parameters of grain size, rainfall, and
temperature, we can construct an extremely simplified
 
UNIT 10: ECOLOGICAL PRINCIPLES 
figure that gives an idea of the multidimensional repre-
sentation of the ecological niche of a species (Fig.).
3. Fundamental Niche and Actual Niche: Each animal
or plant species having its own vital preferences can
develop very well within its adaptive potentialities,
taking into account competitions. In consequence, its
ecological niche corresponds to the hypervolume present
in Figure. This hypervolume is called fundamental
niche.
Most frequently, abiotic conditions, and above all
interspecific competition, ensure that the species does
not express its entire genetic potential. It develops only a
part of its potential. The hypervolume obtained in a
given ecosystem is much smaller than what is possible,
and this is the actual or realized niche.
It is therefore possible to find the same species in
seemingly different abiotic and biotic conditions in the
biosphere. These will nevertheless always be within the
hypervolume of the fundamental niche.
Moreover, when we study a species in a defined eco-
system, we analyse only a very small part of its global
population and that part does not present all of the ge-
netic potential that species may have throughout the
world. This means that its actual ecological niche is not
necessarily the potential ecological niche in the common
sense of the term.
Ecological Equivalents: Species that use similar niches
in different habitats or locations.
Guilds: Group of species with comparable role and
niche dimensions within a community.
4. r- and K-strategists and Ecological Niche
a. The Ecological Niche of r-strategists
The r-strategists support variations of abiotic and biotic
conditions for each parameter, taking into account their
low demands. Their ecological niche, which represents
the hyper volume common to all these parameters, will
therefore be very large, so they have large potentialities.
Just as a worker may know a great deal but do a job in a
mediocre way, the r-strategists exploit their environment
poorly. On the other hand, they are very comfortable in a
INSTITUTE FOR ADVANCED STUDIES, JODHPUR  13
new ecosystem because it meets all their needs, since
there are no competitors.
b. The Ecological Niche of K-strategists
Gradually, the generalist species face competition from
less wasteful species that are better adapted and have
more precise functions. These latter species, the K-strat-
egists, are located only on a very small part of the food
spectrum and therefore rarely compete with each other.
There are thus numerous species within a single ecosys-
tem but the number of individuals is limited by the
biological potential of their ecological niche.
As they reduce their energy losses in the cycling of
matter to a minimum, K-strategists soon occupy the en-
tire ecosystem. The hypervolume that characterizes their
specific ecological niche will be very small. They have a
limited function but they carry it out with maximum effi-
ciency. The K-strategists thus have tiny ecological
niches that are very narrow and close to one another, but
each niche is occupied. It is because of this that a mature
ecosystem is well 'balanced'.
c. The Strategists and the Actual Niche We will show
that the fundamental niche of r-strategists is wide and
that of K-strategists very narrow. In fact, the
fundamental niche of r-strategists is so large that their
actual niche in a given ecosystem may appear different
from that found in another ecosystem. Nevertheless, it
remains within the hypervolume of the species consid-
ered on the global scale.
The conditions of life of a copepod (Acanthodiaptam
us), for example, may be defined by Japanese
researchers in a very precise context in their country.
Those researchers may on the other hand be contradicted
by French or Polish researchers to. the extent that the
actual niche of this copepod in any of these countries
constitutes only a part of its fundamental niche on the
global scale.
Such is not the case with the K-strategists, for which the
fundamental niche is so narrow that there are only a few
variations possible for the actual niche.
5. Ecological Niche and the Gause Principle
According to Gause, it is acknowledged that two species
that possess the same ecological niche cannot live to-
gether in the same ecosystem. This is the principle of
reciprocal exclusion, which is linked to competition. If
this principle seems true' in the absolute, it will
nevertheless probably be inapplicable in nature because
there are a number of dimensions of niches in an
ecosystem.
To prove this, we can reproduce an example from
Blondel and Frochot (Fig.). These authors show that bird
species in a forest whose food corresponds to the same
part of the food spectrum live together in a forest by
exploiting different strata.
Resource partitioning: Extinction of one of two
populations living sympatrically and competing over too
many resources is not the only possible outcome of
interspecific competition. An alternative outcome is the
UNIT 10: ECOLOGICAL PRINCIPLES 

evolution of a divergence of resource needs. Such a sympatric populations of two species than allopatric
divergence is called resource partitioning, and is simply population of the same two species is called character
the ecological version of the idea that it is often easier to displacement. That is, characters diverge presumably in
switch than it is to fight response to interspecific competition, but do not diverge
in populations not subject to the same interspecific
Character displacement: Character displacement is competition. Thus, the character differs between the
presumably a consequence resource partitioning. "The population undergoing interspecific competition and the
tendency for characters to be more divergent in population not undergoing interspecific competition
   
B2. Niche differentiation and Resource Partitioning 
 
The  term  niche  differentiation  (synonymous  with  balanced the suitability of their environment must be 
niche  segregation,  niche  separation  and  niche  in order to allow coexistence. There are limits to the 
partitioning),  as  it  applies  to  the  field  of  ecology,  amount  of  niche  differentiation  required  for 
refers  to  the  process  by  which  natural  selection  coexistence,  and  this  can  vary  with  the  type  of 
drives  competing  species  into  different  patterns  of  resource,  the  nature  of  the  environment,  and  the 
resource use or different niches. This process allows  amount  of  variation  both  within  and  between  the 
two species to partition certain resources so that one  species. 
species does not out‐compete the other as dictated by   
the competitive exclusion principle; thus, coexistence  To answer questions about niche differentiation, it is 
is  obtained  through  the  differentiation  of  their  necessary for ecologists to be able to detect, measure, 
realized ecological niches.  and  quantify  the  niches  of  different  coexisting  and 
  competing  species.  This  is  often  done  through  a 
Niche  differentiation  is  a  process  which  occurs  combination  of  detailed  studies,  controlled 
through  several  different  modes  and  on  multiple  experiments  (to  determine  the  strength  of 
temporal  and  spatial  scales.  In  most  cases,  niche  competition),  and  mathematical  models  (Strong 
differentiation  has  created  a  relationship  between  1982, Leibold 1995). To understand the mechanisms 
two  species  where  current  competition  is  small  or  of  niche  differentiation  and  competition,  much  data 
non‐existent.  Because  of  this,  the  presence  of  niche  must  be  gathered  on  how  the  two  species  interact, 
differentiation  can  be  methodologically  difficult  to  how  they  use  their  resources,  and  the  type  of 
prove or disprove. The lack of evidence for current or  ecosystem  in  which  they  exist,  among  other  factors. 
past  competition  can  blur  the  line  between  1)  two  In  addition,  several  mathematical  models  exist  to 
competitive  species  differentiating  their  niches  to  quantify niche breadth, competition, and coexistence 
allow  coexistence  as  opposed  to  2)  two  non‐ (Bastolla  et  al.  2005).  However,  regardless  of 
competing  species  which  occupy  similar  niches.  It  is  methods  used,  niches  and  competition  can  be 
important  to  keep  in  mind  that  niche  differentiation  distinctly difficult to measure quantitatively, and this 
and  inter‐specific  competition  cannot  always  be  makes  detection  and  demonstration  of  niche 
considered linked.  differentiation difficult and complex. 
   
As an example of resource partitioning, seven Anolis  Development of niche differentiation 
lizards  in  tropical  rainforest  share  common  food   
needs  —  mainly  insects.  They  avoid  competition  by  Over time, two competing species can either coexist, 
occupying  different  sections  of  the  rainforest.  Some  through  niche  differentiation  or  other  means,  or 
live on the leaf litter floor while others live on shady  compete  until  one  species  becomes  locally  extinct. 
branches, thereby  avoiding competition over  food in  Several  theories  exist  for  how  niche  differentiation 
those sections of the forest. All resources are subject  arises or evolves given these two possible outcomes. 
to partitioning, for example; space, food, nesting sites.   
This minimizes competition between similar species.  Current competition 
   
Niche  differentiation:  detection  and  Niche  differentiation  can  arise  from  current 
quantification  competition.  For  instance,  species  X  has  a 
  fundamental niche of the entire slope of a hillside, but 
The  Lotka‐Volterra  equation  states  that  two  its realized niche is only the top portion of the slope 
competing  species  can  coexist  when  intra‐specific  because  species  Y,  which  is  a  better  competitor  but 
(within  species)  competition  is  greater  than  inter‐ cannot  survive  on  the  top  portion  of  the  slope,  has 
specific  (between  species)  competition  (Armstrong  excluded it from the lower portion of the slope. With 
and  McGehee  1980).  Since  niche  differentiation  this  scenario,  competition  will  continue  indefinitely 
concentrates  competition  within‐species,  due  to  a  in the middle of the slope between these two species. 
decrease  in  between‐species  competition,  the  Lotka‐ Because  of  this,  detection  of  the  presence  of  niche 
Volterra  model  predicts  that  niche  differentiation  of  differentiation  (through  competition)  will  be 
any degree will result in coexistence.  relatively easy. It is also important to remember that 
  there  is  no  evolutionary  change  of  the  individual 
In  reality,  this  still  leaves  the  question  of  how  much  species in this case; rather this is an ecological effect 
differentiation is needed for coexistence (Hutchinson  of  species  Y  out‐competing  species  X  within  the 
1959).  A  vague  answer  to  this  question  is  that  the  bounds of species Y’s fundamental niche. 
more  similar  two  species  are,  the  more  finely   

14 INSTITUTE FOR ADVANCED STUDIES, JODHPUR 
UNIT 10: ECOLOGICAL PRINCIPLES 

Via past extinctions  when two competing species with overlapping home 
  ranges partition a resource. Two examples would be 
Another way by which niche differentiation can arise  different species of fish feeding at different depths in 
is  via  the  previous  elimination  of  species  without  a  lake  or  different  species  of  monkey  feeding  at 
realized niches. This asserts that at some point in the  different  heights  in  a  tree.  Geographical 
past,  several  species  inhabited  an  area,  and  all  of  differentiation  is  when  two  competing  species  have 
these  species  had  overlapping  fundamental  niches.  non‐overlapping  home  ranges  and  thus  partition 
However,  through  competitive  exclusion,  the  less  resources. An example might be given with monkeys 
competitive species were eliminated, leaving only the  again;  two  competing  species  of  monkey  using  the 
species  that  were  able  to  coexist  (i.e.  the  most  same  species  of  fruit  trees,  but  in  different  areas  of 
competitive  species  whose  realized  niches  did  not  the forest. 
overlap).  Again,  this  process  does  not  include  any   
evolutionary  change  of  individual  species,  but  it  is   
merely  the  product  of  the  competitive  exclusion   
principle.  Also,  because  no  species  is  out‐competing   
any  other  species  in  the  final  community,  the  Morphological differentiation 
presence  of  niche  differentiation  will  be  difficult  or   
impossible to detect.  The  final  type  of  differential  resource  utilization  is 
  morphological  differentiation  or  niche 
Evolving differences  complementarity.  Morphological  differentiation 
  happens  when  two  competing  species  evolve 
Finally,  niche  differentiation  can  arise  as  an  differing  morphologies  to  allow  them  to  use  a 
evolutionary  effect  of  competition.  In  this  case,  two  resource in different ways. A classic example of this is 
competing  species  will  evolve  different  patterns  of  a  study  detailing  the  link  between  bumblebee 
resource  use  so  as  to  avoid  competition.  Here  too,  proboscis  lengths  and  flower  corolla  lengths  (Pyke 
current  competition  is  absent  or  low,  and  therefore  1982).  In  this  study,  the  long‐proboscis  bee  species 
detection  of  niche  differentiation  is  difficult  or  would  preferentially  feed  on  the  long‐corolla  plants, 
impossible.  the medium‐proboscis bee species would feed on the 
  medium‐corolla  plants,  and  so  on.  By  evolving 
Types of niche differentiation  different  proboscis  lengths,  several  competing  bee 
  species  are  able  to  partition  the  available  resources 
Resource partitioning  and coexist. 
   
When two species partition [divide] a resource based  Conditional differentiation 
on behavioral or morphological variation, it is termed   
differential  resource  utilization  or  resource  The  second  form  of  niche  differentiation  is 
partitioning.  There  are  three  types  of  differential  conditional  differentiation,  which  occurs  when  two 
resource utilization.  competing  species  differ  in  their  abilities  to  use  a 
Temporal partitioning  resource based on varying environmental conditions. 
  One  species  may  be  more  competitive  in  one  set  of 
Temporal  resource  partitioning  is  when  two  species  environmental  conditions,  but  another  species  is 
eliminate  direct  competition  by  utilizing  the  same  more  competitive  in  another  set  of  conditions. 
resource  at  different  times.  This  can  be  on  a  daily  Therefore,  in  a  varying  environment,  each  species  is 
scale (e.g. one species of spiny mouse feeds on insects  sometimes  a  better  competitor  and  they  can  coexist. 
during the day while a second species of spiny mouse  Differentiation based on environmental conditions is 
feeds  on  the  same  insects  at  night,  Kronfeld‐Schor  often  difficult  to  separate  from  resource 
and  Dayan  1999)  or  on  a  longer,  seasonal  scale.  An  differentiation,  and  often  conditional  differentiation 
instance  of  the  latter  would  be  reproductive  includes one or more types of resource partitioning. 
asynchrony,  or  the  division  of  resources  by  the   
separation  of  breeding  periods.  An  example  of  Utilization of two resources (Tilman’s R*) 
reproductive  asynchrony  would  be  two  competing   
species  of  frog  offsetting  their  breeding  periods.  By  The  final  type  of  niche  differentiation  is  based  on 
doing  this  the  first  species’  tadpoles  will  have  Tilman’s  (1990)  notion  that  if  two  species  are 
graduated to a different food resource by the time the  competing  for  the  same  exact  resource  then  the 
tadpoles  of  the  second  species  are  hatching  (Lawler  ultimate winner will be the species which can deplete 
and Morin 1993).  the  resource  the  lowest,  surviving  on  the  lowest 
  amount  of  the  resource.  This  alone  does  not  allow 
Spatial partitioning  coexistence.  However,  if  two  species  rely  on  two 
  resources  differentially,  then  coexistence  is  possible 
Spatial  resource  partitioning  occurs  when  two  when  each  species  can  tolerate  a  lower  amount  of 
competing  species  use  the  same  resource  by  only  one  resource  compared  to  its  competitor.  An 
occupying  different  areas  or  habitats  within  the  example  would  be  if  grass  species  1  is  more  limited 
range  of  occurrence  of  the  resource.  Spatial  by  nutrient  B  than  A  and  grass  species  2  is  more 
partitioning  can  occur  at  small  scales  (microhabitat  limited  by  nutrient  A  than  B.  Then,  if  species  1  uses 
differentiation)  or  at  large  scales  (geographical  more of nutrient B than A and species 2 uses more of 
differentiation).  Microhabitat  differentiation  occurs  nutrient A than B, species 1 can out‐compete species 

INSTITUTE FOR ADVANCED STUDIES, JODHPUR  15
UNIT 10: ECOLOGICAL PRINCIPLES 

2  for  nutrient  A  and  species  2  can  out‐compete  Some competing species have been shown to coexist 

species  1  for  nutrient  B.  Additionally,  the  starting  on the same resource with no observable evidence of 
point for each nutrient’s availability must be roughly  niche  differentiation  and  in  “violation”  of  the 
an  equal  distance  from  each  species  limits  (i.e.  no  competitive  exclusion  principle.  One  instance  is  in  a 
species  can  quickly  lower  one  resource  while  the  group of hispine beetle species (Strong 1982). These 
other  is  still  in  abundance).  Coexistence  is  now  beetle  species,  which  eat  the  same  food  and  occupy 
possible  because  each  species  uses  more  of  the  the  same  habitat,  coexist  without  any  evidence  of 
nutrient  that  limits  its  own  growth,  but  each  still  segregation  or  exclusion.  The  beetles  show  no 
requires  both  nutrients  to  survive.  This  subtle  form  aggression  either  intra‐  or  inter‐specifically. 
of  niche  differentiation  is  dependent  on  two  Coexistence  may  be  possible  through  a  combination 
conditions:  1)  the  habitat  needs  to  be  such  that  one  of  non‐limiting  food  and  habitat  resources  and  high 
species  is  more  limited  by  one  resource,  and  the  rates of predation and parasitism, though this has not 
other  species  is  more  limited  by  the  other  resource,  been demonstrated. 
and  2)  each  species  must  consume  more  of  the   
resource that strongly limits its own growth.  This  example  illustrates  that  the  evidence  for  niche 
  differentiation  is  by  no  means  universal.  Niche 
  differentiation  is  also  not  the  only  means  by  which 
  coexistence  is  possible  between  two  competing 
  species  (see  Shmida  and  Ellner  1984).  However, 
  niche  differentiation  is  a  critically  important 
Coexistence  without  niche  differentiation:  ecological  idea  which  explains  species  coexistence, 
exceptions to the rule  thus  promoting  the  high  biodiversity  often  seen  in 
  many of the world’s biomes. 
 
 
B3. Character displacement 
 
Character  displacement  refers  to  the  phenomenon  involved in speciation, stating, “we believe that it is a 
where  differences  among  similar  species  whose  common  aspect  of  geographical  speciation,  arising 
distributions  overlap  geographically  are  accentuated  most often as a product of the genetic and ecological 
in  regions  where  the  species  co‐occur  but  are  interaction  of  two  (or  more)  newly  evolved,  cognate 
minimized or lost where the species’ distributions do  species  [derived  from  the  same  immediate  parental 
not  overlap.  This  pattern  results  from  evolutionary  species]  during  their  period  of  first  contact”  (1956). 
change  driven  by  competition  among  species  for  a  While character displacement is important in various 
limited  resource  (e.g.  food).  The  rationale  for  scenarios of speciation, including adaptive radiations 
character  displacement  stems  from  the  competitive  like  the  cichlid  fish  faunas  in  the  rift  lakes  of  East 
exclusion  principle,  also  called  Gause's  Law,  which  Africa,  it  also  plays  an  important  role  in  structuring 
contends that to coexist in a stable environment two  communities.  The  results  of  numerous  studies 
competing  species  must  differ  in  their  respective  contribute  evidence  that  character  displacement 
ecological  niche;  without  differentiation,  one  species  often influences the evolution of resource acquisition 
will  eliminate  or  exclude  the  other  through  among members of an ecological guild. 
competition.   
  Competitive  release,  defined  as  the  expansion  of  an 
Character displacement was first explicitly explained  ecological  niche  in  the  absence  of  a  competitor,  is 
by  Brown  and  Wilson  (1956):  “Two  closely  related  essentially  the  mirror  image  of  character 
species  have  overlapping  ranges.  In  the  parts  of  the  displacement.  It  too  was  described  by  Brown  and 
ranges  where  one  species  occurs  alone,  the  Wilson  (1956):  “Two  closely  related  species  are 
populations  of  that  species  are  similar  to  the  other  distinct  where  they  occur  together,  but  where  one 
species and may even be very difficult to distinguish  member of the pair occurs alone it converges toward 
from it. In the area of overlap, where the two species  the  second,  even  to  the  extent  of  being  nearly 
occur  together,  the  populations  are  more  divergent  identical with it in some characters.” 
and  easily  distinguished,  i.e.,  they  “displace”  one   
another  in  one  or  more  characters.  The  characters  Examples:  While  studies  on  character  displacement 
involved can be morphological, ecological, behavioral,  have been performed in a wide variety of taxa, a few 
or  physiological;  they  are  assumed  to  be  genetically  groups  have  disproportionately  contributed  our 
based.”  understanding  of  this  principle:  mammalian 
  carnivores,  Galapagos  finches,  Anolis  lizards  on 
Brown  and  Wilson  (1956)  used  the  term  character  islands, three‐spined stickleback fish and snails. 
displacement  to  refer  to  instances  of  both   
reproductive  character  displacement,  or  In  the  initial  explication  of  character  displacement, 
reinforcement  of  reproductive  barriers,  and  many  of  the  examples  they  set  forth  as  potential 
ecological  character  displacement  driven  by  evidence  for  character  displacement  were 
competition.  As  the  term  character  displacement  is  observations  between  multiple  pairs  of  birds.  These 
commonly  used,  it  generally  refers  to  morphological  included  rock  nuthatches  in  Asia,  Australian  honey‐
differences  due  to  competition.  Brown  and  Wilson  eaters  of  the  genus  Myzantha,  Australian  parrots, 
viewed  character  displacement  as  phenomenon  shearwaters in the Cape Verde Islands, flycatchers of 

16 INSTITUTE FOR ADVANCED STUDIES, JODHPUR 
UNIT 10: ECOLOGICAL PRINCIPLES 

the  Bismarck  Archipelago  and  notably,  Darwin's  islands with the bees. Individual finches that harvest 

finches  in  the  Galapagos.  Lack  (1947)  found  that  nectar are smaller than conspecifics that do not. 
when the two species Geospiza fortis and G. fuliginosa   
occurred  on  large  islands  together,  they  could  be  Introduced  species  have  also  provided  recent 
distinguished  unequivocally  by  beak  size.  When  “natural  experiments”  to  investigate  how  rapidly 
either  one  occurred  by  itself  on  a  smaller  island,  character  displacement  can  effect  evolutionary 
however,  the  beak  size  was  intermediate  in  size  change.  When  American  mink  (Mustela  vison)  was 
relative to when the two co‐occurred.  introduced  in  north‐eastern  Belarus,  the  native 
  European  mink  (Mustela  lutreola)  increased  in  size, 
The  lizard  genus  Anolis  on  the  islands  in  the  and  the  introduced  mink  decreased  in  size.  This 
Caribbean  has  also  been  the  subject  of  numerous  displacement  was  observed  within  a  ten‐year  study, 
studies  investigating  the  role  of  competition  and  demonstrating  that  competition  can  drive  rapid 
character  displacement  in  community  structure.   evolutionary change. 
Lesser  Antilles  islands  can  only  support  Anolis   
species of different sizes, and the relative importance  Conceptual  development:  “Character  displacement 
of character displacement versus size at colonization  is the situation in which, when two species of animals 
in  determining  invasion  success  has  been  explored  overlap geographically, the differences between them 
and debated.  are  accentuated  in  the  zone  of  sympatry  and 
  weakened or lost entirely in the parts of their ranges 
Three spine sticklebacks (Gasterosteus spp.) in post‐ outside  this  zone”.  While  the  term  “ecological 
glacial  lakes  in  western  Canada  have  contributed  character  displacement”  first  appeared  in  the 
significantly  to  recent  research  of  character  scientific literature in 1956, the idea has earlier roots. 
displacement.  Both  observations  of  natural  For example, Joseph Grinnell, in the classic paper that 
populations and manipulative experiments show that  set  forth  the  concept  of  the  ecological  niche  (1917), 
when  two  recently  evolved  species  occur  in  a  single  stated, “It is, of course, axiomatic that no two species 
lake,  two  morphologies  are  selected  for:  a  limnetic  regularly established in a single fauna have precisely 
form that feeds in open water and a benthic form that  the  same  niche  requirements.”  The  existence  of 
feeds  at  the  lake  bottom.  They  differ  in  size,  shape  character  displacement  is  evidence  that  the  two 
and the number and length of gill rakers, all of which  species  do  not  completely  overlap  in  their  niche 
is  related  to  divergence  in  their  diet.  Hybrids  requirement. 
between  the  two  forms  are  selected  against.  When   
only one species inhabits a lake, that fish displays an  Following the dissemination of the concept, character 
intermediate  morphology.  Studies  on  other  fish  displacement  was  viewed  as  an  important  force  in 
species  have  shown  similar  patterns  of  selection  for  structuring  ecological  communities,  and  biologists 
benthic  and  limnetic  morphologies,  which  can  also  identified numerous examples. During the late 1970s 
lead to sympatric speciation.  and  early  1980s,  however,  the  role  of  competition 
  and  character  displacement  in  structuring 
The  Appalachian  salamanders  Plethodon  hoffmani  communities  was  questioned  and  its  importance 
and  P.  cinereus  displayed  no  trophic,  morphological  greatly downgraded. Many found the early examples 
or  resource  use  differences  among  allopatric  unconvincing  and  suggested  it  to  be  a  rare 
populations; when the species occurred in sympatry,  phenomenon. Criticisms with earlier studies included 
however,  they  displayed  morphological  the lack of rigor in statistical analyses and the use of 
differentiation  that  was  associated  with  segregation  poorly  rationalized  characters.  Additionally,  theory 
in prey size. Where these two species co‐occurred, P.  seemed  to  indicate  that  the  conditions  that  allowed 
hoffmani  had  a faster closing jaw required for larger  character  displacement  to  occur  were  limited.  This 
prey,  and  P.  cinereus  had  a  slower,  stronger  jaw  for  scrutiny  helped  motivate  theoretical  and 
smaller  prey.  Other  studies  have  found  Plethodon  methodological advances as well as the development 
salamander  species  that  demonstrate  character  of  a  more  rigorous  framework  for  testing  character 
displacement  from  aggressive  behavioral  displacement. 
interference  rather  than  exploitation.  That  is,   
morphological  character  displacement  between  the  Six  criteria  have  been  developed  to  establish 
two species is due to aggressive interaction between  character  displacement  as  the  mechanism  for 
them  rather  than  the  exploitation  of  different  food  differences  between  sympatric  species.  These 
resources.  It  is  often  assumed  that  closely  related  include:  (1)  differences  between  sympatric  taxa  are 
species  are  more  likely  to  compete  than  are  more  greater  than  expected  by  chance;  (2)  differences  in 
distantly  related  species,  and  hence  many  character states are related to differences in resource 
researchers  investigate  character  displacement  use;  (3)  resources  are  limiting,  and  interspecific 
among congeners. While character displacement was  competition  for  these  resources  is  a  function  of 
originally  discussed  in  the  context  of  very  closely  character similarity; (4) resource distribution are the 
related  species,  evidence  suggests  that  even  same in sympatry and allopatry such that differences 
interactions  among  distantly  related  species  can  in  character  states  are  not  due  to  differences  in 
result in character displacement. Finches and bees in  resource  availability;  (5)  differences  must  have 
the  Galapagos  may  provide  an  interesting  example.  evolved  in  situ;  (6)  differences  must  be  genetically 
Two finch species (Geospiza fuliginosa and G. difficilis)  based. Rigorously testing these criteria necessitates a 
exploit more flower nectar on islands where the lager  synthetic approach, combining areas of research like 
carpenter  bee  (Xylocopa  darwini)  is  absent  than  on  community  ecology,  functional  morphology, 

INSTITUTE FOR ADVANCED STUDIES, JODHPUR  17
UNIT 10: ECOLOGICAL PRINCIPLES 

adaptation,  quantitative  genetics  and  phylogenetic 

systematics, While satisfying all six criteria in a single 
study of character displacement is not often feasible, 
they  provide  the  necessary  context  for  research  of 
character displacement. 

18 INSTITUTE FOR ADVANCED STUDIES, JODHPUR 
UNIT 10: ECOLOGICAL PRINCIPLES 
C1. POPULATION ECOLOGY:
Population ecology studies organisms from the point of
view of the size and structure of their populations A
population ecologist studies the interaction of organisms
with their environments by measuring properties of
populations rather than the behavior of individual
organisms
Properties of populations include
ƒ Population size (size)
ƒ Population density (density)
ƒ Patterns of dispersion (dispersion)
ƒ Demographics (demographics)
ƒ Population growth (growth)
ƒ Limits on population growth (limits)
Note that all of these properties are not those of
individual organisms but instead are properties which
exist only if one considers more than one organism at
any given time, or over a period of time (i.e., they are
emergent properties)
"The characteristics of a population are shaped by the
interactions between individuals and their environments
on both ecological and evolutionary time scales, and
natural selection can modify these characteristics in a
population."
Thus, population ecology also goes beyond
consideration of just population parameters and
additionally considers how the characteristics of
individual organisms impact on population parameters
POPULATION PROPERTIES
(1) Population
A population in an ecological sense is a group of
organisms, of the same species, which roughly occupy
the same geographical area at the same time
Individual members of the same population can either
interact directly, or may interact with the dispersing
progeny of other members of the same population (e.g.,
pollen). Population members interact with a similar
environment and experience similar environmental
limitations
(2) Population size
A population's size depends on how the population is
defined. If a population is defined in terms of some
degree of reproductive isolation, then that population's
size is the size of its gene pool.
If a population is defined in terms of some geographical
range, then that population's size is the number of
individuals living in the defined area
Ecologists typically are more concerned with the latter
means of defining a population since this is both easier
to do and is a more practical measure if one is interested
in determining the impact of a given population on a
given ecosystem, or vice versa
INSTITUTE FOR ADVANCED STUDIES, JODHPUR  19
(3) Population density
Given that a population is defined in terms of some
natural or arbitrarily defined geographical range, then
population density may be defined as simply the number
of individual organisms per unit area.
Different species, of course, exist at different densities in
their environments, and the same species may be able to
achieve one density in one environment and another in a
different environment. Population densities may
additionally be determined in terms of some measure
other than population size per unit area such as
population mass per unit area.
Crude density: is the density (number of biomass) per
unit total space.
Specific or ecological or economic density: is the
density (number of biomass) per unit of habitat space i.e.
available area or volume that can actually be colonized
by the population.
(4) Patterns of dispersion
Individual members of populations may be distributed
over a geographical area in a number of different ways
including
ƒ Clumped distribution (attraction): Clumping
may result either from individual organisms being
attracted to each other, or individual organisms being
attracted more to some patches within a range than they
are to other patches; the net effect is that some parts of
the range will have a large number of individuals
whereas others will contain few or none
ƒ Uniform distribution (repulsion): A uniform
distribution means that approximately the same distance
may be found between individual organisms; uniform
distributions result from individual organisms actively
repelling each other
ƒ Random distribution (minimal
interaction/influence): A random distribution means
that where individual organisms are found is only
minimally influenced by interactions with other
members of the same population, and random
distributions are uncommon; "Random spacing occurs in
the absence of strong attractions or repulsions among
individuals of a population."
Note that both clumping and uniform distributions
suggest that individual organisms are either interacting
with one another (actively seeking each other out or
actively avoiding each other), or are all competing with
one another for the same limited resources, regardless of
the overall population density (as in the case of clumping
which results from geographical patchiness)
(4) Demographics
A population's demographics are its vital statistics,
particularly those statistics which can impact on present
and future population size. Two statistics that are of
particular import are a population's age structure and a
population's sex ratio.
UNIT 10: ECOLOGICAL PRINCIPLES 
Age structure: Age structure refers to the size of
cohorts within a population. Parameters related to age
structure include
Cohort: A cohort is a group of individuals all of whom
have the same age. In a typical population, the size of
cohorts will vary with age. For example, in a typical
population, younger cohorts will be larger (i.e., more
individuals per cohort) than older cohorts, all else being
equal.
Natality (Birth rate): Natality rate is the number of
offspring per female per unit time. There are two type of
natality
ƒ Fecundity/Maximum (absolute or potential)
natality: It is theoretical maximum production of new
individuals under ideal conditions when resources are
not limited. The greater a population's fecundity, all else
held constant, the faster a population will increase in
size. Note that fecundity typically varies with the age of
individuals.
ƒ Ecological or realized natality: It refers to
population increase under actual, existing specific
conditions. This is also designated as fertility rate.
Generation time: Generation time is simply the average
span between the birth of individuals and the birth of
their offspring. "Other factors being equal, a shorter
generation time will result in faster population growth."
Note that species which are capable of reproducing more
than once will display an overlapping of generations
which basically means that parental cohorts and progeny
cohorts can be alive (and potentially competing with one
another) at the same time. Note that another way of
saying this is that when life expectancies exceed the
minimum time between generations, generations will
overlap.
Mortality/ Death rate: Death rate is the rate at which
individuals of a certain age die. Note that death rates
often vary with age with either the very young or the
very old displaying the greatest death rates. Note
additionally that population growth occurs when overall
birth rates exceed overall death rates
Sex ratio: More often than not the rate at which a
population may grow is dependent on the sex ratio in the
population; the fewer females, the slower the rate of
population growth. This, of course, is because uteruses
are limiting and males often can inseminate more than
one female. This generalization falls apart, however,
when males are limited in their ability to inseminate
more than one female, or when males contribute
significantly to the raising of offspring
Survivorship curves: Observing age structure
graphically can provide insights into a species' (or a
population's) ecology. Survivorship curves graph cohort
size against relative age. The typical survivorship curve
shows cohort size declining with age. There exist three
general types of survivorship curves
ƒ Type I, II and III
Note in the following survivorship curves that the y axis
is logarithmic.
20 INSTITUTE FOR ADVANCED STUDIES, JODHPUR
Type II survivorship curves: The simplest type of
decline is exponential, i.e., the death rate for every
cohort is the same. These survivorship curves graph as a
straight line on semi-logarithmic graph paper (i.e., as
presented in a typical survivorship curve). The
individuals in populations that display a type II curve are
those that both do not age and are born as fully fit as
adults, e.g., hydra. Individuals are lost in these
populations mostly to accidents and predation. Other
examples include butterfly, birds, mice, rabbit and most
of holometabolous insects.
Type I survivorship curves: Because individuals tend
to die exponentially due to accidents or predation, it
often is a good strategy to reproduce relatively early in a
life span rather than relatively late. That way individual
achieve reproduction while they still have a reasonable
likelihood of being alive. This is assuming, of course,
that the goal is a Darwinian one, i.e., maximizing one's
reproductive output. Note that how such a strategy works
is complicated if individual fecundity increases with age.
Very often for a given species there will be some age at
which individuals are maximally fecund. Species that
combine maximum fecundity with early ages typically
do so at the expense of their ability to survive long
periods (i.e., this is an example of the principle of
allocation). A survivorship curve of such individuals
may display a relatively shallow slope while individuals
are younger (i.e., maximally robust and maximally
reproductive) but then show an abrupt increase in death
rate at ages that are coincident to declines in fecundity.
Humans, of course, have a type I survivorship curve. It is
displayed by most of higher organisms such as mammals
and even by Drosophila.
Type III survivorship curves: The other side of the
survivorship coin is the degree of investment in
individual progeny. Some organisms invest a great deal
in each offspring and those organisms are (ideally at
least) rewarded with relatively high survivorship at early
ages. Other organisms invest little in individual
offspring, and display very low early-age survivorship
(which they make up for by producing buckets of
offspring). Organisms that produce large numbers of
cheap progeny and which display minimal declines in
fecundity with age, if they survive their youth, display
type III survivorship curves. Examples include sea
turtles, Oysters, pelagic fishes and trees.
That is, type III survivorship species have a very large
rate of mortality when young, but should they survive
their youth, they put significant energy into continued
 
UNIT 10: ECOLOGICAL PRINCIPLES 
survival since the longer they survive, the more progeny
they will produce
Life history: “The traits that affect an organism’s
schedule of reproduction and survival (from birth
through reproduction to death) make up its life history.”
The study of life history characteristics is the detailed
study of those ecological and evolutionary parameters
that impact on survivorship curves.
"In many cases there are trade-offs between survival and
traits such as clutch size (number of offspring per
reproductive episode), frequency of reproduction, and
investment in parental care. The traits that affect an
organism's schedule of reproduction and death make up
its life history. Of course, a particular life history pattern,
like most characteristics of an organism, is the result of
natural selection operating over evolutionary time."
In other words, the Darwinian goal is to maximize
lifetime reproductive output, and this can be achieved
by having babies more rapidly or living longer, or some
combination of the two, as well as by varying many
additional details having to do with survival and
reproduction
Allocation of limited resources:
"Darwinian fitness is measured not by how many
offspring are produced but by how many survive to
produce their own offspring: Heritable characteristics of
life history that result in the most reproductively
successful descendants will become more common
within the population. If we were to construct a
hypothetical life history that would yield the greatest
lifetime reproductive output, we might imagine a
population of individuals that begin reproducing at an
early age, have large clutch sizes, and reproduce many
times in a lifetime. However, natural selection cannot
maximize all these variables simultaneously, because
organisms have a finite energy budget that mandates
trade-offs. For example, the production of many
offspring with little chance of survival may result in
fewer offspring that can compete vigorously for limited
resources in an already dense population."
“The life history we observe in organisms represent a
resolution of several conflicting demands. An important
part of the study of life histories has been understanding
the relationship between limited resources and
competing functions: Time, energy, and nutrients that
are used for one thing cannot be used for something
else."
"These issues can be phrased in terms of three basic
questions:
ƒ How often should an organism breed?
ƒ When should it begin to reproduce?
ƒ How many offspring should it produce during each
reproductive episode?
ƒ percentage of young individuals. In rapidly growing
The way each population resolves these questions results
in the integrated life history patterns we see in nature."
(all one quote starting with (c) but broken up for clarity).
“Many life history issues involve balancing the profit of
immediate investment in offspring against the cost to
INSTITUTE FOR ADVANCED STUDIES, JODHPUR  21
future prospects of survival and reproduction. These
issues can be summarized by three basic “decisions”:
when to begin reproducing, how often to breed, and how
many offspring to produce during each reproductive
episode. The various “choices” are integrated into the
life history patterns we see in nature.”
Semelparity (big bang): Organisms that produce one
clutch of offspring (progeny) per life time are said to be
semelparous (i.e., to display semelparity). The advantage
of semelparity is that at the point of reproduction few if
any resources need be devoted to survival past
reproduction
Iteroparity (repeated reproduction): Organisms that
produce more than one clutch of offspring (progeny) per
life time are said to be iteroparous (i.e., to display
iteroparity). The advantage of iteroparity is that it allows
organisms to display more than one statistical “shot” at
producing a successful litter. “The critical factor in the
evolutionary dilemma of big-bang versus repeated
reproduction is the survival rate of the offspring. If their
chance of survival is poor or inconsistent, repeated
reproduction will be favored.”
(5) Age Structure
In most populations, individuals are of different ages.
The proportion of individuals in wach group is called
age structure of that population. For instance an
understory palm tree populations (Astrocayum
mexicanum) in evergreen forest of Mexico, had 50 %
individuals as seedlings (less than 2 yr old), 19 % as
saplings (8 year old), 5 % as 30 year old adults and so
until 70-year old trees made up less than 2% of the
population.
Age Pyramids: Age distribution is important, as it
influences both, natality and mortality of the population.
The ratio of the various age groups in a population
determines the current reproductive status of the
population, thus anticipating its future. From an
ecological view point there are three major ecological
ages (age groups) in any population. These are, pre-
reproductive, reproductive and post-reproductive. The
relative duration of these age groups in proportion to the
life span varies greatly with different organisms. In man,
the three 'ages' are relatively equal in length, about a
third of his life falling in each class. Many plants and
animals have a very long prereproductive period. Some
animals, particularly insects, have extremely long pre-
reproductive periods, a very short reproductive period
and no post-reproductive period.
The model representing geometrically the proportions of
different age groups in the population of any organism is
called age pyramid. The three hypothetical pyramid
types are shown in Figure which are:
A pyramid with broad base: It indicates a high
young populations birth rate is high and population
growth may be exponential as in yeast, house-fly,
paramecium etc. Under such conditions, each successive
generation will be more numerous than the preceding
one, and thus a pyramid-with broad base would result
(Fig. A).
UNIT 10: ECOLOGICAL PRINCIPLES 
A bell-shaped polygon: It indicates a moderate
proportion of young to old. As the rate of growth
becomes slow and stable, i.e. the pre-reproductive and
reproductive age group: become more or less equal in
size, post-reproductive group remaining as the smallest,
there results a bell shaped structure (Fig. B).
An urn-shaped figure: It indicates a low percentage of
young individuals If the birth rate is drastically reduced
the prereproductive group dwindles in proportion to the
other two groups and it results in an urn-shape, figure
(Fig. C), which indicates that population is dying off.
While density reveals little about difference between
populations in developed and developing countries, age
structure highlights some of the cause of rapid
population growth in the latter. Developing countries
have a much greater preponderance of young people, and
the side of pyramids become concave. Such a shape
means high population growth, not only at present but
also likely in future.
Life tables:
Information on natality and mortality in different ages
and sexes can be combined in the form of life tables.
From these it is possible to estimate the growth or
decline of a population. As with survivorship curves, life
tables are standardised to follow the progress of a cohort.
In each table there are columns for age of individuals;
number surviving to each age; the number dying in each
age group; the proportion dying from the previous age
category; fertility rate; and the number of young born by
each age groups, The information obtained from these
figures provides the net reproductive rate of the
population i.e., offspring left by each individual.
Similarly from life table, mortality in a logarithmic form
is also obtained. These are then used to calculate the rate
of population growth.
(6) Population growth
The simplest case of population growth is that which
occurs when there exist no limitations on growth within
the environment. In such situations two things occur
ƒ The population displays its intrinsic rate of increase
ƒ The population experiences exponential growth
Biotic potential & Intrinsic rate of growth
Each population has the inherent power to grow. When
the environment is unlimited (space, food, other
organisms not exerting a limiting effect), the specific
growth rate (i.e., the population growth rate per
individual) becomes constant and maximum for the
existing conditions. The value of the growth rate under
these favourable conditions is maximal, is characteristic
22 INSTITUTE FOR ADVANCED STUDIES, JODHPUR
of a particular population age structure, and is a single
index of the inherent power of a population to grow. It
may be designated by the symbol r; which is the
exponent in the differential equation for population
growth in an unlimited environment under specific
physical conditions. 'r' represents intrinsic rate of
population increase.
The index r is actually the difference between the
instantaneous specific natality rate (i.e., rate per time per
individual) and the instantaneous specific death rate and
may thus be expressed:
r=b-d
The overall population growth rate under unlimited
environmental conditions (r) depends on the age
composition and the specific growth rates due to
reproduction of component age groups. Thus, there may
be several values of r for a species depending upon
population structure. When a stationary and stable age
distribution exists, the specific growth rate is called the
intrinsic rate of natural increase or r max. The maximum
value of r is often called by the less specific but widely
used expression biotic potential or reproductive
potential. Thus with this term, one is able to put together
natality, mortality and age distribution. The ranges of the
intrinsic rate of natural increase for various kinds of
organism are shown in Table 7.1. Chapman (1928)
proposed the term biotic potential to designate maximum
reproductive power. He defined it as "the inherent
property of an organism to reproduce, to survive, i.e., to
increase in numbers. It is sort of the algebraic sum of the
number of young produced at each reproduction, the
number of reproduction in a given period of time, the sex
ratio and their general ability to survive under given
physical conditions." For details, reader may refer to
Graham (1952). But under natural conditions, this is a
rare phenomenon, since environmental conditions do not
permit unlimited growth of any population. Its size is
kept under natural check. The difference between the
maximum r (biotic potential) and the rate of increase
which occurs in an actual laboratory or field condition is
often taken as a measure of the environmental resistance,
which is the sum total of environmental limiting factors
which prevent the biotic potential from being realised.
Organisms Approx biotic potential,
r (per year)
Large mammals 0.02 - 0.5
Birds 0.05 - 1.5
Small mammals 0.3 - 8
Larger invertebrates 10 - 30
Insects 4 - 50
Small invertebrates 30 - 800
(including large protozoans)
Protozoa & Unicellular algae 600 - 2,000
Bacteria 3,000 - 20,00
Population Dynamics
After having studied the various characteristics of a
population, we attempt to follow the change in it
Population have characteristic patterns of increase which
are called population growth forms. Such growth forms
represent the interaction of biotic potential and
environmental resistance. What are the theoretical bases
that attempt to account for such changes?
 
UNIT 10: ECOLOGICAL PRINCIPLES 
Approaches to study of population dynamics are, (i)
mathematical models (ii) laboratory studies and (iii) field
studies. Mathematical models are of two types:
theoretical models and simulation models. The former
derive equations that may describe population changes;
and the behaviour of such models can be tested by
comparing these with real populations. The latter
(simulation models) use detailed data from real
populations and try to predict future behaviour of these
populations under specific environmental conditions.
Models are of use only if they explain facets of
population dynamics of real populations. Their
predictions are often tested experimentally using
laboratory populations. Mathematical and laboratory
studies are of value only if they help to explain the
behaviour of natural populations. Field studies are made,
though these are complex.
Theory of population growth
Exponential growth: Exponential growth simply means
that a population's size at a given time is equal to the
population's size at an earlier time, times some greater-
than-one number. For example, if a population increased
in size per unit time in the following manner: 1, 2, 4, 8,
16, 32, 64, 128, etc. (or, e.g., 1, 3, 9, 27…, or 1, 5, 25,
125, …, etc.) then the population is displaying
exponential growth, each unit time the population is
increasing by a factor of 2 (or 3 or 5 in the other
examples; note that exponential growth is occurring so
long as the rate of increase per unit time is greater than a
factor of 1, e.g., 2 or 4 or 10 or 1.2, etc.).
When population size is graphed against time (e.g.,
generations) a population growing exponentially
displays a J-shaped curve. Note differences in intrinsic
rates of growth, in this J-shaped curves, that result in
differences in rates of exponential growth (declining
intrinsic growth rates are seen going from left to right in
this graph):
There may be assumed a model of geometric increase
that assumes that there is no environmental constraint on
population growth. A geometrical series is one with a
constant multiplier. The simplest example is binary
fission in Protozoa, where each individual gives two
each time it reproduces. Starting with one individual
(generation 0), there will be 2,4,8,16,32,64... in
succeeding genera-lions if fission is synchronous. To
determine number (N 1) present in any generation (t),
the following equation is used: Nt = No(2t), No is no.
present in generation 0. Thus the number in generation 3
will be N°3 = 1 (23) = 8
This equation is for a population that doubles in each
generation. A more generation equation would be, Nt=
No(ert)
This is the exponential growth equation, where er is
substituted for 2; e is a universal constant, the base of
INSTITUTE FOR ADVANCED STUDIES, JODHPUR  23
natural logarithms and has a value of 2.718; r is a
constant for a particular population under specific
environmental conditions and is called the intrinsic rate
of natural increase. The significance of logarithms in
exponential growth is shown in Figure which compares
the predictions of the binary fission equation on
arithmetic and logarithmic scales. The logarithmic plot is
a straight line, meaning that the number added in any
time period is proportional to the number present at the
beginning of the period (compare with survivorship
curve (diagonal) in Figure 6 which is an exponential
decline).
The intrinsic rate of natural increase (r) is easy to
calculate if natality and mortality rates are known. For
example, for each of the 14 million people present in a
country in 1980, 0.028 were born and 0.008 .~ed during
the year; r = natality rate minus mortality rate, or 0.028-
0.008 per year. Substituting in the exponential growth
equation, the number present in 1990 is predicted as:
Nt = No (ert)
N10 = 14 million x 2.7180.02 x 10 = 17 million
Note this is only strictly correct if the population is
growing exponentially with constant birth and death
rates and age structure, and there is no immigration or
emigration.
[In a rich culture medium
bacteria, grown under aerobic
conditions, achieve a final
concentration of 2-5 x 109
cells per ml in about 12-18
hours. Although plotted on a
different time scale the human
growth curve looks the same;
the human population at
similar points on the growth curve are shown. When
population size is graphed against time (e.g.,
generations) a population growing exponentially
displays a straight line curve when graphed on semi-
logarithmic graph paper (for example, below is a graph
of the exponential increase in the computer processing
power available per dollar—note that on log-linear graph
paper this curve is approximately a straight line):
"The J-shaped curve of exponential growth is
characteristic of populations that are introduced into a
new or unfilled environment, or whose numbers have
been drastically reduced by a catastrophic event and are
rebounding." In other words, a population that is in an
environment lacking limits will grow exponentially
(indeed, a population that is capable of growing will tend
UNIT 10: ECOLOGICAL PRINCIPLES 

to grow exponentially), and the rate at which growth will
occur will be a function of rmax and the degree to which
the environment matches the ideal environment in which
an organism is capable of achieving rmax.
Does it occur in nature? Most populations do not show
exponential increase because their environment prevents
this. In an ecosystem, other species present, a variety of
abiotic conditions etc. act in a way that a given
population will have an upper size limit that cannot be
exceeded. Such a limit is called carrying capacity of the
environment for that population. Carrying capacity is
determined by the availability of resource as food and
space.
population density approaches that dictated by an
environment’s carry capacity for that population. Note
that populations do not typically display the idealized
logistic growth seen with the model. One deviation from
idealized logistic growth is delayed feedback; this can
cause population size overshooting and, in fact, what is
typically observed in real populations is not just effects
of random events but also populations sizes which vary
up and down around the carrying capacity rather than
remaining invariant exactly at the carrying capacity

Limits on population growth: Exponential growth

cannot go on forever; sooner or later any population will
run into limits in their environment.

Carrying capacity (K):

"Populations subsist on a finite amount of available
resources, and as the population becomes more crowded,
each individual has access to an increasingly smaller
share. Ultimately, there is a limit to the number of
individuals that can occupy a habitat. Ecologists define
carrying capacity as the maximum stable population size
that a particular environment can support over a
relatively long period of time. Carrying capacity,
symbolized as K, is a property of the environment, and it
varies over space and time with the abundance of
limiting resources."
In other words, for any given organism, there will be a
maximum number of individuals that the environment
can support without the environment being consequently
degraded to the point where it can no longer support that
number of individuals.
Generally, as population size approaches carrying
capacity, the amount of some key resource declines per
capita to the point where individuals experience either a
higher death rate or a lower fecundity; thus, as
population size approaches carrying capacity, the rate of
population growth declines towards zero.
The logistic equation was devised by Verhulst in 1838 to
describe population growth with an upper limit.
Suppose, environment has a carrying capacity (K) for a
particular population. Then the logistic equation assumes
that the intrinsic rate of natural increase (r). is
progressively reduced as population size increases
toward that carrying capacity. If there are 10 individuals
already present (N = 10) and K = 100 individuals, the
remaining resources can support a further K-N = 90
individuals and the proportion of remaining resources is
K-N/K = 0.9.
The logistic equation simply assumes that r is reduced in
relation to proportion of remaining resources. The
equation is derived from the exponential such that:
⎡K − N ⎤
t
N t = N 0e r ⎢
⎣ K ⎥⎦
the logistic growth equation.

Logistic growth: Logistic growth is a mathematical

description of population growth that employs two
parameters, rmax and K, and two variables, N and t . The
logistic growth curve is S-shaped .That is, the population
grows exponentially at a rate which is determined by rmax
and the suitability of a given environment to an
organism’s needs until population size is sufficient that
the limitations associated with the carrying capacity of
the environment are approached. This slows the rate of
population growth in a way such that the larger the
population becomes, the slower its rate of growth; this
Suppose r = 1; then when N = 10, the growth rate
slowing of the growth transforms the curve from a J-
actually achieved, r (K-N/K) is 1 x 0.9. When N=90, (K-
shaped one to an S-shaped one. Ultimately the rate of
N/K) = 0.1 and the growth rate is 1 x 0.1 and so on,
growth of the population reaches zero at the carrying
giving the progressive decline in realised growth rate.
capacity.
The logistic curve is S-shaped on arithmetic coordinates
and is always less steep than its exponential equivalent
"Because the rate at which a population grows changes
(Fig. A). A comparison of the same curve on semi-
with the density of organisms that are currently in the
logarithmic coordinates is given in Figure 7 B. using
population, the logistic model is said to be density
dependent." That is, population growth grows as

24 INSTITUTE FOR ADVANCED STUDIES, JODHPUR 
UNIT 10: ECOLOGICAL PRINCIPLES 
differential calculus, the logistic equation becomes
dN (K − N )
= rN
dt K there is an upper limit to population density set by the
Growth of laboratory and field populations
From the above account of carrying capacity, one would
expect exponential growth only under circumstances of
no environmental resistance. Such a growth is possible
in laboratory culture if excess organisms are removed to
avoid over crowding. However, in nature, exponential
increase in population experiences catastrophic decline
for insect population. Under adequate food and climate,
population increased, but then declined when these
turned unfavourable.
In the laboratory logistic growth has been observed in
bacteria, yeasts, algae and protozoa. This is shown for
Paramecium. In laboratory cultures, somewhat much less
fit to logistic is usually found in metazoans as beetle,
like azuki bean weevil (Callosobruchus chinensis), but
populations do often fluctuate around a carrying
capacity. There are several reasons as competition, loss
in female fecundity, increased egg mortality etc. at high
density of this beetle. In the field, direct observation of
variatation and limitation of a population are rare.
However, there is a logistic model of wildebeest
(Connochaetes taurinus) in Tanzania and Kenya. The
population levels off at K determined by food
availability following its low density.
Modifications of logistic theory
Following assumptions are implicit in the logistic
equation: (i) environment is constant such that r and K
do not charge, (ii) there arc no time lags in response of
the population to change in its density, (iii) individuals
have identical ecological characteristics in terms of their
responses to density, regardless of age or sex (iv) growth
is continuous and (v) age structure does not change in
successive generations. These assumptions are unlikely
to be met in laboratory and will never obtain in the field.
Therefore, logistic theory has been suitably modified to
make it biologically reasonable. These are (i) inclusion
of a time lag there (ii) there are fluctuations in the
carrying capacity, (iii) there may be chance events for
survival and reproduction of an individual as well as for
their death, due to landslide etc. (iv) discontinuous
growth of some populations which grow in discrete steps
with little or no overlap between successive generations.
Keeping in view the above, it is natural to ask "Are, then
mathematical models of any use? The answer is "Yes"
because
(l) as theoretical abstractions, they set the crude limits to
possible patterns of population growth. Observations on
real populations can be assessed within these limits.
(2) as models become more refined, they more closely fit
the behaviour of real populations. One can make
increasingly accurate predictions of population
behaviour.
(3) From above 1 and 2, it follows that population
models can be applied to real practical problems as
human population growth and the management of plant
and animal populations.
(7) Regulation of population density:
INSTITUTE FOR ADVANCED STUDIES, JODHPUR  25
The logistic model and its derivatives assume that a
population will level off at its carrying capacity, that
environment and that the population is regulated at or
around that level. The significance of regulation in
population has been a subject of controversy among
ecologists for last more than years. What regulates
population? Krebs (1985) presented an account of this
subject.
Nature of factors that influence population density
Population density can only be increased by natality or
immigration and decreased by mortality or emigration.
These factors may be density dependent or density
independent in their effects.
Density-dependent factors: Density-dependent limits
on population growth are ones that stem from
intraspecific competition. Typically, the organisms best
suited to compete with another organism are those from
the same species. Thus, the actions of conspecifics can
very precisely serve to limit the environment (e.g., eat
preferred food, obtain preferred shelter, etc.). Actions of
that serve to limit the environment for conspecifics—
e.g., eating, excreting wastes, using up non-food
resources, taking up space, defending territories—are
those that determine carrying capacity. They are referred
to as density dependent because the greater the density of
the population, the greater their effects. Density-
dependent factors may exert their effect by reducing
birth rates, increasing death rates, extending generation
times, or by forcing the migration of conspecifics to new
regions.
“The impact of disease on a population can be density
dependent if the transmission rate of the disease depends
on a certain level of crowding in the population.” “A
death rate that rises as population density rises is said to
be density dependent, as is a birth rate that falls with
rising density. Density-dependent rates are an example
of negative feedback. In contrast, a birth rate or death
rate that does not change with population density is said
to be density independent… Negative feedback prevents
unlimited population growth.”
Predation can also be density dependent since predators
often can switch prey preferences to match whatever
prey organisms are most plentiful in a given
environment. “Many predators, for example, exhibit
switching behavior: They begin to concentrate on a
particularly common species of prey when it becomes
energetically efficient to do so.
Density-independent factors: Density-independent
effects on population sizes (or structures) occur to the
same extent regardless of population size. These can be
things like sudden changes in the weather.
"Over the long term, many populations remain fairly
stable in size and are presumably close to a carrying
capacity that is determined by density-dependent factors.
Superimposed on this general stability, however, are
short-term fluctuations due to density-independent
factors."
UNIT 10: ECOLOGICAL PRINCIPLES 

C2. Life history Adaptive Strategies of Development (r and K selection) 
 
Logistic  Law:  It  can  be  said  that,  overall,  natural  fluctuating numbers   
selection,  which  enables  the  colonization  of   
ecosystems,  has  put  into  place  two  major  types  of  1.  The  r­strategists:  The  r‐strategists  are  species 
species  (see  Table).  Even  though  there  are  several  with a very high coefficient of growth. They are thus 
models for growth of individuals in a population, the  located near the base of the curve. Among plants, the 
most  commonly  recognized  is  that  of  logistic  law  of  grasses  or  members  of  family  Poaceae  are  typical  r‐
development of populations (Fig.):  strategists. They are annual plants that spend 70% of 
dN (K − N ) their  energy  on  seed  production.  With  their  phe‐
= rN   nomenal  capacity  for  dissemination  and  their  low 
dt K needs,  they  have  a  particular  aptitude  for  colonizing 
spaces that are devoid of all other vegetation. 
Where,   N = number of individuals or biomass at an   
instant t;   Among animals, a small sparrow, the crossbill, is also 
               r = coefficient (intrinsic rate) of growth; and   representative of this group. It feeds on conifer seeds 
K = maximum number of individuals the biotope can  and  in  the  spring  produces  two  broods  of  6  to  8 
hold as a function of its resources.  young each, which are capable of reproducing in that 
same  autumn.  The  crossbill  lives  only  two  years  on 
average  but  each  pair  can  in  good  conditions  have 
about 40 offspring. 
 
