Bio 313 PDF

COURSE
GUIDE
BIO 313
ANIMAL ECOLOGY
Course Team O.A. Olajuyigbe -Adeyemi Colledge of

Education,
Miss F.C. Olakolu - Nigerian Institute for
Oceanography and Marine Research,
Mrs H.O. Omogoriola -Nigerian Institute for
Oceanography and Marine Research ((Team
Writers )
Mrs. Ajetomobi (Editor) - Lagos State Polytechnic
Dr.Maureen N. Chukwu (Course Coordinator) -
NOUN
Dr. Esenowo Imeh kokoete (Course Reviewer)-
University of Uyo
NATIONAL OPEN UNIVERSITY OF NIGERIA

BIO 313 COURSE GUIDE
© 2023 by NOUN Press

National Open University of Nigeria
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All rights reserved. No part of this book may be reproduced, in any form
or by any means, without permission in writing from the publisher.
Printed 2022, Reprinted 2023
ISBN: 978-978-58-055-7
Reviewed 2022
ii
CONTENTS
Introduction………………………………………………….. iv
Course competencies…………………………………..……. iv
Course objectives………………………………………….…. iv
Working through this course…………………………………. iv
Study units…………………………………………………... iv
References and further readings……………………………… v
Presentation schedule………………………………………... v
Assessment…………………………………………………... vi
How to get the most from the course………………………… vi
Online facilitation……………………………………………. vii
iii
INTRODUCTION
Animal Ecology (BIO 313) is a 2 credit-hours course for undergraduate

students of Biology that deals with the historical concept of ecology,
ecology of local and aquatic animals. The other contents are growth rate,
age structure of animal population, Natality and Mortality, Survivorship
Curves, Life Tables and K-Factor Analysis, Competition, The Natural
Regulation of Animal Numbers, Population Cycles, Dynamics of
Predator-Prey Systems, Ecology of African Bats and Behavioural
Ecology.
COURSE COMPETENCIES
This course aims to provide the basic knowledge in Ecology to enable the
sudent know/understand the relationship between animals in their
ecosystem and with the environment
COURSE OBJECTIVES
The Comprehensive Objectives of the Course are to;
1. Explain the historical background of animal ecology,

2. Explain the basic fundamentals of ecology and its components,
3. Explain the various relationships influencing the ecological
community,
4. State and explain the different types of ecosystem components to
include the levels of energy flow in ecosystem and ways by which
population can be changed
WORKING THROUGH THIS COURSE
To successfully complete this course, you are required to read each study
unit, read the textbooks and other materials provided by the National
Open University.
Reading the reference materials can also be of great assistance. Each unit
has self –assessment exercise which you are advised to do.
There will be a final examination at the end of the course. The course
should take you about 8 weeks to complete.
This course guide provides you with all the components of the course,
how to go about studying and how you should allocate your time to each
unit so as to finish on time and successfully
iv
STUDY UNITS
The course is divided into 3 modules and study units in this course are
given below:
Module 1 Introduction to Animal Ecology
Unit 1 Historical Background of Animal Ecology

Unit 2 Fundamentals of Ecology
Unit 3 Interactions in Animals
Unit 4 Ecology of Aquatic Animals
Unit 5 Ecology of Terrestrial Animals
Module 2 Ecology of Animal Population
Unit 1 Growth rate and Age structure of animal population

Unit 2 Factors affecting Animal Population
Unit 3 Measurement of Population dynamics
Unit 4 Life tables and K-values
Unit 5 Key-Factor Analysis
Module 3 Population Cycle in Animal Ecology
Unit 1 Competition in Animal Ecology

Unit 2 Population Cycle
Unit 3 Population Cycles in a Predator-Prey System
Unit 4 Ecology of African Bat
Unit 5 Behavioural Ecology
REFERENCES AND FURTHER READINGS
You would be required to read the recommended references and

textbooks in each unit of the course materials.
PRESENTATION SCHEDULE
There is a time-table prepared for the early and timely completion and
submissions of your TMAs as well as attending the tutorial classes. You
are required to submit all your assignments at the stipulated date and time.
v
ASSESSMENT
There are three aspects to the assessment of this course. The first one is
the in-text questions and the second is self-assessment exercises, while
the third is the written examination or the examination to be taken at the
end of the course. Review the exercises or activities in the unit by
applying the information and knowledge you acquired during the course.
The work submitted to your tutor for assessment will account for 30% of
your total work. At the end of this course you will have to sit for a final
or end of course examination of about a two hour duration and this will
account for 70% of your total course mark.
HOW TO GET THE MOST FROM THE COURSE
In this course, you have the course units and a course guide. The course
guide will tell you briefly what the course is all about. It is a general
overview of the course materials you will be using and how to use those
materials. It also helps you to allocate the appropriate time to each unit so
that you can successfully complete the course within the stipulated time
limit.
The course guide also helps you to know how to go about your in-text
questions and Self-assessment questions which will form part of your
overall assessment at the end of the course. Also, there will be tutorial
classes that are related to this course, where you can interact with your
facilitators and other students. Please I encourage you to attend these
tutorial classes.
This course exposes you to Animal Ecology, a sub-discipline and very

interesting field of Biological Science.
ONLINE FACILITATION
Eight weeks are provided for tutorials for this course. You will be notified
of the dates, times and location for these tutorial classes.
As soon as you are allocated a tutorial group, the name and phone number
of your facilitator will be given to you.
The duties of your facilitator is to monitor your progress and provide any
necessary assistance you need.
Do not delay to contact your facilitator by telephone or e-mail for

necessary assistance if
• You do not understand any part of the study in the course material.
vi
• You have difficulty with the self-assessment activities.

• You have a problem or question with an assignment or with the
grading of the assignment.
It is important and necessary you attend the tutorial classes because this
is the only chance to have face to face contact with your facilitator and to
ask questions which will be answered instantly. It is also a period where
you can point out any problem encountered in the course of your study.
Course Information
Course Code: BIO 313
Course Title: ANIMAL ECOLOGY
Credit Unit: 2
Course Status: ELECTIVE
Course Blub: This course provides students with the basic knowledge in
Ecology to enable them understand the relationship between animals in
their ecosystem and with the environment
Semester: FIRST SEMESTER
Course Duration: 13 WEEKS
Required Hours for Study : 65 hours
Ice Breaker
Dr. Esenowo, Imeh Kokoete is a Senior Lecturer of Ecology and

Environmental Biology in the Department of Animal and Environmental
Biology, University of Uyo. Dr. Esenowo moderate and facilitate courses
in the National Open University. He has supervised student projects and
seminar review in the Department of Biology, Faculty of Science.
Dr. Esenowo research interests are physico-chemical aspects of
freshwater and terrestrial ecosystem, fish biology and environmental
toxicology.
vii
MAIN
COURSE
CONTENTS
Module 1 Introduction to Animal Ecology ……………… 1
Unit 1 Historical Background of Animal Ecology ……… 1

Unit 2 Fundamentals of Ecology……………………….. 9
Unit 3 Interactions in Animals…………………………… 14
Unit 4 Ecology of Aquatic Animals……………………… 24
Unit 5 Ecology of Terrestrial Animals………………… 36
Module 2 Ecology of Animal Population……………….. 46
Unit 1 Growth rate and Age structure of animal

population …………………………………….. 46
Unit 2 Factors affecting Animal Population………….. 58
Unit 3 Measurement of Population dynamics ……….. 71
Unit 4 Life tables and K-values ………………………. 80
Unit 5 Key-Factor Analysis ……………………….. 94
Module 3 Population Cycle in Animal Ecology …… 107
Unit 1 Competition in Animal Ecology …………. 107

Unit 2 Population Cycle…………………………… 119
Unit 3 Population Cycles in a Predator-Prey System.
Unit 4 Ecology of African Bat…………………….. 124
Unit 5 Behavioural Ecology ……………………….. 132
BIO 313 ANIMAL ECOLOGY
Module 1 Introduction to Animal Ecology
Module Introduction
In Module One, unit one deals with the history and current understanding
of Animal Ecology and how organisms and its environment relate and
influence one another in their various ecosystems. You are taught about
the fundamentals of ecology; interaction in animals and ecology of
aquatic and terrestrial animals.

Unit 4 Ecology of Aquatic animal
Unit 5 Ecology of Terrestrial animal
Glossary
Contents
1.1 Introduction
1.2 Intended Learning Outcomes (ILOs)
1.3 Historical background of Ecology
1.4 Summary
1.5 References/Further Readings/Web Sources
1.6 Possible Answers to Self-Assessment Exercises
1.1 Introduction
Ecology (from Greek: οἶκος,"house"; -λογία, "study of") is the study

of living organisms relating with each other and their surroundings.
Ecosystems can be defined as a web, community, or network of
individuals that arrange into a self-organized and complex hierarchy of
pattern and process. Ecosystems create a biophysical feedback between
living (biotic) and non-living (abiotic) components of an environment
that generates and regulates the biogeochemical cycles of the planet.
Ecosystems provide goods and services that sustain human societies
and general well-being. Ecosystems are sustained by biodiversity
within them Biodiversity is the full-scale of life and its processes,
including genes, species and ecosystems forming lineages that
1
integrate into a complex and regenerative spatial arrangement of types,

forms, and interactions.
Ecology is a sub-discipline of biology, the study of life. The word

"ecology" ("oekologie") was coined in 1866 by the German scientist
Ernst Haeckel (1834–1919). Haeckel was a zoologist, artist, writer, and
later in life a professor of comparative anatomy. Ancient philosophers
of Greece, including Hippocrates and Aristotle, were among the
earliest to record notes and observations on the natural history of plants
and animals; the early rudiments of modern ecology. Modern ecology
mostly branched out of natural history, science that flourished in the
late 19th century. Charles Darwin's evolutionary treatise and the
concept of adaptation as it was introduced in 1859 is a pivotal
cornerstone in modern ecological theory.
Ecology is not synonymous with environment, environmentalism,

natural history or environmental science. Ecology is closely related to
the biological disciplines of physiology, evolution, genetics and
behaviour. An understanding of how biodiversity affects ecological
function is an important focus area in ecological studies. Ecosystems
sustain every life supporting function on the planet, including climate
regulation, water filtration, soil formation (pedogenesis), food, fibers,
medicines, erosion control, and many other natural features of
historical, spiritual or scientific value. Ecologists seek to explain:
1. life processes and adaptations

2. distribution and abundance of organisms
3. the movement of materials and energy through living
communities
4. the successional development of ecosystems. and
5. the abundance and distribution of biodiversity in context of the
environment.
There are many practical applications of ecology in conservation

biology, wetland management, natural resource management
(agriculture, forestry, fisheries), city planning (urban ecology),
community health, economics, basic and applied science, and it
provides a conceptual framework for understanding and researching
human social interaction (human ecology).
2
At the end of this unit, the student should be able to
 explain the historical background of animal ecology and the

various contributions of scientists to the development of
animal ecology
1.3 Historical Background of Ecology
Studies of animal distribution began in the nineteenth century, but the

formal development of animal ecology did not occur until the 1920s.
British zoologist Charles Elton, whose field research emphasized the
study of populations in the wild, was perhaps the most influential
figure. Elton's work, often involving northern fur-bearing animals of
commercial value, made a number of concepts part of the naturalist's
vocabulary, including the ecological niche, the food chain, and the
pyramid of numbers, that is, the decrease in numbers of individual
organisms, or total quantity (weight) of organisms, at each successive
stage in a food chain, from plants and plant-eating animals at the
bottom to large carnivores at the top. Just as with plant ecology, diverse
schools of animal ecology emerged in Europe and the United States
during the first half of the twentieth century. Some schools, like Elton's,
focused on empirical studies of predator-prey interactions and
population fluctuations, others focused on animal community
organization, still others on broader patterns of distribution and
abundance.
Although some of the early work in animal ecology, particularly in the

United States, attempted to model itself on plant ecology, by the 1930s
animal ecology had emerged as an independent field. There was little
overlap or interaction between the work of animal and plant ecologists.
Effective impetus for an integrated perspective in ecology came from
work in aquatic biology, best exemplified in the late nineteenth century
by Karl Möbius's studies of the depleted oyster bank off Germany's
north coast and the pioneering limnological (freshwater) studies of
François Alphonse Forel on Swiss lakes. This work was continued and
refined in the early twentieth century by many researchers, including
August Thienemann in Germany and Einar Naumann in Sweden.
Möbius's concept of the “biocenosis,” the integrated community
consisting of all living beings associated with a given habitat or a
particular set of environmental conditions, was adopted widely by
German and Russian ecologists in the 1920s and 1930s. An integrative
3
perspective also emerged in soil science, as in Sergei Winogradsky's

turn-of-the-century studies of soil microbiology, and in studies of
biogeochemical cycles, as in the work of Russian geochemist Vladímir
Vernadsky, who introduced the term “biosphere” in 1914. However,
the integrative concept that had the broadest appeal and played a central
role in bringing together the many different strands of ecological
science was that of the “ecosystem,” introduced by British botanist
Arthur G. Tansley in 1935 but first used effectively in an aquatic
setting.
Tansley was Britain's foremost plant ecologist and the founder in 1913
of the British Ecological Society, the first such national organization,
formed two years earlier than its American counterpart. A pioneer in
vegetation surveys, a critic of Clements's idea of the climax
community, a passionate conservationist, and a student of Sigmund
Freud, Tansley brought his broad experience and erudition to bear on
the problem of identifying the ideal ecological unit of study. He
suggested that the term “ecosystem” captured this concept best without
implying any mysterious vital properties. The new term received its
fullest early treatment in a seminal paper published in 1942 by a young
American limnologist, Raymond Lindeman. Making use of the concept
of ecological succession, Elton's pyramid of numbers and food chains,
earlier studies of energy flow in aquatic systems, and Clements's notion
of the stable climax community, Lindeman traced the flow of energy
through the different trophic (feeding) levels (producers, primary
consumers, secondary consumers) in a small Minnesota pond as a way
to mapping its structure as an ecosystem and to demonstrate its
progress in development toward a stable, equilibrium state.
World War II proved to be a watershed for ecology. Although earlier

preoccupations with community classification and structure,
population dynamics, and patterns of distribution continued in the post-
war years, newer methodologies, practices, and conceptual schemes
took hold, and ecology as a science and a profession grew in size,
status, and organization. In the post-war period, Lindeman's ground
breaking work on ecosystem ecology found a home among biologists
funded by the U.S. Atomic Energy Commission, who used
radionuclides to trace the flow of materials and energy through natural
ecosystems. Ecosystem research soon expanded from its base in the
Atomic Energy Commission. It also prospered among a small group of
Tansley's followers at the new Nature Conservancy in Britain. It
became an essential feature of modern ecological science, a message
conveyed to several generations of students worldwide through the
successive editions of Eugene P. Odum's Introduction to Ecology, first
published in 1953. Meanwhile, the pre-war synthesis of Darwinian
natural selection theory with Mendelian genetics resulted in the gradual
4
post-war emergence of a more strongly Darwinian perspective in

population and community ecology.
The post-war years also saw a shift toward quantitative aspects of

ecology. Mathematical techniques developed in the United States,
Europe, and the Soviet Union during the interwar period joined with
war-born techniques involving information systems and cybernetics to
produce a movement toward mathematical modelling and computer
simulation of populations, communities, and ecosystems. Much of this
modelling and its techniques came under attack during the last decades
of the twentieth century. Some ecologists abandoned model building
for empirical studies, others worked on refining and improving the
models, and many called into question the underlying notions of
stability and equilibrium upon which most of the models were based.
The devastation brought by World War II also contributed to greater post-

war interest in the conservation of natural resources, protection of
wildlife, and preservation of natural environments, a trend that, when
linked in the 1960s with social criticism, blossomed into an international
environmental movement that drew heavily upon concepts and theories
of ecology. As had occurred before the war in a more limited way among
a few visionaries, ecology now came to be widely viewed not only as the
source of remedies for environmental ills but also as the scientific
underpinning for a new social order. This proved to be a mixed blessing
for ecologists. On the one hand, funding for ecological research increased
considerably, and many more people were drawn into the field. On the
other hand, the theoretical framework of ecological science, being neither
unified nor consistent, could not provide easy, unambiguous solutions to
environmental problems, let alone unified and consistent social visions.
Toward the end of the twentieth century, this disagreement and
uncertainty among ecologists was used as fuel in legislative and legal
debates arguing against the protection of endangered species and the
maintenance of pristine nature reserves. This situation encouraged the
further refinement and integration of ecological science toward the
incorporation of human disturbance and the notion of managed
ecosystems.The Studies of animal distribution began in which century?
Who is the founder of British Ecological

Society?
Self-Assessment Exercises 1
Attempt these exercises to measure what you have learnt so far.
This should not take you more than 5 minutes.
1. The British zoologist Charles Elton works involves many
concepts which include what type of animal?
2. How has post-World War II impacted on the ecosystem?
5
1.4 Summary
You have been introduced to the field of animal ecology including

some of its most fundamentals; early history, and current understanding
of its development. Animal Ecology asks questions about how
organisms and its environment relate and influence one another in their
various ecosystems.
Donald Worster, Nature's Economy: A History of Ecological Ideas, 2d

ed. (1994).
Eugene Cittadino, Nature as the Laboratory: Darwinian Plant Ecology

in the German Empire, 1880–1900 (1990).
Leslie A. Real and James H. Brown, eds., Foundations of Ecology

(1991).
Gregg Mitman, The State of Nature: Ecology, Community, and

American Social Thought, 1900–1950 (1992).
Michael Shortland, ed., Science and Nature: Essays in the History of

the Environmental Sciences (1993).
Robert P. McIntosh,The Background of Ecology (1985).
Sharon Kingsland, Modeling Nature: Episodes in the History of

Population Ecology, 2d ed. (1995).
Stephen Bocking, Ecologists and Environmental Politics: A History of

Contemporary Ecology (1997).
Pascal Acoted., The European Origins of Scientific Ecology (1800–

1901), 2 vols. (1998).
https://www.bing.com/search?q=Historical+Background+of+Anima
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https://www.bing.com/videos/search?q=Historical+Background+of+
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1.6 Possible Answers to SAEs
Answers to SAEs 1
1. Elton's work, often involving northern fur-bearing animals of

commercial value, made a number of concepts part of the
naturalist's vocabulary, including the ecological niche, the food
chain, and the pyramid of numbers, that is, the decrease in numbers
of individual organisms, or total quantity (weight) of organisms, at
each successive stage in a food chain, from plants and plant-eating
animals at the bottom to large carnivores at the top.
2. The devastation brought by World War II also contributed to

greater post-war interest in the conservation of natural resources,
protection of wildlife, and preservation of natural environments, a
trend that, when linked in the 1960s with social criticism,
blossomed into an international environmental movement that
drew heavily upon concepts and theories of ecology.
8
Contents
2.1 Introduction
2.3 Ecological Community
2.3.1 Composition and Diversity
2.3.2 Habitat and Ecological Niche
2.4 Summary
2.1 Introduction
Fundamentally, ecology is like a society, while ecologists are like

economists that investigate the economy of nature instead of human. We
will be talking about ecology in three basic scales after the introductory
section: population, community, and ecosystem. They are stacked and
connected one by one, each acts as the building block of the superior
system, until finally we arrive at the global level, with the most dynamic,
diversified, and ultimate living things system on the planet, the biosphere
At the end of this unit, the student should be able to;
 Explain the basic fundamentals of ecology and its components

 Explain the various relationships influencing the ecological
community
2.3 Ecological Community
A community is comprised of all the various populations interacting in

an area. An example of a community is a coral reef where numerous
populations of fishes, crustacea and corals exist and interact. Ecologists
try to know at this level how different relationships like predation and
competition are influencing the organization and evolution of a
community.
9
In-Text Question(s)
What is Community?
Answer: A community is comprised of all the various populations
interacting in an area.
2.3.1 Composition and Diversity
Communities distinguish from each other by two characteristics:

composition and diversity. The composition of a community is simply a
listing of the various species in the community. The diversity digs deeper
than mere composition in that it involves both species richness (the
number of species) as well as evenness (the relative abundance of
different species).
Figure 1: Community Individualistic Model (www. econguru.com)
A species' spreading range is based on its tolerance for such abiotic

factors in the environment as temperature, light, water availability,
salinity, and so forth. To determine a species' tolerance range, we plot
the species' ability to survive and reproduce under a particular gradient
of environmental conditions, resulting a bell-shaped curve. If we put
together several species' range in one graph, a community is formed
because their tolerance ranges simply overlap.
However, other models exist such as interactive model to explain

community and most ecologists supported it for many years, because it
bases its hypothesis not only on species' responses to abiotic factors but
also biotic factors. Search the web if you want to learn more about it.How
can communities be distinguish from each other?
10
Attempt this exercises to measure what you have learnt so far. This
should not take you more than 5 minutes.
1. Differentiate between habitat and Niche?
2. 3. 2 Habitat and Ecological Niche
Just as Elton John sings the Circle of Life, each species plays its role in a
community, eats or be eaten, lives or let live. They occupy particular
positions both in a spatial sense (where to live) and a functional sense
(what is the part). A habitat is an environment wherein an organism lives
and reproduces, while the ecological niche is the functional role the
organism plays in its community, including its habitat as well as the
interactions with other organisms. Niche includes everything e.g.
resources an organism needs to meet its energy, nutrient, and survival
demands) and every aspects of the way(e.g. the environmental features it
needs to hunt and to escape successfully) an organism live with the
environment, since it's difficult to delve into one niche completely, most
observations concentrate on certain aspects of it.
Since an organism's niche is affected by abiotic factors (such as climate

and habitat) and biotic factors (such as competitors, parasites, and
predators) simultaneously, usually two types of niches are looked at
separately by ecologist, the fundamental niches and the realized niches.
The fundamental niche of an organism comprises all conditions where
under, it can potentially survive and reproduce; the realized niche is the
set of conditions where under, it exists in nature.What is a Habitat?
Attempt this exercise to measure what you have learnt so far. This
should not take you more than 5 minutes
What is Diversity?
2.4 Summary
You are taught about the fundamentals of ecology; Composition and

diversity, habitat and ecological niche.
11
Begon, M., J. L. Harper and C. R. Townsend, 1990. Ecology -

Individuals, Populations and Communities, Blackwell Scientific
Publ., London, UK, 2nd edition. Chapter 4.
Chapman, J. L. and Reiss, M. J. (2010), Ecology, Principles and Applications,

2nd ed. Cambridge University Press.
Fundamental of ecology, author: EconGuru, introduction to

ecosystems and community. Ecological population; Chapter 1
to chapter 3.
Miller, G. T. and Spoolman, S. E. (2009), Essentials of Ecology, 5th ed.

Brooks/Cole, Cengage Learning.
Miller, G. T. Jr. (1988), Environmental Science, 2nd ed. Wadsworth,

Belmont.
Odum, E. P. and Barrett, G. W. (2004), Fundamentals of Ecology, 5th ed.

