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Last Update: 6 December 2017 Part II

E.T - 1
It is also called ecological succession. When a new area is destroyed by some disturbances. After
sometime plants appear in these area. At 1st step some algae, lichen, fungi occur then moss & fern herbs
& grasses shrubs Tress. She area again covered by vegetation & animals also occurs in the area. So, a
bare area courted into a stable ecosystem. In doing so, the ecosystem passes from an initial less complex
stage to a mature or more complex stage i.e. at 1st biodiversity is low & then final stage.
Initial stage of succession is called pioneer stage and final stage is called climax stage and inter
mediate stages are known as shrubs stage.
Pioneer stage Seral
climax stage Ecosystem tends towards maturity and in doing so it passes

from less complex pioneer stage to more stable state which is climax stage. There is a progressive change
but not unidirectional change.
Definition :-
Ecological succession is a directional progressive change in which there is an orderly replacement of
one community and in doing so it passes from a simple pioneer stage to a complex & stable climax stage
through several intermediate seral stage. However, Margalef (1968) prefers to replace the term climax by
term more mature ecosystem.
In our biosphere most mature ecosystem are replacement by coral reef, tropical rain forest etc. So,
coral reef & tropical rain forest is the e.g. of most mature eco-system.
During eco-system various factors are involve these are biotic factor, climatic factor, physiographic
factor & geologic factor. These influence the development of ecosystem.
Classification of Succession :-
1) Depending upon the role played by above these factor or pre dominant factor succession may
be classified-
a) Biosere
b) Cliosere
c) Eosere
d) Geosere
a) Biosere :- If biotic factors are predominating in succession, i.e. organisms are responsible.
b) Cliosere :- When climate is primarily responsible for bringing about succession.
c) Eosere :- If physiogeographic factors are primarily responsible (i.e.- topograph).
d) Geosere :- If geological properties of earth are primarily bringing about the succession.
If geological properties of earth are primarily bringing about the succession.
Last 3 type succession (cliosere, eosere, geosere) are called palaeoeology because these 3 requires
considerable time (involves 100,000 years) for succession.
Biosere involve less period of time. Here biotic factor is more important.
e.g.- Land dure appears by mud
2) Depending upon their origin succession can be classified
a) Hydrosere
b) Halosere
c) Xerosere
a) Hydrosere :- If a succession is start from a fresh water body, that type of succession is called
b) Halosere :- When succession is start from a salt marsh that type of succession is called Halosere.
c) Xerosere :- When succession start from rock, soil, that type of succession is called Xerosere.
3) Now have few other kinds of succession. It can classified-
a) Primary & secondary succession.
b) Autotrophic & heterotrophic succession.
c) Autogenic & allogenic succession.
d) Unidirectional & cyclic succession.
a) Primary & secondary succession :- Primary succession are those succession which takes place in
area devoid of organism or which not previously occupy by organism and environment is
predominantly inorganic in nature. Here no living thing and have no organic matter.
e.g. Newly exposed sand dune, recent lava flow.
b) Secondary succession :- Secondary succession is the succession which takes in environment and
which previously occupied by living thing or organic matter.
e.g.- In deforested area, In harvested crop land.
Slash & burn cultivation is called shifting cultivation.
Autotrophic succession :- Autotrophic succession which is characterized by early & continued dominance
of autotrophic elements. In this succession community metabolism is autotrophic in nature. i.e. 1
P = Community production.
R = Community respiration.
Environment is inorganic in nature.
e.g.- Autotrophs succession i.e. plant succession.
Heterotrophic Succession :- In this, succession is characterized by early and continued predominance of
heterotrophic elements. It is predominantly inorganic in nature. Heterotrophic succession follows
Autotrophic community metabolism is heterotrophic in nature.
i.e. - 1
e.g. Fungi
Autogenic Succession :- In this succession internal forces is involved. It is self driven succession.
Environmental changes are brought about by biotic factor. Causal factors are located within the succession
decomposition takes place.
Allogenic succession :- Allogenic successions are those succession in which factors are located outside the
environment (i.e.- climate physiochemical properties of soil, light, rainfall, temperature). So, these allogenic
factors cause allogenic succession. Initial phase is allogenic & latter phase is autogenic succession.
Unidirectional & cyclic succession :- Most of the successions are unidirectional. 1st pioneer change & then
climax change through seral stage. But sometime it cyclically repeated.
e.g.- There are some shallow which is seasonal in nature that means every year it is drying & with it. So, its
all organism also die and after rain alive from pioneer stage it starts succession. This is the cyclic succession.