Characteristics 
 
ƒ Size of individuals: The r‐strategists are small, and 
there  seems  to  be  an  explanation  for  their  size.  For 
example,  a  spherical,  unicellular  alga  has  a  volume 
(cubic function) corresponding to the classic formula 
V  =  4/3  πr3  The  zone  of  exchange  with  the  medium, 
for  photosynthesis,  respiration,  or  absorption,  is  the 
cell  membrane,  the  surface  of  which  is  a  square 
function, since it follows the formula S =πr2. When the 
size  of  the  organism  increases,  the  shape  being 
constant,  the  SN  ratio  diminishes.  Its  surface  of 
exchanges grows proportionately less rapidly than its 
  volume. Its metabolic activity is affected to the extent 
The curve always has the same shape, but, the greater  that  it  absorbs  nutrients  less  quickly.  Transfers  of 
the value of coefficient r, the more abruptly the curve  gases and nutrients are not as efficient as required by 
rises.  On  the  other  hand,  when  N  approaches  K,  the  the increase in volume. The r‐strategists, which have 
value (N ‐ K) is close to 0, the product rN(O) =‐0, the  an  active  metabolism,  thus  develop  well  when  they 
growth is thus null, the environment is saturated, and  remain small. 
its  potential  does  not  allow  it  to  advantageously   
support any more individuals.  ƒ  Productivity:  Productivity  is  very  high.  The 
Table:  comparative  table  of  r­strategist  and  k­ species that follow the r‐strategy, not requiring much 
strategist  energy,  establish  themselves  right  away  and  grow 
              r­strategist                k­strategist  rapidly. Such is the case with 'weeds' that invade any 
ƒ Small size  ƒ Large size  abandoned field. 
ƒ High  ƒ Low productivity   
productivity  ƒ Long  period  of  sexual  ƒ  Prolificity: The r‐strategists have a high prolificity 
ƒ Sexual precocity  immaturity  because  of  their  high  sexual  precocity  and  the  large. 
ƒ High mortality  ƒ Low mortality  number  of  seeds  or  descendants  they  produce.  This 
ƒ Short life span  ƒ Long life span  high  prolificity  is  accompanied  by  enormous 
ƒ Considerable  ƒ Economical  use  of  expenditure  of  energy,  much  of  it  purely  loss,  which 
waste of energy  energy  leads to considerable waste. 
ƒ 'Generalist'  ƒ 'Specialist' species   
species  ƒ Population  density  ƒ  Life span: Mortality is very high and the enormous 
ƒ Population  closely  dependent  on  expenditures  devoted  to  reproduction  that 
density  independent  of  environmental variations  characterize r‐strategists are accompanied  by a  very 
environmental variations  ƒ Demographic  short life span. There are many descendants, but only 
ƒ Demographic  regulation  linked  to  some  survive  and  become  adults  capable  of. 
regulation  not  linked  to  population density  reproduction. 
population density  ƒ Species  closely   
ƒ Pioneering  and  associated with climax   ƒ  Mode  of  life:  The  r‐strategists  can  only  satisfy 
colonizing species  ƒ Low  fluctuation  in  their energy needs by having a wide range of feed. In 
ƒ Highly  numbers 

26 INSTITUTE FOR ADVANCED STUDIES, JODHPUR 
UNIT 10: ECOLOGICAL PRINCIPLES 

abiotic as well as biotic terms, they have a generalist  ƒ Population density: The number of descendants is 
mode of life.  highly  dependent  on  the  potential  of  the 
  environment.  Population  density  will  therefore  vary 
ƒ  Population  density:  Because  of  their  wide  considerably  according  to  the  biotope.  The 
tolerance,  these  ubiquitous  species  are  independent  demographic regulation is thus here totally linked to 
of environmental fluctuations, which affect them only  the demography. 
slightly.  Similarly,  since  they  act  as  pioneer  species   
that often colonize new or virgin environments, their  If  the  environmental  conditions  remain  stable,  fluc‐
demographic  regulation  is  not  dependent  on  the  tuations in numbers will remain limited. 
density  of  their  population.  Also,  their  numbers   
fluctuate  greatly,  as  appropriate  for  colonizing  3.  Why  Do  the  Less  Competitive  r­strategists  not 
species.  Disappear? 
   
Since  nature  always  optimizes  the  flows  of  matter  All ecosystems are not 'stable'. A single cataclysm can 
and energy, these wasteful species are quite soon re‐ disturb  the  K‐strategists,  which,  like  all  'specialists', 
placed  by  others  that  are  more  economical  and  are  effective  only  when  the  environment  is  stable. 
efficient and that have K‐ strategies.  The least disturbance is a catastrophe for them. They 
  are  thus  very  fragile  and  sensitive  to  the  vagaries  of 
2.  The  K­strategists:  Unlike  the  r‐strategists,  the  K‐ nature.  And  nature  never  ceases  to  produce  climatic 
strategists are species that are highly demanding but  accidents  (flood,  drought,  frost)  or  geological 
well adapted and profiting fully from the potential of  accidents  (earthquakes,  landslides,  erosion),  to  cite 
their  environments.  Under  the  logistic  law,  only  the  major  disturbances.  Just  as  an  economic 
environments favor ‘K’ over ‘r’. The K‐strategists are  disaster  rapidly  creates  an  army  of  unemployed,  an 
at the top of the curve that characterizes equilibrium  ecological  disturbance  leads  to  the  disappearance  of 
between  the  population  and  the  possibilities  of  the  K‐strategists. 
biotope. In plants, the type example is the oak, which   
lives for several centuries but fructifies quite slowly.  Their place does not long remain unoccupied, since r‐
This  tree  spends  only  7%  of  its  energy  in  producing  strategists, which are tolerant of various living condi‐
acorns.  Among  animals,  the  large  albatross  of  the  tions,  adapt  themselves  to  these  disturbances  and 
Australian seas attains puberty only at the age of 10.  populate  the  ecosystem  until  it  becomes  stable  and 
The  female  lays  a  single  egg  every  two  years,  but  as  perhaps  newly  occupied  by  efficient  K‐strategists. 
the individuals live an average of 80 years, this gives  When  lightning  strikes  an  old  tree  in  a  primeval 
a  potential  of  40  descendants  (the  same  number  as  forest,  it  creates  a  clearing  that  provides  favourable 
for the crossbill).  conditions for the growth of raspberry bushes. These 
  bushes  were  stunted  in  the  shade  of  the  large  tree 
Characteristics  and regain vitality when the tree is gone. 
   
ƒ Size  of  individuals:  The  K‐strategists  are  always  The r‐strategists thus prove to be valuable in case of 
large. All the same, at the same trophic level,  a large  disruptions.  They  are  pioneering  and  colonizing 
bacterium  behaving  as  a  K‐strategist  will  always  be  species, but they never attain the K threshold because 
smaller than a tiny r‐strategist mammal.  they  find  themselves  competing  with  the  more 
  competitive  K‐strategists  when  the  environment 
ƒ Productivity: Since they are very precisely adaptea  stabilizes. There is in fact a gradient of evolution with 
to  their  environmental  conditions,  the  K‐strategists  intermediate strategies. 
by themselves have a low productivity. But since, like   
other  species,  they  exploit  other  trophic  sectors,  the  Note: The human species is not free of this law. In the 
biotope as a whole‐is used and the total productivity  northern  or  'rich'  countries,  the  environment  is 
is optimal.  nearly stable for the large majority of the population. 
  Food is abundant and nutritious, health is monitored, 
ƒ Prolificity: The K‐strategists have a low prolificity.  conditions of life convenient (with heated houses, for 
They have in fact a long period of sexual immaturity  example).  The  life  span  is  increasingly  long  and  the 
and the size of litters or broods is limited.  average  family  has  two  children  who  are  well 
  educated  and  well  cared  for.  These  criteria 
ƒ Life span: Their mortality is very low. The animals  correspond  to  those  that  have  been  defined  for  K‐
raise  only  a  few  young,  but  they  give  them  much  strategists. 
attention.  They  teach  the  young  to  feed,  hunt,  and   
defend  themselves,  so  well  that  almost  all  become  It is very different in developing countries, where the 
adults. The life span is therefore long.   environment  is  unstable.  Drought,  poverty,  and 
  political,  religious,  or  ethnic  conflicts  keep  it  in  an 
ƒ Mode  of  life:  The  K‐strategists,  unlike  the  r‐ unstable  condition.  This  leads  the  human  species  to 
strategists, are economical with energy. They feed on  adopt r‐strategies. The population is high. Births are 
only  a  small  part  of  the  food  chain  but  waste  little.  numerous,  but  infant  mortality  is  excessive,  as  a 
One  can  call  them  specialist  species  that  are  highly  result  of  famine,  disease,  and  parasites  associated 
adapted  to  the  nutritional  and  ambient  abiotic  with lack of health care, poor living conditions, and a 
conditions.  lack of potable water. 
 

INSTITUTE FOR ADVANCED STUDIES, JODHPUR  27
UNIT 10: ECOLOGICAL PRINCIPLES 

 
C3. Concept of  Metapopulations 
 
A  metapopulation  consists  of  a  group  of  spatially  equation  suggested  that  the  relationship  between 
separated  populations  of  the  same  species  which  predators  and  their  prey  would  result  in  population 
interact at some level. The term metapopulation was  oscillations over time based on the initial densities of 
coined by Richard Levins in 1969 to describe a model  predator  and  prey.  Gause's  early  experiments  to 
of population dynamics of insect pests in agricultural  prove  the  predicted  oscillations  of  this  theory  failed 
fields, but the idea has been most broadly applied to  because  the  predator‐prey  interactions  were  not 
species  in  naturally  or  artificially  fragmented  influenced  by  immigration.  However,  once 
habitats.  In  Levins'  own  words,  it  consists  of  "a  immigration  was  introduced,  the  population  cycles 
population of populations".   accurately  depicted  the  oscillations  predicted  by  the 
  Lotka‐Volterra  equation,  with  the  peaks  in  prey 
A  metapopulation  is  generally  considered  to  consist  abundance  shifted  slightly  to  the  left  of  the  peaks  of 
of several distinct populations together with areas of  the  predator  densities.  Huffaker's  experiments 
suitable  habitat  which  are  currently  unoccupied.  In  expanded  on  those of  Gause  by  examining  how  both 
classical  metapopulation  theory,  each  population  the  factors  of  migration  and  spatial  heterogeneity 
cycles  in  relative  independence  of  the  other  lead to predator‐prey oscillations. 
populations  and  eventually  goes  extinct  as  a   
consequence  of  demographic  stochasticity  Huffaker's  experiments  on  predator­prey 
(fluctuations  in  population  size  due  to  random  interactions (1958): 
demographic events); the smaller the population, the   
more prone it is to extinction.  In  order  to  study  predation  and  population 
  oscillations,  Huffaker  used  mite  species,  one  being 
Although  individual  populations  have  finite  life‐ the predator and the other being the prey. He set up a 
spans, the metapopulation  as  a whole is often  stable  controlled experiment using oranges, which the prey 
because  immigrants  from  one  population  (which  fed on, as the spatially structured habitat in which the 
may,  for  example,  be  experiencing  a  population  predator  and  prey  would  interact.  At  first,  Huffaker 
boom)  are  likely  to  re‐colonize  habitat  which  has  experienced  difficulties  similar  to  those  of  Gause  in 
been  left  open  by  the  extinction  of  another  creating  a  stable  predator‐prey  interaction.  By  using 
population.  They  may  also  emigrate  to  a  small  oranges  only,  the  prey  species  quickly  went  extinct 
population  and  rescue  that  population  from  followed  consequently  with  predator  extinction. 
extinction (called the rescue effect).  However, he discovered that by modifying the spatial 
  structure  of  the  habitat,  he  could  manipulate  the 
The  development  of  metapopulation  theory,  in  population  dynamics  and  allow  the  overall  survival 
conjunction  with  the  development  of  source‐sink  rate  for  both  species  to  increase.  He  did  this  by 
dynamics, emphasised the importance of connectivity  altering  the  distance  between  the  prey  and  oranges 
between seemingly isolated populations. Although no  (their  food),  establishing  barriers  to  predator 
single population may be able to guarantee the long‐ movement,  and  creating  corridors  for  the  prey  to 
term survival of a given species, the combined effect  disperse. These changes resulted in increased habitat 
of many populations may be able to do this.  patches and in turn provided more areas for the prey 
  to  seek  temporary  protection.  When  the  prey  would 
Metapopulation  theory  was  first  developed  for  go extinct locally at one habitat patch, they were able 
terrestrial  ecosystems,  and  subsequently  applied  to  to  reestablish  by  migrating  to  new  patches  before 
the  marine  realm  (Kritzer  &  Sale  2006).  In  fisheries  being  attacked  by  predators.  This  habitat  spatial 
science,  the  term  "sub‐population"  is  equivalent  to  structure of patches allowed for coexistence between 
the  metapopulation  science  term  "local  population".  the predator and prey species and promoted a stable 
Most  marine  examples  are  provided  by  relatively  population  oscillation  model.  Although  the  term 
sedentary  species  occupying  discrete  patches  of  metapopulation  had  not  yet  been  coined,  the 
habitat, with both local recruitment and recruitment  environmental  factors  of  spatial  heterogeneity  and 
from  other  local  populations  in  the  larger  habitat  patchiness  would  later  describe  the 
metapopulation.  Kritzer  &  Sale  have  argued  against  conditions  of  a  metapopulation  relating  to  how 
strict  application  of  the  metapopulation  definitional  groups  of  spatially  separated  populations  of  species 
criteria that extinction risks to local populations must  interact  with  one  another.  Huffaker's  experiment  is 
be non‐negligible. The most important contributor to  significant  because  it  showed  how  metapopulations 
metapopulation  theory  is  the  Finnish  biologist,  Ilkka  can directly affect the predator‐prey interactions and 
Hanski, of the University of Helsinki.  in turn influence population dynamics. 
   
Predation and Oscillations  Aggregation  effect  and  stability  of  predator­prey 
  interactions 
The  first  experiments  with  predation  and  spatial   
heterogeneity  were  conducted  by  G.F.  Gause  in  the  The  stability  of  predator‐prey  interactions  in  patchy 
1930's,  based  on  the  Lotka‐Volterra  equation,  which  environments  is  reinforced  when  an  aggregation 
was  formulated  in  the  mid‐1920s,  but  no  further  effect  occurs.  Aggregation  effect  is  when  predators 
application  had  been  conducted.  The  Lotka‐Volterra  are  more  greatly  attracted  to  habitat  patches  with 

28 INSTITUTE FOR ADVANCED STUDIES, JODHPUR 
UNIT 10: ECOLOGICAL PRINCIPLES 

high  prey  densities,  therefore,  areas  with  low  prey   

densities are relieved from the pressure of predation  Stochastic patch occupancy models (SPOMs) 
and  are  able  to  generate  a  larger  population  and   
avoid extinction.  One major drawback of the Levins model is that it is 
  deterministic,  whereas  the  fundamental 
The Levins model  metapopulation  processes  are  stochastic. 
  Metapopulations  are  particularly  useful  when 
Levins'  original  model  applied  to  a  metapopulation  discussing  species  in  disturbed  habitats,  and  the 
distributed  over  many  patches  of  suitable  habitat  viability of their populations, i.e., how likely they are 
with  significantly  less  interaction  between  patches  to become extinct in a given time interval. The Levins 
than  within  a  patch.  Population  dynamics  within  a  model cannot address this issue. 
patch  were  simplified  to  the  point  where  only   
presence and absence were considered. Each patch in  For  conservation  biology  purposes,  metapopulation 
his model is either populated or not.  models  must  include  (a)  the  finite  nature  of 
  metapopulations (how many patches are suitable for 
Let  N  be  the  fraction  of  patches  occupied  at  a  given  habitat), and (b) the probabilistic nature of extinction 
time.  During  a  time  step,  each  occupied  patch  can  and  colonisation.  Also,  note  that  in  order  to  apply 
become unoccupied  with  an extinction probability  e.  these models, the extinctions and colonisations of the 
Additionally,  1  −  N  of  the  patches  are  unoccupied.  patches must be asynchronous. 
Each of these may become populated by colonization.   
Let  c  be  a  constant  rate  of  propagule  generation  for  Stochasticity and Metapopulations 
each  of  the  N  occupied  patches.  This  yields  a   
probability  of  cN  for  each  unoccupied  patch  to  be  Huffaker's  studies  of  spatial  structure  and  species 
colonized.  So  for  each  time  step,  the  change  in  the  interactions are an example of early experimentation 
proportion of occupied patches, dN/dt, is  in  metapopulation  dynamics.  Since  the  experiments 
of  Huffaker  and  Levins,  models  have  been  created 
  which integrate stochastic factors. These models have 
This  takes  on  a  sigmoid  shape  similar  to  the  logistic  proven  that  the  combination  of  environmental 
model.  The  equilibrium  value  of  N  can  be  calculated  variability  (stochascity)  and  relatively  small 
by setting dN to be equal to zero. Solving for N gives  migration  rates  cause  indefinite  or  unpredictable 
either N = 0 or  persistence.  However,  Huffaker's  experiment  almost 
guaranteed  infinite  persistence  because  of  the 
  controlled immigration variable. 
This  result,  that  N  is  always  less  than  one,  implies 
that some fraction of a species habitat will always be 
unoccupied. 
 
D. Species interactions:  
Types of interactions, Interspecific competition, Herbivory, Carnivory, Pollination, Symbiosis.  
 
Biological interactions result from the fact that ecosystems, species may affect each other through
organisms in an ecosystem interact with each other, in intermediaries such as shared resources or common
the natural world, no organism is an autonomous entity enemies.
isolated from its surroundings. It is part of its
environment, rich in living and non living elements all of
which interact with each other in some fashion. An
organism's interactions with its environment are
fundamental to the survival of that organism and the Interactions categorized by effect
functioning of the ecosystem as a whole. Sign-mediated
interactions in which molecules serve as signs are the Effect Type of interaction
characteristic feature of communicative interactions.
0 0 Neutralism
In ecology, biological interactions are the relationships - 0 Amensalism
between two species in an ecosystem. These + 0 Commensalism
relationships can be categorized into many different - - Competition
classes of interactions based either on the effects or on + + Mutualism
the mechanism of the interaction. The interactions + - Predation or Parasitism
between two species vary greatly in these aspects as well
as in duration and strength. Species may meet once in a Some types of relationships listed by the effect they have
generation (e.g. pollination) or live completely within on each partner. '0' is no effect, '-' is detrimental, and '+'
another (e.g. endosymbiosis). Effects may range from is beneficial.
one species eating the other (predation), to mutual
benefit (mutualism). Terms which explicitly indicate the quality of benefit or
harm in terms of fitness experienced by participants in
The interactions between two species need not be an interaction are listed below. There are six possible
through direct contact. Due to the connected nature of combinations, ranging from mutually beneficial through

INSTITUTE FOR ADVANCED STUDIES, JODHPUR  29
UNIT 10: ECOLOGICAL PRINCIPLES 
neutral to mutually harmful interactions. The level of
benefit or harm is continuous and not discrete, such that
an interaction may be trivially harmful through to
deadly, for example. It is important to note that these
interactions are not always static. In many cases, two
species will interact differently under different
conditions. This is particularly true in, but not limited to,
cases where species have multiple, drastically different
life stages.
Neutralism
Neutralism describes the relationship between two
species which do interact but do not affect each other. It
is to describe interactions where the fitness of one
species has absolutely no effect whatsoever on that of
other. True neutralism is extremely unlikely and
impossible to prove. When dealing with the complex
networks of interactions presented by ecosystems, one
cannot assert positively that there is absolutely no
competition between or benefit to either species. Since
true neutralism is rare or nonexistent, its usage is often
extended to situations where interactions are merely
insignificant or negligible.
Amensalism
Amensalism between two species involves one impeding
or restricting the success of the other without being
affected positively or negatively by the presence of the
other. It is a type of symbiosis. Usually this occurs when
one organism exudes a chemical compound as part of its
normal metabolism that is detrimental to another
organism.
The bread mold Penicillium is a common example of
this; penicillium secrete penicillin, a chemical that kills
bacteria. A second example is the black walnut tree
(Juglans nigra), which secrete juglone, a chemical that
harms or kills some species of neighboring plants, from
its roots. This interaction may still increase the fitness of
the non-harmed organism though, by removing
competition and allowing it access to greater scarce
resources. In this sense the impeding organism can be
said to be negatively affected by the other's very
existence, making it a +/- interaction.
Antibiosis or allelopathy also explain similar
interactions.
Competition
Competition is an interaction between individuals or
populations that is mutually detrimental.
Synnecrosis is a particular case in which the interaction
is so mutually detrimental that it results in death, as in
the case of some parasitic relationships. It is a rare and
necessarily short-lived condition as evolution selects
against it. The term is seldom used.
Antagonism
In antagonistic interactions one species benefits at the
expense of another. Predation is an interaction between
organisms in which one organism captures biomass from
another. It is often used as a synonym for carnivory but
in its widest definition includes all forms of one
30 INSTITUTE FOR ADVANCED STUDIES, JODHPUR
organism eating another, regardless of trophic level (e.g.
herbivory), closeness of association (e.g. parasitism and
parasitoidism) and harm done to prey (e.g. grazing).
Other interactions that cannot be classed as predation
however are still possible, such as Batesian mimicry,
where an organism bears a superficial similarity of at
least one sort, such as a harmless plant coming to mimic
a poisonous one.
Ecological facilitation
The following two interactions can be classed as
facilitative. Facilitation describes species interactions
that benefit at least one of the participants and cause no
harm to either. Facilitations can be categorized as
mutualisms, in which both species benefit, or
commensalisms, in which one species benefits and the
other is unaffected. Much of classic ecological theory
(e.g., natural selection, niche separation, metapopulation
dynamics) has focused on negative interactions such as
predation and competition, but positive interactions
(facilitation) are receiving increasing focus in ecological
research.
Commensalism
Commensalism benefits one organism and the other
organism is neither benefited nor harmed. It occurs when
one organism takes benefits by interacting with another
organism by which the host orgaism is not affected. A
good example is a remora living with a shark. Remoras
eat leftover food from the shark. The shark is not
affected in the process as remoras eat only leftover food
of the shark which doesn't deplete the sharks resources.
Mutualism
Mutualism is an interaction between two or more
species, where species derive a mutual benefit, for
example an increased carrying capacity. Similar
interactions within a species are known as co-operation.
Mutualism may be classified in terms of the closeness of
association, the closest being symbiosis, which is often
confused with mutualism. One or both species involved
in the interaction may be obligate, meaning they cannot
survive in the short or long term without the other
species. Though mutualism has historically received less
attention than other interactions such as predation, it is
very important subject in ecology. Examples include
cleaner fish, pollination and seed dispersal, gut flora and
nitrogen fixation by fungi.
Interactions classified by mechanism
Symbiosis
The term symbiosis (Greek: living together) can be used
to describe various degrees of close relationship between
organisms of different species. Sometimes it is used only
for cases where both organisms benefit, sometimes it is
used more generally to describe all varieties of relatively
tight relationships, i.e. even parasitism, but not
predation. Some even go so far as to use it to describe
predation. It can be used to describe relationships where
one organism lives on or in another, or it can be used to
describe cases where organisms are related by mutual
stereotypic behaviors.
 
UNIT 10: ECOLOGICAL PRINCIPLES 
In either case symbiosis is much more common in the
living world and much more important than is generally
assumed. Almost every organism has many internal
parasites. A large percentage of herbivores have
mutualistic gut fauna that help them digest plant matter,
which is more difficult to digest than animal prey. Coral
reefs are the result of mutalisms between coral
organisms and various types of algae that live inside
them. Most land plants and thus, one might say, the very
existence of land ecosystems rely on mutualisms
between the plants which fix carbon from the air, and
Mycorrhyzal fungi which help in extracting minerals
from the ground. In fact the evolution of all eukaryotes
(plants, animals, fungi, protists) is believed to have
resulted from a symbiosis between various sorts of
bacteria: endosymbiotic theory.
Competition
D2. Competition
Organisms grow, reproduce and die. They are affected
by the conditions in which they live, and by the
resources that they obtain. But no organism lives in
isolation. Each, for at least part of its life, is a member of
a population composed of individuals of its own species.
Individuals of the same species have very similar
requirements for survival, growth and reproduction; but
their combined demand for a resource may exceed the
immediate supply. The individuals then compete for the
resource and, not surprisingly, at least some of them
become deprived. This topic is concerned with the nature
of such intraspecific competition, its effects on the
competing individuals and on populations of competing
individuals.
We begin with a working definition: ‘competition is an
interaction between individuals, brought about by a
shared requirement for a resource, and leading to a
reduction in the survivorship, growth and/or
reproduction of at least some of the competing
individuals concerned’. We can now look more closely
at competition.
Competition can be defined as an interaction between
organisms or species, in which the fitness of one is
lowered by the presence of another. Limited supply of at
least one resource (such as food, water, and territory)
used by both is required. Competition both within and
between species is an important topic ecology, especially
in community ecology. Competition is one of many
interacting biotic and abiotic factors that affect
community structure. Competition among members of
the same species is known as intraspecific competition,
while competition between individuals of different
species is known as interspecific competition.
Competition is not always a straightforward, direct
interaction either, and can occur in both a direct and
indirect fashion.
Consider, initially, a simple hypothetical community: a
thriving population of grasshoppers (all of one species)
feeding on a field of grass (also of one species). To
INSTITUTE FOR ADVANCED STUDIES, JODHPUR  31
Competition can be defined as an interaction between
organisms or species, in which the fitness of one is
lowered by the presence of another. Limited supply of at
least one resource (such as food, water, and territory)
used by both is required.[12] Competition is one of
many interacting biotic and abiotic factors that affect
community structure. Competition among members of
the same species is known as intraspecific competition,
while competition between individuals of different
species is known as interspecific competition.
Competition is not always a straightforward, direct
interaction either, and can occur in both a direct and
indirect fashion.
According to the competitive exclusion principle,
species less suited to compete for resources should either
adapt or die out. According to evolutionary theory, this
competition within and between species for resources
plays a critical role in natural selection.
provide themselves with energy and material for growth
and reproduction, grasshoppers eat grass; but in order to
find and consume that grass they must use energy. Any
grasshopper might find itself at a spot where there is no
grass because some other grasshopper has eaten it.
The grasshopper must then move on and expend more
energy before it takes in food. The more grasshoppers
there are, the more often this will happen. An increased
energy expenditure and a decreased rate of food intake
may all decrease a grasshopper’s chances of survival,
and also leave less energy available for development and
reproduction. Survival and reproduction determine a
grasshopper’s contribution to the next generation.
Hence, the more intraspecific competitors for food a
grasshopper has, the less its likely contribution will be.
As far as the grass itself is concerned, an isolated
seedling in fertile soil may have a very high chance of
surviving to reproductive maturity. It will probably
exhibit an extensive amount of modular growth, and will
probably therefore eventually produce a large number of
seeds. However, a seedling that is closely surrounded by
neighbors (shading it with their leaves and depleting the
water and nutrients of its soil with their roots) will be
very unlikely to survive, and if it does, will almost
certainly form few modules and set few seeds.
We can see immediately that the ultimate effect of
competition on an individual is a decreased contribution
to the next generation compared with what would have
happened had there been no competitors. Intraspecific
competition typically leads to decreased rates of resource
intake per individual, and thus to decreased rates of
individual growth or development, or perhaps to
decreases in the amounts of stored reserves or to
increased risks of predation. These may lead, in turn, to
decreases in survivorship and/or decreases in fecundity,
which together determine an individual’s reproductive
output.
Exploitation and interference: In many cases,
competing individuals do not interact with one another
UNIT 10: ECOLOGICAL PRINCIPLES 
directly. Instead, individuals respond to the level of a
resource, which has been depressed by the presence and
activity of other individuals.
The grasshoppers were one example. Similarly, a
competing grass plant is adversely affected by the
presence of close neighbors, because the zone from
which it extracts resources (light, water, nutrients) has
been overlapped by the ‘resource depletion zones’ of
these neighbors, making it more difficult to extract those
resources. In such cases, competition may be described
as exploitation, in that each individual is affected by the
amount of resource that remains after that resource has
been exploited by others. Exploitation can only occur,
therefore, if the resource in question is in limited supply.
In many other cases, competition takes the form of
interference. Here individuals interact directly with each
other, and one individual will actually prevent another
from exploiting the resources within a portion of the
habitat. For instance, this is seen amongst animals that
defend territories and amongst the sessile animals and
plants that live on rocky shores. The presence of a
barnacle on a rock prevents any other barnacle from
occupying that same position, even though the supply of
food at that position may exceed the requirements of
several barnacles. In such cases, space can be seen as a
resource in limited supply.
Another type of interference competition occurs when,
for instance, two red deer stags fight for access to a
harem of hinds. Either stag, alone, could readily mate
with all the hinds, but they cannot both do so since
matings are limited to the ‘owner’ of the harem.
Thus, interference competition may occur for a resource
of real value (e.g. space on a rocky shore for a barnacle),
in which case the interference is accompanied by a
degree of exploitation, or for a surrogate resource (a
territory, or ownership of a harem), which is only
valuable because of the access it provides to a real
resource (food, or females). With exploitation, the
intensity of competition is closely linked to the level of
resource present and the level required, but with
interference, intensity may be high even when the level
of the real resource is not limiting.
Whether they compete through exploitation or
interference, individuals within a species have many
fundamental features in common, using similar resources
and reacting in much the same way to conditions. None
the less, intraspecific competition may be very one
sided: a strong, early seedling will shade a stunted, late
one; an older and larger bryozoan on the shore will grow
over a smaller and younger one. Hence, the ultimate
effect of competition is far from being the same for
every individual. Weak competitors may make only a
small contribution to the next generation, or no
contribution at all. Strong competitors may have their
contribution only negligibly affected.
Finally, note that the likely effect of intraspecific
competition on any individual is greater the more
competitors there are. The effects of intraspecific
competition are thus said to be density dependent.
According to the competitive exclusion principle,
species less suited to compete for resources should either
32 INSTITUTE FOR ADVANCED STUDIES, JODHPUR
adapt or die out. According to evolutionary theory, this
competition within and between species for resources
plays a critical role in natural selection.
Types of Competition
By mechanism: The following terms describe
mechanisms by which competition occurs, which can
generally be divided into direct and indirect. These
mechanisms apply equally to intraspecific and
interspecific competition.
Male-male competition in red deer during rut is an
example of interference competition within a species.
• Interference competition - occurs directly between
individuals via aggression etc. when the individuals
interfere with foraging, survival, reproduction of others,
or by directly preventing their physical establishment in
a portion of the habitat.
• Exploitation competition - occurs indirectly through
a common limiting resource which acts as an
intermediate. For example the use of the resource(s)
depletes the amount available to others, or they compete
for space.
• Apparent competition - occurs indirectly between
two species which are both preyed upon by the same
predator. For example, species A and species B are both
preys for predator C. The increase of species A will
cause the decrease of species B because the increase of
As would increase the number of predator Cs which in
turn will hunt more of species B.
Intraspecific competition: Intraspecific competition
occurs when members of the same species vie for the
same resources in an ecosystem (e.g. food or nutrients,
light or space). For example, two trees growing close
together will compete for light above ground, and water
and nutrients in the soil. Therefore, getting less
resources, they will usually perform less well than if
they grew by themselves. Adaptations to such an
environment include growing taller (especially in
forests), or developing a larger root system.
Interspecific competition: Interspecific competition
may occur when individuals of two separate species
share a limiting resource in the same area. If the resource
cannot support both populations, then lowered fecundity,
growth, or survival may result in at least one species.
Interspecific competition has the potential to alter
populations, communities and the evolution of
interacting species. An example among animals could be
the case of cheetahs and lions; since both species feed on
similar prey, they are negatively impacted by the
presence of the other because they will have less food. In
fact, lions sometimes steal prey items killed by cheetahs.
Evolutionary strategies: In evolutionary contexts,
competition is related to the concept of r/K selection
theory, which relates to the selection of traits which
promote success in particular environments. The theory
originates from work on island biogeography by the
ecologists Robert MacArthur and E. O. Wilson.
In r/K selection theory, selective pressures are
hypothesised to drive evolution in one of two
stereotyped directions: r- or K-selection. These terms, r
and K, are derived from standard ecological algebra, as
 
UNIT 10: ECOLOGICAL PRINCIPLES 
illustrated in the simple Verhulst equation of population
dynamics:
where r is the growth rate of the population (N), and K is
the carrying capacity of its local environmental setting.
Typically, r-selected species exploit empty niches, and
D3. Predation
In ecology, predation describes a biological interaction
where a predator organism feeds on another living
organism or organisms known as prey. Predators may or
may not kill their prey prior to feeding on them, but the
act of predation always results in the (ecologically
significant) death of the prey. The other main category
of consumption is detritivory, the consumption of dead
organic material (detritus). It can at times be difficult to
separate the two feeding behaviors, for example where
parasitic species prey on a host organism and then lay
their eggs on it for their offspring to feed on its decaying
corpse. The key characteristic of predation however is
the predator's direct impact on the prey population. On
the other hand, detritivores simply eat what is available
and have no direct impact on the "donor" organism(s).
Classification of predators
The unifying theme in all classifications of predation is
the predator lowering the fitness of its prey, or put
another way, it reduces its prey's chances of survival,
reproduction, or both. Ways of classifying predation
surveyed here include grouping by trophic level or diet,
by specialization, and by the nature of their interaction
with prey.
Functional classification: Classification of predators by
the extent to which they feed on and interact with their
prey is one way ecologists may wish to categorize the
different types of predation. Instead of focusing on what
they eat, this system classifies predators by the way in
which they eat, and the general nature of the interaction
between predator and prey species. Two factors are
considered here: How close the predator and prey are
physically (in the latter two cases the term prey may be
replaced with host). Additionally, whether or not the
prey are directly killed by the predator is considered,
with the first and last cases involving certain death.
True predation: A true predator is one which kills and
eats another organism. Whereas other types of predator
all harm their prey in some way, this form results in their
instant death. Predators may hunt actively for prey, or sit
and wait for prey to approach within striking distance, as
in ambush predators. Some predators kill large prey and
dismember or chew it prior to eating it, such as a jaguar,
while others may eat their (usually much smaller) prey
whole, as does a bottlenose dolphin or any snake, or a
duck or stork swallowing a frog. In some cases the prey
organism may die in the mouth or digestive system of
the predator. Baleen whales, for example, eat millions of
microscopic plankton at once, the prey being broken
down well after entering the whale. Seed predation is
another form of true predation, as seeds represent
potential organisms. Predators of this classification need
not eat prey entirely, for example some predators cannot
digest bones, while others can. Some may merely eat
INSTITUTE FOR ADVANCED STUDIES, JODHPUR  33
produce many offspring, each of whom has a relatively
low probability of surviving to adulthood. In contrast, K-
selected species are strong competitors in crowded
niches, and invest more heavily in much fewer offspring,
each of whom has a relatively high probability of
surviving to adulthood.
only part of an organism, as in grazing, but still
consistently cause its direct death.
Grazing: Grazing organisms may also kill their prey
species, but this is seldom the case. While some
herbivores like zooplankton live on unicellular
phytoplankton and have no choice but to kill their prey,
many only eat a small part of the plant. Grazing
livestock may pull some grass out at the roots, but most
is simply grazed upon, allowing the plant to regrow once
again. Kelp is frequently grazed in subtidal kelp forests,
but regrows at the base of the blade continuously to cope
with browsing pressure. Animals may also be 'grazed'
upon; female mosquitos land on hosts briefly to gain
sufficient proteins for the development of their offspring.
Starfish may be grazed on, being capable of regenerating
lost arms.
Parasitism: Parasites can at times be difficult to
distinguish from grazers. Their feeding behavior is
similar in many ways, however they are noted for their
close association with their host species. While a grazing
species such as an elephant may travel many kilometers
in a single day, grazing on many plants in the process,
parasites form very close associations with their hosts,
usually having only one or at most a few in their
lifetime. This close living arrangement may be described
by the term symbiosis, 'living together,' but unlike
mutualism the association significantly reduces the
fitness of the host. Parasitic organisms range from the
macroscopic mistletoe, a parasitic plant, to microscopic
internal parasites such as cholera. Some species however
have more loose associations with their hosts.
Lepidoptera (butterfly and moth) larvae may feed
parasitically on only a single plant, or they may graze on
several nearby plants. It is therefore wise to treat this
classification system as a continuum rather than four
isolated forms.
Parasitoidism: Parasitoids are organisms living in or on
their host and feeding directly upon it, eventually leading
to its death. They are much like parasites in their close
symbiotic relationship with their host or hosts. Like the
previous two classifications parasitoid predators do not
kill their hosts instantly. However, unlike parasites, they
are very similar to true predators in that the fate of their
prey is quite inevitably death. A well known example of
a parasitoids are the ichneumon wasps, solitary insects
living a free life as an adult, then laying eggs on or in
another species such as a caterpillar. Its larva(e) feed on
the growing host causing it little harm at first, but soon
devouring the internal organs until finally destroying the
nervous system resulting in prey death. By this stage the
young wasp(s) are developed sufficiently to move to the
next stage in their life cycle. Though limited mainly to
the insect order Hymenoptera, parasitoids make up as
much as 10% of all insect species.
UNIT 10: ECOLOGICAL PRINCIPLES 
Degree of specialization: Among predators there is a
large degree of specialization. Many predators specialize
in hunting only one species of prey. Others are more
opportunistic and will kill and eat almost anything
(examples: humans, leopards, and dogs). The specialists
are usually particularly well suited to capturing their
preferred prey. The prey in turn, are often equally suited
to escape that predator. This is called an evolutionary
arms race and tends to keep the populations of both
species in equilibrium. Some predators specialize in
certain classes of prey, not just single species. Almost all
will switch to other prey (with varying degrees of
success) when the preferred target is extremely scarce,
and they may also resort to scavenging or a herbivorous
diet if possible.
Trophic level: Predators are often another organism's
prey, and likewise prey are often predators. Though blue
jays prey on insects, they may in turn be prey for cats
and snakes, which may themselves be the prey of hawks.
One way of classifying predators is by trophic level.
Organisms which feed on autotrophs, the producers of
the trophic pyramid, are known as herbivores or primary
consumers; those that feed on heterotrophs such as
animals are known as secondary consumers. Secondary
consumers are a type of carnivore, but there are also
tertiary consumers eating these carnivores, quartary
consumers eating them, and so forth. Because only a
fraction of energy is passed on to the next level, this
hierarchy of predation must end somewhere, and very
seldom goes higher than five or six levels, and may go
only as high as three trophic levels (for example, a lion
that preys upon large herbivores such as wildebeest
which in turn eat grasses). A predator at the top of any
food chain (that is, one that is preyed upon by no
organism) is called an apex predator; examples include
the orca, tiger, and crocodile and even omnivorous
humans and American black bears. An apex predator in
one environment may not retain this position if
introduced to another habitat, such as dogs among
alligators.
Many organisms (of which humans are prime examples)
eat from multiple levels of the food chain and thus make
this classification problematic. A carnivore may eat both
secondary and tertiary consumers, and its prey may itself
be difficult to classify for similar reasons. Organisms
showing both carnivory and herbivory are known as
omnivores. Even supposedly strict herbivores may
supplement their diet with meat. Carnivorous plants
would be very difficult to fit into this classification,
producing their own food but also digesting anything
that they may trap. Organisms which eat detritivores or
parasites would also be difficult to classify by such a
scheme.
Predation as competition: An alternative view offered
by Richard Dawkins is of predation as a form of
competition: the genes of both the predator and prey are
competing for the body (or 'survival machine') of the
prey organism. This is best understood in the context of
the gene centered view of evolution.
Ecological role
34 INSTITUTE FOR ADVANCED STUDIES, JODHPUR
Predators may increase the biodiversity of communities
by preventing a single species from becoming dominant.
Such predators are known as keystone species and may
have a profound influence on the balance of organisms
in a particular ecosystem. Introduction or removal of this
predator, or changes in its population density, can have
drastic cascading effects on the equilibrium of many
other populations in the ecosystem. For example, grazers
of a grassland may prevent a single dominant species
from taking over.
The elimination of wolves from Yellowstone National
Park had profound impacts on the trophic pyramid.
Without predation, herbivores began to over-graze many
woody species, affecting the area's plant populations.
Additionally, wolves often kept animals from grazing in
riparian areas, which protected beavers from having their
food sources encroached upon. The removal of wolves
had a direct effect on beaver populations, as their habitat
became territory for grazing.
Adaptations and behavior
The act of predation can be broken down into a
maximum of four stages: Detection of prey, attack,
capture and finally consumption. The relationship
between predator and prey is one which is typically
beneficial to the predator, and detrimental to the prey
species. Sometimes, however, predation has indirect
benefits to the prey species,[9] though the individuals
preyed upon themselves do not benefit. This means that,
at each applicable stage, predator and prey species are in
an evolutionary arms race to maximize their respective
abilities to obtain food or avoid being eaten. This
interaction has resulted in a vast array of adaptations in
both groups.
General: One adaptation helping both predators and
prey avoid detection is camouflage, a form of crypsis
where species have an appearance which helps them
blend into the background. Camouflage consists of not
only color, but also shape and pattern. The background
upon which the organism is seen can be both its
environment (e.g. the praying mantis to the right
resembling dead leaves) other organisms (e.g. zebras'
stripes blend in with each other in a herd, making it
difficult for lions to focus on a single target). The more
convincing camouflage is, the more likely it is that the
organism will go unseen.
Mimicry is a related phenomenon where an organism
has a similar appearance to another species. One such
example is the drone fly, which looks a lot like a bee, yet
is completely harmless as it cannot sting at all. Another
example of batesian mimicry is the io moth, (Automeris
io), which has markings on its wings which resemble an
owl's eyes. When an insectivorous predator disturbs the
moth, it reveals its hind wings, temporarily startling the
predator and giving it time to escape. Predators may also
use mimicry to lure their prey, however. Female fireflies
of the genus Photuris, for example, copy the light signals
of other species, thereby attracting male fireflies which
are then captured and eaten.
Predator: While successful predation results in a gain of
energy, hunting invariably involves energetic costs as
well. When hunger is not an issue, most predators will
 
UNIT 10: ECOLOGICAL PRINCIPLES 
generally not seek to attack prey since the costs
outweigh the benefits. For instance, a large predatory
fish like a shark that is well fed in an aquarium will
typically ignore the smaller fish swimming around it
(while the prey fish take advantage of the fact that the
apex predator is apparently uninterested). Surplus killing
represents a deviation from this type of behaviour. The
treatment of consumption in terms of cost-benefit
analysis is known as optimal foraging theory, and has
been quite successful in the study of animal behavior.
Costs and benefits are generally considered in energy
gain per unit time, though other factors are also
important, such as essential nutrients that have no caloric
value but are necessary for survival and health.
Size-selective predation involves predators preferring
prey of a certain size. Large prey may prove troublesome
for a predator, while small prey might prove hard to find
and in any case provide less of a reward. This has led to
a correlation between the size of predators and their
prey. Size may also act as a refuge for large prey, for
example adult elephants are generally safe from
predation by lions, but juveniles are vulnerable.
It has been observed that well-fed predator animals in a
lax captivity (for instance, pet or farm animals) will
usually differentiate between putative prey animals who
are familiar co-inhabitants in the same human area from
wild ones outside the area. This interaction can range
from peaceful coexistence to close companionship;
motivation to ignore the predatory instinct may result
from mutual advantage or fear of reprisal from human
masters who have made clear that harming co-
inhabitants will not be tolerated. Pet cats and pet mice,
for example, may live together in the same human
residence without incident as companions. Pet cats and
pet dogs under human mastership often depend on each
other for warmth, companionship, and even protection,
particularly in rural areas.
Anti-predator adaptations: Antipredator adaptations
have evolved in prey populations due to the selective
pressures of predation over long periods of time.
Aggression: Animals themselves predatory often use
their usual methods of attacking prey (often claws and
teeth) to inflict or to threaten to inflict grievous injury
upon potential predators. Such makes a rattlesnake or a
badger a less-than-desirable prey item. Many prey
creatures not themselves predatory, such as a zebra, can
give one of its usual predators (such as a lion or a
spotted hyena) a strong kick that can break the jaw of its
enemy and cause eventual starvation.
Mobbing behavior: Mobbing behavior occurs when a
species turns the tables on their predator by
cooperatively attacking or harassing it. This is most
frequently seen in birds, though it is also known to occur
in other social animals. For example, nesting gull
colonies are widely seen to attack intruders, including
humans. Costs of mobbing behavior include the risk of
engaging with predators, as well as energy expended in
the process; mockingbirds can effectively force a cat or
dog to seek something less troublesome. One
mockingbird might fly in front of the cat or dog, enticing
it to lunge, while another pecks at the cat or dog from
behind. While mobbing has evolved independently in
INSTITUTE FOR ADVANCED STUDIES, JODHPUR  35
many species, it only tends to be present in those whose
young are frequently preyed on, especially birds. It may
complement cryptic behavior in the offspring
themselves, such as camouflage and hiding. Mobbing
calls may be made prior to or during engagement in
harassment.
Mobbing behavior has functions beyond driving the
predator away. Mobbing draws attention to the predator,
making stealth attacks impossible. Mobbing also plays a
critical role in the identification of predators and inter-
generational learning about predator identification.
Reintroduction of species is often unsuccessful because
the established population lacks this cultural knowledge
of how to identify local predators. Scientists are
exploring ways to train populations to identify and
respond to predators before releasing them into the wild.
Mobbing can be an interspecies activity: it is common
for birds to respond to mobbing calls of a different
species. Many birds will show up at the sight of mobbing
and watch and call, but not participate. It should also be
noted that some species can be on both ends of a
mobbing attack. Crows are frequently mobbed by
smaller songbirds as they prey on eggs and young from
these birds' nests, but these same crows will cooperate
with smaller birds to drive away hawks or larger
mammalian predators. On occasion, birds will mob
animals that pose no threat.
Black-headed Gulls are one species which aggressively
engages intruding predators, such as Carrion Crows.
Experiments on this species by Hans Kruuk involved
placing hen eggs at intervals from a nesting colony, and
recording the percentage of successful predation events
as well as the probability of the crow being subjected to
mobbing. The results showed decreasing mobbing with
increased distance from the nest, which was correlated
with increased predation success. Mobbing may function
by reducing the predator's ability to locate nests, as
predators cannot focus on locating eggs while they are
under direct attack.
Advertising unprofitability: Once a predator has
detected its prey, one would expect it to pursue it.
However, it is not always profitable for the predator to
do so. Consider the example of a Thomson's Gazelle
being spotted by a predator. Giving chase to prey
requires a sacrifice in energy. If, however, there is some
way the prey species can convey the information that it
is unprofitable, energy will be saved by both organisms.
Thomson's Gazelles are hunted by species such as lions
and cheetahs. When they see the predator approach, they
may start to run away, but then slow down and stot.
Stotting describes a behavior involving jumping into the
air with the legs kept straight and stiff, and the white rear
fully visible. Obviously this behavior is maladaptive if
they hope to outrun the predator, so it must serve some
other purpose.
Although other hypotheses have been put forward,
evidence supports the proposition that they stot to signal
an unprofitable chase. For example, cheetahs abandon
more hunts when the gazelle stots, and in the event they
do give chase, they are far less likely to make a kill.
UNIT 10: ECOLOGICAL PRINCIPLES 
Aposematism, where organisms are brightly colored as a
warning to predators, is the antithesis of camouflage.
Some organisms pose a threat to their predators - for
example they may be poisonous, or able to harm them
physically. Aposematic coloring involves bright, easily
recognizable and unique colors and patterns. Upon being
harmed (e.g. stung) by their prey, the appearance in such
an organism will be remembered as something to avoid.
Population dynamics
It is fairly clear that predators tend to lower the survival
and fecundity of their prey, but on a higher level of
organization, populations of predator and prey species
also interact. It is obvious that predators depend on prey
for survival, and this is reflected in predator populations
being affected by changes in prey populations. It is not
so obvious, however, that predators affect prey
populations. Eating a prey organism may simply make
room for another if the prey population is approaching
its carrying capacity.
The population dynamics of predator-prey interactions
can be modelled using the Lotka–Volterra equations.
These provide a mathematical model for the cycling of
predator and prey populations.
Evolution of predation
Predation appears to have become a major selection
pressure shortly before the Cambrian period - around
550 million years ago - as evidenced by the almost
simultaneous development of calcification in animals
and algae, and predation-avoiding burrowing. However,
predators had been grazing on micro-organisms since at
least 1,000 million years ago.
Humans and predation
In conservation: Predators are an important
consideration in matters relating to conservation.
D4. Symbiosis 
The term symbiosis (from the Greek: σύν syn "with";
and βίωσις biosis "living") commonly describes close
and often long-term interactions between different
biological species. The term was first used in 1879 by
the German mycologist, Heinrich Anton de Bary, who
defined it as: "the living together of unlike organisms".
The definition of symbiosis is in flux and the term has
been applied to a wide range of biological interactions.
The symbiotic relationship may be categorized as being
mutualistic, parasitic, or commensal in nature. Others
define it more narrowly, as only those relationships from
which both organisms benefit, in which case it would be
synonymous with mutualism.
Symbiotic relationships included those associations in
which one organisms lives on another (ectosymbiosis,
such as mistletoe), or where one partner lives inside
another (endosymbiosis, such as lactobacilli and other
bacteria in humans or zooxanthelles in corals).
Symbiotic relationships may be either obligate, i.e.,
necessary to the survival of at least one of the organisms
36 INSTITUTE FOR ADVANCED STUDIES, JODHPUR
Introduced predators may prove too much for
populations which have not coevolved with them,
leading to possible extinction. This will depend largely
on how well the prey species can adapt to the new
species, and whether or not the predator can turn to
alternative food sources when prey populations fall to
minimal levels. If a predator can use an alternative prey
instead, it may shift its diet towards that species in a
behavior known as functional response, while still eating
the last remaining prey organisms. On the other hand the
prey species may be able to survive if the predator has
no alternative prey - in this case its population will
necessarily crash following the decline in prey, allowing
some small proportion of prey to survive. Introduction of
an alternative prey may well lead to the extinction of
prey, as this constraint is removed.
Predators are often the species endangered themselves,
especially apex predators who are often in competition
with humans. Competition for prey from other species
could prove the end of a predator - if their ecological
niche overlaps completely with that of another the
competitive exclusion principle requires only one can
survive. Loss of prey species may lead to coextinction of
their predator. In addition, because predators are found
in higher trophic levels, they are less abundant and much
more vulnerable to extinction.
Biological pest control: Predators may be put to use in
conservation efforts to control introduced species.
Although the aim in this situation is to remove the
introduced species entirely, keeping its abundance down
is often the only possibility. Predators from its natural
range may be introduced to control populations, though
in some cases this has little effect, and may even cause
unforeseen problems. Besides their use in conservation
biology, predators are also important for controlling
pests in agriculture. Natural predators are an
environmentally friendly and sustainable way of
reducing damage to crops, and are one alternative to the
use of chemical agents such as pesticides.
involved, or facultative, where the relationship is
beneficial but not essential to survival of the organisms.
Physical interaction
Endosymbiosis is any symbiotic relationship in which
the symbiote lives within the tissues of the host; either in
the intracellular space or extracellularly. Examples are
nitrogen-fixing bacteria (called rhizobia) which live in
root nodules on legume roots, Actinomycete nitrogen-
bacteria called Frankia which live in Alder tree root
nodules, single-celled algae inside reef-building corals,
and bacterial endosymbionts that provide essential
nutrients to about 10%–15% of insects.
Ectosymbiosis, also referred to as exosymbiosis, is any
symbiotic relationship in which the symbiont lives on
the body surface of the host, including the inner surface
of the digestive tract or the ducts of exocrine glands.
Examples of this include ectoparasites such as lice,
commensal ectosymbionts, such as the barnacles that
attach themselves to the jaw of baleen whales, and
mutualist ectosymbionts such as cleaner fish.
 