Cengage Learning
http://gdcganderbal.edu.in/Files/a8029a93-30ad-4933-a19a-
59136f648471/Link/EcologyandEnvironment_44344ff9-021a-
4e6b-ab1f-cf8a148398f2.pdf
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12
Answers to SAEs 1
1. A habitat is an environment wherein an organism lives and
reproduces, while the ecological niche is the functional role the
organism plays in its community, including its habitat as well as
the interactions with other organisms. Niche includes everything
e.g. resources an organism needs to meet its energy, nutrient, and
survival demands) and every aspects of the way(e.g. the
environmental features it needs to hunt and to escape successfully)
an organism live with the environment, since it's difficult to delve
into one niche completely, most observations concentrate on
certain aspects of it.
Answers to SAEs 2
1. Diversity involves both species richness (the number of species) as
well as evenness (the relative abundance of different species).
13
Contents
3.1 Introduction
3.3 Interactions
3.3.1 Competition among Populations
3.3.2 The Competition Eclusive Principle
3.3.3 Predator – Prey Interactions
3.3.4 Symbiotic Relationships
3.4 Summary
3.1 Introduction
Animals interact with each other in numerous, complex ways. Of the

various types of interactions between species, most involve resources and
consumers. We shall be considering the different types of interaction that
exist in nature.
At the end of this unit, the student should be able to;

 Explain the various relationships influencing the ecological
community
 Explain the different types of interaction in animals
3.3 Interactions
A resource, in ecological terms, is something (such as food, water, habitat,

sunlight, or prey) that is required by an organism to perform a vital
function such as growth or reproduction. A consumer is an organism that
consumes a resource (such as predators, herbivores, or detritivores). Most
interactions between animals involve one or more competitor species
vying for a resource.
Competition for resources, predator-prey, parasite-host, and other

types of interactions integrate species into a system of dynamic
interacting populations.
14
Table 1: Showing the different types of Interaction among animals

Species Interactions
Type of Interaction Expected Outcome
Competition - - Decrease in both species
Predation + - Predator increases, prey decreases
Symbiotic
Relationships
Parasitism + - Parasite increases, host decreases
Commensalism + 0 One increases, the other not affected
Mutualism + + Increase in both species
Source: (www. econguru.com)
As indicated in the above table, competition for limited resources between

two species has a negative effect on the population abundances of both
species. In predation and parasitism, the abundances of predator and
parasite are expected to increase at the expense of that of prey and host,
since predators feed on prey and the parasites obtain nutrients from the
host. In commensalism one species is benefited whereas the other is not
harmed. In mutualism, two species help one another and both species are
benefited. List the different types of interaction in nature
3.3.1 Competition among Populations
Interspecific competition occurs when members of different species try to

utilize the same resource like light, space, or nutrients that is in limited
supply, or when their niches overlap. If it is unlimited, no competition
would have been triggered. Competition leads to several possible
outcomes and one of them, is the extinction of one of the competitors.
Competition can cause species to evolve differences in traits. This occurs

because the individuals of a species with traits similar to competing
species always experience strong interspecific competition. These
individuals have less reproduction and survival than individuals with traits
that differ from their competitors. Consequently, they will not contribute
many offspring to future generations. For example, the finches previously
15
discussed can be found alone or together on the Galapagos Islands. Both

species' populations actually have more individuals with intermediate-
sized beaks when they live on islands without the other species present.
However, when both species are present on the same island, competition
is intense between individuals that have intermediate-sized beaks of both
species because they all require intermediate sized seeds. Consequently,
individuals with small and large beaks have greater survival and
reproduction on these islands than individuals with intermediate-sized
beaks.
Studies show that when G. fortis and G. fuliginosa are present on the same
island, G. fuliginosa tends to evolve a small beak and G. fortis tends to
evolve a large beak. The observation that competing species' traits are
more different when they live in the same area than when competing
species live in different areas is called character displacement. For the two
finch species, beak size was displaced: Beaks became smaller in one
species and larger in the other species. Studies of character displacement
are important because they provide evidence that competition plays a very
important role in determining ecological and evolutionary patterns in
nature.
What is interspecific competition?
Self-Assemeement Exercises 1
1. Can competition cause species to evolve differences in
traits?
3.3.2 The Competition Exclusive Principle
To explain how species coexist, in 1934 G. F. Gause proposed the

competitive exclusion principle: species cannot coexist if they have the
same niche. The word "niche" refers to a species' requirements for
survival and reproduction. These requirements include both resources
(like food) and proper habitat conditions (like temperature, pH). Gause
reasoned that if two species had identical niches (required identical
resources and habitats) they would attempt to live in the exact same area
and would compete for the exact same resources. If this happened, the
species that was the best competitor would always exclude its competitors
from that area. Therefore, species must at least have slightly different
niches in order to coexist.
Peter Grant and colleagues tested Gause's principle by studying seed-
eating finches (birds) that live on the Galapagos Islands in the Pacific
Ocean. They found that different finch species can coexist if they have
traits that allow them to specialize on particular resources. For example,
16
two finch species, Geospiza fuliginosa and Geospiza fortis, vary in a key
trait: beak size. Beak size is a critical trait because it determines the size
of a seed that a finch can eat: Individuals with small beaks eat small seeds,
individuals with intermediate sized beaks can eat intermediate size seeds
and individuals with large beaks can eat large seeds. G. fuliginosa and G.
fortis do compete for intermediate sized seeds because each species has
some individuals with intermediate sized beaks. However, G.
fuliginosa specializes upon smaller seeds because it has more individuals
with small beaks. Conversely, G. fortis specializes upon larger seeds
because it has more individuals with large beaks. Thus, these species
niches differ slightly because a specific trait, beak size, allows them to
specialize upon a particular seed size.
Joe Connell also tested Gause's principle by studying barnacles (shelled
marine organisms) that live on rocks along European coastlines. In 1961,
Connell found that two barnacle species, Balanus and Chthamalus, can
coexist because they differ in two traits: growth rate and vulnerability
to desiccation. Balanus 's growth is rapid, which allows it to smother and
crush the slower-growing Chthamalus. Balanus, however, dies close to
shore because it gets too dry during low tide. In
contrast, Chthamalus tolerates these dry conditions. Consequently, even
though Balanus is a better competitor for space, these barnacles coexist
because Chthamalus can survive in areas that Balanus cannot survive.
These and many other examples support the competitive exclusion
principle: Species can only coexist if they have different niches
Figure 1: Competition to Extinction (Source: www.econguru.com)
17
When grown alone in pure culture, both A and B exhibit a somewhat

logistic growth pattern, expanding their colonies rapidly till reaching the
carrying capacity of the environment. However, when the two species
come to mixed culture together, A is the better competitor who drives B
out in the end.
You might wonder, now that competitions lead to extinctions, why the
world is still filled with myriads of living things that share the resources.
Good question, but you are partly right, and partly wrong. Extinction is
not the only result of competition, two species can both survive the
competition, but they have to change, or more technically, their niches
have to adapt. In the light of competitive exclusion principle that no two
species can concurrently occupy the same niche, either one of the species
die out or both shift their niches. One of the embodiment of niche shift,
or niche partitioning is resource partitioning. Resource partitioning
decreases competition between two species, and it is more observable
than other subtle forms of niche partitioning. Example of Resource
Partitioning are when three species of ground finches of the Galapagos
Islands occur on isolated islands, their bills tend to be the same
intermediate size, enabling them to feed on a wider range of seeds.
Where they co-occur, selection has favoured divergence in beak size
because the size of the beak affects the kinds of seeds that can be eaten.
In other words, competition has led to resource partitioning.
The tendency for characteristics to be more divergent when populations

belong to the same community than when they live separately is termed
character displacement. And it is often used as evidence for competition
and how resource partitioning have taken place. Who proposed the
competition exclusive principle?
1. What was Gause reasoned on the Competition Exclusive
Principle?
3.3.3 Predator-Prey Interactions
In predation, one organism, called the predator feeds on another,

called the prey. With common sense, there should be no dispute on that
the relationship between lion and zebra is that of predation. But what
is the relationship that herbivorous deer feeds on trees and bushes? This
might be a little bit surprising, but in a broader sense, predaceous
18
consumers include not only animals but also herbivores that feed on
plants.
By observation, we have revealed the interacting pattern between the

populations of predator and prey, cycles of fluctuation that one drives
the other and vice versa. Population of the prey increases as that of the
predator decreases, since fewer prey are being eaten. At the carrying
capacity of the environment, the number of prey reaches its summit
and stops growing. The predators now are provided with plenty of prey
to feed on, so the population increases as that of prey decreases. Again,
the predators' increased number overconsume the prey, as the prey
population declines, so does the prey population. See the graph below.
Figure 2: Predator-Prey Population Dynamics (source:

www.econguru.com)
Notice that the predator population is smaller than that of prey, and that
it fluctuates lagging behind the prey also.
Most predator-prey population cycles are like what we have discussed,

probably with more elaboration of the dynamics and curves. However,
their interactions involve more than just population cycles, other
behaviours like prey defences might also interest you. If so, do check the
picked links below. What is Predator- prey Interaction?
3.3.4 Symbiotic Relationships
A symbiotic relationship, or symbiosis is one in which members of two

populations interact very closely. As indicated by the figure 3 in section
3.1.3, three types of symbiotic relationships exist and by the way at least
one species benefits from such a relationship while the other is harmed
or unaffected or benefited.
Parasitism resembles predation in that an organism called a parasite
derives nourishment from another called the host (just as the predator
19
derives nourishment from its prey), though parasites also take hosts as
habitats and springboards to transmit themselves to other hosts.
Parasites appear in all kingdoms of life. Some of the frequently heard of

parasites include viruses (e.g., HIV), bacteria (e.g., strep infection),
protists (e.g., malaria), fungi (e.g., rusts and smuts), plants (e.g.,
mistletoe) and animals (e.g., leeches).
Commensalism is a symbiotic relationship wherein one species is

benefited and the other is neither benefited nor harmed. Well known
instances are those in which one species provides a habitat or a means of
transportation for the other.
1. Example of Commensalism
Animalia: Barnacles attach themselves to the backs of whales and the

shells of horseshoe crabs to get a free home and ticket for transportation.
Remoras are fishes that attach themselves to the bellies of sharks by
means of modified dorsal fin acting as a suction cup.
Plantae: Epiphytes grow in branches of tree in order to receive light, but

not to take nourishment from the trees. Instead, their roots obtain
nutrients and water from the air.
Mutualism is a symbiotic relationship in which both species benefit. In

many cases, mutualistic relationships help organisms obtain food or
avoid predation. As with parasitism, it is possible to find examples of
mutualism in all kingdoms.
2. Example of Mutualism
Human and Bacteria: Human cannot synthesize vitamins by

themselves, but can benefit from some bacteria residing in their intestinal
tract that make vitamins. Meanwhile, bacteria are provided with food.
Termites and Protozoa: Termites rely on the protozoa in their intestinal

tract to digest wood.
To sum up, symbiotic relationships do occur between species, but the

three patterns we provided may be too simple to embrace all the natural
forms of symbiosis. We were just skimming roughly. Many other
derivative forms of symbiosis are developed, look for other materials if
you are interested. What is Symbiosis?
20
1. List examples of mutualism in animals?
3.4 Summary
You are taught about interactions, competition among populations,

predator-prey. Interactions, Symbiotic Relationships

Connell, J. H., 1970. A predator-prey system in the marine intertidal

region. I. Balanus glandula and several predatory species of
Thais. Ecological Monographs 40: 49-78.
Chapman, J. L. and Reiss, M. J. (2010), Ecology, Principles and Applications,

2nd ed. Cambridge University Press.
Fundamental of ecology, author: EconGuru, introduction to ecosystems

and community. Ecological population; Chapter 1 to chapter 3.
Miller, G. T. and Spoolman, S. E. (2009), Essentials of Ecology, 5th ed.
Brooks/Cole, Cengage Learning.
Miller, G. T. Jr. (1988), Environmental Science, 2nd ed. Wadsworth,

Belmont.
Odum, E. P. and Barrett, G. W. (2004), Fundamentals of Ecology, 5th

ed. Cengage Learning
http://gdcganderbal.edu.in/Files/a8029a93-30ad-4933-a19a-
59136f648471/Link/EcologyandEnvironment_44344ff9-021a-
4e6b-ab1f-cf8a148398f2.pdf
https://www.bing.com/search?q=fundametal+of+Ecology+&FORM=H
DRSC1#
21
https://www.bing.com/videos/search?q=fundametal+of+Ecology+&&v
iew=detail&mid=C50BC92935E3D16219C7C50BC92935E3D1
6219C7&&FORM=VRDGAR&ru=%2Fvideos%2Fsearch%3Fq
%3Dfundametal%2Bof%2BEcology%2B%26FORM%3DHDR
SC3
https://www.bing.com/videos/search?q=fundametal+of+Ecology+&&v
iew=detail&mid=D4FE9EFC126F94339E64D4FE9EFC126F94339E6
4&&FORM=VRDGAR&ru=%2Fvideos%2Fsearch%3Fq%3Dfundame
tal%2Bof%2BEcology%2B%26FORM%3DHDRSC3
https://study.com/academy/lesson/symbiotic-relationships-mutualism-
commensalism-amensalism.html
22
Answer to SAEs 1
1. Competition can cause species to evolve differences in traits. This

occurs because the individuals of a species with traits similar to
competing species always experience strong interspecific
competition. These individuals have less reproduction and survival
than individuals with traits that differ from their competitors.
Competition can cause species to evolve differences in traits. This
occurs because the individuals of a species with traits similar to
competing species always experience strong interspecific
competition. These individuals have less reproduction and survival
than individuals with traits that differ from their competitors.
Answer to SAEs 2
1. Gause reasoned that if two species had identical niches (required
identical resources and habitats) they would attempt to live in the
exact same area and would compete for the exact same resources.
If this happened, the species that was the best competitor would
always exclude its competitors from that area. Therefore, species
must at least have slightly different niches in order to coexist.
Answer to SAEs 3
1. Answer: Human and Bacteria: Human cannot synthesize vitamins
by themselves, but can benefit from some bacteria residing in their
intestinal tract that make vitamins. Meanwhile, bacteria are
provided with food.
Termites and Protozoa: Termites rely on the protozoa in their intestinal

tract to digest wood
23
Unit 4 Ecology of Aquatic Animals
Contents
4.1 Introduction
4.3 Ecologya and Biology of Aquatic species
4.3.1 Clupeidae
4.3.2 Carangidae
4.3.3 Polyynemidae
4.3.4 Sciaenidae
1.3.5 Sparidae
4.3.6 Penaeid shrimps
4.4 Summary
4.1 Introduction
Animal ecology is an important area of study for scientists. It is the study

of animals and how they relate to each other as well as how they relate
to their environment. There are various forms of animal ecology. By
studying this information, you learn more about what makes these
animals prosper or what potentially holds them back. In animal ecology,
there are many factors that is currently threatening the existence of these
animals which are caused by human activities.
Perhaps the best examples are in the water. A look at areas like lakes,
coastlines and even marine life will show you just how much human
environmental damage has hurt these animals. Animal ecology has
changed drastically in recent time. Here are some examples of how the
environment and human interaction has changed the scope of many
animals.
1. Animal habitats in many marine areas have ceased to exist. Coral
reefs and other very delicate ecosystems have been harmed by
human presence.
2. In the arctic regions, melting ice has limited the lifespan of polar
bears, which make the ice their home. Additionally, sea lions and
other marine life that use the ice to rest on have been unable to do
so.
3. Dams and other waterway changes have hurt animal ecology
throughout the country. Animals are no longer able to get to the
source of water they need.
24
4. Deforestation in jungles and other habitats has caused many of the

only locations for animals to live to be wiped away. Urban city
development has also pushed animals farther and farther out of
their natural habitats.
There are many other ways that animal ecology has changed. The goal
of scientists is to find out what is happening and why it is happening that
way. It is often very much a worry when animal species are dying or are
unable to evolve naturally because of the drastic changes in their
lifestyles and living areas. Through study of animal ecology, scientists
hope to understand better what really is happening and what effect it will
have both in the short and in the long term.
At the end of this unit, the student should be able to explain the ecology
of different fish species
4.3 Ecology and Biology of Aquatic Animals
An aquatic animal is any animal whether invertebrate or vertebrate that

lives in water for most or all of its lifetime. Many insects such as
mosquitoes, mayflies, dragonflies and caddisflies have aquatic larvae,
with winged adults. Aquatic animals may breathe air or extract oxygen
from water through specialised organs called gills or directly through the
skin.
The term aquatic can be applied to animals that live in either fresh
water or salt water. However, the adjective marine is most commonly
used for animals that live in saltwater, i.e. in oceans, seas, etc.
Aquatic animals (especially freshwater animals) are often of special
concern to conservationists because of the fragility of their environments.
Aquatic animals are subject to pressure from overfishing, destructive
fishing, marine pollution and climate change. Many habitats are at risk
which puts aquatic animals at risk as well. Aquatic animals play an
important role in the world. The biodiversity of aquatic animals provide
food, energy, and even jobs.
Fresh water creates a hypotonic environment for aquatic organisms. This
is problematic for some organisms with pervious skins or
with gill membranes, whose cell membranes may burst if excess water is
not excreted. Some protists accomplish this using contractile vacuoles,
while freshwater fish excrete excess water via the kidney. Although most
25
aquatic organisms have a limited ability to regulate their osmotic balance

and therefore can only live within a narrow range of
salinity, diadromous fish have the ability to migrate between fresh water
and saline water bodies. During these migrations they undergo changes to
adapt to the surroundings of the changed salinities; these processes are
hormonally controlled. The eel (Anguilla anguilla) uses
the hormone prolactin, while in salmon (Salmo salar) the
hormone cortisol plays a key role during this process.
Freshwater molluscs include freshwater snails and freshwater bivalves.
Freshwater crustaceans include freshwater crabs and crayfish.
In addition to water breathing animals, e.g., fish, most mollusks etc., the
term "aquatic animal" can be applied to air-breathing aquatic or sea
mammals such as those in the orders Cetacea (whales) and Sirenia (sea
cows), which cannot survive on land, as well as the pinnipeds (true seals,
eared seals, and the walrus). The term "aquatic mammal" is also applied
to four-footed mammals like the river otter (Lontra canadensis) and
beavers (family Castoridae), although these are technically amphibious
or semiaquatic. There are up to one million types of aquatic animals and
aquatic species.
Amphibians, like frogs (the order Anura), while requiring water, are
separated into their own environmental classification. The majority of
amphibians (class Amphibia) have an aquatic larval stage, like a tadpole,
but then live as terrestrial adults, and may return to the water to mate.
Certain fish also evolved to breathe air to survive oxygen-deprived water,
such as Arapaima (family Osteoglossidae) and walking catfish.
Most mollusks have gills, while some fresh water ones have
a lung instead (e.g. Planorbidae) and some amphibious ones have both
(e.g. Ampullariidae). Many species of aquatic animals lack a backbone or
are invertebrates.
Aquatic animals play an important role for the environment as well as
human's daily usage. The importance of aquatic animals comes from the
fact that they are organisms that provide humans with sources such as
medicine, food, energy shelter, and raw materials that are used for daily
life.
Each aquatic species plays a different role to help us make every day
easier, healthier, and also more productive. They also help with
the atmospheric pressure and global climate change. What is Aquatic
animal?
26
1. What kind of condition does freshwater create for aquatic
organism?
4.3.1 Clupeidae
Most clupeid species are marine, but some are anadromous (shads) and
Ethmalosa fimbriata (bonga) are adopted to withstand low salinities
particularly in the rainy season.
a. Bonga (Ethmalosa fimbriata)

Bonga is the most important clupeid species in the coastal inshore waters
of Nigeria. This species rarely goes below 20 m. It is more euryhaline
than the flat sardinella and it is found in estuaries, the sea, lagoons and
also in places that are liable to have great variations in salinity. It prefers
warm and turbid waters. Because of these ecological preferences, it tends
to replace the flat sardinella, and even more clearly the round sardinella,
in those sectors without upwelling but with strong surface desalination.
Its biology and migrations seem small in extent and limited to estuaries
and the adjacent coastal areas (Longhurst, 1960).
Ethmalosa is a non-selective filter-feeder subsisting mainly on large

diatoms and phytoplankton. The species migrates into and out of the
estuaries following seasonal changes in salinity as well as with the
abundance of plankton in the estuaries during the dry season. Ethmalosa
tends to be more abundant in Nigerian estuaries during the period
October-April. Its migration is possibly due to spawning and feeding.
Juveniles are definitely more abundant in rivers and in estuaries, while
young spawners and adults can be found both in estuaries and at sea. This
pelagic fish is a target species for the artisanal gillnet and beach seine
fisheries.
Fig 1: Bonga (Ethmalosa fimbriata)
b.Shad (Iisha africana)

27
Shad is an anadromous clupeid in habiting inshore waters, sand beaches

and estuaries (in almost all fresh waters). Ilisha africana has a maximum
length (L∞) of about 22 cm and it has a good preference for crustacea and
small fishes (juveniles). It may be caught at the surface or near the bottom
down to about 25 m. Hence it can be a target species for beach seine,
gillnet, purse seine and inshore trawl fisheries.
c. Sardine (Sardinella spp)

The flat sardinella is found from Mauritania to Angola. It is coastal fish,
more euryhaline, most often found to be abundant near the outlet of water
courses. It prefers warmer waters with a temperature above 24°C and
seems to avoid waters that are not clear. It is not very abundant in areas
without upwelling where the warm and low saline superficial layer is
permanently present as in the Baight of Biafra and a large portion of the
Nigerian shelf.
In Nigeria Sardinella spp. are caught by canoe fishermen using ringnets,

castnets, gillnets, beach seines and also by trawlers. But Sardinella is not
a target species of any of the main fisheries. What is another name for
Bonga fish?
1. Write a little ecological notes on Sardine?
4.3.2 Carangidae
The following carangid species are fairly abundant in Nigerian waters:

Caranx spp., Chloroscombrus chrysurus, Decapterus rhonchus amd
Trachurus spp. There are mostly schooling species distributed on the
continental shelf but some occur in brackish waters especially when
young.
a. Various jacks (Caranx spp.)
Caranx spp. have wide distribution along the West African coast from
Senegal to Angola. Some species inhabit inshore waters and estuaries and
the others are located in deeper waters (over 100–m depth). Hence, this
fish group can be vulnerable to both artisanal and industrial fleets. Caranx
spp. feed mainly on fish but also on shrimps, some crabs and
invertebrates. This fish species group is caught in pelagic and bottom
trawls, seines, set and ring gillnets and sometimes on line gear.
28
Fig 2: Various jacks (Caranx spp.)
b. Atlantic pumber (Chloroscombrus chrysurus)

Chloroscombrus chrysurus occurs along the West African coast from
Mauritania to Angola. This schooling pelagic species inhabits the
Nigerian continental shelf at depths of 10–50 m. It also occurs in estuaries
and the mangrove fringed lagoons and brackishwater areas. Its juveniles
are sometimes located offshore in association with jellyfish. Atlantic
bumper can be a target species of the artisanal fleet using set gillnets and
seines as well as for the industrial fleets using trawls and operating in
waters of 10– 50 m depth.
c. False scad (Decapterus rhonchus)

This is a schooling carangid species inhabiting near bottom waters, mostly
between 30 m and 50 m but can be located in waters over 200 m depth. It
feeds on small fish and invertebrates. This species is mostly exploited by
industrial fleets using trawls, but it can also be fished by artisanal
motorized canoes using gillnets.
d. Horse mackerel (Trachurus spp.)

