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Paper No.

: 12 Principles of Ecology

Module : 05 Population Characteristics (Part 1)

Development Team
Principal Investigator: Prof. Neeta Sehgal
Head, Department of Zoology, University of Delhi

Paper Coordinator: Prof. D. K. Singh


Department of Zoology, University of Delhi

Content Writer: Dr. Kapinder


Kirori Mal College, University of Delhi

Content Reviewer: Prof. K. S. Rao


Department of Botany, University of Delhi

Principles of Ecology
ZOOLOGY
Population Characteristics (Part 1)
Description of Module

Subject Name ZOOLOGY

Paper Name Principles of Ecology

Module Name/Title Population Characteristics (Part 1)

Module Id 5; Population Characteristics (Part 1)

Keywords Population, population characteristics, natality, mortality,


immigration, emigration, age pyramids, sex ratio.

Contents
1. Learning Outcomes
2. Introduction
3. Population Characteristics
3.1 Population density
3.1.1 Methods of measuring population density
3.2 Natality
3.2.1 Types of natality
3.3 Mortality
3.3.1 Types of mortality
3.4 Age structure
3.4.1 Age pyramids
3.5 Sex ratio
4. Summary

Principles of Ecology
ZOOLOGY
Population Characteristics (Part 1)
1. Learning Outcomes

After studying this module, you shall be able to

 Understand the meaning of term ‘population’ and various types of population in the nature.
 Learn about characteristics of a Population
 Learn how various factors affect population density
 Evaluate how age structure will estimate the future population projection
 Understand the importance of sex ratio and its variation in different populations of different species

2. Introduction

The word population is derived from the Latin word ‘populus’ which means people. Organisms in
nature rarely grow as separate from each other. Each species in an ecosystem exist as a population.
Ecologically, a population is an assemblage of individuals of a species which potentially interbreed
and occupying a particular space at the same time. For example, grasshopper in the field, deer
population in the forest, pine trees in a forest etc. Sometimes individuals of some species live solitary
but they interact with other members of the same species or others at different times in their lives.
Clarke (1954) distinguished the population into two types.

1. Monospecific population in which group of individuals belong to the same species.


2. Polyspecific population or Mixed population in which groups of individual belongs to different
species. The term community is often used for the polyspecific population.

The populations of species have numerous local populations or subpopulations known as demes. It
can be defined as a smallest collective unit of animal population which is able to interbreed. These
individuals share a common gene pool. The gene flow between populations occurs through
emigration and immigration. The natural selection acts on individuals of a population leading to its
evolution due to change in gene frequency over a period of generations. It may change the physical
expression of organisms in the population.

3. Population Characteristics

The population ecologist Thomas park expressed that a population has several characteristics or
biological attributes which can be best expressed as statistical functions. These functions are unique to
the group and cannot be applied to the individuals in the group. Some of these properties are density,
natality, mortality, age distribution, biotic potential, dispersion, dispersal and growth forms. A brief
description of the characteristic of the population is given below.

3.1. Population density

It is defined as number of individuals in relation to a definite unit of space. It is generally expressed as


number of individuals or biomass/area or volume e.g. 100people/ square km or 4000 crustaceans/m3
of water.
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Principles of Ecology
ZOOLOGY
Population Characteristics (Part 1)
The density of the organisms in the given area can vary with the food supply, seasons and climatic
conditions but always has definite upper and lower limits. The upper limit to density is imposed by
the size of the organism and its trophic level. If the animal is smaller, greater is its abundance per unit
area. For example, a forest is able to support more wood lice than deer population. The lower limit of
density is not well defined but homeostatic mechanisms help to keep the density within limit. The
density is one of the important characteristic of population which is dependent upon a number of
factors like resource availability, productivity of the population, energy flow, utilization,
physiological stress and dispersal

The important factors responsible for change in the population density are shown in figure 1:

MORTALITY
NATALITY
POPULATION
DENSITY
IMMIGRATION EMIGRATION

Population density Population density


increases decreases

Figure 1: Factors responsible for change in population density

Population density is categorized into two types:

1. Crude density: It is defined as the total number of individuals or biomass per unit of the total
space e.g. the number of Rhinoceros living in the Kaziranga National Park.
2. Specific or ecological density: It is defined as the number of individuals or biomass per unit of
the habitat space. It is the available area or volume which is actually colonized by the population.
For example Rhinoceros do not occupy all area of Kaziranga national park but it may avoid some
of the area due to the lack of food, shelter and human habitations. Therefore the area inhabited by
the Rhinoceros actually will be its ecological density.