UNIT 10: ECOLOGICAL PRINCIPLES 
Mutualism
Mutualism is a biological interaction between
individuals of two different species, where both
individuals derive a fitness benefit, for example
increased survivorship. Similar interactions within a
species are known as co-operation. It can be contrasted
with interspecific competition, in which both species
experience reduced fitness, and exploitation, in which
one species benefits at the expense of the other.
Mutualism and symbiosis are sometimes used as if they
are synonymous, but this is strictly incorrect: symbiosis
is a broad category, defined to include relationships
which are mutualistic, parasitic or commensal.
Mutualism is therefore only one type. Mutualism plays a
key part in ecology and evolutionary biology. For
example, mutualistic interactions are vital for terrestrial
ecosystem function as more than 70% of land plants rely
on mycorrhizal relationships with fungi to provide them
with inorganic compounds and trace elements. In
addition, mutualism has driven the evolution of much of
the biological diversity we see, such as flower forms
(important for pollination mutualisms) and co-evolution
between groups of species. However mutualism has
historically received less attention than other interactions
such as predation and parasitism.
Measuring the exact fitness benefit to the individuals is
not always straightforward, particularly when the
individuals can receive benefits from a range of species,
for example most plant-pollinator mutualisms. It is
therefore common to categorise mutualisms according to
the closeness of the association, using terms such as
obligate versus facultative. Defining "closeness",
however, is also problematical. It can refer to mutual
dependency (the species cannot live without one
another) or the biological intimacy of the relationship in
relation to physical closeness (e.g. one species living
within the tissues of the other species).
Types of relationships: Mutualistic interactions can be
thought of as a form of "biological barter" in which
species trade resources (for example carbohydrates or
inorganic compounds) or services such as gamete,
offspring dispersal, or protection from predators.
Resource-resource relationships:Resource-resource
interactions, in which one type of resource is traded for a
different resource, are probably the most common form
of mutualism; for example mycorrhizal associations
between plant roots and fungi, with the plant providing
carbohydrates to the fungus in return for primarily
phosphate but also nitrogenous compounds. Other
examples include rhizobia bacteria which fix nitrogen
for leguminous plants (family Fabaceae) in return for
energy-containing carbohydrates.
Service-resource relationships: Service-resource
relationships are also common, for example pollination
in which nectar or pollen (food resources) are traded for
pollen dispersal (a service) or ant protection of aphids,
where the aphids trade sugar-rich honeydew (a by-
product of their mode of feeding on plant sap) in return
for defence against predators such as ladybird beetles.
INSTITUTE FOR ADVANCED STUDIES, JODHPUR  37
Service-service relationships: Strict service-service
interactions are very rare, for reasons that are far from
clear. One example is the relationship between sea
anemones and anemonefish in the family Pomacentridae:
the anemones provide the fish with protection from
predators (which cannot tolerate the stings of the
anemone's tentacles) and the fish defend the anemones
against butterfly fish (family Chaetodontidae) which eat
anemones. However, in common with many mutualisms,
there is more than one aspect to the biological barter: in
the anemonefish-anemone mutualism, waste ammonia
from the fish feed the symbiotic algae that are found in
the anemone's tentacles.
Therefore what appears to be a service-service
mutualism in fact has a service-resource component. A
second example is that of the relationship between some
ants and trees in the genus Acacia, such as the Whistling
Thorn and Bullhorn Acacia. The ants nest inside the
plant's thorns. In exchange for shelter, the ants protect
acacias from attack by herbivores (which they frequently
eat, introducing a resource component to this service-
service relationship) and competition from other plants
by trimming back vegetation that would shade the
acacia. In addition, another service-resource component
is present, as the ants regularly feed on lipid-rich food-
bodies called Beltian bodies that are on the Acacia plant.
Humans and mutualism: Humans also engage in
mutualisms with other species, including our gut flora
(without which we would not be able to digest food
efficiently) and domesticated animals such as horses,
which provide transportation in return for food and
shelter. In traditional agriculture, many plants will
function mutualistically as companion plants, providing
each other with shelter, soil fertility and the repelling of
pests. For example, beans may grow up cornstalks as a
trellis, while fixing nitrogen in the soil for the corn, as
exploited in the Three Sisters gardening technique. The
question how and why species might cooperate has been
addressed philosophically by a number of writers. Gilles
Deleuze, for example, was interested in the way this
questioned the conception of evolutionism and the
notion of linear historical progress.
A large percentage of herbivores have mutualistic gut
fauna that help them digest plant matter, which is more
difficult to digest than animal prey. Coral reefs are the
result of mutualisms between coral organisms and
various types of algae that live inside them. Most land
plants and land ecosystems rely on mutualisms between
the plants which fix carbon from the air, and
Mycorrhyzal fungi which help in extracting minerals
from the ground.
Another example is the goby fish, which sometimes lives
together with a shrimp. The shrimp digs and cleans up a
burrow in the sand in which both the shrimp and the
goby fish live. The shrimp is almost blind leaving it
vulnerable to predators when above ground. In case of
danger the goby fish touches the shrimp with its tail to
warn it. When that happens both the shrimp and goby
fish quickly retract into the burrow.
One of the most spectacular examples of obligate
mutualism is between the siboglinid tube worms and
symbiotic bacteria that live at hydrothermal vents and
UNIT 10: ECOLOGICAL PRINCIPLES 
cold seeps. The worm has no digestive tract and is solely
reliant on their internal symbionts for nutrition. The
bacteria oxidize either hydrogen sulfide or methane
which the host supplies to them. These worms were
discovered in the late 1980s at the hydrothermal vents
near the Galapagos Islands and have since been found at
deep-sea hydrothermal vents and cold seeps in all of the
world's oceans.
Symbiosis and evolution : While historically, symbiosis
has received less attention than other interactions such as
predation or competition, it is increasingly recognised as
an important selective force behind evolution, with many
species having a long history of interdependent co-
evolution. In fact the evolution of all eukaryotes (plants,
animals, fungi, protists) is believed to have resulted from
a symbiosis between various sorts of bacteria.
Symbiogenesis: The biologist Lynn Margulis, famous
for the work on endosymbiosis, contends that symbiosis
is a major driving force behind evolution. She considers
Darwin's notion of evolution, driven by competition, as
incomplete, and claims evolution is strongly based on
co-operation, interaction, and mutual dependence among
organisms. According to Margulis and Dorion Sagan,
"Life did not take over the globe by combat, but by
networking."
Co-evolution: Symbiosis played a major role in the co-
evolution of flowering plants and the animals that
pollinate them. Many plants that are pollinated by
insects, bats or birds, have very specialized flowers
modified to promote pollination by a specific pollinator
that is also correspondingly adapted. The first flowering
plants in the fossil record had relatively simple flowers.
Adaptive speciation quickly gave rise to many diverse
groups of plants, and at the same time, corresponding
speciation occurred in certain insects groups. Some
groups of plants developed nectar and large sticky pollen
while insects evolved more specialized morphologies to
access and collect these rich food sources. In some taxa
of plants and insects the relationship has become
dependent,[32] where the plant species can only be
pollinated by one species of insect.
Biotic Pollination: a best example of mutualism
Pollination in angiosperms and gymnosperms is the
process that transfers pollen grains, which contain the
male gametes (sperm) to where the female gamete(s) are
contained within the carpel; in gymnosperms the pollen
is directly applied to the ovule itself. The study of
pollination brings together many disciplines, such as
botany, horticulture, entomology, and ecology.
Pollination is a necessary step in the sexual reproduction
of flowering plants, resulting in the production of
offspring that are genetically divers. It is important in
horticulture and agriculture, because fruiting is
dependent on fertilisation, which is the end result of
pollination.
Types: The process of pollination requires pollinators:
agents that carry or move the pollen grains from the
anther to the receptive part of the carpel or pistil. The
receptive part of the carpel is called a stigma in the
flowers of angiosperms. The receptive part of the
38 INSTITUTE FOR ADVANCED STUDIES, JODHPUR
gymnosperm ovule is called the micropyle. The various
flower traits that attract different pollinators are known
as pollination syndromes. Methods of pollination, with
common pollinators or plants, are:
Biotic pollination, occurs when pollination is mediated
by an organism, termed a pollinator. There are roughly
200,000 varieties of animal pollinators in the wild, most
of which are insects.
Entomophily, pollination by insects, often occurs on
plants that have developed blue petals and a strong scent
to attract insects such as, bees, wasps and occasionally
ants (Hymenoptera), beetles (Coleoptera), moths and
butterflies (Lepidoptera), and flies (Diptera).
In Zoophily, pollination is done by vertebrates such as
birds and bats, particularly, hummingbirds, sunbirds,
spiderhunters, honeyeaters, and fruit Bats.
Pollinators and pollenizers: Pollination also requires
consideration of pollenizers. (The terms "pollinator" and
"pollenizer" are often confused: a pollinator is the agent
that moves the pollen, whether it be wind, bees, bats,
moths, or birds; a pollenizer is the plant that provides the
pollen.) Some plants are self-fertile or self-compatible
and can pollinate themselves. Other plants have
chemical or physical barriers to self-pollination and need
to be cross-pollinated: with these self-infertile plants, not
only pollinators must be considered but pollenizers as
well. In pollination management, a good pollenizer is a
plant that provides compatible, viable and plentiful
pollen and blooms at the same time as the plant that is to
be pollinated.
Pollination can be cross-pollination with a pollinator and
an external pollenizer, self-pollenization with a
pollinator, or self-pollination without any pollinator:
• Cross-pollination (syngamy): pollen is delivered to a
flower of a different plant. Plants adapted to outcross or
cross-pollinise have taller stamens than carpels to better
spread pollen to other flowers.
• Self-pollenization (autogamy): pollen moves to the
female part of the same flower, or to another flower on
the same individual plant. This is sometimes referred to
as self-pollination, but this is not synonymous with
autogamy. Clarity requires that the term "self-
pollination" be restricted to those plants that accomplish
pollination without an external pollinator (example: the
stamens actually grow into contact with the pistil to
transfer the pollen). Most peach varieties are
autogamous, but not truly self-pollinated, as it is
generally an insect pollinator that moves the pollen from
anther to stigma. Plants adapted to self-fertilize have
similar stamen and carpel length.
• Cleistogamy: pollination that occurs before the
flower opens is always self-pollination. Some
cleistogamous flowers never open, in contrast to
chasmogamous flowers that open and are then
pollinated. Cleistogamous flowers must of necessity be
self-compatible or self-fertile plants. Other plants are
self-incompatible. These are end points on a continuum,
not absolute points.
Hybridization is effective pollination between flowers of
different species of the same genus, or even between
flowers of different genera (as in the case of several
 
UNIT 10: ECOLOGICAL PRINCIPLES 
orchids). Peaches are considered self-fertile because a
commercial crop can be produced without cross-
pollination, though cross-pollination usually gives a
better crop. Apples are considered self-incompatible,
because a commercial crop must be cross-pollinated.
Remember that most fruits are grafted clones,
genetically identical. An orchard block of apples of one
variety is in effect all one plant. Growers now consider
this a mistake. One means of correcting this mistake is to
graft a limb of an appropriate pollenizer (generally a
variety of crabapple) every six trees or so.
Commensalism
In ecology, commensalism is a kind of symbiotic
relationship between two organisms where one benefits
and the other is not significantly harmed or helped (like
a bird living in a tree). The term derives from the
English word commensal, meaning "sharing of food" in
human social interaction; that word derives from the
Latin com mensa, meaning "sharing a table".
Types: As with all ecological interactions,
commensalisms vary in strength and duration from
intimate, long-lived symbioses to brief, weak
interactions through intermediaries. It is one of the three
kinds of symbiotic relationships. Originally it was used
to describe the use of waste food by second animals, like
the carcass eaters that follow hunting animals but wait
until they have finished their meal. Other forms of
commensalism include:
• Phoresy: One animal attaching to another animal for
transportation only. This concerns mainly arthropods,
examples of which are mites on insects (such as beetles,
flies, or bees), pseudoscorpions on mammals and
millipedes on birds. Phoresy can be either obligate or
facultative (induced by environmental conditions).
• Inquilinism: Using a second organism for housing.
Examples are epiphytic plants (such as many orchids)
which grow on trees, or birds that live in holes in trees.
• Metabiosis: A more indirect dependency, in which
the second organism uses something the first created,
however after the death of the first. An example is the
hermit crabs that use gastropod shells to protect their
bodies.
The question of whether the relationship between
humans and some types of our gut flora is commensal or
mutualistic is still unanswered.
Some biologists argue that any close interaction between
two organisms is unlikely to be completely neutral for
either party, and that relationships identified as
commensal are likely mutualistic or parasitic in a subtle
way that has not been detected. For example, epiphytes
are "nutritional pirates" that may intercept substantial
amounts of mineral nutrients that would otherwise go to
the host plant. Large numbers of epiphytes can also
cause tree limbs to break or shade the host plant and
reduce its rate of photosynthesis. Similarly, the phoretic
mites in the image above may hinder their host by
making flight more difficult, which may affect its aerial
hunting ability or cause it to expend extra energy while
carrying these passengers.
Parasitism
INSTITUTE FOR ADVANCED STUDIES, JODHPUR  39
Parasitism is a type of symbiotic relationship between
organisms of different species in which one, the parasite,
benefits from a prolonged, close association with the
other, the host, which is harmed. In general, parasites are
much smaller than their hosts, show a high degree of
specialization for their mode of life and reproduce more
quickly and in greater numbers than their hosts. Classic
examples of parasitism include the interactions between
vertebrate hosts and such diverse animals as the
tapeworms, flukes, Plasmodium species and fleas.
The harm and benefit in parasitic interactions concern
the biological fitness of the organisms involved.
Parasites reduce host fitness in many ways, ranging from
general or specialized pathology (such as castration),
impairment of secondary sex characteristics, to the
modification of host behaviour. Parasites increase their
fitness by exploiting hosts for food, habitat and
dispersal.
Although the concept of parasitism applies
unambiguously to many cases in nature, it is best
considered part of a continuum of types of interactions
between species, rather than an exclusive category.
Particular interactions between species may satisfy some
but not all parts of the definition. In many cases, it is
difficult to demonstrate that the host is harmed. In
others, there may be no apparent specialization on the
part of the parasite, or the interaction between the
organisms may be short-lived. In medicine, only
eukaryotic organisms are considered parasites, to the
exclusion of bacteria and viruses. Some branches of
biology, however, do regard members of these groups to
be parasitic.
Types of parasitism: Parasites are classified based on a
variety of aspects of their interactions with their hosts
and on their life cycles.
ƒ Those that live inside the host are called
endoparasites (e.g., hookworms) and those that live on
its surface are called ectoparasites (e.g., some mites).
ƒ An epiparasite is one that feeds on another parasite.
This relationship is also sometimes referred to as
"hyperparasitism".
ƒ Parasitoids are organisms that cause the host to die
as a result of parasitism. Thus, the interaction between
the parasitoid and the host is fundamentally different
than true parasites and their host, and shares some
characteristics with predation.
ƒ Social parasites take advantage of interactions
between members of social organisms such as ants or
termites. In kleptoparasitism, parasites appropriate food
gathered by the host. An example is the brood parasitism
practiced by many species of cuckoo. Many cuckoos use
other bird species as "babysitters", depositing their eggs
in the nest of the host species, which raise the cuckoo
young as one of their own.
Parasitism can take the form of isolated cheating or
exploitation among more generalized mutualistic
interactions. For example, broad classes of plants and
fungi exchange carbon and nutrients in common
mutualistic mycorrhizal relationships; however, a few
plants species (known as myco-heterotrophs) "cheat" by
taking carbon from a fungus rather than donating it.
UNIT 10: ECOLOGICAL PRINCIPLES 
Evolutionary aspects: Biotrophic parasitism is an
extremely common mode of life that has arisen
independently many times in the course of evolution.
Depending on the definition used, as many as half of all
animals have at least one parasitic phase in their life
cycles, and it is also frequent in plants and fungi.
Moreover, almost all free-living animals are host to one
or more parasite taxa.
Parasites evolve in response to defense mechanisms of
their hosts. Examples of host defenses include the toxins
produced by plants to deter parasitic fungi and bacteria,
the complex vertebrate immune system, which can target
parasites through contact with bodily fluids, and
behavioural defenses. An example of the latter is the
avoidance by sheep of open pastures during spring, when
roundworm eggs accumulated over the previous year
hatch en masse. As a result of these and other host
defenses, some parasites evolve adaptations that are
specific to a particular host taxon and specialize to the
point where they infect only a single species. Such
narrow host specificity can be costly over evolutionary
time, however, if the host species becomes extinct. Thus,
many parasites are capable of infecting a variety of host
species that are more or less closely related, with varying
success.
Host defenses also evolve in response to attacks by
parasites. Theoretically, parasites may have an
advantage in this evolutionary arms race because of their
more rapid generation time. Hosts reproduce less quickly
than parasites, and therefore have fewer chances to adapt
than their parasites do over a given span of time.
E. COMMUNITY ECOLOGY 
1. Concept    Species diversity
Community: An association of interacting populations,
usually defined by the nature of their interaction or the
place in which they live. "community" can be used in a
more limited sense, such as the plant community, animal
community, or even the bird community, the insect
community, etc.
Holistic concept: The view that communities are super
organisms that can only be understood as wholes
(Clements)
Individualistic concept: the idea that communities are
merely functions of populations, and are not organized in
any way (Gleason).
Most ecologists today are between these two extremes.
Keystone species: a species which influences the entire
community so much that its removal will cause
ecosystem collapse (can be a keystone predator).
2. Characteristics of a Community
Like a population, a community too has its own
characteristics, which are not shown by its individual
component species. These characteristics which have
meaning only with reference to community level of
organisation, are as follows:
40 INSTITUTE FOR ADVANCED STUDIES, JODHPUR
In some cases, a parasite species may coevolve with its
host taxa. In theory, long-term coevolution should lead
to a relatively stable relationship tending to
commensalism or mutualism, in that it is in the
evolutionary interest of the parasite that its host thrives.
A parasite may evolve to become less harmful for its
host or a host may evolve to cope with the unavoidable
presence of a parasite to the point that the parasite's
absence causes the host harm. For example, although
animals infected with parasitic worms are often clearly
harmed, and therefore parasitized, such infections may
also reduce the prevalence and effects of autoimmune
disorders in animal hosts, including humans.
The presumption of a shared evolutionary history
between parasites and hosts can sometimes elucidate
how host taxa are related. For instance, there has been
dispute about whether flamingos are more closely related
to the storks and their allies or to ducks, geese and their
relatives. The fact that flamingos share parasites with
ducks and geese is evidence these groups may be more
closely related to each other than either is to storks.
Parasitism is part of one explanation for the evolution of
secondary sex characteristics seen in breeding males
throughout the animal world, such as the plumage of
male peacocks and manes of male lions. According to
this theory, female hosts select males for breeding based
on such characteristics because they indicate resistance
to parasites and other disease.
Each community is made up of much different
organisms-plants, animals, microbes, which differ
taxonomically from each other. The number of species
and population abundance in community also vary
greatly.
There are two levels of species diversity: (i) regional
diversity of whole nations or parts of continents within
which many different communities exist, and (ii) local
diversity in a given nation where different communities
exist at different latitudes.
Growth form and structure
Community is described in terms of major growth forms
as trees, shrubs, herbs, mosses etc. In each growth form
as in trees, there may be different kinds of plants as
broad-leaved trees, evergreen trees etc. These different
growth forms determines the structural pattern of a
community. According to the mode of arrangement of
the various growth forms, the communities exhibit: (i)
zonation-horizontal layering) as well as, (ii)
stratification-vertical layering).
Dominance: In each community all ,the species are not
equally important. There are relatively only a few of
these, which determine the nature of the community.
These few species exert a major controlling influence on
the community. Such species are known as dominants.
 
UNIT 10: ECOLOGICAL PRINCIPLES 
Dominance may be due to number, biomass or energy
flow.
Trophic structure (Self-sufficiency)
Nutritionally, each community, a group of autotrophic
plants as well as heterotrophic animals, exists as a self-
sufficient, perfectly balanced assemblage of organisms.
Each community has its own composition, structure and
developmental history.
Composition: Communities may be large or small.
Larger ones extend over areas of several thousands of
square kilometers, as forests, others such as deserts etc.
are comparatively smaller with dimensions in hundreds
of kilometers, and still others such as meadows, rivers,
ponds, rocky plateaus etc. occupying a more restricted
area. Very small-sized communities are the groups of
microorganisms in such microhabitats as leaf surface,
fallen log, litter, soil etc. The number of species and
population abundance in communities vary greatly.
In each community there are diverse species. All these
species are not equally important but these are only a
few overtopping species which by their bulk and growth
modify the habitat and control the growth of other
species of the community, thus forming a sort of
characteristic nucleus in the community. These species
are called the dominants. Generally in most of the
communities, only a single species, due to being
particularly conspicuous, is dominant, and in such case
the community is called by the name of dominant
species as for example, spruce forest community. In
other communities, there may be more than one
dominants as in oak-hickory forest community.
INSTITUTE FOR ADVANCED STUDIES, JODHPUR  41
Structure: Besides composition and dominance, the
communities exhibit a structure or recognizable pattern
in the spatial arrangement of their members. Thus
structurally, a community may be divided horizontally
into' 'sub-communities', which are units of homogeneous
life-form and ecological relation.
This horizontal, division constitutes the zonation in the
community. Figure 1 shows zonation of different distinct
vegetational types on a mountain. Latitudinal as well as
altitudinal zonations of vegetation have also been shown
in relation with climatic factors. In shallow ponds,
zonation is very little. However, in deep ponds and
lakes, there may be recognised three zones (Fig. 2), viz,
littoral zone, limnetic zone and profundal zone. In each
zone, organisms differ from each other.
Another aspect of structure that is more common, is
stratification which involves vertical rather than
horizontal changes within the community. In each
horizontal zone, there may be recognised distinct vertical
storeys. Sometimes the stratification is very complex
where community possesses a number of vertical layers
of species, each made up of a characteristic growth form.
In grassland communities, there is a subterranean floor,
containing basal portions of the vegetation such as
rhizomes of grass covered by litter and debris of plants
as well as animals, and herbaceous substratum consisting
of upper parts of the grasses and herbs with a
characteristic fauna.
However, stratification in a forest community (Fig. 3) is
most complicated where as many as five vertical sub
divisions may be recognised. These vertical subdivisions
are: (i) subterranean subdivision, (ii) forest floor, (iii)
herbaceous vegetation, (iv) shrubs, and (v) trees. In some
tropical rain forests, there may be as many as eight
vertical strata. Thus, based upon the light and relative
UNIT 10: ECOLOGICAL PRINCIPLES 
humidity requirements, we find stratification in above
ground parts. Similar stratification may also be found in
the underground parts, roots, rhizome or other structures
below the soil.
Origin and development: A community with its
particular environment constitutes an entity which has its
origin and development. The details of community
development will be given in the next section
(Ecological Succession). However, for the present
purpose, a brief outline of the process is being given
here. In a barren area there reach the seeds and
propagules of the species. This is known as migration.
The seed.s or propagules after germination develop into
the seedlings which then develop into adults. But only a
few of these survive as by this time the new environment
does not suit to all of them and thus only a few are
capable of successful growth, and this process of
seedling establishment and successful growth is called
ecesis. As a result of migration and subsequent ecesis,
species colonise the new areas - colonization. By this
time with the changing environment due to plants'
growth, several other species of both plants and animals
start colonising the area and sooner or later the area is
colonised by a definite community.
This community constitutes the biotic environment of
that particular area. For quite sometime, species live
together in a community under a particular set of
environmental conditions. But time and place affect the
living organism very much, as a result of which there are
chiefly two major types of relationships between the
organisms of a community and their environment. These
are as follows:
Interrelations between organisms themselves: Such
relations are chiefly in respect of food and space. These
include:
Competition: In the process of rapid colonisation,
individuals become aggregated at a place, and there
develop inter-as well as intraspecific competition for
space and food. As a result of these actions and
interactions on the habitat, the pioneer species modify
their own environment. Those, which are unable to adapt
themselves to the changing environmental conditions,
disappear from the area. This changed habitat is now
colonised by newer species. This community may lead to
another, entirely different type of community. This
orderly change, which leads to the development of
vegetation, is called succession. Sooner or later this
change comes to a stop, when a particular community
comes to stay and both, community and the habitat have
reached a stabilization. This final community is called
the cUmax community.
Stratification: The vertical strata in a community have
already been explained. For example, in forests we find
several storeys or layers of plant species and in each
layer plant species differ in size, form, behaviour etc.,
Various plants in each stratum are adapted to that
particular set of local conditions and moreover, the
plants of one layer modify the environment, which is
suitable for species in another layer of the community.
Stratification thus is the result of interdependencies of
species, as for example, lianas and epiphytes grow on
other plants.
42 INSTITUTE FOR ADVANCED STUDIES, JODHPUR
Cohabitation: There are reports of a number of
chemicals secreted by vegetative organs of plants of a
community which modify the edaphic conditions.
Similarly, chemicals secreted by fungi, bacteria and
actinomycetes in soil affect the root system as well as
shoots of higher plants.
Relationships between species and the environment:
Organisms and the environment interact with each other
and bring about several modifications. This is one of the
chief reasons that a particular community changes to
another type sooner or later in a particular area.
Fluctuation: 
 
Once  the  population  complete  their  growth, 
population  density  tends  to  fluctuate  above  and 
below the steady state level (carrying capacity) even 
in  population  subject  to  various  forms  of  feedback 
control. Often the fluctuation result form the seasonal 
or annual changes in availability of resources, or they 
may  be  stochastic(random).  However,  some 
population  oscillate  so  regularly  that  they  can  be 
classified as as cyclic. 
 
Seasonal  Fluctuation  in  population  size,  largely 
controlled  by  life  history  adaptations  coupled  with 
swasonal  changes  in  the  environment  factors.  Eg.  At 
certain times of year mosquito will be abundant. 
 
Annual Fluctuation may be considered as under two 
sub‐headings  i)  fluctuation  controlled  primarily  by 
annual  differences  in  extrinsic  factors  such  as 
temperature  and  rainfall.  ii)  intrinsic  factors, 
oscillation  is  controlled  primarily  by  population 
dynamics  (biotic  factors  such  as  food  or  energy 
awailability, disease etc.) 
Year to year changes in the abundance seems clearly 
correlated  with  variation  in  one  or  more  major 
extrinsic  limitating  factors  are  called  cyclic 
fluctuation. 
Characters Used in Community Structure
It is pointed out above that each community is
characterised by its species diversity, growth forms and
structure, dominance, successional trend etc. To study
the details of these aspects of any community there are
taken into consideration a number of characters
(parameters). These are then used to express the
characteristics of a community. For instance, if we say
that this particular species is dominant in the community,
what may be the possible characters on the basis of
which such a conclusion can be drawn.
These various characters used for such purposes are
broadly classified into two major categories: (i)
analytical characters, these are generally expressed in
terms of 5-point scale and (ii) synthetic characters are
actually computed from analytical characters.
 
UNIT 10: ECOLOGICAL PRINCIPLES 
Analytical characters
These are quantitative, which are expressed in
quantitative terms, and qualitative, expressed only in
qualitative way although finally grouped under 5-point
scale.
Quantitative characters: These include such characters
as frequency, density, abundance, cover and basal area
etc.
Frequency: Various species of the community are
recorded by different phytosociological methods, by
taking any sampling unit like quadrat, transect and point.
Details of these methods will be described later.
Frequency is the number of sampling units (as %) in
which a particular species occurs. Thus frequency of
each species is calculated as follows :
Frequency (%) =
No. of Quadrates in which species of occurredx100
Total no. of sampling units studied
After determining the percentage frequency of each
species, various species are distributed among
Raunkiaer's (1934) five frequency classes depending
upon their frequency values as follows:
Frequency % Frequency class
0-20 A the number of organisms sometimes may not give
21-40 B correct idea of the species. If we base our conclusion on
41-60 C number, a single or few trees in a grassland, or few
61-80 D grasses in a forest should be of little value. But if we
81-100 E consider the species on the basis of area occupied or
Density: Frequency does not give correct idea of the
distribution of any species, unless it is correlated with
other characters, such as density etc. Density represents
the numerical strength of a species in the community.
The number of individuals of the species in any unit area
is its density.
Density gives an idea of degree of competition. It is
calculated as follows:
Density = Total no. of individuals of the species in all
the sampling units /Total no. of sampling units studied.
The value thus obtained is then expressed as number of
individuals per unit area.
Abundance: This is the number of individuals of any
species per sampling unit of occurrence. It is calculated
as follows:
Abundance= Total no. of individuals of the species in all
the sampling units/ Total no. of sampling units studied
But, abundance thus obtained in quantitative terms gives
little idea of the distribution of the species.
Cover and basal area: Quantitative values of frequency,
density etc., do not give correct information on the
distribution of a species. Since, different species differ in
their growth forms, such values' are to be supplemented
with other characters like cover and basal area.
Cover, although sometimes used in general sense for the
area occupied by a plant, (which may be the herbage
INSTITUTE FOR ADVANCED STUDIES, JODHPUR  43
cover or the cover or basal area), it is generally used for
above ground parts. Thus cover or herbage cover
signifies primarily the area of ground occupied by the
above ground parts of plants, such as leaves, stems and
inflorescence as viewed from above. It is a good
measure of the herbage availability. It is estimated by
chart-quadrat, line intercept or point frame.
Basal area refers to the ground actually penetrated by the
stems and is readily seen when the leaves and stems are
clipped at the ground surface. It is one of the chief
characteristics to determine dominance. It is measured
either 2.5 cm above ground or actually on the ground
level. It is measured by callipers, line-interception or
point-centered quadrat method.
Since plants differ in their growth form the relationship
between herbage cover and basal area differs in different
types of plants. Based on their cover values, species are
grouped into six groups as follows: Group 1 (less than
5%), 2 (5-25), 3 (25-50%), 4 (50-75%), 5(75-95%) and 6
(95-100%). Braun-Blanquet (1932) included only five
groups.
Dominance: Infact, dominance is a synthetic character.
However, Daubenmire (1959) considered it as a
analytical character and expressed its six types. Infact
weight (biomass), the situation may be different. Thus in
dominance, cover is included as important character. The
single tree in grassland may occupy fairly a large area
and may have much mass.
Qualitative characters
These include physiognomy, phenology, stratification,
abundance, sociability or gregariousness, vitality and
vigour, life form (growth form) etc., most of which are
described, and may be grouped in point scales.
Physiognomy: This is the general appearance of
vegetation as determined by the growth form of
dominant species. Such a characteristic appearance can
be expressed by single term. For example, a simple look
to such a community of 'Plants where larger trees are
dominant with some shrubs, would tell, that it is a forest.
Similarity on the basis of apperarnace it may be
grassland, desert etc.
Phenology: It also includes periodicity, aspection etc.
Phenology is the scientific study of seasonal change i.e.
the periodic phenomena of organisms in relation to their
climate. Different species have different periods of seed
germination, vegetative growth, flowering and fruiting,
leaf fall, seed and fruit dispersal etc. Such data for
individual species are recorded. A study of the date and
time of these events is phenology. In other words,
phenology is the calendar of events in the life history of
the plant. These events are shown by phenograms. Thus
phenology of different species may differ from each
other and in a community, we find species with different
phenology, changing the composition of community.
UNIT 10: ECOLOGICAL PRINCIPLES 
Stratification: As pointed elsewhere, stratification is the
way in which plants of different species are arranged in
different vertical layers in order to make full use of the
available physical and physiological requirements. Data
on stratification are obtained by using a bisect. Vertical
projection is constructed and stature of the species may
be plotted to find out distribution pattern of stem, roots
etc. of different species.
Abundance: Although related with density, abundance
may not be expressed generally in 'quantitative terms.
Organisms, particularly plants are not found uniformly
distributed in an area. They are found in smaller patches
or groups, differing in number at each place. Abundance
is divided in five arbitrary groups, depending upon the
number of plants. The groups are, very rare, rare,
common, frequent and very much frequent.
Sociability: It denotes the proximity of plants to one
another. Plants generally grow as isolated individuals, in
grow better when nearer to each other and produce thick
population. Others become weak or die in such an
association. Fruit and seed dispersal etc. and nature of
reproduction are affected by the way in which they are
aggregated. Species with same density, may differ in
sociability values. Thus sociability expresses the degree
of association between species. Some divide the
sociability into as many as ten classes, but Braun -
Blanquet (1932) used only five sociability groups as
follows:
ƒ S1-Plants (stems) found quite separately from each
other, thus growing singly.
ƒ S2-A group of 4-6 plants at one place
ƒ S3-Many smaller scattered groups at one place
ƒ S4-Several bigger groups of many plants at one
place
ƒ S5-A large group occupying larger area.
Vitality: This is the capacity of normal growth and
reproduction, which are important for successful survival
of a species. The vitality depends upon weight of
species. In plant, stem height. root length, leaf area, leaf
number, number and weight of flowers, fruits, seeds etc.
determine the vitality.
On this basis plants are grouped into different vitality
groups. R. Misra and G. S. Purl (1954) gave three groups
as (i) well developed plants completing their life cycles
regularly, prodqcing viable seeds, (ii) plants reproducing
vegetatively and (iii) ephemerals, short lived. Braun-
Blanquet (1932) gave four groups.
Daubenmire (1968) gave vitality groups as follows:
ƒ V1-Plants whose seedlings die
ƒ V2-Seedlings grow, but unable to reproduce
ƒ V3-Reproduce only vegetatively
ƒ V4-Reproduce sexually, but uncommon
ƒ V5-Reproduce sexually and grow regularly.
Life-form (Growth form): In past, ecologists used the
general appearance of plant communities (as determined
by the stature of the plant species, their spread and
characters of life forms, as the basis of the study of
communities. These methods are known as
physiognomic methods. Of these, Raunkiaer's (1934)
life-form method has been commonly employed. A life-
form is "the sum of the adaptation of the plant to
44 INSTITUTE FOR ADVANCED STUDIES, JODHPUR
climate". He considered that the way in which different
species overcome the adverse environmental conditions
determines their limits of distribution. Thus the plants'
climate can be expressed by the statistical distribution of
life-forms in the flora of a particular region. On the basis
of the position of perennating buds on plants and the
degree of their protection during adverse conditions,
Raunkiaer classified plants into five broad life-form
categories which are shown in Figure 4.
These are as follows:
ƒ Phanerophytes: Their buds are naked or covered
with scale, and are situated high up on the plant.
These life-forms include trees, shrubs and climbers
generally common in tropical climates. Depending
upon the size, they may be further subdivided into
megaphanerophytes (over 30 meters high), meso-
phanerophytes (8-30 meters high), micro-
phanerophttes (2-8 meters high), and nano-
phanerophytes (under 2 meters). Epiphytes are
either included in phanerophytes, or sometimes
included under a separate life-form.
ƒ Chamaephytes: Their buds are situated close to the
ground surface. They are common at high altitudes.
Example-Trifolium repens.
ƒ Hemicryptophytes: These are mostly found in cold
temperate zone. Their buds are hidden under soil
surface, protected by the soil itself. Their shoots
generally die each year. Examples-most of the
biennial and perennial herbs.
ƒ Cryptophytes or Geophytes: Their buds are
completely hidden in the soil, as bulbs and
rhizomes. Most of them are found in arid zones.
Hydrophytes are the cryptophytes whose buds are
found below the water surface.
ƒ Therophytes: These are seasonal plants, completing
their life cycle in a single favourable season, and
remain dormant throughout the rest unfavourable
period of year in the form of seeds. They are
common in deserts. Braun-Blanquet (1951), on the
basis of Raunkiaer's life-forms, divided plants into
as many as ten categories.
 
UNIT 10: ECOLOGICAL PRINCIPLES 
Synthetic characters
These are determined after computing the data on the
quantitative and qualitative characters of the community.
For comparing the vegetation of different areas,
community comparison needs the calculation of their
synthetic characters. These are determined in terms of:
Presence and constance: It expresses the extent of
occurrence of the individuals of a particular species in
the community Le. how uniformly a species occurs in a
number of stands of the same type of community. The
species on the basis of its percentage frequency may
belong to any of the following five presence classes:
ƒ Rare-present in 1 to 20% of the sampling units
ƒ Seldom present-present in 21-40% of the sampling
units
ƒ Often present-present in 41-60% of the sampling
units
F. Ecological succession: Types; mechanisms; changes involved in succession; concept of climax.
Ecological Succession
Environment is always kept on changing over a period of
tome due to i) variation in climatic and physiographic
factors and ii) the activities of species of communities
themselves. These influences bring about marked
changes in the dominants of existing community, which
in turn sooner or later replaced by another community at
the same place. This process continues and successive
communities develops one after another over the same
area , until the final community again becomes more or
less stable for a period of time. This occurrence of
relatively different sequence of communities over a
period of time in same area is known as ecological
succession.
INSTITUTE FOR ADVANCED STUDIES, JODHPUR  45
ƒ Mostly present-present in 61-80% of the sampling
units
ƒ Constantly present-present in 81-100% of the
sampling units.
Fidelity: This is the degree with which a species is
restricted in distribution to one kind of community. Such
species are sometimes known as indicators. The species
have been grouped into five fidelity classes as follows:
ƒ Fidelity 1 - Plants appearing accidentally.
ƒ Fidelity 2 - Indifferent plants, may occur in any
community.
ƒ Fidelity 3 - Occur in many communities, but
predominant in one (Prefer-ants).
ƒ Fidelity 4 - Specially present in one community but
may occasionally occur in either community as well
(Selectives).
ƒ Fidelity 5 - Occur only in one particular community
and not in others (Exclusives).
Dominance: It has been described under quantitative
characters. Here, the dominance is expressed in synthetic
form. On the basis of density, frequency and dominance
(cover) values, there has been proposed the idea of
importance value index (IVI). IVI of a species in the
community gives the idea of its relative importance. For
IVI, values of relative density, relative frequency and
relative dominance (cover basis) are obtained as follows:
Relative frequency= Frequency of the species x 100
Total frequency of all the species
Relative density = Density of the species x l00
Total density of all the species
Relative dominance= Dominance of the species x 100
Total dominance of all the species
Now for IVI, three values are added. IVI values of
different species are then arranged in decreasing order.
Other synthetic characters: Besides the above
mentioned ones, there have also been proposed some
other characters. These have been quite useful in
comparative studies on communities. Such characters
include, interspecific association and association index,
index of similarity, dominance index, species diversity
and diversity index etc. It is beyond the scope of the text
of the book to go into details of such characters.
  
1 Trends of Succession:
1. An orderly continuous change in kind of plants and
animals.
2. Tends to increase in diversity of species.
3. An increase in organic matter and biomass
supported by the awailable energy flow(reverse in
heterotropic succession).
4. Decrease in net community productivity or annual
yield.
2 TYPES OF SUCCESSION
Autotrophic vs. heterotrophic
ƒ Autotrophic succession: Succession in which most of
the organic matter is fixed by autotrophs; most living
biomass is in plants.
UNIT 10: ECOLOGICAL PRINCIPLES 
ƒ Heterotrophic (Degradative) succession: Succession
on a degradable source; most living biomass is animal,
fungal, or microbial material.
Primary vs. Secondary
ƒ Primary succession (glossary): sequence of
communities developing in a newly exposed habitat
devoid of life (eg. succession on bare rock, newly
deposited sand).
ƒ Secondary succession (different from glossary):
sequence of communities taking place on sites that have
already supported life (eg. oldfield succession, clearcut
forests, burned areas, etc.).
Perhaps the most useful distinction between primary and
secondary succession is that the latter originates with
soil.
Autogenic vs. allogenic (heterogenic)
ƒ In autogenic succession environment is modified by
species of community itself such that conditions become
unfavorable for that community and more favorable for
upcoming communities.
ƒ In allogenic succession replacement of community
is due to other external factors
3 THREE MECHANISMS OF SUCCESSION
a. Facilitation: The organisms at a given successional
stage make the environment more suitable for later
successional stages. Examples: lichens breaking down
rock into soil, nitrogen-fixing plants improve fertility of
soil, nurse plants
b. Tolerance: The organisms of a given successional
stage have little impact on later successional stages.
Example: Oldfield succession (possibly)- species of all
stages get started at the same time, but are dominant at
different times because of different life histories.
c. Inhibition: The organisms at a given stage resist
invasion by organisms of later stages. Succession
proceeds when the individuals of a given stage die.
Example: allelopathy
4. CLIMAX THEORIES
• Monoclimax theory (Clements): all vegetation within
a region will converge to the same vegetation type,
which is regulated by homeostasis.
• Polyclimax theory (Tansley): there are a number of
types of climax within a region, each of which reflects a
different soil type, local climate, etc.
• Climax-Pattern theory (Whittaker): Succession will
result in a continuum of climax types, varying along
environmental gradients.
• Nonequilibrium hypotheses: Climax rarely, if ever,
occurs; change is continual. History and random events
are important. Even if equilibrium is theoretically
possible, disturbance is too frequent to allow it.
• Shifting mosaic steady state: Although the vast
majority of sites in a landscape are changing (recovering
from disturbance), the landscape may be in a steady
state. This is because there is, through time, a reasonably
constant portion of the landscape in each successional
stage. A shifting mosaic steady state is a special case of
a Dynamic Equilibrium.
5 General Process of Succession:
46 INSTITUTE FOR ADVANCED STUDIES, JODHPUR
i. Nudation: This is development of a bare area without
any form of life. This area may develop due to number
of causes such as landslide, erosion, deposition or other
catastrophic agency. The cause of nudation may be:
a. topographic: Due to soil erosion by gravity, water or
wind, the existing community may disappear. Other
causes may be seoposition of sand etc., landslide,
volcanic activity and other factors.
b. Climatic: Glaciers, dry period, hails and storm, frost,
fire etc., may also destroy the community.
c. Biotic: Man is most important, responsible for
destruction of forests, grasslands for industry,
agriculture, housing etc. Other factors are disease
epidemics due to fungi, viruses etc. which destroy the
whole population.
ii. Invasion: This is the successful establishment of a
species in a bare area. The species actually reaches this
new site from any other area. This whole process is
completed in following three successive stages:
a. Migration (dispersal): The'seeds, spores, or other
propagules of the species reach the bare area. This
process, known as migration, is generally brought about
by air, water, etc.
b. Ecesis (establishment): After reaching to 'new area,
the process of successful establishment of the species, as
a result of adjustment with the conditions prevailing
there, is known as ecesis. 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. Thus as a result of ecesis, the
individuals of species become established in the area
c. Aggregation: After ecesis, as a result of reproduction,
theindividuals of the species increase in number, and
they come close to each other. This process is known as
aggregation.
iii. Competition and coaction: After aggregation of a
large number of individuals of the species at the limited
place, there develops competition (inter-as well as
intraspecific) mainly for space and nutrition. Individuals
of a species affect each other's life in various ways and
this is called coaction. The species, if unable to compete
with other species, if present, would be discarded. To
withstand competition, reproductive capacity, wide
ecological amplitude etc. are of much help to the species.
iv. 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. Due to all these the
environment is modified, becoming unsuitable for the
existing community which sooner or later is replaced by
another community (seral community). The whole
sequence of communities that replaces one another in the
given area is called a sere, and various communities
constituting the sere, as seral communities, seral stages
or developmental stages. The pioneers are likely to have
low-nutrient requirements, more dynamic and able to
take minerals in comparatively more complex forms.
They are small-sized and make less demand from
environment.
 
UNIT 10: ECOLOGICAL PRINCIPLES 
v. Stabilization (climax): Finally, there occurs a stage
in the process, when the final terminal community
becomes more or less stabilised for a longer period of
time and it can maintain itself in equilibrium with the
climate of the area. This final community is not replaced,
and is known as climax community and the stage as
climax stage.
6. Hydrosere (Succession in water body):
1. Phytoplankton stage: Blue green algae, diatoms,
bacteria
2. Rooted submerged stage: Hydrilla, vallisneria,
potamogeton, utricularia.
3. Rooted floating stage: Trapa, Nymphea, Lemna.
4. Reed swamp stage: Typha, Sagitarria, Phragmites
5. Sedge meadow stage; Cyperus, Carex, Juncus.
6. Woodland stage: salix. Populus, Almus
7. Forest Stage: Almus, Acer, Quercus.
7. Lithosere(Succession on barren rock):
1. Crustose Lichen stage: Rhizocarpon, Rinidina,
Lecanora
2. Foliose Lichen Stage: Parmelia, Dermatocarpon.
3. Moss Stage: Polytrichum, Tortula, Grimmia.
4. Herb Stage: Artisida, Poa, Festula.
5. Shurb Stage: Rhus, Phytocarpus
6. Forest Stage: Mespphytic trees.
8 Heterotropic/ Microbial/ Microsere Succession:
INSTITUTE FOR ADVANCED STUDIES, JODHPUR  47
It begins on habitat rich in organic matter; autogenic
succession. It involves opposite progression i.e. High
energy content (more organic matter) to nil energy
Primary colonizer(Feeds on senescent Tissues)
Stage 1 (a) Weak parasites. Eg, Bacteria.
Stage 1 Primary saprophytic sugar fungi, living on
sugars and carbon compounds. Eg. Mucoraceous
phycomycetes.
Secondary colonizers(Feeds on dead tissues)
Stage 2. Cellulose decomposers and associated
secondary saprophytic sugar fungi sharing products of
cellulose decomposition. Eg. Ascomycetes and some
mucorales
Stage 3. Lignin decomposers and associated fungi Eg.
Basidiomycetes.
Climax Concept in Succession
The final, tenninal and more or less stabilised
community in succession that is able to establish some
sort of equilibrium with the environmental conditions of
that area, was termed climax by Clements (1916). The
subject of climax has been much controversial, and is
reviewed from time to time by many ecologists.
According to Clements (1916, 1935), climax has the
following three principal characteristics:
Unity: Climax is a unit, and index of the climate of area.
Life or growth forms of plants indicate the climate type.
Unless all the species are not taken as an organised unit,
climax would not indicate the climate.
Stability: The form of climax community is more or less
stable with the climate. This climax community can not
be replaced through competition by any other group of
species. In other words, according to Clements, in a
particular climate area there may develop climax
communities only with a few characteristic dominant
species.
Origin and phylogenetic relations: Climax community
is to be treated equivalent to an organism, like which it
takes birth, grows and develops and becomes mature. In
this organisrnicconcept of climax community, Clements
went to the extent of designating it a superorganism. In
any climatic region, the developmental stages of the
climax community have their own characteristics, which
reflect the type of climate. Similar to the development of
organism with changing age, climax communities have
also simultaneously undergone changes with changing
UNIT 10: ECOLOGICAL PRINCIPLES 
climate. Thus phylogenetic relations may be established
between different climax communities of the world.
There have been put three popular theories about the
climax concept in ecology.
Monoclimax theory:
As evident from the above account, F.E. Clements
emphasized the importance of only climate in the
stabilisation of the climax community. According to the
monoclimax theory, within a given region all land
surfaces eventually tend to be occupied by a single kind
of community which is climax. The climax is determined
by the regional climate. Given a stable climate the
climax community is stable indefinitely. However, his
views were strongly criticised by many ecologists.
Cowles, disagreeing with the idea of stability, stated that
'equilibrium state is never reached and succession is
infact a variable approaching a variable rather than a
constant.
According to Cooper, all progressive as well as
retrogressive changes in communities reflect succession.
Thus he considered climax state as stage of minimum
change, rather than finally changed state of succession.
Climax may not be taken equivalent to an organism,
from which it differs in several respects. Therefore,
community may not be considered as a unit
It is common to observe that in the same climate, climax
communities are different. Because, depending upon
their primary stages and habitat characteristics, their
successional stages are also different. Also, in same
climate, a lithosere and hydrosere beginning with
different pioneer communities and proceeding through
different successive developmental stages, might
develop into similar climax communities. Similarly, it
may not always be true that successions with similar
pioneer communities and seral communities would
develop into similar climax communities.
The most objectionable point and subject of controversy
about the climax (with Clements view point) has been its
intimate relationship with the climate. In many
conditions, in an area of uniform climate, it is common
to observe different types of climax communities
according to the soil, topography and other factors.
Under such situations it would not be correct to consider
only the climate as determining factor in climax. For
example, Hult (1885, 1887) in Finland, long before the
Clements concept of climax, described as many as seven
different habitat types under similar climatic conditions.
Tansley (1935) too disagreeing with Clements put forth
his concept that climax communities are infact
controlled by more than one factor and not solely by
climate. In this way, the climatic-climax concept of
Clements became popularly known as monoclimax
theory, although Clements did not use this term.
Polyclimax theory:
It is indicated above, how the climatic-climax concept of
Clements was strongly opposed by ecologists including
Tansley (1935), who believed that climax is controlled
by many (and not one climate) factors. Accordingly
48 INSTITUTE FOR ADVANCED STUDIES, JODHPUR
Tansley's concept became popularly known as
polyclimax theory. Clements, although agreed with the
possible control of factors other than climate on climax,
but he thought that these communities would sooner or
later develop into climatic-climax types. Consequently,
with an attempt to accommodate these stages in his own
hypothesis, Clements introduced in literature a number
of new terms. These are as follows:
The stage in succession just preceding the climatic
climax community was called a sub-climax. The
community, which became stabilised at any of the seral
stages of succession due to microclimate, or effects of
factors as soil, fire etc. was called a sereclimax.
A disclimax (disturbed climax) was called the
community which became more or less stabilised due to
recurrent disturbance by man or other biotic factors,
which prevent the establishment of climatic climax
community in the area. At the same place, under similar
climatic conditions, at smaller distances, due to
differences in water and soil characters, there may
develop different climax communities. Under such
circumstances, Clements (1936) coined two more terms-
(i) preclimax. It is the community with life forms lower
than those in the expected climatic climax, which might
develop in a habitat drier than that of the expected
climatic climax, and (ii) postclimax, a strip of
community with life-forms higher than those in the
expected climatic climax and which develops under
more moist or colder habitats.
As a result, followers of Clements coined a number of
new terms. For instance, coclimax, superclimax,
quasiclimax, paraclimax, anti-climax, conclimax,
peniclimax, metaclimax, pseudoclimax etc.
The only way to come out of the jargoons of Clements
tenninology was the polyclimax theory (Tansley, 1935,
1939). There is evidence that even under primeval
conditions it was difficult to find large areas of uniform
vegetation. According to this theory the climax stage
may be controlled by any factor of the environment and
not only by climate. Accordingly, the climax stage is to
be named, depending upon the nature of the factor in
stabilisation. Thus, in addition to climatic climax,
controlled by climate, there may develop:
Edaphic climax: On an underdeveloped soil, it develops
due to edaphic effects.
Biotic climax: Developed due to biotic disturbances.
Some ecologists make further categorisation into
anthropogenic climax (due to man), grazing climax (due
to grazing effects) and zootic climax (due to animals).
Topographic climax: Due to differences in topographic
factors at mountains, hills, mounds etc.
Fire climax: Due to repeated effects of fire.
According to Daubenmire (1968), if one accepts
monoclimax theory, it means to admit that other factors
of the environment are of secondary value.
Climax-pattern hypothesis
 
UNIT 10: ECOLOGICAL PRINCIPLES 
This concept about climax was put by R.H. Whittaker
(1953) who rejected the classification approach. On the
basis of community gradient analysis, it was postulated
that the communities developing at a particular place are
in accordance with all the factors of environment. In this
area, the vegetation may not be divided into smaller basil
units. In fact different climax-types are orderly organised
parallel to the environmental gradient According to this
hypothesis, there is, in a sense only one big community
that changes according to soil, slope, and other habitat
factors.
Biome
As pointed out earlier in the discussion on concept of
climax, in certain regions, besides the climax
community, there are also associated some other
communities in earlier stages of succession as well as
communities of a different type, controlled by special
local conditions (soil and topographic conditions).
Climax communities are different for different
G1. THE CONCEPT OF ECOSYSTEM 
1. Preliminary Definitions
In any natural system, two major components can be
distinguished:
ƒ a set of physical and chemical conditions, relatively
homogeneous in a given geographic area, at a time t
(climatic conditions, for example, clearly varying
according to the hours of the day and the seasons), which
constitutes the biotope;
ƒ a certain number of living things inhabiting this
biotope-bacteria, fungi, plants, animals-which we call
biocenose.
For the ecologist, the basic unit of the biocenose is a
Population, which is defined as a set of organisms be-
longing to the same species. A settlement is the set of
populations. There is some ambiguity between the terms
‘settlement’ and 'biocenose'. In current usage the term
biocenose' is reserved for the group of organisms that
habit a biotope, whereas the term 'settlement' specifies
the nature of the species considered. We therefore talk
of, for example, a settlement of herbivorous mammals,
Gramineae, or ichthyophagous fish.
2. The Ecosystem-A Network of Interactions
From these definitions, we could attempt, as have num-
erous authors since Tansley (1935), to define ecosystem
by the following equation:
Ecosystem = Biotope + Biocenose
This formula now seems totally wrong, as can be seen
from the following example (admittedly a caricature) of
a Zoological park in Northern Europe.
For this park, a meteorological station can provide year-
round data on climatic conditions such as wind speed
and direction, air temperature, and rainfall. These consti-
tute the parameters of a biotope, which we call abiotic
parameters, because they are not influenced by the living
things inhabiting the system. On the basis of these
measurements, we observe that the temperature in winter
is -20°C, that the wind blows at 100 km/h, and that
INSTITUTE FOR ADVANCED STUDIES, JODHPUR  49
geographical areas. In 1939, E.E. Clements and V.E.
Shelford described the biome system for dealing with the
geographical distribution of communities. They defined
biome as "biotic community of geographical extent
characterised by distinctiveness in the life forms of the
important climax species". The biome is not based on the
species of plants and animals present but rather it is
based, on the life form of the most important plants (i.e.
trees, grasses, scrubs) that give to the landscape its
special character. The biome is viewed as a biotic
community which, in this sense, means that the plants
and animals are considered together as an interacting
unit that gives to it a special look. The chief character of
a biome is depicted by climax communities with their
dominant life forms. Seral communities within the
region of a biome are considered part of that biome.
Thus biomes, a bigger unit than community, constitute
the great regions of the world distinguished on an
ecological basis, such as tundra biomes, forest biomes,
grasslands and deserts etc.
precipitations occur for several months in the form of
snow.
Using a guide to identify fauna, we can easily identify
the following species in this park: hyena, antelope,
giraffe, elephant, lion, and rhinoceros. Does this mean
that such animals normally live in a geographic area in
which the temperature is -20°C and in which it snows in
winter, since we have not observed any seasonal
migration among them? Obviously not!
In fact, the error introduced by the equation given above
arises from the mathematical sign +. Ever since we
learned to count, we have known this sign and known
that 1 + 1 = 2, whereas that is not the case in an ecosys-
tem, which cannot be reduced to the simple addition of a
biotope to a biocenose.
In the case of the zoological park, we know very well
that these animals, mostly of African origin, subsist only
with human intervention. The park managers place them
in heated cages during winter and feed them artificially
throughout the year. As Margalef (1975) has remarked,
if we were to open the cages in a zoo, the biocenose
would very rapidly be simplified, being limited to one
lion, obese from overeating but destined to die of
hunger, and one rhinoceros, which is inedible.
UNIT 10: ECOLOGICAL PRINCIPLES 
Left to itself, the biocenose of a zoological park is not
viable, unlike a natural biocenose. The only difference
between the two types of communities lies in the
artificial input of matter (feed) and energy (calorific and
chemical) by humans in the case of a zoo. In a natural
environment, the biocenose has only solar energy and
local food resources at its disposal.
Because of the imposition of multiple interactions
between biotope and biocenose, detailed in the rest of
this work, and the network of biocenose among organ-
isms, a functional system or ecosystem can be described
as in the diagram in Figure 1.
3. Concept of Eco-complexes
The idea of the functional network of Interactions is in
fact found in living matter at all levels of organization. A
cell is not simply the addition of macromolecules; an or-
ganism is not the sum of cells or organs; a population is
not just the juxtaposition of individuals that have no rela-
tion with each other or with neighboring populations.
The ecosystem is not moreover the highest level of
integration known in nature. Ecosystems are in effect
largely interdependent. For example, a lake is an ecosys-
tem of stagnant water and the sloped basin that
surrounds it. On the basin, two types of ecosystems can
be distinguished:
ƒ terrestrial ecosystems (prairies, forests, moors, etc.)
ƒ one or more aquatic ecosystems of running water (one
or several affluent rivers).
G2. Tropic Levels (Structure of community)
Trophic levels are the feeding position in a food chain,
the sequence of consumption and energy transfer
through the environment
1. Producers (first trophic level) are mostly
photosynthetic autotrophs (self feeders), such as plants
and cyanobacteria, that extract energy from sunlight and
chemical building blocks from the inorganic world, e.g.,
water, carbon dioxide gas, etc.
2. Consumers (secondary trophic level) are organisms
that consume live organic matter and transform it to
ensure the constitution and functioning of their own
organs. They are conventionally divided into three major
categories, herbivores, carnivores, and parasites.
50 INSTITUTE FOR ADVANCED STUDIES, JODHPUR
The various ecosystems found in a territory are not
entirely independent of one another. Interactions also
occur between them in their physical and chemical
factors. Similarly, many species are not limited to a
single ecosystem and their lives may depend on different
environments that are more or less far apart. In each
territory, there is a set of interdependent ecosystems,
exchanging matter and living organisms with one
another. Such sets of interdependent ecosystems
exchanging matter and living organisms between them is
called an eco-complex. (Blandin and Lamotte, 1988)
In fact, by the pressure of movements of air and water
caused by physical processes that will be described later,
and because of movements that most living things are
capable of (migration, transport of pollen, seeds and
fruits), all ecosystems are closely interrelated in the vast
network called the biosphere.The biosphere can be
defined as the space of the planet covering all living
things and in which life is always possible. The term
therefore covers, partly, the three major compartments
that make up the earth: the lithosphere, the hydrosphere,
and the atmosphere.
a. Herbivores: They are primary consumers (C1) as
they consume plants; they make up the second level of
the food chain. Their size varies widely, from the small
organisms that nibble on algae, to most insects and
rodents, to giraffes and elephants.
b. Carnivores: They are consumers at several degrees.
Carnivores most often capture prey. By definition they
are therefore predators. They thus transform animal
biomass. According to the degree of predation, we have:
ƒ Consumers of herbivores, or secondary consumers
(C2), for example, a mole that eats a beetle grub, a lion
that captures a gazelle, or an owl that eats a field mouse.
ƒ Consumers of carnivores or tertiary consumers (C3),
for example, an adder that eats a shrew (which is an
insectivore). As they ingest predators, they are often
called super predators.
 