Horse mackerel occurs in schools in sandy bottom localities and usually
at 100–200 m depth. Since the main fishing grounds are on the continental
shelf, the species is not normally caught by artisanal fishermen. It is
usually a target species of the offshore trawl and purse seine fisheries and
sometimes it can be caught with longlines. It appears that the Nigerian
industrial fisheries can exploit Trachurus capensis (Cape horse mackerel)
and Trachurus trecae (Cunene horse mackerel).
What is another name for Trachurus spp?
4.3.3 Polynemidae
a. Lesser African threadfin (Galeoides decadactylus)

Galeoides decadactylus does not appear to penetrate below the
thermolcline. It occurs in inshore waters adjacent to sandy beaches. The
species is known to develop female gonads by passage through a
29
nonfunctional hermaphroditic stage arising from a normal male.

Understanding its reproductive and recruitment strategy appear to be vital
in the managing of this fish species.
Galeoides prefers silty and sand-silty bottoms. It is a semi-diadromous

fish with spawning migration into estuaries and lower reaches of rivers. It
feeds on benthic organisms such as crustacea and polychaetes. It is a
target species for the artisanal fishery using gillnets and beach seines as
well as the industrial fleets employing trawls in the inshore areas.
Fig 3: Lesser Africana threadfin (Galeoides decadatylus)
b. Royal threadfin (Pentanemus quinquarius)

Pentanemus quinquarius has a normal reproductive cycle. It occurs on
sandy bottoms down to a depth of 50 m. It is caught by the artisanal gillnet
fishery on nearshore sandy bottoms but the species is also harvested
offshore by the industrial fleet using trawls. Additionally Pentanemus can
be caught with beach seines.
c. Giant African threadfin (Polydactylus quadrifilis)

The giant African threadfin (Polydactylus quadrifilis) can grow up to
lengths 150–200 cm. The species inhabits inshore and offshore sandy
bottoms up to a depth of 50 m. It also occurs in estuaries and lagoons
fringed by mangrove. This fish species is jointly harvested by the artisanal
fishermen and industrial fleets. Its attractive size has made it extremely
vulnerable to gillnet and beach seine fisheries.
The giant African threadfin (Polydactylus quadrifilis) can grow up to

what size?
4.3.4 Sciaenidae
The croakers and drums are the important sciaenid species in Nigeria.
This fish species group is primarily marine but also occurs seasonally in
brackishwater areas. Most of the species inhabit sandy and muddy
bottoms in coastal areas with large river flows.
30
a. Bobo croaker (Pseudotolithus elongates)

Pseudotolithus elongatus prefers surroundings that are less saline. In
fact, commercial concentrations correspond to the great estuaries in the
gulf of guinea where the species can be caught in large quantities in
certain seasons.
They inhabit mud bottoms in coastal waters up to 50-m depth but also
enter estuaries and coastal lagoons. This species, with maximum length
of about 45 cm, moves further offshore to spawn during the rainy season.
P. elongatus is jointly harvested by the artisanal and industrial fleets. It
can be caught with bottom trawls, setnets, beach seines and longlines.
Fig 4: Bobo croaker (Pseudotolithus elongatus)
b. Longneck croaker (Pseudotolithus typus)

Pseudotolithus typus grows to a larger size than P. elongatus. It attains
a maximum length (L∞) of 100 cm and fish of 50-cm length are
common in the catch. The main fishing ground for this species is from
the Gulf of Guinea to the Congo. It is the most important commercial
sciaenid species in Nigeria.
It inhabits mud and sandy bottoms up to a depth of 150 m but it is more

abundant in waters of less than 60 m and temperature above 18°C. It
also occurs in estuaries. Hence, it is fished by artisanal and industrial
fleets using bottom trawls, bottom set nets and longlines.
c. Boe drum (Pteroscion peli)

Pteroscion peli occurs along the west coast of Africa, from Senegal to
Angola. It inhabits mud and sandy-mud bottoms in coastal waters
extending to 200-m depth. But it is most common in waters of less than
50-m depth. This species is more accessible to the industrial fisheries
using trawls and hook on line than to the artisanal fisheries using
gillnets and beach seines. What is another name for Bobo croaker?
31
1. Write a little ecological notes on Longneck croaker?
4.3.5 Sparidae
The seabreams occur in fairly deep waters of the continental shelf and
off the slope. The small young individuals do occur in shallow waters
but mostly at a depth greater than 15 m, forming aggregations. The adult
seabreams are more solitary. The most common species are Dentex
angolensis and Pagellus bellottii.
a. Angola dentex (Dentex angolensis)
Dentex angolensis occurs along the West African coast from Morocco
33°N to Angola. It inhabits various bottoms on the continental shelf and
the slope from about 15 m to about 300-m depth. It is a protogynic
hermaphrodite with most individuals beginning as females and changing
to males at a length 18–23 cm.
They are known to occur in Nigerian waters but the species is not an
important element of the artisanal fisheries. It is caught by the trawl
fishery but separate statistics are not available. Angola dentex is a
carnivorous species feeding on crustacea, small fish, molluscs and other
invertebrates. It can be caught in bottom trawls, bottom setnets and
longlines.
Fig 5: Angola dentex (Dentex angolensis)
b. Red pandora (Pagellus bellottii)

The geographical distribution of P. bellottii extends from the straits of
Gibraltar to Angola and also around the Canary Islands. It is a protogynic
hermaphrodite (the majority of individual are first females), then become
32
males. Red pandora is omnivorous with a predominantly carnivorous

diet consisting of crustacea, cephalopods, small fish and worms.
This is one of the most abundant sparid species in the CECAF area but
it is not a target species of artisanal fisheries in Nigeria. It is possibly
caught by the trawl fishery but separate catch data are not reported. What
is another name for Angola dentex?
4.3.6 Penaeid shrimps
Three commercially important penaeid shrimps occur in Nigerian

waters. Penaeus notialis (the pink shrimp) is by far the most dominant
species. It occurs in the lagoons, estuaries, creeks and open sea.
Parapenaeopsis atlantica (Guinea shrimp) is also fairly abundant in the
open sea depth 10–16 m. The estuarine white shrimp (Palaemon
hastatus) occurring in brackish waters and open sea is mainly exploited
by artisanal fishermen.
The coastal penaeid shrimps have interesting recruitment features. The

first phase in the life of coastal penaeid shrimps takes place at sea
between three weeks, and one month and thereafter in the coastal zones,
in bays, estuaries, in mangrove swamps which are rich in food, or in
submerged vegetation. As their development progresses the shrimps
migrate toward greater and greater depths. When the areas of distribution
of juveniles and adults are clearly separated geographically, a true
migration seaward occurs after which spawning takes place.
Since the types of exploitation (and the operational zones of the various
gears) are extremely diversified, there are in fact several successive
recruitment phases: when the shrimps leave the nursery edges and
become accessible to artisanal fisheries; when they reach the large bays
where they are accessible to small trawlers; during migration, when they
are caught by fixed nets; when they reach the sea and are caught by
industrial trawlers.
The entry process into the different fisheries is associated with the
development stage of the shrimps. If recruitment is defined as the
probability of a shrimp of a given size to be found in the fishing area this
probability can be expressed for shrimps of each size as the percentage
of shrimps at that size, in the total population that is present in that area.
If the percentages are plotted against size a recruitment curve will be
obtained. What is another name for Penaeus notialis?
33
4.4 Summary
The ecology of animals integrate the organisms (animal) and their

environment dependently, animals inhabit community and interact
with the biotic and abiotic factors in the environment to survive. Fish
species have their favourable environmental condition at their best
active. Some of these is discuss in this this unit.

Longhurst, 1960. A summary of the food of West African demersal fish.
Bull.Inst.Fondam.Afr.Noire (A Sci.Nat.), 22:276–82
National Marine Fisheries Service and U.S Fish and Wildlife

Service.(1998). Recovery Plan for U.S Pacific Populations of the
Olive Ridley Turtle (Lepidochelys olivacea). National Marine
Fisheries Service, Silver Spring, MD.
Barbour, Roger, Ernst, Carl, & Jeffrey Lovich. (1994). Turtles of the
United States and Canada. Washington, DC: Smithsonian
Institution Press.
Department of the Environment, Water, Heritage and the Arts (2010).
Lepidochelys olivacea in Species Profile and Threats Database,
Department of the Environment, Water, Heritage and the Arts,
Canberra. Available from: http://www.environment.gov.au/sprat.
Accessed 20 Apr 2010.
https://byjus.com/biology/scientific-name-of-bear/
https://www.monkeyworlds.com/
https://thetinyphant.com/terrestrial-animals/
https://www.bing.com/videos/search?q=what+is+terrestrial+animals&
&view=detail&mid=CFD2D59C5AF7A73BF04DCFD2D59C5
AF7A73BF04D&&FORM=VRDGAR&ru=%2Fvideos%2Fsear
ch%3Fq%3Dwhat%2Bis%2Bterrestrial%2Banimals%26FORM
%3DHDRSC3
https://www.bing.com/videos/search?q=Terrestrial+and+Aquatic&&vi
ew=detail&mid=9AC4FC5D4310CDE831239AC4FC5D4310C
DE83123&&FORM=VRDGAR&ru=%2Fvideos%2Fsearch%3F
q%3DTerrestrial%2Band%2BAquatic%26FORM%3DRESTAB
34
Answers to SAEs 1
1. Fresh water creates a hypotonic environment for aquatic
organisms. This is problematic for some organisms with pervious
skins or with gill membranes, whose cell membranes may burst if
excess water is not excreted. Some protists accomplish this
using contractile vacuoles, while freshwater fish excrete excess
water via the kidney. Although most aquatic organisms have a
limited ability to regulate their osmotic balance and therefore can
only live within a narrow range of salinity, diadromous fish have
the ability to migrate between fresh water and saline water bodies.
Answers to SAEs 2
1. The flat sardinella is found from Mauritania to Angola. It is coastal
fish, more euryhaline, most often found to be abundant near the
outlet of water courses. It prefers warmer waters with a
temperature above 24°C and seems to avoid waters that are not
clear. It is not very abundant in areas without upwelling where the
warm and low saline superficial layer is permanently present as in
the Baight of Biafra and a large portion of the Nigerian shelf.
Answers to SAEs 3
1. Pseudotolithus typus grows and attains a maximum length (L∞) of
100 cm and 50-cm in length. The main fishing ground for this
species is from the Gulf of Guinea to the Congo. It is the most
important commercial sciaenid species in Nigeria. It inhabits mud
and sandy bottoms up to a depth of 150 m but it is more abundant
in waters of less than 60 m and temperature above 18°C. It also
occurs in estuaries. Hence, it is fished by artisanal and industrial
fleets using bottom trawls, bottom set nets and longlines
35
Unit 5 Ecology of Terrestrial Animals
Contents
5.1 Introduction
5.3 Ecology and Biology of Terrestrial species
5.3.1 Terestrial Animals
5.3.2 Terrestrial Birds
5.3.3 Aerial Animals
5.3.4 Arboreal Animals
5.3.5 Amphibious Animals
5.4 Summary
5.1 Introduction
Animal ecology is an important area of study for scientists. It is the study

of animals and how they relate to each other as well as how they relate to
their environment. There are various forms of animal ecology. By
studying this information, you learn more about what makes these animals
prosper or what potentially holds them back. In animal ecology, there are
many factors that is currently threatening the existence of these animals
which are caused by human activities.
At the end of this unit, the student should be able to explain the ecology
of different terrestrial species
5.3 Ecology and Biology of Terrestrial species
The word “Terrestrial,” which means earthly, is borrowed from the Latin
word ‘terra,’ meaning earth, land, or ground, with the suffix -al. The
terrestrial animals are the animals that live predominantly or entirely on
land, and they also grow and reproduce on land. Having successfully
adapted to dry environments, animals belonging to this habitat have
completely abandoned their need for an aquatic phase in their lifespan.
36
Sometimes it’s hard to differentiate or judge whether an animal is

“terrestrial” or “aquatic” since there are no universally accepted criteria
to decide. Many animals considered terrestrial have a lifecycle, which is
partly dependent on being in the water, just like in the case of penguins,
seals, and walruses who sleep on land, but feed on the ocean and are yet
considered terrestrial. Even some species of crabs are also considered
terrestrial species. What are Terrestrial animal?
5.3.1 Terrestrial Animals
i.) Lions (Panthera leo)
The lion (Panthera leo) is a large cat of the genus Panthera native to
Africa and India. Deep-chested build, tiny round ears, a fuller mane, and
a heavy furry tuft at the ledge are the main physical characteristics of a
lion. The fur of lions varies in color from light buff to silver-gray,
yellowish red, and dark brown.
The males have a prominent mane, typically brownish and tinged with
yellow, rust, and black hair that grows downward and backward, covering
most of the head, neck, shoulders, and muscular chest. The average head
to body length of a male lion is about 190-210 cm, whereas the female
ones have a length of 160-189 cm.
Lions are deadly predators who have a short and powerful attack, and they
catch their prey with a fast rush and final leap. Lions usually hunt zebra,
giraffe, African buffalo, and blue wildebeest.
They are mostly active at night, and they prefer habitat consisting of
grassland, savanna, dense scrub, open woodland. Lions are found in sub-
Saharan Africa, and a small population of Asiatic lions is found in India’s
Gir forest.
Fig 1: Lions (Panthera leo)
The lion (Panthera leo) are native to which locality?
37
ii.) Dogs (Canis sp.)
The dog or domestic dog (Canis familiaris or Canis lupus familiaris) is a

domesticated descendant of the wolf which is characterized by an
upturning tail. The dog is derived from an ancient extinct wolf, and the
modern wolf is the dog's nearest living relative. These adorable fuzzballs
are one of the most ubiquitous domestic animals in the world. The
anatomy of dogs differs from breed to breed; they have powerful muscles,
a cardiovascular system supporting sprinting and endurance, and teeth to
capture, hold and tear.
They regulate their body temperature through panting and sweating via
their paws. They have an incredible hearing ability – the frequency range
of dog hearing is between 16-40 Hz and up to 45-60 Hz. The amazing part
is the number of smell-sensitive receptors is forty times more than
humans.
How do dog regulate their body temperature?
iii.) Monkeys
Monkey is a common name that may refer to most mammals of the

infraorder Simiiformes, also known as the simians. Monkeys are
mammals; some monkeys live on the ground, and some inhabit the trees.
Many small monkeys are kept as exotic pets for their adorable look, a
common example being pygmy marmosets.
Their bodies are designed to provide them both strength and agility, which
makes them flexible and fast. Monkeys are extremely intelligent and
social creatures since socialization is important for them to survive in their
environment. There are more than 350 species of primates found all over
from the habitats in Asia, Africa, and South America. What is the function
of their body?
iv.) Giraffes (Giraffa sp.)
Giraffes are regarded as the tallest living terrestrial animals, with long
necks, legs, and a unique coat pattern that serves as camouflage.
The interesting feature in their body is the ossicones that are usually
mistaken as horns. Their diet consists of leaves, fruits, and flowers of
woody plants like acacia. They have a long purple colored Prehensile
tongue of about 45cm in length. The average height of a Giraffe is around
5m; they are mostly found in the sub-Saharan region of Africa. What is
an interesting feature in body of Giraffes?
38
v.) Hippopotamus (Hippopotamus amphibious)
Deriving their name from the Greek word, translating to ‘river horse’
hippos are one of the heaviest terrestrial animals on earth. These creatures
are the heaviest animals on the land, next to elephants. Although they are
also termed as ‘semi-aquatic,’ they are generally leagued into the
terrestrial category.
These mammals are mostly herbivorous with a lifespan of 40-50 years,

and although they are exaggeratingly heavy and fleshy, they can run at a
speed of 48 km/hour on the land. Seemingly serene and calm, these 8000
pounds creatures shouldn’t be messed with because they are just as
dangerous as a wild elephant that’s gone frenzy
What is another common name for Hippos?
vi.) Bears (Ursus sp.)
Bears are the large solitary, stocky mammals equipped with

nonretractable claws, shaggy fur and are about 8 feet long and weigh
about 60-1600 pounds. They are extensively found in America, Europe,
and Asia.
They can hibernate, which is an effective survival solution. Bears are

great swimmers, which allows them to survive in near the coldest of
oceans. Despite the bulky build that bears possess, they are incredible
climbers and adept runners.
Where are bears found?
vii.) Snakes
Snakes are limbless reptiles with long, cylindrical bodies, scaly skin,
lidless eyes, and a forked tongue. Most species are not poisonous, some
are mildly poisonous, and others produce a deadly poison. The term
venom is commonly used to describe the poison produced by a snake.
Although controversial, snakes that have adapted to both land and water
are majorly considered terrestrial animals. These infamous reptiles known
for their toxic venoms or constrictions around their prey’s body are found
all over the world except for a few parts like New Zealand, Antarctic, and
Ireland, etc.
All snakes are carnivores (meat-eaters) and cold blooded, meaning their
body temperature is determined by the environment rather than being
internally regulated. For this reason, snakes are found mainly in tropical
and temperate regions, and are absent in cold climate zones. The 2,700
species of snakes fall into four superfamilies: Boidae (boas, anacondas,
and pythons), Elapidae (cobras, coral snakes, mambas, and kraits),
39
Colubridae (king snakes, water snakes, garter snakes, black snakes, and
adders, to name only a few) and Viperidae (true vipers and pit vipers).
Snakes have extremely poor sight and hearing. They detect their prey
primarily by means of vibrations, heat, and chemical signals they detect
with their other senses. For example, a snake's flicking, forked tongue acts
as a chemical collector, drawing chemical "smells" into the mouth. Those
smells are then analyzed by two chemical sensors known as Jacobson's
organs on the roof of its mouth. This mechanism also allows male snakes
to detect females in the reproductive state. The legless carnivorous
reptiles that come in absolutely mesmerizing patterns and colours from
jet black to light green are ectothermic creatures. What is the feeding
habits of snake?
Attempt this exercises to measure what you have learnt so

far. This should not take you more than 5 minutes.
1. List some characteristics features of snake?
5.3.2 Terrestrial Birds
Terrestrial birds are those that forage, roost, and predominantly spend a
significant portion of their lives on the ground. When these types of birds
are threatened, then they don’t fly away; rather, they freeze, walk or run.
Some of these birds have a unique cumbersome flight style, which is less
suited for long flights, and when flying, they generally stay low above the
ground while the others are flightless. Examples of terrestrial birds are;
The Barred Buttonquail, Female Black Francolin, The Cheer Pheasant,
The Chestnut-bellied Sandgrouse, The Eurasian Thick-knee, The
Ferruginous Partridge, The Grey Francolin, The Indian Courser, The Kalij
Pheasant, The Masai Ostrich. What are terrestrial birds?
5.3.3 Aerial Animals
Animals that thrive when airborne, accomplishing other activities like

feeding, drinking, and bathing, spending much of their life in flight are
called Aerial Animals. There are several adaptations that were adapted by
these animals to fly swiftly in the air.
These adaptations include their lightweight skeletons and slender,

streamlined bodies, which increase the flight efficiency and make lengthy
flights easier for them, the long and pointed wings serve to aid their flight
agility, and their long tail helps them to steer while flying in the air easily.
40
Since these animals have an amazing flight style, aeronautics and

mechanical engineers are studying their flying patterns and developing
myriad flying vehicles and machines from commercial aircraft to remote-
controlled drones. Examples are; Albatross, Vultures, Butterflies, Moth,
Sugar Gliders, Hummingbird, Geese, Bats, Gliding lizard, Eagles. What
are Aerial birds?
5.3.4 Arboreal Animals
The arboreal animals are the tree-living animals who spend most part of
their life cycle on trees and branches, mainly in the rainforest. These
animals have amazing adaptations that not only help them to live and
move about in trees but also help them to survive and hide in the
environment.
To deal with the mechanical challenges of moving through their habitats,

the arboreal animals have elongated limbs, which help them to cross gaps,
testing the firmness of support ahead, and to reach food and other
resources.
They use their prehensile tails to grasp the branches of trees. Animals like
spider monkeys and crested geckos have an adhesive pad at the tip of the
tail to provide increased friction.
Their small body size provides them with advantages like increased
stability, lower mass, controlled gliding, and gives them the ability to
move through more cluttered habitat amidst branches, stems, leaves, etc.
the following are example; Sloths, Koalas, Beckos, Tree Snakes,
Possums, Orangutans, Parrots, Woodpeckers, Chameleons, Red Pandas.
What are Arboreal Animals?
1. Give examples of arboreal animals.
5.3.5 Amphibious Animals
To live a double life – that succinctly describes Amphibians, in a nutshell.

The amphibious animals are cold-blooded vertebrate animals who are
born in water and breathe through gills in their larva stage, and as they
grow adult, their lungs develop the ability to breathe air, allowing them to
live on both land and water.
41
They have skin glands that help to transport water, oxygen, and carbon
dioxide into or out of them, producing useful proteins. They are also
frequently used as a defense to fight against bacteria or fungal infections.
The toxic amphibians are mostly bright coloured to warn the potential
predators from coming in close vicinity. Their skin needs special
environmental conditions in order to allow them to survive, and too much
exposure to the sun can damage their skin cells.
Wind can also dry their skin and dehydrate them, which is why most
amphibians are nocturnal, surfacing, and preying only in the dark. The
amphibians inhabit a variety of habitats, including terrestrial, fossorial,
freshwater ecosystems. Examples are; American Bullfrogs, Geckos,
Salamanders, Toads, Newts, Tortoise, Alligator Newts, Worms, Axolotls,
Green Tea Frogs. What are amphibious animals?
Attempt this exercises to measure what you have learnt so far.

1. Where are amphibious animals found and list examples
5.4 Summary
The ecology of animals integrate the organisms (animal) and their

environment dependently, animals inhabit community and interact with
the biotic and abiotic factors in the environment to survive. Terrestrial
species have their favourable environmental condition at their best active.
Some of these is discuss in this this unit.
Longhurst, 1960. A summary of the food of West African demersal

fish. Bull.Inst.Fondam.Afr.Noire (A Sci.Nat.), 22:276–82
National Marine Fisheries Service and U.S Fish and Wildlife

Service.(1998). Recovery Plan for U.S Pacific Populations of
the Olive Ridley Turtle (Lepidochelys olivacea). National
Marine Fisheries Service, Silver Spring, MD.
42
Barbour, Roger, Ernst, Carl, & Jeffrey Lovich. (1994). Turtles of the
United States and Canada. Washington, DC: Smithsonian
Institution Press.
Department of the Environment, Water, Heritage and the Arts (2010).