The difference between crude and ecological density can be explained by taking the example of a fish
density in pond (Kahl, 1964; Figure 2) The crude density of the fish in the pond goes down as the
water level drops during the summer season but the ecological density in the contracting pond
increases as the fish are crowded into smaller water area. It becomes very easy for the predatory bird
to catch fish at this time of the year as the ecological density of the fish is at its peak.

Principles of Ecology
ZOOLOGY
Population Characteristics (Part 1)
Ecological density

Crude density

Water depth

Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Jan

Figure 2: Relation between Crude density and Ecological density of fish population in pond
Months
3.1.1. Methods of measuring population density

Population density of organisms can be measured by the following methods:

1. Total counts: It is the total counts of all the organisms in a given area especially with large or
conspicuous organisms. Eg. Conducting census of human population, whales in an area of sea etc.
2. Sampling methods: In this method, a small proportion of the population is counted which is used
to estimate the total population. Sampling can be done in two ways.
i. Use of quadrate: In this method, count all the individuals on several quadrates of known
size and extrapolate the average to the whole area. The shape and size of quadrate vary with
type of organisms studied. It can be circular, triangular or square type (figure 3). The method
is extensively used for plant populations and many invertebrates.

A B C

Figure 3: Types of Quadrate; A: Circular, B: Square, C: Triangle

ii. Capture- recapture method: The simple type of population method is known as Peterson
method as it was developed by scientist C.G.J. Peterson in 1898. This method is used to count
larger animals like birds, fish etc. The animals are captured, marked and released in the
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Principles of Ecology
ZOOLOGY
Population Characteristics (Part 1)
environment. On subsequent day animals were recaptured to calculate the population density.
The population at any time will have marked and unmarked individuals. The proportion of
marked individuals in a later sample is used to determine the total population. For eg, suppose
100 fishes are captured, marked and released again into the pond. After some time again 120
fishes were captured this includes 30 marked fishes. The size of the population was counted
by the formula:

Marked animal in second sample Marked animal in first sample


=
Total caught in second sample Total population size
30/120 = 100/N
Thus total population size estimated is 400.

Sometime estimation of absolute density of a population is very difficult. So, it is simple to determine
or N= 400
the relative density. The relative density never provides an estimate of density but it is rather an index
of abundance that is more or less accurate. Relative density can be measured by the following
methods:

Trap: It is generally used in capture- recapture methods but also used to determine relative density as
the number of individuals’ caught/ day/trap.

Number of fecal pellets: By counting the number of fecal pellets in an area, index of population size
can be determined. For eg., population of rabbits, deer etc.

Vocalization frequency: By analyzing the calls of animals in particular time duration, index of
population size can be determined. The number of pheasant calls heard/15 minutes in the early
morning has been used as an index of the size of the pheasant population.

Feeding capacity: Population density can be counted by calculating the amount of food consumed by
animals before and after treatment. For eg., the amount of bait consumed by rat/mice before and after
poisoning and can be used determines an index of change in density.

3.2. Natality

It is defined as the birthrate of a population. In general, Natality covers the production of new
individuals which arises from hatching, germination or arises by fission.

3.2.1. Types of natality

Natality can be categorizes into two types:

1. Absolute or physiological natality: It is defined as the physiological ability of organisms to


produce maximum number of new individuals under optimum conditions. It is always constant for
a given population. The absolute natality is very difficult to achieved in the wild populations but it
can be observed only under favorable conditions or when major limiting factors are temporarily
non-operative, e.g. If a small number of paramecia is placed in the culture medium, under
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Principles of Ecology
ZOOLOGY
Population Characteristics (Part 1)
favorable conditions they achieved maximum natality by multiplying at maximum reproductive
rate. It is beneficial for two reasons.
a) It provides a standard for comparison with realized natality.
b) It is useful in setting up equations to calculate the rate of increase in a given population.

2. Ecological or realized natality: It is defined as increase in the population size under ecological
or actual environmental conditions. It is never constant for a population but varies with the age,
composition, size and physical environmental conditions of the population.
Natality can be expressed as: Δ Nn/ Δt = Absolute/ crude natality/birth rate
Or Δ Nn/ N Δt = specific natality/ birth rate
i.e. the number of new individuals/ unit time/ unit population
N = Total population or the reproductive part of Population
Nn = New individuals added into the population
Δt = time lapsed during change in population.

Since increase in population depends upon the number of females in a population, the age specific
birth schedule counts only females giving rise to females. The age specific schedule is obtained by
determining the mean number of females borne in each group of females. Crude natality / birth rate is
expressed in terms of population size e.g. 40 births /1000 individuals of the population. The specific
birth rate will be 4 percent. The natality rate is expressed as the number of offsprings produced per
female per unit time. The magnitude of natality rate is dependent on the type of organisms being
studied. Some species breed once in a year or some throughout year; some produce few offsprings or
some produces enormous. For eg. Fish commonly lays few thousand eggs, frog produces few
hundred, birds lays 1 to 20 eggs and mammals produces only few offsprings (between 1to10). The
birth rate is usually inversely related to parental care.