UNIT 10: ECOLOGICAL PRINCIPLES 
It is possible to distinguish trophic levels of the fourth,
fifth, even sixth order, especially in the marine environ-
ment, where food chains are long. The same animal may
belong to several levels. A buzzard is a simple C2 when
it feeds on crickets, which it very commonly does in the
summer. It becomes a C3 when it captures a mole or a
shrew, and it is a C4 when it catches an adder, which
also consumes insectivores.'
c. Parasites: Parasites run along the food chain. A
raptor, which is a C4, may carry ticks (parasite acarids)
that suck its blood. These arthropods are therefore C5.
But the ticks may also be victims of a superparasitism
because they often harbour pathogenic bacteria or virus.
That is how they transmit pyroplasmosis to their hosts.
The bacteria, which are thus C6, may in turn be subject
to viral parasitism by bacteriophages. These viruses are
C7.
3. The Detritivores and Decomposers are Specific
Consumers: It is not clear how organisms that slowly
decompose cadavers and plant wastes should be
categorized. As they use dead organic matter to ensure
their metabolism, growth, and reproduction, they behave
as consumers. The result of their action is the
mineralization of various elements. They thus playa
significant role in the cycling of matter, which allows
them to recover, in passing, the energy they need to live.
This cycling is done in two stages and involves two
categories of living things:
ƒ detritivores, which feed on the remains of dead
organisms fallen on the soil, and
ƒ decomposers, which complete the transformation
begun by the detritivores in the process of mineralization
that makes organic matter into mineral matter.
a. Detritivores:
As their name suggests, detritivores represent all indi-
viduals that feed on animal or plant detritus. Their main
action, which is the first step in the transformation of
dead organic matter, is the fragmentation of debris into
finer elements that other transformers find in their
excrement or in faecal pellets.
There are several categories of detritivores according to
the place or type of consumption.
ƒ NECROPHAGES feed only on animal corpses. This
is the case of necrophores, coleopteran insects often
found on carcasses of birds, mammals, or other animals
after a few days, along with other adult insects or with
fly maggots.
ƒ COPROPHAGES are organisms that eat excrement.
The example of dung-beetles (coleopteran insects) is
well known.
G3. Ecological Pyramids
Ecological pyramids are the graphical representation of
tropic structure and function at successive tropic level of
an ecosystem which may be shown in terms of their
number, biomass or energy content.
INSTITUTE FOR ADVANCED STUDIES, JODHPUR  51
ƒ SAPROPHAGES consume decomposing plant
elements. They include woodlice (terrestrial
crustaceans), which find their food under rotting bark.
ƒ GEOPHAGES are soil animals that play an essential
role, in humification. The best known are the
earthworms, which 'eat their way through the soil'. While
doing this, they digest the fragments of plant matter
buried in or fallen on the soil.
b. Decomposers
Once the first stage of degradation of dead organic
matter is completed by detritivores, the microscopic
organisms-bacteria, fungi, or protozoa-accomplish the
second stage of this transformation. These micro-
organisms are responsible for actual mineralization. By
aerobic processes in some cases and by anaerobic
processes in others, they trigger fermentation.
Fermentation breaks down the organic molecules still
present in the soil in the form of glucids, protids or
lipids, such as humic acids, into much smaller mineral
molecules, the mineral salts.
4. Role of Detritivores and Decomposers in the
Functioning of Ecosystems
By successive stages, the joint heterotrophic metabolic
activity of detritivores and decomposers brings about the
remineralization of organic matter that was created in the
first place by the producers and that passed through all
the trophic levels of the food chain.This role is important
in the functioning of ecosystems since it corresponds to
the completion of the cycle of matter. It again makes
available to plants the nutrients they need for
photosynthetic activity.
In terrestrial ecosystems, the production-mineralization
cycle has only a few stages, since on an average 80% of
the plant biomass produced is not consumed by
herbivores. Such is not the case in aquatic ecosystems,
where most of the plant biomass produced by unicellular
algae is ingested after it is formed. In the soil,
detritivores and decomposers play an often silent role in
the functioning of ecosystems, that of initiating new food
chains. While mineralizing organic matter, they produce
their own biomass by heterotrophy. They are preyed
upon by several predators that are part of the trophic
network of soil.
Finally, the cycling that they accomplish is essential for
soil fertility. We will see in the next chapter that this
mineralization may happen sooner or later and that it
contributes to the proper functioning of what is called
the argilo-humic complex.
UNIT 10: ECOLOGICAL PRINCIPLES 
1. Pyramid of Numbers: To construct a pyramid of
numbers, the number of individuals present in each
trophic link of a given environment is calculated, in a
prairie in the example described in Figure. On paper, the
organization looks like a triangle pointing upward.
Each level is drawn as a rectangle the size of which is
proportionate to the number of individuals calculated at
that level. All the rectangles have the same height; only
the base changes. The longer the chains, the more
pointed and tall the pyramid. This imaginary
representation enables us to understand some elementary
principles at a glance.
ƒ The number of individuals: When we go upward
from the base of a trophic system to its summit, we
observe that there are progressively fewer individuals, in
the predator chains, from one link to the next. The
largest rectangle corresponds to the primary producers
(P1). There are a large number of plants, especially in a
prairie, even though there are few species. The
population of herbivores (C1) is much smaller, which is
expressed in a smaller rectangle. That of carnivores (C2)
is still smaller and that of higher predators (C3) is
reduced to a few units.
ƒ The size of individuals: Although the pyramid of
numbers does not show the size of individuals, it is
obvious that the reduction of numbers, as one goes up
the pyramid, is compensated by an increase in size.
Generally, the herbivores are larger than the grasses they
consume and the carnivores are larger than their prey.
ƒ Prolificity: The grasses have a tremendous power of
reproduction, since each plant can produce a profusion
of grains. The grasshoppers that consume them, in a very
simple imaginary chain, lay only a few hundred eggs.
The amphibians that eat grasshoppers in turn produce a
few tadpoles. The reptiles that consume amphibians have
only a few descendants. As for the blackbreasted snake
eagle, which eats only reptiles, it lays only one egg a
year and its period of sexual immaturity is very long (3-5
years). This allows us to understand why predators are
the first animals that tend to disappear in cases of
disequilibrium.
The study of pyramids of numbers, because it does not
show directly the criteria of mass and size and em-
phasizes only the number of individuals, must be com-
plemented by other pyramids.
Note: The pyramid of numbers for parasites is the
inverse of that for predators. The number of parasites
that infest a host is most of the time very high. For
example, when a dog has fleas, it generally has many
fleas. It is the same for lice infesting human heads.
These parasites are themselves victims of
hyperparasitism by acarids or viruses. Here also, the
numbers of parasites are higher than, those of their hosts.
Thus, the inverted pyramid has narrow bottom (the
primary host) and a very wide top, (the viruses).
2. Pyramid of Biomass
52 INSTITUTE FOR ADVANCED STUDIES, JODHPUR
a. In terrestrial ecosystems: For this other
representation of biocenoses, we draw, pyramid in which
the superposed rectangles have a surface proportionate to
the biomass of individuals present in each link. For
grassland and forest it is upright.
For parasites, contrary to the preceding representation,
the pyramid of biomass is not inverted. Even though
there are many in relation to the host, the parasites have
a low biomass. Thousands of fleas have a negligible
weight in relation to that of the dog that carries them.
b. In aquatic ecosystems: Even though phytoplankton
has a rate of renewal higher than that of the zooplankton
that feeds on it, the level of primary producers (base of
the pyramid) is narrower than that of the herbivores
(zooplankton). The rest of the pyramid has the usual
shape (Fig.).
The phytoplankton is in fact consumed to the extent that
it is produced. Its biomass therefore does not
accumulate. In the sea, this is frequently the case
because the zooplankton consumes all the phytoplankton
produced in a day. The same phenomenon occurs in the
lakes at the end of spring, where it assists in a
proliferation of zooplankton. This over-consumption
thus leads to a decrease in the turbidity of the water,
called the clear water period. The pyramid of biomass is
thus more useful than that of numbers. All the same, the
pyramid does not take into account the presence of water
in the tissues. For the same dry weight, plants are much
more hydrated than animals. It is also necessary to
calculate the biomass in dry matter and not in living
matter.
3. Pyramid of Energies: The pyramid of energies is the
most satisfactory representation because it is the only
one that really indicates the metabolic losses. According
to the laws of thermodynamics, it has the same shape as
the pyramids of numbers and biomass. It is upright
(Fig.). To construct this pyramid, the biomass of tissues
must be converted into energy values. Generally, the unit
used is the kilojoule. Sometimes, the older measure,
kilocalorie, is also used, and 1 kcal is equivalent to 4.18
kJ.
4. Decomposers and their Place in the Pyramids:
What is lacking in the schematic representation of Elton
is the place of detritivores and decomposers. It must be
noted that their biomass is lower than that of primary
and secondary producers, which develop living organic
matter. Still, they have a very high metabolism. Figure
 
UNIT 10: ECOLOGICAL PRINCIPLES 
shows that, for Silver Springs, the decomposers have an
energy potential of 460 kcal, higher than that of herbiv-
ores, which have an energy potential of only 383 kcal.
Much more than their quantitative importance is the role
of decomposers in the buckling of the biomass cycle,
which is the capital in the framework of the ecological
pyramids. The primary producers to create their own
matter, must draw from the soil the mineral salts located
around their roots. But these nutrients are not
inexhaustible. The proof of this is the enormous quantity
of fertilizers that farmers must use to improve their soils
and replace the soil minerals used up by harvests. All
living beings in the ecosystem, by their metabolism,
eliminate a significant quantity of energy dissipated in
the form of heat (Fig.).
When primary producers and various consumers die,
their carcasses are exploited by detritivores and
decomposers. Ultimately, the carcasses are mineralized
in the form of mineral salts that are again absorbed by
plants. This turnover is considerable if one limits it to
energy references.
G4. Food chain and food web
1. Food chains: It’s a sequence of organisms through
which energy and nutrients move. The general sequence
of food chain is: From producer to primary consumer to
secondary consumer to tertiary consumer to quaternary
consumer to detritus feeder. Primary consumers are
herbivores. All others are meat eaters (carnivores) or
both eaters (omnivores). For instance, a typical food
chain in a field ecosystem might be:
grass Æ grasshopper Æ mouse Æ snake Æ hawk
1.1 Grazing vs. Detritus Food Chains: Grazing food
chains based on producers (photosynthesis) while
Detritus food chains based on detritovores (breaking
down detritus and wastes). Grazing food chains are
important in marine and aquatic ecosystems while
Detritus chains play important in forest litter.
2. Food web: The real world, of course, is more
complicated than a simple food chain. Food chains are
not found isolated; rather they are interconnected to form
food webs.“The greater number of alternative channels
INSTITUTE FOR ADVANCED STUDIES, JODHPUR  53
5. Biological Magnification: It is the tendency of
pollutants to become concentrated in successive trophic
levels. Often, this is to the detriment of the organisms in
which these materials concentrate, since the pollutants
are often toxic. In order for a pollutant to biomagnify,
the following conditions must be met:
ƒ The pollutant must be long-lived.
ƒ The pollutant must be concentrated by the
producers.
ƒ The pollutant must be fat-soluble.
Biomagnification occurs when organisms at the bottom
of the food chain concentrate the material above its
concentration in the surrounding soil or water.
Producers, as we saw earlier, take in inorganic nutrients
from their surroundings. The problem comes up when a
pollutant, such as DDT or mercury, is present in the
environment. Chemically, these pollutants resemble
essential inorganic nutrients and are brought into the
producer's body and stored "by mistake". This is the first
step in biomagnification; the pollutant is at a higher
concentration inside the producer than it is in the
environment.
The second stage of biomagnification occurs when the
producer is eaten. Not only are they taken up by the
producers, but they are absorbed and stored in the bodies
of the consumers. This often occurs with pollutants
soluble in fat such as DDT or PCB's. These materials are
digested from the producer and move into the fat of the
consumer. If the consumer is caught and eaten, its fat is
digested and the pollutant moves to the fat of the new
consumer. In this way, the pollutant builds up in the fatty
tissues of the consumers.
through which energy can flow, the greater the stability
of the food web and the ecosystem.”
The food webs you see here are grazing food chains
since at their base are producers which the herbivores
then graze on. While grazing food chains are important,
in nature they are outnumbered by detritus-based food
chains. In detritus-based food chains, decomposers are at
the base of the food chain, and sustain the carnivores
which feed on them. In terms of the weight (or biomass)
of animals in many ecosystems, more of their body mass
can be traced back to detritus than to living producers.
3. Energy Flow across the Food Web
UNIT 10: ECOLOGICAL PRINCIPLES 

for the metabolism of carnivores and will be eliminated

When energy passes from one level of production to the
next in a food web, the transfer of energy always
involves enormous losses. Moreover, a flow is defined
for the energy that transits each of the elementary levels
that make up a web (Fig.). This notion of flow differs
from that of energy yields seen earlier.
Thus, at each link (P1, C1, C2, C3, decomposers), the
flow corresponds to the quantity of energy assimilated
by the beings that make up that link. The flow takes into
account not only the energy fixed in the living organic
matter, but also the catabolic losses that enabled this
fixation.
in the form of respiratory losses, R3.
If PS2 represents the energy gained that is added to the
existing biomass, the flow at the level of carnivores is
the following: A2 = PS2 + R3
Flow = Energy fixed by the C2 + Respiratory losses
The process runs in the same manner if the food chain
extends to the level of supercarnivores. In view of the
enormous losses that appear at each trophic level, it is
clear that the food chains are always short. Detritivores
and decomposers intervene also in energy flow across
the ecosystem. It is these organisms that recover the
energy stored in all the unused matter (NU1, NU2,
NU3...). They draw from this energy all that is necessary
for their metabolism and the increase of their biomass,
and like other consumers they lose energy by respiratory
catabolism or fermentation (Rd). As was explained in the
preceding paragraphs, each trophic level in the food
chain is the site of a significant dissipation of the energy'
it receives in the form of heat. This implies that the
structure of biocenoses is graded.

The most significant of these energy losses occurs at the

3.1. Flow at the Level of Primary Producers (P1): For
plants, the flow is equivalent to the real photosynthesis
(assimilated energy), which we have already defined as
the gross production (PB) and which is added to the
energy stored in the existing biomass. In fact, apparent
photosynthesis or net production (PN) does not take into
account respiratory losses of plants (R1). We can thus
write:
PB = PN + R1
Flow = Net production of P1 + Respiratory losses
3.2. Energy Flow at the Level of Herbivorous
Consumers (C1): Only one portion of the plant
production is ingested by the herbivores, which we call
PI1 (the part ingested by primary consumers). The rest is
available to detritivores and decomposers when the
plants die. As it is not used by the herbivores, we call it
NU1.
level of primary production. Photosynthesis uses, by
transformation of photons, only 1 %, on an average, of
the solar radiation for formation of plant biomass. After-
wards, during passage from trophic level n to level n + 1,
these losses continue to occur, although at a lesser rate.
There are two reasons for this. The first reason is that all
the biochemical energy contained in the biomass of a
trophic level is not consumed by the higher level. The
second lies in the fact that because of metabolic losses
the total gross production of a given level is not
available for the higher level. Any trophic relation, like
that of the consumption of prey by a predator, is of an
energy type. The best yields are never 100%, since yield
does not go beyond 10% on average from C1 onward.
We can also say that, in thermodynamic terms,
ecosystems are constantly dissipating energy.

What is consumed (PI1) is not absorbed totally by

digestion. Only a part is assimilated, or A 1 (flow at the
level of C1). The rest is rejected in the form of
excrements or secretions, which is the NA 1 fraction (not
assimilated). We must also take into account respiratory
catabolic losses, R2. The flow at this level is, if PS1
represents the energy fixed which is added to the energy
already stored in the biomass of herbivores:
A1 = PS1 + R2
Flow = Energy fixed by the C1 + Respiratory losses

3.3. Flow at the Level of Carnivorous Consumers

(C2): Only part of the energy fixed by the primary
consumers will be used for the functioning of
carnivorous organisms or secondary consumers. This
part is cal)ed PJ2. As for the first level of consumption,
many prey die of old age and their carcasses will be put
to good use by decomposers. This fraction is called BU2.
Only part of the biomass consumed will be assimilated,
A2. All that is eliminated in the faeces and various
secretions corresponds to NA2. As in the preceding
level, much of the energy fixed by digestion will serve

54 INSTITUTE FOR ADVANCED STUDIES, JODHPUR 
UNIT 10: ECOLOGICAL PRINCIPLES 
G5. ENERGY FLOW AND THE FUNCTIONING OF ECOSYSTEMS
In the flow of energy and inorganic nutrients through the
ecosystem, a few generalizations can be made:
1. The ultimate source of energy (for most
ecosystems) is the sun
2. The ultimate fate of energy in ecosystems is for it to
be lost as heat.
3. Energy and nutrients are passed from organism to
organism through the food chain as one organism eats
another.
4. Decomposers remove the last energy from the
remains of organisms.
5. Inorganic nutrients are cycled, energy is not.
Many forms of energy are indispensable for the
functioning of ecosystems and it is well known that
photosynthesis is not the only metabolic process
sensitive enough to produce organic matter from mineral
elements. However, solar radiation is undoubtedly the
main source of energy flow in most ecosystems.
1. Solar Energy
1.1. Spectrum of Wavelength of Solar Radiation
Solar radiation reaching the upper levels of the atmos-
phere consists of wavelengths between10-4 µm and sev-
eral kilometres. But 99% of the energy is transported by
wavelengths of 0.2 to 4 µm, and 42% is in the visible
portion of the light spectrum, 0.38-0.75 µm..
1.2. Incident Energy
The quantity of incident energy, measured at the top of
the atmosphere, is on an average 350 W/m2, or 30 x 106
kilojoules/m2/day. There are two reasons why a signifi-
cant portion of this energy does not reach the earth's
surface.
The first has to do with ultraviolet (UV) rays, of wave-
length 0.2 to 0.38 µm, that are practically absorbed by
the ozone layer (03) located at a height of 25 km. Even
though these UV rays represent only 9% of the total en-
ergy, the photons they associate with have high energy
and alter organic molecules. That is why, if they were to
completely reach the surface of the earth, all life would
be impossible. It is likely that life first appeared in the
aquatic medium because UV was absorbed while passing
through several millimetres depth of water. The ozone
layer, which is very thin, was formed because of oxygen
produced by aquatic photosynthetic organisms of the
PreCambrian era (algae and some bacteria) and allowed
the appearance of life in terrestrial ecosystems.
The second reason is that air molecules, dust, aerosols,
and water drops that constitute clouds absorb and diffuse
a significant amount of that light in all directions. On an
average, although the values vary considerably with the
altitude, only 50% of the light energy measured at the
top of the atmosphere reaches the surface of the planet.
The energy received is:
ƒ mostly reflected by the receiving surface, this fraction
constituting albedo, or
ƒ absorbed by that surface.
INSTITUTE FOR ADVANCED STUDIES, JODHPUR  55
Infrared rays, of wavelength 0.75 to 4 pm, are especially
absorbed by water, in which they can travel only a few
centimetres. That has three consequences:
ƒ a warming up of the receiving environment inducing
water evaporation;
ƒ movements of air and water caused by the unequal
distributions of these rises in temperature as a function
of altitude and season;
ƒ emission of radiation by the surface and receiving
particles.
This radiation generally consists of wavelengths (10 to
15 µm) greater than those that have been absorbed.
These infrared radiations, called thermics, are rapidly
absorbed by the water drops in clouds and carbon diox-
ide in the air. The absorption results in a warming up of
the atmosphere that is known as the greenhouse effect.
Average energy contribution varies from 80-90 W/m2 at
the poles to 200-290 W/m2 at the equatorial regions.
2. Biological Roles of Light Energy
2.1. Photosynthesis
In the biosphere, most autotrophic living things, those
that can synthesize their own organic matter from
mineral elements and from energy, are plants. Because
of their chlorophyll, these organisms capture solar
energy through a chemical reaction whose universal
equation is simply:
CO2 + H2O Æ (HCHO) + O2↑
The term (HCHO) represents the basal element of a
glucid (Sugars). In the chemical sense of the term, this
oxido-reduction requires the input of 470 kilojoules per
mole (12 g) of carbon assimilated (or 44 g of carbon
dioxide) and (32 g) of free oxygen. Nevertheless,
chlorophyll does not use all the solar energy received by
the surface of the plant. Overall, radiation that can be
used in photosynthesis is only 21 % of the total radiation
hitting the surface of the earth.
2.1.1. Absorption Spectrum and Spectrum of
Chlorophyll Action
White light, visible to our eyes and emitted by a light
source such as the sun or an electric lamp, is composed
of a spectrum of many rays whose wavelength varies
from 0.38 to 0.75 µm. By placing a prism in a light
beam, we can break up the white light into a spectrum of
rays seen in a rainbow: violet, indigo, blue, green,
yellow, orange, red. All these rays do not have the same
efficacy vis-a-vis photosynthesis.
ABSORPTION SPECTRUM: The absorption by
chlorophyll pigments is essentially concentrated in the
blue part of the spectrum, between 380 and 440 nm with
a maximum at 430 nm, and in the red part, between 620
and 680 nm with a maximum at 630 nm. The
wavelengths corresponding to the yellow and above all
the green are practically not absorbed and cross the
leaves without being stopped by the chloroplasts. That is
why vegetation appears mostly green to us.
UNIT 10: ECOLOGICAL PRINCIPLES 
In fact, although chlorophyll is the only pigment to carry
out photosynthesis, other pigments like carotenes,
xanthines, xanthophylls also favour the absorption of
light energy. These pigments play the role of 'reflector'
in transmitting a part of the energy that they themselves
have captured to the chlorophyll by resonance. That
energy leads to the true absorption spectrum, for some
plant cells, which differs from that of strong chlorophyll.
Thus, the absorption spectrum in naturally occurring
algae shows an increase in the use of available light in
the blue area of the spectrum relative to that observed for
the red part. Most of the secondary pigments capture
energy associated with short wavelength light.
ACTION SPECTRUM: Place some plants of Hydrilla
under a glass funnel in a petri dish full of water. The aim
is to recover and measure the quantity of oxygen freed in
the tube by these aquatic plants. In order to do this,
illuminate the Hydrilla with a beam of light, and
successively place in its path coloured filters that
correspond to rays of the white light spectrum.
G6. TRANSFER OF ENERGY AND MATTER
The energy flow caused by the impact of solar radiation
In the earth's surface is the origin of two closely comple-
mentary phenomena. It enables first of all a contact be-
tween plant cells and dissolved minerals in the oceans,
lakes or saline solutions of soil. This leads logically to
cellular assimilation. It then ensures an input of energy
needed by chlorophyll to synthesize organic matter, and
thus living matter.
Plants absorb only about 1.2 % of the sunlight that
strikes them, and about a third of that energy becomes
incorporated in organic molecules that another organism
might eat. These organic molecules constitute the
biomass of the plant. Organisms at higher trophic levels
are able to incorporate about 10 % of ingested biomass
as biomass of their own. Thus a herbivore that eats 10 kg
of plant matter may gain 1 kg in weight.
1. Primary Production
Primary production conveys the rate per unit of time at
which a given quantity of organic matter is formed from
mineral matter and energy. It is expressed in mass of as-
similated carbon per unit of time. Primary production is
carried out by autotrophs or primary producers, which
constitute the first link or trophic level of what is
commonly called the food chain.
The best-known autotrophs are plants that synthesize
their matter by photosynthesis, thus from photons. These
are the phototrophs. There are also autotrophic micro-
organisms drawing the energy needed for their synthesis
from chemical reactions. These are called
chemoautotrophs or chemosynthetic organisms.
2. Productivity
Most authors define productivity incorrectly as being the
quantity of biomass produced per unit of surface area
during a particular unit of time, which amounts to an an-
nual net primary production. Productivity is actually
56 INSTITUTE FOR ADVANCED STUDIES, JODHPUR
Then count the bubbles of oxygen released during a
particular period of time with each colored filter. We
logically observe that we will have the maximum
efficacy of photosynthesis, measured by a maximum of
bubbles, with the rays that have been absorbed by the
chlorophyll. This corresponds to wavelengths in the 630
nm for red light and in the 430 nm for the blue light. The
green rays simply cross the chlorophyll without
'working'.
evaluated by the ratio between the production during a
given time and the biomass present in the habitat.
Productivity = P/B
It is expressed as a unit that is the inverse of a time (t-1).
For plankton, for example, in an aquatic environment
where the primary production is calculatwd in
milligrams of carbon assimilated per cubic metre of
water per hour, productivity is expressed as follows:
Productivity = mg of C/m3/h
mg of C/m3
Its inverse is the rate of renewal of biomass, or turnover:
Rate of renewal = B/P
In the preceding example, the rate of renewal for plank-
ton is:
Rate of renewal = mg of C/m3
mg of C/m3/h
The productivity of plankton is very high. When con-
ditions are favourable, cells can divide in one day. Their
rate of renewal is therefore 1/day. For terrestrial plants,
this turnover is much lower and varies according to the
organ and the plant. It is thus 1.1 /year for leaves and
0.02/ year for wood.
3. Primary Production: Only a small fraction of the
sunlight striking the earth is converted to chemical
energy by primary producers. That sunlight energy that
is converted to chemical energy, over a given period, is
termed primary production.
3.1 Gross Primary Production: Gross primary
production is all of the light energy that is converted to
chemical energy by producers
3.2 Net primary production: Net primary production is
all of the light energy that is converted to chemical
 
UNIT 10: ECOLOGICAL PRINCIPLES 
energy and that is subsequently stored by the primary
producer (i.e., the gross primary productivity minus that
employed to run the primary producer's metabolism).
The ratio of net primary productivity to gross primary
production gives an indication of the cost of keeping the
organism going, with large ratios indicative of relatively
few costs (e.g., algae, ~50%) and smaller ratios
associated with many costs (e.g., complex plants such as
trees, ~10%)
Net Primary Production = Gross P - metabolic losses
(respiration, excretion)
When the conditions allowing photosynthetic activity are
combined, the gross primary production is always
positive. On the other hand, net production can be either:
ƒ positive when the synthesis of organic molecules by
photosynthesis is greater than the nocturnal consumption
and various metabolic losses (respiration, excretion), or
ƒ negative in the contrary case.
3.3 Order of Magnitude of Production
In ecological literature there is a considerable amount of
data on the primary production of various types of eco-
5. Energy Efficiency or Outputs
Energy efficiency is defined, for a trophic level, as the
ratio between the energy released and the energy con-
sumed, that is, the ratio between energy fixed and energy
received.
5.1. Efficiency of Consumption: The consumption
efficiency is the ratio between the energy integrated by a
consumer and energy contained in the field available to
it (Figure). For a predator, this will be that part of energy
INSTITUTE FOR ADVANCED STUDIES, JODHPUR  57
systems. These figures have an extremely limited value,
because they are calculated over a few hours and ex-
trapolated to the year. Their results thus vary consider-
ably as a function of abiotic conditions for the same eco-
system.
Abiotic conditions, in terms of solar radiation, tem-
perature, or rainfall, have highly significant inter-annual
fluctuations. Some authors have attempted an overall
mean estimate for the major types of ecosystems on the
planet. In light of the inevitably approximate nature of
such estimations, the authors obtain varying values.
4. Secondary Production
The term 'secondary production' applies to all biomass
produced per unit of time by organisms called consum-
ers. Consumers only transform organic matter that is
synthesized by the primary producers. The consumers
are thus heterotrophic organisms.
contained in a prey which will be effectively integrated.
All the net production available to the consumer is not
exploited. A herbivore does not use all the net
production of a plant. For example, it leaves behind roots
and woody stems and prefers tender shoots. A carnivore
leaves behind the bones and shell, which contain energy.
The consumption efficiency varies considerably
according to the ecosystems.
5.2. Assimilation Efficiency: Food ingested by a
consumer is subjected to mechanical and chemical
digestive processes during its transit through the
UNIT 10: ECOLOGICAL PRINCIPLES 
digestive tract. Only part of that food, the nutrients,
passes through the intestinal mucus into the bloodstream.
The rest is not assimilated and is eliminated with the fae-
ces. The assimilation efficiency therefore is the ratio be-
tween the energy assimilated (A) of nutrients and energy
E (assimilation) = A/I
It varies considerably according to the nature of the
consumer organisms. In the aquatic environment,
whenthere are massive inputs of mineral elements such
as nitrogen and phosphorus, the phytoplankton grows en
masse, and this growth is called the flowering of water
or 'bloom'. In such conditions, the filtering zooplankton,
which is integrating by filtering the biomass present,
ingests much more organic matter than it needs for its
metabolism.
G7. Limitating factors
Abiotic and biotic regulators determine the distribution
and success of living organisms. There is range of
tolerance for each of the physical and biological
components of environment by organisms and when this
range is passed by any of factor it becomes Limitating.
1 Leibigs Law of Minimum: Under steady state
conditions, the essential material awailable in amount
most closely approaching the minimum needs tends to
become limitating one.
It is strictly applicable under steady state conditions.
There should not be any Factor Interaction: High
concentration of same substance or action of some other
factor other then minimum one may modify rate of
utilization of other. Eg. Utilization of strontium in place
of zinc by mollusks in shell.
2 Shelford Law of tolerance: Not only too little of
something be a Limitating factor but also too much as in
case of factors like heat, light etc. can be Limitating.
Thus organisms have an ecological maximum and
minimum range in between which represents the limit of
tolerance.
ƒ In aquatic organisms oxygen is generally Limitating
factor.
ƒ In open sea nutrients are Limitating factor.
58 INSTITUTE FOR ADVANCED STUDIES, JODHPUR
A large part of the material ingested is liberated in the
form of faeces that settles in the deeper zones, where it is
consumed by other organisms. A large part of this faeces
contains phytoplanktonic cells that are not destroyed by
the digestive enzymes of zooplankton and thus the
growth is even stimulated by nitrogen and phosphorus
liberated during the enzymatic processes.
Note: All that is assimilated does not contribute to the
growth of the consumer's biomass. A significant part of
the energy assimilated is not fixed because it is used to
meet metabolic needs of respiration, excretion, or
homeothermy. Because of this, we can define "another
form of yield, the net production efficiency, which
corresponds to the ratio of fixed energy, that is net
energy (N) over assimilated energy (A):
E (net production) = N/A
5.3. Ecological Efficiency: Ecological efficiency is the
most relevant one to our discussion because it measures
the yield of a trophic relation. It is commonly called
ecological yield or feed efficiency or ecological growth
yield. It is the only ratio that indicates the energy losses
undergone when passing from one feed level to another.
It thus has a considerable advantage for the purpose of
understanding the functioning of biocenoses. It is
defined as the ratio between energy fixed (N) in the net
production of a consumer and energy ingested (I)
contained in the feed consumed.
E (ecological) = N/I
Ecological efficiency is, for example, the ratio between
the net production of a predator and the net production
of the prey captured. It involves the ratio between energy
values and not between weights of matter.
ƒ Generally nitrates and phosphates become limitating
in agricultural lands/fields.
ƒ Reproduction is usually critical period when
environmental factors are most likely to be Limitating.
3 Factor Compensation: Organisms in nature adapt
themselves and may even modify the physical
environment so as to reduce the limitating effect of
temperature, light, water etc. Such factors compensation
is effective particularly at community level but also
occur within species. Eg. Ecotypes (Locally adapted
populations/races).
 
UNIT 10: ECOLOGICAL PRINCIPLES 

G8. Biogeochemical cycles in nature (Chemical Cycling in an Ecosystem)

In previous sections we have emphasized the importance

of transfers of matter and energy within the ecosystem.
Each living organism represents potential energy. All its
organic constituents, glucids, protids, and lipids, liberate
a significant amount of energy when they are catabolized
through respiratory mechanisms or fermentation, while
they return to the mineral state. On the other hand,
mineral salts are fixed by various autotrophic living
things to elaborate new cells and keep them functioning.
As for heterotrophic individuals, parasites or predators,
they draw the constructive elements of their molecules
from their organic environment and not from their
mineral environment.
Thus, matter constantly passes from a mineral state to an
organic form and vice versa. The producers manufacture
and store organic matter while the decomposers ensure
the return to the mineral state. In this gigantic and
constant cycle, each element participates in its own
biogeochemical cycle. We will study those of carbon, ni-
trogen, phosphorus, sulphur, and oxygen, which are of
greatest interest, but the cycle of other elements also
deserves some attention.
1. Carbon Cycle
The most important of the chemical elements character-
izing the living world is' carbon. In the mineral form, it
is present on the globe in carbon dioxide, hydrogen car-
bonates, and carbonates. In the organic form, it consti-
tutes glucids, protids, and lipids.
1.2. Biochemical and Physicochemical Exchanges
between the Three Reservoirs
a. Between the Atmosphere and the Continental
Biosphere: The exchanges between these two sources
are essentially linked to the activities of living things.
The primary producers, autotrophs by photosynthesis or
chemosynthesis, fix CO2 to ensure the synthesis of their
organic constituents. The consumers then transfer these
constituents into various food webs. Each year, by
means of these biochemical exchanges, autotrophs keep
up a gigantic flow of carbon of the order of 100 Gt,
which is extracted from the atmosphere (Fig).
On the other hand, other biochemical exchanges balance
these extractions by emitting into the atmosphere an
identical flow of 100 Gt of carbon (Fig. 89). These re-
leases of mineral carbon result from catabolism of all
living things. To carry on their various functions, these
beings consume energy, which is provided by cellular
respiration for animals as well as for plants, or by
fermentation. In the soil, the most common mineralizers
are the lactic, alcoholic, and butyric fermenting agents.
These decomposers enrich the atmosphere in CO2.
b. Between the Atmosphere and the Oceans: The flow
of carbon between the air and the oceans is equally

1.1. The Three Natural Reservoirs of Carbon

On the global scale, there are three reservoirs of carbon:

the atmosphere, the oceans, and the continental bio-
sphere. Only the atmosphere contains purely mineral
carbon. In the other two reservoirs the two forms of
carbon are associated: the organic form in the biomass
of living things and the mineral form in hydrogen
carbonates or carbonate sediments.

a .Carbon Dioxide Stored in the Atmosphere: Stored

carbon dioxide occupies around 0.03% of the volume of
the atmosphere. It is estimated at 700 gigatonnes (1 Gt
= 109 t). Its equilibrium is regulated by constant
exchanges between the three reservoirs.
b. Carbon Stored in the Oceans: Living things in the
seas and oceans represent only a small part of the
carbon that is in the waters. Most of the carbon is
present in the water in the form of soluble elements,
HCO3- or monohydrogen carbonates, CO2 and
balanced and of the same order. Physicochemical
carbonate ions CO32-. The carbon stored in the surface
exchanges at the interfaces cause a release of 100 Gt of
waters is estimated at 600 Gt and the water in deep seas
C02 into the atmosphere and an identical absorption of
is estimated at 39,000 Gt. Over geological time,
100 Gt of this gas by the water. The quantity of carbon
sedimentation and compaction led to the formation of
dioxide dissolved in the oceans is around 50 times as
carbonate rocks such as chalk or dolomites, which
much as that present in the atmosphere. Also, the entire
represent a formidable reserve of 30,000,000 Gt.
atmospheric CO2 is renewed every 8 years by exchanges
c. Carbon Stored in the Continental Biosphere: Carbon
that conform to the following equilibriums between
in the continental biosphere is estimated at 700 Gt for
these two reservoirs.
living things and 1100 Gt for the soil, totalling 1800 Gt.
In addition, there is the carbon contained in fossil fuels,
The first equilibrium, with reversible reactions, is
such as petroleum, coal, and peat, of around 6000 to
established between dissolved carbon dioxide and
10,000 Gt, not counting bituminous schists, which may
constitute sizeable reserves.

INSTITUTE FOR ADVANCED STUDIES, JODHPUR  59
UNIT 10: ECOLOGICAL PRINCIPLES 
hydrogen carbonate ions (HCO3-), which may represent
90% of carbon in the oceans.
CO2 + 2H2O↔ H3O+ + HCO3-
The second is established between hydrogen carbonates
and carbonate ions (CO3-2)
HCO3-+ H2O↔ H3O+ + CO3-2
c. Human Activities and Carbon Emissions: For our
industrial, domestic, and agricultural activities, we
constantly burn increasing amounts of fossil fuels
(petroleum, coal) in boilers and motors. We exploit
chalk, incinerating it to fabricate cement or lime. Certain
crops such as rice harbour symbiotic root bacteria that
are responsible for a significant emission of methane.
Moreover, because of the digestive activity of our
herbivorous livestock, the quantity of methane released
into the air has quadrupled in the past few decades.
Ultimately, human activities are responsible each year
for the emission of 5 Gt of CO2.Some 2 Gt should be
added to that for the consequences of deforestation in the
tropical forests, which limits their photosynthetic role.
Two thirds of this additional 7 Gt of human origin are
observed by the ocean and the soil. Assessment of this
cycle thus makes it clear that there is an annual increase
of 2 to 3 Gt in atmospheric carbon.
12.1.3. Fossilization of Organic Carbon
In beat bogs, the activity of mineralizing micro-
organisms is interrupted by cold, humidity, and the
acidity of the medium. The organic residues, still rich in
energy, accumulate in the form of peat. In the course of
geological time, an analogous mechanism of
carbonification has led to formation of coal from buried,
partly decomposed plants. Petroleum is similarly formed
60 INSTITUTE FOR ADVANCED STUDIES, JODHPUR
Calcium carbonates and magnesium carbonates, which
are poorly soluble, precipitate and accumulate in the
sediments. Over geological periods of time, they thus
store formidable reserves of mineral carbon in the
carbonate rocks.
by suspension of the mineralization of marine organisms
accumulated in reducing abysses.
2 The Nitrogen Cycle
Nitrogen is the element best known to farmers, who
intervene actively in the nitrogen cycle by adding
nitrogen fertilizer or by certain cultural practices.
Mineral nitrogen is abundant on our planet, since its
gaseous form, N2, atmospheric nitrogen, represents most
of the air we breathe. On the other hand, nitrate ions,
NO3-, the only form of mineral nitrogen that can be
assimilated by plants, and ammonium ions are highly
unstable and easily leached.
Organic nitrogen constitutes all animal and plant cells. It
also plays an important role as a metabolite; since it is
found in amino acids, the basic links of all proteins. It is
also involved in the construction of nucleic acids in the
form of puric and pyramidal bases.
When living things die, the agents of putrefaction,
mostly aerobic bacteria, mineralize the organic nitrogen
according to the process of nitrification, which happens
in several stages (Fig.). By the mediation of symbiotic
bacteria, certain plants, especially the Leguminosae, can
fix atmospheric nitrogen, which biochemical reactions
reduce to ammoniacal nitrogen, and then to proteins.
 
UNIT 10: ECOLOGICAL PRINCIPLES 
2.1. Fixation and Transformation of Atmospheric
Nitrogen
a. Fixation by Symbiotic Bacteria and Cyanobacteria:
Gaseous molecular nitrogen can be reduced to ammonia
by micro-organisms. These are sometimes located in the
rhizosphere of roots, as with free fixers such as Azo-
tobacter(aerobic bacteria) or Clostridium (anaerobic bac-
teria). They are also found on nodules located on the
roots or stems of certain floristic groups, as with
symbiotic fixers such as Rhizobium (on Leguminosae) or
Frankia (symbiotic bacteria of Casuarina and alder).
Micro-organisms can also live in the aquatic envi-
ronment, such as cyanobacteria (also called Cyano-
phyceae). Some of these organisms float by means of
b. Transformation of N2 by Meteorological Phenomena:
At high altitudes, multiple rays transform atmospheric
nitrogen into NH3+ and nitrogen oxides. This also
happens during storms because the colossal amount of
energy liberated by lightning acts in the same way.
Added to these photochemical and electric effects is the
influence of certain precipitations. European farmers say
that snow in February is as good as fertilizer, because the
slowly failing flakes capture nitrogen, which is then
incorporated in the soil.
2.2. Mineralization of Nitrogen
a. Nitrification: When living things die, micro-
organisms that are autotrophic with respect to nitrogen
progressively degrade the proteins into ammoniacal
INSTITUTE FOR ADVANCED STUDIES, JODHPUR  61
specialized cells, the heterocysts, and thus fix nitrogen in
the air to balance their NIP ratio in an optimal way.
The reduction of atmospheric nitrogen leads to a
synthesis of ammonia molecules that immediately par-
ticipate in the elaboration of amino acids and then pro-
teins.
N2 ⎯⎯→⎯ HH=NH ⎯⎯→ ⎯ H2N-NH2 ⎯⎯→
⎯
2H 2 H 2 H
2NH3+
dimide hydrazine ammonia
This reduction is catalysed by a bacterial enzyme,
nitrogenase, which is inactivated by oxygen. Also, the
nodules, large consumers of oxygen, must be
apportioned by the transporters, the leghaemoglobins,
which facilitate the diffusion of O2 under very low
pressure without inactivating the nitrogenase. .
mineral nitrogen, nitrous nitrogen, and finally nitric
nitrogen. These three stages set off the oxidation of these
various substrates by aerobic bacteria, which draw from
it the energy required for chemosynthesis. The result of
these redox pairs is similar to that which results from
photosynthesis, that is, C3 molecules that are taken up
again by various metabolisms. Here it is called
chemosynthesis because the energy comes from
chemical reactions and not from photons.
CO2 + 2H+ + energy Æ C3 molecules Æ C6 molecules
AmmonificationÆAction of ammonifying bacteria: If
the soil is sufficiently aerated, bacterial enzymes cata-
lyse the oxidation of amino acids contained in the
decomposing proteins which releases ammonia or NH4+
ions:
UNIT 10: ECOLOGICAL PRINCIPLES 

RCHNH2COOH + O2 Æ RH + 2CO2 + NH3+ (ammonia) 3.1. Phosphorus In Terrestrial Ecosystems: The

RCHNH2COOH + O2 + H+ Æ RH + 2CO2 + NH4
 +            dissolution of phosphate rocks makes mineral carbon
(ammonium ion)  available to plant roots, which they can absorb and
NitrosationÆAction of nitrous bacteria: Other
aerobic autotrophic bacteria, such as Nitrosomonas
europea, use ammonia or ammonium in a redox pair and
transform them into nitric acid or into NO2- ions
according to the following reactions:
2NH3+ + 3O2 -+ 2HNO2 + H2O + energy
NitrationÆaction of nitric bacteria: Highly unstable
nitrous acid. or NO 2- ions from nitrosatioo are
recovered by bacteria of the Nitrobacter type, which
oxidize them into nitric acid or into NO3- ions.
Thesecombine quickly with soil cations to give nitrates,
which the plants can then absorb.
2HNO2 + O2 Æ 2HNO3 + energy
2NO2- + O2 Æ 2NO3- + energy
b. Denitrification: When soils are not aerated, the
nitrates are lost to the plants because anaerobic bacteria
metabolize them into nitrites, nitrogen oxide, or free
nitrogen N2. The bacteria Pseudomonas transforms
nitrates into ammonia, Nitrococcus denitrificans
transforms them into nitrogen oxide. Thiobacillus
denitrificans, which is autotrophic with respect to
sulphur and nitrogen, degrades sulphur and nitrates
respectively into SO4 2- and N2.
incorporate in their biomass. Phosphorus is a major ele-
ment in the metabolism of living things. It is especially
involved in the constitution of essential molecules
involved in the major functions of organisms. It is
involved in the elaboration of tissues (nervous system,
skeleton) and controls heredity and energy transport.
Above all, phosphorus is found in the nucleic acids,
DNA and the RNAs, in the phospholipids of the mem-
branes and the brain, in the energy-rich molecules ATP,
ADP, and AMP, and associated with sugars, glucose 6
phosphate, ribulose diphosphate, and other molecules.
Phosphorus circulates in biocenoses through the food
webs. It is first absorbed by plants and then incorporated
in herbivores, carnivores, parasites, predators and finally
decomposers. When living things die, micro-organisms
mineralize the organic phosphorus molecules into phos-
phates. Some of these may be newly absorbed by plants
and keep up a more or less complete cycle on land. But
most phosphate ions are leached and run off into the
aquatic ecosystems. .

2.3. Synthesis of Fertilizers and the Nitrogen Cycle

Industry, by making numerous synthetic fertilizers avail-

able to farmers, intervenes to some extent in the nitrogen
cycle. Many nitrogen fertilizers are in fact synthesized
by reduction of the nitrogen in the air. Since they must
release either NO3- or NH4+ ions, their production
triggers chemical reactions using the sulphuric acid or
nitric acid. For example:
H2SO4 + 2NH3+ Æ (NH4)2SO4 (ammonium sulphate)
HNO3 + NH3+ Æ NH4NO3 (ammonium nitrate)

These two fertilizers, with calcium nitrate, Ca(NO3)2

obtained chemically, are the most commonly used.
Those that directly input nitrates act very quickly but are
a significant source of pollution because they are highly
soluble. Basic slow-acting fertilizers are also used, such
as urea, CO(NH2)2 of calcic cyanamide CaCN2.
Ammonitrates simultaneously input fast-acting nitric ni-
trogen and ammoniacal nitrogen, which acts more
slowly.
3. The Phosphorus Cycle
On the scale of the biosphere, the phosphorus cycle is
incomplete or open. Like the other elements of
biogeochemical cycles, phosphorus exists in the mineral
and organic form. The stored mineral phosphorus avail-
able to biocenoses is entirely contained in the rocks. It is
found in the lithosphere in apatites or fossilized
sediments such as phosphorites.
This relatively rare element in the biosphere has a cycle
that has two phases: one that acts in terrestrial eco-
systems, the other in aquatic ecosystems (Fig.).
3.2. Phosphorus in Aquatic Ecosystems: Because of
run-off, there are dissolved phosphates (PO43-) in fresh
water and in the oceans. Phytoplankton captures and
uses them in their development. In normal conditions,
the phosphorus is insufficient to ensure optimal
photosynthesis of algae and their development is rather
limited. Sometimes, on the other hand, because of
human activities, there is excessive phosphorus in the
water. The disturbance of the N/P ratio leads to an
abnormal proliferation of certain algae or bacteria and
eutrophication of waters.
As in the continental biosphere, mineral phosphorus
captured by the algae is incorporated in the organic form
in aquatic food webs. It passes from phytoplankton to
zooplankton, then concentrates in the molluscs, fish,
mammals, and aquatic birds.A small part of the

62 INSTITUTE FOR ADVANCED STUDIES, JODHPUR 
UNIT 10: ECOLOGICAL PRINCIPLES 
phosphorus in oceans returns to the land because of
fishing or through the excrement of aquatic birds. Fish
are one source of high amounts of phosphorus. As for
bird excrement, it is collected from the rock cliffs on
which the birds build their nests, especially in Chile, and
used as guano, a phosphate fertilizer.
Another part, mineralized in the form of sediments from
cadavers of aquatic creatures and accumulated in
shallow water, rises by the phenomenon of upwelling
(ascending currents). It can also be incorporated in the
food webs by phytoplankton and thus goes through a
closed cycle. But the greatest quantity of this
mineralized phosphorus is lost in very deep sediments. It
thus escapes cycling and represents the open part of the
phosphorus cycle. It ultimately fossilizes in the form of
phosphate rocks within the oceanic plates.
4.1. Sulphur in the Atmosphere: Sulphur in the air
comes from various sources. The four major sources are
the following:
ƒ Gases emitted by volcanoes, the sulphatres,
release a hot water vapour that is highly
charged with S02 and above all H2S. Around
the mouth of a volcano, the suphurous gases
are deposited regularly at the vent openings in
the form of magnificent yellow flakes of
sulphur.
ƒ Combustion of hydrocarbons and coal in
factories, heating of domestic stoves, and
motor vehicle emissions release fumes rich in
SO2. . These sediments, when compacted, constitute rock
ƒ Erosion and dissolution of soils by rain water
also enriches the atmosphere in sulphurous
elements.
ƒ Finally, sulphur bacteria, which are
autotrophic with respect to sulphur and
profuse in marshes and reducing mediums that
are normally hostile to life, metabolize sulphur
INSTITUTE FOR ADVANCED STUDIES, JODHPUR  63
4. The Sulphur Cycle: The biogeochemical cycle of
Sulphur is best known by the phenomenon of acid rain,
in which it combines with the action of nitrous oxides.
Like other elements, Sulphur is present in two forms,
mineral and organic.The organic form appears only in
the constitution of proteins that contain sulphurate amino
acids: cystine, cystein, and methionine. In the molecules,
it is characterized most often in disulphide bridges and
thiol groups.
The mineral form, on the other hand, is quite well
represented in the atmosphere and terrestrial and aquatic
ecosystems. In the air, sulphur is mostly present in the
form of sulphurous or sulphur dioxide (SO2) sulphurate
hydrogen (H2S), or sulphates (SO4-2). In the water and.
sediments, mostly sulphides (S2-) are seen. In the soil it
is found in minerals such as iron sulphur or pyrite (FeS)
or calcium sulphate or gypsum (CaSO4), from which
plaster is made. The fossil fuels, coal and petroleum,
also contain sulphur (Fig).
and draw energy from it to reduce CO2 into
organic molecules. They also release
sulphurous anhydride or sulphurate hydrogen
into the atmosphere.
4.2 Sulphur in the Sediments
The largest reserve of sulphur is found in sediments. It
accumulates here when conditions are reducing,
combining with iron, phosphorus, calcium, magnesium,
and other elements. This sedimentation is constantly
being produced in marshes, but mostly in the sea
bottoms, in euxenic abysses.
minerals such as gypsum and pyrite. In the same way,
when petroleum or coal is formed over geological
periods time, sulphurate sediments are added to organic
matter, which is fossilized. This explains why the
combustion. If these fuels is so polluting and is partly
responsible for acid rain.
UNIT 10: ECOLOGICAL PRINCIPLES 

5 The Oxygen Cycle

5.1. Production of Molecular Oxygen: The oxygen

cycle is closely linked to those of carbon and water since
the essential source of free molecular oxygen is
photosynthesis. It is because of green plants that the air
we breathe contains 21% of O2 in volume. This reserve
is maintained by the forest, which each year releases
about 15 t of oxygen per hectare, and also by the oceans
(Fig.). In the aquatic environment, phytoplankton plays
the role of trees. It maintains an equilibrium that
compensates the consumption of dissolved oxygen by
biocenoses.

12.5.2. Degradation of Molecular Oxygen: Oxygen is

consumed by respiration and fermentation, which
characterize the catabolism of living things, and also by
various combustions linked to human activity. The
Krebs cycle of cellular oxidation is dependent on energy
provided to cells in the form of ATP. Since it is a
powerful oxidant, oxygen nevertheless remains toxic to
organisms. Also, its obligatory cellular utilization
necessitates the intervention of appropriate molecular Note: If all the fossilized organic molecules formedover
transporters such as haemoglobin in blood or geological time were burnt, there would no longer be
leghemoglobins in plants. oxygen on earth.