Example of Succession
Heterotrophic Succession :- Major Community situated in some minor community. These minor
Community provided substratum for their own type of succession. These substratum are predominantly
organic in nature. So, it is heterotrophic in nature. Such succession is characterized by early and continued
dominance of heterotrophic elements. Therefore these serve as an excellent example of heterotrophic
It was studied by Winston (1956). He studied the succession in a fallen Acron in the forest field. It is
Oat like fruit & the shell is hard.
Succession in Acron is start when it hangs from a tree & fruit is invaded by Acron weevil (curculio
rectus). The adult female burrows through the surface of the Acron and lays egg inside the fruit
(endosperm). From the egg larva hatches out & it starts feeding on the embryo. Sometimes though happens
that some pathogenic hole which has been created by Acron weevil & they also start consuming the embryo.
It pathogenic fungi like Penicillium & Fusarium. They destroy the embryo which is brown in color. And
such larva of weevil become stunted in growth. This feature represent the pioneer stage. These group Of
feature are followed by another group Of feature. These are fungivores and scavengers and include cheese
mite and Acron moth (Velentinia glandenella).
Cheese mite belonging to family. Tryophagus & Phyzoglypheus. These are fungivores in nature.
Acron moth belonging to Velentinia glandenella. It lay egg in the surface of the Acron or in the exit hole

created by curculso rectus. The larva of Velentinia after entering through the exit hole turns round and spins
a tough web to seal the exit hole. Cheese mites are consuming the remainder of endosperm and also feeds on
the faces of the previous occupants (Curculso rectus).
So, the pioneer stage is replaced by second group or organisms.

What happens?
The acron is attacked by cellulose and liguine consuming fungi which mostly attacked the surface of
the Acron and then invade inside also. These are another group Of organism.
These are followed by fungivores mites of the soil & collembola. They will feed on the cellulose &
liguine consuming fungi & the surface of acron.
These are followed by predatory mites belonging to genus gamasellus. They invade acron be cause
they are extremely flattened mite & enter through cracks and crevices. They consuming fungivores and
predatory mites. So, invaded group Of organisms are replaced by another group Gradually replaced.
Ultimately outer surface be comes very thin large amount of soil enter into the cavity and Acron
collapsed and then it is invaded by generalized soil fauna e.g.- Centriped, milliped, etc. so, there represents
the final stage.
This type of succession occur in an environment which is predominantly characterized by early &
continued predominance of heterotrophs. So, this is an e.g. of heterotrophic succession.

Fundamental aspects of Succession :-

1) There is an orderly and directed change of both environment and community. Both undergo
continuous reciprocal influence and adjustment.
2) Change in environment is brought about by organisms themselves. So, it is an e.g. of autotrophic
3) Exploitation of the environment makes the habitat unfavourable for the survival of pioneer organism
and instead create a favourable environment from other group of organisms.
4) Kinds of animals and plants change continuously during such process.
5) Since supply of sp is limited. Therefore succession can not go no for ever. As such succession is an
asymmetric process.
6) Succession calumniates (stop) with the establishment of relatively stable ecosystem.
7) The ecological succession is community control. The physical environment determines the pattern of
succession but does not cause it. It is caused by the biotic factor. It is a community control
8) Time require for succession depends on geography, climate, substrate, other physical factor and
species Involved.