Lepidochelys olivacea in Species Profile and Threats Database,
Department of the Environment, Water, Heritage and the Arts,
Canberra. Available from:
http://www.environment.gov.au/sprat. Accessed 20 Apr 2010.
https://byjus.com/biology/scientific-name-of-bear/
https://www.monkeyworlds.com/
https://thetinyphant.com/terrestrial-animals/
https://www.bing.com/videos/search?q=what+is+terrestrial+animals&&
view=detail&mid=CFD2D59C5AF7A73BF04DCFD2D59C5AF7A73B
F04D&&FORM=VRDGAR&ru=%2Fvideos%2Fsearch%3Fq%3Dwhat
%2Bis%2Bterrestrial%2Banimals%26FORM%3DHDRSC3
https://www.bing.com/videos/search?q=Terrestrial+and+Aquatic&&vie
w=detail&mid=9AC4FC5D4310CDE831239AC4FC5D4310CDE83123
&&FORM=VRDGAR&ru=%2Fvideos%2Fsearch%3Fq%3DTerrestrial
%2Band%2BAquatic%26FORM%3DRESTAB
43
Answer to SAEs 1
1. Snakes are limbless reptiles with long, cylindrical bodies, scaly
skin, lidless eyes, and a forked tongue. Most species are not
poisonous, some are mildly poisonous, and others produce a
deadly poison. The term venom is commonly used to describe the
poison produced by a snake.
Answer to SAEs 2
2. Sloths, Koalas, Beckos, Tree Snakes, Possums, Orangutans,
Parrots, Woodpeckers, Chameleons, Red Pandas.
Answer to SAEs 3
3. The amphibians inhabit a variety of habitats, including terrestrial,
fossorial, freshwater ecosystems. Examples are; American
Bullfrogs, Geckos, Salamanders, Toads, Newts, Tortoise,
Alligator Newts, Worms, Axolotls, Green Tea Frogs.
Glossary
°C = degrees Celsius
cm = centimeters
CO2 = Carbondioxide
CECAF= Fishery Committee for the Eastern Central Atlantic
CH4 = methane (CH4)
ft = feets
°F = degree Fahrenheit
Hz = Hertz
in = inches
Ib = pounds
Kgs = kilograms.
km = kilometers
m = meters
mm = millimeters
NaCl = Sodium Chloride
O2 = Oxygen
oz = ounce
pH = Hydrogen ion concentration
TEDs = turtle exclusion devices
L = length
HIV = Human immunodeficiency virus
44
% = percentage
g = grams
spp = species
End of the Module Questions
1. Who is Charles Elton?

2. Diferentiate between Habitat and Niche
3. What is Mutualism?
4. What is Aquatic animal?
5. What is Terrestrial animal?
45
Module 2: Ecology in Animal Population
Module Introduction
Module Two is concerned with the growth and regulation of population

size, as well as the factors influencing them. Populations are not stable
and always exhibit up and down variations in response to changes in
environmental or intrinsic factors. The measurement of population
dynamics is very important in ecological study of animals. Life table and
K- factor analysis has been applied to a variety of animal species.
Unit 1 Growth rate and Age structure of animal population

Unit 2 Factors affecting Animal Population
Unit 3 Measurement of Population dynamics
Unit 4 Life tables and K-values
Glossary
Unit 1: Example 1
Contents
1.1 Introduction
1.3 Animal Population
1.3.1 Growth Patterns
1.3.2 Age Structure of Animals
1.3.3 Population Growth Curve
1.4 Summary
1.5 References/Further Readings/ Web Sources
1.1 Introduction
A population is a collection of individuals of the same species that live

together in a region. Population ecology is the study of populations
(especially population abundance) and how they change over time.
Crucial to this study are the various interactions between a population and
its resources. A population can decline because it lacks resources or it can
decline because it is prey to another species that is increasing in numbers.
46
Populations are limited by their resources in their capacity to grow; the

maximum population abundance (for a given species) an environment can
sustain is called the carrying capacity. As a population approaches its
carrying capacity, this has led to overcrowding which means that there
are fewer resources for the individuals in the population and leading to
reduction in the birth rate. A population with these features is said to be
density dependent. Of course most populations are density dependent to
some extent, but some grow (almost) exponentially and these are, in
effect, density independent.
At the end of this unit, a student should be able to:
 Explain the term population growth, age distribution and

population size
 Describe the growth pattern in animal population
 Estimate the size of a population using two different techniques.
1.3 Animal Population
Population density is the number of individuals of a certain species per

unit area or volume, and population distribution is the pattern of
dispersal of individuals within that area. They are indispensable
variables for ecologists to analyse and discover the spreading pattern of
a certain species within a certain area and time. Consider calculating the
average density of people in Nigeria, but we know very well that most
people live in cities where the number of people per unit area is
dramatically higher than that in the country. Therefore, basing ecological
population models solely on density can be misleading.
The density and distribution of a population changes with time, due to

abiotic factors (inorganic factors) as well as biotic factors (organic
factors). Abiotic factors that could have an influence on a population
include temperature, rainfall, type of soil and so forth, while biotic
factors are those that are related to other living things. For example, a
particular kind of plant pervading only in a particular area is very likely
to affect the density and distribution of a population of an animal that
feeds only on it. In these situations, limiting factors are those that
particularly determine whether an organism lives in an area.
An example of a limiting factor is found in mountainous regions and high

latitudes where timberline is the limit of tree growth. Trees cannot grow
47
above the high timberline because water remains frozen at the low
temperature for most of the year. In this case, timberline, or more
specifically, temperature is the limiting factor for tree density and
distribution.
Population size is the number of individuals in a population.

Technically, genetic relationship is used to distinguish whether an
individual belongs to a population or not. Instead of simply counting, it
is necessary to estimate the present population size, using methods which
vary with the kind of species in question.
Just as density and distribution, population size fluctuates with time.
There are generally four sources of contribution to the fluctuation of a
population size, natality (rate of birth), mortality (rate of death),
immigration and emigration. How they each changes the population size
is indicated below.
Immigration
Natality Population Size Mortality
Emigration
Fig 1: Factors that Affect Population Size
Usually it is acceptable to assume that immigration and emigration are

about equal and therefore only necessary to consider natality and
mortality. What population density? What is Population size?
1. What are limiting factors?
1.3.1 Growth Patterns
Theoretically, there exist two distinct and simple growth patterns, or

mathematical models for population growth. In the first one, only one
reproductive chance is given to members of the population during their
entire lifespan. Once mission accomplishes, they die. Many insects and
48
annual plants reproduce in this manner. In the other model, members

experience many reproductive events throughout their lifetime. Most
vertebrates and trees have this pattern of reproduction.
Expressed in mathematical equations and graphs, the two growth
patterns can be referred to as the exponential growth pattern and the
logistic growth pattern respectively. We are not going to resort to
number and equation here, but a glance at the graphs below will be
enough for this introductory course.
Fig 2: Population Exponential Growth (Source: www.econguru.com)
Two major features of the curve are:
Lag phase: in which growth is slow because population base is small

and
Exponential growth phase: in which growth is accelerating, that is, the
rate of growth itself grows.
Fig 3: Population Logistic Growth (Source: www.econguru.com)
Four major features of the curve are:
A. Lag phase: in which Growth is slow because the population base

is small
49
B. Exponential growth phase: in which Growth is accelerating,

that is, the rate of growth itself grows.
C. Deceleration phase: in which the rate of population growth
slows down.
D. Stable equilibrium phase: in which little growth because births
and death are about equal.
Population growth patterns require an assumption that members of a

population are all identical individuals. However, individuals are in their
different stages of lifespan. In a given period of time, some are born and
some die. See if you can figure out the data in this table:
Table 1: Showing A Life table for a Poa annua (a grass) Cohort
Age Number Proportion Number Mortality Avg. Number

(Month) Observed Surviving Dying Rate Per of
Alive Capita seeds/individual
0-3 843 1,000 121 0.143 0
3-6 722 857 195 0.271 300
6-9 527 625 211 0.400 620
9-12 316 375 172 0.544 430
12-15 144 171 95 0.626 210
15-18 54 64 39 0.722 60
18-21 15 17.8 12 0.800 30
21-24 3 3.6 3 1.000 10
24 0 - - - -
(Source: www.econguru.com)
A cohort of a population, is the total number of new births of it at the

same time. As you can see in table 1, the number of grass observed
alive at the beginning is 843, and data are noted down every 3 months.
The grass are gradually dying out, and the life stages at which they
perish vary. From another perspective, let's look at the number that
survives instead of focusing on the number dying in each period. After
the first observational period, 722 survive the first 3 months.
Survivorship is the probability of new born individuals of a cohort
surviving to particular ages. Plotting the number surviving against
percent of life span, we get survivorship curves that show the number
of individuals of a cohort still living over time.
50
For the sake of discussion, we will establish three types of

representative survivorship curves hypothetically, displayed in the
graph of the following image. Curve I is characteristic of a population
in which most individuals survive will pass the midpoint. On the
contrary, Curve III typifies populations wherein most individuals die
young. In the type II curve, survivorship decreases at a constant rate
throughout the lifespan.
Fig 4: The different types of Survivorship curve (Source:

www.econguru.com)
As shown by the other three graphs, the survivorship curves of natural

populations do not fit these three idealized curves congruously.
However, similarities exist to help understanding. Survivorship curve for
gulls fits the type II curve somewhat. The survivorship curves of human
males and females differ slightly but both resemble the type I curve fairly
well. The survivorship curve for Poa annua seems to be a combination
of the type I and type II curves.
Survivorship curves denote mortality patterns of a certain population

over a certain period of time. It may vary in abnormal conditions, but in
most cases the pattern stay predictable. Mention the two growth
patterns? What is Survivorship?
1. What are the major features of the logistic growth curve?
1.3.2 Age Structure of Animals
51
This is the number of individuals in different age classes; pre-

reproductive, reproductive, and post-reproductive. Short-lived organisms
increase rapidly, with short span between generations long-lived
organisms, increase slowly, and have long span between generations.
Determining individual ages of animals with the following;
A. Marking of young individuals through time

B. Examining a representative sample of carcasses of individuals
wear
C. Replacement of teeth in deer and other ungulates.
D. Annual growth rings in the horns of sheep
E. Plumage changes and wear in birds and
F. Growth rings in scales of fishes
Age Pyramids is comparing of the percentages population in different

age groups. The pyramids with a broad base of young suggest growing
populations. Pyramids with a narrow base of young and even ratios,
suggest a declining or aging population and depict changing dynamics
of a population.
Fig 5: Showing the different age structure (Source: www.econguru.com)
In-Text Question(s)
The age structure of animals is divided into how many parts or groups?
Answer: The age structure of animals is divided into three groups, pre-
reproductive, reproductive, and post-reproductive
1.3.3 Population Growth Curve
This shows the net result of births, deaths, and dispersion. It usually
shows three to five phases. Most organisms show 3 phases: lag phase,
exponential growth phase, and equilibrium phase
52
i) Lag phase - slow growth because the process of growth and

reproduction of offspring takes time
ii) Exponential growth phase - characterized by more organisms
undergoing reproduction, so that the population begins to
increase at very fast rate; birth rate exceeds the death rate
iii) Equilibrium phase - characterized by the birth rate and death rate
that are equal to one another; population will stop growing and
reach a relatively stable population.
The net result of births, deaths, and dispersion can take a number of
forms:
i) J-shaped or exponential growth form – density occurs when

there is Increases in a geometric fashion until the population
runs out of some resource or encounters some other limitation
(Fig 5).
ii) S-shaped or Sigmoid growth - limiting factors resulting from

crowding provide negative feedback that reduces the rate of
growth more and more as density increases. If the limitation is
linearly proportional to density, the growth form will be a
symmetrical sigmoid curve with density levelling off as to reach
the carrying capacity; the carrying capacity represents the
maximum sustainable density (Fig 6).
Fig 5: Showing the J-shaped growth form (Source: www.econguru.com)
53
Fig 6: Showing the S-shaped or Sigmoid growth (Source:

www.econguru.com)
Fig 7: showing the J and S population curve (Source:

www.econguru.com)
The Basic Concepts of Rate are shown below:
i) Population dynamics - the study of changes in the relative

number of organisms in populations and the factor explaining
these changes.
ii) Rate – obtained by dividing the change in some quantity by the

period of time elapsed during the change
54
iii) Equation ∆N/∆t = average rate of change in the number of

organisms per time.
iv) Instantaneous rate: rate at a particular time (rate of change

when ∆ t approaches 0,
v) d= derivative dN/dt = i.e rate of change in the number of

organisms per time at a particular instant).
vi) Point of inflection – point where growth rate is maximum
What is Exponential growth phase?
1.4 Summary
A population is defined as the organism belonging to the same species

located in the same place at the same time and ecology is the study of
those organisms in relation to their environments. At this ecological level,
the interest is in the growth and regulation of population size, as well as
the factors behind them.
Chapman, J. L. and Reiss, M. J. (2010) Ecology, Principles and Applications, 2nd

ed. Cambridge University Press.

region. I. Balanus glandula and several predatory species of
Thais. Ecological Monographs 40: 49-78.
Fundamental of ecology, author: EconGuru, introduction to

ecosystems and community. Ecological population; Chapter 1
to chapter 3.
https://ibiologia.com/population/
http://faunaofindia.nic.in/PDFVolumes/spb/005/index.pdf
https://www.researchgate.net/publication/233797058_Estimating_anima
l_population_density_using_passive_acoustics
55
https://www.bing.com/videos/search?q=animal+population+lecture+not
es&&view=detail&mid=B1073E47AB9137135B05B1073E47AB91371
35B05&&FORM=VRDGAR&ru=%2Fvideos%2Fsearch%3Fq%3Dani
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56
Answers to SAEs 1
1. Limiting factors are those that particularly determine whether an
organism lives in an area. An example of a limiting factor is
found in mountainous regions and high latitudes where
timberline is the limit of tree growth. Trees cannot grow above
the high timberline because water remains frozen at the low
temperature for most of the year. In this case, timberline, or
more specifically, temperature is the limiting factor for tree
density and distribution.
2. The Four major features of the curve are:

A. Lag phase: in which Growth is slow because the population base
is small
B. Exponential growth phase: in which Growth is accelerating,
that is, the rate of growth itself grows.
C. Deceleration phase: in which the rate of population growth
slows down.
D. Stable equilibrium phase: in which little growth because births
and deaths are about equal.
57
Unit 2 Factors Affecting Animal Population
Contents
2.1 Introduction
2.3 Factors affecting animal population
2.3.1 Birth rate or Natality rate
2.3.2 Death or Mortality rate
2.3.3 Migration
2.3.4 Age distribution (Age composition)
2.3.5 Abiotic and Biotic Factors
2.4 Summary
2.1 Introduction
There are various factors affecting population size: natality, mortality,

immigration, and emigration etc. Natality refers to the birth rate.
Mortality refers to the death rate. Immigration occurs when individuals
from one population of a species join another population of the same
species. Emigration occurs when individuals leave a population. The
intrinsic population growth rate (r) = (births + immigration) – (deaths +
emigration). Zero population growth occurs when the birth rate and
immigration are balanced over the long term by the death rate and
emigration.
At the end of this unit, the student should be able to state and explain
the different factors affecting animal population
2.3 Factors Affecting Animal Population
All species of organisms have a great capacity to reproduce. A theoretical

growth curve reflects population growth at its maximum rate of increase,
per individual, under ideal conditions. Without restraints, the size of any
population would increase at an exponential rate. The result is a J-shaped
curve. As long as birth rates remain even slightly higher than death rates,
a population will grow exponentially. The size and density of a
58
population are affected by various factors. Some of the important ones

are discuss below.
2.3.1 Birth rate or Natality rate
Natality rate is the rate at which new individuals are added to a particular
population by reproduction (birth of young ones or hatching of eggs or
germination of seeds/spores).
It is generally expressed as number of births per 1,000 individuals of a
population per year.
'Absolute', 'physiological or maximum natality', refers to the theoretical

maximum production of new individuals under ideal conditions. But it is
never realised because of environmental resistance (factors like inter and
intra specific competitions, availability of food, space etc.)
'Actual birth rate' being achieved under existing conditions, which is

much lesser than 'absolute natality is called realized natality. Higher
realized natality rate increases the population size and population
density. What is Birth rate?
Natality rate is the rate at which new individuals are added to a particular
population by reproduction (birth of young ones or hatching of eggs or
germination of seeds/spores)
2.3.2 Death or Mortality rate
Mortality rate is the rate at which the individuals die or get killed. It is
the opposite of natality rate.
Mortality rate is generally expressed as number of deaths per 1,000

individuals of a population per year.
Lowest death rate for a given species in most favourable conditions is

called potential mortality while the actual death rate being observed in
existing conditions is called realized mortality.
Realized mortality decreases the population size and population density.
The percentage ratio of natality over mortality expressed in percentage

is called Vital index.
Vital index determines the normal rate of growth of a population.
59
Table 1: Showing the Differences between Natality rate and Mortality rate
What is death rate?
2.3.3 Migration
The movement of organisms in large numbers from one place to another

place is termed as migration. There are many reasons for migration. The
term migration is mainly used to define regular and periodic movements
of the population away from or back to their place of origin. These
migrations are of various types. Some may span for an entire lifetime and
it happens to be a single migration of an individual. This is in the case for
the Pacific salmon. On another hand, an individual may migrate
frequently, like many of the migratory birds and other mammals. Animals
usually travel in groups along well-known routes or may travel in separate
gathering for breeding.
Immigration is the permanent entry of new individuals of same species

into a population from outside. It increases the size of local population.
Emmigration is the permanent movement/departure of individuals of same

species out of the local population due to several reasons such as lack of
food, scarcity of space (overcrowding), etc. Emmigration decreases the
size of local population, but the species spread to new areas.
If more individuals are added than lost, then the population will show
positive growth. If more individuals are lost than added, then the
population will show negative growth. But if the two rates are equal, then
the population will become stationary and is called zero growth.
Population growth = (Birth + Immigration) - (Death + Emmigration)
Many factors play their role in the initiation of migration. External factors
like climate, natural disasters, drought, shelter, food shortage, etc may
cause animals to migrate to seek better conditions.
Seasonally, the timing of migration is influenced by internal “clocks” that
are influenced by day length and perhaps also weather. For example –
Consider a species of the deer that live in a certain park. They would
60
migrate to warmer places during winter if the park they reside is harsh for
them to survive. But, if they are living near a place, where they can graze
throughout the year, then there is no need for their migration.
Over shorter time scales, the exact timing of migratory movements is
greatly influenced by the weather. Birds tend to move when conditions
favour flying in the direction they need to go (e.g., when they have a
tailwind, when air turbulence is low, when it is not raining).
It is noted that many birds migrate at night and there are possible reasons
for this action.
1. The atmosphere is more stable at night than during the day because
there are fewer thermals (“updrafts” caused by warming of the
Earth’s surface).
2. The air temperature is lower, which may make thermoregulation
easier; remember that flight generates a lot of heat that birds need
to offload.
3. Predation risk may be lower than during the day.
But some species do migrate by day. In particular, large soaring birds
such as hawks, storks and pelicans move during the daytime. These birds
use thermals to help them travel.
In birds, the migratory instinct is closely linked to the cycle of
enlargement of the reproductive system in spring. There are several
species of birds that migrate to long ranges, flying to the north in the
springtime to breed in the warm conditions and migrate back to their
origin.
Bird migration is the natural seasonal movement, often south and north
along a flyway, in between the breeding grounds and wintering grounds.
Migratory birds fly several kilometres in search of the best environmental
specifications and habitats for food, breeding and raising their young
ones. Migration is motivated primarily by the unavailability of food and
nesting locations. It occurs mainly in the northern hemisphere, where
birds travel in search of warmer places such as the Caribbean Sea or the
Mediterranean Sea. What is Migration? What do you understand as
positive growth?
1. Write on the timing of migration influenced by internal clocks.
61
2.3.4 Age distribution (Age composition)
The relative abundance of the organisms of various age groups in the

population is called age distribution of population.
With regard to age distribution, there are three kinds of populations.
1. Rapidly growing population is a population, which has high

birth rate and low death rate, so there are more number of young
individuals in the population.
2. Stationary population is a population, which has equal birth and
death rates, so population shows zero population growth.
3. Declining population is a population, which has higher death
rate than birth rate, so the population has more numbers of older
individuals.
What is stationary population?
2.3.5 Biotic and Abiotic factors
Biotic and abiotic are the two essential factors responsible for shaping
the ecosystem. The biotic factors refer to all the living organisms present
in an ecosystem from animals and humans, to plants, fungi, and bacteria,
and the abiotic factors refer to all the non-living components like
physical conditions (temperature, pH, wind, humidity, salinity, sunlight,
etc.) and chemical agents (different gases and mineral nutrients present
in the air, water, soil, etc.) in an ecosystem. Therefore, both the abiotic
and biotic resources affect survival and reproduction process.
Biotic factors like organisms of other species living in the same area
affect the population, as they involve in different types of food
relationships. For e.g., if the population happens to increase in size, it is
brought down by an increase in its predators number or decrease in the
amount of available food. Different populations have different ability to
tolerate changes in weather, physico-chemical and biotic factors. This is
called resilience. In nature, factors like predators, diseases, food scarcity
etc. prevent a population to sour towards infinity. The sum of all these
factors, which prevent a population from growing at its maximum rate,
is called environmental resistance or population regulation.
The following are examples of Abiotic factors that affect population.
1. Water
Wetland conditions such as shallow water, high plant productivity, and

anaerobic substrates provide a suitable environment for important
62
physical, biological, and chemical processes. Because of these processes,

wetlands play a vital role in global nutrient and element cycles.
The rate of diffusion of carbon dioxide and oxygen is approximately

10,000 times slower in water than it is in air. When soils become flooded,
they quickly lose oxygen and transform into a low-concentration
(hypoxic -with less than 2 mg O2l−1) environment and eventually become
completely (anoxic) environment where anaerobic bacteria thrive among
the roots. Water also influences the spectral composition and amount of
light as it reflects off the water surface and submerged particles.
Aquatic plants exhibit a wide variety of morphological and physiological

adaptations that allow them to survive, compete and diversify these
environments. For example, the roots and stems develop large air spaces
(Aerenchyma) that regulate the efficient transportation gases (for
example, CO2 and O2) used in respiration and photosynthesis. In drained
soil, microorganisms use oxygen during respiration. In aquatic
environments, anaerobic soil microorganisms use nitrate, manganese
ions, ferric ions, sulfate, carbon dioxide and some organic compounds.
The activity of soil microorganisms and the chemistry of the water
reduces the oxidation-reduction potentials of the water. Carbon dioxide,
for example, is reduced to methane (CH4) by methanogenic bacteria.
Salt water plants (or halophytes) have specialized physiological

adaptations, such as the development of special organs for shedding salt
and osmoregulate their internal salt (NaCl) concentrations, to live in
estuarine, brackish, or oceanic environments. The physiology of fish is
also specially adapted to deal with high levels of salt through
osmoregulation. Their gills form electrochemical gradients that mediate
salt excresion in saline environments and uptake in fresh water.
2. Gravity
The shape and energy of the land is affected to a large degree by

gravitational forces. On a larger scale, the distribution of gravitational
forces on the earth are uneven and influence the shape and movement of
tectonic plates as well as having an influence on geomorphic processes
such as orogeny and erosion. These forces govern many of the
geophysical properties and distributions of ecological biomes across the
Earth. On an organism scale, gravitational forces provide directional
cues for plant and fungal growth (gravitropism), orientation cues for
animal migrations, and influence the biomechanics and size of animals.
Ecological traits, such as allocation of biomass in trees during growth are
subject to mechanical failure as gravitational forces influence the
position and structure of branches and leaves. The cardiovascular
systems of all animals are functionally adapted to overcome pressure and
63
gravitational forces that change according to the features of organisms