Part of the oxygen in air is lost in the oxidation of 6 Hydrologic Cycle or Water Cycle
minerals, as for iron:
4FeO + O2 Æ 2Fe2O3 Saltwater evaporates from sun's energy producing fresh
Molecular oxygen is also involved in the formation of water in clouds; leaves salts in ocean. Water vapor cools
ozone by reaction with UV rays: and condenses to precipitation over oceans and land.
O2 + O Æ O3 Runoff forms fresh water lakes, streams, ponds,
A small part is finally associated with carbon monoxide groundwater, and is held in plants and transpired. Gravity
from volcanoes or the combustion of hydrocarbons to returns water to oceans but some moves slowly through
produce CO2 water table and porous aquifers between rock layers.
2CO + O2 Æ2CO2 Although the water cycle shows water to be a renewable
resource, the 3% of water that is fresh may be polluted or

H. Biogeography: Major terrestrial biomes; theory of island biogeography; biogeographical zones 
of India.  

Biogeography is the science that documents and
attempts to explain the patterns of distribution of
populations, species, and ecological communities across
Earth.
Earth’s Biogeographic Regions
Explaining species’ distributions might seem to be a
simple matter, because the question of why a species is
or is not found in a certain location has only a few
possible answers:
• If a species occupies a particular area, either it
evolved there, or it evolved elsewhere and dispersed to
the area.
• If a species is not found in a particular area,
either it evolved elsewhere and never dispersed to the
area, or it was once present in the area but no longer
lives there.
Determining which of these possible answers is correct
requires information about the evolutionary histories of
species, which comes from fossils and from knowledge
of their phylogenetic relationships. It also requires
information about changes in Earth’s geography (such as
continental drift, glacial advances and retreats, sea level
changes, and mountain building) that occurred as the
organisms were evolving. Such geological information
can tell us whether organisms evolved where they are
currently found or dispersed and colonized new areas
from a distant area of origin.
The biotas of the continents differ enough to allow the
division of Earth into several major biogeographic
regions. Biogeographic regions are based on the
taxonomic similarities of the organisms living in them.
The boundaries of biogeographic regions are set where
species compositions change dramatically over short
distances (Figure 1).
The biotas of the biogeographic regions differ because
oceans, mountains, deserts, and other barriers restrict the
dispersal of organisms. Although there has been
dispersal of organisms between adjacent biogeographic
regions, such interchanges have not been frequent
enough to eliminate the striking differences that have

64 INSTITUTE FOR ADVANCED STUDIES, JODHPUR 
UNIT 10: ECOLOGICAL PRINCIPLES 
resulted from speciation and extinction within each
region.
A species found only within a certain region is said to be
endemic to that region. Remote islands typically have
distinctive endemic biotas because water barriers greatly
restrict immigration. If the islands are large enough or
form part of an archipelago, allopatric speciation often
produces unique species and communities. For example,
nearly all the tracheophytes and vertebrates of
Madagascar, a large island off the eastern coast of
Africa, are endemic to that island. Madagascar by itself
could be called a biogeographic region, but because
dozens of islands would qualify as biogeographic
History and Biogeography
Before 1850, most people, including biogeographers,
believed in a relatively unchanging Earth that was too
young for long-term processes to account for the
diversity and distribution of life. Linnaeus (1758), for
example, believed that all organisms had been created in
one place, which he called Paradise, from which they
later dispersed.
Indeed, because most people believed that the continents
were fixed in their positions, the only way to account for
the current distributions of organisms was to invoke
massive dispersal.
The notion that the continents might have moved was
not seriously considered until 1912, when Alfred
Wegener, a German meteorologist, argued that the
continents had drifted over time. Wegener based his
theory on several observations:
• The shapes of continents (the outlines of
Africa and South America seem to fit together like
pieces of a puzzle)
• The alignment of mountain chains, rock strata,
coal beds, and glacial deposits on different continents
• The distributions of closely related species that
were shared between Africa and South America, which
were difficult to explain if the continents had always
been where they are now
INSTITUTE FOR ADVANCED STUDIES, JODHPUR  65
regions on the basis of the distinctness of their biotas,
islands are treated differently from continents.
Most species are confined to a single biogeographic
region. The most widespread species today is probably
Homo sapiens, but a few other species—for example, the
great egret, the osprey, the peregrine falcon, and the barn
owl—are found on all continents except Antarctica.
Next we will discuss the influence of speciation,
extinction, and other historical processes on
biogeographic patterns, and then consider the influence
of processes operating today.
When Wegener proposed that the continents had moved,
few scientists took him seriously. There were no known
mechanisms to move continents, and no convincing
geological evidence of such movements existed.
Geological evidence and plausible mechanisms were
eventually discovered. The broad pattern of continental
movement, which continues today, is now clear. About
280 million years ago, the continents were united to
form a single land mass, called Pangaea. The continents
then began to separate from one another, but when the
continents were still very close to one another (about 245
mya), many groups of terrestrial and freshwater
organisms, such as insects, freshwater fishes, frogs, and
tracheophytes, had already evolved. The ancestors of
some organisms that live on widely separated continents
today were probably present on those land masses when
they were part of Pangaea.
By 100 mya, continental drift had separated Pangaea into
northern (Laurasia) and southern (Gondwana) land
masses, and the southern continents were moving away
from each other. Over time, India separated from Africa
and slammed into southern Asia, Australia moved closer
to Southeast Asia, and South America, which had drifted
as an island for 60 million years, came into contact with
North America. Throughout the history of life,
continental drift has both separated and combined biotas,
thus greatly influencing the distribution and evolution of
species.
UNIT 10: ECOLOGICAL PRINCIPLES 
Biogeographers convert phylogenies to “area
phylogenies”
As the age of Earth, the geological processes that shaped
it, and the mechanics of evolution became better
understood, biogeographers were able to ask questions
such as, Where and when did evolutionary lineages
originate? How did they spread? What do the present-
day distributions of organisms tell us about their past
histories?
A technique that was developed to help answer these
questions was the creation of area phylogenies. To
generate an area phylogeny, biogeographers alter a
taxonomic phylogeny by replacing the names of the taxa
with the names of the places where those taxa live or
lived. For example, an area phylogeny suggests that
horses speciated as they moved from Asia to Africa,
Vicariant events and dispersal both influence
distributions
The appearance of a barrier that splits the range of a
species is called a vicariant event. Avicariant event
divides the population of a species even though no
individuals have dispersed to new areas. If, however,
members of a species cross an already existing barrier
and establish a new population, the species’ disjunct
range is a result of dispersal. By studying a single
evolutionary lineage, a biogeographer may discover
evidence suggesting that the distribution of an ancestral
species was influenced by a vicariant event such as a
change in sea level or mountain building (see Figure 3).
If that inference is correct, then species in other lineages
are likely to have been influenced by the same event;
that is, a number of lineages may have similar
distribution patterns. Differences in distribution patterns
among lineages may indicate that the lineages responded
differently to the same vicariant events, that the lineages
separated at different times, or that the lineages have had
very different dispersal histories. By analyzing such
similarities and differences, biogeographers can discover
the relative roles of vicariant events and dispersal in
determining today’s distribution patterns.
66 INSTITUTE FOR ADVANCED STUDIES, JODHPUR
whereas the speciation of zebras took place entirely in
Africa (Figure 2). Biogeographers use several
approaches to infer the approximate times of separation
of taxa within a lineage. First, if a molecular clock has
been ticking at a relatively constant rate, the amount of
difference in the molecules of species should be strongly
correlated with the length of time their lineages have
been evolving independently.
Second, fossils can help to show how long a taxon has
been present in an area and whether its members
formerly lived in areas where they are no longer found.
A third valuable source of information is the distribution
of living species. Much more information can be
gathered on current distributions than will ever be
available from fossils. Similarities in the distributions of
many lineages of organisms provide clues about past
events that affected them.
The longer an area has been isolated from other areas by
a vicariant event, such as continental drift, the more
endemic taxa it is likely to have, because there has been
more time for evolutionary divergence to take place.
Australia, which has been separated the longest from the
other continents (about 65 million years), has the most
distinctive biota of any continent.
South America has the next most distinctive biota,
having been isolated from other continents for nearly 60
million years. North America and Eurasia, which were
joined together for much of Earth’s history, have very
similar biotas. That is why the early European travelers
felt more at home in North America than in Australia.
When several hypotheses can explain a pattern, scientists
typically prefer the most parsimonious one—the one that
requires the smallest number of unobserved events to
account for it. We saw how the parsimony principle is
used in the reconstruction of phylogenies. To see how it
is applied to biogeography, consider the distribution of
the New Zealand flightless weevil Lyperobius huttoni, a
species that is found in the mountains of South Island
and on sea cliffs at the extreme southwestern corner of
North Island (Figure 3). If you knew only its current
 
UNIT 10: ECOLOGICAL PRINCIPLES 
distribution and the current positions of the two islands,
you might surmise that, even though this weevil cannot
fly, it had somehow managed to cross Cook Strait, the
25-kilometer body of water that separates the two
islands. However, more than 60 other animal and plant
species, including other species of flightless insects, live
on both sides of Cook Strait. Although organisms do
cross marine and terrestrial barriers, it is unlikely that all
of these species made the same ocean crossing. In fact,
we do not need to make that assumption. Geological
evidence indicates that the present day southwestern tip
of North Island was formerly united with South Island.
Therefore, none of the 60 species need have made a
water crossing. Asingle vicariant event, the separation of
the northern tip of South Island from the remainder of
the island by the newly formed Cook Strait, could have
split all of the distributions.
As we have just seen, the distributions of species today
have been determined, in part, by history. However,
because all organisms must be able to survive in today’s
environmental conditions, Earth’s climates also exert a
powerful influence on the distributions, abundances, and
evolution of species.
Ecology and Biogeography
The climate of a region is the average of the
atmospheric conditions (temperature, precipitation, and
wind velocity) found there over time. Climates vary
greatly from place to place on Earth, primarily because
different places receive different amounts of solar
energy. We will first examine how these differences in
solar energy input determine atmospheric and oceanic
circulation. Then we will show how climates influence
the geographic distributions of organisms.
Solar energy inputs drive global climates
INSTITUTE FOR ADVANCED STUDIES, JODHPUR  67
Every place on Earth receives the same total number of
hours of sunlight each year—an average of 12 hours per
day—but not the same amount of energy. The rate at
which solar energy arrives per unit of Earth’s surface
depends primarily on the angle of sunlight. If the sun is
low in the sky, a given amount of solar energy is spread
over a larger area (and is thus less intense) than if the
sun is directly overhead.
In addition, when the sun is low in the sky, sunlight must
pass through more of Earth’s atmosphere, so more of its
energy is absorbed and reflected before it reaches the
ground. Thus, at higher latitudes (closer to the poles),
there is greater variation in both day length and the angle
of arriving solar energy over the course of a year than at
latitudes closer to the equator. On average, mean annual
air temperature decreases about 0.4°C for every degree
of latitude (about 110 kilometers) at sea level. Air
temperature also decreases with elevation. As a parcel of
air rises, it expands, its molecules move farther apart, its
pressure and temperature drop, and it releases moisture.
When a parcel of air descends, it is compressed, its
pressure rises, its temperature increases, and it takes up
moisture. Earth’s climates are strongly influenced by
global air circulation patterns, which result from the
global variation in solar energy input that we have just
described and from the spinning of Earth on its axis. Air
rises when heated by the sun. Warm air rises in the
Tropics, which receive the greatest solar energy input.
This rising air is replaced by air that flows toward the
equator from the north and south. The coming together
of these air masses produces the intertropical
convergence zone. Typically, heavy rains fall in a region
when it is close to the intertropical convergence zone as
the rising air releases its moisture. The intertropical
convergence zone shifts latitudinally with the seasons,
following the shift in the zone of greatest solar energy
input. This shift results in a characteristic latitudinal
pattern of distribution of rainy and dry seasons in
tropical and subtropical regions (Figure 4).
The air that moves into the intertropical convergence
zone to replace the rising air is replaced, in turn, by air
UNIT 10: ECOLOGICAL PRINCIPLES 
from aloft that descends at roughly 30° north and south
latitudes after having traveled away from the equator at
great heights. This air, which cooled and lost its moisture
while rising at the equator, now descends, warms, and
takes up rather than releasing moisture. Many of Earth’s
deserts, such as the Sahara and the Australian deserts,
are located at these latitudes where dry air descends.
At about 60° north and south latitudes, air rises again
and moves either toward or away from the equator. At
the poles, where there is little input of solar energy, air
descends. The black arrows around the edge of Figure 5
show these vertical air circulation patterns. These
movements of air masses are responsible, in part, for
global wind patterns.
The spinning of Earth on its axis also influences surface
winds because Earth’s velocity is rapid at the equator,
where its diameter is greatest, but relatively slow close
to the poles.
An air mass at a particular latitude has the same velocity
as Earth has at that latitude. As an air mass moves
toward the equator, it confronts an increasingly faster
spin, and its rotational movement is slower than that of
Earth beneath it. Conversely, as an air mass moves
poleward, it confronts an increasingly slower spin, and it
speeds up relative to Earth beneath it. Therefore, air
masses moving latitudinally are deflected to the right in
the Northern Hemisphere and to the left in the Southern
Hemisphere. Winds blowing toward the equator from the
north and south veer to become the northeast and
southeast trade winds, respectively.
Winds blowing away from the equator also veer and
become the westerly winds that prevail at mid-latitudes.
The broad arrows in Figure 5 show the average
directions of these surface winds.
When wind patterns bring air into contact with a
mountain range, the air rises to pass over the mountains,
cooling as it does so. Because cool air cannot hold as
much moisture as warm air, clouds frequently form and
release moisture as rain or snow. On the leeward side of
the mountain range, the now-dry air descends, warms,
and once again picks up moisture.
68 INSTITUTE FOR ADVANCED STUDIES, JODHPUR
This pattern often results in a dry area called a rain
shadow on the leeward sides of moutain ranges (Figure
6).
Global oceanic circulation is driven by wind patterns
The global pattern of wind circulation drives the
circulation of ocean water. Ocean water generally moves
in the direction of the prevailing winds (Figure 7).
Winds blowing toward the equator from the northeast
and southeast cause water to converge at the equator and
move westward until it encounters a continental land
mass. At that point the water splits, some of it moving
north and some of it moving south along continental
shores. This poleward movement of ocean water that has
been warmed in the Tropics is a major mechanism of
heat transfer to high latitudes. As it moves toward the
poles, the water veers right in the Northern Hemisphere
and left in the Southern Hemisphere. Thus, water
flowing toward the poles turns eastward until it
encounters another continent and is deflected laterally
along its shores. In both hemispheres, water flows
toward the equator along the west sides of continents,
continuing to veer right or left until it meets at the
equator and flows westward again.
The climates created by these atmospheric and oceanic
circulation patterns play key roles in determining what
kinds of organisms can live in a given region, as we’ll
see in the next section.
Terrestrial Biomes
In addition to recognizing biogeographic regions,
ecologists also classify communities of organisms into
biomes, ecosystem types that are based on the structure
of their dominant vegetation. The vegetation of a biome
has a similar appearance wherever on Earth that biome is
found, but the plant species in these communities,
despite their physical similarities, may not be
evolutionarily closely related. Biomes are named for and
identified by their characteristic vegetation, sometimes
supplemented by a description of their location or
climate, but each biome contains many species in all
other taxonomic groups.
 
UNIT 10: ECOLOGICAL PRINCIPLES 
The distribution of biomes on Earth is strongly
influenced by annual patterns of temperature and rainfall
(Figure 8). In some biomes, such as temperate deciduous
forest, precipitation is relatively constant throughout the
year, but temperature varies strikingly between summer
and winter. In other biomes, both temperature and
precipitation change seasonally.
INSTITUTE FOR ADVANCED STUDIES, JODHPUR  69
In still other biomes, such as tropical rainforest,
temperatures are nearly constant, but rainfall varies
seasonally. In the Tropics, where seasonal temperature
fluctuations are small, annual cycles are dominated by
wet and dry seasons (see Figure 4). It is easiest to grasp
the similarities and differences among terrestrial biomes
by means of a combination of photographs and graphs of
temperature, precipitation, and biological activity,
supplemented by a few words that describe the species
UNIT 10: ECOLOGICAL PRINCIPLES 

richness and other attributes of those biomes. We use summers are cool and dry. These forests are home to
this method in the following pages to describe the major Earth’s tallest trees.
terrestrial biomes of the world.
• Each biome is represented by a map showing its Boreal forests have only a few tree species. The
locations and two photographs that illustrate either dominant animals (e.g., moose, hares) eat leaves. The
thebiome at different times of year or representatives of seeds in the cones of conifers support a fauna of rodents
the biome in different places on Earth. and birds.
• One set of graphs plots seasonal patterns of
temperature and precipitation at a site in the biome.
• Other graphs show how active different kinds of
organisms are during the year. (For high-latitude biomes,
patterns in the Southern Hemisphere are six months out
of phase with those shown, which represent the Northern
Hemisphere.) Levels of biological activity, shown by the
width of the horizontal bars, change either because
resident organisms become more or less active (produce
leaves, come out of hibernation, hatch, or reproduce) or
because organisms migrate into and out of the biome at
different times of the year.
• A small box describes the growth forms of the plants
that dominate the vegetation in the biome and its patterns
of species richness.

These descriptions are very general and cannot describe

the variation that exists within each biome.

Tundra is found at high latitudes and in high

mountains

The tundra biome is found in the Arctic and high in

mountains at all latitudes. Arctic tundra vegetation,
which consists of short perennial plants, is underlain by
permafrost—soil whose water is permanently frozen.
The top few centimeters of soil thaw during the short
summers, when the sun shines 24 hours a day. Even
though there is little precipitation, lowland Arctic tundra
is very wet because water cannot drain down through the
permafrost. Plants grow for only a few months each
year. Most Arctic tundra animals either migrate into the
area only for the summer or are dormant for most of the
year.

Tropical alpine tundra is not underlain by permafrost, so

photosynthesis and most other biological activities
continue (albeit slowly) throughout the entire year. More
plant growth forms are present in tropical alpine than in
arctic tundra vegetation.

Most boreal forests are dominated by evergreen trees

Boreal forest is found equatorward from tundra and at

lower elevations on temperate-zone mountains. Boreal
forest winters are long and very cold; summers are short
(although often warm). The shortness of the summers
favors trees with evergreen leaves because these trees
are ready to photosynthesize as soon as temperatures
warm in spring.

The boreal forests of the Northern Hemisphere are

dominated by evergreen coniferous gymnosperms. In the
Southern Hemisphere the dominant trees are southern
beeches (Nothofagus), some of which are evergreen.
Evergreen forests also grow along the west coasts of
continents at middle to high latitudes in both
hemispheres, where winters are mild but very wet and

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UNIT 10: ECOLOGICAL PRINCIPLES 

Deciduous trees, which dominate these forests, lose their

leaves during the cold winters and produce leaves that
photosynthesize rapidly during the warm, moist
summers. Many more tree species live here than in
boreal forests. The temperate forests richest in species
are in the southern Appalachian Mountains of the United
States and in eastern China and Japan—areas that were
not covered by glaciers during the Pleistocene. Many
genera of plants and animals are shared among the three
geographically separate deciduous forest biomes.

Temperate deciduous forests change with the seasons

The temperate deciduous forest biome is found in

eastern North America, eastern Asia, and Europe.
Temperatures in these regions fluctuate dramatically
between summer and winter. Precipitation is relatively
evenly distributed throughout the year.

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UNIT 10: ECOLOGICAL PRINCIPLES 
Temperate grasslands are widespread
The temperate grassland biome is found in many parts
of the world, all of which are relatively dry for much of
the year. Most grasslands, such as the pampas of
Argentina, the veldt of South Africa, and the Great
Plains of North America, have hot summers and
relatively cold winters. Most of this biome has been
converted to agriculture. In some grasslands, most of the
precipitation falls in winter (California grasslands); in
others, the majority falls in summer (Great Plains,
Russian steppe). Grassland vegetation is structurally
simple, but it is rich in species of perennial grasses,
sedges, and forbs. Grasslands are often riots of color
when forbs are in bloom. Grassland plants are adapted to
grazing and fire. They store much of their energy
underground and quickly resprout after they are burned
or grazed.
72 INSTITUTE FOR ADVANCED STUDIES, JODHPUR
Cold deserts are high and dry
The cold desert biome is found in dry regions at middle
to high latitudes, especially in the interiors of large
continents in the rain shadows of mountain ranges.
Seasonal changes in temperature are great.
Cold deserts are dominated by a few species of low-
growing shrubs. The surface layers of the soil are
recharged with moisture in winter, and plant growth is
concentrated in spring. Because soils dry rapidly in
spring, annual primary production is low.
Cold deserts are relatively poor in species of most
taxonomic groups, but the plants of this biome tend to
produce large numbers of seeds, supporting a rich fauna
of seed-eating birds, ants, and rodents.
 
UNIT 10: ECOLOGICAL PRINCIPLES 
Hot deserts form around 30° latitude
The hot desert biome is found in two belts, centered
around 30° north and 30° south latitudes, where air
descends, warms, and picks up moisture. Hot deserts
receive most of their rainfall in summer, but they also
receive winter rains from storms that form over the mid-
latitude oceans. The driest large regions, where summer
and winter rains rarely penetrate, are in the center of
Australia and the middle of the Sahara Desert of Africa.
Except in these driest regions, hot deserts have richer
and structurally more diverse vegetation than cold
deserts. Succulent plants that store large quantities of
water in their expandable stems are conspicuous in some
hot deserts. Annual plants germinate in abundance and
grow when rain falls. Pollination and dispersal of fruits
by animals are common. Rodents, termites, and ants are
often remarkably abundant, and lizards and snakes
typically are rich in species and abundant.
INSTITUTE FOR ADVANCED STUDIES, JODHPUR  73
The chaparral climate is dry and pleasant
The chaparral biome is found on the west sides of
continents at moderate latitudes (around 30o), where cool
ocean waters flow offshore. Winters in this biome are
cool and wet; summers are warm and dry. Such climates
are found in the Mediterranean region of Europe, coastal
California, central Chile, extreme southern Africa, and
southwestern Australia. The dominant plants of
chaparral vegetation are low growing shrubs and trees
with tough, evergreen leaves. The shrubs carry out most
of their growth and photosynthesis in early spring, when
insects are active and birds breed. Annual plants are
abundant and produce copious seeds that fall onto the
soil. This biome thus supports large populations of small
rodents, most of which store seeds in underground
burrows. Chaparral vegetation is naturally adapted to
survive periodic fires. Many shrubs of Northern
Hemisphere chaparral produce bird-dispersed fruits that
ripen in the late fall, when large numbers of migrant
birds arrive from the north.
UNIT 10: ECOLOGICAL PRINCIPLES 

Thorn forests and savannas have similar climates

Thorn forests are found on the equatorial sides of hot

deserts. The climate is semiarid; little or no rain falls
during winter, but rainfall may be heavy during summer.
Thorn forests contain many plants similar to those found
in hot deserts. The dominant plants are spiny shrubs and
small trees, many of which drop their leaves during the
long dry winter. Members of the genus Acacia are
common in thorn forests worldwide.

The dry tropical and subtropical regions of Africa, South

America, and Australia have extensive areas of
savannas— expanses of grasses and grasslike plants
with scattered trees.

The largest savannas are found in central and eastern

Africa, where the biome supports huge numbers of
grazing and browsing mammals and many large
carnivores that prey on them. The grazers and browsers
maintain the savannas. If savanna vegetation is not
grazed, browsed, or burned, it typically reverts to dense
thorn forest.

Tropical deciduous forests occur in hot lowlands

As the length of the rainy season increases toward the

equator, tropical deciduous forests replace thorn
forests. These forests have taller trees and fewer
succulent plants than thorn forests, and they are much
richer in plant and animal species.

Most of the trees, except for those growing along rivers,

lose their leaves during the long, hot dry season. Many
of them flower while they are leafless, and most species
are pollinated by animals. During the hot rainy season,
biological activity is intense.

The soils of the tropical deciduous biome are some of

the best soils in the tropics for agriculture, because they
are less leached of nutrients than the soils of wetter
areas. As a result, most tropical deciduous forests have
been cleared for agriculture
and cattle grazing. Restoration efforts are underway on
several continents.

Tropical evergreen forests are species-rich

Tropical evergreen forests are found in equatorial

regions where total rainfall exceeds 250 cm annually and
the dry season lasts no longer than 2 or 3 months. They
are the richest of all biomes in number of species of both
plants and animals, with up to 500 species of trees per
km2. Along with their immense species richness,
tropical evergreen forests have the highest overall
productivity of all ecological communities.

However, most mineral nutrients are tied up in the

vegetation. The soils usually cannot support agriculture
without massive applications of fertilizers. On the slopes
of tropical mountains, trees are shorter than lowland
tropical trees. Their leaves are smaller, and there are
more epiphytes (plants that grow on other plants,
deriving their nutrients and moisture from air and water
rather than soil).

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INSTITUTE FOR ADVANCED STUDIES, JODHPUR  75
UNIT 10: ECOLOGICAL PRINCIPLES 

Aquatic Biogeography currents (Figure 9). These temperature changes, in

combination with seasonal changes in the amount of
Three-fourths of Earth’s surface is covered by water, daylight, determine the seasons of maximum primary
most of it in the oceans. Earth’s oceans form one large, production. Species of marine algae photosynthesize
interconnected water mass with only partial barriers to either in summer or in winter, but not during both
dispersal. Fresh waters, in contrast, are divided into river seasons.
basins and thousands of relatively isolated lakes. For
organisms that cannot survive out of water, terrestrial
habitats between bodies of water are barriers to
dispersal. However, some aquatic species have flying
adults that can disperse widely among water bodies.

Others have windborne, desiccation-resistant spores and

seeds. Still others are small enough to be transported by
mud on the feet of birds.

Freshwater environments have little water but many

species

Although only about 2.5 percent of Earth’s water is

found in ponds, lakes, and streams, about 10 percent of
all aquatic species live in freshwater habitats. Many
freshwater taxa that are capable of dispersing across
terrestrial barriers are found over several continents.
Prominent among freshwater taxa are the more than
25,000 species of insects that have at least one aquatic
stage in their life cycle. Typically, eggs and larvae are
aquatic; the adults have wings. Some of these insects,
such as dragonflies, are powerful flyers, but mayflies
and some other species are weak flyers, desiccate rapidly
in air, and live no longer than a few days. As you would
expect, oceanic islands have few, if any, species of these
weak flyers. Similarly, fishes unable to live in salt water
can disperse only within the connected rivers and lakes
of a river basin. Most families of freshwater fishes that
cannot tolerate salt water are restricted to a single
continent.
Deep ocean waters prevent the dispersal of marine
organisms that live only in shallow water. The distance
that eggs and larvae of many marine organisms can be
carried by ocean currents is determined in large part by
the time it takes for larvae to metamorphose into
sedentary adults. Relatively few species have eggs and
larvae that survive long enough to disperse across wide
barriers of deep water. As a result, the richness of
shallow-water species in the intertidal and subtidal
zones of isolated islands in the Pacific Ocean decreases
with distance from the larger islands of Indonesia
(Figure 10).

Those families with species distributed on both sides of Regional Patterns of Species Richness
major saltwater barriers are believed to be ancient
lineages whose ancestors were distributed widely in Local species richness is often positively correlated with
Laurasia or Gondwana. both productivity and disturbance level. Other patterns
of species richness appear at larger spatial scales. As we
Water temperature defines marine biogeographic increase the area we are sampling, the number of species
regions we record increases slowly (Figure 10). However, if our
sampling area crosses a biogeographic boundary, the rate
As we saw in Figure 7, ocean water moves in great at which we add new species suddenly increases.
circular patterns—clockwise in the Northern Hemisphere
and counterclockwise in the Southern Hemisphere. Even
organisms with limited swimming abilities can move
long distances simply by floating with ocean currents.
Nevertheless, most marine organisms have restricted
ranges. Why is this true?

The oceans may be connected, but water temperatures,

salinities, and food supplies all may change spatially.
Living successfully in different regions of the ocean
requires different physiological tolerances and
morphological attributes.

Ocean temperatures, for example, can be barriers to

dispersal because many marine organisms function well
in only a relatively narrow range of temperatures. The
main biogeographic divisions of the ocean coincide with
regions where the surface water temperatures and
salinities change relatively abruptly as a result of
horizontal and vertical ocean

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UNIT 10: ECOLOGICAL PRINCIPLES 
At that point, we have added to our sample another
biogeographic region with a different evolutionary
history and a different biota. One of the first geographic
patterns of species richness observed by biologists was
that more species are found in low-latitude than in high-
latitude regions.
Species richness on islands and peninsulas is always less
than that in an equivalent area on the nearest mainland.
On islands, species richness is positively correlated with
island size, but inversely correlated with distance from
the mainland (Figure 11). An influential model relates
this pattern to the island’s history of immigrations and
extinctions.
Island biogeography:
Species richness is related to rates of immigration
and extinction
Over periods of a few hundred years (during which
speciation is unlikely), the species richness of an area is
influenced by the immigration of new species and the
extinction of species already present. It is easiest to
visualize the effects of these two processes if we
consider, as did Robert MacArthur and Edward O.
Wilson, an oceanic island that initially has no species.
Imagine a newly formed oceanic island that receives
colonists from a mainland area. The list of species on the
mainland that might possibly colonize the island is
called the species pool. The first colonists to arrive on
the island are all “new” species because no species live
there initially. As the number of species on the island
increases, a larger fraction of colonizing individuals will
be members of species already present. Therefore, even
if the same number of species arrive as before, the rate
of arrival of new species should decrease, until it reaches
zero when the island has all the species in the species
pool. As we will see, however, the process is unlikely to
proceed that far.
Now consider extinction rates. At first there will be only
a few species on the island, and their populations may
grow large. As more species arrive and their populations
increase, the resources of the island will be divided
among more species. Therefore, the average population
INSTITUTE FOR ADVANCED STUDIES, JODHPUR  77
size of each species will become smaller as the number
of species increases.
The smaller a population, the more likely it is to become
extinct. In addition, the number of species that can
possibly become extinct increases as species accumulate
on the island. Furthermore, new arrivals on the island
may include pathogens and predators that increase the
probability of extinction for other species. For all these
reasons, the rate of extinction increases as the number of
species on the island increases.
Because the rate of arrival of new species decreases and
the extinction rate increases as the number of species
increases, eventually the number of species on the island
should reach an equilibrium at which the rates of arrival
and extinction are equal (Figure 12a). If there are more
species than the equilibrium number, extinctions should
exceed arrivals, and species richness should decline. If
there are fewer species than the equilibrium number,
arrivals should exceed extinctions, and species richness
should increase.
The equilibrium is dynamic because if either rate
fluctuates, as they generally do, the equilibrium number
of species shifts up and down. MacArthur and Wilson’s
model can also be used to predict how species richness
should differ among islands of different sizes and
different distances from the mainland. We expect
extinction rates to be higher on small islands than on
large islands because species’ populations are, on
average, smaller there. Similarly, we expect fewer
immigrants to reach islands that are more distant from
the mainland. Figure 12b gives hypothetical relative
species richnesses for islands of different sizes and
distances from the mainland. As you can see, the number
of species should be highest for islands that are relatively
large and relatively close to the mainland.
The MacArthur-Wilson model has been tested
Major disturbances, which serve as “natural
experiments,” sometimes permit colonization and
extinction rates to be estimated directly. In August 1883,
Krakatau, an island in the Sunda Strait between Sumatra
and Java, was devastated by a series of volcanic
eruptions that destroyed all life on the island’s surface.
After the lava cooled, Krakatau was colonized rapidly by
plants and animals from Sumatra to the east and Java to
the west. By 1933, the island was again covered with a
tropical evergreen forest, and 271 species of plants and
27 species of resident land birds were found there.
UNIT 10: ECOLOGICAL PRINCIPLES 
During the 1920s, when a forest canopy was developing,
there were high rates of colonization by both birds and
plants (Table 1). Birds probably brought the seeds of
many plants because, between 1908 and 1934, both the
percentage (from 20% to 25%) and the absolute number
(from 21 to 54) of plant species with bird-dispersed
seeds increased.
Today the numbers of species of plants and birds are not
increasing as fast as they did during the 1920s, but
colonizations and extinctions continue, as predicted by
the model.
Biogeography and Human History
The distributions of land masses and species on Earth
have had a strong influence on human history. Humans
first evolved in Africa, but eventually dispersed
throughout the world. In recent times, human cultures
from Eurasia came to dominate other cultures.
Biogeography gave these cultures a number of
advantages.
Eurasia happened to have a large number of species of
plants and animals that were suitable for domestication.
Eurasia was home to 39 species of large-seeded grasses,
many more than were found in Africa or the Americas. It
also had 72 species of large mammals, compared with 51
in sub- Saharan Africa and 24 in the Americas. Thirteen
of these species, including pigs, horses, cattle, sheep,
goats, and camels, were domesticated in Eurasia. None
were domesticated in Africa, and only one, the llama, in
the Americas.
To be amenable to domestication, large mammals had to
have three important social characteristics: They had to
live in herds, have well-developed male dominance
hierarchies, and not defend territories. These traits
enabled humans to tame the animals, exert behavioral
dominance over them, and keep them in herds. All of the
large mammals of Africa lacked one or more of these
traits.
Besides providing people with food, the domestication
of large mammals had other important influences on
human history. Many human diseases, such as smallpox
and measles, were acquired from domesticated
mammals.
Eurasian people acquired immunity to these diseases, but
people on other continents did not. Thus, when
Europeans colonized the New World, they brought with
them diseases that devastated the indigenous people.
78 INSTITUTE FOR ADVANCED STUDIES, JODHPUR
Those unfortunate people transmitted no fatal diseases to
the Europeans. In addition, the Europeans had horses,
the only domesticated mammals capable of carrying a
person at high speeds.
Throughout human history, cultures with horses have
defeated and dominated cultures without them. In
Eurasia, most mountain ranges are oriented in an
eastwest direction. Therefore, dispersal of people and
their domesticated plants and animals was relatively
easy, and dispersing individuals were always within
climates with similar temperatures and day lengths.
Humans dispersed only recently into North America
across the high-latitude Bering Land Bridge. They
brought with them no domesticated plants or animals,
except dogs. North America had few species of grasses
with large seeds. Maize, the grass that came to dominate
American agriculture, was difficult to domesticate.
Its eventual spread northward from its center of
domestication in Mexico was possible only after
extensive genetic changes that adapted the plants to the
very different day lengths and climates of temperate
North America. Human history would have been very
different if the continents and their biotas had been
distributed differently. Thus, the study of biogeography
can help us understand ourselves as well as other
species.
BIOGEOGRAPHICAL ZONES IN INDIA
To a common man the term wildlife means living things
that are neither humans nor domesticated, especially
mammals, birds and fishes hunted by man. An ecologist
uses the term biodiversity for the variety of species of
all living plants, animals and microbes living in their
natural habitats.
India occupies a major part of South and SouthEast Asia,
the oriental realm, one of the six biogeographic realms
of the world (biosphere). Besides this Himalayan region
of India falls in palaearctic realm. Geological events in
the landmass of India provided conditions for high levels
of biodiversity. India's unique geographical position
between three different centres of evolution (beginning
from Gondwanaland, then shifting to northern Euratia
and finally influx from Africa and Ethiopia) led to
diversity in its climate, soil, flora and fauna. India is thus
very rich in biodiversity, and considered to be one of the
twelve centres of origin and diversity of several plant
species in the world. India is one of the twelve mega-
biodiversity countries of the world. The other countries
are, Mexico, Columbia, Brazil, Peru, Ecuador,. Zaire,
Madagascar, Indonesia, Malaysia, China and Australia.
India's biological wealth becomes apparent from the fact
that it represents ten biogeographic zones (Table 1,
Fig.1)
The Indian region is one of the most diverse
biogeographic regions of the world, embracing a wide
range of topography; from perpetually snow covered
high Himalayan ranges to plains at sea level low lying
swamps and mangroves, island systems, tropical
evergreen rain forests, fertile alluvial plains hot deserts
and high altitude cold deserts. There are almost rainless
areas, as well as the world's highest rainfall areas. The
climate ranges from tropical and subtropical in Indo
 
UNIT 10: ECOLOGICAL PRINCIPLES 
Gangetic plains and peninsular regions to temperate and
arctic in the Himalayan region. The habitat types vary
from the humid tropical Western Ghats to the hot deserts
of Rajasthan, from cold deserts of Ladakh to the long
warm coast line stretches of peninsular India. Thus, a
great variety of climatic and altitudinal variations
coupled with varied ecological habitats have contributed
immensely to the rich vegetational wealth, and varied
flora and fauna generating very unique biodiversity in
India.
Wildlife helps in promoting economic activities that
brings money through tourism. It contributes towards
maintenance of biodiversity. The colorful birds, animals
and other life forms in the forests are important in
maintaining the ecobalance. Disappearance of forests or
its reduction in area will cause disappearance of wildlife.
Table1. Major biogeographic zones of India (ZSI, 1997).
Thus, Conservation of forests and wildlife go side by
side. Establishment of national parks and wildlife
sanctuaries helps conservation of wildlife. Many
countries have through legislation banned killing of birds
and other animals. In India, restrictions are imposed on
killings of lions, tigers, deers, chital, Great Indian
bustard and peacocks etc.
Biogeographic Zones of India
We have already considered the soil, flora and fauna of
the country earlier in separate chapter. A brief account
of ecological sub-divisions of India and their faunna and
flora is presented in this section.
India is divided into the following three ecological sub-
regions :
Himalayan mountain system
INSTITUTE FOR ADVANCED STUDIES, JODHPUR  79
There are altitudinal as well as East-West variations in
this region. There are three distinct sub-zones each with
its characteristic fauna and flora.
Himalaya foothills: These extend from the eastern
frontiers of Kashmir to Assam.
Western Himalayas: In includes higher altitudes in
Himalayas from Kashmir, including Ladakh to Kumaon.
Eastern Himalayas: It includes regions of Sikkim and
extends in the east up to NEF A.
Peninsular-Indian sub-region: This is the raised
plateau land of the Deccan extending into the flood
plains of the Indo-Gangetic basin westwards into the
Great Thar desert of Rajasthan.
Tropical evergreen forests or Indo-Malayan sub-region
This is the region of heavy rainfall comprising north-
eastern India and the Western Ghats in South including
Malabar Coast.
Himalayan Mountain System
Fauna is typical of both the oriental and palaearctic
regions. Altitude affects the wildlife distribution.
Besides altitude, east-west variations also occur due to
decrease in monsoon rainfall towards the west. The west
Himalayas have low rainfall, heavy snowfall (temperate
conditions), whereas in east Himalayas, there is heavy
rainfall, snowfall only at very high altitudes, at lower
altitudes conditions are like tropical rain forests. Flora
and fauna of both Himalayas differ.
Himalaya foothills
These have characteristic bhabar and tarai formations
and the Siwalik ranges in the South.
Flora
UNIT 10: ECOLOGICAL PRINCIPLES 
Natural monsoon evergreen and semi-evergreen forests;
dominant species are sal, silk-cotton trees, giant
bamboos; tall grassy meadow with savannahas in tarai.
Fauna
Big mammals of North India like elephant, sambar,
swamp deer, cheetal, hog deer, barking deer, wild
ooar tIger, panther, wild dogs, hyena, black bear, sloth
bear, porcupine, Great Indian one-horned rhinoceros,
wild buffalo, gangetic gharial and golden langur.
Western Himalayas (High altitude region)
Flora
Coniferous-pine forests, rhododendrons, dwarf hill
bamboo and birch forests mixed with alpine pastures.
Fauna
Wild ass, wild goats (thar, markhor, ibex) and sheep
(Nayan, Marcopolo's sheep, bharal or blue sheep);
antelopes (chiru and Tibetan gazelle), deers (hangul or
Kashmir stag and shou or Sikkim stag, musk deer);
smaller mammals like marmots and pikas or mouse
hares; golden eagle, snow cocks, snow partridges; snow
leopard, wolf, fox, cats, black and brown bears; birds
like Himalayan monal pheasant, western trogopan,
Koklass, whitecrested khalij cheer pheasant; Griffon
vultures, lammergiers, choughs and ravens.
Eastern Himalayas
Flora
Oaks, magnolias, laur~ls and birches covered with moss
and ferns; coniferous forests of pine, fir, yew and
junipers with undergrowth of scrubby rhododendrQns
and dwarf bamboos; lichens, moses, orchids and other
epiphytes dominant (due to high humidity and high
rainfall).
Fauna. Red panda, hog badgers, ferret badgers, crestless
porcupines; goal antelopes (Serow, Goral, Takins).
Peninsular-Indian Sub-region
It has two zones, (i) peninsular India and its extension
into the drainage basin of the Ganges river system, and
(ii) desert region of Rajasthan-the Thar of Indian desert
region.
Peninsular India: It is home of tropical moist deciduous
to tropical dry deciduous and scrub vegetation depending
upon the variation in rainfall and humidity.
Flora
Sal in north and east extensions (higher rainfall) and teak
in southern plateau are dominant trees. West Ghats have
evergreen vegetation (flora and fauna similar to
evergreen rainforests of north-eastern India). In dry areas
of Rajasthan desert and Aravalli hills, trees are scattered
and thorny scrub species predominate. The forests give
way to more open savannah habit.
Fauna
Elephant, wild boar, deers (cheetal or axis deer, hog
deer, swamp deer or barasinga, sambar, muntjak or
barking deer), antelopes (four-homed antelope, nilgai,
blackbuck, chinkara gazelle), wild dog or dhole, tiger,
80 INSTITUTE FOR ADVANCED STUDIES, JODHPUR
leopard, cheetah, lion, wild pig, monkey, striped hyena,
jackal, gaur (a bull).
Indian desert
Thar desert of Rajasthan has unique flora and fauna.
Flora
Thorny trees with reduced leaves; cacti, other succulents
are the main plants.
Fauna
Animals are mostly burrowing ones. Among mammals
rodents are the largest group. The Indian desert gerbils
are mouse like rodents. Other animals are, wild ass,
black buck, desert cat, caracal, red fox; reptiles (snakes,
lizards and tortoise) well represented. Desert lizards
include agarnids, lacertids and geckos. Among birds the
most discussed is Great Indian bustard.
Tropical Rain Forest Region
Flora
Extensive grassy stands interspersed with densely
forested gorges of evergreen vegetation known as sholas
occur in the Nilgiris (an offshoot of Western ghats).
Sholas also occur in Annamalai and palni hills. The rain
forests of the Western ghats have dense and lofty trees
with much species diversity (more than 100 spp).
Mosses, ferns, epiphytes, orchids, lianas and vines,
herbs, shrubs make diverse habitat. Dipterocarpus spp
predominate in these forests. A variety of tropical
orchids are found. Stratification in rain forests is very
distinct-three horizontal layers are distinguished. The
lower storey has seedlings of palms, cardamom and
ginger.
Fauna
It is very rich with all kinds of animals. There are wild
elephants, gaur and other larger animals. Most species
are tree dwellers. The most prominent are hoolock
gibbon (only ape found in India), golden langur, capped
langur or leaf monkey, Assam macaque and the pig-
tailed macaque, lion-tailed macaque, Nilgiri langur,
stender loris, bats, giant squirrel, civets, flying squirrels,
Nilgiri mongoose, spiny mouse.
Andaman and nicobar Islands
Flora
These are home of tropical rain forests. Giant
Dipterocarpus, Terminalia and Lagerstroemia are
dominant. There are mangroves also on coastal areas.
Fauna
Many species (about 35) of mammals, a number of
reptiles and marine animals occur in this area. Among
mammals, bats and rats predominate (3/4th of the total
mammals on islands); Andaman pig, crab-eating
macaque, palm civet and deers (spotted deer, barking
deer, hog deer, sambar). Among marine mammals there
are dugong, false killer whale, dolphin. A~ong birds-
(about 250 species and sub species) are rare Narcondum
bombill, Nicobar pigeon and megapode. There are also
other birds like whitebellied sea-eagle, white- breasted
swiftlet and several fruit pigeons.
 
UNIT 10: ECOLOGICAL PRINCIPLES 

There are salt water crocodiles, a number of marine
turtles, coconut crab, lizards (the largest being water
monitor), 40 species of snakes including cobra, viper,
voral and sea snakes and pythons.
Mangrove swamps of sunderbans
Sunderbans are delta of the Ganges where both the
Brahmaputra and Ganges join and drain into the Bay of
Bengal.
Flora
The lower tidal zones have pioneer trees like Sonneratia
and Avicennia. Above this zone (middle level) there are
Rhizophora, Bruguiria and Excaecaria-Cereops forests
(covering nearly 70% of the mangrove forests). Above
this level (high land level) there are supporting forests of
Phoenix, pure or in association with Excaecaria. There
are Reriteria forests in the highest portion with thick
undergrowth of Phoenix and Neepa palms.
Temperate or montane zone
It is represented from 5,000 to 11,675 ft. above sea level.
At about 5,500 ft. Pinus longifolia is generally replaced
by P. excelsa. From 5,500 ft. to 6,000 ft. Cedrus
deodara is quite abundant forming pure forest stands. At
these altitudes Quercus incana also grows as separate
patches. In the inner Himalayas in Kashmir, Betula
(birch), Salix (cane) and Populus (poplar) are abundant
on~rtain soil types. At higher altitudes, Aesculus indica
(horse chestnut), Quercus semecarpifolia, Q. dilatata
alongwith the conifers such as Abies pindrow, Picea
morinda, Cupressus torulosa, Taxus boccata etc. are
most common components of vegetation. Rhododendron
companulatum grows at higher altitudes. In inner valleys
on dry mountains, Pinus gerardiana is also found. In dry
areas of Punjab, wheat and barely are cultivated, while in
wet valley of Kashmir, rice is the common crop. Other
common plants grown in Kashmir are saffron (Croccus
sativus), apples, peaches, walnuts, almonds etc.

Fauna
Ash (mud skippers or semi-terrestrial gobies), small
crabs land crabs, fiddler crabs, and the Dorippie the one
that has unusual association with sea anemone. Other
animals are weaver ants. In the higher regions of
mangrove, there are spotted deer, pigs, monitor lazards,
monkeys etc. The most interesting animal is the man-
eater Sunderbans tige

FLORISTIC (BOTANICAL) REGIONS OF INDIA

As Indian sub-continent is characterised with a variety of

climate types, flora of the country is a1so
correspondingly of different types in different parts. For
the study of flora, the country has been dividee into the
following nine floristic regions (Fig. 2) (i) Western
Himalayas, (ii) Eastern Himalayas (iii) West Indian
Deserts, (iv) Gangetic plains (v) Assam, (vi) Central
India, (vii) Malabars (viii) The Deccan, and (ix)
Andamans.

Western Himalayas

It extends from central region of Kumaon to north west

region of Kashmir. Altitudinally there are three zones of
vegetation corresponding to three climatic belts.
Submontane or lower region (tropical and
subtropical)
It is represented from about 1,000 to 5,000 ft, above sea
level in regions of Siwaliks and adjacent areas. The
forest is dominated by timber trees of Shorea robusta. In
riverain regions trees of Dalbergia sissoo are dominant,
while in more moist soils, dominants are Cedrela toona,
Ficus glomerata and Eugenia jambolana. In isolated
patches of grasses, there are present trees of Acacia
catechu and Butea monosperma. In dry belts towards
west, Shore robusta is replaced by such xerophytes as
Zizyphus, Carissa, Acacia, etc. with thorny succulent
euphorbias on slopes. Pinus roxburghii begins to appear
at 3,000 to 5,000 ft. Ground vegetation is poor.
Alpine zone
It is the limit of tree growth at about 12,000 ft. known as
timber or tree line, where the plant height is considerably
reduced. Plants are mostly dwarfed and cushion shaped
shrubs and grasses. At about 15,000 ft. and above-snow
line, plant growth is almost nil. On lower levels of this
zone, some rhododendrons, Betula utilis and small
junipers are present. Above this zone there are present
many types of herbs, with short period of vegetative
growth and flowering. These include Primula,
Potentilla, Polygonum, Geranium, Saxifraga, Aster etc.
Eastern Himalayas
It consists of regions of Sikkim and extends in the east
upto NEF A. In its vegetational zones, it is similar to the
western Himalayas. On the whole, the eastern Himalayas
have more tropical elements, greater variety of oaks and
rhododendrons and less of conifers than the western

INSTITUTE FOR ADVANCED STUDIES, JODHPUR  81
UNIT 10: ECOLOGICAL PRINCIPLES 
Himalayas. The chief differences are the higher rainfall
and warmer conditions in this part of Himalayas. The
tree and snow lines are higher by about 1,000 ft. than the
corresponding lines on western Himalayas. Species
diversity and vegetation density are higher in the east.
This region is also divided into three zones.
Submontane zone
Due to warm and humid weather it is typically tropical
with dense forests of Shorea rubusta. It extends from the
plain foot of the hill upto 6,000 ft. altitude. In riverain
area there are forests of Dalbergia sissoo and Acacia
catechu. Mixed forests of deciduous trees like
Sterospermum, Cedrela toona, Bauhinia, Anthocephalus
cadamba, Lagerstroemia parviflora are pre-dominant.
Tall trees like Albizzia procera, Salmalia, Artocarpus
chaplasha, bamboo (Dendrocalamus) are important.
Temperate zone
It ranges between 6,000 to 12,000 ft. altitude above sea
level. The lower region has several species of oaks, such
as Quercus lemellosa and Q. lineata, Michelia, Cedrela
and Eugenia. The upper region which is cooler, has such
conifers as Juniperus, Cryptomeria, Picea, Abies, and
Tsuga. One bamboo, Arundinaria sp. is also common.
Some rhododendrons are also common at higher
elevations.
Alpine zone
It is above 12,000 ft. where vegetation is devoid of trees.
Shrubby growth of Juniperus and Rhododendron is
found in grassy areas.
West Indian Desert (Indian Plains)
This region consists of parts of Rajasthan, Kutch, Delhi
and part of Gujarat. The climate is characterised by very
hot and dry summer, and cold winter. Rainfall is less
than 70 cm. The plants are mostly xerophytic, such as
Acacia nelotica, Prosopis spicifera, P. juliflora,
Salvadora oleo ides, S. persica. Tecomella, Capparis
aphylla, Tamarix dioica. and Zizyphus nummularia. The
ground vegetation is mostly represented by small
Calotropis sp., Panicum antidotale, Eleusine sp.,
Tribulus terrestris etc. Some common species used in
plantations are Saccharum munja, Panicum antidotale,
Cenchrus ciliaris, Capparis aphylla, Tamarix articulata,
Prosopis spicifera, P. juliflora, Acacia leucophloea and
A. senegal.
Gangetic plain
This region comprising Uttar Pradesh, Bihar and Bengal
is most fertile region. The chief climatic factors, the
temperature and rainfall together are responsible for
distinct type of vegetation. Rainfall is less than 70 cm in
west U.P., being more than 150 cm in Bengal.
Vegetation is chiefly of tropical moist and dry deciduous
forest type. In north-western D.P., near foothills of the
Himalayas, Dalbergia sissoo and Acacia nelotica are
most common. In south-west U.P., there are desert areas,
where characteristic species are Capparis aphylla.
Saccharum munja, Acacia nelotica etc. In eastern V.P.,
Butea monosperma (dhak) , Madhuca indica (mahua),
Terminalia arjuna (arjun), Buchanania lanzan
82 INSTITUTE FOR ADVANCED STUDIES, JODHPUR
(chiraunji), Diospyros melanoxylon (tendu), Cordia
myxa (lisora), Sterculia urens, Boswellia serrata (salai),
Acacia catechu (Khair), Azadirachta indica (neem),
Mangifera indica (mango), Ficus bengalensis (bargad),
F. religiosa (pi pal) are most dominant trees. Besides
them, some weeds and grasses like Xanthium
strumarium, Cassia tora, Argemone mexicana.
Amaranthus sp., Peristrophe bicalyculata. Dichanthium
annulatum. Bothriochloa pertusa etc. are also present. In
Gangetic delta region extreme swampy and halophytic
vegetation is common, where dominant species are
Rhizophora mucronata. R. conjugata, Acanthus
ilicifolius. Kandelia rheedii, Bruguiera gymnorhiza.
Ceriops roxburghiana etc.
Assam
This region receives the heaviest rainfall, with
Cherrapunji as much as more than 1000 cm. The
temperature and wetness are very high, which are
respons.ible for dense tropical evergreen forests. Some
of the important trees are Dipterocarpw macrocarpus.
Mesua ferrea, Michelia champaca,Shorea robusta,
Artocarpus chaplasha, Alstonful scholaris, Sterculia
alata, Lagerstroemia flosregina, Ficus elastica etc.
Some bamboos, as Bambusal pallida, Dendrocalamus
hamiltonii. Calamus sp. grasses as Imperata cylindrica,
Saccharum arundinaceum. Themeda sp., Phragmites sp.,
and insectivorous plants like Nepenthes sp. are also
present. In northern cooler regions, Alnus nepalensiJ,
Rhododendron arboreum, Betula sp. are also found in
hilly tracts, some conifers like Pinus khasiya and P.
insularis are also present.
Central India
It comprises Madhya Pradesh, parts of Orissa, and
Gujarat. Depending upon the amount of rainfall, forests
have developed into thorny, mixed deciduous and sal
types. The forest vegetation is chiefly constituted by
Tectona grandis, Diospyros melanoxylon, Butea
monosperma. Terminalfu tomentosa and Dalbergia
latifolia. The thorny vegetation consists of Carissa spina
rum, Zizyphus rotundifolia, Acacia leucophloea, A.
catechu. Butea
frondosa etc.
Malabar
This region comprises the western coast of India
extending from Gujarat in the north to the Cape Camorin
in the south. Rainfall is heavy. The vegetation is of four
types-tropical moist evergreen forests, mixed deciduous
forests, subtropical or temperate evergreen forests and
the mangrove forests.
The tropical wet evergreen forests are very luxuriant and
multi storeyed, with such tall trees as Dipterocarpus
indicus, Sterculia alata, Cedrela toona, Tectona grandis
and Dalbergia latifolia,Bamboos, like Dendrocalamus
strictus and Bambusa arundinacea are also present.
In the Nilgiri hills, there are temperate evergreen forests
of such trees as Eurya japonica, Michelia nilagirica and
Gordonia obtusa known as the sholas.
 
UNIT 10: ECOLOGICAL PRINCIPLES 
The Deccan
This region is drier with rainfall of about 10 cm. It
includes Andhra Pradesh, Tamilnadu and Kamataka. It
has a central hilly plateau with forests of Boswellia
serrata, Tectona grandis and Hardwickia pinnata, and
the [ow eastern dry Coromandal coast, with tropical dry
evergreen forests of Santalum album (chandan), Cedrela
toona and plants like Capparis, Phyllanthus, Euphorbia
sp.
Andamans
It has a wide range of spreading coastal vegetation like
mangroves, beech forests and in the interior evergreen
forests of tall trees. There are some pockets of dry areas
also. Important species of the island are Rhizophora,
Manusops, Calophylium, Dipterocarpus, Lagerstroemia
and Terminalia. Most of the area is now cleared for
paddy and sugarcane cultivation.
Vegetation of India
This subcontinent has been affected worst by human
influences since long lime. Due to this, climax
fonnations are very much altered and/or destroyed for
agriculture and other similar purposes. The present
vegetation has suffered immensely from effects of
plants, animals, soil, climate and man. Therefore, the
vegetation of the country that we see arond us are much
interfered. The most important factors used in the
classification of vegetation are rainfall, temperature,
biotic influences, and life forms. In India, there are two
most common types of plant formations, (i) forest and ii)
grassland.
Forest vegetation
Indian forests have generally been classified on the basis
of temperature into four major types (i) tropical, (ii)
montane subtropical, (iii) temperate, and (iv) alpine.
Tropical forests
Common in the warmer plains, ranging from very dense,
multistoreyed of diverse trees, shrubs and lianas in areas
of high rainfall to dry, scrub jungles of thorny bushes in
isolated patches in dry areas. Thus, they are of two types
- moist tropical and dry tropical forests.
Moist tropical forests
On the basis of degree of wetness these are of the
following three principal types:
(i) Tropical wet evergreen forests, where annual rainfall
is over 250 cm., as in West Coast, Assam, Bengal and
Andaman islands. They are multi storeyed, made up of
small trees, shrubs, epiphytes, lianas and dense ground
vegetation. The dominant members are such trees as
Dipterocarpus, Hopea, Artocarpus, Mangifera, Emblica,
Michelia, Ervatamia, Lagerstroemia, Ixora and some
climbers.
(ii) Tropical moist semi-evergreen forests, better
developed in northern than the southern region of
country. Trees shed their leaves for brief period. In
north, they develop in north Assam and Bengal and parts
INSTITUTE FOR ADVANCED STUDIES, JODHPUR  83
of Orissa receiving heavy rainfall. There are some
evergreen plants also as Artocarpus, Michelia, and
Eugenia. The deciduous trees are Terminalia, Terameles
and Shorea.
(iii) Tropical moist deciduous forests, in which some
trees shed leaves for brief period, some are evergreen
and semi-evergreen. They are common in moist areas of
Kerala, Kamataka and south M.P. in south and parts of
north U.P., M.P., Bihar, Bengal and Orissa in north. The
well known teak and sal forests belong to this category.
In south India, moist deciduous forests are dominated by
species of Terminalia, Grewelia, Gariya, Salmalia,
Tectona grandis, Adina cordifalia, Melia, Albizzia,
Dalbergia etc. In the northern half, Shorea robusta is
dominant in Gorakhpur and Tarai regions of U.P.,
Assam and north Bengal. Other associates of sal are
Terminalia tomentosa, Dillenia sp., Eugenia sp. etc.
Dry tropical deciduous forests
Most of their trees remain leafless for several weeks in
dry season. In north such forests are found in the Punjab,
U.P., Bihar and Orissa in the regions which are neither
wet nor too dry. Trees are of moderate size (2.5 meters
tall), with sparse canopy. Thorny scrubs, grasses and
some bamboos are also present in some regions. In
Punjab and west U.P. forests, Anogeissus latifolia,
Acacia catechu, Terminalia tomentosa, Bosweliia
serrata are dominants with subdominance of
Dendrocalamus strictus, Emblica officina lis,
Woodfordia floribunda etc. Forests of Shorea robusta
are also found as scattered in wet areas. In south, such
forests are trees with scattered patches of densely
growing grasses intermixed with shrubs. Terminalia.
Anogeissus latifolia, Tectona grandis, Diospyros
melanoxylon, Boswellia sp. fonn the top layer. followed
by smaller plants like Dendrocalamus, Bambusa,
Lantana, Helicteris etc. Common grasses are Panicum,
Andropogon and Heteropogon.
Montane subtropical forests
These are found on. hills of south India, as Nilgiri,
Mahabaleshwar and Pachmarhi, between an altitude of
3,000 to 5,600 ft. These are cooler than the tropical, and
warmer than the temperate forests. In southern parts the
common trees are Eugenia, Actinodaphne, Canthium,
Mangifera and Ficus, and climbers are Piper
trichostachyon, Gnetum scandens, Smilax macrophylla
etc. The northern areas have rather tail trees. In eastern
Himalayas due to higher humidities, bamboos, many
epiphytes including orchids, and ferns become abundant.
Most of the trees are evergreen. The floristic description
of eastern and western Himalayas has already been
given.
Temperate forests
They occur above 5,300 ft altitude, chiefly on mountains
of Himalayas and Nilgiri. In Himalayas, oaks and
conifers are abundant. Their distribution has already
been described. Oaks fonn relatively stable evergreen
pure stands. The southern temperate vegetation is chiefly
represented by the sholas near Ootacamund, Nilgiri hills
in Tamilnadu. The forests are very dense with extensive
growth of grasses and evergreen tall trees, like
UNIT 10: ECOLOGICAL PRINCIPLES 
Balanocarpus utilis, Hopea parviflora, ,Artocarpus
hirsuta, Salmalia malbaricum etc. Their branches are
clothed with mosses, many woody climbers, ferns and
other epiphytes.
Alpine forests
Sometimes they are subdivided as sub-alpine forests,
alpine scrubs, moist alpine scrubs and dry alpine scrubs.
They are extensive throughout the Himalayas above
11,000 ft. The tree height becomes lesser with increasing
altitude, being replaced finally by sparse growth of small
plants like Sedum, Primula, Saxifraga and lichens etc.
The details of the flora are already given in description
of the Himalayas.
Grassland Vegetation
In India, natural grasslands are hardly present, and most
of them are maintained in their present seraI stages due
to biotic influences. Thus, grasslands are not climax
fonnations but have developed secondarily by the
forests' destruction. The two major factors that resulted
into their secondarily development are edaphic and
biotic. The grasslands of India are of three major types:
(i) Xerophilous-occur in dry regions of northwest India
under semi-desert conditions, (ii) mesophilous- called as
savannahs, are extensive grass. flats, typical of moi.st
deciduous forests of D.P., and (iii) hygrophilous - called
as wet savannahs. All these are being controlled under
biotic influences.
Indian grasslands have been classified by Whyte et al
(1954) into eight major types which are shown in Table
2.
Diverse Fauna of India
Like flora our fauna (animal life) is also equally rich and
varied. There are about 77,000 known species. These
I1. Applied ecology: Environmental pollution; global environmental change;
POLLUTION
Pollution is the introduction of contaminants into an
environment that causes instability, disorder, harm or
discomfort to the physical systems or living organisms
they are in. Pollution can take the form of chemical
substances, or energy, such as noise, heat, or light
energy. Pollutants, the elements of pollution, can be
foreign substances or energies, or naturally occurring;
when naturally occurring, they are considered
contaminants when they exceed natural levels. Pollution
is often classed as point source or nonpoint source
pollution.
Sometimes the term pollution is extended to include any
substance when it occurs at such unnaturally high
concentration within a system that it endangers the
stability of that system. For example, water is innocuous
and essential for life, and yet at very high concentration,
it could be considered a pollutant: if a person were to
drink an excessive quantity of water, the physical system
could be so overburdened that breakdown and even
death could result. Another example is the potential of
84 INSTITUTE FOR ADVANCED STUDIES, JODHPUR
include about 53,000 species of insects, 1200 birds, 453
reptiles, 372 mammals, 5000 molluscs, 2,500 fish and
450 reptiles.
Among the mammals we have elephant found in hot wet
equatorial dense forests of Assam, KeraIa and
Kamataka, with heavy rainfall. Camel and wild asses are
found in hot and arid deserts. Camels are common in
Thar desert, the wild asses are confined to arid areas of
the Rann of Kachch. One-homed rhinoceros lives in
swampy and marshy lands of Assam and North Bengal.
An interesting group-Indian bison, the Indian buffalo
and the Nilgai is also found. Chousingha (four-homed
antelope), black buck (Indian antelope), gazel and deer
are an unique group of Indian animals. The deer species
include Kashmir stag, swamp deer, spotted deer, musk
deer and mouse deer.
Indian lion is most distinct, which is found only in India
and African continent in the world. It occurs in Git
forests of Saurashtra in Gujarat. Tiger, the most
powerful species of the forest is also found in India.
Bengal tiger is found in the Sunderbans. Other animals
of cat family are leopards, clouded leopards, and snow
leopards. The latter are confined to the upper reaches of
Himalayas.
Several interesting animals live in the Himalayan ranges.
Chief species include wild sheep, mountain goats, ibex,
shrew and tapir. Panda and snow leopard are also found
here. There are several species of monkey in India, of
which the langur is the most common. Lion-tailed
macaque has hair around the face. Bird life in India is
both rich and colourful. Tiger is the national animal,
peacock is national bird. Pheasants, geese, ducks,
mynahs, parakeets, pigeons, cranes, hornbills and sun
birds are found in forests and wetlands.
 
excessive noise to induce imbalance in a person's mental
state, resulting in malfunction and psychosis;
Growing evidence of local and global pollution and an
increasingly informed public over time have given rise
to environmentalism and the environmental movement,
which generally seek to limit human impact on the
environment.
Pollution control
Pollution control is a term used in environmental
management. It means the control of emissions and
effluents into air, water or soil. Without pollution
control, the waste products from consumption, heating,
agriculture, mining, manufacturing, transportation and
other human activities, whether they accumulate or
disperse, will degrade the environment. In the hierarchy
of controls, pollution prevention and waste minimization
are more desirable than pollution control.
 