Trends of Autogenic succession :-

Basic ecology by Odum (1983).
A) Energetics :-
1) Biomass & organic detritus increase.
2) Gross production in primary production.
3) Net production decrease.
4) Respiration increase.
5) Production/respiration i.e. ratio moves towards unity from more than one or less than 1.
1 or 1
6) ratio increases and ratio decreases where B = Biomass & P = Production.
B) Nutrient cycle :-
7) Elements cycle are increasing closed. (two pole- Biological & non biological)

8) Turn over time and storage of essential elements increase. (Time require to release of the soil to
some amount of nutrient observed by plant).
9) Cycling ratio (ratio between the recycle and through put) increases.
10) Nutrient retention and conservation increase.
C) Species and Community Structure :-
11) Species Composition Changes.
12) Diversity & Richness Increases.
13) Evenness increases. (It is the allotment of individuals in the species It increase where the
species have even number & individuals and decrease with the increasing variance.
14) R = strategist largely replaced by K-strategist (Initial phase of succession species Following r
strategies and in mature stage species Follows K strategies.
15) Life cycle increase in length and complexity.
16) Size of organism & of propagules (seeds) increases.
17) Mutalistic, symbiosis increases.
Stability :-
18) Resistance increases.
19) Resilience decreases.
Over-all strategy :-
20) Increasing efficiency of energy and nutrient utilization.
Models of succession :-
I) Pioneer concept or model.
II) Connel-strtyer model.
III) Mechanistic model.
I) Pioneer model :- The earliest model of succession was put forth by F.E. elements (1916) and this
was called pioneering model. It was the 1st model. According to elements succession consists of 6 phases.
These are -
1. Nudation
2. Migration
3. Ecesis
4. Competition
5. Reaction
6. Stabilization
Nudation :- According to Clements succession is an autogenic process and it begins with nudation. The
literal meaning of nudation is disturbance and during nudation a new bare ground is exposed.
Migration :- This is followed by migration in which propagules (the juvenile stages of plants seeds, spares)
these invade the area from other area.
Ecesis :- After migration the invading species establish in the new area and in the absence of any
competetion they start growing. This growth is called Ecesis.

4. Competition :- Ecesis is followed by competition when new individuals grow, they grow at the
expense of resource. There is competition for food, space, light in case of plant also.
5. Reaction :- As a result of competition the different species and individuals interact with each other.
This stage is called reaction.
6. Stabilization :- Reaction permit replacement of one gr of species by another group of species until a
species complex is established in the area which is a more or less stabilized in nature.
This species complex is now known as biotic community. Clements considered species complex as
some roof of a super organism. In this way organismic concept of elements was developed. But this view
was challenged by H.L. Gleason (1917) who put forth the individualistic concept of the biotic community.
He regarded biotic community as a group of species remaining similar environmental condition for their

II. Connell Statyer model :- (1977)

This model was put forth by Connell & Statyer. This concept consists of 3 different models.

1. Facilitation model
2. Inhibition model
3. Tolerance model
1. Facilitation model :- According to the 1st model process of succession is autogenic where one group
of species exploites the environment in such a way. So, as to facilitate it for the existence of another group
of organism. That is why this model is called facilitation model.
2. Inhibition model :- This model holds that different species compete for each other & same groups
inhibit the growth of other groups & therefore they became dominant in the area. So, here emphasis is given
on inhibition.
3. Tolerance model :- This is the model. According to this model all the different group of organism
invade the area simultaneously & their existence is tolerated but due to the differential rate of growth. Some
group became dominant with their early phase & other group become dominant with their early phase &
other group become dominant with their early phase & other group become dominant in the later phase. This
happens because the rate of growth in different in different species
III. Mechanistic model :- It was put forth by Tilman (1937) and Houstan & Smith (1987). According to
this model the process of species replacement consists of a array of factors which includes competition,
species specific life history traces and physiological response of plants to different factors such as light,
nutrient and moisture of soil etc. This model combines both autogenic aspects as well as the allogenic
Concept of climax :- During succession the sere develop & it culminates in the establishment of an
equilibrium or steady state with the physical & biological factors of the environment. This is called climax
& climax is stable and self replicating.
Three theories have been put forth to explain the concept of climax. These are-
1. Mono climax / climatic climax theory.
2. Polyclimax theory (Tansley, 1938)
3. Climax pattern theory (Whittaker, 1903).
1. Mono climax theory :- It was put forth by clement (1916). According to this theory the climax is
achieved as a result of the prevailing climate of the area and the climate prefers only single type of climax.
e.g.- Monsoon prefer formation of rain forest.
2. Polyclimax theory :- According to this theory the climax is the result of a mosaic of vegetation or
climaxes depending upon a numbers of factors which includes the climate of the area, topography, condition
of soil (including nutrient & moisture content) and the response of animal species
3. Climax pattern theory :- According to this theory the climax is the result of the total characteristics
of the ecosystem. This includes autogenic factors as well as allogenic factors. So, this is the combination of
the 1st two theory.