(e.g., height, size, shape), their behaviour (e.g., diving, running, flying),
and the habitat occupied (e.g., water, hot deserts, cold tundra).
3. Pressure
Climatic and osmotic pressure places physiological constraints on

organisms, such as flight and respiration at high altitudes, or diving to
deep ocean depths. These constraints influence vertical limits of
ecosystems in the biosphere as organisms are physiologically sensitive
and adapted to atmospheric and osmotic water pressure differences.
Oxygen levels, for example, decrease with increasing pressure and are a
limiting factor for life at higher altitudes. Water transportation through
trees is another important ecophysiological parameter where osmotic
pressure gradients factor in. Water pressure in the depths of oceans
requires that organisms adapt to these conditions. For example,
mammals, such as whales, dolphins and seals are specially adapted to
deal with changes in sound due to water pressure differences. Different
species of hagfish provide another example of adaptation to deep-sea
pressure through specialized protein adaptations.
4. Light intensity
It may be interesting to note that even something as simple as the level

of light present in an environment or ecosystem can greatly affect the
organisms in that ecosystem. As you know, photosynthesis is the main
form of energy consumption for plants, which means that all plants need
a certain level of light to create their own food. Some plants in dark
ecosystems have evolved to a point where they make do with minimal
amounts of light, such as the plants found deep in the oceans where light
doesn't reach.
Some plants have evolved for optimum growth in bright sunlight. An

example of this is a cactus houseplant. Cacti originally come from deserts
where they grow in bright sunlight. Other plants have evolved to grow
in shade.
Many orchids, which are also kept as houseplants, grow on trees in the
rainforest and have evolved for optimum growth in darker conditions. If
you were to put an orchid on a bright windowsill and a cactus in a dark
corner of your room neither plant would grow well.
5. Temperature
Temperature is a limiting factor for photosynthesis - and low temperature

therefore limits growth of plants. In cold climates, the number of plants
64
is usually low - which limits the number of herbivores that can live there.
Both animals and plants have evolved to grow healthily at their optimum
temperatures. If you planted either your cactus or orchid houseplants
outside in cold temperatures, they would die. Similarly, animals that have
evolved to live at the North Pole, such as the polar bear, could not survive
in warmer conditions.
6. Moisture levels
More people kill houseplants by overwatering than by under-watering

them. Many plants cannot survive in waterlogged soils. Their roots are
unable to respire, they rot and the plant dies. Other plants, such as pitcher
plants, grow best in bogs where the moisture levels are high. Soil moisture
meters can accurately determine how wet an area is.
7. Soil pH content
The pH balance of an ecosystem refers to the general level of acidity or

alkalinity present in the environment. Pure water has a pH of 7, meaning
it is neutral. Acidic mixtures have a pH balance of less than 7 while
alkaline mixtures have a pH balance of more than 7. This also affects the
organisms in an environment, because many creatures or plants, or
microorganisms cannot survive in certain pH ranges.
The pH of soils can have a huge effect on the plants that are able to grow
in them. Some plants, like azaleas, grow best in acidic soils and will
quickly die if planted in alkaline soils. Others, like clematis, prefer
alkaline soils. Some, like the hydrangea, can grow in both. These plants
are unusual in that their flower colour changes in different soils. Just like
universal indicator paper, hydrangea flowers are pink in acidic soils and
blue in alkaline soils.
The pH of water can also affect the aquatic organisms that are found there.
Different species have evolved to survive at different pH levels found
within water.
65
Fig 1: The graph highlights the various pH tolerance of different species

found in water
8. Wind and turbulence

The architecture of inflorescence in grasses is subject to the physical
pressures of wind and shaped by the forces of natural selection
facilitating wind-pollination (or anemophily).
Turbulent forces in air and water have significant effects on the

environment and ecosystem distribution, form and dynamics. On a
planetary scale, ecosystems are affected by circulation patterns in the
global trade winds. Wind power and the turbulent forces it creates can
influence heat, nutrient, and biochemical profiles of ecosystems. For
example, wind running over the surface of a lake creates turbulence,
mixing the water column and influencing the environmental profile to
create thermally layered zones, partially governing how fish, algae, and
other parts of the aquatic ecology are structured. Wind speed and
turbulence also exert influence on rates of evapotranspiration rates and
energy budgets in plants and animals. Wind speed, temperature and
moisture content can vary as winds travel across different land features
and elevations.
9. Soil mineral content
Many plants require high levels of soil minerals to grow well. An example
of this is magnesium, which is required to produce chlorophyll. Plants
with unnaturally yellow leaves may have a magnesium deficiency.
Carnivorous plants, such as pitcher plants, have evolved to catch insects
to supplement the low levels of minerals found in the soils in which they
grow.
10. Carbon dioxide levels for plants
Carbon dioxide is a reactant in photosynthesis which means plants need

it to survive. Areas with higher levels of carbon dioxide are more likely
to have healthy plants growing. Farmers often release carbon dioxide
within their greenhouses to maximise their crop yield. Woodlands often
have higher carbon dioxide levels than open grassland, so many plants
living in open areas have evolved mechanisms to overcome a shortage of
carbon dioxide.
11. Oxygen levels for aquatic animals
Oxygen from the air and oxygen produced by aquatic plants dissolves in
water. Without this, aquatic animals would suffocate and die. Healthy
lakes and rivers have high levels of oxygen, and polluted waters often
have low levels of oxygen. This pollution means that only
66
certain species can survive there such as sludge worms. These are bio-
indicator species because their presence or absence informs us about the
condition of the habitat.
12. Fire
Forest fires modify the land by leaving behind an environmental mosaic

that diversifies the landscape into different seral stages and habitats of
varied quality. Some species are adapted to forest fires, such as pine trees
that open their cones only after fire exposure.
Approximately 350 million years ago (near the Devonian period) the
photosynthetic process brought the concentration of atmospheric oxygen
above 17%, which allowed combustion to occur. Fire releases CO2 and
converts fuel into ash and tar. Fire is a significant ecological parameter
that raises many issues pertaining to its control and suppression in
management.
Fire creates environmental mosaics and a patchiness to ecosystem age and

canopy structure. Native North Americans were among the first to
influence fire regimes by controlling their spread near their homes or by
lighting fires to stimulate the production of herbaceous foods and basketry
materials. The altered state of soil nutrient supply and cleared canopy
structure also opens new ecological niches for seedling establishment.
Most ecosystem are adapted to natural fire cycles. Plants, for example, are
equipped with a variety of adaptations to deal with forest fires. Some
species (e.g., Pinus halepensis) cannot germinate until after their seeds
have lived through a fire. This environmental trigger for seedlings is called
serotiny. Some compounds from smoke also promote seed germination.
Fire plays a major role in the persistence and resilience of ecosystems.
13. Natural Disaster

A drastic change in the environment destabilizes or even exterminates a
population. Natural calamities such as earthquake, volcanic eruptions etc.
cause drastic changes in the environment leading to the destruction of the
resources. What is the important of Moisture levels in plant population?
1. Write on the important of oxygen levels for aquatic animals.
67
2.4 Summary
Populations are not stable and always exhibit up and down variations in
response to changes in environmental or intrinsic factors. The irregular
variations in size of populations are called fluctuations or population
cycle. This irregularity may be as result of living or non-living factors
affecting the organisms. As explained in this unit. Some of the example
of the factors are natality and mortality rate, immigration and emigration,
abiotic and biotic factors and carrying capacity etc.
Bell, H.L. 1971. Effect of low pH on the survival and emergence of

aquatic insects. Water Research 5, 313-319.
Cairns, J. and J.R. Pratt 1993. A history of biological monitoring using

benthic macroinvertebrates. Pages 10-27, In: D. M. Rosenberg
and V.R. Resh (eds.) Freshwater Biomonitoring And Benthic
Macroinvertebrates, Chapman & Hall, N.Y.
Conca, J. L. and Wright, J. 1992. Diffusion and Flow in Gravel, Soil, and
Whole Rock. Applied Hydrogeology, Hanover.
Faith, D.P. and R.H. Norris 1989. Correlation of environmental variables

with patterns of distribution and abundance of common and rare
freshwater macroinvertebrates. Bio. Conserv. 50, 77-98.
Foster, G.N. 1991. Aquatic beetle population changes associated with

recreating a trout fishery by limiting a lake catchment. Arch.
Hydrobiol. 122(3), 313-322.
Fryer, G. 1980. Acidity and species diversity in freshwater crustacean

faunas. Freshwater Bio. 10, 41-45
https://people.clas.ufl.edu/gillooly/files/predicting_mortality.pdf
https://www.demographic-research.org/volumes/vol15/14/15-14.pdf
https://www.dhnature.org/uploads/2/5/7/0/25708496/animal_migration
s.pdf
https://www.researchgate.net/publication/299888982_Patterns_of_anim
al_migration
68
https://www.bing.com/videos/search?q=animal+migration+pdf&&view
=detail&mid=93BE3075757D684903C293BE3075757D684903C2&&
FORM=VRDGAR&ru=%2Fvideos%2Fsearch%3Fq%3Danimal%2Bm
igration%2Bpdf%26FORM%3DHDRSC3
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69
Answers to SAEs 1
1. Seasonally, the timing of migration is influenced by internal
“clocks” that are influenced by day length and perhaps also
weather. For example – Consider a species of the deer that live in
a certain park. They would migrate to warmer places during winter
if the park they reside is harsh for them to survive. But, if they are
living near a place, where they can graze throughout the year, then
there is no need for their migration.
Answers to SAEs 1
1. Oxygen from the air and oxygen produced by aquatic plants
dissolves in water. Without this, aquatic animals would suffocate
and die. Healthy lakes and rivers have high levels of oxygen, and
polluted waters often have low levels of oxygen. This pollution
means that only certain species can survive there such as sludge
worms. These are bio-indicator species because their presence or
absence informs us about the condition of the habitat.
70
Unit 3 Measurement of Population Dynamics
Contents
3.1 Introduction
3.3 Population Dynamics
3.4 Summary
3.1 Introduction
There are a number of methods to describe populations from field

measurements such as, dynamic and static life tables, and transition
matrices. All are based on using census on individuals in groups,
categorized according to their state (age, size, or stage). They are used to
explain population dynamics in relation to demographic processes, and
to predict the fate of the population. The methods differ because it
depends on the following;
a) convenience of data collection,

b) basic assumptions and
c) the way they derive population growth rate.
Because this also depends on the general biology and life cycle of the
organism, some methods are better for some kinds of populations, but
not for others.
At the end of this unit, student should be able to

 explain the measurement of population dynamics and the
survival ship curve
3.3 Population Dynamics
Dynamic life tables describe survivorship and fecundity (production of

eggs, seeds, or young) at different ages of a cohort (a sample of
individuals recruited approximately at the same time). Usually the cohort
is followed until the last member dies. Survivorship is calculated as the
71
relative change in the number of individuals of the cohort between two

successive censuses. The census is very simple, since individuals need
only be recognized to differentiate them from other cohorts in the same
population, and thus are only counted. No marking is needed, because
for an individual to be counted means it survived since last census, and
determines its age unambiguously. Transition models, as will be
discussed later, require more elaborate censuses.
Age may be convenient to use in cohort studies, but it does not always
play a primary role in the development and life cycle of the organisms.
This is probably true for all plant and most animal taxa, except birds and
mammals. The latter's determinate growth (an individually fixed adult
size) and hormonal clock, combined with their ability to learn, make age
a suitable attribute to describe population dynamics. In most other cases,
size is biologically more important, as it determines resource acquisition,
competitive ability, survivorship and sexual maturity and reproductive
output. Only if size and age are strongly correlated in such a population,
age may be preferable.
Stage is very useful in cohort studies of annual organisms if surviving

organisms all pass through relatively short stages, so that stages hardly
overlap (occur at the same time), and stage and age are correlated.
However, if size or stage is attained at different rates for different
individuals in the population, as is often the case, they are not useful for
constructing life tables. Then, transition models are more appropriate.
What is Dynamics Life table? Dynamic life table is used for what kind
of animals?
i) Survivorship Curve
Two kinds of information are derived from these cohort studies:

survivorship curves and population growth rates. Survivorship curves
show how mortality varies with age of the individuals of the cohort or
is a graph that measures the proportion of individuals in a given species
that are alive at different ages. Age-specific mortality, as well as age-
specific fecundity, is due to changing susceptibilities and capabilities
of the individual, and the variation in its environmental exposure.
Depending on the age at which most of the mortality takes place,
organisms can have different survivorship curves. Deevey (1947)
described three types of curves (Fig. 1):
1. A type I survivorship curve shows individuals that have a high
probability of surviving through early and middle life but have
a rapid decline in the number of individuals surviving into late
life. This basically means that most of the individuals will make
it to adulthood but the proportion surviving into old age is
72
greatly decreased. A type I survivorship curve is plotted as a

convex curve on a graph.
2. A type II survivorship curve shows a roughly constant mortality
rate for the species through its entire life. This means that the
individual's chance of dying is independent of their age. Type II
survivorship curves are plotted as a diagonal line going
downward on a graph.
3. A type III survivorship curve depicts species where few
individuals will live to adulthood and die as they get older
because the greatest mortality for these individuals is
experienced early in life. This type of survivorship curve is
drawn as a concave curve on a graph
Fig.1. Three types of survivorship curves (Begon et al. 1990, Deevey

1947).
A classic example of a type I survivorship curve is the human

population. Advances in medicine and technology have made the
chances of surviving through early and middle life highly probably for
humans. The highest levels of mortality appear in late life. Type II
survivorship curves indicate that the chance if dying is independent of
age. Type II survivorship curves are used for animals, such as birds,
who have many random chances of being killed or dying at all stages
of their life. Type III individuals initially have a rather low chance of
survival. Those that do survive may live to an advanced age. Examples
include many fish and other marine organisms
73
Table 1: Showing the diffeeces in Survivorship strategies between

Type 1 and Types III
Type I Type III
Habitats Stable Disturbed
Life span Long Short
Reproductive age Later in Life Early in Life
Gestation period Long Short
Number of Offspring Low High
Care of Offspring Greater Little
Infant Mortality Low High
Dispersal Low High
Growth Curve Logistic-K Exponential - r
(Source: Begon et al. 1990)
What is survivorship curve?
1. What does Survivorship curve type I indicate?
II) Population growth rate
Population growth rate of a cohort is defined as the basic reproductive

rate R0 for a generation over its lifetime. Using dynamic life tables is best
suited for semelparous animals include many insects or monocarpic
plants (those that flower, set seeds and then die), especially annual
organisms, as in the study on the annual plant species Phlox drummondii
by Leverich and Levin (1979) (Table 2). These have a single cohort per
year, and thus have no overlapping generations.
Perennial iteroparous /polycarpic organisms (reproductive strategy) do
have overlapping generations, which makes the method less
straightforward, though often still useful. Connell (1970) used this
method on the long-lived barnacle Balanus glandula (Table 3). Basic
reproductive rate R0 measures the mean number of offspring that an
individual of the cohort produces in its lifetime. Offspring denotes the
number of individuals in the first stage of the life cycle (zygotes or
young, depending on the organism).
74
Table 2. A cohort life-table for Phlox drumondii (after Leverich and Levin
1979).
Table 3. A cohort life-table and fecundity schedule for the barnacle

Balanus glandula at Pile Point, San Juan Island, Washington
(Conell, 1970). (* - estimated by interpolation from the survivorship

curve).
There are two ways to arrive at the basic reproductive rate.

The first method is R0= ∑ F x /a0, where ∑ F x, the sum of the Fx's, is
the number of offspring over the life span of the cohort, and a0 is the first
stage. The ratio shows the relative change in population size / density
from one generation of the cohort to the next. R0 is also measured as R0=∑
l x m x, the sum of l x, the chance of an individual surviving to age x (the
time from the birth of the cohort to the census x, not necessarily in years),
times m x, the number of offspring produced during the time from age x-
1 to x. The advantage of this model is, that it is explicit about the relation
between overall population growth and the actual demographic processes
that take place through time in the cohort. For annual organisms this
sequence of processes tracks the changing of the seasons.
75
In cohort studies, R0 is determined over the generation time. If the

organism is an annual plant or animal, R0 also denotes population growth
rate R, defined per year: R = R0. In perennial semelparous or monocarpic
organisms, R0 should be corrected for generation time T (>1 year).
Since R0=RT, so that lnR= (ln R0)/T.

However, for organisms with overlapping generations, it is difficult to
estimate annual growth rate accurately since generation time T is in fact
unknown. Instead of T, R is given by lnR = (ln R0)/Tc.
R0 is corrected by Tc, the average cohort lifespan, which is the average

time from the birth of an individual to the birth of one of its offspring,
calculated as the sum of lengths of time of the offspring of all individuals
divided by the total number of offspring:
Tc= ( ∑x. lx m x)/ ∑ lx m x ∑x. lx m x)/ R0.
The presence of three generations at the same time, i.e. if some individuals
have produced offspring themselves while their parents are still alive,
cannot be incorporated in the equation. What is population growth rate?
1. How is R0 determined In cohort studies?
III) Static life tables
A static life table is similar to a cohort life table but introduces a few
complications. A static life table contains the age groups in a population
at one particular period of time. Thus, cohorts are not followed in time,
but reconstructed using one-time observations. These can be used to
calculate population growth only if an assumption is made. The
assumption is that the mortality experienced by the cohort at any age stays
constant in time. In other words, birth rates and age-specific survivorship
are assumed to be independent of the actual year in which the observations
are made. Only rarely is this assumption truly justified. Therefore, the
conclusions tell us how a cohort should behave, if we would have
observed it and if conditions are constant between years. An example is
the study of red deer by Lowe (1969) (Table 4), described in Begon et al.
(1990).
76
A second, less problematic application of static life tables is using them in

order to reconstruct past events, as Crisp and Lange (1976) did with the
desert shrub Acacia burkitii (Begon et al. 1990, page 144 and 145). They
were able to show, among others, the effects of grazing, because they
compared two stands using one as a control
Table 3. A static life-table for red deer hinds on the island of Rhum, based
on the reconstructed age-structure of the population in 1957 (After Lowe,
1969)
What is Static life table?
3.4 Summary
The measurement of population dynamics is very important in ecological

study of animals. To be able to assess the status of any ecosystem there
is necessity for continuous measurement of the animal population, either
quantitatively or qualitatively.


region. I. Balanus glandula and several predatory species of Thais.
Ecological Monographs 40: 49-78.
77
Crisp, M. D. and R. T. Lange, 1976. Age structure distribution and

survival under grazing of the arid zone shrub Acacia burkitii.
Oikos 27: 86-92.
Deevey, E. S., 1947. Life tables for natural populations of animals.

Quarterly Review of Biology 22: 283-314.
Leverich, W. J. and D. A. Levin, 1979. Age-specific survivorship and

reproduction in Phlox drummondii. American Naturalist 113:
881-903.
Lowe, V. P. W., 1969. Population dynamics of the red deer (Cervus

elaphus L.) on Rhum. Journal of Animal Ecology 38: 425-457.
Richards, O. W. and N. Waloff, 1954. Studies on the biology and

population dynamics of British grasshoppers. Anti-Locust
Bulletin 17: 1-182.
https://dba.stackexchange.com/questions/141404/what-are-static-
tables-dynamic-
tables#:~:text=Static%20tables%20are%20the%20master%20ta
bles%20that%20are,of%20data%20populated%20in%20them%
20that%20hardly%20changes.
http://studyoflife.org/lifetables.pdf
https://www.youtube.com/watch?v=7aJ5rtRm_t8
https://www.bing.com/videos/search?q=Dynamic+life+table&&view=d
etail&mid=9BDB32079B2FFFED6FD59BDB32079B2FFFED6FD5&
&FORM=VRDGAR&ru=%2Fvideos%2Fsearch%3Fq%3DDynamic%
2Blife%2Btable%26FORM%3DHDRSC3
78
Answer to SAEs 1
1. A type I survivorship curve shows individuals that have a high
probability of surviving through early and middle life but have a
rapid decline in the number of individuals surviving into late life.
This basically means that most of the individuals will make it to
adulthood but the proportion surviving into old age is greatly
decreased. A type I survivorship curve is plotted as a convex curve
on a graph.
Answer to SAEs 2
1. In cohort studies, R0 is determined over the generation time. If the
organism is an annual plant or animal, R0 also denotes population
growth rate R, defined per year: R = R0. In perennial semelparous
or monocarpic organisms, R0 should be corrected for generation
time T (>1 year).
79
Unit 4 Life Tables and K-values
Contents
4.1 Introduction
4.3 Life Tales
4.4 K-values
4.5 Summary
4.1 Introduction
A population’s growth potential has much to do with how often

individual members reproduce. Some species (e.g., most invertebrates)
have only one reproductive event in their lifetime, while others (e.g.,
most birds and mammals) are capable of multiple events over an
extended portion of their lives. The former are called semelparous and
the latter, iteroparous life cycles. There is a large amount of variation,
however, within these broad categories. For example, some semelparous
species have overlapping generations of young so that, at any one time,
there may one-, two-, and three-year-old individuals present in the
population. A common form of semelparity in insects of temperate
regions is an annual species.
In this case, the insect overwinters as an egg or larval resting stage until
spring, then grows throughout the warm months and emerges into the
reproductive adult. Adults mate and lay eggs that, again, remain dormant
throughout the winter. Still other semelparous species complete several
generations each summer. It is easy to imagine, then, how the frequency
of reproductive events, the number of young produced in each event, and
the length of each generation can greatly influence how fast a population
can grow.
At the end of this unit the student should be able to

 explain with examples the different type of life tables and life
expectancy in hypothetical population
80
4.3 Life Table
A life table is a record of survival and reproductive rates in a population,

broken out by age, size, or developmental stage (e.g., egg, hatchling,
juvenile, adult). Wildlife population ecologists have found life tables
useful in understanding patterns and causes of mortality, predicting the
future growth or decline of populations, and managing populations of
Threaten & Endangered species. Constructing a life table is often a
simple method for keeping track of births, deaths, and reproductive
output in a population of interest. Basically, there are three methods of
constructing such a table:
1. the cohort life table follows a group of same-aged individuals

from birth (or fertilized eggs) throughout their lives,
2. a static life table is made from data collected from all ages at one
particular time it assumes the age distribution is stable from
generation to generation, and
3. a life table can be made from mortality data collected from a
specified time period and also assumes a stable age distribution.
Note: For organisms that have separate sexes, life tables frequently
follow only female individuals.
Predicting the growth and decline of a wildlife population is an important

application of life table use. As you might expect, whether the population
increases or decreases depends in part on reproductive rate (i.e., litter or
clutch size) and longevity, especially how long the females produce
young. But it may surprise you to learn that population growth or decline
is also driven by the age at which the females begin to have their young.
A major part of this exercise will explore the effects of changing patterns
of survival and reproduction on population dynamics
Another use of life tables is in species conservation efforts, such as in the

case of loggerhead sea turtles (Caretta caretta), which occur off the coast
of the southeastern United States. We will explore loggerhead sea turtle
population dynamics in more detail in problem set 3, but generally
speaking, loggerhead populations are declining in part because egg and
hatchling mortality is high. This situation has led conservation biologists
to advocate the protection of nesting beaches. When these measures
proved ineffective in halting the population decline, compiling and
analyzing a life table for loggerheads indicated that reducing mortality
of older turtles would have a greater probability of reversing the
population decline. Therefore, management 2 efforts shifted to
persuading fishermen to install turtle exclusion devices (TEDs) on their
81
nets to prevent older turtles from drowning. Examples of life tables are
shown below. What is a Life table?
i.Stage-dependent life –table
Stage-dependent life tables are used mainly for insects and other
terrestrial invertebrates. Stage-dependent life tables are built in the
cases when;
1. The life-cycle is partitioned into distinct stages (e.g., eggs,

larvae, pupae and adults in insects)
2. Survival and reproduction depend more on organism stage
rather than on calendar age
3. Age distribution at particular time does not matter (e.g., there is
only one generation per year)
Specific features of stage-dependent life tables are as follow:
1. There is no reference to calendar time. This is very convenient

for the analysis of poikilothermous organisms.
2. Gypsy moth development depends on temperature but the life
table is relatively independent from weather.
3. Mortality processes can be recorded individually and thus, this
kind of life table has more biological information than age-
dependent life tables.
Table 1: Life table of Gypsy moth (Lymantria dispar L.) in New