UNIT 10: ECOLOGICAL PRINCIPLES 
Pollution control devices
• Dust collection systems
o Cyclones
o Electrostatic precipitators
o Baghouses
• Scrubbers
o Baffle spray scrubber
o Cyclonic spray scrubber
o Ejector venturi scrubber
o Mechanically aided scrubber
o Spray tower
o Wet scrubber
• Sewage treatment and Wastewater treatment
o API oil-water separators
o Sedimentation (water treatment)
o Dissolved air flotation (DAF)
o Activated sludge biotreaters
o Biofilters
o Powdered activated carbon treatment
• Vapor recovery systems
Major forms of pollution and major polluted areas
The major forms of pollution are listed below along with
the particular pollutants relevant to each of them:
• Air pollution, the release of chemicals and particulates
into the atmosphere. Common gaseous air pollutants
include carbon monoxide, sulfur dioxide,
chlorofluorocarbons (CFCs) and nitrogen oxides
produced by industry and motor vehicles. Photochemical
ozone and smog are created as nitrogen oxides and
hydrocarbons react to sunlight. Particulate matter,
characterized by size PM10 to PM2.5, is produced from
natural sources such as volcanoes or as residual oil fly
ash from power plants. Diesel particles are another class
of airborne particulate matter.
• Water pollution, by the release of waste products and
contaminants into surface runoff into river drainage
systems, leaching into groundwater, liquid spills,
wastewater discharges, eutrophication and littering.
• Soil contamination occurs when chemicals are
released by spill or underground leakage. Among the
most significant soil contaminants are hydrocarbons,
heavy metals, MTBE, herbicides, pesticides and
chlorinated hydrocarbons.
• Radioactive contamination, resulting from 20th
century activities in atomic physics, such as nuclear
power generation and nuclear weapons research,
manufacture and deployment. (See alpha emitters and
actinides in the environment.)
• Noise pollution, which encompasses roadway noise,
aircraft noise, industrial noise as well as high-intensity
sonar.
• Light pollution, includes light trespass, over-
illumination and astronomical interference.
• Visual pollution, which can refer to the presence of
overhead power lines, motorway billboards, scarred
landforms (as from strip mining), open storage of trash
or municipal solid waste.
• Thermal pollution, is a temperature change in natural
water bodies caused by human influence, such as use of
water as coolant in a power plant.
The Blacksmith Institute issues annually a list of the
world's worst polluted places. In the 2007 issues the ten
top nominees are located in Azerbaijan, China, India,
Peru, Russia, Ukraine and Zambia.
INSTITUTE FOR ADVANCED STUDIES, JODHPUR  85
Sources and causes
Motor vehicle emissions are one of the leading causes of
air pollution. China, United States, Russia, Mexico, and
Japan are the world leaders in air pollution emissions;
however, Canada is the number two country, ranked per
capita. Principal stationary pollution sources include
chemical plants, coal-fired power plants, oil refineries,
petrochemical plants, nuclear waste disposal activity,
incinerators, large livestock farms (dairy cows, pigs,
poultry, etc.), PVC factories, metals production factories,
plastics factories, and other heavy industry.
Some of the more common soil contaminants are
chlorinated hydrocarbons (CFH), heavy metals (such as
chromium, cadmium--found in rechargeable batteries,
and lead--found in lead paint, aviation fuel and still in
some countries, gasoline), MTBE, zinc, arsenic and
benzene. In 2001 a series of press reports culminating in
a book called Fateful Harvest unveiled a widespread
practice of recycling industrial byproducts into fertilizer,
resulting in the contamination of the soil with various
metals. Ordinary municipal landfills are the source of
many chemical substances entering the soil environment
(and often groundwater), emanating from the wide
variety of refuse accepted, especially substances illegally
discarded there, or from pre-1970 landfills that may have
been subject to little control in the U.S. or EU. There
have also been some unusual releases of polychlorinated
dibenzodioxins, commonly called dioxins for simplicity,
such as TCDD.
Pollution can also be the consequence of a natural
disaster. For example, hurricanes often involve water
contamination from sewage, and petrochemical spills
from ruptured boats or automobiles. Larger scale and
environmental damage is not uncommon when coastal
oil rigs or refineries are involved. Some sources of
pollution, such as nuclear power plants or oil tankers,
can produce widespread and potentially hazardous
releases when accidents occur.
In the case of noise pollution the dominant source class
is the motor vehicle, producing about ninety percent of
all unwanted noise worldwide.
Effects
Human health: Adverse air quality can kill many
organisms including humans. Ozone pollution can cause
respiratory disease, cardiovascular disease, throat
inflammation, chest pain, and congestion. Water
pollution causes approximately 14,000 deaths per day,
mostly due to contamination of drinking water by
untreated sewage in developing countries. Oil spills can
cause skin irritations and rashes. Noise pollution induces
hearing loss, high blood pressure, stress, and sleep
disturbance. Mercury has been linked to developmental
deficits in children and neurologic symptoms. Lead and
other heavy metals have been shown to cause
neurological problems. Chemical and radioactive
substances can cause cancer and as well as birth defects.
Ecosystems
• Sulfur dioxide and oxides of nitrogen can cause acid
rain which reduces the pH value of soil.
UNIT 10: ECOLOGICAL PRINCIPLES 
• Soil can become infertile and unsuitable for plants.
This will affect other organisms in the food web.
• Smog and haze can reduce the amount of sunlight
received by plants to carry out photosynthesis.
• Invasive species can out compete native species and
reduce biodiversity. Invasive plants can contribute debris
and biomolecules (allelopathy) that can alter soil and
chemical compositions of an environment, often
reducing native species competitiveness.
• Biomagnification describes a situation where toxins
may pass through trophic levels, becoming exponentially
more concentrated in the process.
• Ocean acidification, the ongoing decrease in the pH of
the Earth's oceans.
• Global warming.
AIR POLLUTIION
Air pollution is the human introduction into the
atmosphere of chemicals, particulate matter, or
biological materials that cause harm or discomfort to
humans or other living organisms, or damages the
environment. Air pollution causes deaths and respiratory
disease. Air pollution is often identified with major
stationary sources, but the greatest source of emissions is
mobile sources, mainly automobiles. Gases such as
carbon dioxide, which contribute to global warming,
have recently gained recognition as pollutants by climate
scientists, while they also recognize that carbon dioxide
is essential for plant life through photosynthesis.
The atmosphere is a complex, dynamic natural gaseous
system that is essential to support life on planet Earth.
Stratospheric ozone depletion due to air pollution has
long been recognized as a threat to human health as well
as to the Earth's ecosystems.
Pollutants
There are many substances in the air which may impair
the health of plants and animals (including humans), or
reduce visibility. These arise both from natural processes
and human activity. Substances not naturally found in
the air or at greater concentrations or in different
locations from usual are referred to as pollutants.
Pollutants can be classified as either primary or
secondary. Usually, primary pollutants are substances
directly emitted from a process, such as ash from a
volcanic eruption, the carbon monoxide gas from a
motor vehicle exhaust or sulfur dioxide released from
factories.
Secondary pollutants are not emitted directly. Rather,
they form in the air when primary pollutants react or
interact. An important example of a secondary pollutant
is ground level ozone - one of the many secondary
pollutants that make up photochemical smog.
Note that some pollutants may be both primary and
secondary: that is, they are both emitted directly and
formed from other primary pollutants.
Major primary pollutants produced by human activity
include:
• Sulfur oxides (SOx) especially sulfur dioxide are
emitted from burning of coal and oil.
86 INSTITUTE FOR ADVANCED STUDIES, JODHPUR
• Nitrogen oxides (NOx) especially nitrogen dioxide
are emitted from high temperature combustion. Can
be seen as the brown haze dome above or plume
downwind of cities.
• Carbon monoxide is colourless, odourless, non-
irritating but very poisonous gas. It is a product by
incomplete combustion of fuel such as natural gas,
coal or wood. Vehicular exhaust is a major source of
carbon monoxide.
• Carbon dioxide (CO2), a greenhouse gas emitted
from combustion.
• Volatile organic compounds (VOC), such as
hydrocarbon fuel vapors and solvents.
• Particulate matter (PM), measured as smoke and
dust. PM10 is the fraction of suspended particles 10
micrometers in diameter and smaller that will enter
the nasal cavity. PM2.5 has a maximum particle size
of 2.5 µm and will enter the bronchies and lungs.
• Toxic metals, such as lead, cadmium and copper.
• Chlorofluorocarbons (CFCs), harmful to the ozone
layer emitted from products currently banned from
use.
• Ammonia (NH3) emitted from agricultural
processes.
• Odors, such as from garbage, sewage, and industrial
processes
• Radioactive pollutants produced by nuclear
explosions, war explosives, and natural processes such
as the radioactive decay of radon.
Secondary pollutants include:
• Particulate matter formed from gaseous primary
pollutants and compounds in photochemical smog,
such as nitrogen dioxide.
• Ground level ozone (O3) formed from NOx and
VOCs.
• Peroxyacetyl nitrate (PAN) similarly formed from
NOx and VOCs.
Minor air pollutants include:
• A large number of minor hazardous air pollutants.
Some of these are regulated in USA under the Clean
Air Act and in Europe under the Air Framework
Directive.
• A variety of persistent organic pollutants, which
can attach to particulate matter.
Sources
Sources of air pollution refer to the various locations,
activities or factors which are responsible for the
releasing of pollutants in the atmosphere. These sources
can be classified into two major categories which are:
Anthropogenic sources (human activity) mostly
related to burning different kinds of fuel
• "Stationary Sources" as smoke stacks of power plants,
manufacturing facilities, municipal waste incinerators.
• "Mobile Sources" as motor vehicles, aircraft etc.
• Marine vessels, such as container ships or cruise
ships, and related port air pollution.
• Burning wood, fireplaces, stoves, furnaces and
incinerators .
• Oil refining, and industrial activity in general.
• Chemicals, dust and controlled burn practices in
agriculture and forestry management.
 
UNIT 10: ECOLOGICAL PRINCIPLES 
• Fumes from paint, hair spray, varnish, aerosol sprays
and other solvents.
• Waste deposition in landfills, which generate
methane.
• Military, such as nuclear weapons, toxic gases, germ
warfare and rocketry.
Natural sources
• Dust from natural sources, usually large areas of land
with little or no vegetation.
• Methane, emitted by the digestion of food by animals,
for example cattle.
• Radon gas from radioactive decay within the Earth's
crust.
• Smoke and carbon monoxide from wildfires.
• Volcanic activity, which produce sulfur, chlorine, and
ash particulates.
WATER POLLUTION
Water pollution is the contamination of water bodies
such as lakes, rivers, oceans, and groundwater caused by
human activities, which can be harmful to organisms and
plants which live in these water bodies.
Although natural phenomena such as volcanoes, algae
blooms, storms, and earthquakes also cause major
changes in water quality and the ecological status of
water, water is typically referred to as polluted when it
impaired by anthropogenic contaminants and either does
not support a human use (like serving as drinking water)
or undergoes a marked shift in its ability to support its
constituent biotic communities. Water pollution has
many causes and characteristics. The primary sources of
water pollution are generally grouped into two categories
based on their point of origin. Point-source pollution
refers to contaminants that enter a waterway through a
discrete "point source". Examples of this category
include discharges from a wastewater treatment plant,
outfalls from a factory, leaking underground tanks, etc.
The second primary category, non-point source
pollution, refers to contamination that, as its name
suggests, does not originate from a single discrete
source. Non-point source pollution is often a cumulative
effect of small amounts of contaminants gathered from a
large area. Nutrient runoff in storm water from sheet
flow over an agricultural field, or metals and
hydrocarbons from an area with high impervious
surfaces and vehicular traffic are examples of non-point
source pollution. The primary focus of legislation and
efforts to curb water pollution for the past several
decades was first aimed at point sources. As point
sources have been effectively regulated, greater attention
has come to be placed on non-point source contributions,
especially in rapidly urbanizing/suburbanizing or
developing areas.
The specific contaminants leading to pollution in water
include a wide spectrum of chemicals, pathogens, and
physical or sensory changes. While many of the
chemicals and substances that are regulated may be
naturally occurring (iron, manganese, etc) the
concentration is often the key in determining what is a
natural component of water, and what is a contaminant.
Many of the chemical substances are toxic. Pathogens
can produce waterborne diseases in either human or
INSTITUTE FOR ADVANCED STUDIES, JODHPUR  87
animal hosts. Alteration of water's physical chemistry
include acidity, electrical conductivity, temperature, and
eutrophication. Eutrophication is the fertilization of
surface water by nutrients that were previously scarce.
Water pollution is a major problem in the global context.
It has been suggested that it is the leading worldwide
cause of deaths and diseases, and that it accounts for the
deaths of more than 14,000 people daily.
Contaminants
Contaminants may include organic and inorganic
substances.
Some organic water pollutants are:
• Insecticides and herbicides, a huge range of
organohalides and other chemicals
• Bacteria, often is from sewage or livestock operations
• Food processing waste, including pathogens
• Tree and brush debris from logging operations
• VOCs (volatile organic compounds), such as
industrial solvents, from improper storage
• DNAPLs (dense non-aqueous phase liquids), such as
chlorinated solvents, which may fall at the bottom of
reservoirs, since they don't mix well with water and are
more dense
• Petroleum Hydrocarbons including fuels (gasoline,
diesel, jet fuels, and fuel oils) and lubricants (motor oil)
from oil field operations, refineries, pipelines, retail
service station's underground storage tanks, and transfer
operations. Note: VOCs include gasoline-range
hydrocarbons.
• Detergents
• Various chemical compounds found in personal
hygiene and cosmetic products
• Disinfection by-products (DBPs) found in chemically
disinfected drinking water
Some inorganic water pollutants include:
• Spill of oil over the seas is the biggest danger.
• Heavy metals including acid mine drainage
• Acidity caused by industrial discharges (especially
sulfur dioxide from power plants)
• Pre-production industrial raw resin pellets, an
industrial pollutant
• Chemical waste as industrial by products
• Fertilizers, in runoff from agriculture including
nitrates and phosphates
• Silt in surface runoff from construction sites, logging,
slash and burn practices or land clearing sites
Macroscopic, that is, large visible items polluting the
water are termed marine debris, and can include such
items as:
• Nurdles, small ubiquitous waterborne plastic pellets
• Shipwrecks, large derelict ships
GLOBAL WARMING
The term "global warming" refers to the warming in
recent decades and its projected continuation, and
implies a human influence.The United Nations
Framework Convention on Climate Change (UNFCCC)
uses the term "climate change" for human-caused
change, and "climate variability" for other changes. The
term "climate change" recognizes that rising
temperatures are not the only effect. The term
UNIT 10: ECOLOGICAL PRINCIPLES 
"anthropogenic global warming" is sometimes used
when focusing on human-induced changes.
Causes
Components of the current radiative forcing as estimated
by the IPCC Fourth Assessment Report.Main articles:
Attribution of recent climate change and Scientific
opinion on climate change. The Earth's climate changes
in response to external forcing, including variations in its
orbit around the Sun (orbital forcing), changes in solar
luminosity, volcanic eruptions, and atmospheric
greenhouse gas concentrations. The detailed causes of
the recent warming remain an active field of research,
but the scientific consensus is that the increase in
atmospheric greenhouse gases due to human activity
caused most of the warming observed since the start of
the industrial era. This attribution is clearest for the most
recent 50 years, for which the most detailed data are
available. Some other hypotheses departing from the
consensus view have been suggested to explain most of
the temperature increase. One such hypothesis proposes
that warming may be the result of variations in solar
activity.
None of the effects of forcing are instantaneous. The
thermal inertia of the Earth's oceans and slow responses
of other indirect effects mean that the Earth's current
climate is not in equilibrium with the forcing imposed.
Climate commitment studies indicate that even if
greenhouse gases were stabilized at 2000 levels, a
further warming of about 0.5 °C (0.9 °F) would still
occur.
Greenhouse gases in the atmosphere
The greenhouse effect was discovered by Joseph Fourier
in 1824 and was first investigated quantitatively by
Svante Arrhenius in 1896. It is the process by which
absorption and emission of infrared radiation by
atmospheric gases warm a planet's lower atmosphere and
surface.
Recent increases in atmospheric carbon dioxide (CO2).
The monthly CO2 measurements display small seasonal
oscillations in an overall yearly uptrend; each year's
maximum is reached during the Northern Hemisphere's
late spring, and declines during the Northern
Hemisphere growing season as plants remove some CO2
from the atmosphere.Existence of the greenhouse effect
as such is not disputed. Naturally occurring greenhouse
gases have a mean warming effect of about 33 °C (59
88 INSTITUTE FOR ADVANCED STUDIES, JODHPUR
°F), without which Earth would be uninhabitable. On
Earth, the major greenhouse gases are water vapor,
which causes about 36–70 percent of the greenhouse
effect (not including clouds); carbon dioxide (CO2),
which causes 9–26 percent; methane (CH4), which
causes 4–9 percent; and ozone, which causes 3–7
percent. The issue is how the strength of the greenhouse
effect changes when human activity increases the
atmospheric concentrations of some greenhouse gases.
Human activity since the industrial revolution has
increased the concentration of various greenhouse gases,
leading to increased radiative forcing from CO2,
methane, tropospheric ozone, CFCs and nitrous oxide.
Molecule for molecule, methane is a more effective
greenhouse gas than carbon dioxide, but its
concentration is much smaller so that its total radiative
forcing is only about a fourth of that from carbon
dioxide. Some other naturally occurring gases contribute
small fractions of the greenhouse effect; one of these,
nitrous oxide (N2O), is increasing in concentration
owing to human activity such as agriculture. The
atmospheric concentrations of CO2 and CH4 have
increased by 31% and 149% respectively since the
beginning of the industrial revolution in the mid-1700s.
These levels are considerably higher than at any time
during the last 650,000 years, the period for which
reliable data has been extracted from ice cores. From
less direct geological evidence it is believed that CO2
values this high were last attained 20 million years ago.
Fossil fuel burning has produced approximately three-
quarters of the increase in CO2 from human activity over
the past 20 years. Most of the rest is due to land-use
change, in particular deforestation.
The present atmospheric concentration of CO2 is about
385 parts per million (ppm) by volume. Future CO2
levels are expected to rise due to ongoing burning of
fossil fuels and land-use change. The rate of rise will
depend on uncertain economic, sociological,
technological, and natural developments, but may be
ultimately limited by the availability of fossil fuels. The
IPCC Special Report on Emissions Scenarios gives a
wide range of future CO2 scenarios, ranging from 541 to
970 ppm by the year 2100. Fossil fuel reserves are
sufficient to reach this level and continue emissions past
2100, if coal, tar sands or methane clathrates are
extensively used.
Feedbacks
One of the most pronounced feedback effects relates to
the evaporation of water. Warming by the addition of
long-lived greenhouse gases such as CO2 will cause
more water to evaporate into the atmosphere. Since
water vapor is a greenhouse gas, the atmosphere warms
further; this warming causes more water vapor to
evaporate (a positive feedback), and so on until other
processes stop the feedback loop. The result is a much
larger greenhouse effect than that due to CO2 alone.
Although this feedback process causes an increase in the
absolute moisture content of the air, the relative
humidity stays nearly constant or even decreases slightly
because the air is warmer. This feedback effect can only
be reversed slowly as CO2 has a long average
atmospheric lifetime.
 
UNIT 10: ECOLOGICAL PRINCIPLES 
Feedback effects due to clouds are an area of ongoing
research. Seen from below, clouds emit infrared
radiation back to the surface, and so exert a warming
effect; seen from above, clouds reflect sunlight and emit
infrared radiation to space, and so exert a cooling effect.
Whether the net effect is warming or cooling depends on
details such as the type and altitude of the cloud. These
details are difficult to represent in climate models, in
part because clouds are much smaller than the spacing
between points on the computational grids of climate
models.
Northern Hemisphere ice trends
Southern Hemisphere ice trends.A subtler feedback
process relates to changes in the lapse rate as the
atmosphere warms. The atmosphere's temperature
decreases with height in the troposphere. Since emission
of infrared radiation varies with the fourth power of
temperature, longwave radiation emitted from the upper
atmosphere is less than that emitted from the lower
atmosphere. Most of the radiation emitted from the
upper atmosphere escapes to space, while most of the
radiation emitted from the lower atmosphere is re-
absorbed by the surface or the atmosphere. Thus, the
strength of the greenhouse effect depends on the
atmosphere's rate of temperature decrease with height: if
the rate of temperature decrease is greater the
greenhouse effect will be stronger, and if the rate of
temperature decrease is smaller then the greenhouse
effect will be weaker. Both theory and climate models
indicate that warming will reduce the decrease of
temperature with height, producing a negative lapse rate
feedback that weakens the greenhouse effect.
Measurements of the rate of temperature change with
height are very sensitive to small errors in observations,
making it difficult to establish whether the models agree
with observations.
Another important feedback process is ice-albedo
feedback. When global temperatures increase, ice near
the poles melts at an increasing rate. As the ice melts,
land or open water takes its place. Both land and open
water are on average less reflective than ice, and thus
absorb more solar radiation. This causes more warming,
which in turn causes more melting, and this cycle
continues.
Positive feedback due to release of CO2 and CH4 from
thawing permafrost, such as the frozen peat bogs in
Siberia, is an additional mechanism that could contribute
to warming. Similarly a massive release of CH4 from
methane clathrates in the ocean could cause rapid
warming, according to the clathrate gun hypothesis.
The ocean's ability to sequester carbon is expected to
decline as it warms. This is because the resulting low
nutrient levels of the mesopelagic zone (about 200 to
1000 m depth) limits the growth of diatoms in favor of
smaller phytoplankton that are poorer biological pumps
of carbon.
INSTITUTE FOR ADVANCED STUDIES, JODHPUR  89
Temperature changes
Two millennia of mean surface temperatures according
to different reconstructions, each smoothed on a decadal
scale. The unsmoothed, annual value for 2004 is also
plotted for reference. Global temperatures have
increased by 0.75 °C (1.35 °F) relative to the period
1860–1900, according to the instrumental temperature
record. This measured temperature increase is not
significantly affected by the urban heat island effect.
Since 1979, land temperatures have increased about
twice as fast as ocean temperatures (0.25 °C per decade
against 0.13 °C per decade). Temperatures in the lower
troposphere have increased between 0.12 and 0.22 °C
(0.22 and 0.4 °F) per decade since 1979, according to
satellite temperature measurements. Temperature is
believed to have been relatively stable over the one or
two thousand years before 1850, with possibly regional
fluctuations such as the Medieval Warm Period or the
Little Ice Age.
Sea temperatures increase more slowly than those on
land both because of the larger effective heat capacity of
the oceans and because the ocean can lose heat by
evaporation more readily than the land. The Northern
Hemisphere has more land than the Southern
Hemisphere, so it warms faster. The Northern
Hemisphere also has extensive areas of seasonal snow
and sea-ice cover subject to the ice-albedo feedback.
More greenhouse gases are emitted in the Northern than
Southern Hemisphere, but this does not contribute to the
difference in warming because the major greenhouse
gases persist long enough to mix between hemispheres.
Based on estimates by NASA's Goddard Institute for
Space Studies, 2005 was the warmest year since reliable,
widespread instrumental measurements became
available in the late 1800s, exceeding the previous
record set in 1998 by a few hundredths of a degree.
Estimates prepared by the World Meteorological
Organization and the Climatic Research Unit concluded
that 2005 was the second warmest year, behind 1998.
Temperatures in 1998 were unusually warm because the
strongest El Niño-Southern Oscillation in the past
century occurred during that year.
Anthropogenic emissions of other pollutants—notably
sulfate aerosols—can exert a cooling effect by increasing
the reflection of incoming sunlight. This partially
UNIT 10: ECOLOGICAL PRINCIPLES 

accounts for the cooling seen in the temperature record
in the middle of the twentieth century,[62] though the
cooling may also be due in part to natural variability.
James Hansen and colleagues have proposed that the
effects of the products of fossil fuel combustion—CO2
and aerosols—have largely offset one another, so that
warming in recent decades has been driven mainly by
non-CO2 greenhouse gases.
have resulted in more warming than observed if not for
these dimming agents.
Ozone depletion, the steady decline in the total amount
of ozone in Earth's stratosphere, is frequently cited in
relation to global warming. Although there are areas of
linkage, the relationship between the two is not strong.

Related climatic issues

A variety of issues are often raised in relation to global

warming. One is ocean acidification. Increased
atmospheric CO2 increases the amount of CO2 dissolved
in the oceans. CO2 dissolved in the ocean reacts with
water to form carbonic acid, resulting in acidification.
Ocean surface pH is estimated to have decreased from
8.25 near the beginning of the industrial era to 8.14 by
2004, and is projected to decrease by a further 0.14 to
0.5 units by 2100 as the ocean absorbs more CO2. Since
organisms and ecosystems are adapted to a narrow range
of pH, this raises extinction concerns, directly driven by
increased atmospheric CO2, that could disrupt food
webs and impact human societies that depend on marine
ecosystem services.

Global dimming, the gradual reduction in the amount of

global direct irradiance at the Earth's surface, may have
partially mitigated global warming in the late 20th
century. From 1960 to 1990 human-caused aerosols
likely precipitated this effect. Scientists have stated with
66–90% confidence that the effects of human-caused
aerosols, along with volcanic activity, have offset some
of the global warming, and that greenhouse gases would
Figure: Recent increases in atmospheric carbon dioxide
(CO2). The monthly CO2 measurements display small
seasonal oscillations in an overall yearly uptrend; each
year's maximum is reached during the Northern
Hemisphere's late spring, and declines during the
Northern Hemisphere growing season as plants remove
some CO2 from the atmosphere.

90 INSTITUTE FOR ADVANCED STUDIES, JODHPUR 
UNIT 10: ECOLOGICAL PRINCIPLES 
Biodiversity-status, monitoring and documentation; major drivers of biodiversity change;
biodiversity management approaches.
Introduction
The biosphere (the web of life that lives within and
depends upon the inorganic spheres) constitutes a vital
life support system for man and its existence in a healthy
and functional state is essential for the existence of
human race. It is the collection of innumerable
organisms, the biological diversity, which makes our life
pleasant and possible. The term biodiversity was coined
by Walter and Rosen (1985) and is the abbreviated word
for Biological Diversity. Life originated on earth almost
four billion years ago and nature took more than 1 billion
years to develop this wide and complex spectrum of life
on earth. Scientists believe that the total number of
species on earth is in between 10-80 million (Wilson
1988) of which 1.4 million species have been enlisted so
far. However, we are losing this heritage of millions of
years at a very fast rate. The reduction in diversity in life
forms is bound to have grave consequences for the entire
living world. It has become extremely important to study
the various life forms on earth and the causes of their
destruction.
Biodiversity is the total variety of life on our planet. The
total number of races, varieties or species i.e., the sum
total of various types of microbes, plants and animals
present in a system is referred to as biodiversity. The
word biodiversity is now very widely used not only by
the scientific community, but also by the common
people, environmental groups, conservationists,
industrialists and economists. So it is very important to
have clear idea about the definition of biodiversity which
is recognized as a separate science with its own
principles.
Some of the important definitions of biodiversity are
given here-
(i) Biodiversity is the variety of life in all its forms,
levels and combinations. It includes species
diversity, genetic diversity and ecosystem diversity
(International Union for Conservation of Nature and
Natural Resources-lUCN).
(ii) United Nations Earth Summit in Rio de Janeiro
defined biodiversity as - The variability among
living organisms from all sources, including
terrestrial, marine and other aquatic ecosystems and
the ecological complexes of which they are part.
This includes diversity within species, between
species and of ecosystems.
(iii) According to U.S. Congressional Biodiversity Act -
Biological Diversity is the variety and variability
among living organisms and the ecological
complexes in which they occur and encompasses
ecosystem diversity, species diversity and genetic
diversity.
(iv) In the simplest terms, biological diversity is the
variety of life and its processes and' it includes the
variety of living organisms, the genetic differences
among them and the communities and ecosystems
in which they occur. In this article we will present
some fundamental aspects of biodiversity and its
conservation.
INSTITUTE FOR ADVANCED STUDIES, JODHPUR  91
Types of biodiversity
Biodiversity is usually studied at three different levels-
Species diversity, Genetic diversity and Ecosystem
diversity. .
(i) Species diversity: Evolution of species diversity has
probably been possible because of habitat diversity on
earth. It refers to the variety of species within a region.
This diversity could be measured on the basis of number
of species in a region. The term biodiversity is
commonly used as a synonym of species diversity. It
actually refers to species richness, in terms of number of
species in a site or habitat.
Global diversity is typically represented in terms of total
number of species of different taxonomic groups. As
mentioned before, an estimated 1.4 million species have
been identified to date.
Species diversity, again, is studied at three levels:
ƒ alpha diversity (number of species coexisting
at a site),
ƒ beta diversity (difference in species
complement between patches) and
ƒ gamma diversity (number of species in a large
area, e.g. a country).
This series can further be extended to delta diversity for
biomes and omega diversity for the entire biosphere.
Some authors call it taxon diversity (variety of taxa
within a community of an area). It is generally studied at
the species level and hence called species diversity.
When the taxonomic levels such as genus and family are
considered, toe term taxon diversity is more
appropriate.This term is similar to taxic diversity.
(ii) Genetic diversity: Within a species there are a
number of subspecies, varieties (subspecies and varieties
are recognizable morphological variations within a
species), forms (form is generally used to recognize and
describe sporadic variations in a single morphological
feature) or strains' which slightly differ from each other.
These differences are due to slight variations in their
genetic organization. This diversity in the genetic make
up of a species is referred to as genetic diversity.
A species with a large number of varieties or- strains is
considered to be rich and diverse in its genetic
organization. Genetic variations arise in individuals of a
species by genic or chromosomal mutations. Genetic
variation within populations is considered a "prerequisite
for adaptation and evolutionary change", and as such an
important aspect of biodiversity. Genetic variation is
often expressed in terms of alleles (genes occupying the
same locus in a chromosome) and is mainly studied at
the population level. Genetic variations can be measured
by different recent techniques such as allozyme analysis,
DNA fingerprinting, polymerase chain reaction,
restriction site mapping and DNA sequencing.
UNIT 10: ECOLOGICAL PRINCIPLES 
Diversities go on increasing at the microlevel.
Differences in the level of varieties are followed by
differences among the subspecies, varieties and species.
Accumulation of these differences at infra-specific level
will automatically lead to distinctive character at the
species level.
(iii) Ecosystem diversity: In ecosystem, there may exist
different land-forms, each of which supports different
and specific vegetation. Ecosystem diversity in contrast
to genetic and species diversity is difficult to measure
since the boundaries of the communities which constitute
the various sub-ecosystems are not distinct. Ecosystem
diversity could best be understood if one studies the
communities in various ecological niches within the
given ecosystem, each community is associated with
definite complexes. These complexes are related to
composition and structure of biodiversity. Loss of
ecosystem diversity may be considered as ultimate cause
of loss of species and genetic diversity. Community
diversity is a synonym of ecosystem diversity and is
defined as the diversity of community types within larger
areas (ecological units). It should not be confused with
habitat variety which is an expression mainly used for
different species of animals which have different
habitats.
Evolution of Biodiversity
Darwin (1859) proposed that species compete and only
the fittest survive in nature. It is inferred that under a
strong pressure of natural selection the less fit species are
eliminated. From this concept has arisen the competitive
exclusion principle (Hardin 1960), which is based on the
idea that no two species can be exactly equally fit. It
states that if two or more species exist in the same
habitat, ultimately all but one of them will be excluded.
This is the paradox of biodiversity: we expect few
species but we find many' in nature. Mechanisms that
may be responsible for preventing loss of species by
competitive exclusion, and allow species diversity to be
maintained are as follows (Newman 2000):
(1) Each species has an exclusive ecological niche, and
subjected to conditions where it is fitter than, its
competitors.
(2) A perfect balance is maintained between species loss
and gain. The slightly less fit species are eliminated by
competitive exclusion, but this process is so slow that
there will be time for other' species to arise by evolution
or to invade from other region.
(3) Competition is reduced or prevented, because the
main controls on abundance are physical disturbance,
stresses (e.g. low temperature, toxic substances),
predation and disease, hence competitive exclusion does
not occur.
Factors that promote high diversity
Biodiversity varies greatly from site to site, over both
large and small distances. This variation is due to certain
factors that we discuss here.
(i) Favourable environmental conditions: It is quite
natural to think that species diversity would be greater
where the conditions for growth are very favourable for
plants and animals. But this is not universally true.
Curriae (1991) studied the numbers of species of trees,
92 INSTITUTE FOR ADVANCED STUDIES, JODHPUR
mammals, birds, reptil~s and ampl}ibians across the
USA and Canada in relation to environmental conditions.
Species richness' of each group showed positive
correlation with temperature and solar influx. In relation
to these two factors biodiversity increases with
increasing favorableness. The results from the long-term
Park Grass Experiment at Rothamsted, England, showed
that the more favourable the nutrient regime for plant
growth - the lower was the biodiversity. Grime (1973)
proposed the highest diversity at intermediate stress or
favourableness. Therefore, it can be inferred that there is
no universal relationship between species-diversity and
the prevalence of favourable condition.
(ii) Reducing soil fertility: There is a negative
relationship between diversity and soil fertility
(Newman, 2000). It is necessary to reduce soil fertility to
achieve high species diversity in grasslands. The low
divers~ty in high productivity grasslands is because a
few species grow tall, there is intense competition for
light and low growing species are eliminated. .
(iii) Disturbance: Minor disturbances help to maintain
local species diversity. Disturbance of forests by felling
of trees, fires affects the subsequent species composition.
Grazing animals in grasslands can be considered
disturbance and they can increase diversity. In the past
tropical rainforests had been used for shifting cultivation.
The diversity was initiated as a response to such
disturbance. The present day forest represent a mosaic of
small patches at different stages of succession following
disturbance that provides niches and contributes to
diversity (Connell 1979). The conclusion is that
disturbance can augment biodiversity and diversity
managers need to consider carefully what disturbance to
allow or introduce.
(iv) Heterogeneity of the environment: Environmental
heterogeneity increases β-divershy but has. no such
effect on α-diversity (Newman 2000). On a landscape
scale patches and mosaics of varying vegetation can be
related to differences in exposure, steepness, soil depth,
wetness, rock type affecting soil properties and other
factors of microclimate and soil. Each species responds
differently to the environmental factors and so the
proportions of species change. Whittaker (1956) showed
that each woody species in the Great Smoky Mountains
had a different distribution to altitude and exposure. So if
we want to promote β-diversity we should pay attention
to heterogeneity in the physical environment.
(v) Plant species diversity may promote insect
diversity(Co-evolution): High plant diversity may
.promote insect diversity. This is primarily because of
coevolution between plants and insects involving
secondary chemicals (Harbome 1993). The most of the
secondary chemicals in plants such as alkaloids,
terpenoids and flavonoids, are poisonous to most
animals. However, there are examples where one insect
species being tolerant to one secondary chemical. This
gives the insect the ability to eat something that most
other insects cannot eat and it may then specialize in
eating one plant species. Thus many insects eat only one
.or a few plant species. Other herbivorous animals tend
to show less specificity in their diet. They show
preferences between plant species but rarely confine their
feeding to one plant species. Thus plant species diversity
is likely to promote diversity of insects, but not
necessarily of other animals.
 
UNIT 10: ECOLOGICAL PRINCIPLES 

Global biodiversity (ii) Total number of species of animals

1.4 million species of various life-forms have been
enlisted so far. These include 300,000 species of vascular
plants; 40,000 species of vertebrates; 800,000 species of
insects; 360,000 species of microorganisms. However,
these estimates are increasing all the time. A
conservative estimate of the total figure would be 14
million. According to some recent estimates, the number
of insects alone may be as high as 10 million. This means
that only 13% of the total biodiversity on earth has been
described.
(i) Total number of species of plants and microbes
Invertebrates as a whole includes little over one million
species, distributed among nine major phyla. These
include 800,000 species of insects, 5,000 species of
sponges, 38,000 species of molluscs and 9,000 species of
crustaceans (Table 1). The number of species of
vertebrates in the world is about 40,000. These include
2,400 species of amphibians, 6,100 species of fish, 6,300
species of reptiles, 9,035 species of birds and 4,230
species of mammals.
Table 1: Detailed distribution of species spread
worldwide

ƒ Viruses-It is evident from Table 1 that about S. No Group Number of Species

4,000 different viruses are known to date. A
conservative estimate of the total number of
1 Virus 4,000
viruses is 400,000. Thus, we know only 1 % of
the existing viruses.
ƒ Bacteria-About 5,300 species of bacteria 2 Bacteria 3,600
(including 1,700 species of cyanobacteria)
have been recorded so far. According to an 3 Cyanobacteria 1,700
estimate total number of bacterial species on
the Earth is around 400,000. 4 Fungi 46,983
ƒ Fungi and Lichens-More than. 70,000 species
of fungi (including about 17,000 species of 5 Lichens 17,000
lichens) are known to us. The conservative
figure for existing fungal species on the Earth 6 Algae 40,000
is about 1.5 million.
ƒ Algae-So far 40,000 species have been 7 Bryophytes 19,000
described and another 300,000 species are
believed to exist on the Earth. Algae yet to be 8 Pteridophytes 15,000
described are likely to come from open ocean
as well as from polar regions. 9 Gymnosperms 750
ƒ Bryophytes-So far 19,000 species of
bryophytes are known, of Which about 10,000
10 Angiosperms 300,000
species are mosses, 8,000 species liverworts
and 800 species of horn worts. The number of
bryophyte species is likely to increase to about 11 Insects 800,000
30,000, if more areas are explored.
ƒ Pteridophytes-The Pteridophytes are vascular 12 Sponges 5,000
land plants and together with Gymnosperms
and Angiosperms dominate terrestrial 13 Crustaceans 9,000
communities of the world. There are about
15,000 species of Pteridophytes known to us. 14 Molluscs 38,000
ƒ Gymnosperms-Most species of Gymnosperms
are trees, although a few species are shrubs. 15 Starfishes 50,000
There are about 750 species of this group.
ƒ Angiosperms-The angiosperms or flowering 16 Fishes 6,100
plants constitute the most diverse group of
vascular plants. There are about 300,000 17 Amphibians 2,400
species described so far. This group is very
recent (35 million years) to evolve in the
18 Reptiles 6,300
geological history but have become the most
dominant of all plants because of their great
evolutionary capabilities. Plant families vary 19 Birds 9,035
considerably in the number of their constituent
species, e.g. the Orchidaceae contains 25,000- 20 Mammals 4,230
35,000 species and the Leguminosae about
15,000 species. A total of 31 families comprise
62% of the known angiosperm species; 36
families like Adoxaceae are monospecific. (iii) The mega diversity countries of the world:

Warm and humid regions in between the Tropics of

Cancer and Capricorn are provided with a rich and
diverse plant, animal and microbial life. In this wide belt

INSTITUTE FOR ADVANCED STUDIES, JODHPUR  93
UNIT 10: ECOLOGICAL PRINCIPLES 

around the globe occur more than half of the total some disadvantage to the organism to survive in the
number of species present on our planet. Countries which existing environmental condition are eliminated while
happen to lie in this zone are referred to as Megadiversity those changes which are advantageous are retained.
There should always be a sustained effort to conserve the
countries, as they possess a wide variety of plants and
endemic species and their habitat.
animal species. These countries are Brazil, Columbia,
Mexico, Indonesia, Peru, Malaysia, Ecuador, India, Areas where high levels of species richness, threat and
China, Zaire, Madagascar -and Australia, Table 2 endemism coincide are termed as hotspots. Myers (1988)
provides the approximate number of flowering plant first identified' ten tropical forest 'hotspots' characterized
species recorded in the mega diversity countries on the both by high level of plant endemism and by serious
Earth. level of habitat loss. In 1990 Myers added further eight
'hotspots', including four Mediterranean-type
ecosystems. In 1996 Conservation International (CI)
Table 2: The mega diversity countries and the number of
decided to undertake a reassessment of the hotspots
flowering plants species concept, including an examination of whether some key
areas had been overlooked. Three years later (1999) an.
No. of flowering extensive global review was undertaken for the
S. No. Country
plant species designation of biodiversity hotspots and 25 biodiversity
hotspots were identified (Myers et al. 2000).
1 Brazil 55,000
To be identified as a hotspot, a region must meet two
2 Columbia 45,000 strict criteria:
(i) it must contain at least 1,500 species of vascular
plants (>0.5 % of the world's total) as endemics, and
3 China 30,000 (ii) it has to have lost at least 70% of its original habitat.

4 Mexico
5 Australia
6 Indonesia
7 Peru
8 India
9 Malaysia
10 Ecuador
-25,000 These areas (25 hotspots) contain 44% of the world's
plants as endemics and 35% of the terrestrial land
22,500 vertebrates in an area of 1.4% of Earth's land surface.
Recently, 34 biodiversity hotspots .have been identified
20,000 covering only 2.3% of the Earth's land surface. Over
50% of the world's plant species and 42% of all
terrestrial vertebrate species are endemic to these 34
20,000
designated hotspots. The 34 hotspots, distributed in
different regions of the world are listed below:
17,500
North and Central America
15,000 1. California Floristic Province, 2. Caribbean Islands, 3.
Madrean Pine-Oak Woodlands,
15,000 4. Mesoamerica.

South America
11 Zaire 10,000
5. Atlantic Forest, 6. Cerrado, 7. Chilean Winter
Rainfall-Valdivian Forests,
12 Madagascar 10,000 8. Tumbes-Choco-Magdalena, 9. Tropica Andes

Europe and Central Asia

(iv) Endemism and 'Hot-spots' of biological diversity
Endemic species can be defined as those species which
are confined only to a particular locality. There are two
important categories of endemic species-
Palaeoendemics and Neoendemics. Palaeoendemics are
phylogenetic ally distinct taxa which arose in Tertiary
and bound to islands or environmentally isolated
situations. Neoendemics are those which have arisen
relatively recently. Endemism represents a unique step in
the process of evolution which could be perpetuated and
sustained only in the locality concerned depending on the
environmental quality. It is the environment which is
instrumental in the operation of the process of natural
selection. This makes the habitats in which endemic
species thrive'very important. The importance of the
habitat is further highlighted by the fact that in most of
the cases such localities possess a number of endemic
species distributed in several taxonomic categories. In
the process of natural selection, changes which confer
10. Caucasus, 11. Irano-Anatolian, 12. Mediterranean
Basin, 13. Mountains of Central Asia.
Africa
14. Cape Floristic Region, 15. Coastal Forests of Eastern
Africa, 16. Eastern Afromontane,
17. Guinean Forests of West Africa, 18. Horn of Africa,
19. Madagascar and the Indian Ocean Islands, 20.
Maputaland-Pondoland-Albany, 21. Succulent Karoo.
Asia-Pacific
22. East Melanesian Islands, 23. The Himalaya, 24. Indo-
Burma, 25. Japan,
26. Mountains of Southwest China, 27. New Caledonia,
28. New Zealand, 29. Philippines,
30. Polynesia-Micronesia, 31. Southwest Australia, 32.
Sundaland, 33. Wallacea,
34. Western Ghats and Sri Linka. .

94 INSTITUTE FOR ADVANCED STUDIES, JODHPUR 
UNIT 10: ECOLOGICAL PRINCIPLES 
Biodiversity of India
India is very rich in all aspects of biodiversity and is one
of the most significant biodiversity centres of the world.
Varying physiographic and climatic conditions at
different parts of India are the reasons for this high level
of biodiversity. India, with a geographical area of 329
million hectare has almost all kinds of ecological zones
found in the world. India is the meeting ground for three
major global biogeographic region: (i) Indo-Malayan, (ii)
Eurasian and (iii) Afro-tropical. It has two of the 34
recognized biodiversity 'hotspots' in the world -The
Himalaya and the Western Ghats.
48,000 species of plants which represent 11 % of the
total world flora and 80,000 species of animals
representing 6.4% of the world's fauna are found in
India. India's floral diversity includes 17,500 species of
angiospenns, over 64 species of Gymnospenns, 1,022
species of Pteridophytes, 2,843. of Bryophytes, 1,600
species of lichens, 23,000 species of fungi and 2,500
species of Algae (Table 3). Similarly, the faunal diversity
of India includes 5,000 species of molluscs, 60,000
species of insects, 1,693 species of fishes, 205 species of
amphibian, 420 species of reptiles, 1,200 species of birds
and 372 species of mammals (Table 4). According to an
estimate 35% of the flowering plants and 18% of the
total flora are endemic to India. The endemism among
the animals is high and it may be as high as 62% in case
of Amphibia. Many of the biologically rich area in India
are either unexplored or under-explored. The present
knowledge regarding the diversity of microorganisms,
wetland flora and fauna and marine life is scanty.
Table 3: Floristic diversity of India
S. No. Number of Species
1 Angiosperms 17,500
2 Gymnosperms 64
3 Pteridophytes 1,022
4 Bryophytes 2,843
5 Lichens 1,600
6 Fungi 23,000
7. Algae 2,500
It will be quite relevant to mention here that the Indian
sub-continent is one of the 12 centres of origin and
diversity of crop plants. It has given to the world nearly
167 species of crop plants. The following crops arose in
India and spread throughout the world: rice, sugarcane,
Asiatic vignas, jute, mango, citrus, banana, several
millets, spices, medicinals, aromatics and ornamentals.
The rich germplasm reserve of Indian sub-continent
includes 51 species of cereals and millets, 104 species of
fruits, 27 species of spices and condiments, 55 species of
vegetables and pulses, 24 species of fibre crops, 12
species of oilseeds and various strains of wild tea,
INSTITUTE FOR ADVANCED STUDIES, JODHPUR  95
tobacco and sugarcane. Besides above, 1,000 species of
plants are of ethno-botanical interest and 2,000 species
are used in various traditional system of medicine. India's
indigenous livestock diversity includes about 27 breeds
of cattles, 40 breeds of sheep, 22 of goats, 8 of camels, 6
of horses, 2 of donkeys and 18 types of poultry. Eight
indigenous breeds of buffalo in India represent the entire
range of genetic diversity on the earth. These prized
plants and animals together with their ancestral wild
forms are used by plant and animal breeders for the
development of improved varieties.
Table 4: Faunistic diversity of India
S. No. Group Number of Species
1 Mammals 372
2 Birds 1,200
3 Reptiles 420
4 Amphibians 205
5 Fish 1,693
6 Molluscs 5,000
7 Insects 60,000
Measures of biodiversity
Biodiversity can be measured in different ways. Two
main factors taken into account when measuring
diversity are richness and evenness. Species richness is
the number of different species present in an area.
However, diversity depends not only on richness, but
also on evenness. Evenness compares the similarity of
the population size of each of the species present.
Evenness is a measure of the 'relative abundance of the
different species making up the richness of an area.
Let us consider, two communities. A and B of species 1
and 2, both with 100 individuals:
No, of Individuals No. of Individuals
of Species 1 of Species 2
Community A 99 1
Community B 50 50
The species richness of community B would equal that of
community A. However, community B has more
evenness than A. Community B must be considered more
diverse: one is more likely to get both species there than
in community A, A community dominated by one or two
species is considered to be less diverse than one in which
several different species have a similar abundance. As
species richness and evenness increase, the diversity
increases.
UNIT 10: ECOLOGICAL PRINCIPLES 

Diversity indices Putting the values into the formula for Simpson's index

There are a great many types of diversity indices but we

can divide them into two broad categories: dominance
D=
∑ n(n − 1)
indices and information-statistic indices. N ( N − 1)
Dominance indices: Dominance indices are weighted
toward the abundance of the commonest species. A 64
widely used dominance index is Simpson's diversity D= = 0.3 (Simpson Index)
index. It takes into account both richness and evenness. 15(15 − 1)

Simpson's diversity indices: The term "Simpson's
diversity index" can actually refer to anyone of 3 closely
related indices.
Simpson's index (D): Simpson's index measures the
probability that any two individuals drawn at random
from an infinitely large community will belong to same
species. There are two versions of the formula for
calculating D. Either is acceptable but is to be consistent:
D = ∑ (n / N ) 2
Then, Simpson's index of diversity 1 - 0 = 0.7 and
Simpson's reciprocal index l/D = 3.3.
All these three values represent the same biodiversity. It
is, therefore, important to ascertain which index has
actually been used in any comparative studies of
biodiversity. The disadvantage of Simpson's index is that
it is heavily weighed toward the most abundant species,
as are in all dominance indices. The addition of rare
species with one individual will fail to change the index.
As a result, Simpson's index is of limited value in

D=
∑ n(n − 1) conservation biology if an area has many rare species
with just one individual.
N ( N − 1)
Information-statistic indices:
where, n = the total number of individuals of each Information-statistic indices can take into account rare
species, N = the total number of organisms of all species. species in a community. Information-statistic indices are
based on the rationale that diversity in a natural system
The value of D ranges between 0 and 1. can be measured in a way that is similar to the way
information contained in a code or message is measured.
With this index, a represents infinite diversity and 1, no By analogy, if we know how to calculate the uncertainty
diversity. That is, the bigger the value of D, the lower the of the next letter in a coded message, then we can use the
diversity. This does not sound logical, so to get over this same technique to calculate the uncertainty of the next
problem, D is often subtracted from 1 or the reciprocal of species to be found in a community.
the index is taken.
Shannon index: A widely used diversity index is
Simpson's index of diversity (l-D): This index Shannon index. The index is given by
represents the probability that two individuals randomly s
selected from a community will belong to different
species. The value of this index also ranges between 0
H s = ∑ pi ln pi
i =1
and 1, but here, the greater the, value, the greater the
diversity. where, pi is the proportion of individuals found in the ith
species and ln denotes natural logarithm. The following
Simpson's reciprocal index (1/D) : The value of this table gives an example:
index starts with 1 as the lowest possible figure. This
figure would represent a community containing only one Species Abundance Pi Pi ln Pi
species. The higher the value, the greater would be the A 50 0.5 - 0.347
diversity. The maximum value is the number of species B 30 0.3 - 0.361
in the sample. For example, if there are, five species in C 10 0.1 - 0.230
the sample, then maximum value is 5. D 9 0.09 - 0.217
E 1 0.01 - 0.046
The name Simpson's diversity index is often very loosely Total 5 100 1.0 - 1.201
applied and all three related indices described above
(Simpson's index, Simpson's index of diversity and Even the rare species with one individual (species E)
Simpson's reciprocal index) have been quoted under contributes some value to the Shannon index, so if an
term, depending on authors. area has many rare species, their contributions would
accommodate. Shannon index has a minus sign in the
As an example, let us consider the following table: calculation, so the index actually becomes 1.201, not -
1.201. Values of Shannon index for real communities are
Species Number(n) n(n - 1)
A 2 2 often found to fall between 1.5 and 3.5. The value
B 8 56 obtained from a sample is in itself of no significance.
C 1 0 The index becomes useful only while comparing two or
D 1 0 more sites.
E 3 6
Total (N) 15 64

96 INSTITUTE FOR ADVANCED STUDIES, JODHPUR 
UNIT 10: ECOLOGICAL PRINCIPLES 

Brillouin index: A second information-statistic index, Biodiversity values

designed to reflect species abundance, is the Brillouin
index and is given by The convention on biological diversity, signed at Rio de
ln( N !) − ∑ ln(ni!) Janeiro by 157 nations, emphasized the value of
HB = biodiversity in the environmental, genetic, social,
N economic scientific, cultural, recreational and aesthetic
contexts. We discuss below some of the definitions and
where, N is the total number of individuals in the brief explanation of values of biodiversity.
community, ni is the number of individuals in the ith
species. The following table gives an example: (a) Total Environmental Value (TEn V): UNEP (1995)
defined this as a function of primary value and total
Species No. of individuals In(n !)
economic value. Scientists and economists working
together arrived at a surrogate evaluation of all
A 5 4.79 environmental goods and services. It amounts to $33
trillion worldwide per year and thus is larger than the
B 5 4.79 global economy of $29 trillion (1997 figures). In other
word~, global natural resources are more valuable than
C 5 4.79 global national products.