3.3. Mortality

It is defined as the rate of death of individuals in a population.

3.3.1. Types of mortality

Two types of mortality can be recognized.

1. Ecological or realized mortality: It is the actual loss of individuals under a given environmental
conditions. It is not constant for a population but varies with the environment conditions and
population such as predator, disease (Figure 4) etc.

2. Theoretical/Minimum mortality: It is the number of death of individuals under ideal or non-


limiting conditions. Therefore, even if the conditions are ideal for individuals, they would die
because of old age determined by their physiological longevity. It may be constant for the
population. Minimum mortality is always greater than ecological longevity. The organisms in
nature actually become old aged due to the process of ageing or senescence.

Principles of Ecology
ZOOLOGY
Population Characteristics (Part 1)
Figure 4: Mortality in fishes due to diseases

Mortality is the number of deaths of individuals in a population per thousand in a given period of
time. Generally mortality is expressed as either the probability of dying or as death rate. Death rate
can be defined as the number of deaths during a given interval of time divided by the average
population. For example, a population with initial size of 500 individuals in which individuals
survived at the end of the period is 400. The average size of the population for the period will be
900/2 = 450. The total number of individuals died is 100, so the death rate is 100/450 = 0.22. The
probability of death of individuals is the number that died during a given time interval divided by the
number of individuals at the beginning of the period, i.e.100/500=0.2. The complement of probability
of dying is the probability of surviving i.e. the number of survivors divided by the number alive at the
beginning of the period. As we are more interested in the number of survivors, so mortality can be
expressed in terms of life expectancy. It is the average number of years to be lived in future by the
members of a given age in the population.

3.4. Age distribution

It is the total number or percentage of individuals in a given population in different age group. The
nature of a population is influenced by its age structure. Age distribution influences natality, mortality
and the ratio of various age groups in a population which indicates its reproductive status. Age is
generally expressed in days, months or years but it also considered in other categories such as pre-
reproductive, reproductive and post reproductive. The age structure also determines the future
population projections.

1. The population with high reproductive phase individuals than post reproductive phase shows an
expanding population. However, some of the plants and animal including insects and fishes etc do
not have post reproductive phase. For example, bamboo shows mass flowering and subsequently
die after producing seeds.
2. A population has more or less even distribution of post-reproductive and reproductive
individuals showed stable population.

Principles of Ecology
ZOOLOGY
Population Characteristics (Part 1)
3. A declining population will be having large proportion of older or post reproductive individuals.

Usually a population tries to maintain a stable age distribution. The ratio of various age groups in a
population indicates its current reproductive status. Usually the ratio of young to adult in a relatively
stable population of most animals is approximately 2:1. Once a stable age distribution is reached,
changes in natality and mortality affect population stability temporarily and it tries to return to the
stable condition at the earliest.

3.4.1. Age pyramids

The age distribution in a population can be represented in the form of polygons or age pyramids
where the relative width of the successive horizontal bars represents the number of individuals or the
percentage in different age classes. There are three types of pyramids recognized (Figure 5).

1. Bell shaped pyramid having broad base represent an expanding population.


2. Dome shaped pyramid represent a moderate proportion of young and old individuals.
3. Urn shaped pyramid symbolize a declining population in which pre-reproductive individuals are
less in number.

Post-Reproductive

Reproductive

Pre-Reproductive

A B C

Figure 5: Pyramids of A) Expanding Populations (Bell Shape); B) Stable Populations (Dome Shape); C)
Declining Populations (Urn Shape)

Age distribution has practical value in the wild life management. A low ratio of pre-reproductive to
reproductive phase indicates a poor reproductive season. However, changes in age distribution alone
do not imply changes in survivorship or fecundity and it alone should not be used to predict
population trends.

3.5. Sex ratio

The individuals of a population differ themselves in a number of ways such as age, sex, breeding
condition and health. The eukaryotic organisms reproduce through sexual reproduction. There are
some organisms which have the capacity to reproduce asexually but still they have provisions for
sexual reproduction to maintain genetic variability in the population. Sex ratio can be defined as the
percentage of male and female individuals in a population.

The sex ratio can be categorized into two types:


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Principles of Ecology
ZOOLOGY
Population Characteristics (Part 1)
i) Primary sex ratio: It is the ratio of male and female at the time of conception and it tends to be
1:1.
ii) Secondary sex ratio: It is the ratio of male and female at the time of birth. This ratio often
favours males in younger age but in the older age it shifted towards females.