Pattern of Spatial Distribution

Distribution of animal can be described in various way. It can be discussed in term of time as
space. There are two aspects of distribution.
1. Temporal distribution Distribution is time.
2. Spatial distribution Distribution is space.
Now, there are two aspect of spatial pattern of distribution.
i) Vertical distribution :- When distribution in relation to depth is called vertical distribution.
ii) Horizontal distribution :- When distribution occur in surface of the plane is called Horizontal
Vertical distribution :- In terrestrial ecosystem the different species prefer to live in different attitude.
Within the same population there may be vertical in distribution.
e.g.- Human being

Vertical distribution is more important in aquatic ecosystem where the creatures show dial variation
in distribution. The same may be true for soil organism. They also show dial pattern of distribution with
respect to their vertical distribution.
Similarly seasonal variation is also important.
Horizontal distribution :- Main type of distribution is Horizontal spatial type of distribution. Here is micro
distribution pattern. In any area many species may be distributed into 3 basic pattern of distribution.
a) Randomly distribution.
b) Regularly distribution.
c) Contagious distribution.
But, to which, one more pattern of distribution, is
d) colonial or strongly clump distribution.
0 0 0 0 0 0 0
0 0 0 0 0 0 0
0 0 0 0 0 0 0 Regular
0 0 0 0 0 0 0
0 0 0 0 0 0 0

Randomly distribution :- Here individuals are randomly distributed. Distribution of one individual is
independent of the distribution of other individual. This is not influenced by other individual. It is possible if
the environment is uniform. But in earth environment is not uniform. Therefore this type of distribution is
more in nature. But still it is form.
e.g.- Clam (Mulinia lateralis)
which is found in intertidal zone.


Regular Distribution :- In this distribution all individuals are distributed at regular intervals that means
individuals are distributed at even distance. In order to have in regular distribution environment should be
homogeneous. This type of distribution is very rare in nature particularly in animal. In commonly seen in
man made plantation and agro-ecosystem.
Clumped distribution :- This is also called aggregated distribution. This type of distribution individuals are
clumped or aggregated together in some area where the environment condition is most favourable since
environment is not homogeneous. Here, individuals are also present in between clumped.
Colonial or strongly clumped distribution :- In colonial type of distribution individuals make some sort of
colony or individuals are strongly aggregated or clumped. There is no individuals in between colony. This
type of distribution mainly seen in sedentary animals or social animals.
How we judge the distribution?
Distribution can be solved with the help statistical analysis. In horizontal pattern of distribution to
sample and study the relationship between variance (S2) & mean ( X or m). Then me can analysis the
horizontal pattern of distribution. When-

S 2 m, then 1
same as, S 2 m, 1
S 2 m, 1
If, S2=m, that type of distribution is called random distribution.
If, S2>1, that type of distribution is clumped. If S2<1, that type of distribution is regular.
e.g.- 1, 2, 3, 4 & 5 are 5 sample and their mean have same i.e.
2=5 X 5 & S2 0
X X 1
So, random distribution
But there is a problem. For, this purpose, different type of indices are available which are clumping
index. All index of clumping are based on variance and mean ratio.
1) Fischers Co-efficient of dispersion :-
It was developed by Fischer (1960).

CD X = Number of individual
X N 1
X = mean
N = Number of sample.
This is simple ratio. If CD tends to 1 then that type of distribution is called random distribution.
If CD>1, i.e. contagious distribution, if CD<1, i.e. Negative contagious (tending towards regularity). If there
is only mean and variance, then this is absolutely regular distribution. If plot graphically by putting X in x-
axis & S2 in Y-axis.