D 5 4.79 (b) Primary Value (PV): This is defined as the value of

the system characteristics upon which all ecosystem
E 5 4.79 functions depend (UNEP, 1995). It is called primary
value because the structured ecosystem produces
N = 25 ∑ In(n!) = 23.95 functions that have secondary value. The secondary
value will exist as long as the ecosystem retains its
health, existence, homeostasis, operation and
maintenance.
Putting the values into the formula for Brillouin index,
we get,
(c) Total Economic Value (TEV): Also called Total
ln(25!) − 23.95 58 − 23.95 Value (TV) or simply Value (V). TEV denotes the sum
HB = = = 1.362 total of all kinds of values attached to biodiversity minus
25 25
the primary value. It is the function of use and non-use
values. Total economic value by itself will underestimate
This index describes a known population. There is no
the true value of ecosystems. It has to be considered
room for uncertainty while using ihis index. It places
along with the primary value.
more emphasis on species richness and is moderately
sensitive to sample size.
(d) Use Value (UV): This represents the value arising
from an actual use made of a given component of
The species area relationship and the theory of island biodiversity. It is often a function of Direct and Indirect
biogeography
Use values.
There is a good correspondence between the number of
(e) Direct Use Value (DUV): This is also known as
species and the extent of area they are found in.
direct value. It is defined as actual uses especially in
Generally, the larger the area, more the number of
consumption. It represents the economic values derived
species found in that area. The species-area relationship
from direct use or interaction with a biological resource
has been shown to follow the Arrhenius equation. This
or resource system. DUV is relatively easily measured by
equation can be written as :
assigning market prices.
Log S = C + Z log A,
where S = number of species, A = area of study, and C, Z
(I) Indirect Use Value (IUV) : It is defined as benefits
are constants.
arising from an ecosystem function. It represents the
economic value derived from the role of resources and
MacArthur and Wilson (1967) took this idea a step
system in supporting or protecting activities whose
forward in their Theory of 'Island Biogeography'. The
outputs have direct value in production on consumption
main points of this theory is that the number of species
(UNEP, 1995). Indirect contributions of biodiversity to
on an island is determined by a balance between
human welfare are said to have this value.
immigration and extinction and that this balance is
Biogeochemical cycles, photosynthesis, climate
dynamic. Some of the predictions made by the theory
regulation, prevention of soil erosion, pollutant
are:
degradation are the phenomena which contribute
(1) The number of species on an island should eventually
indirectly to biodiversity.
become constant through time.
(2) Large islands should support more species than small
(g) Ethical and Aesthetic Values: People with good
islands. This theory in fact is exploited to explain the
cultural background have deep concern for biodiversity.
species-area relationship in other areas as well.
They derive ethical benefit from biodiversity.
Biodiversity is considered to have great value on cultural
and religious grounds specially in India and East Asian
countries. Ethical values differ from place to place,
culture to culture, time to time and differ between

INSTITUTE FOR ADVANCED STUDIES, JODHPUR  97
UNIT 10: ECOLOGICAL PRINCIPLES 
different components of biodiversity. For example, the
ethical value attached to sacred basil is not accorded to
cactus in India. Most people value certain species more
than others subconsciously.
The aesthetic value of biodiversity is very well known.
Most people react more aesthetically towards plants that
are appealing, visually or otherwise. Most cultured
societies have attached great value to the effect that plant
and animal beauty have on human mind and emotions.
Poets, writers and artists from various cultures have
given expression to the aesthetic appeal of plants and
animals. Roses, for instance, kindle the aesthetic sense
much more than cacti and carnivorous plants, although
the latter have their own admirers. Such relative aesthetic
judgments could compel greater concern for certain
biodiversity elements than for others.
Molecular assessing of plant diversity
One of the reasons for understanding inter-and
intraspecific genetic variation is that global gene pool
represents all the information on the planet's biological
processes. Genetic markers are observable traits and put
into five groups: Morphological, cytological, chemical,
protein and DNA. The 'great importance of genetic
variation and the availability of powerful marker systems
have initiated widespread application of marker
technologies to issues related to biodiversity. Some of
the important marker systems for characterizing
biological diversity are:
(i) Study of allozymes,
(ii) Restriction fragment length polymorphism (RFLP)-it
uses restriction enzymes to detect variation in primary
DNA structure, followed by southern blotting and a
suitable method to reveal variation in plant genome,
(iii) Randomly amplified polymorphic DNA (RAPD)
analysis,
(iv) Polymerase chain reaction-restriction fragment
length polymorphism (PCR-RFLP). Combining
sequences and PCR-RFLP analyses is effective for initial
polymorphism identification and subsequent screening,
(v) Amplified fragment length polymorphism (AFLP),
(vi) Study of simple sequence repeats (SSR).
Molecular marker data must be placed within the context
of the organisms biology. One should understand the
basis of the variation and then only the value of
molecular markers to identify diversity will be properly
understood.
Uses and importance of biodiversity
Each species in biosphere has its own role and
importance towards the benefit of population. It is a
combined effort of different kinds of living organisms
which enables the biosphere to sustain the human life.
Biodiversity is vital to biosphere's health, stability and
proper functioning.
(i) Biodiversity provides the natural resource: Plants
and animals have been exploited by man since time
immemorial for food, clothing, shelter and a number of
useful products. Man has domesticated a number of such
economically important plants and animal species. At the
sametime a large number of useful products also come
from
98 INSTITUTE FOR ADVANCED STUDIES, JODHPUR
wild plants and animals. The most important contribution
of plants is to provide food to ever increasing human
population. Of the 300,000 flowering plant species,
75,000 are edible but to date only about 3,000 are
regarded as source of food. Of these, around 200 plant
species have been domesticated with 15-20 constituting
crops of major economic value. Plants have several uses
but only the most important ones are mentioned here.
Many species of plants are used as fodder. They are
either used directly from the wild, as in pastures and
rangelands, or domesticated. Grasses and legumes are the
most important . fodder sources.
Wood, the source of timber, is one of the most utilized
commercial plant products throughout the world. Plants,
which provide wood, are predominantly harvested from
the wild. At the same time monoculture plantations under
agro- and social-forestry programmes are increasingly
being raised as a source of timber. Wild sources of
timber, especially from hardwoods, are predominantly
tropical.
Plants are very important in health care. In.developing
countries, many people still rely on traditional medicines
obtained from local plants. Some 200 chemicals
extracted in pure form from about 90 plant species are
used in medicine throughout the world. About half of the
world's medicinal compounds are still derived from plant
sources. Many of these chemicals cannot be synthesized.
Therefore, medicinal plants are of great significance to
both developed and developing countries.
A number of species like cotton, linen, jute, sisal, hemp,
and coconut yield fibres of great value for cloth and other
industrial purposes. Plants are very efficient sources of
renewable energy. Plants offer a good source of fuel,
either as wood or its transformed product, charcoal. Plant
biomass from any source can also be converted into fuel.
Natural rubber, latex, gums, resins, dyes, essential oils
and beverages are some of the other products of
commercial value obtained from plants.
Microbes are of immense importance in the
pharmaceutical industry. Their capacity to ferment
various substrates has led to the production of a number
of clinically and otherwise important antibiotics.
Microbes are also good sources of various medicinally
important enzymes (streptokinase and asparginase),
toxins (botulin), immunomodulators (Cyclosporin A) etc.
Vaccines such as BCG, typhoid, he:Jatitis B, and
alkaloids such as ergot are also derived from some
microbes. Single cell protein, microbic ides, pesticides,
insecticides, flavouring agents, alcohol, acetone, butanol,
glycerol and certain organic acids such as citric acid,
fumaric, acetic and lactic, are also derived from the
activity of microbes.
Several microorganisms (Bacteria and Cyanobacteria)
are highly useful in the agriculture industry, either as bio-
fertilizers due to their capacity to fix atmospheric
nitrogen or in phosphate solubilization. Other important
roles of microorganisms include biomining,
bioremediation, biosorption, biogass production,
harnessing solar energy etc.
(ii) Biodiversity provides the genetic resource: Drastic
global climatic changes may cause large-scale shifts in
 
UNIT 10: ECOLOGICAL PRINCIPLES 
natural vegetation and agricultural crops. Hence there is
urgent need to protect genetic resources of food plants to
maintain crop productivity in different climatic
conditions. There are several species of useful plants in
the tropics alone that could be used as an alternative
source. Their uses could be extended from emergency
sustenance in isolated locations or disaster areas to fully
exploitable alternative sources of food.
There are many instances when useful genes in wild
species or in old traditional varieties were used to
improve the strains we cultivate today. It was from a
wild melon growing in U.P., India, that genes for
resistance to powdery mildew were obtained to be
introduced in crops of musk melon grown in California.
The kans grass, Saccharum spontaneum, from Indonesia
has provided genes for resistance to red rot disease of
sugar cane. A wild variety of rice collected from V.P.,
India, in 1963, saved about 30 million hectares of paddy
from Grassy stunt virus. Nearly 20 cultivars of rice
grown in rice growing countries of the world contain
useful genes from wild varieties of Kerala, India. Fungus
blighted leaves of com grown in plains of the VSA,
caused as much as 50% damage in some regions.
Resistant genes. from Mexican varieties imparted blight
resistance to halt the damage. Old traditional varieties
and the wild relatives of domesticated plants and animals
constitute a vital genetic resource for us. Under the
circumstances we should have a collection of gene-pool
as large as possible. It is only from this gene pool that we
can synthesize the cultivars of future.
(iii) Biodiversity maintains a stable ecosystem: In an
ecosystem all the components are related to one another
and it occurs in a state of dynamic equilibrium. This
system of checks and balances is of fundamental
importance in an ecosystem which is maintained in a
functional state by the activity of a large number of
organisms. So in a complicated ecosystem with several
trophic levels, each of which is composed of several
species, elimination of single species does not create any
problem. There are several species or alternatives which
can take over and keep the system in a functional state.
But in a simple system loss of a single or a few species
could be catastrophic because of lack of alternatives.
Thus diversity imparts stability to an ecosystem.
Each species in the complex biosphere has its own
importance. Earthworms killed by pesticides do not
aerate our soils. Mangroves, destroyed to supply
firewood, have stopped protecting our coastline from
erosion. At one time oysters were so numerous in
Chespeak Bay, USA, that they could filter the entire
water in just three days. It takes about a year to filter the
water now as oy~ter population has declined by 99%.
This has caused the water of Chespeak Bay to be
increasingly muddied and oxygen deficient. An adult
frog can consume insects equal to its own weight in a
day. Diminishing frog population has been associated
with increased rate of pest damage to crops and
recurrence of malaria in India.
(iv) Biodiversity ensures optimum utilization and
conservation of abiotic resources in an ecosystem:
Tropical regions are having the richest biodiversity in the
world, most of the nutrients are lodged in its biotic
community. Due to warm and humid conditions
INSTITUTE FOR ADVANCED STUDIES, JODHPUR  99
decomposition of organic matter and regeneration of
nutrients take place rapidly. A. wide variety of plants
which include trees, herbs, shrubs, grasses and climbers
quickly absorb all the nutrients which are made available
to them as a result of mineralization. The rich
biodiversity in the tropics is sustained largely by recycled
nutrients. In ecosystem of low biodiversity, the uptake of
nutrient is not so efficient. The minerals remain in the
soil while organic matter lies on the forest floor.
As most of the minerals are used by the biotic
community of an ecosystem the nutrient loss is
prevented. Mineral nutrients are not lost to the flowing
waters, rains and floods which are frequent in the tropics.
Luxuriant growth of vegetation hinders rapid flow of
water, binds soil particles together and prevents soil
erosion. Loss of biodiversity reduces the efficiency of
this vital machinery which in turn results in rapid losses
of nutrients and degradation of soil.
Loss of biodiversity
The loss qf biodiversity is a global crisis. There is hardly
any region on the earth that is not facing ecological
catastrophes. Of the 1.4 million species known to. inhabit
the earth, one-fourth is likely to become extinct within
the next few decades. Biological extinction has been a
natural phenomenon in geological history. But the rate of
extinction was perhaps one species every 1,000 years.
But man's intervention has hastened the extinction rates.
Between 1600 and 1950, about 30 species of higher
animals were lost and now we are probably losing one
species every year.
The destruction of the world's tropical forests, v,:hich are
disappearing at an alarming rate, is one of today's most
urgent global environmental issues. A rich species
diversity is slowly being lost for ever. Tr,opical forests
are estimated to contain 50 to 90% of the world's
biodiversity. The report, based on studies carried out by
FAO and wcs, found the tropical forest is shrinking at the
rate of 0.8% each year (during 1980 to 1990). About 154
mha of tropical forests equivalent to almost three times
the size of France have been destroyed. If the current rate
of deforestation continues scientists estimate that 5 to
10% of tropical forest species may face extinction,
within next one or two decades.
Rainforests, the home to half of the world's life forms,
continue to be destroyed at the rate of over 100,000 sq.
km every year. This loss of biodiversity has immediate
and long term effects on human survival. The majority of
the world's human population still depends on wild plant
and animals for their daily food, medicine, housing and
household material, agriculture. fodder, fuel wood and
intellectual stimulation.
The loss is even more direct in the case of domesticated
biodiversity. Traditional farmers of the world have
developed an incredible variety of crops and livestock.
This too has been eroded over the last few decades, as
thousands of traditional crop strains and hundreds of
domesticated livestock breeds being replaced by a
handful of laboratory generated hybrids or by dominant
cash crops.
UNIT 10: ECOLOGICAL PRINCIPLES 
Major causes for the loss of biodiversity
Disturbances of any type in an ecosystem tend to reduce
its biological diversity. As human population rises, an
increasing demand for raw material, food and space is
placed on natural ecosystem while enormous quantities
of wastes and spoils are introduced into the environment
collectively damage the biotic components of natural
systems partially or completely. Major causes for the loss
of biodiversity can be summarized as follows:
(i) Destruction of habitat: The natural habitat may be
destroyed by man for his settlement, agriculture, mining,
industries, highway construction, dam building etc. As a
consequence, the species must either adapt to the
changes ,in the environment, move elsewhere or may
succumb t.o predation, starvation or disease and
eventually die. Several rare butterfly species are facing
extinction due to habitat destruction in the Western
Ghats. Of the 370 butterfly species available in the
Ghats, around 70 are at the brink of extinction.
(ii) Hunting: Wild animals are hunted for the
commercial utilization of their products such as hides
and skin, tusk, fur, meat, pharmaceuticals, cosmetics,
perfumes and decoration purposes. In Africa, in recent
years 95% of the black rhino population have been
exterminated in Africa by poachers for their horn. Today,
rhino horn fetches more than $15,000 in the
pharmaceutical market. In the last one decade, over one-
third of Africa's elephants have been killed to collect
3,000 tonnes of ivory. International regulations have, to a
great extent, reduced illegal trading and poaching of
African Tuskers. In 1987, the Indian Govt also banned
the trade in Indian ivory. The scarlet macaw, once
common throughout South America, has been eliminated
from most of its range in Central America. Several
species of spotted cats such as ocelot and Jaguar have
been jeopardized , by the demand for their fur. In 1962,
nearly 70,000 whales were slaughtered. However,
international trade in whale products is banned now.
In India, rhino is hunted for its horns, tiger for bones and
skin, musk deer for musk (medicinal value), elephant for
ivory, Gharial and crocodile for skin and jackal for fur
trade in Kashmir. One of the most publicized commercial
hunts is that on whale. Convention on International
Trade in Endangered Species of Wild Fauna and Flora
(CITES) listed 9 Indian animal species which have. been.
severely depleted due to international trade. These are
Fin whale (Balenoptera physalus), Himalyan Musk
deer(Moschus moschiferus), Green Turtle (Chelonia
mydas), Hawksbill Turtle (Eretmochelya imbricata),
Olive Ridley Turtle (Dermochelys olivacea), Salt-water
Crocodile (Crocodylus porosus), Desert Monitor Lizard
(Varanus griseus), Yellow Monitor Lizard (V.
flavesoens) and Bengal Monitor Lizard (V. bengalensis).
Officials of Trade Record Analysis of Flora and Fauna in
Commerce (TRAFFIC-India) say, poaching of the Indian
tiger has risen because of increasing demand from south
east Asian countries and China, where pharmaceutical
factories consume the bones of 100 tigers each year.
Such demand has decimated the tiger population in
China and brought the Russian tiger to the brink of
extinction. As a result, in recent years much of the
demand has been met by poachers in India. One kg of
100 INSTITUTE FOR ADVANCED STUDIES, JODHPUR
tiger bones fetches $ 90 in India and $300 in the
international market. Hunting for sport is also a factor for
loss of animal biodiversity.
(iii) Exploitation of selected species: Exploitation of
medicinally important plants has resulted in their
disappearance from many of their natural habitat. The
pitcher plants, Nepenthes khasiana, Drosera sp., Gnetum
sp., Psilotum sp. Isoetes sp. are ruthlessly sought and
collected for teaching and laboratory work. They have
already become rare. Medicinal plants like Podophyllum
sp., Coptis sp., Aconitum sp., Rouvolfia sp., Saussura
lappa, Atropa acuminata, Dioscorea deltoidea etc. are
also disappearing rapidly as a consequence of merciless
over-collection. Similarly, the natural populations of a
number of economically important trees like Pterocarpus
santalum, Dysoxylon malabaricum, Santalum album
which yield valuable timber are fast dwindling. In the
category of over-exploited plants may also be placed a
number of orchids producing world's most showy
flowers. Plants like Paphiopedilum fairieyanum,
Cymbidium aloiflium, Aerides crispum etc. are in great
demand but their natural populations have almost
disappeared.
Today, only nine varieties of wheat occupy more than
half of United States wheat fields. Almost 95% of the old
strains of wheat grown in Greece before the Second
World War (1939-1945) have disappeared. They are
replaced 'by a few new hybrid varieties. Only four
varieties provide almost 72% of the entire potato harvest
of the United States. Over 2,000 varieties of apples were
under cultivation during the earlier century. Today,
three-fourth of entire apple production of France consists
of' North American varieties of which nearly 70%
happens to be the Golden variety. Indonesia has lost
nearly 1,500 strains of rice ana nearly three fourth of its
rice production comes from varieties discussed from a
single maternal stock. Practically all varieties of
Sorghum grown in South Africa have disappeared
following introduction of high yielding hybrid varieties
from Texas. In India, an estimated 50-60 thousand
varieties of rice were cultivated before independence,
most of which are being dropped in, favour of a few high
yielding varieties. All over the world traditional varieties
which together constituted a diverse mosaic, are being
dropped one by one 'being replaced by a few high
yielding strains. The reduction of generic diversity
among the cultivated species and the disappearance of
their wild relatives, drastically limit possibilities of
creating new cultivar in the future.
(iv) Habitat fragmentation: Habitat fragmentation may
be defined as an "unnatural detaching or separation of
expansive tracts of habitats into spatially segregated
fragments that are too limited to maintain their different
species for an infinite future. This phenomenon was
observed as early as 1885 when de Candolle noticed that
'the breakup of a landmass into smaller unfts would
necessarily lead to the extinction or local extennination
of one or more species and the differential preservation
of others.
Habitat fragmentation is one of the most s~rious causes
of erosion of biodiversity. Fragmentation leads to
artificially created 'terrestrial islands'. Such fragments
experience microclimatic effects markedly different from
 
UNIT 10: ECOLOGICAL PRINCIPLES 
those that existed in the large tracks of habitats before
fragmentation. Air temperature at the edges of fragments
can be significantly higher than that found in the interior;
light can penetrate deep into the edge, thereby affecting
the growth of existing species. Fragmentation promotes
the migration and colonization of alien species. Such
substantial and continuous colonization, profoundly
affect the survival of native species. The most serious
effect of fragmentation is segregation of larger
populations of a species into more than one smaller
populations. There is considerable evidence that the
number of species in a fragmented habitat will decrease
over time, although the probable rates at which it will
happen are variable. In fact, actual data on rain forests
show that forest fragments have lower species richness
and fewer populations compared with continuous
undisturbed forests.
An example of loss of biodiversity as the result of the
fragmentation is that of the Western forest of Ecuador,
which were largely undisturbed till 1960, where newly
Constructed network of roads led to rapid human
settlements and clearance of much of the forest area,
have been fragmented into small patches of one to few
square kilometers. Such a patch, about 0.8 square
kilometers in area at Rio Palenque Biological station now
contains only about 1,033 plant species many of which
are represented by a single specimen only and are
endemic to the locality. Before 1960 the intact forest had
thousands of species as found in any other tropical
regions of the world.
(v) Collection for zoo and research: Animals and plants
are collected throughout the world for zoos and
biological laboratories for study and research in science
and medicine. For example, primates such as monkeys
and chimpanzees are sacrificed for research as they have
anatomical, genetic and physiological similarities to
human beings.
(vi) Introduction of exotic species: Any species which
is not a natural inhabitant of the locality but is
deliberately or accidentally introduced into the system
may be designated as an exotic species. Native species
are subjected to competition for food and space due to
the introduction of exotic species. There are many
instances when introduction of exotic species has caused
extensive damage to natural biotic community of the
ecosystem. The introduction of Nile perch from north in
Lake Victoria, Africa's largest lake, mas driven almost
half of the 400 original fish species of the lake to near
extinction. Both Eu.calyptus and Casuarina are plants
introduced in India from Australia. The remarkably fast
growth of these plants has made them valuable source of
rough timber. However, these plants appear to be
ecologically harmful as they tend to suppress the original
species of the locality.
While economically useful plants are deliberately
introduced a large number of exotic weeds are
transferred from one locality to another accidentally. The
wheat imported to India from the USA under PL-480
scheme were contaminated with seeds of Parthenium
hysterophorus, the congress grass and Agrostemma
githago, the corn cockle. Both of these plants have
spread throughout India as a pernicious weed in wheat
fields. Parthenium was first observed growing on a
INSTITUTE FOR ADVANCED STUDIES, JODHPUR  101
rubbish heap in Pune in 1960. It is an aggressive plant
which matures rapidly and produces thousands of seeds.
The native .grasses and other herbs are crowded out of
existence. Water hyacinth, Eichomia crassipes, was
introduced in 1914 in West Bengal. The first appearance
of Alligator weed, Altemanthera philexeroides, was
reported near Calcutta air port in 1965, while Salvinia
molesta was brought in India by an aquarist. These plants
grow vigorously and result in the formation of thick mat
on the water surface. They impede run off in streams and
promote water logged conditions. A number of useful
water plants are displaced by these vigorous but useless
plants. There is an overall reduction in biodiversity
wherever these exotic weeds migrate.
(vii) Pollution: Pollution alters the natural habitat. Water
pollution especially injurious to the biotic components of
estuary and coastal ecosystems. Toxic wastes entering
the water bodies disturb the food chain and so the aquatic
ecosystems. Insecticides, pesticides, sulphur and nitrogen
oxides, acid rain, ozone depletion and global warming
too, affect adversely the plant and animal species. The
impact of coastal pollution is also very important. It is
seen that coral reefs are being threatened by pollution'
from indusralization, oil transport and offshore mining
along the coastal areas.
Noise pollution is also the cause of wildlife extinction.
This has been evidenced by the study by the Canadian
Wildlife Protection Fund. According to a study, Arctic
Whales are seen on the verge of extinction as a result of
increasing noise of ships, particularly ice-breakers and
tankers.
(viii) Control of pests and predators: Predator and pest
control measures, generally kill predators that are a
component of balanced ecosystem and may also
indiscriminately kill non-target species.
(ix) Natural calamities: Natural calamities, such as
floods, draught, forest fires, earth-quakes, volcanic
eruptions, epidemics etc, sometimes take a heavy toll of
plant and animal life. Floods are frequent in moist
tropical regions of the world which inundate much of the
ground vegetation, trap a large number of animals while
leading away soil nutrients. Failure of monsoon in
succession for two or three years dries' up ground
vegetation and as the subsurface water table recedes trees
are also affected. With plant life animals also suffer.
Forest fires in densely wooded localities often reduce to
ashes a large number of plant and animal species and so
do earthquakes. Volcanic eruptions may at times
completely destroy plant and animal life in its
surrounding areas. Epidemics sometimes destroy large
portions of a natural population. In nature such episodes
are usually confined to specific plant or animal
populations as the pathogen is often specific to particular
species or group of species.
(x) Other factors: Other ecological factors that may also
contribute to the extinction of plant and animal diversity
are as follows
(a) Distribution range-The smaller the range of
distribution, the greater the threat of extinction.
(b) Degree of specialization-The more specialized an
organism is, the more vulnerable it is to extinction.
UNIT 10: ECOLOGICAL PRINCIPLES 

(c) Position of the organism in the food chain-The Table 6: Country-wise threatened species of plants
higher the organism in food chain, the more susceptible it
becomes. S. No. Country No. of threatened species
(d) Reproductive rate-Large organisms tend to produce
fewer off springs at widely intervals. 1 USA 2,262
Listing of threatened biodiversity
2 Australia 2,024
Red Data Book (RDB) is the name given to the book
dealing with threatened plants and animals of any region. 3 Turkey 1,994
Many countries have prepared their own Red Data Book
(e.g. Britain and New Zealand). On the global level, the 4 India 1,336
lUCN published RDB in two volumes. Its opposite is the
Green Book,. which lists rare plants growing in protected 5 South Africa 1,016
areas like botanic gardens. A mimeographed Green Book
for India has been brought out for hundred rare plant 6 Spain 936
species growing in the garden of Botanical Survey. of
India (BSI). The Bsr has also compiled three volumes of 7 Mexico 883
Red Data Book having information of endangered plant
species. The UNEP has compiled endangered species of
8 Cuba 860
the world under the title "Blue Book". The IUCN has
defined the Red Data Categories that specify the state of
ex.tinction process. 9 Jordan 752

Threatened Categories: The major IUCN threatened 10 Panama 549

categories (IUCN Red List Categories, 1995) currently
recognized, together with their definitions are: 11 Malaysia 522

(i) Extinct (EX): Species not definitely located in the 12 Costa Rica 419
wild during the past fifty years but which may survive in
cultivation (e.g. Franklinia aloetamha). Some authors 13 Peru 360
suggest that 'Extinct' should denote those taxa that have
been totally lost and that the terminology 'Extinct in the
14 China 350
wild' should be used to refer to species lost in the wild,
while living under cultivation (dubbed by IUCN as EW).
15 Vietnam 338
(ii) Endangered (EN): Species in danger of extinction
(within a few decades) and whose survival is unlikely if 16 Columbia 327
the causal factors continue to operate (Areca concinna,
Euphorbia obdelkuri). In this category are included those 17 Brazil 318
taxa whose numbers have been reduced to a critical level
or whose habitats have been so drastically reduced that 18 Iran 301
they are deemed to be in immediate danger of extinction.
Also included are taxa that may now be extinct even 19 Chile 284
though seen in the wild in the past 50 years. The other
criteria are 50% decline in the last 10 years; < 5,000 km2
20 Ecuador 256
area of occupancy or < 500 km2 in fragmented areas;
2,500 individuals or subpopulation of 250 mature
individuals. The category Critically Endangered (CR)
includes species that face an extremely high risk of (iv) Rare (R) : Taxa with small populations that are not
extinction in the wild in the immediate future. These are endangered or vulnerable at present but are at risk are
characterized by 80% decline in the last 10 years, 100 included under this category (Lactuca sa ligna, Salvia
km2 occupapcy or 10 km2 in fragmented areas. saxicola). A species may be rare because of restricted
geographical range, high habitat specificity and small
(iii) Vulnerable (VU): Taxa likely to move into the local population size, or thinly scattered over a more
endangered category in near future if the causal factors extensive range, or due to a combination of two or more
continue to operate (Ranunculus ophioglossifolius). of these characteristics. Rare species have a population
Included in this category are taxa in which most or all of less than 20,000 individuals. Some species are
populations decrease in size because of overexploitation, naturally rare and have never occurred in greater
extensive destruction of habitat or other environmental numbers, yet they are able to maintain these numbers.
disturbances. Also included are taxa with populations Other species become rare through man's action or other
that are still abundant but under threat from severe natural forces.
adverse factors throughout their distribution range. The
other criteria include 50% decline in last 20 years; < (v) Indeterminate (I) : Species considered definitely to
20,000 km2 occupancy or < 2,000 km2 in fragmented be endangered, vulnerable or rare but for which
populations; 10,000 individuals or subpopulation of information is insufficient to categorically assign them to
1,000 mature individuals. any of these three categories.

102 INSTITUTE FOR ADVANCED STUDIES, JODHPUR 
UNIT 10: ECOLOGICAL PRINCIPLES 
Census of threatened species: Raven (1987) mentioned
that of the 300,000 species of vascular plants, 170,000
are tropical and subtropical; of the 130,000 tropical plant
species, some 60,000 are threatened and at risk of
extinction within the next halfcentury. Of the 80,000
temperate plant species, about 8,000 are threatened. In a
more recent estimate, a total of 26,106 species of plants
were considered threatened by the World Conservation
Monitoring Centre (WCMC, 1994). This number account
for 8% of the world's plant species. Of these, 3,632
species of plants belong to the category Endangered,
5,687 to Vulnerable, 11,485 to Rare and 5,302 to
Indeterminate (WCMC 1994). Table 6 provides data on
the total number of threatened species in the major
countries of the world.
Conservation and management of biodiversity
Conservation implies trying to keep things the way they
are. Things cannot remain unchanged everywhere,
because there is an increasing human population. This is
generating strong pressure to convert some areas of
natural vegetation to other uses, such as fannland,
forestry plantations, roads or towns. It is, therefore,
inevitable that there will be conflicts between wildlife
conservation and other demands and desires of people.
So we need to consider what should be the aims and
priorities for conservation.
(i) Aims for conservation: (a) To preserve ecosystem
processes e.g. primary productivity, water balance,
energy balance, nutrient cycling, slow soil erosion. (b)
To maintain all the genetic diversity within living things.
(c) To prevent extinction of any species. (d) To preserve
particular species.
High priority is given to preserve ecosystem processes.
For example, destruction of forest and converting it to
other vegetation can bring around climatic changes and
increase soil erosion. However, the ecosystem processes
are likely to be primarily influenced by characteristics of
the mo~t abundant plant species and much less by plants
and animals of sporadic occurrence.
Maintenance of genetic diversity is also highly desirable.
Various populations within each species and each
individual within a population may exhibit genetic
diversity. This enables the species to overcome future
environmental change (e.g. climatic change) and against
sudden catastrophes (e.g. disease epidemic). A gene may
carry a useful character that we can later insert into a
farm crop or animal, or the ability to produce a useful
chemical such as an antibiotic.
There should always be efforts to prevent any species
from becoming extinct. The preservation of any species
is an absolute, which must take priority over anything
else. Every species is unique, and once it is extinct it is
lost for ever.
(ii) Giving priority to particular species: Some species
must be given priority in conservation. We should first
concentrate on conserving species that have strong and
widespread public appeal called flagship species. As
there will be widespread support for conserving areas
~here they live, many other wild species will also be
saved. Examples of flagship species are the Florida
INSTITUTE FOR ADVANCED STUDIES, JODHPUR  103
panther, the giant panda, the African elephant and in
New Zealand the takahe (a flightless bird). A serious
limitation of using flagship species as the centre of
conservation planning is that there is no guarantee that
preserving them will preserve many other species. An
example of this is the northern spotted owl in the forests
of us Pacific Northwest. The plan for preserving forest
patches was specifically geared to the needs of the owl
and was clearly not suited to other species, for example
some. fish and amphibians.
This has led to the suggestion of giving priority to the
preservation of umbrella species, which are chosen
because in order to preserve them we shall also preserve
suitable habitat for many other species. The difficulty is
that we know too little about the ecology of most of the
species in any community to have a sound basis for
choosing umbrella species.
A third suggestion is to identify and conserve a keystone
species which remains at the top of an arch. If it is
removed the whole arc~ will fall down. The keystone
species is defined as a species whose impacts on its
community or ecosystems are large and much larger than
would be expected from its abundance (Power and Mills
1995). If a keystone species disappears many other
species will be affected and will perhaps disappear too.
The converse is redundant species, whose disappearance
has little or no effect on the remaining species. It has
been suggested that keystone species should have high
priority in conservation and redundant species low
priority. This distinguishes between keystones and
dominants. The elephant is an example of a species that
is sometimes endangered and which can have a major
effect on savanna vegetation. By destroying trees
elephants can influence the balance between trees and
grassland. Elephant could certainly be classed as a
keystone species, and a flagship species too.
Another approach to setting priorities for conservation is
to decide which species are in danger of becoming
extinct soon and to concentrate our conservation
activities on these. Lists of species at risk have been
drawn up, notably the Red Lists and Red Data Books of
IUCN (now called the World Conservation Union).
(iii) In situ and ex situ conservation: In situ
conservation is conservation of biodiversity in its natural
habitat. The strategy of in situ conservation involves the
establishment of protecled areas (an area of land and/or
sea specially dedicated to the protection and maintenance
of biodiversity). There are many types of protected areas
with different degrees of protection, permanency and
purpose. The range of protected areas can be classified
broadly into the following: National Park, Sanctuary,
Protected landscapes, and seascapes, biosphere reserves,
World Heritage Sites etc. Today, there are approximately
9,832 protected areas (lucN-1995), including 1,508
national parks of 9.25 million km2 or about 8.2% of the
Earth's land surface. A further 40,000 smaller protected
areas cover another 5% of the land area (UNEP 1995).
The recommendation by IUCN is preservation of a cross-
section of all major ecosystems to the extent of 13
million km2 or about 10-12% of the Earth's surface. In
India, 4.2% of the total land area earmarked for in situ
conservation includes 448 wildlife sanctuaries, 85
national parks, 5 world heritage sites, 10 biosphere
UNIT 10: ECOLOGICAL PRINCIPLES 
reserves, 6 Ramsar sites, 15 mangrove areas, 23 tiger
reserves and 17 wetland areas.
Ex-situ conservation is the maintenance of species away
from their natural habitat. It is considered a means of
static conservation (opposite to dynamic in situ
conservation). Ex-situ conservation can be applied to
both wild and domesticated species. A botanic garden is
the most important form of ex-situ conservation. There
are more than 1,500 botanic gardens in the world (UNEP
1995) of which 582 have a seed bank facility. In India,
there are 33 botanic gardens. Zoos of the world
contributed significantly to ex situ conservation of
endangered wild life. Many species maintained in zoos
have already become rare and some even extinct in the
wild.
A seed bank is one of the most efficient methods of ex
situ conservation for angiosperms. Dried seeds (5-8%
moisture content) can be stored for 5-25 years at 00C to
50 C or for up to 100 years if stored at -100 C to -200 C.
The total number PGRFA (Plant Genetic Resources for
Food and Agriculture) stored in seed banks worldwide is
3,610,428 (FAO 1996). In vitro methods are applied
throughout the world for conservation of plant
germplasm. Around 1,500 wild taxa have been stored in
vitro in various institutions of the world.
SPECIES RICHNESS CURVES
Species-individual curve: a plot of the cumulative
number of species encountered vs. the cumulative
number of individuals captured.
Species-area curve: a plot of the (cumulative) number of
species encountered as a function of area. Species-area
curves can be used to compare different regions
PATTERNS OF SPECIES RICHNESS
Species richness is not constant through space.
ƒ Species Richness (for plants and animals) is
o negatively related to:latitude and altitude
104 INSTITUTE FOR ADVANCED STUDIES, JODHPUR
o positively related to: area and
environmental variability
o and have a complex relationship with:
time since disturbance, nutrients,
predation rate, productivity
COMPETITIVE EXCLUSION PRINCIPLE IN
CONTEXT TO BIODIVERSITY
Gause's principle, Gause's law, competitive exclusion
principle, or CEP (modified from glossary): "No two
species can coexist indefinitely on the same limiting
resource". The CEP has been experimentally verified in
simple systems, as well as in mathematical models.
However, we often observe not just two, but hundreds of
species coexisting in a community. One of the major
goals of community ecology is to determine the nature of
coexistence in the face the CEP. This is the paradox of
the plankton.
Competitive exclusion: the situation in which at least one
species is driven to local extinction (that is, excluded
from a community) by interspecific competition.
The Reformulation of the Competitive Exclusion
Principle, with its conditions
The Principle: Given a suite of species, interspecific
competition will result in the exclusion of all but one
species.
Conditions:
1. time has been sufficient to allow exclusion
2. the environment is temporally constant
3. the environment has no spatial variation
4. growth is limited by one resource
5. rarer species are not disproportionately favored in
terms of survivorship, reproduction, or growth
6. species have the opportunity to compete
7. there is no immigration
Corollary: The greater the degree to which these
conditions are broken, the greater the number of species
which can coexist.
Examples of Species Richness Hypotheses
Environmental variability hypothesis: higher SR is
expected where there is more environmental variability
(violates condition #3)
Niche specialization hypothesis: higher SR is expected if
species are specialists (violates #4)
Mass effect: Higher SR is expected if a region is
surrounded by different habitats, because of constant
dispersal from these other habitats (violates #7)
INTERMEDIATE DISTURBANCE HYPOTHESIS
High species richness is expected in regions with
intermediate levels of disturbance (violates #1, #2, #3,
and sometimes #5)
 
UNIT 10: ECOLOGICAL PRINCIPLES 

REASONS FOR HIGH DIVERSITY AT

INTERMEDIATE DISTURBANCE:

1. More heterogeneity within patch at

intermediate disturbance
2. On a landscape scale, many kinds of patches at
intermediate disturbance, therefore many
species
3. Overlap of early successional species and late
successional species
4. Disturbance prevents competitive exclusion
and thus allows high niche overlap at
intermediate disturbance. However, few
species can cope with high disturbance.
 
THE CONVENTION ON BIOLOGICAL
DIVERSITY (CBD)

The Convention on Biological Diversity (CBD) was

signed by over 150 governments at the 1992 Earth
Summit in Rio de Janeiro, and became effective as
international law in December 1993. It is the first
international agreement committing governments to
comprehensive protection of the Earth's biological
resources. As of February, 2004 188 countries and the
European Union had ratified the agreement; the U.S. was
one of the last nations to sign.

The CBD has three overall goals:

ƒ the conservation of biological diversity,
ƒ the sustainable use of its components, and
ƒ the fair and equitable distribution of benefits
derived from "genetic resources".
By signing the CBD, participating governments agree to
carry out various measures to conserve biodiversity.

The measures include (among other things):

ƒ to create national plans for protection of
biodiversity.
ƒ to identify ecosystems, species and genomes
important for conservation and sustainable use
of biodiversity
ƒ to monitor biological diversity and any factors
that might have impacts on it.
ƒ to establish a system of protected areas
ƒ to manage biological resources to ensure
conservation and sustainable use
ƒ to rehabilitate and restore ecosystems
ƒ to take measures for ex situ conservation.

The CBD has also published a useful Global Biodiversity

Outlook.
 

INSTITUTE FOR ADVANCED STUDIES, JODHPUR  105
UNIT 10: ECOLOGICAL PRINCIPLES 

  
J1. Principle and Need for Conservation of Natural resources:
 
Natural  resources  are  naturally  occurring  fish  (iv)  land‐use  (v)  water;  (vi)  wildlife  and  (vii) 
substances  that  are  considered  valuable  in  their  energy resources. 
relatively unmodified (natural) form. A commodity is   
generally  considered  a  natural  resource  when  the  Need for Conservation: 
primary  activities  associated  with  it  are  extraction   
and  purification,  as  opposed  to  creation.  Thus,  Expanding  human  population  resulted  into 
mining,  petroleum  extraction,  fishing,  and  forestry  expanding needs of man. With scientific progress and 
are generally considered natural‐resource industries,  technological  development  man  started  utilising 
while agriculture is not. The term was introduced to a  natural  resources  at  much  larger  scale.  Continuous 
broad audience by E.F. Schumacher in his 1970s book  increase  of  population  caused  an  increasing  demand 
Small Is Beautiful.  for resources. This created a  situation when the non 
  renewable resources may come to an end after some 
Natural resources are often classified in several ways  time. In order to have maximum production, we have 
  started  even  taking  loans  from  the  resources  meant 
1.  Based  on  continual  utility:  Some,  resources  are  for future. That  can  not be paid  back. As a result we 
exhausted  soon,  whereas  others  last  a  long  period.  would  be  using  all  those  resources  which  are  infact 
Thus  depending  upon  the  availability  of  resources,  the  property    of  future  generation.  It  is  a  matter  of 
during  our  continuous  use,  a  resource  may  be  much  concern.  There  must  be  some  sort  of  balance 
renewable  (inexhaustible),  non‐renewable  between the population growth and the utilisation of 
(exhaustible) or cyclic. Resources which can renewed  natural resources. 
along with their exploitation are always available for   
use.  Hence  they  are  called  renewable  resources.  For  The non‐availability of resources and their price‐rise 
instance,  forests  are  renewable.  However,  formation  are  having  an  adverse  effect  on  the  economies  of 
of some resources like iron ore, coal, mineral oil‐etc.  countries  world  over.  During  1980s  the  world  has 
has taken several thousand years. Once they are used  experienced a state of imbalance between the growth 
in  unlimited  way  they  can  not  be  easily  replaced.  rates  of  population  and  economic  development.  The 
Therefore, if exploited at large scale they will deplete  prices of resources as petroleum, after 1973 showed 
fast. Hence such resources are called  non‐renewable  abrupt  hike.  Hence  the  growth  rates  of  food 
resources or exhaustible. For certain resources these  production  and  economic  development  suffered 
is no final use as they can be used continuously. For  setbacks. 
example,  water  used  in  industry  and  domestic  way   
lean  be  cleaned  and  used  again  for  similar  or  other  Conservation  is  also  concerned  with  complete 
purposes. Such resources that can be used again and  elimination  of  some  unique  species  for  which  there 
again are called cyclic resources.  may  by  no  alternative  at  all.  One  can  not  imagine  a 
  situation if  Penicillium had been eliminated from the 
2.  Based  on  origin:  On  the  basis  of  their  origin  nature before mankind made use of it as an antibiotic, 
resources  maybe  biotic  (organic)  or  abiotic  (in  or  if  Cinchona  become  extinct  before  quinine  was 
organic).  Biotic  resources  are  obtained  from  the  discovered as a cure for malaria. It is therefore in our 
biosphere.  Forest  and  forest  products,  crops,  birds,  own  interest  to  conserve  our  plant,  animal  and 
animal, fish, and other marine life forms are examples  microbial  wealth.  There  is  global  realization  about 
of  biotic  resources.  Coal  and  mineral  oil  also  belong  the urgent need to conserve the biological diversity. 
to  this  category  since  they  originate  from  organic   
matter.  Some  biotic  resources  like  forest  and  Conservation  has  been  defined  as  "the  management 
livestock  are  renewable,  whereas  coal  and  oil  are  for  the  benefit  of  all  life  including  humankind  of  the 
non‐renewable.  Resources  composed  of  non‐living  biosphere  so  that  it  may  yield  sustainable  benefit  to 
inorganic  matter  are  called  abiotic  resources.  Land,  the  present  generation  while  maintaining  its 
water  and  minerals  like  iron,  copper,  lead  and  gold  potential  to  meet  the  needs  and  aspirations  of  the 
are  abiotic  resources.  Some  abiotic  resources  are  future generations".  
found as nodules in oceans as nodules of copper and   
manganese.   Living  resources  conservation  has  three  specific 
  objectives:  
3. Based on utility: Every resource has some utility.  (i) to  maintain  essential  ecological  processes  and 
For  example,  some  are  used  as  food,  some  as  raw  life support system;  
materials and others as sources of energy.   (ii) to preserve biologicaJ diversity; and  
  (iii) to  ensure  that  any  utilisation  of  species  and 
Though  serious  concern  has  already  been  felt  in  all  ecosystems is sustainable.  
nations of the world for a need to conserve their own   
natural  resources  we  would  examine  this  subject  Conservation  therefore  makes  important 
with  reference  to  our  country.  The  various  types  of  contributions to social and economic development. 
natural  resources  may  be  considered  as  follows.  (i)   
soil,  climate,  flora  and  fauna  of  India  (ii)  forests  (iii)  The  Indian  region  (80‐300  N  and  600‐97.5°E)  with  a 
total  area  of  32  million  hectares  is  very  rich  in 

106 INSTITUTE FOR ADVANCED STUDIES, JODHPUR 
UNIT 10: ECOLOGICAL PRINCIPLES 

biological  diversity.  In  fact,  India  is  one  of  the  12  Methods  of  ex­situ  conservation:  Zoos,  botanical 
centres  of  diversity  and  origin  of  several  cultivated  gardens  and  arboretums  are  the  most  conventional 
plants in the world.   methods  of  ex‐situ  conservation,  all  of  which  house 
  whole,  protected  specimens  for  breeding  and 
ƒ It  is  estimated  that  about  45,000  species  of  reintroduction  into  the  wild  when  necessary  and 
plants  occur  in  India.  The  flowering  plants  comprise  possible.  These  facilities  provide  not  only  housing 
15,000  species  of  which  several  hundred  (5000‐ and  care  for  specimens  of  endangered  species,  but 
7500) species are endemic to India.   also  have  an  educational  value.  They  inform  the 
  public of the threatened status of endangered species 
ƒ The region is also rich in fauna, containing about  and of those factors which cause the threat, with the 
65,000  species  of  animals.  Among  these  more  than  hope  of  creating  public  interest  in  stopping  and 
50,000 species of insects, 4,000 of mollusks, 6,500 of  reversing  those  factors  which  jeopardize  a  species' 
other invertebrates, 2,000 of fish, 140 of amphibians,  survival in the first place.  
420  of  reptiles,1,200  of  birds  and  340  of  mammals   
are recorded from India.   Endangered  plants  may  also  be  preserved  in  part 
  through  seed  banks  or  germplasm  banks.  The  term 
This  richness  in  biological  diversity  is  due  to  seed bank sometimes refers to a cryogenic laboratory 
immense  variety  of  climatic  and  altitudinal  facility  (at  ‐1960  C  using  liquid  Nitrogen  as  cryo‐
conditions  coupled  with  varied  ecological  habitats.  preservent  and  DMSO  as  cryo‐protectant)  in  which 
These vary from the humid tropical western Ghats to  the  seeds  of  certain  species  can  be  preserved  for  up 
the  hot  desert  of  Rajasthan,  from  the  cold  desert  of  to  a  century  or  more  without  losing  their  fertility.  It 
Ladakh  and  the  icy  mountains  of  Himalayas  to  the  can  also  be  used  to  refer  to  a  special  type  of 
warm costs of peninsular India.  arboretum where seeds are harvested and the crop is 
  rotated.  For  plants  that  cannot  be  preserved  in  seed 
We  have  so  far  given  attention  to  preserve  large  banks,  the  only  other  option  for  preserving 
mammals,  crocodiles,  birds  etc.  ignoring  plants,  germplasm  is  in‐vitro  storage,  where  cuttings  of 
particularly  forest  trees.  We  have  to  take  steps  to  plants are kept under strict conditions in glass tubes 
keep  the  species  away  from  decline.  Our  late  Prime  and vessels. 
Minister  Smt.  Indira  Gandhi  emphasized  our  ancient   
care, conservation  and  worship of trees and animals  Endangered  animal  species  are  preserved  using 
while  launching  the  World  Conservation  Strategy  in  similar  techniques.  The  genetic  information  needed 
India  on  6th  March,1980.  In  her  own  words  "The  in the future to reproduce endangered animal species 
interest  in  conservation  is  not  sentimental  one  but  can  be  preserved  in  genebanks,  which  consist  of 
the  rediscovery  of  a  truth  well  known  to  our  sages.  cryogenic  facilities  used  to  store  living  sperm,  eggs, 
The Indian tradition teaches  us  that all forms of life,  or embryos.  
human,  animal  and  plants  are  so  closely  linked  that   
disturbance  in  one  gives  rise  to  imbalance  in  the  Drawbacks  to  ex­situ  conservation:  Ex‐situ 
others."  conservation, while helpful in man's efforts to sustain 
  and  protect  our  environment,  is  rarely  enough  to 
There are two main categories of conservation: in situ  save  a  species  from  extinction.  It  is  to  be  used  as  a 
conservation and ex situ conservation.  last resort, or as a supplement to in‐situ conservation 
  because it cannot recreate the habitat as a whole: the 
1.  In  situ  conservation:  This  is  the  conservation  of  entire  genetic  variation  of  a  species,  its  symbiotic 
genetic  resources  through  their  maintenance  within  counterparts,  or  those  elements  which,  over  time, 
natural  or  even  human‐made  ecosystems  in  which  might  help  a  species  adapt  to  its  changing 
they  occur.  This  is  an  ideal  system  for  genetic  surroundings.  Instead,  ex‐situ  conservation  removes 
resources  conservation.  This  type  includes  a  system  the  species  from  its  natural  ecological  contexts, 
of  protected  areas  of  different  categories,  managed  preserving it under semi‐isolated conditions whereby 
with  different  objectives  to  bring  benefit  to  the  natural evolution and adaptation processes are either 
society. National parks, Sanctuaries, Nature Reserves,  temporarily  halted  or  altered  by  introducing  the 
Natural  Monuments,  Cultural,  Landscapes,  and  specimen  to  an  unnatural  habitat.  In  the  case  of 
Biosphere  Reserves  etc.  belong  to  this  type  of  cryogenic storage methods, the preserved specimen's 
conservation.  In  situ  conservation,  therefore  is  not  adaptation  processes  are  frozen  altogether.  The 
practicable for domesticates.  downside  to  this  is  that,  when  re‐released,  the 
  species  may  lack  the  genetic  adaptations  and 
2.  Ex­situ  conservation:  It  means  literally,  "off‐site  mutations which would allow it to thrive in its ever‐
conservation".  It  is  the  process  of  protecting  an  changing natural habitat. 
endangered species of plant or animal by removing it   
from an unsafe or threatened habitat and placing it or  Furthermore,  ex‐situ  conservation  techniques  are 
part  of  it  under  the  care  of  humans.  While  ex‐situ  often  costly,  with  cryogenic  storage  being 
conservation  is  comprised  of  some  of  the  oldest  and  economically  infeasible  in  most  cases  since  species 
best  known  conservation  methods  known  to  man,  it  stored  in  this  manner  cannot  provide  a  profit  but 
also  involves  newer,  sometimes  controversial  instead  slowly  drain  the  financial  resources  of  the 
laboratory methods.  government  or  organization  determined  to  operate 
  them.  Seed  banks  are  ineffective  for  certain  plant 
genera  with  recalcitrant  seeds  that  do  not  remain 

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UNIT 10: ECOLOGICAL PRINCIPLES 

fertile  for  long  periods  of  time.  Diseases  and  pests 

foreign  to  the  species,  to  which  the  species  has  no 
natural  defense,  may  also  cripple  crops  of  protected 
plants  in  ex‐situ  plantations  and  in  animals  living  in 
ex‐situ  breeding  grounds.  These  factors,  combined 
with  the  specific  environmental  needs  of  many 
species,  some  of  which  are  nearly  impossible  to 
recreate  by  man,  make  ex‐situ  conservation 
impossible  for  a  great  number  of  the  world's 
endangered flora and fauna. 
 
However, when the extinction of a species is eminent, 
ex‐situ  conservation  becomes  the  only  option  left  to 
humanity. It is simply better to preserve a species in 
part than to let it die out completely. 

108 INSTITUTE FOR ADVANCED STUDIES, JODHPUR 
UNIT 10: ECOLOGICAL PRINCIPLES 

J2. Environmental Impact Assessment: 
 
Environment impact can be defined as any change in   
the  environment  that  is  caused  by  an  activity  or  Impacts of different human activities on environment 
factor  which  may  be  physical,  chemical,  biological,  which is assessed by EIA: 
social,  or  economical.  Environment  Impact  1. Impact  due  to  River  Valley  Projects,  Dam 
assessment  (EIA)  consists  in  establishing  quantative  constructions, Thermal Power station 
values  for  selected  parameters,  which  indicate  the  2. Impact of Irrigation by Canals. 
quality  of  the  environment  before,  during  and  after  3. Impact  of  Power  lines,  Roads,  Mining  & 
the  action.  EIA  is  the  documentation  of  an  mineral based industries. 
environmental  analysis,  which  includes  4. Impact  of  establishment  of  large  industries 
identification,  interpretation,  prediction  and  in remote areas. 
mitigation  of  impacts  caused  by  a  proposed  action  5. Impact of forest fires & desertification. 
and project.   6. Establishment  of  Biosphere  reserves, 
  National parks, sanctuaries etc. 
To  accomplish  this  analysis,  it  is  first  necessary  to   
develop  a  complete  understanding  of  the  proposed  There are two study designs routinely used in EIA: 
action,.  What  is  to  be  done?  What  kind  of  material,  1. Time  based  design:    It  is  based  on 
manpower & or resources are involved. Second, it  is  comparison  of  the  same  sites  at  different  times.  If 
necessary  to  gain  a  complete  understanding  of  the  measurements  of  environmental  components  of 
affected  environment,  i.e,  what  is  nature  of  interest  perturbation  occurred,  these  values  may  be 
biophysical  or  socioeconomic  characteristics  that  compared  with  the  measurement  taken  at  or  more 
may  be  changed  by  the  action.  Third,  it  is  necessary  times  after  perturbation.  This  simple  “before‐after” 
to project the proposed action into the environmental  or  baseline  design  is  frequently  used  in  assessing 
characteristics;  quantifying  the  changes  whenever  plans. 
possible.  Fourth,  it  is  the  analysis  of  the  probable  2. Spatial  Designs:  Instead  of  making 
environmental  consequences  of  the  proposed  action  comparison  over  time,  sites  in  different  location  can 
may be used in the decision making process.  be compared at a single time. Samples are taken from 
  sites within perturbed area and from sites that were 
The concept of EIA has originated in order to give an  not perturbed.  
idea  of  the  possible  impacts  at  any  developmental 
plan it is going to have on our environment. 
 
J3. Sustainable Development: 
 
For  society  to  continue  developing  in  the  way  it  has  stem  solely  from  financial  growth.  Friends  and 
done  in  the  past,  we  need  to  pay  more  attention  to  families,  culture,  religion  and  ethics  are  important 
our environment. How this is best achieved is often a  elements  of  society,  that  are  not  primarily  based  on 
matter  of  opinion  rather  than  fact,  dependent  upon  exchanging goods and services, but contribute to the 
different  perspectives  of  the  environment  and  views  overall  quality  of  life.  Society,  in  turn,  exists  entirely 
of  nature.  Recently,  a  concept  has  emerged  that  has  within the environment. Our basic requirements ‐ air, 
attempted to bring together the best aspects of these  food  and  water  ‐  come  from  the  environment,  as  do 
different  viewpoints,  and  to  harmonize  the  the  energy  and  natural  resources  for  housing, 
development  of  mankind  with  the  protection  of  transportation  and  the  products  we  depend  on. 
nature.  This  is  the  concept  of  "Sustainable  Protection  of  the  environment,  therefore,  resides  at 
Development".  Sustainable  development  involves  the  core  of  Sustainable  Development.  In  the  1980s, 
maintaining  our  current  rate  of  development  whilst  increasing  concern  about  the  effects  of  economic 
leaving  suitable  resources  behind  for  later  development  on  health,  natural  resources  and  the 
generations  to  continue  to  develop.  In  this  context  environment  led  the  United  Nations  to  release  the 
then,  environmental  problems  must  be  tackled  by  Brundtland  Report.  This  defines  sustainable 
considering  their  relationship  with  the  state  of  the  development as 'development which meets the needs 
economy  and  the  wellbeing  of  society.  In  fact,  the  of  the  present  without  compromising  the  ability  of 
environment,  the  economy  and  society  taken  future generations to meet their own needs.' 
together, include everything that we need to consider   
for a healthy, prosperous and stable life.  Definition:  Sustainable  development  is  a 
  development  strategy  that  manages  all  aspects  of 
Although  sustainable  development  is  about  natural  resources  and  human  resource  as  well  as 
integrating  the  environment,  society  and  economy,  financial and physical assets for increasing long term 
the  economy,  and  in  turn  society  exist  within  the  wealth and well being. Sustainable development as a 
wider  context  of  the  environment.  The  economy  goal rejects policy and practices that support current 
exists entirely within society, because all parts of the  living  standards  by  depleting  the  productive  base, 
human  economy  require  interaction  among  people.  including the natural resource and that leaves future 
However,  society  is  much  more  than  just  the  generation with poor prospects and greater risk then 
economy.  Happiness,  pleasure  and  well  being  to  not  our own. 