England (modified from Campbell 1981)
Stage Mortality Initial No. of Mortality Survival K-value

Factor no. of deaths (d) (s) {In(s)}
insect
Egg Predation, etc. 450.0 67.5 0.150 0.850 0.1625
Egg Parasites 382.5 67.5 0.176 0.824 0.1942
Larvae I-III Dispersion, etc. 315.0 157.5 0.500 0.500 0.6932
Larvae IV- Predation, etc. 157.5 118.1 0.750 0.250 1.3857
VI
Larvae IV- Disease 39.4 7.9 0.201 0.799 0.2238
VI
Larvae IV- Parasites 31.5 7.9 0.251 0.749 0.2887
VI
Prepupae Desiccation,etc. 23.6 0.7 0.030 0.970 0.0301
82
Pupae Predation 22.9 4.6 0.201 0.799 0.2242

Pupae Other 18.3 2.3 0.126 0.874 0.1343
Adults Sex ratio 16.0 5.6 0.350 0.650 0.4308
Adult 10.4
females
TOTAL 439.6 97.69 0.0231 3.7674
What are the specific features of stage- dependent life table?
ii.Static (Vertical) Life Table Based on Living Individuals
Most organisms have more complex life histories than those found in the
above example, and while it is possible to follow a single cohort from
birth to death, it is often too costly or time-consuming to do so. Another,
less accurate, method is the static, or vertical, life table. Rather than
following a single cohort, the static table compares population size from
different cohorts, across the entire range of ages, at a single point in time.
Static tables make two important assumptions: 1) the population has a
stable age structure Ñ that is, the proportion of individuals in each age
class does not change from generation to generation, and 2) the
population size is, or nearly, stationary.
iii.Static (Vertical) Life Table Based on Mortality Records
Static life tables can also be made from knowing, or estimating, age at
death for individuals from a population. This can be a useful technique
for secretive large mammals (e.g., moose) from temperate regions where
it is difficult to sample the living members. Because the highest mortality
of large herbivores occurs during the winter, an early spring survey of
carcasses from starvation and predator kills can yield useful information
in constructing a life table. Keep in mind, however, that all static tables
suffer from the same two assumptions stated above.
Because we keep good birth and death records on humans, static life
tables can also be used to answer questions concerning our populations.
For instance, we know that females today have a larger mean life
expectancy than men. But, if we may ask, was this true for our
population 100 years ago? We can use data collected from cemetery
grave markers to construct a static life table and reveal interesting
features of human populations from past generations. The following data
were collected from a random sample of 30 females and 30 males off
grave markers located in an Ann Arbor cemetery.
83
Table 2. Male and female age at death frequencies from a random

sample of 60 Ann Arbor grave markers of individuals born prior to
1870. (From G. Belovsky, unpubl.)
Age at death Females Males
0-5 1 2
6-10 0 0
11-15 1 0
16-20 2 1
21-25 1 1
26-30 0 2
31-35 0 0
36-40 1 2
41-45 1 0
46-50 2 1
51-55 1 0
56-60 2 3
61-65 0 4
66-70 0 4
71-75 1 3
76-80 6 1
81-85 4 1
86-90 7 3
91-95 0 0
96-100 0 2
Besides R0, the basic reproductive rate, several other population

characteristics can be determined from life tables. Some of the most
common features are the cohort generation time (Tx), life expectancy
(ex), and the intrinsic growth rate (r). Cohort generation time is quite
easy to obtain from our first example, a semelparous annual life cycle
(Tx = 1 year), but generation time is less obvious for more complex life
cycles. Generation time can be defined as the average length of time
between when an individual is born and the birth of its offspring.
Therefore, it can be calculated by summing all the lengths of time to
offspring production for the entire cohort divided by the total offspring
produced by the survivors:
Life expectancy is a useful way of expressing the probability of living

(x) number of years beyond a given age. We usually encounter life
expectancy in newspaper articles comparing the mean length of life for
individuals of various populations. However, this value is actually the
life expectancy at birth. One can also calculate the mean length of life
84
beyond any given age for the population. Life expectancy is a somewhat
complicated calculation. Because lx is only the proportion surviving to
the beginning of a particular age class, we must first calculate the average
proportion alive at that age (Lx):
Lx = lx + lx + 1
-----------
----
2
Next, the total number of living individuals at age x and beyond (Tx) is:
Tx = Lx + Lx + 1 +¼+ Lx + n
Finally, the average amount of time yet to be lived by members surviving

to a particular age (ex) is:
Ex
= Tx
-------
lx
The following example shows life expectancy changes in a hypothetical

population that experienced 50% mortality at each age:
Table3. Life expectancy in a hypothetical population.
Age (years) Ix Lx Tx Ex (years)

0 1.0 0.75 1.375 1.375
1 0.5 0.375 0.625 1.25
2 0.25 0.1875 0.25 1.0
3 0.125 0.0625 0.0625 0.5
4 0.0 - - -
The basic reproduction rate (R0) converts the initial population size to
the new size one generation later as:
N T = N 0 x R0
If R0 remains constant from generation to generation, then we can also
use it to predict population size several generations in the future. To
predict population size at any future time, it is more convenient to use a
parameter that already takes generation time into account. This term is
85
ÔrÕ, the intrinsic rate of natural increase, and it can be calculated (or
approximated for complex life cycles) by the following equation:
r @ lnR0
-----
Tc
The term, r, is used in mathematical models of population growth
discussed later
How is Static (Vertical) Life table on Mortality records estimated?

1. What is life expectancy?
4.4 K-value
K-value is just another measure of mortality. The major advantage of k-

values as compared to percentages of died organisms is that k-values are
additive: the k-value of a combination of independent mortality
processes is equal to the sum of k-values for individual processes.
Mortality percentages are not additive. For example, if predators alone

can kill 50% of the population, and diseases alone can kill 50% of the
population, then the combined effect of these process will not result in
50+50 = 100% mortality. Instead, mortality will be 75%
Survival is a probability to survive, and thus we can apply the theory of
probability. In this theory, events are considered independent if the
probability of the combination of two events is equal to the product of
the probabilities of each individual event. In our case event is survival.
If two mortality processes are present, then organism survives if it
survives from each individual process. For example, an organism
survives if it was simultaneously not infected by disease and not captured
by a predator.
Assume that survival from one mortality source is s1 and survival from
the second mortality source is s2. Then survival from both processes,
s12, (if they are independent) is equal to the product of s1 and s2:
This is a "survival multiplication rule". If survival is replaced by 1

minus mortality [s=(1-d)], then this equation becomes:
86
For example, if mortality due to predation is 60% and mortality due to

diseases is 30%, then the combination of these two death processes
results in mortality of d = 1-(1-0.6)(1-0.3)=0.72 (=72%).
Varley and Gradwell (1960) suggested to measure mortality in k-value
which is the negative logarithms of survival:
k = -ln(s)
We use natural logarithms (with base e=2.718) instead of logarithms

with base 10 used by Varley and Gradwell. The advantages of using
natural logarithms will be shown below. It is easy to show that k-values
are additive:
The k-values for the entire life cycle (K) can be estimated as the sum of
k-values for all mortality processes:
In the life table of the gypsy moth (see above), the sum of all k-values
(K = 3.7674) was equal to the k-value of total mortality.
Fig 1: Showing the relations between mortality and the K-value
From the above, when mortality is low, then the k-value is almost equal
to mortality. This is the reason why the k-value can be considered as
another measure of mortality. However, at high mortality, the k-value
grows much faster than mortality. Mortality cannot exceed 1, while the
k-value can be infinitely large.
The following example shows that the k-value represents mortality better
than the percentage of dead organisms: One insecticide kills 99% of
cockroaches and another insecticide kills 99.9% of cockroaches. The
87
difference in percentages is very small (<1%). However the second

insecticide is considerably better because the number of survivors is 10
times smaller. This difference is represented much better by k-values
which are 4.60 and 6.91 for the first and second insecticides,
respectively.
If k-values are estimated for a number of years, then the dynamics of k-

values over time can be compared with the dynamics of the generation
K-value. The following graph shows the dynamics of k-values for the
winter moth in Great Britain.
Fig 2: Showing the dynamics of K-values for the winter moth in

Great Britain
It is seen that the dynamics of winter disappearance (k1) is most

resembling the dynamics of total generation K-value. The conclusion
was made that winter disappearance determines the trend in population
numbers (whether the population will grow or decline), and thus, it can
be considered as a "key factor". T
The key-factor analysis was often considered as a substitute for
modeling. It seems so easy to compare time series of k-values and to find
key-factors without the hard work of developing models of ecological
processes. However, reliable predictions can be obtained only from
models.
This critique does not mean that life-tables have no value. Life-tables are
very important for gathering information about ecological processes
which is necessary for building models. It is the key-factor analysis that
has little sense.
88
A population that experience constant mortality during a specific stage

(e.g., larval stage of insects) change in numbers according to the
exponential model with a negative rate r. We cannot call r intrinsic rate
of natural increase because this term is used for the entire life cycle, and
here we discuss a particular stage in the life cycle. According to the
exponential model:
Population numbers decrease and thus, Nt < N0. Survival is: s = Nt/N0 .
Now we can estimate the k-value:
k = -r t
Instantaneous mortality rate, m, is equal to the negative exponential

coefficient because mortality is the only ecological process considered
(there is no reproduction):
m = -r,
k=mt
Exponential coefficient r is negative (because population declines), and

mortality rate, m, is positive.
We proved that if mortality rate is constant, then k-value is equal to the

instantaneous mortality rate multiplied by time. This is analogs to
physics: distance is equal to speed multiplied by time. Here,
instantaneous mortality rate is like speed, and k-value is like distance. K-
value shows the result of killing organisms with specific rate during a
period of time. If the period of time when mortality occurs is short then
the effect of this mortality on population is not large.
If instantaneous mortality rate changes with time, then the k-value is

equal to its integral over time. In the same way, in physics, distance is
the integral of instantaneous speed over time.
Example. Annual mortality rates of oak trees due to animal-caused bark

damage are 0.08 in the first 10 years and 0.02 in the age interval of 10-
20 years. We need to estimate total k-value (k) and total mortality (d) for
the first 20 years of oak growth.
k = 0.08 × 10 + 0.02 × 10 = 1.0
d = 1 – exp (-k) = 0.63
Thus, total mortality during 20 years is 63%.
89
The limitation of the k-value concept is that all organisms are assumed
to have equal dying probabilities. In nature, dying probabilities may vary
because of spatial heterogeneity and individual variation (both inherited
and non-inherited).
i) Estimation of k-values in natural populations
Estimation of k-values for individual death processes is difficult because

these processes often go simultaneously. The problem is to predict what
mortality could be expected if there was only one death process. In order
to separate death processes it is important to know the biology of the
species and its interactions with natural enemies. Below you can find
several examples of separation of death processes.
Example 1. Insect parasitoids oviposit on host organisms. Parasitoid

larva hatches from the egg and starts feeding on host tissue. Parasitized
host can be alive for a long period. Finally, it dies and parasitoid emerges
from it. Insect predators usually don't distinguish between parasitized
and non-parasitized prey. If an insect was killed by a predator, then it is
usually impossible to detect if this insect was parasitized before. Thus,
mortality due to predation is estimated as the ratio of the number of
insects numbers destroyed by predators to the total number of insects,
whereas mortality due to parasitism is estimated as the ratio of the
number of insects killed by parasitoids to the number of insects that
survived predation. In this example, predation masks the effect of
parasitism, and thus, insects killed by predators are ignored in the
estimation of the rate of parasitism. The effect is the same as if predation
occurred before parasitism in the life cycle. Thus, in the gypsy moth life
table, predation was always considered before parasitism. Diseases also
mask the effect of parasitism and thus they are considered before
parasitism.
Example 2. It is often possible to distinguish between organisms

destroyed by different kinds of predators. For example, small mammals
and birds open sawfly cocoons in a different way. Suppose, 20% of
cocoons were opened by birds, 50% were opened by mammals, and
remaining 30% were alive. The question is what would be the rate of
predation if birds and mammals were acting alone. We assume that
sawfly cocoons have no individual variation in predator attack rate, and
that cocoons destroyed by one predator cannot be attacked by another
predator. First, we estimate total k-value for both predator groups: k12 =
-ln(0.3) = 1.204. Second, we subdivide the total k-value into two portions
proportionally to the number of cocoons destroyed by each kind of
predator. Thus, for birds k1 = 1.204×20/(20+50) = 0.344, and for
mammals k2 = 1.204×50/(20+50) = 0.860. The third step is to convert k-
values into expected mortality if each predator was alone: for birds d1 =
90
1- exp. (-0.344) = 0.291, and for mammals d2 = 1- exp. (-0.860) = 0.577.

What is K-value?
What are the limitations of the k-values concept?
1. Why is the estimation of K-values in natural population
difficult?
4.5 Summary
K-values analysis has been applied to a variety of animal species in order

to assess the role of natural enemies in population fluctuations. In general,
this technique is not applicable to tropical insects because in most species
the generations overlap.

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sciences: Biographical supplement (Vol. 18). New York:
Macmillan.
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Answer to SAEs 1
1. Life expectancy is a useful way of expressing the probability of
living (x) number of years beyond a given age.
Answer to SAEs 2
1. Estimation of k-values for individual death processes is difficult
because these processes often go simultaneously. The problem is
to predict what mortality could be expected if there was only one
death process. In order to separate death processes it is important
to know the biology of the species and its interactions with
natural enemies
93
Contents
5.1 Introduction
5.3 Key-Factor Analysis
5.3.1 Type of factor analysis
5.4 Summary
5.1 Introduction
Key-factor analysis has been applied to a variety of animal species in

order to assess the role of natural enemies in population fluctuations. In
general, this technique is not applicable to tropical insects because in most
species the generations overlap. However, a unique feature in the life
system of Andraca bipunctata Walker is that it has four generations that
are fairly well differentiated in a year, and it does not have a fixed
seasonal peak. These features make the species amenable to analysis
using life-tables.
At the end of this unit the student should be able to
 explain with examples the different type of Factor Analysis

and crieteria for determining of number of factors.
5.3 Factor Analysis
Factor analysis is a statistical method used to describe variability among

observed variables in terms of a potentially lower number of unobserved
variables called factors. In other words, it is possible, for example, that
variations in three or four observed variables mainly reflect the variations
in a single unobserved variable, or in a reduced number of unobserved
variables. Factor analysis searches for such joint variations in response
to unobserved latent variables. The observed variables are modelled as
linear combinations of the potential factors, plus "error" terms. The
information gained about the interdependencies between observed
94
variables can be used later to reduce the set of variables in a dataset.

Factor analysis originated in psychometrics, and is used in behavioural
sciences, social sciences, marketing, product management, operations
research, and other applied sciences that deal with large quantities of
data.
Factor analysis is related to principal component analysis (PCA), but the

two are not identical. Because PCA performs a variance-maximizing
rotation of the variable space, it takes into account all variability in the
variables. In contrast, factor analysis estimates how much of the
variability is due to common factors ("communality"). The two methods
become essentially equivalent if the error terms in the factor analysis
model (the variability not explained by common factors, see below) can
be assumed to all have the same variance.
,
Definition
Suppose we have a set of p observable random variables,
with means .
Suppose for some unknown constants lij and k unobserved random
variables Fj, where
and , where k < p,
we have
Here, they are independently distributed error terms with zero mean
and finite variance, which may not be the same for all i Let
so that we have
In matrix terms, we have
If we have n observations, then we will have the dimensions

, and . Each column of x and F denote values for one particular
observation, and matrix L does not vary across observations.
Also we will impose the following assumptions on F.

1. F and are independent.
2. E(F) = 0
3. Cov(F) = I
Any solution of the above set of equations following the constraints for F
is defined as the factors, and L as the loading matrix.
95
Suppose Cov(x − µ) = Σ. Then note that from the conditions just imposed
on F, we have
or
or
Note that for any orthogonal matrix Q if we set L = LQ and F = QTF, the
criteria for being factors and factor loadings still hold. Hence a set of
factors and factor loadings is identical only up to orthogonal
transformations.
The following example is a fictionalized simplification for expository

purposes, and should not be taken as being realistic. Suppose a
psychologist proposes a theory that there are two kinds of intelligence,
"verbal intelligence" and "mathematical intelligence", neither of which
is directly observed. Evidence for the theory is sought in the examination
scores from each of 10 different academic fields of 1000 students. If each
student is chosen randomly from a large population, then each student's
10 scores are random variables. The psychologist's theory may say that
for each of the 10 academic fields, the score averaged over the group of
all students who share some common pair of values for verbal and
mathematical "intelligences" is some constant times their level of verbal
intelligence plus another constant times their level of mathematical
intelligence, i.e., it is a linear combination of those two "factors". The
numbers for a particular subject, by which the two kinds of intelligence
are multiplied to obtain the expected score, are posited by the theory to
be the same for all intelligence level pairs, and are called "factor
loadings" for this subject. For example, the theory may hold that the
average student's aptitude in the field of amphibiology is
{10 × the student's verbal intelligence} + {6 × the student's

mathematical intelligence}.
The numbers 10 and 6 are the factor loadings associated with

amphibiology. Other academic subjects may have different factor
loadings.
Two students having identical degrees of verbal intelligence and
identical degrees of mathematical intelligence may have different
aptitudes in amphibiology because individual aptitudes differ from
average aptitudes. That difference is called the "error" — a statistical
term that means the amount by which an individual differs from what is
average for his or her levels of intelligence (see errors and residuals in
statistics).
96
The observable data that go into factor analysis would be 10 scores of

each of the 1000 students, a total of 10,000 numbers. The factor loadings
and levels of the two kinds of intelligence of each student must be
inferred from the data.
Mathematical model of the same example
In the example above, for i = 1, ..., 1,000 the ith student's scores are
where
1. xk,i is the ith student's score for the kth subject
2. µ k is the mean of the students' scores for the kth subject (assumed
to be zero, for simplicity, in the example as described above,
which would amount to a simple shift of the scale used)
3. vi is the ith student's "verbal intelligence",
4. mi is the ith student's "mathematical intelligence",
5. are the factor loadings for the kth subject, for j = 1, 2.
6. εk,i is the difference between the ith student's score in the kth
subject and the average score in the kth subject of all students
whose levels of verbal and mathematical intelligence are the same
as those of the ith student,
In matrix notation, we have
Where:
1. N is 1000 students
2. X is a 10 × 1,000 matrix of observable random variables,
3. µ is a 10 × 1 column vector of unobservable constants (in this
case "constants" are quantities not differing from one individual
student to the next; and "random variables" are those assigned
to individual students; the randomness arises from the random
way in which the students are chosen),
4. L is a 10 × 2 matrix of factor loadings (unobservable constants,
ten academic topics, each with two intelligence parameters that
determine success in that topic),
97
5. F is a 2 × 1,000 matrix of unobservable random variables (two

intelligence parameters for each of 1000 students),
6. ε is a 10 × 1,000 matrix of unobservable random variables.
Observe that by doubling the scale on which "verbal intelligence"—the

first component in each column of F—is measured, and simultaneously
halving the factor loadings for verbal intelligence makes no difference to
the model. Thus, no generality is lost by assuming that the standard
deviation of verbal intelligence is 1. Likewise for mathematical
intelligence. Moreover, for similar reasons, no generality is lost by
assuming the two factors are uncorrelated with each other. The "errors"
ε are taken to be independent of each other. The variances of the "errors"
associated with the 10 different subjects are not assumed to be equal.
Note that, since any rotation of a solution is also a solution, this makes
interpreting the factors difficult. See disadvantages below. In this
particular example, if we do not know beforehand that the two types of
intelligence are uncorrelated, then we cannot interpret the two factors as
the two different types of intelligence. Even if they are uncorrelated, we
cannot tell which factor corresponds to verbal intelligence and which
corresponds to mathematical intelligence without an outside argument.
The values of the loadings L, the averages µ, and the variances of the
"errors" ε must be estimated given the observed data X and F (the
assumption about the levels of the factors is fixed for a given F). What
is Factor Analysis?
5.3.1 Types of Factor Analysis
There are two types of factor analysis and are listed below:
1. Exploratory factor analysis: (EFA) is used to uncover the

underlying structure of a relatively large set of variables. The researcher's
a priori assumption is that any indicator may be associated with any
factor. This is the most common form of factor analysis. There is no prior
theory and one uses factor loadings to intuit the factor structure of the
data.
2. Confirmatory factor analysis: (CFA) seeks to determine if the

number of factors and the loadings of measured (indicator) variables on
them confirm to what is expected on the basis of pre-established theory.
Indicator variables are selected on the basis of prior theory and factor
analysis is used to see if they load as predicted on the expected number
of factors. The researcher's à priori assumption is that each factor (the
number and labels of which may be specified à priori) is associated with
a specified subset of indicator variables. A minimum requirement of
98
confirmatory factor analysis is that one hypothesizes beforehand the