In the case of humans also male’s ratio is more than females at birth (52%) but as age of parent’s
increases, the ratio of females increases than males. Similarly in birds, cattle and rabbits, the ratio of
males is higher than females but in domestic chicken, sheep and horse males constitute 49% of the
population.

However, the secondary sex ratios of vertebrates show greater variation. For example, in Alaskan fur
seal one male always dominate a breeding group of 30 females and the male sex ratio exceeds 3%.
The other male adults live at the periphery of the breeding groups and suffer high mortality. In some
birds, mortality of female increases then males particularly in nesting season.

The shifting of sex ratios from 1:1 to unequal during the later age may be due to physiological and
behavioral patterns, which may affect mortality in different sexes differently. In birds, generally the
female help males in building the nest, defending the territory, incubate the eggs, brood and feed the
youngs which make female more vulnerable to predation and other dangers than males. The adult
males in mammals and adult females in birds expend more energy and are more vulnerable to
predation and this may be responsible for imbalance in the sex ratio in the older age groups.

In sexually mature animals, there is a selection process of choosing mate. This process is called
mating system. The mating system is categorized as monogamy, involves the pairing of male and
female and the relation is maintained for longer duration of time. Other mating system is promiscuity
where males and females mate with more than one of the opposite sex but does not form any relation
between them. Monogamy is most commonly found in birds and rarely among mammals. In
monogamy, both parents show cooperative behavior to successively grow their young ones. Most
species of birds are monogamous during the breeding season, because their young one are helpless
and need food, warmth, and protection. Among mammals, females produce milk for providing food to
the young. Males often can contribute little or nothing to the survival of the young, so it is to their
advantage to mate with as many females as possible.

Among many species of monogamous birds, such as bluebirds (Sialia sialis), the female or male may
“cheat” by mating with others while maintaining the reproductive relationship with the primary mate
and caring for the young to increase their chances in contributing their gene into gene pool. Polygamy
is the acquisition by an individual of two or more mates. It can involve one male and several females
or one female and several males. A pair bond exists between the individual and each mate. The
individual having multiple mates is generally not involved in caring for the young. Environmental and
behavioral conditions result in various types of polygamy. In polygyny, an individual male pairs with
more than one female. In polyandry, an individual female pairs with more than one male. Polyandry is
interesting because it is the exception rather than the rule. This system is best developed in three
groups of birds, the jacanas, phalaropes and some sandpipers. The female competes for and defends

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Principles of Ecology
ZOOLOGY
Population Characteristics (Part 1)
resources for the male. The female produces multiple clutches of eggs, each with a different male. The
male begins incubation and becomes sexually inactive (Figure 6).

Figure 6: The male African jacana showing polyandry after the female lays the eggs, the male incubates
the eggs and cares for the young while the female seeks additional mates

4. Summary

A population is an assemblage of individuals of a species which potentially interbreed and occupying


a particular space at the same time. The population has several characteristics whose functions are
unique to the group. Some of these properties are density, natality, mortality, age distribution and
growth forms.

Population density is defined as number of individuals in relation to a definite unit of space.


Population density is categorized as Crude density which means total number of individuals per unit
of the total space and Ecological density as the number of individuals per unit of the habitat space.
Natality is another important characteristic of the population which tells the production of new
individuals which arises from hatching, germination or arises by fission. It is further explained as
Absolute Natality which means the physiological ability of organisms to produce maximum number
of new individuals under optimum conditions and Ecological Natality which is defined as increase in
the population size under actual environmental conditions. It varies with the age, composition, size
and physical environmental conditions of the population.

Mortality can be explained as the rate of death of individuals in a population. Mortality can be
recognized as two types. Ecological mortality is the actual loss of individuals under a given
environmental conditions. It also varies with the environment conditions and population such as
predator, diseases etc and Minimum mortality means number of death of individuals under ideal
conditions. Therefore, the individual would die because of old age determined by their physiological
longevity and is constant for the population.

Age distribution is another component of population characteristics which includes the total number
or percentage of individuals in a given population in different age group. It explains the projection of

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Principles of Ecology
ZOOLOGY
Population Characteristics (Part 1)
future population. The age distribution influences natality, mortality and the ratio of various age
groups in a population which indicates its reproductive status. Sex ratio is the percentage of male and
female individuals in a population. The primary sex ratio is constant which indicate equal proportion
od male and female in the population but actually or in real condition the ratio vary in population of
different species. In sexually mature animals, there is a selection process of choosing mate.
Monogamy involves the pairing of male and female whereas Polygamy is the acquisition by an
individual of two or more mates. Environmental and behavioral conditions result in various types of
polygamy. In polygyny, an individual male pairs with more than one female. In polyandry, an
individual female pairs with more than one male.

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Principles of Ecology
ZOOLOGY
Population Characteristics (Part 1)

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