1 2 3 4 5 6

2) For unity significance test :-

2 2 N N 1
N=Number of sample.
If taking 5 sample any value which is deviating by range 1 1.52 should be random. (-0.48 to 2.52)
is random distribution, greater than 2.52 it is contagious distribution (>2.52) and less than 0.48 is regular
distribution (<0.48). This Co-efficient is suitable only is large no of sample.
This has been modified by different author. This concept is dependent only no of individual. The
modifications are.
a) Early modification which was suggested by Neyman (1939) and it is contagious index.
S2 X
b) Index of clumping was developed by David and Moore (1954).
d 2 1 , where 2
c) Index of clumping :-
IC 1
When m=0, then O represent random pattern of distribution. Maximum regularity and this is
clumping distribution.
d) Further modified by green (1966) and this is greens index.
n 1
IC = Index of clumping
n = Number of individual.
e) L. Loyed (1967) further modified and put another index which is called mean crowding
* S2 *
X X 1 X = mean crowding
= X IC IC = index of clumping
L. Loyed compared mean & mean crowding X

When, X X , then the pattern of distribution is random.
When, X X , then the distribution is regular
X X , then the distribution is contagious
X X , then the distribution is random.
If, Value exceed 1, i.e. Contagious
Value < 1, i.e. Negatively Contagious
Value = 1, i.e. random.
S2 m
f) Hills index which was developed by Hill (1973).
n X
g) Finally Marisita index (1973) which is Id
n 1 X

It was modification of L-Loyd index According to Ludwig & Reynolds (1988) Greens index is most
suitable. But Fischers Co-efficient is most simple distribution.

Why should a species Contagiously distributed?

1) In these area where environment is favourable individuals are prefer to occupy or aggregated
2) These species which lay large number of eggs or youngs they are initially aggregated &
distributed in random manner depending upon their locomotory power.
Some important factors also govern the horizontal distribution, such as-
Availability of foods (environmental condition)
Locomotory availability.
Other mechanism of dispersion.

Co-efficient similarity :-
Diversity is of 3 types - , & .
- diversity Diversity within a habitat.
- diversity Diversity between habitats in the same region.

- diversity Diversity between two region.
In order to study degree of similarity or Co-efficient of similarity is analysed.
Earliest Co-efficient of similarity proposed by Jaccards index of similarity (1907).
Jaccards Co-efficient = cj =
ab j
cj , where,
ab j
a = number of species in the 1st habitat
b = number of species in the 2nd habitat.
J = number of species which are common to both habitats.
The index varies from 0 to 1. 1 indicating total similarity and 0 indicating no similarity. Increase in
similarity the index gradually increase.
Subsequently this is modified by Sorensens and he developed quotient of similarity.
or Cs = Sorensons quotient of similarity.
The index varies from 0 to 1. If the index exceed 0.5 then the sites are considered as similar. If the
index less than 0.5 then they are considered as dissimilar.
Thus the another modification was developed Kulezenskis Co-efficient of similarity
1 j
Ck Cs =
2 ab
Here, the index vary from 0 to 0.5.
In these 3 type of similarity most ecologist prefer the Cs. But all these indices have disadvantage &
too much emphasis given a presence or absence of similarities of species or organisms.
Therefore, puts too much emphasis on rare species where capture depends on chance. In order to
eliminate this draw back Bray & Curtis (1957) have modified the index, bringing into consideration in
abundance of species also.
2 jN
aN bN
N = number of individuals
aN = total number of individual of all species present in the 1st habitat.
bN = Total number of individual of all species present in the 2nd habitat.
jN = Sum of lesser value of individual of both habitat where both species were found.
Here index varies from 0 to 1.
0 No similar.
1 Similar.
0.5 Cut of value.
< 0.5 Dissimilar.
> 0.5 Similar.
Such studies relating to diversity may be important in biological assessment of environmental
stress. Suppose environmental stress is compared after one year of industry starting.
1. e.g.
Species Habitat (A) Habitat (B)
1 10 5
2 5 0
3 0 5
4 3 5
5 7 10
35 25

2 jN 2 5 3 10 36
CN 0.6
aN bN 35 25 60
2. e.g.- Suppose you are comparing two habitat arbitrarily.
Sample 1 2 3 4 5 6 7 8 9 Total
Habitat A *47 41 23 15 5 *2 *1 0 0 134
Habitat B 50 *28 *5 *6 *1 0 3 6 1 100

* = lesser value
Total lesser value = 88
Whether two habitats are similar or dissimilar.
a = Total number of species present in 1st habitat = 7
b = Total number of species present in 2nd habitat = 8
j = 6 species are common to both the habitat
So, Jaccards Co-efficient cj
a b j
cj 0.6
2j 2 6 12
cs 0.8
a b 7 8 15
1 1
c k c s 0.8 0.4
2 2

Inter specific association :-

Under what circumstances two species live together?
i) If they require similar environmental condition for survive.
ii) If there is a some inter specific association between two species
iii) Combination of this two.
(Inter specific association means symbiotic relationship, prey-predator relationship nutritional reamirement,
host-parasitic relation etc.)