number of factors in the model, but usually also the researcher will posit
expectations about which variables will load on which factors. The
researcher seeks to determine, for instance, if measures created to
represent a latent variable really belong together.
i) Different Factoring in Factor analysis
1. Principal component analysis (PCA): The most common form of

factor analysis, PCA seeks a linear combination of variables such
that the maximum variance is extracted from the variables. It then
removes this variance and seeks a second linear combination which
explains the maximum proportion of the remaining variance, and
so on. This is called the principal axis method and results in
orthogonal (uncorrelated) factors.
2. Canonical factor analysis (CFA), also called Rao's canonical
factoring, is a different method of computing the same model as
PCA, which uses the principal axis method. CFA seeks factors
which have the highest canonical correlation with the observed
variables. CFA is unaffected by arbitrary rescaling of the data.
3. Common factor analysis, also called principal factor analysis
(PFA) or principal axis factoring (PAF), seeks the least number of
factors which can account for the common variance (correlation)
of a set of variables.
4. Image factoring: based on the correlation matrix of predicted
variables rather than actual variables, where each variable is
predicted from the others using multiple regression.
5. Alpha factoring: based on maximizing the reliability of factors,
assuming variables are randomly sampled from a universe of
variables. All other methods assume cases to be sampled and
variables fixed.
ii) Terminology used in Factor analysis
1. Factor loadings: The factor loadings, also called component

loadings in PCA, are the correlation coefficients between the
variables (rows) and factors (columns). Analogous to Pearson's r,
the squared factor loading is the percent of variance in that
indicator variable explained by the factor. To get the percent of
variance in all the variables accounted for by each factor, add the
sum of the squared factor loadings for that factor (column) and
divide by the number of variables. (Note the number of variables
equals the sum of their variances as the variance of a standardized
variable is 1.) This is the same as dividing the factor's eigenvalue
by the number of variables.
99
2. Interpreting factor loadings: By one rule of thumb in confirmatory

factor analysis, loadings should be .7 or higher to confirm that
independent variables identified a priori are represented by a
particular factor, on the rationale that the .7 level corresponds to
about half of the variance in the indicator being explained by the
factor. However, the .7 standard is a high one and real-life data may
well not meet this criterion, which is why some researchers,
particularly for exploratory purposes, will use a lower level such as
.4 for the central factor and .25 for other factors call loadings above
.6 "high" and those below .4 "low". In any event, factor loadings
must be interpreted in the light of theory, not by arbitrary cut off
levels.
In oblique rotation, one gets both a pattern matrix and a structure matrix.
The structure matrix is simply the factor loading matrix as in orthogonal
rotation, representing the variance in a measured variable explained by a
factor on both a unique and common contributions basis. The pattern
matrix, in contrast, contains coefficients which just represent unique
contributions. The more factors, the lower the pattern coefficients as a rule
since there will be more common contributions to variance explained. For
oblique rotation, the researcher looks at both the structure and pattern
coefficients when attributing a label to a factor.
3. Communality (h2): The sum of the squared factor loadings for all
factors for a given variable (row) is the variance in that variable
accounted for by all the factors, and this is called the communality.
The communality measures the percent of variance in a given
variable explained by all the factors jointly and may be interpreted
as the reliability of the indicator.
4. Spurious solutions: If the communality exceeds 1.0, there is a
spurious solution, which may reflect too small a sample or the
researcher has too many or too few factors.
5. Uniqueness of a variable: 1-h2. That is, uniqueness is the
variability of a variable minus its communality.
6. Eigenvalues:/Characteristic roots: The eigenvalue for a given
factor measures the variance in all the variables which is accounted
for by that factor. The ratio of eigenvalues is the ratio of
explanatory importance of the factors with respect to the variables.
If a factor has a low eigenvalue, then it is contributing little to the
explanation of variances in the variables and may be ignored as
redundant with more important factors. Eigenvalues measure the
amount of variation in the total sample accounted for by each
factor.
7. Extraction sums of squared loadings: Initial eigenvalues and
eigenvalues after extraction (listed by SPSS as "Extraction Sums
100
of Squared Loadings") are the same for PCA extraction, but for
other extraction methods, eigenvalues after extraction will be lower
than their initial counterparts. SPSS also prints "Rotation Sums of
Squared Loadings" and even for PCA, these eigenvalues will differ
from initial and extraction eigenvalues, though their total will be
the same.
8. Factor scores: Also called component scores in PCA, factor
scores are the scores of each case (row) on each factor (column).
To compute the factor score for a given case for a given factor, one
takes the case's standardized score on each variable, multiplies by
the corresponding factor loading of the variable for the given
factor, and sums these products. Computing factor scores allows
one to look for factor outliers. Also, factor scores may be used as
variables in subsequent modelling
iii) Criteria for determining the number of factors
1. Comprehensibility: Though not a strictly mathematical criterion,

there is much to be said for limiting the number of factors to those
whose dimension of meaning is readily comprehensible. Often this
is the first two or three. Using one or more of the methods below,
the researcher determines an appropriate range of solutions to
investigate. For instance, the Kaiser criterion may suggest three
factors and the scree test may suggest 5, so the researcher may
request 3-, 4-, and 5-factor solutions and select the solution which
generates the most comprehensible factor structure.
2. Kaiser criterion: The Kaiser rule is to drop all components with
eigenvalues under 1.0. The Kaiser criterion is the default in SPSS
and most computer programs but is not recommended when used
as the sole cut-off criterion for estimating the number of factors.
3. Scree plot: The Cattell scree test plots the components as the X
axis and the corresponding eigenvalues as the Y-axis. As one
moves to the right, toward later components, the eigenvalues drop.
When the drop ceases and the curve makes an elbow toward less
steep decline, Cattell's scree test says to drop all further
components after the one starting the elbow. This rule is sometimes
criticised for being amenable to researcher-controlled "fudging".
That is, as picking the "elbow" can be subjective because the curve
has multiple elbows or is a smooth curve, the researcher may be
tempted to set the cut-off at the number of factors desired by his or
her research agenda.
4. Horn's Parallel Analysis (PA): A Monte-Carlo based simulation
method that compares the observed eigenvalues with those
obtained from uncorrelated normal variables. A factor or
component is retained if the associated eigenvalue is bigger than
the 95th of the distribution of eigenvalues derived from the random
101
data. PA is one of the most recommendable rules for determining

the number of components to retain, but only few programs include
this option.
5. Variance explained criteria: Some researchers simply use the
rule of keeping enough factors to account for 90%; (sometimes
80%) of the variation. Where the researcher's goal emphasizes
parsimony (explaining variance with as few factors as possible),
the criterion could be as low as 50%
Before dropping a factor below one's cut-off, however, the researcher

should check its correlation with the dependent variable. A very small
factor can have a large correlation with the dependent variable, in which
case it should not be dropped
iv) Different types of Rotation methods
Rotation serves to make the output more understandable and is usually

necessary to facilitate the interpretation of factors.
1. Varimax rotation is an orthogonal rotation of the factor axes to

maximize the variance of the squared loadings of a factor
(column) on all the variables (rows) in a factor matrix, which has
the effect of differentiating the original variables by extracted
factor. Each factor will tend to have either large or small loadings
of any particular variable. A varimax solution yields results which
make it as easy as possible to identify each variable with a single
factor. This is the most common rotation option.
2. Quartimax rotation is an orthogonal alternative which minimizes
the number of factors needed to explain each variable. This type
of rotation often generates a general factor on which most
variables are loaded to a high or medium degree. Such a factor
structure is usually not helpful to the research purpose.
3. Equimax rotation is a compromise between Varimax and
Quartimax criteria.
4. Direct oblimin rotation is the standard method when one wishes
a non-orthogonal (oblique) solution – that is, one in which the
factors are allowed to be correlated. This will result in higher
eigenvalues but diminished interpretability of the factors.
5. Promax rotation is an alternative non-orthogonal (oblique)
rotation method which is computationally faster than the direct
oblimin method and therefore is sometimes used for very large
datasets. What is Principal component analysis?
102
1. What is Factor Loading?
2. What is Eigenvalues:/Characteristic roots?
5.4 Summary
Key-factor analysis has been applied to a variety of animal species in

order to assess the role of natural enemies in population fluctuations. In
general, this technique is not applicable to tropical insects because in
most species the generations overlap.

Bryant and Yarnold (1994). "Principal components analysis and

exploratory and confirmatory factor analysis". In: Grimm and
Yarnold, Reading and understanding multivariate analysis.
American Psychological Association Books. ISBN 978-1-
55798273-5.

Gorsuch, R. L. (1983) Factor Analysis. Hillsdale, NJ: Lawrence

Erlbaum. ISBN 089859-202-X
Sheppard, A. G. (1996). The sequence of factor analysis and cluster

analysis: Differences in segmentation and dimensionality through
the use of raw and factor scores. Tourism Analysis, 1 , 49-57.
Sternberg, R.J.(1990). The geographic metaphor. In R.J. Sternberg,

Metaphors of mind: Conceptions of the nature of intelligence (pp.
85–111). New York: Cambridge.
103
Stills, D.L. (Ed.). (1989). International encyclopedia of the social

sciences: Biographical supplement (Vol. 18). New York:
Macmillan.
https://www.studocu.com/en-ie/document/university-college-
dublin/act-maths-1/3-the-life-table-lecture-notes/1695739
populations/238151
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104
Answers to SAEs 1
1. The factor loadings, also called component loadings in PCA, are
the correlation coefficients between the variables (rows) and
factors (columns). Analogous to Pearson's r, the squared factor
loading is the percent of variance in that indicator variable
explained by the factor. To get the percent of variance in all the
variables accounted for by each factor, add the sum of the squared
factor loadings for that factor (column) and divide by the number
of variables. (Note the number of variables equals the sum of their
variances as the variance of a standardized variable is 1.) This is
the same as dividing the factor's eigenvalue by the number of
variables.
2. The eigenvalue for a given factor measures the variance in all the
variables which is accounted for by that factor. The ratio of
eigenvalues is the ratio of explanatory importance of the factors
with respect to the variables. If a factor has a low eigenvalue, then
it is contributing little to the explanation of variances in the
variables and may be ignored as redundant with more important
factors. Eigenvalues measure the amount of variation in the total
sample accounted for by each factor
Glossary
cm = centimeters
CO2 = Carbondioxide
CH4 = methane (CH4)
ft = feets
Hz = Hertz
in = inches
Ib = pounds
Kgs = kilograms.
km = kilometers
m = meters
mm = millimeters
O2 = Oxygen
oz = ounce
105

L = length
% = percentage
g = grams
spp = species
1. What is Population density?

2. State the two growth pattern in animal population?
3. What is migration in animal?
4. What is survivorship curve?
5. What is Life table?
106
Module 3 Population Cycle in Animal Ecology
Module Introduction
Introduce the module and state the units under the module.

GLossary
Contents
1.1 Introduction
1.3 Competition in Animal
1.3.1 Types of competition
1.3.2 Resources contributing to competition among organisms
1.3.3 Evolutionary strategies
1.4 Summary
1.1 Introduction
Competition is 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 in ecology, especially community ecology.
Fig 1: Showing Hyena competing for food (Resources)
107
At the end of this unit, students should be able to:
 Define competition?
 List factors/resources that contribute to competition among
organisms
 Explain evolution strategies and state the formula selection theory
1.3 Competition in Animal
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 straightforward, 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, however, competition
may play less of a role than expansion among larger groups such as
families
1.3.1 Types of Competition
1.3.1.1Competiton 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.
Fig 2: Male-male competition in red deer during rut is an example of

interference competition within a species.
108
1. 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.
2. Exploitation competition Occurs indirectly through a common

limiting resource which acts as an intermediate. For example, use
of resources depletes the amount available to others, or they
compete for space. Also known as exploitative competition.
3. 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 prey of 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
1.3.1.2Competition by Species
1. Intraspecific competition: Intraspecific competition occurs when

members of the same species vie for the same resources in an
ecosystem. 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. Although in this situation it
may actually be more useful to think in terms of resource
availability than competition. Adaptations to such an environment
include growing taller, (where the specific prediction provided by
the competition model is that all species in such a situation will
grow as tall as possible) or developing a larger root system (where
the specific prediction is that all species in the system will develop
very deep root systems).
2. Interspecific compeptition: 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, however they still
persist together, despite the prediction that under competition one will
displace the other. In fact, lions sometimes steal prey items killed by
109
cheetahs. Potential competitors can also kill each other, and this
phenomenon is called 'intraguild predation'. For example, in southern
California coyotes often kill and eat gray foxes and bobcats, all three
carnivores sharing the same stable prey (small mammals).
Competition has been observed between individuals, populations and

species, but there is little evidence that competition has been the driving
force in the evolution of large groups. For example, between reptiles and
mammals. Mammals lived beside reptiles for many millions of years of
time but were unable to gain a competitive edge until dinosaurs were
devastated by the K-T Extinction. What is Competition?
Define Apparent competition.
1.3.2 Resources contributing to competition among organisms
1.3.2.1 Territory (Animal)
In ethology the term territory refers to any sociographical area that an

animal of a particular species consistently defends against conspecifics
(and, occasionally, animals of other species). Animals that defend
territories in this way are referred to as territorial.
1.3.2.2 Classic territories
Territorial animals defend areas that contain a nest, den or mating site
and sufficient food resources for themselves and their young. Defence
rarely takes the form of overt fights: more usually there is a highly
noticeable display, which may be visual (as in the red breast of the robin),
auditory (as in much bird song, or the calls of gibbons) or olfactory,
through the deposit of scent marks. Many territorial mammals use scent-
marking to signal the boundaries of their territories; the marks may be
deposited by urination, by defecation, or by rubbing parts of the bodies
that bear specialised scent glands against the substrate. For example,
dogs and other canids scent-mark by urination and defecation, while cats
scent-mark by rubbing their faces and flanks against objects, as well as
by the notoriously persistently smelly spraying of urine by tomcats.
Many prosimians use territorial marking; for example, the Red-bellied
Lemur creates territories for groups of two to ten individuals in the
rainforests of eastern Madagascar by scent marking: the male Diademed
Sifaka also scent marks defended territories in some of these same
rainforests. The male Western fence lizard defends a territory by
posturing and combat, but less intensely after the mating season.
Invertebrates which show territorality include some ants and bees, and
the owl limpet.
110
1.3.2.3 Spraying
Spraying (also known as territorial marking) is behaviour used by

animals to identify their territory. Most commonly, this is scent marking,
accomplished by depositing strong-smelling chemicals such as urine at
prominent locations within the territory. Often the scent contains carrier
proteins, such as the major urinary proteins, to stabilize the odours and
maintain them for longer.
Not only does the marking communicate to others of the same species,
but it is also noted by prey species and avoided. For example felids such
as leopards and jaguars mark by rubbing themselves against vegetation.
Some prosimians, such as the Red-bellied Lemur, also use scent marking
to establish a territory. Many ungulates, for example the Blue
Wildebeest, use scent marking from two glands, the preorbital gland and
a scent gland in the hoof.
Fig 3: A wolf marking its territory.
1.3.2.4 Defence
Territories may be held by an individual, a mated pair, or a group.

Territoriality is not a fixed property of a species: for example, robins
defend territories as pairs during the breeding season and as individuals
during the winter, while some nectarivores defend territories only during
the mornings (when plants are richest in nectar). In species that do not
form pair bonds, male and female territories are often independent, in the
sense that males defend territories only against other males, and females
only against other females; in this case, if the species is polygynous, one
male territory will probably contain several female territories, while in
some polyandrous species such as the Northern Jacana, this situation is
reversed.
Quite often territories that only yield a single resource are defended. For
example, European Blackbirds may defend feeding territories that are
distant from their nest sites, and in some species that form leks, for
111
example the Uganda kob (a grazing antelope), males defend the lek site
(which is used only for mating)
Territoriality is only shown by a minority of species. More commonly, an

individual or a group of animals will have an area that it habitually uses
but does not necessarily defend; this is called its home range. The home
ranges of different groups often overlap, and in the overlap areas the
groups will tend to avoid each other rather than seeking to expel each
other. Within the home range there may be a core area that no other
individual group uses, but again this is as a result of avoidance rather than
defense.
Behavioural ecologists have argued that food distribution determines

whether a species will be territorial or not. This however, though true as
far as it goes, is too narrow a point of view. As mentioned above, there
are several kinds of territoriality; for example, the defence of lek areas
by kob has nothing to do with food. Many other examples of territorial
defence, including fish, birds or even invertebrates, are related to
competition for mates or safe lairs, rather than food. Territoriality will
emerge where there is a focused resource that provides enough for the
individual or group, within a boundary that is small enough to be
defended without the expenditure of too much effort.
Many birds, particularly seabirds, though they nest in dense

communities, are none the less territorial in that they defend their nesting
site to within the distance that they can reach while brooding. This is
necessary to prevent attacks on their own chicks or nesting material from
neighbours. Commonly the resulting superimposition of the short-range
repulsion onto the long range attraction characteristically leads to the
well-known roughly hexagonal spacing of nests. Interestingly, one gets
a similar hexagonal spacing resulting from the territorial behaviour of
gardening limpets such as species of Scutellastra. They vigorously
defend their gardens of particular species of algae,that extend for perhaps
1–2 cm around the periphery of their shells.
Territoriality is least likely with insectivorous birds, where the food
supply is plentiful but unpredictably distributed. Swifts rarely defend an
area larger than the nest. Conversely, other insectivorous birds that
occupy more constrained territories, such as the ground-nesting
Blacksmith Lapwing may be very territorial, especially in the breeding
season, where they not only threaten or attack many kinds of intruders,
but have stereotyped display behaviour to deter conspecifics sharing
neighbouring nesting spots.
Conversely, large solitary (or paired) carnivores, such as bears and the
bigger raptors require an extensive protected area to guarantee their food
112
supply. This territoriality will only break down when there is a glut of
food, for example when Grizzly Bears are attracted to migrating salmon.
1.3.2.5 Sunlight
Sunlight, in the broad sense, is the total frequency spectrum of

electromagnetic radiation given off by the Sun. On Earth, sunlight is
filtered through the Earth's atmosphere, and solar radiation is obvious as
daylight when the Sun is above the horizon. Sunlight is a key factor in
photosynthesis, a process vital for life on Earth.
Fig 4: Sunlight shining through the large tree in the forest.
Rainforests around the globe have a remarkably consistent pattern of tree

sizes. Now researchers have found that the reason for this structure has
to do with the competition for sunlight after a large tree falls and leaves
an opening in the canopy.
In a new study using data from a rainforest in Panama, researchers

determined that competition for sunlight is the underlying cause of this
common structure, which is observed in rainforests around the globe
despite differences in plant species and geography.
After a large tree falls, many small individuals are able to grow due to
an increase in available sunlight (T=1). Once they have grown to touch
one another (T=2), they begin to overtop one another and leave
individuals behind in the understory (T=3). (below)
113
Fig 5: Showing the effects of sunlight of trees in a rainforest
The researchers found that the rainforest structure stems from what
happens after a tall tree falls and creates a gap in the canopy. The gap
enables sunlight to reach the forest floor and fuel the rapid growth of
small trees. Over time, the trees’ crowns grow to fill the gap until the
point where not all of the trees can fit in the sunlit patch. Some will be
left behind in the shade of their competitors.
Also, Sunlight can only filter through the water at about 30 meters or
100 feet below the water’s surface. Therefore only the upper layer of
the aquatic life zone will support photosynthesis. Sea life that depends
on photosynthesis such as algae can only survive at this level. What is
Territory? What do you understand by a Territory?
1.3.3 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 (1967).
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
illustrated in the simple Verhulst equation of population dynamics:
114
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 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. Explain the concept of r/K
selection theory.
1. What is Intraspecific competition?
2. What is Exploitation competition?
1.4 Summary
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, however, competition
may play less of a role than expansion among larger groups such as
families.
Begon, M.; Harper, J. L.; Townsend, C. R. (1996) Ecology: Individuals,

populations and communities Blackwell Science.
Fedriani, J. M., T. K. Fuller, R. M. Sauvajot and E. C. York. 2000.

Competition and intraguild predation among three sympatric
carnivores. Oecologia, 125:258-270.
MacArthur, R. and Wilson, E. O. (1967). The Theory of Island

Biogeography, Princeton University Press (2001 reprint), ISBN
0-691-08836-5M.
Pianka, E. R. (1970). On r and K selection. American Naturalist '104,

592-597.
115
Sahney, S., Benton, M.J. and Ferry, P.A. (2010). "Links between global
taxonomic diversity, ecological diversity and the expansion of
vertebrates on land" (PDF). Biology Letters 6 (4): 544–547.
doi:10.1098/rsbl.2009.1024. PMID 20106856.
http://rsbl.royalsocietypublishing.org/content/6/4/544.full.pdf+ht
ml.
Verhulst, P. F. (1838). Notice sur la loi que la population pursuit dans

son accroissement. Corresp. Math. Phys. '10, 113-121.
Branch,G.M., Griffiths,C., Beckley,L.E., Branch,M.L.; Two Oceans.
Pub. Struik, 2010. ISBN13: 978-1770077720.
http://www.sspca.org/TerritorialMarking.html
Hurst JL, Robertson DHL, Tolladay U, Beynon RJ (May 1998).

"Proteins in urine scent marks of male house mice extend the
longevity of olfactory signals". Anim Behav 55 (5): 1289–97.
doi:10.1006/anbe.1997.0650. PMID 9632512.
Michael Hogan .C, (2008) "Western fence lizard (Sceloporus

occidentalis)", Globaltwitcher, ed. Nicklas Stromberg .
Stimson J. (1969). "Territoriality of the owl limpet Lottia gigantea".