INSTITUTE FOR ADVANCED STUDIES, JODHPUR  109
UNIT 10: ECOLOGICAL PRINCIPLES 

  of  neither  other  communities  nor  it  can  export  its 

Objectives:  own  waste  to  create  pollution  in  other  societies. 
  Interdependence  among  sustainable  societies  if  any 
1. Renewability:  Renewable  resources  can  be  is  always  balanced  meaning  thereby  export  and 
utilized  at  permissible  rate  that  is,  the  rate  of  import should be equal. 
utilization of such resources should be lower then or   
equal  to  rate  of  generation  of  resource.  Renewable   
resources  that  have  already  been  depleted  due  to  4. Substitution:  Sustainable  development  ensures 
overuse  should  be  given  complete  protection  substitution  of  use  of  non‐renewable  resources  by 
temporarily.  They  can  be  utilized  for  reaching  a  the  use  of  renewable  natural  resources;  however, 
quantity of self sustainable.  complete  substitution  is  extreme  case  of  sustainable 
  development.  Eg.  Inorganic  fertilizer  by  organic 
2. Adaptability:  A  sustainably  developing  society  fertilizer;  substitution  of  fossil  fuel  energy  by  wind, 
has  a  ability  to  adapt  in  changing  environment.  The  solar, tidal energy. 
society have high resistance ability can adapt quickly   
under  stress  without  causing  harm  to  existing   5. Institutional  Commitment:  This  includes 
natural  resource  and  society  is  also  capable  of  political  support  constitutional  provisions,  legal 
creating  opportunities  for  survival  and  recreation  framework, coordination between legal institutes and 
through  innovation  manifestation  in  research  and  above  all  the  ability  to  understand  the  need  of 
development activities.  sustainable  development  in  a  society.  Proper 
  education should be imparted to all in the society so 
3. Interdependence: A sustainable society neither  that  at  large,  accept  values  and  practices  of 
import  resources  from  outside  through  deprivation  sustainable development.  

J4.STATUS OF BIODIVERSITY IN INDIA

India occupies only 2.4% of the world’s land area but its
contribution to the world’s biodiversity is approximately
8% of the total number of species, which is estimated to
be 1.75 million. Of these, 126 188 have been described
in India. The species recorded includes flowering plants
(angiosperms), mammals, fish, birds, reptiles, and
amphibians, constitute 17.3% of the total whereas nearly
60% of India’s bio-wealth is contributed by fungi and
insects. Such a distribution is similar to that found in the
tropics and the subtropics. Biogeographically, India is
situated at the tri-junction of three realms namely afro-
tropical, Indo-Malayan and Paleo-Arctic realms, and
therefore, has characteristic elements from each of them.
This assemblage of three distinct realms probably is a
fact which is believed to partly account for its rich and possesses, while its record in agro-biodiversity is very
unique in biological diversity. Based on the available impressive as well. There are 166 crop species an
data, India ranks tenth in the world and fourth in Asia in numerous wild relatives of domesticated animals.
plant diversity, and ranks tenth in the number of endemic Overall India ranks seventh in terms of contribution to
species of higher vertebrates in the world. There are 10 world agriculture. 
biogeographical zones in India. They can be classified as
under: The Biogeographic classification of India (Rodgers
and Pawar 1990)
India’s contribution to agro-biodiversity has been
impressive. India stands seventh in the world as far as the • Trans-Himalayas. An extension of the
number of species contributed to agriculture and animal Tibetan plateau, harboring high-altitude cold
husbandry is concerned. In qualitative terms too, the desert in Laddakh (J&K) and Lahaul Spiti
contribution has been significant, as it has contributed (H.P) comprising 5.7 % of the country’s
such useful animal species as water buffalo and camel landmass.
and plant species such as rice and sugarcane. India has • Himalayas. The entire mountain chain running
also been a secondary centre of domestication for animal from north-western to northeastern India,
species such as horse and goat, and such plant species as comprising a diverse range of biotic provinces
potato and maize. Animal species, which are reported to and biomes, 7.2 % of the country’s landmass.
be threatened in India, have been listed in Table . • Desert. The extremely arid area west of the
Aravalli hill range, comprising both the salty
India has 47 000 species of flowering and non-flowering desert of Gujarat and the sand desert of
plants representing about 12% of the recorded world’s Rajasthan. 6.9% of the country’s landmass.
flora. Out of 47 000 species of plants, 5 150 are endemic • Semi-arid. The zone between the desert and
and 2532 species are found in the Himalayas and the Deccan plateau, including the Aravalli hill
adjoining regions and 1 782 in the peninsular India. India range. 15.6 % of the country’s landmass.
is also rich in the number of endemic faunal species it

110 INSTITUTE FOR ADVANCED STUDIES, JODHPUR 
UNIT 10: ECOLOGICAL PRINCIPLES 
• Western ghats. The hill ranges and plains
running along the western coastline, south of
the Tapti river, covering an extremely diverse
range of biotic provinces and biomes. 5.8% of
the country’s landmass.
• Deccan peninsula. The largest of the zones,
covering much of the southern and southcentral
plateau with a predominantly deciduous
vegetation. 4.3 % of the country’s landmass.
• Gangetic plain. Defined by the Ganges river
system, these plains are relatively
homogenous. 11% of the country’s landmass.
• North-east India. The plains and non-
Himalayan hill ranges of northeastern India,
Apart from the biogeographic classifications described
above ecosystems can also be demarcated on the basis of
purely geographical or geological features like
mountains, islands, valleys, plateaux, oceans; on the
basis of vegetative cover like forests, grasslands,
mangroves and deserts; on the basis of climatic
conditions like arid and semi-arid areas, permanently
snow-bound areas, high rainfall areas; on the basis of soil
characteristic and other such criteria. In some
descriptions the biomes/ecosystems are clubbed together
into very general habitat classifications. The main natural
habitat types are:
1. Forests
The forest cover of the country is placed at 633 397 sq
km according to the forest survey of India assessment
(1997). This presents 19.27% of India’s total
geographical areas. India is endowed with diverse forest
types ranging from the Tropical wet evergreen forests in
North-Eastern to the Tropical thorn forests in the Central
and Western India.
2. Grasslands
In India the spread of grassland and shrubland is put at
12% of the total landmass while the planning
commission (1989) and Grasslands and Fodder research
Institute, Jhansi gives an estimate of about 3.7 to 3.9%.
The diversity of grasslands in India is high ranging from
semi-arid pastures of the western part of the Deccan
INSTITUTE FOR ADVANCED STUDIES, JODHPUR  111
with a wide variation of vegetation. 5.2% of
the country’s landmass.
• Islands. The Andaman and Nicobar Islands in
the Bay of Bengal, with a highly diverse set of
biomes. 0.03% of the country’s landmass.
• Coasts. A large coastline distributed both to
the west and east, with distinct differences
between the two; Lakshadeep islands are
included in this with the percent area being
negligible.
peninsula, the humid, semi-waterlogged tall grassland of
the Terai belt, the rolling shola grasslands of the western
ghat hilltops, and the high-altitude alpine pastures of the
Himalayas.
3. Wetlands
Wetlands cover 3% of the Indian landmass, or nearly 100
000 sq. Km. Wetlands in India harbor a vast variety of
life forms that are a part of the complex food of these
transitional ecosystems. About 320 species of birds are
associated with the Indian Wetlands. Apart from birds,
the wetlands support a diverse population of plants and
animals including 150 species of amphibians. Wetlands
are the habitat of some of the world’s endangered and
threatened flora and fauna. The Western and Central
flock of Siberian crane, one of the most endangered
cranes in the world, uses Keoladeo as its winter site. The
brown antlered deer (Cervus eldi eldi) or ‘sangai’ is
found only in phumadis (floating landmasses) of Lok
Tak Lake. Gahirmatha beach is a major breeding site of
olive ridley turtles. Chilka is the habitat of many
threatened species such as green sea turtle, Hawksbill
turtle, dugong, and blackbuck.
4. Mangroves
Government of India estimated mangrove cover of 674
000 ha, which is about 7% of the world’s mangrove.
Mangroves are salt-tolerant ecosystems in tropical and
subtropical regions. These ecosystems are largely
UNIT 10: ECOLOGICAL PRINCIPLES 
characterized by assemblage of unrelated tree genera that
share the common ability to grow in saline tidal zone.
India harbours some of the best mangroves swamps in
the world, located in the alluvial deltas of Ganga,
Mahanadi, Godavari, Krishna, and Cauveri rivers and on
the Andaman and Nicobar group of Islands. The largest
stretch of mangroves in the country lies in the
Sunderbans in West Bengal covering an area of about
4,200 sq. km. The predominant mangroves species are
Avicennia officinalis Excoecaria agallocha, Heritiera
fomes, Bruguiera parviflora, Ceriops decandra,
Rhizophora mucronata and Xylocarpus granatum.
Mangroves also harbour a number of molluscs,
polychaetes and honeybees. The Indian mangroves are
host to 105 species of fish, 20 kinds of shellfish, and 229
crustacean species. The Royal Bengal tiger is found in
the Sunderban mangroves. Different species of monkeys,
otters, deer, fishing cats, snakes and wild pigs are
common. A total of 117 species of migratory and
residential birds have been reported. The most common
birds are flamingos, storks, sea eagles, kites, kingfishers,
sandpipers, bulbuls, and whistlers.
5. Coral reefs
Accurate estimates of coral reef extent in the world are
not available. A rough estimate puts it at 600 000 sq Km
(Smith 1978) out of which 60% occurs in the Indian
Ocean region and most of it in south-east Asia. The
coral reef cover in Indian waters is roughly estimated
upto 19,000 sq. Km (Wafar 1992). Indian reefs belong to
the following categories:
1. PalkBay and Gulf of Mannar : Fringing
2. Gulf of Kachchh : Fringing, Patchy
3. Andaman and Nicobar Islands : Fringing
4. Lakshadeep Islands : Atolls
5. Central West coast : Patchy
The diversity of the Indian coral reefs is very impressive
with about 200 coral species belonging to 71 genera. The
richest being Andaman and Nicobar Islands which alone
harbors 179 species.
6. Deserts
In India, deserts extend over about 2% of the landmass.
Three kinds of deserts are noticeable in India:
• The sand desert of western Rajasthan and
neighbouring areas.
• The vast salt desert of Gujarat
• The high-altitude cold desert of Jammu and
Kashmir and Himachal Pradesh.
Desert fauna in India is also quite diverse, with about
1200 sp. of animals reported from Thar region of which
440 are vertebrates and 755 are invertebrates. Desert fox,
Desert cat, Houbara Bustard and some Sandgrouse
species are restricted to the Thar area (Rodgers and
Pawar 1988). In the remote part of Great Rann, Gujarat
lies the nesting ground of Flamingoes and the only
known population of Asiatic wild ass.
The cold deserts in India cover a vast area of 109 990 sq.
Km, about 87,780 sq. km in Laddakh (Jammu and
Kashmir) and 22,210 sq. Km in Lahaul -Spiti (Himachal
Pradesh). The diversity of the high altitude cold deserts
has been studied only recently with many insect species
being endemic. Interestingly the cold desert harbors
Kiang a close relative of the Indian wild ass found the
112 INSTITUTE FOR ADVANCED STUDIES, JODHPUR
Rann of Kachchh. Other distinctive animals include
Snow leopard, Yak, Tibetan antelope, Ibex, Blue sheep,
Tibetan gazelle, Woolly hare etc.
Biodiversity hotspots of India
In order to concentrate resources on those areas that are
most vulnerable, conservationists have identified certain
areas as biodiversity “hotspots.” The term was first used
by British ecologist Norman Myers in 1988 to designate
areas in which there is a disproportionate number of
endemic species (species that are found nowhere else)
and which are losing habitat at a high rate. In a February
2000 article in Nature, Myers and other ecologists
identified 25 “hotspots” that together comprise only 1.4
percent of the Earth’s surface yet contain 44 percent of
all species of higher plants and 35 percent of all land
vertebrate species.
Hotspots are defined according to their plant vegetation.
According to Myers’s definition, a hotspot has to contain
at least 0.5 percent of the world’s 300,000 plant
species as endemics. Plants are important because
vegetation is what determines the primary productivity of
an ecosystem. Most, but not all, of the hotspots are in
tropical areas; many are in developing countries where
populations rely on species-rich ecosystems for food,
firewood, cropland, and income from timber. In
Madagascar, for example, about 85 percent of the plants
and animals are found nowhere else in the world
Biodiversity hotspots are areas that are unusually rich in
species, most of which are endemic, and are under a
constant threat of being overexploited. Among the 25 hot
spots in the world, two are found in India. These are two
distinct areas: the Eastern Himalayas and the Western
Ghats and are also depicted in the National forest
vegetation map of India.
1. Eastern Himalayas
Phytogeographically, the Eastern Himalayas forms a
distinct floral region and comprises Nepal, Bhutan,
neighbouring states of east and north-east India, and a
contiguous sector Yunnan province in south western
China. In the whole of Eastern Himalayas, there are an
estimated 9000 plant species, with 3500 (i.e. 39%) of
them being endemic. In India’s sector of the area, there
occur some 5800 plant species, roughly 2000 (i.e. 36%)
of them being endemic. At least 55 flowering plants
endemic to this area are recognized as rare, for example,
the pitcher plant (Nepenthes khasiana). The area has
long been recognized as a rich centre of primitive
flowering plants and the area is recognized as ‘Cradle of
Speciation’.
Species of several families of monocotyledons,
Orchidaceae, Zingiberaceae and Arecaceae abound in the
area. Gymnosperms and pteridophytes (ferns) are also
well represented in the area. The area is also rich in wild
relatives of plants of economic significance, e.g. rice
banana, citrus, ginger, chilli, jute and sugarcane. The
region is regarded as the centre of origin and
diversification of five palms of commercial importance
namely, coconut, arecanut, palmyra palm, sugar palm
and wild date palm. Tea (Thea sinensis) is reported to
be in cultivation in this region for the last 40,000 years.
 
UNIT 10: ECOLOGICAL PRINCIPLES 
Many wild and allied species of tea, the leaves of which
are used as substitute of tea, are found growing in the
North East in the natural habitats. The ‘taxol’ plant
Taxus wallichiana is sparsely distributed in the region
and has come under red data category due to its over
exploitation for extraction of a drug effectively used
against cancer.
As regards faunal diversity, 63% of the genera of land
mammals in India are know from this area. During the
last four decades, two new mammals have been
discovered from the region: Golden Langur from
Assam – Bhutan region, and Namdapha flying
squirrel from Arunachal Pradesh indicating the
species richness of the region. The area is also a rich
centre of avian diversity – more than 60% of the
Indian birds are recorded in the North East. The region
also has two endemic genera of lizards, and 35 endemic
reptilian species, including two turtle. Of the 204 Indian
amphibians, at least 68 species are known from North
East, 20 of which are endemic. From Namdapha National
Park itself, a new genus of mammal, a new subspecies of
bird, 6 new species of amphibia, four new species of fish,
at least 15 new species of beetles and 6 new species of
flies have been discovered.
2. Western Ghats
The Western Ghats region is considered as one of the
most important biogeographic zones of India, as it is one
of the richest centres of endemism. Due to varied
topography and micro-climatic regimes, some areas
within the region are considered to be active zones of
speciation.
The region has 490 arborescent taxa, of which as many
as 308 are endemics this endemism of tree species shows
a distinct trend, being the highest (43%) in 8N-10°30’N
location and declining to 11% in 16N - 16°30’N location.
About 1 500 endemic species of dicotyledonous plants
are reported from the Western Ghats. 245 species of
orchids belonging to 75 genera are found here, of which
112 species in 10 genera are endemic to the region.
As regards the fauna, as many as 315 species of
vertebrates belonging to 22 genera are endemic, these
include 12 species of mammals, 13 species of birds, 89
species of reptiles, 87 species of amphibians and 104
species of fish. The extent of endemism is high in
amphibian and reptiles. There occur 117 species of
amphibians in the region, of which 89 species (i.e. 76%)
are endemic. Of the 165 species of reptiles found in
Western Ghats, 88 species are endemic. Many of the
endemics and other species are listed as threatened.
Nearly 235 species of endemic flowering plants are
considered endangered. Rare fauna of the region
includes: Lion Tailed Macaque, Nilgiri Langur, Nilgiri
Tahr, Flying Squirrel, and Malabar Gray Hornbill.
Response
The Ministry of Environment and Forests (MoEF) is the
nodal agency in the Government of India for planning,
promotion, coordination, and overseeing the
implementation of the environmental and forestry
programmes. The MoEF is also the focal point for
implementation of the Convention on Biological
INSTITUTE FOR ADVANCED STUDIES, JODHPUR  113
Diversity. The mandates of the Ministry interalia include
survey of flora, fauna, forests and wildlife, and
conservation of natural resources. These objectives are
supported by legislative and regulatory measures. A
number of institutions affiliated with the Ministry are
involved in the work related to various aspects of
biological diversity. Survey and inventorization of the
floral and faunal resources are carried out by the
Botanical Survey of India (BSI) established in 1890, and
the Zoological Survey of animals species have been
recorded by the BSI and ZSI respectively. The Survey
organizations have published over the years, documents
on flora and fauna at country, state and in some cases
district levels and for selected ecosystems. Besides,
extensive reports on inventories of resources indicating
level of biodiversity in selected areas have also been
brought out. The Surveys have also published Red Data
Books (Originally printed by International council for
conservation of nature and natural resources) on
endangered species. The voucher specimens are
preserved in Central National Herbarium (CNH) of BSI
and National Zoological Collection (NZC) of ZSI. The
Forest Survey of India publishes every three years, a
State of Forest in India report based on remote sensing
and ground truth data.
Existing policy response In situ conservation (within
natural habitat)
India has a network of 85 national parks and 448
sanctuaries, covering 4.2% of the its land area [MoEF
1998]. This number has progressively increased over the
last twenty-five years. Areas having significant
biodiversity value are declared national parks or
sanctuaries under the Wild Life (Protection) Act of 1972
(henceforth referred to as WL Act), as amended in 1991.
Before this act, national parks and sanctuaries were
being set up but under various state or area specific acts.
With the coming of this act, all areas notified under any
other act became parks or sanctuaries notified under this
act, Under the WL Act, national parks are given a
higher level of protection and no human use activity is
permitted within them: The act specifies that: "No person
shall destroy, exploit or remove any wild life from a
National Park or destroy or damage the habitat of any
wild animal or deprive any wild animal of its habitat
within such National Park except under and in
accordance with a permit granted by the Chief Wild Life
Warden and no such permit shall be granted unless the
State Government, being satisfied that such destruction,
exploitation or removal of wild life from the National
Park is necessary for the improvement and better
management of wild life therein, authorises the issue of
such permit." [Section 35(6) of the Act]. Also, no private
land holding or right is allowed within a national park.
Sanctuaries are accorded a lesser level of protection,
and grazing and some community or individual rights
can be permitted. Under the WL Act, national parks are
fully protected from all human disturbance and,
consequently, correspond to the revised category Ia
(Scientific Reserves) of the IUCN categorisation system
for protected areas. A sanctuary, where grazing and
various other rights can be permitted, corresponds to
IUCN category IV Habitat and Wildlife Management
Area).
UNIT 10: ECOLOGICAL PRINCIPLES 
Some important measures taken are as follows:
• A programme entitled “Eco-development” for in situ
conservation of biological diversity involving local
communities has been initiated in recent years. The
concept of eco-development integrates the ecological and
economic parameters for sustained conservation of
ecosystems by involving the local communities with the
maintenance of earmarked regions surrounding protected
areas. The economic needs of the local communities are
taken care of under this programme through provision of
alternative sources of income and a steady availability of
forest and related produce. To conserve the respective
ecosystems, a Biosphere Reserve Programme is being
implemented. Twelve biodiversity rich areas of the
country have been designated as Biosphere Reserves
(Table 9.5) applying the diversity and genetic integrity of
plants, animals and microorganisms in their totality as
part of the natural ecosystems, so as to ensure their self-
perpetuation and unhindered evolution of the living
resources.
• Programmes have also been launched for scientific
management and wise use of fragile ecosystem. Specific
programmes for management and conservation of
wetlands, mangroves, and coral reef systems are also
being implemented. 21 wetlands, 15 mangrove areas and
4 coral reef areas have been identified for management.
National and sub-national level committees oversee and
guide these programme to ensure strong policy and
strategic support.
• Six internationally significant wetlands of India have
been declared as “Ramsar Sites” under the Ramsar
Convention. To focus attention on urban wetlands
threatened by pollution and other anthropogenic
activities, State Governments were requested to identify
lakes that could be include the National Lake
Conservation Plan. The activities of the NLCP include
formulation of perspective plans for conservation based
on resource survey using remote sensing technology and
GIS studies on biodiversity and related ecological
matters, prevention of pollution from point and non-point
sources, treatment of catchment, desilting and weed
control.
• Wild Life Protection Act is in the final stage of
revision and provisions have been made for conservation
reserves and com- munity reserves to allow restrictive
use to make it more people oriented. Presently
Biodiversity Act which is in the final stage, has got the
component of National
Biodiversity Authority to control access to genetic
resources form international community. There will also
be State Biodiversity Boards to control access to
domestic consumers. Under the World Heritage
Convention, five natural sites have been declared as
“World Heritage Sites”, the name of which are under:
• The Tura Range in Gora Hills of Meghalaya is a gene
sanctuary for preserving the rich native diversity of wild
Citrus and Musa species.
• Sanctuaries for rhododendrons and orchids have been
established in Sikkim.
114 INSTITUTE FOR ADVANCED STUDIES, JODHPUR
• Large mammal species targeted protection based on
the perception of threat to them have been under
implementation
• Project Tiger. A potential example of an highly
endangered species is the Indian Tiger (Panthera tigris)
The fall and rise in the number of Tiger’s in India is an
index of the extent and nature of conservation efforts. It
is estimated that India had about 40 000 tigers in 1900,
and the number declined to a mere about 1 800 in 1972.
Hence, Project Tiger was launched in 1973 with the
following objectives:
• To ensure maintenance of available
population of Tigers in India for scientific,
economic, aesthetic, cultural and ecological
value
• To preserve, for all times, the areas of such
biological importance as a national heritage for
the benefit, education and enjoyment of the
people
• At present there are 25 Tiger Reserves
spreading over in 14 states and covering an
area of about 33 875 sq km and the Tiger
population has more than doubled now due to a
total ban on hunting and trading tiger products
at national and international levels and the
implementation of habitat improvement and
anti-poaching measures (MoEF 2000)
• Project Elephant was launched in 1991-92 to assist
States having free ranging population of wild elephants
to ensure long term survival of identified viable
populations of elephants in their natural habitats. Major
activities of Project Elephant are:
• Ecological restoration of existing natural
habitats and migratory routes of elephants.
Development of scientific and planned
management for conservation of elephants
habitats and value population of wild Asiatic
elephants in India
• Promotion of measures for mitigation of man-
elephant conflict in crucial habitats and
moderating pressures of human and domestic
stock activities in crucial elephant habitats
• Strengthening of measures for protection of
wild elephants from poachers and unnatural
caused of death. Research on Project Elephant
management related issues Public education
and awareness programmes • Eco-
development • Veterinary care
• Rhinos have been given special attention in selected
sanctuaries and national parks in the North East and
North-west India. All these programmes, though
focussed on a single species, have a wider impact as they
conserve habitats and a variety of other species in those
habitats. The Ministry of Environment and Forests
constituted the National Afforestation and Eco-
development Board (NAEB) in August 1992. National
Afforestation and Eco-development Board has evolved
specific schemes for promoting afforestation and
management strategies, which help the states in
developing specific afforestation and management
strategies and eco-development packages for augmenting
biomass production through a participatory planning
process of Joint Forest Management and microplanning.
 
UNIT 10: ECOLOGICAL PRINCIPLES 
Ex-situ conservation (outside natural habitats)
To complement in situ conservation, attention has been
paid to ex-situ conservation measures. According to
currently available survey, Central Government and State
Government together run and manage 33 Botanical
Gardens. Universities have their own botanical gardens.
There are 275 zoos, deer parks, safari parks, aquaria etc.
A Central Zoo Authority was set up to secure better
management of zoos. A scheme entitled Assistance to
Botanic Gardens provides one-time assistance to botanic
gardens to strengthen and institute measure for ex-situ
conservation of threatened and endangered species in
their respective regions.
• Maximum biodiversity among vertebrates:
Pisces (osteochythes/bony fishes)
• Maximum biodiversity among invertebrates:
Arthropods (insecta)
J5. CRYOPRESERVATION
Cryopreservation (Gr. Kryos means frost) refers to
‘preservation in the frozen state’. It means storage at
very low temperature such as over solid carbon dioxide
(- 790C), in deep freezers (- 800C), in vapour phase
nitrogen (- 1500C or in liquid nitrogen (- 196°C).
The plant material is generally preserved and maintained
in liquid nitrogen. There are several techniques for
preservation of microbial cells including
cryopreservation too. Moreover, cryopreservation of
animal stock cells and cells lines is preferred to protect
them from genetic drift (arising due to genetic
instability), microbial contamination, cross
contamination by other cell lines, incubator failure,
senescence, etc.
A. CRYOPRESERVATION OF PLANT STOCK
CELLS
Due to gradual disappearance of economic and rare plant
species the necessity for storage of genetic resources of
plant realm in general and agricultural plants in particular
was realized by the biologists. The conventional methods
of storage rail to prevent from losses caused by:
(i) attack of pathogens and pests,
(ii) climatic disorders,
(iii) natural disorders, and
(iv) political and economic causes.
However, the conventional methods could not save the
viability of short lived seeds of economic plants, for
example, oil palm (Elaeis guineensis), rubber (Hevea
brasiliensis), Citrus sp. and Coffes sp.
These materials are stored at low temperature, due to
which growth-rate of cells retards consequently
biological activities are conserved for long time.
Cryobiology deals with the stud: of metabolic activities
and their responses in plant materials (and animal cells)
stored at low temperature (-196°C) by using liquid
nitrogen in the presence of cryoprotectants.
Dodds am Roberts (1985) have discussed three principal
methods used for growth suppression in plant tissue
culture:
INSTITUTE FOR ADVANCED STUDIES, JODHPUR  115
• Maximum biodiversity occurs in: tropics (5O to
25O North South)
• Countries with maximum biodiversity: Brazil,
Columbia, China
• In India maximum biodiversity occurs along
Western Ghats.
• Wild life which become extinct in near past:
Dodo (Mauritus/1680), Yellow headed Macaw
(Jamaica/1765), Stellar sea cow (Bering sea
/1767) and many Australian animals like
Wallabies, Tasmanian emus, Tasmanian Tiger,
Rat Kangaroo etc.
• Red Data book which have list of Threatened
organism is published by Union of Nature and
Natural resources (IUCN)
(i) the alteration of physiological conditions of culture
i.e. temperature or gas composition within the vessel;
(ii) changing the composition of basal medium e.g. using
sub or supra-optimal concentrations of nutrients (some
factors essential for normal growth may be either omitted
or employed at a reduced level) and
(iii) supplementing the medium with growth retardants
(e.g. abscisic acid) or osmoregulatory compounds (e.g.
mannital, sorbital, etc.).
Storage at reduced temperature has been very effective
for tissue culture of most of the plant species such as
potato, cassava (Manihot esculentum), pea (Pisum
sativum), chickpea (Cicer arietinum), rice (Oryza sativa),
wheat (Triticum vulgare), coconut (Cocos nucifera), oil
palm (E. guineensis) strawberry (Fragaria vesca) and
sugarcane (Saccharum officinarum).
1. Difficulties in Cryopreservation: A number of
reviews available during the last two decades illustrate
the significant progress made in this field and also the
outline of the existing problem. The difficulties are:
(i) high specific feature of plant cells, such as their large
size, strong vacuolization and abundance of water,
(ii) cell damage during freezing and subsequent thawing
caused by ice cyrstals formed inside the cells and by cell
dehydration, and
(iii) gradual formation of large crystals of more than 0.1
µm whose facets rupture many cell membranes.
However, in the presence of cryoprotectants (the
chemicals decreasing cryo-destruction) and reduced
temperature, free water has enough time to leave the
cells. Therefore, it can freeze on the crystal surface in the
solution. This results in marked dehydration and
protoplast shrinkage. Excessive time and degree of
plasmolysis are the reasons of cell destruction during
slow freezing, since they cause irreversible contraction of
the plasmalemma.
2. Methods of Cryopreservation: The freezing-storage-
thawing cycle is an external procedure which is
accomplished in the following basic stages:
2.1 Selection of Materials: For selecting the plant
materials a number of factors are taken into account; the
UNIT 10: ECOLOGICAL PRINCIPLES 
important ones are, nature and density of cells in the
vials/ampules to be cryopreserved; because the
cryoability of the cell cultures depends on these. Young
meristematic, highly cytoplasmic and small cells which
are non-vacuolated and thin walled and in small
aggregates, are good materials to be selected for this
purpose. Cell density in vials or ampules should be high,
as it shows prolonged survival at high cell density.
The ability of explant to survive at -196°C is influenced
by the ability of their morphological and physiological
conditions. Different types of explants which are used in
cryopreservation are apical meristem and plant organs,
ovules, anther/pollen, embryos, protoplasts, etc.
Log phase cell suspension of many plant species has
successfully been cryopreserved. The cultured cells are
not the ideal system that can be cryopreserved. In
constrast the organised structures (e.g. shoot pieces,
young plantlets, embryos) are preferred for
cryopreservation.
2.2 Addition of Cryoprotectants: During freeze
preservation the two strong sources cause cell damage:
formation of large ice crystals inside the cells that
rupture cell organelles and cells itself, and intracellular
concentration of solutes that increases to toxic levels due
to dehydration before or during storage.
Cryoprotectors are the chemicals which decrease
cryodestruction. These are sugars, glycols, sugar
alcohols, alcohols, polyvinylpyrrollidone, polyethylene
glycol (PEG), polyethylene oxide (PEO), dextrans,
hydroxy starch, glycerine, sucrose, and some amino acids
(e.g. proline). Bajaj (1987) has advised to use a mixture
of two or three cryoprotectants at low concentrations
rather than a single cryoprotectant at a high concentration
as it could be toxic. During treatment, the cultures should
be maintained in ice to avoid deleterious effects.
Dimethyl sulfoxide (DMSO), sucrose, glycerol and
proline are most frequently used cryoprotectants. DMSO
has proved an excellent cryoprotectant because:
(i) it has low molecular weight,
(ii) it is easily miscible with solvent,
(iii) it is toxic at low concentration,
(iv) it is easily permeable into cells and easily
washable.
The material to be preserved is put in culture medium
and treated with a cryoprotectant. Such material is
transferred to sterile cryovials or ampoules which are
made up of polypropylene. Gradually 5-10%
cryoprotectant is added into the ampoules. It is tightly
closed with a screw cap.
2.3 Freezing. Freezing should be done in such a way that
it does not cause intracellular freezing and crystal
formation. To avoid this problem, regulated rate of
cooling or pre-freezing is done. Moreover, freezers have
also been developed which allow the uniform
temperature decrease at a desired rate, commonly not
less than 1°C per minute. In 1987, the Institute of
Cryobiology and Cryomedicine of the Ukrainian
Academy of Sciences (erstwhile U.S.S.R.) devised the
programme freezer which envisaged lower rate (0.5°C
116 INSTITUTE FOR ADVANCED STUDIES, JODHPUR
per minute) of temperature decrease. The following types
of freezing can be done:
i. Rapid freezing: This method is simple and easy to
handle. After placing the plant material, the cryovials are
put into liquid nitrogen which causes a decrease in
temperature. Freezing is done quickly so that there
should be least change or development of intracellular
crystals. Ultra cooling prevents ice crystals. To achieve
this objective, dry ice (C02) can be used instead of
nitrogen. Rapid freezing of several plant materials has
been done, for example, somatic embryos and shoot tips
of Brassica napus, strawberry, potato, etc.
ii. Slow freezing: In this method the rate of freezing is
slow i.e. 0.1 -10° C per minute. This facilitates the flow
in water from inside to outside. Therefore, extracellular
ice crystals are formed but not intracellular crystals.
Consequently, cell cytoplasm gets concentrated to cause
cellular dehydration. Such dehydrated cells survive for a
longer duration. Generally slow freezing has been done
by using computer-controlled freezers. Meristems of
potato, cassava, strawberry, etc. have successfully been
cryopreserved.
iii. Stepwise freezing: It includes the earlier described
two methods. In this method temperature gets lowered by
-20 to -40°C. It allows protective freezing of the cells.
Further, freezing is stopped for 30 minutes. Thereafter, it
is rapidly freezed in liquid nitrogen to get -196°C. By
doing such stepwise decline in temperature, formation of
big crystal is increased and good results are obtained.
Excellent results have been obtained with suspension
cultures and strawberry by adopting this method.
2.4 Storage in Liquid Nitrogen: If the cells are not
stored at sufficiently low temperature, an additional
injury to the cultures may be caused. The storage
temperature should be such that it stops all metabolic
activity and prevents biochemical injury. Prolonged
storage of frozen materials is possible only when the
temperature is lower than -130°C. This can be simply
achieved with the help of liquid nitrogen, which keeps
the temperature -196°C. Popove (1988) stored the
cultures of carrot cells for about 5 years by doing so.
2.5 Thawing: Thawing is the process of releasing the
vials containing cultures from the frozen state to elevate
the temperature between 35 and 45°C. It should be done
quickly but without overheating. As soon as the last ice
crystals disappear, the vials are transferred into a water
bath maintained at 20-25°C. During the course of
freezing and thawing, major biophysical changes occur
in the cell. The freshly thawed cells need suitable
nourishment because they are prone to further damage.
2.6 Washing and Reculturing: Washing of plant
materials is done to remove the toxic cryoprotectants.
When low toxic or non-toxic cryoprotectants are used,
the cultures should not be washed, but simply re-
cultured. Washing becomes necessary only when
cryoprotectants have toxic effects on cells. Washing
follows the following procedure: dilution, resuspension,
centrifugation and removal of cells. It is, however,
possible that some cells die due to storage stress and the
most stable ones survive. Therefore, determination of
cell viability by culturing them on growth medium is
essential.
 
UNIT 10: ECOLOGICAL PRINCIPLES 

cryopreserved. Some of the observations made are as

Cell viability test can be done by using FDA staining and below
growth measurements. The parameters of growth (i) Cryopreservation of cell lines: For example, cell
measurements are counting cell number, dry and fresh suspensions (soyabean, tobacco, dhatura, carrot) and
weight, mitotic index, etc. Mitotic index (MI) is counted somatic hybrid protoplasts (rice x pea, wheat x pea).
by using the following formula: (ii) Cryopreservation of pollen and pollen embryos: For
example, fruit crops, trees, mustard, carrot, peanut, etc.
MI = Number of cells destined to cell division (iii) Cryopreservation of excised meristems: For
Total number of cells both dividing and undividing example, potato, sugarcane, chickpea, peanut, etc.
(iv) Cryopreservation of germplasm of vegetatively
2.7 Regeneration of Plantlets. The viable cells are propagated crops: potato, sugarcane, etc.
cultured on non-specific growth media to regenerate into (v) Cryopreservation of recalcitrant seeds and embryos:
plantlets. Bajaj (1987) has given an extensive list of Large sized seeds that are short lived and abortive, such
works on cryopreservation of cells, tissue, and organ as oil palm, coconut, walnut, mango and cocoa.
culture of various plants e.g. potato, cassava, sugarcane,
soyabean, groundnut, carrot, cotton, citrus, coconut, etc. B. CRYOPRESERVATION OF ANIMAL STOCK
CELLS
3. Plant Cell Bank/Germplasm Bank/Cell Cryobank:
Cryopreservation of genetic stock i.e. germplasm (or
vegetatively propagated crops, recalcitrant producing
plants, rare plant species, medicinal, horticultural and
forest plants, and VAM fungi) is a novel approach for
their conservation in liquid nitrogen on a long term basis.
To achieve this goal, a plant cel1 bank (= germplasm
bank and cell cryobank) has been suggested by Bajaj.
Suggestions have also been made that germplasm bank
should be attached to some of the International Research
Institutes (e.g. IRRI) that would hold responsibility for
the storage, maintenance, distribution (at national and
international level), and exchange of these disease free
germplasm of the important plants. Fig. shows the
potential and prospects of cryopreservation of plant cell,
tissue and organ and establishment of germplasm bank.
Facilities for storage of genetic stock of plants can be
developed in large sized cylinders (30- 50 litres capacity)
where liquid nitrogen does not require refilling for 6-8
months.
Development of animal cell line is expensive, time con-
suming and labour intensive. It is essential to protect this
Thus, germplasm bank is such a device where facilities
considerable investment by preserving the cell line so
of cryopreservation of genetic resources of a variety of
that it can further be used whenever required. In
plants are available and on demand, the germplasm can
continuous culture, cell lines are prone to variation due to
be supplied nationally and internationally.
selection in early passage culture, senescensce in finite
cell lines, genetic instability in continuous cell lines,
4. Pollen Bank: Besides germplasm bank, the storage of
cross-contamination, equipment failure, etc.
pollen grains in liquid nitrogen and establishment of
pollen bank have also been suggested to retain their
1. Selection of Cell Line and Standaradisation of
viability for various lengths of time. The freeze storage
Culture Conditions: A cell line possess different
of pollen would enable: (i) hybridization between plants
properties. Therefore, a cell line of different properties is
with flowers at different times, (ii) growth at different
selected. Continuous cell lines are cloned and suitable
places, (iii) reducing the dissemination of diseases by
clone is selected and grown to get sufficient bulk of cells
pollination vectors, and (iv) maintenance of gennplasm
required for freezing. Similarly, the finite cell line is also
and enhancement of longevity.
grown to about fifth population doubling to get sufficient
amount of cells. Before freezing, the cells need to be kept
5. Achievement Made Through Cryopreservation:
under standard culture conditions, characterised and
Various forms of plant materials viz. cell suspensions
checked for contamination. Continuous cell lines offer
clones, callus, tissues, somatic embryos, root/shoot tips,
several advantages over finite cell lines such as: (i) they
propagules (tubers) pollen grains, etc. have been
survive indifinitely, (ii) they grow more rapidly, (iii) they
preserved in liquid nitrogen for prolonged time and
can be cloned more easily but they may be less stable
tested for their survival and regeneration potential.
genetically. Usually the finite cell lines are diploid and
No doubt, in most of the cases, the cells/tissues, organs
stable, but harder to clone. They grow more slowly and
regenerated into plants. Bajaj (1987) has described a
eventually die or transfer (Table 1)
number of plant species that have been successfully
Table 1: Requirement before freezing
Acquistion Finite cell line Freeze at early passage (<5 subcultures)
Continuous cell line Clone, select and characterize, amplify
Standardisation Medium Select optimal medium and adher to it

INSTITUTE FOR ADVANCED STUDIES, JODHPUR  117
UNIT 10: ECOLOGICAL PRINCIPLES 
Serum (if used)
Substrate
Ralidation Provenance
Authentication
Transformation
Contamination
Cross-contamination
The selection and their phenotypic expression of
different cell types are influenced by the type of medium
used. The best method to eliminate serum variation is to
use serum-free medium. Unfortunately serum-free
medium is not available for all cell types. Conversion is
costly and time consuming also. However, if serum is
required, a batch is selected and used throughout every
stage of cryopreservation.
Table 2. Stages in cell line preservation.
Stage Source No. of ampules
Token freeze Originator 1-3
Seed stock Originator stock token freeze 12
Distribution stock Test thaw from seed stock 50-100
User Stock Distribution stock - None
The seed stock of most cell lines should be protected. It
should not be made available for general use. A
distribution stock should be frozen and ampoules from
this stock for a long period, should freeze down their
own user stocks. When the work is over, it should be
discarded. The distribution stock should be replenished
from the seed stock after being over. When the seed
stock falls below five ampoules, it should be replenished
before issuing any other ampoules.
The cells suspension used for cryopreservation should
have high cell density (106-107 cell/ml). By using
glycerol (10-15%) or DMSO (5-10%) as preservative,
cells are frozen slowly at about 1°C per minute. When
temperature is below -70oC, the ampoules are rapidly
transferred to liquid nitrogen. The ampoules are put in
liquid nitrogen or in gas phase above the liquid. Then the
ampoules cool rapidly to about -50oC but cooling rate
falls after that. Therefore, the ampoules should be left
J6. Biosphere Reserves of India
The programme of Biosphere Reserve was initiated
under the 'Man & Biosphere' (MAB) programme by
UNESCO in 1971.
The purpose of the formation of the biosphere reserve is
to conserve in situ all forms of life, along with its
support system, in its totality, so that it could serve as a
referral system for monitoring and evaluating changes in
natural ecosystems.
The first biosphere reserve of the world was established
in 1979, since then the network of biosphere reserves has
increased to 531 in 105 countries across the world
(MAB, 2008). Presently, there are 14 existing biosphere
reserves in India.
118 INSTITUTE FOR ADVANCED STUDIES, JODHPUR
Select a batch for use at all stages
Standardise on one type and supply
Record details of life history and properties
Check characteristics of cells line against provenance
Determine transformed status
Test for microbial contamination i.e. sterility test of material
Adopt criteria to confirm identity e.g. using DNA fingerprinting
2. Stages of Cryopreservation
When the cells are available from subculturing a primary
culture or acquired cell lines, a few cryovials or
ampoules are frozen. It is called token freeze. A seed
stock is stored frozen after producing the propagated cell
lines with the desired characteristics and free of
contamination. The various stages in cell line
preservation are given in Table 2.
Distribution Validation
None Prevenance only
None Viability, authenticity,
transformation, contamination
User including Viability contamination,
other laboratory cross-contamination
Viability, contamination,
overnight at -70oC before transferring to liquid nitrogen.
However, when the cells are to be used, they are thawed
rapidly and re-seeded at a high concentration to optimise
their recovery.
3. Cell Banks: Several cell banks have been established
for the secure storage and distribution of validated cell
lines. Several cell lines may come under patent
restriction (like hybridomas and other genetically
modified cell lines); hence, they should be provided
patent repositories with limited access. One must obtain
the initial seed stock from a renowned cell bank where
characterization and quality control of cell lines are done.
One must deposit the valuable cultures to a cell bank
besides maintaining one's frozen stock. This will help to
protect the loss of cell lines and its distribution to others
too. Data bank of cell lines has also been built up. This
type of data bank provides a vast increase in the amount
of available materials.
The Indian government has established 14 Biosphere
Reserves of India, (categories roughly corresponding to
IUCN Category V Protected areas), which protect larger
areas of natural habitat (than a National Park or Animal
Sanctuary), and often include one or more National
Parks and/or preserves, along buffer zones that are open
to some economic uses.
Protection is granted not only to the flora and fauna of
the protected region, but also to the human communities
who inhabit these regions, and their ways of life.
Four of the fourteen biosphere reserves are a part of the
World Network of Biosphere Reserves, based on the
UNESCO Man and the Biosphere (MAB) Programme
list.
 
UNIT 10: ECOLOGICAL PRINCIPLES 

• Nilgiri Biosphere Reserve  

• Nanda Devi Biosphere Reserve
• Sundarbans Biosphere Reserve
• Gulf of Mannar Biosphere Reserve  

   

Biosphere reserves of India (area wise)

Area
Year Name Location State Type
(km²)

Indian part of Gulf of Mannar between India

1 1989 Gulf of Mannar Tamil Nadu Coasts 10500
and Sri Lanka

Part of delta of Ganges and Barahamaputra Gangetic

2 1989 Sunderbans West Bengal 9630
river system Delta

Parts of Chamoli District, Pithoragarh District West

3 1988 Nanda Devi Uttranchal 5860
& Almora District Himalayas

Part of Wynad, Nagarhole, Bandipur and

Nilgiri Biosphere Tamil Nadu, Kerala Western
4 1986 Mudumalai, Nilambur, Silent Valley and 5520
Reserve and Karnataka Ghats
Siruvani Hills

East
5 1998 Dehang Debang Part of Siang and Debang valley Arunachal Pradesh 5112
Himalayas

Pachmarhi Biosphere Parts of Betul District, Hoshangabad District

6 1999 Madhya Pradesh Semi-Arid 4926
Reserve and Chhindwara District

Deccan
7 1994 Simlipal Part of Mayurbhanj district Orissa 4374
Peninsula

Achanakamar - Madhya Pradesh,

8 2005 Part of Annupur, Dindori and Bilaspur districts 3835
Amarkantak Chattisgarh

Part of Kokrajhar, Bongaigaon, Barpeta, East

9 1989 Manas Assam 2837
Nalbari, Kamrup and Darrang District Himalayas

East
10 2000 Kanchanjunga Parts of Kanchanjunga Hills Sikkim 2620
Himalayas

Agasthyamalai Neyyar, Peppara and Shenduruny Wildlife Western

11 2001 Kerala 1828
Biosphere Reserve Sanctuary and their adjoining areas ghats

Great Nicobar Southern most islands of Andaman and Andaman and

12 1989 Islands 885
Biosphere Reserve Nicobar Islands Nicobar Islands

East
13 1988 Nokrek Part of Garo Hills Meghalaya 820
Himalayas

Part of Dibrugarh District and Tinsukia East

14 1997 Dibru-Saikhowa Assam 765
District Himalayas

J7. Project Tiger

INSTITUTE FOR ADVANCED STUDIES, JODHPUR  119
UNIT 10: ECOLOGICAL PRINCIPLES 
Project Tiger is a wildlife conservation project initiated
in India in 1972 to protect the Bengal Tigers. It was
launched on April 1, 1973 and has become one of the
most successful wildlife conservation ventures. The
project aims at tiger conservation in specially constituted
tiger reserves representative of various biogeographical
regions throughout India. It strives to maintain a viable
tiger population in their natural environment.
In 2007, there were more than 40 Project Tiger wildlife
reserves covering an area of 37,761 km². Project Tiger
helped increase the population of these tigers from 1,200
in the 1970s to 3,500 in 1990s. However, a 2008 census
held by Government of India revealed that the tiger
population had dropped to 1,411. Since then the
government has pledged US$153 million to further fund
the project, set-up a Tiger Protection Force to combat
poachers, and has relocated more than 200,000 villages
to minimize human-tiger interaction. The efforts did
pay-off when in July 2008, the Sariska Tiger Reserve,
whose tiger population was nearly wiped out in 2005,
had a recorded tiger population of 21.
History
At the turn of the 20th century, one estimate of the tiger
population in India placed the figure at 45,000. The first
ever all-India tiger census was conducted in 1972 which
revealed the existence of only 1827 tigers. A recent
report published by the National Tiger Conservation
Authority estimates only 1411 adult tigers in existence in
India (plus uncensused tigers in the Sundarbans). The
project was launched in 1973, and various tiger reserves
were created in the country based on a 'core-buffer'
strategy. Management plans were drawn up for each
tiger reserve based on the principles outlined below:
• Elimination of all forms of human exploitation and
biotic disturbance from the core area and rationalization
of activities in the buffer zone.
• Restricting the habitat management only to repair the
damages done to the eco-system by human and other
interferences so as to facilitate recovery of the eco-
system to its natural state.
• Monitoring the faunal and floral changes over time
and carrying out research about wildlife.
Global organizations, such as the World Wildlife Fund
(WWF), contributed much funding to Project Tiger.
Eventually, however, it was discovered that the project's
field directors had been manipulating tiger census
numbers in order to encourage more donations. In fact,
the numbers were so exaggerated as to be biologically
impossible in some cases.
In addition, Project Tiger's efforts were damaged by
poaching, as well as the Sariska debacle and the latest
Namdapha tragedy, both of which were reported
extensively in the Indian media.
In the wake of these incidents, tiger activists and
environmentalists like Valmik Thapar and Belinda
Wright have demanded that the Prime Minister establish
an independent census to determine the actual number of
the tigers in India. (The country is already conducting a
nationwide tiger census, but its objectivity has been
120 INSTITUTE FOR ADVANCED STUDIES, JODHPUR
questioned by some because it relies on persons known
to have fudged previous census results.)
Initially, 9 tiger reserves were established in different
States during the period 1973-74, by pooling the
resources available with the Central and State
Governments. These nine reserves covered an area of
about 13,017km² -- viz Manas (Assam), Palamau
(Bihar), Similipal (Orissa), Corbett (U.P.), Kanha
(M.P.), Melghat (Maharashtra), Bandipur (Karnataka),
Ranthambhore (Rajasthan) and Sunderbans (West
Bengal). The World Wildlife Fund For Nature has given
Project Tiger assistance in the form of equipment,
expertise and literature worth US $ 1 million. There are
28 tiger reserves in India. Project Tiger was a pet project
of Indira Gandhi, the then Prime Minister of India. The
main achievements of this project are excellent recovery
of the habitat and consequent increase in the tiger
population in the reserve areas, from a mere 268 in 9
reserves in 1972 to above one thousand in 28 reserves in
2006. Tigers, being at the apex of the food chain, can be
considered as the indicator of the integrity of the
ecosystem. Thus, 'Project Tiger' is basically the
conservation of the entire ecosystem and apart from
tigers, all other wild animals' population have also
increased in the project areas.
In the meantime, the struggle to save the Indian tiger
remains difficult as poachers kill the endangered animal
to meet an international demand for tiger parts. Wright,
for example, has cited a strong demand for tiger skins in
China and Tibet as serious problem in tiger preservation.
According to the latest census figures as of 2007, the
tiger population in India has further dropped; particularly
in the state of Madhya Pradesh, where 65 percent of the
tigers have vanished, and the states of Chattisgarh,
Maharashtra and Rajasthan, where there are less than
100 tigers each. The primary reason attributed to this
increasing drop in numbers is poaching. However, the
method used to count the number of tigers earlier was by
identifying pugmarks, which could have resulted in
figures larger than actual. The latest method which
includes analysing habitat, prey base densities and
camera trappings, is more precise and scientific.
Present Organization and aims
The Project Tiger was meant to identify the limiting
factors and to mitigate them by suitable management.
The damages done to the habitat were to be rectified so
as to facilitate the recovery of the ecosystem to the
maximum possible extent. The overall administration of
the project is monitored by a 'Steering Committee'.A
'Field Director' is appointed for each reserve, who is
assisted by the field and technical personnel. At the
Centre, a full-fledged 'Director' of the project
coordinates the work for the country.
Wireless communication system and outstation patrol
camps have been developed within the tiger reserves,
due to which poaching has declined considerably. Fire
protection is effectively done by suitable preventive and
control measures. Voluntary Village relocation has been
done in many reserves, especially from the core area..
Livestock grazing has been controlled to a great extent in
 
UNIT 10: ECOLOGICAL PRINCIPLES 

the tiger reserves. Various compensatory developmental
works have improved the water regime and the ground
and field level vegetation, thereby increasing the animal
density
Future plans
Wildlife protection and crime risk management in the
present scenario requires a widely distributed
Information Network, using state-of-the-art information
and communication technology. This becomes all the
more important to ensure the desired level of protection
in field formations to safeguard the impressive gains of a
focused project like 'Project Tiger'. The important
elements in Wildlife protection and control are:
Mapping/plotting the relative spatial abundance of wild
animals, identification of risk factors, proximity to risk
factors, sensitivity categorization, crime mapping and
immediate action for apprehending the offenders based
on effective networking and communication. Space
technology has shown the interconnectivity of natural
and anthropogenic phenomena occurring anywhere on
earth. Several Tiger Reserves are being linked with the
Project Tiger Directorate in the GIS domain for Wildlife
Crime Risk Management. A Tiger Atlas of India and a
Tiger Habitat & Population Evaluation System for the
country is being developed using state-of-the-art
technology. This involves:
• Mapping , data acquisition and GIS modeling
• Field data collection and validation
• Data Maintenance , Dissemination and Use
The following potential tiger habitats in the country are
being covered:
• Shivalik-Terai Conservation Unit (Uttaranchal, UP,
Bihar, West Bengal, Nepal)
• North east Conservation Unit
• Sunderbans Conservation Unit
• Central Indian Conservation Unit
• Eastern Ghat Conservation Unit
• Western Ghat Conservation Unit
developmental priorities of the Government. This calls
for a cross-sectoral and cross-disciplinary approach.
Tigers now need a "preservationist" approach. Regional
planning is important around Tiger Reserves to foster
ecological connectivity between protected areas through
restorative inputs with integrated landuse planning. The
management plan of a Tiger Reserve, therefore, needs to
be integrated in larger regional management plans.
Criticism
In an Animal Planet special named Tiger Zero, Indian
naturalist Valmik Thapar has said that for several years,
Project Tiger officials have inflated India's wild tiger
numbers so as to save their jobs. Project Tiger itself is
now being threatened by government activities like the
newly passed Tribal Bill, which allows tribal population
to reside inside designated tiger sanctuaries.
Furthermore, all the tigers in Sariska Tiger Reserve have
been poached, showing the ineffectiveness in this case of
Project Tiger. A recent article in Tehelka dwells on the
odds faced by under-equipped and under-paid forest
rangers in the fight to save the Tigers in India.
Tiger Reserves in India 
State Tiger Year of Total
Reserves Estd. Area
(km2)
Assam Kaziranga 2006 859
Assam Manas 1973-74 2840

Satellite data is being used and classified into vegetation Assam Nameri 1999-2000 344
and land use maps on a 1:50,000 scale, with digitized
data relating to contour, villages, roads, drainage,
administrative boundaries and soil. The spatial layers Arunachal Namdapha 1982-83 1985
would be attached with attribute data, viz. human Pradesh
population, livestock population, meteorological data,
agricultural information and field data pertaining to
wildlife, habitat for evolving regional protocols to
monitor tiger and its habitat. Arunachal Pakhui 1999-2000 862
Pradesh
Future activities

Conservation of tigers and their prey species faces Andhra Nagarjunsagar 1982-83 3568
challenges from the need for income, lack of awareness,
Pradesh -Srisailam
and lack of land use policy in landscapes having Tiger
Reserves. These landscapes should be viewed as a
mosaic of different land use patterns, viz, tiger
conservation and preservation, forestry, sustainable use Bihar Valmiki 1989-90 840
and development, besides socio-economic growth.

Tiger habitats exist in environments of thousands of Chhattishgarh Indravati 1982-83 2799

indigenous communities that depend on them. Therefore
we cannot view these protected areas in isolation from
the surrounding socio-economic realities and

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UNIT 10: ECOLOGICAL PRINCIPLES 

Jharkhand Palamau 1973-74 1026 Orissa Simlipal 1973-74 2750

Karnataka Bandipur 1973-74 866 Orissa Sunabeda 2008-09 856

Karnataka Nagarhole 1999-2000 643 Rajasthan Ranthambhore 1973-74 1334

(extension)

Rajasthan Sariska 1978-79 866

Karnataka Bhadra 1998-99 492

Tamil Nadu Kalakad- 1988-89 800

Kerala Periyar 1978-79 777 Mundathurai

Madhya Bandhavgarh 1993-94 1162 Uttar Pradesh Dudhwa 1987-88 811

Pradesh

Uttar Pradesh Katerniaghat 1999-2000 551

Madhya Bori-Satpura 1999-2000 1486 (extension)
Pradesh

Uttar Pradesh Pilibhit 2008-09 1089

Madhya Kanha 1973-74 1945
Pradesh
Uttaranchal Corbett 1973-74 1316

Madhya Panna 1994-95 542

Pradesh West Bengal Buxa 1982-83 759

Madhya Pench 1992-93 758 West Bengal Sunderbans 1973-74 2585

Pradesh

Madhya Ratapani 2008-09 674

Pradesh

Maharashtra Melghat 1973-74 1677

Maharashtra Pench 1992-93 257

Maharashtra Tadoba- 1993-94 620

Andhari

Maharashtra Shahayadri 2008-09 569

Mizoram Dampa 1994-95 500

122 INSTITUTE FOR ADVANCED STUDIES, JODHPUR