When two species are not expected to be present together?

i) Environmental requirements are different.
ii) If there is ammensalism or antibiotic relationship. i.e. ve relationship.
How can we analysis the association of two species :-
Association between two species can be analysed with the help of index of association. Different
scientist proposed the different index, of them two are-

1. Index of association which was proposed by Whittaker & Fairbark (1959) and subsequently modified
by south wood (1966).

Index of association by Southwood is Iai

I ai 2 0.5
This based in the number of individual, not sample.
A = Total number of individual of 1st species of all the sample.
B = Total number of individual of 2nd species of all the sample.
Ji = Total number of individual of both the species in these sample in which they occur together.
e.g. A & B are two species and take 5 sample.
Number A B Ji
1 7 5 12

- 10 -
2 3 5 8
3 0 1 X
4 2 1 3
5 3 3 0
Total 15 15 23
I ai 2 0.5
15 15
The index ranges from 1 to +1
+1 stands for complete +Ve association
-1 stands for complete -Ve association
0 stands for presence of two species by chance.
According to Southwood (1966) under 4 circumstances positive association results-
1. If both the species require exactly similar environmental condition.
2. If there is some mutualism between two species
3. Both the above two points jointly acts on.
4. If there is a prey-predator relationship.
1. The 2nd method developed by Cole (1949) and it is a modified chi-square method. This method based
on the number of samples. Here + ve stands for presence of species and ve stands for absence of species
If both species are present then It is regarded as a and when both species are absent than it is regarded as
d. If species II is present and species II-I is absent it regarded as b. If species II-I is present and species II is
absent it regarded as C. So, occurrence of species I & species II is compared to a t2-table.
+ -
+ a b b+a
- c d c+d
a + c b + d a + b + c+d=x

Now + ve Maximum positive association

-ve Maximum Negative association
0 association is expected by chance.
Depending upon the situation the formula is used, the co-efficient of association (C) is calculated
If, bc ad & d a
ad bc
c (i )
a b a c
If, bc > ad and a > d
ad bc
c (ii )
b d c d
If, ad bc and cb
ad bc
c (iii )
a b b d
If, ad bc and b c
ad bc
c (iv )
a c c d
Expected value calculated by
a b a c [ T = total number = n ]
a b b d

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c d a c Degrees of freedom df = (2-1) = 1
c d b d
Formula of chi-square test is-

0 e2 [ e = expected value ]
Yets correction is needed.
Significance of deviation is judged by t2-table
Problem :- A survey of two species of deer mice. (Peromyscus leucopus & P. maniculatus). Among 192
sample P. maniculatus occur exclusively at 133 & P. leucopus is 14 and both species is 26 & none 19.
Ans :- Here,
a = 26
b = 133
c = 14
d = 19
ad bc
c , because bc>ad, a>d.
b d c d
26 19 133 14

133 19 14 19
494 1862 1368
= -0.273
152 133 5016
So, association is negative but whether is significant or not?
a b a c 26 14 26 133 33
T 192
b d a b 14 19 26 14 6.88 7
T 192

cd ac
133 19 26 133
T 192
c d b d 133 19 14 19 26
T 192

Obsertece value (0) Expect value (e) 0-e (0-e-0.8) (0-e-0.5)2

26 33 -7 56.25 1.7045
14 7 7 42.25 6.0357
133 126 7 42.25 0.3353
19 26 -7 56.25 2.1635

t 2
o e 0.52 10.239
df = 1
t 02.05 (1) 3.84
Significant P < 0.05
If, any fe < 5.
Df = 1
Yates Correction is to be applied.

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