Ecology 51 (1): 113–118. doi:10.2307/1933604.
http://jstor.org/stable/1933604.
Walther, F. R., E. C. Mungall, G. A. Grau. (1983) Gazelles and their

relatives : a study in territorial behavior Park Ridge, N.J. : Noyes
Publications 239 p. ISBN 0-8155-0928-6
Stokes, A. W. (editor) (1974) Territory Stroudsburg, Pa., Dowden,

Hutchinson & Ross 398 p. ISBN 0-87933-113-5 Klopfer, P. H.
(1969) Habitats and territories; a study of the use of space by
animals New York, Basic Books 117 p. Sunlight
https://www.studocu.com/en-us/document/university-of-
pittsburgh/ecology/competition-1-lecture-notes/10399621
populations/238149
populations/238149
116
https://www.bing.com/videos/search?q=competition+in++animal++cla
ss&&view=detail&mid=AA90EFC74AD8F288FE47AA90EFC
74AD8F288FE47&&FORM=VRDGAR&ru=%2Fvideos%2Fse
arch%3Fq%3Dcompetition%2520in%2520%2520animal%2520
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Manage%2520Your%2520Search%2520History%2525E%26sp
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1%26pq%3Dcompetition%2520in%2520animal%2520class%26
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117
Answers to SAEs 1
1. Intraspecific competition occurs when members of the same
species vie for the same resources in an ecosystem. 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. Although in this situation it may actually be more
useful to think in terms of resource availability than competition.
Adaptations to such an environment include growing taller, (where
the specific prediction provided by the competition model is that
all species in such a situation will grow as tall as possible) or
developing a larger root system (where the specific prediction is
that all species in the system will develop very deep root systems).
2. Exploitation competition Occurs indirectly through a common

limiting resource which acts as an intermediate. For example, use
of resources depletes the amount available to others, or they
compete for space. Also known as exploitative competition.
118
Contents
2.1 Introduction
2.3 Population Cycle
2.4 Summary
2.1 Introduction
A population is a group of individuals of the same species living and

interbreeding within a geographical area. The area that is used to define
the population is such that inter-breeding is possible between any pair
within the area and more probable than cross-breeding with individuals
from other areas. Normally breeding is substantially more common within
the area than across the border.
At the end of this unit, students should be able to define population

cycle and List factors that causes or contribute to population cycle
2.3 Population Cycle
A population cycle in zoology is a phenomenon where populations rise

and fall over a predictable period of time. There are some species where
population numbers have reasonably predictable patterns of change
although the full reasons for population cycles is one of the major
unsolved ecological problems. There are a number of factors which
influence population change such as availability of food, predators,
diseases and climate
Olaus Magnus, the Archbishop of Uppsala in central Sweden, identified

that species of northern rodents had periodic peaks in population and
published two reports on the subject in the middle of the 16th century. In
North America, the phenomenon was identified in populations of the
snowshoe hare. In 1865, trappers with the Hudson's Bay Company were
catching plenty of animals. By 1870, they were catching very few. It was
119
finally identified that the cycle of high and low catches ran over
approximately a ten year period.
The most well-known example of creatures which have a population
cycle is the lemming. The biologist Charles Elton first identified in 1924
that the lemming had regular cycles of population growth and decline.
When their population outgrows the resources of their habitat, lemmings
migrate, although contrary to popular myth, they do not jump into the
sea.
Lemmings have period population booms and then disperse in all

directions, seeking food and shelter their natural habitats cannot provide.
The Norway lemming and brown lemming are two of the few vertebrates
which reproduce so quickly that their population fluctuations are chaotic,
rather than following linear growth to a carrying capacity or regular
oscillations. Why lemming populations fluctuate with such great
variance roughly every four years, before numbers drop to near
extinction, is not known. Lemming behaviour and appearance are
markedly different from those of other rodents, which are
inconspicuously coloured and try to conceal themselves from their
predators. Lemmings, by contrast, are conspicuously coloured and
behave aggressively toward predators and even human observers. The
lemming defence system is thought to be based on
aposematism (warning display). Fluctuations in the lemming population
affect the behaviour of predators, and may fuel irruptions of birds of prey
such as snowy owls to areas further south. For many years, the
population of lemmings was believed to change with the population
cycle, but now some evidence suggests their predators' populations,
particularly those of the stoat, may be more closely involved in changing
the lemming population.
While the phenomenon is often associated with rodents, it does occur in
other species such as the ruffed grouse. There are other species which
have irregular population explosions such as grasshoppers where
overpopulation results in locust swarms in Africa and Australia.
There is also an interaction between prey with periodic cycles and

predators. As the population expands, there is more food available for
predators. As it contracts, there is less food available for predators,
putting pressure on their population numbers.
Define Population cycle.
1. Write on Lemmings as a case study of population cycle?
120
2.4 Summary
There are some species where population numbers have reasonably

predictable patterns of change although the full reasons for population
cycles is one of the major unsolved ecological problems.
Chapman JL and Reiss MJ (2010), Ecology, Principles and Applications, 2nd

ed. Cambridge University Press
Miller GT and Spoolman SE (2009), Essentials of Ecology, 5th ed.

Brooks/Cole, Cengage Learning
Miller GT, Jr. (1988), Environmental Science, 2nd ed. Wadsworth,

Belmont.
National Parks Service Document on population cycles
Odum EP and Barrett GW (2004), Fundamentals of Ecology, 5th ed.

Cengage Learning
Raven PH, Johnson GB, Mason KA, Losos JB, and Singer SR (2011),
Biology, 9th ed. McGraw-Hill.
Smith TM and Smith RL (2009), Elements of Ecology, 7th ed. San

Francisco CA: Pearson Benjamin Cummings.
Paper by Professor Stenseth, University of Oslo on lemming cycles
Paper by Baltensweiler, W. & Fischlin, A., 1988. The larch bud moth in
the Alps
Encyclopædia Britannica Online 25 August 2005 "Population Ecology"

article section on Population Cycles
Errki Korpimaki and Charles J Krebs "Predation and Population Cycles

of Small Mammals" Bioscience November 1996 Volume 46,
Number 10
Alan Berryman, Population Cycles, Oxford University Press US, 2002

ISBN 0-19514098-2
121
Hartl, Daniel (2007). Principles of Population Genetics. Sinauer

Associates. p. 45. ISBN 978-087893-308-2.
https://royalsocietypublishing.org/doi/10.1098/rspb.2017.2841
https://en.wikipedia.org/wiki/Population_cycle
https://www.bing.com/videos/search?q=what+is+Population+cycle+pd
f&&view=detail&mid=113A80778052A4059F23113A80778052A405
9F23&&FORM=VRDGAR&ru=%2Fvideos%2Fsearch%3Fq%3Dwhat
%2Bis%2BPopulation%2Bcycle%2Bpdf%26FORM%3DHDRSC3
https://www.bing.com/videos/search?q=what+is+Population+cycle+pd
f&&view=detail&mid=7B480BC3F49F187389E07B480BC3F49F187
389E0&&FORM=VRDGAR&ru=%2Fvideos%2Fsearch%3Fq%3Dwh
at%2Bis%2BPopulation%2Bcycle%2Bpdf%26FORM%3DHDRSC3
122
Answers to SAEs 1
1. Lemmings have period population booms and then disperse in all
directions, seeking food and shelter their natural habitats cannot
provide. The Norway lemming and brown lemming are two of the
few vertebrates which reproduce so quickly that their population
fluctuations are chaotic, rather than following linear growth to a
carrying capacity or regular oscillations. Why lemming
populations fluctuate with such great variance roughly every four
years, before numbers drop to near extinction, is not
known. Lemming behaviour and appearance are markedly
different from those of other rodents, which are inconspicuously
coloured and try to conceal themselves from their predators.
Lemmings, by contrast, are conspicuously coloured and behave
aggressively toward predators and even human observers. The
lemming defence system is thought to be based on
aposematism (warning display). Fluctuations in the lemming
population affect the behaviour of predators, and may fuel
irruptions of birds of prey such as snowy owls to areas further
south. For many years, the population of lemmings was believed
to change with the population cycle, but now some evidence
suggests their predators' populations, particularly those of the
stoat, may be more closely involved in changing the lemming
population.
123
Contents
3.1 Introduction
3.3 Modelling Predator-Prey Interactions
3.4 Evolutionary Dynamics of Predator-Prey Systems: An Ecological
Perspective
3.5 Summary
3.1 Introduction
There is also an interaction between prey with periodic cycles and

predators. As the population expands, there is more food available for
predators. As it contracts, there is less food available for predators, putting
pressure on their population numbers.
Figure 1: Population cycles in a Swedish forest community

(www.encoguru,com)
The top figure (a) shows changes in population size for voles and small
game. The striped arrows indicate years in which voles consumed tree
124
bark as a marginal food. The bottom figure (b) illustrates how predator
populations change in relation to prey abundance.
Some of the most notable examples of population changes occur in

species that experience large, cyclic swings in population size. Quite
often, these cycles co-occur with population cycles of other species in
the same location. For example, red foxes (Vulpes vulpes) in northern
Sweden prey on voles, grouse, and hares. Studies of these species have
demonstrated linked population cycles in each of the prey species, with
population peaks every 3-4 years (Figure 1). What drives these cycles?
Grouse, hares, and voles feed on vegetation, and the availability of their
preferred foods will influence the population size of each. The
availability of food acts as a bottom-up control that affects population
size. In years when their preferred food items are abundant, populations
will grow. When preferred foods are scarce, individuals must turn to less
desirable foods to prevent starvation. They grow more slowly, reproduce
less, and populations decline. When vole populations peak and
competition for food is strongest, they turn to bark as a marginal food,
and this shift in foraging behaviour coincides with a population decline
(Figure 1a). Grouse and hare populations cycle in a manner comparable
to those of voles, which suggests that food availability plays a role in
regulating populations of these herbivores.
Foxes prefer to consume voles and other small rodents, but will
occasionally eat grouse and hares when voles are less abundant. We
would expect that the number of foxes in the population would increase
as availability of their preferred food increases, and studies have
demonstrated that this does, in fact, occur (Figure 1b). Owl populations
cycle in a similar manner, closely following the abundance of voles.
As predator populations increase, they put greater strain on the prey

populations and act as a top down control, pushing them toward a state
of decline. Thus both availability of resources and predation pressure
affect the size of prey populations. We cannot easily determine the extent
to which each of these controls drives population cycles in the Swedish
boreal forest, because this system is not amenable to caging experiments,
but studies show that food and predation work together to regulate
population sizes.
125
At the end of this unit, students should be able to explain Population

cycles in animal and predator-prey interaction using Lotka-Volterra
models with Vole-fox system
3.3 Modeling Predator-Prey Interactions
Figure 2: Graphical view of the Lotka-Volterra model

(www.encoguru.com)
Predator and prey populations cycle through time, as predators decrease

numbers of prey. Lack of food resources in turn decrease predator
abundance, and the lack of predation pressure allows prey populations to
rebound.
To survive and reproduce, individuals must obtain sufficient food

resources while simultaneously avoiding becoming food for a predator.
The snowshoe hare study demonstrates the role of both predator
avoidance and food availability on population sizes. The trade-off
between food intake and predator avoidance is not easily addressed in
the field, and ecologists have turned to mathematical models to better
understand foraging behaviour and predator-prey dynamics, just as
economists and atmospheric scientists do.
Lotka-Volterra models provide a useful tool to help population

ecologists understand the factors that influence population dynamics.
They have been particularly useful in understanding and predicting
predator-prey population cycles. Although the models greatly simplify
126
actual conditions, they demonstrate that under certain circumstances,

predator and prey populations can oscillate over time (Figure 2) in a
manner similar to that observed in the populations described above.
Few systems oscillate in the cyclical manner of those described thus far.
In reality, predator-prey systems are complex; they often involve
multiple predators and multiple types of prey. What factors influence the
type of prey an individual predator takes? What influences the foraging
behaviour of prey species? Under ideal circumstances, an individual will
encounter high-quality food items on a regular basis. These preferred
foods provide the most nutritional benefit with the fewest costs. Costs
for an organism may be handling time (e.g., time required to catch prey
or remove a nut from its shell) or presence of chemicals, such as tannins,
that reduce the nutritional quality of the food item.
When preferred foods are scarce, organisms must switch to other, less-
desirable alternatives. The point at which an organism should make this
shift is not easy to predict. It depends upon many factors, including the
relative abundance of each of the foods, the potential costs associated
with each food, and other factors, such as the risk of exposure to
predators while eating.
Consider the vole-fox system described in the first section. Field voles
(Microtus agrestis) and bank voles (Clethrionomys glareolus)
preferentially consume forbs and grasses, but they will turn to the bark
from trees when their preferred foods become scarce. Bark contains
poorer quality nutrients than do grasses and forbs. In addition, voles must
venture into the open to approach trees to feed on bark, making them
more vulnerable to predation by foxes, which rely on sight to find their
prey. Only when the preferred foods are very difficult to find—as occurs
during times of population peaks—do voles switch to bark.
3.4 Evolutionary Dynamics of Predator-Prey Systems: An

Ecological Perspective
Evolution takes place in an evolutionary setting that typically involves

interactions with other organisms. To describe such evolution, a structure
is needed which incorporates the simultaneous evolution of interacting
species. Here a formal framework for this purpose is suggested,
extending from the microscopic interactions between individuals- the
immediate cause of natural selection, through the mesoscopic population
dynamics responsible for driving the replacement of one mutant
phenotype by another, to the macroscopic process of phenotypic
evolution arising from many such substitutions. The process of
coevolution that results from this is illustrated in the predator-prey
127
systems. With no more than qualitative information about the

evolutionary dynamics, some basic properties of predator-prey
coevolution become evident. More detailed understanding requires
specification of an evolutionary dynamic; two models for this purpose
are outlined, one from our own research on a stochastic process of
mutation and selection and the other from quantitative genetics. Much of
the interest in coevolution has been to characterize the properties of fixed
points at which there is no further phenotypic evolution. Stability
analysis of the fixed points of evolutionary dynamical systems is
reviewed and leads to conclusions about the asymptotic states of
evolution rather than different from those of game-theoretic methods.
These differences become especially important when evolution involves
more than one species.What information does the Lotka-Volterra models
provide?
1. What happen when preferred foods are scarce in Predator-
prey relationship?
3.5 Summary
Species interactions occur on many levels, as part of a complex, dynamic

system in ecological communities. Predators, prey, plants, and parasites
all influence changes in population sizes over time. Simple systems may
undergo large, cyclical changes, but communities with more complex
food webs are likely to experience more subtle shifts in response to
changes in parasite load, predation pressure, and herbivore. Consider,
however, that humans have impacted many ecological communities by
removing predators or reducing the availability of resources. How will
such changes affect population fluctuations in the rest of the community?
128
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Adult Stages of the Wood Frog (Rana Sylvatica) Ecology 71,
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and the stability of coralreef fish communities. Proceedings of the
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Evolution of Genetic Polymorphism by Small Mutations in a
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129
R. Law & U. Dieckmann, 1997. "Symbiosis without Mutualism and the

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https://biologydictionary.net/predator-prey-relationship/
https://www.sciencedirect.com/topics/agricultural-and-biological-
sciences/predator-prey-relationships
https://www.bing.com/videos/search?q=Predator-
prey+relationship&&view=detail&mid=9EF844124B48D856
ABED9EF844124B48D856ABED&&FORM=VRDGAR&ru
=%2Fvideos%2Fsearch%3Fq%3DPredator-
prey%2Brelationship%26FORM%3DHDRSC3
https://www.youtube.com/watch?v=cCmpg8Yg8Vc
130
Answers to SAEs 1
1. When preferred foods are scarce, organisms must switch to other,
less-desirable alternatives. The point at which an organism should
make this shift is not easy to predict. It depends upon many factors,
including the relative abundance of each of the foods, the potential
costs associated with each food, and other factors, such as the risk
of exposure to predators while eating
131
Contents
4.1 Introduction
4.3 Classification and Evolution of Bat
4.4 Summary
4.1 Introduction
Bats are flying mammals in the order Chiroptera. The forelimbs of bats
are webbed and developed as wings, making them the only mammals
naturally capable of true and sustained flight. By contrast, other
mammals said to fly, such as flying squirrels, gliding possums and
colugos, glide rather than fly, and can only glide for short distances.
Bats do not flap their entire forelimbs, as birds do, but instead flap
their spread out digits, which are very long and covered with a thin
membrane or patagium. Chiroptera comes from two Greek words, cheir
(χείρ) "hand" and pteron (πτερόν) "wing."
There are about 1,100 bat species worldwide, which represent about
twenty percent of all classified mammal species. About seventy percent
of bats are insectivores. Most of the rest are frugivores, or fruit eaters. A
few species such as the Fish-eating Bat feed from animals other than
insects, with the vampire bats being the only mammalian parasite
species. Bats are present throughout most of the world and perform
vital ecological roles such as pollinating flowers and dispersing fruit
seeds. Many tropical plant species depend entirely on bats for the
distribution of their seeds.
The smallest bat is the Kitti's Hog-nosed Bat, measuring 29–34 mm

(1.14–1.34 in) in length, 15 cm (5.91 in) across the wings and 2–2.6 g
(0.07–0.09 oz) in mass, The largest species of bat is the Giant Golden-
crowned Flying-fox, which is 336–343 mm (13.23–13.50 in) long,
has a wingspan of 1.5 m (4 ft 11 in) and weighs approximately 1.1–1.2
kg (2–3 lb).
132
At the end of unit, students should be able to explain the classification

and evolution of bat and its general characteristics.
4.0 Classification and Evolution of Bat
Bats are mammals. Sometimes they are mistakenly called "flying

rodents" or "flying rats", and they can also be mistaken for insects and
birds. There are two traditionally recognized suborders of bats:
1. Megachiroptera (megabats)
2. Microchiroptera (microbats / echolocating bats)
Not all megabats are larger than microbats. The major distinctions
between the two suborders are:
1. Microbats use echolocation: megabats do not with the
exception of Rousettus and relatives.
2. Microbats lack the claw at the second toe of the forelimb.
3. The ears of microbats do not close to form a ring: the edges are
separated from each other at the base of the ear.
4. Microbats lack underfur: they are either naked or have guard hairs.
Megabats eat fruit, nectar or pollen while most microbats eat insects;
others may feed on the blood of animals, small mammals, fish, frogs,
fruit, pollen or nectar. Megabats have a well- developed visual cortex
and show good visual acuity, while microbats rely on echolocation for
navigation and finding prey.
i) Fossil bats
There are few fossilized remains of bats, as they are terrestrial and light-
boned. An Eocene bat, Onychonycteris finneyi, was found in the fifty-
two-million-year-old Green River Formation in South Dakota, United
States, in 2004. It had characteristics indicating that it could fly, yet
the well-preserved skeleton showed that the cochlea of the inner ear
lacked development needed to support the greater hearing abilities of
modern bats. This provided evidence that flight in bats developed well
before echolocation. The team that found the remains of this species,
named Onychonycteris finneyi, recognized that it lacked ear and throat
features present not only in echolocating bats today, but also in other
known prehistoric species. Fossil remains of another Eocene bat,
Icaronycteris, were found in 1960.
133
The appearance and flight movement of bats 52.5 million years ago were
different from those of bats today. Onychonycteris had claws on all five
of its fingers, whereas modern bats have at most two claws appearing on
two digits of each hand. It also had longer hind legs and shorter forearms,
similar to climbing mammals that hang under branches such as sloths
and gibbons. This palm-sized bat had broad, short wings suggesting that
it could not fly as fast or as far as later bat species. Instead of flapping
its wings continuously while flying, Onychonycteris likely alternated
between flaps and glides while in the air. Such physical characteristics
suggest that this bat did not fly as much as modern bats do, rather flying
from tree to tree and spending most of its waking day climbing or hanging
on the branches of trees.
ii) Habitats
Flight has enabled bats to become one of the most widely distributed
groups of mammals. Apart from the Arctic, the Antarctic and a few
isolated oceanic islands, bats exists all over the world. Bats are found
in almost every habitat available on Earth. Different species select
different habitat during different seasons — ranging from sea-sides to
mountains and even deserts — but bat habitats have two basic
requirements: roosts, where they spend the day or hibernate, and places
for foraging. Bat roosts can be found in hollows, crevices, foliage, and
even human-made structures; and include "tents" that bats construct by
biting leaves.
iii) Echolocation
Bat echolocation is a perceptual system where ultrasonic sounds are
emitted specifically to produce echoes. By comparing the outgoing
pulse with the returning echoes the brain and auditory nervous system
can produce detailed images of the bat's surroundings. This allows bats
to detect, localize and even classify their prey in complete darkness. At
130 decibels in intensity, bat calls are some of the most intense airborne
animal sounds.
To clearly distinguish returning information, bats must be able to

separate their calls from the echoes they receive. Microbats use two
distinct approaches.
1. Low Duty Cycle Echolocation: Bats can separate their calls and
returning echos by time. Bats that use this approach time their short calls
to finish before echoes return. This is important because these bats
contract their middle ear muscles when emitting a call so that they can
avoid deafening themselves. The time interval between call and echo
allows them to relax these muscles so they can clearly hear the returning
134
echo. The delay of the returning echos provide the bat with the ability to
estimate range to their prey.
2. High Duty Cycle Echolocation: Bats emit a continuous call and

separate pulse and echo in frequency. The ears of these bats are sharply
tuned to a specific frequency range. They emit calls outside of this range
to avoid self-deafening. They then receive echoes back at the finely
tuned frequency range by taking advantage of the Doppler shift of their
motion in flight. The Doppler shift of the returning echos yield
information relating to the motion and location of the bat's prey. These
bats must deal with changes in the Doppler shift due to changes in their
flight speed. They have adapted to change their pulse emission frequency
in relation to their flight speed so echoes still return in the optimal
hearing range.
iv) Behaviour
Most microbats are nocturnal and are active at twilight. A large portion
of bats migrate hundreds of kilometres to winter hibernation dens,
some pass into torpor in cold weather, rousing and feeding when warm
weather allows for insects to be active. Others retreat to caves for winter
and hibernate for six months. Bats rarely fly in rain as the rain interferes
with their echo location, and they are unable to locate their food.
The social structure of bats varies, with some bats leading a solitary
life and others living in caves colonized by more than a million bats.
The fission-fusion social structure is seen among several species of
bats. The term "fusion" refers to a large numbers of bats that
congregate together in one roosting area and "fission" refers to
breaking up and the mixing of subgroups, with individual bats
switching roosts with others and often ending up in different trees and
with different roostmates.
70% of bat species are insectivorous, locating their prey by means of

echolocation. Of the remainder, most feed on fruits. Only three species
sustain themselves with blood. Some species even prey on vertebrates:
these are the leaf-nosed bats (Phyllostomidae) of Central America and
South America, and the two bulldog bat (Noctilionidae) species, which
feed on fish. At least two species of bat are known to feed on bats: the
Spectral Bat, also known as the American False Vampire bat, and the
Ghost Bat of Australia. One species, the Greater Noctule bat, catches
and eats small birds in the air. Predators of bats include bat hawks and
bat falcons.
v) Reproduction
Most bats have a breeding season, which is in the spring for species
living in a temperate climate. Bats may have one to three litters in a
135
season, depending on the species and on environmental conditions such

as the availability of food and roost sites. Females generally have one
offspring at a time, which could be a result of the mother's need to fly
to feed while pregnant. Female bats nurse their youngsters until they
are nearly adult size; this is because a young bat cannot forage on its
own until its wings are fully developed.
Female bats use a variety of strategies to control the timing of pregnancy

and the birth of young, to make delivery coincide with maximum food
ability and other ecological factors. Females of some species have delayed
fertilization, in which sperm are stored in the reproductive tract for several
months after mating. In many such cases, mating occurs in the fall, and
fertilization does not occur until the following spring. Other species
exhibit delayed implantation, in which the egg is fertilized after mating,
but remains free in the reproductive tract until external conditions
become favorable for giving birth and caring for the offspring. In
yet another strategy, fertilization and implantation both occur but
development of the fetus is delayed until favorable conditions prevail. All
of these adaptations result in the pup being born during a time of high
local production of fruit or insects.
At birth the wings are too small to be used for flight. Young microbats
become independent at the age of 6 to 8 weeks, while megabats do not
until they are four months old.
A single bat can live over 20 years, but the bat population growth is
limited by the slow birth rate.
vi) Feeding
Newborn bats rely on the milk from their mother’s nipples for
sustenance. When they are a few weeks old, bats are expected to fly and
hunt on their own. It is up to them to find and catch their prey, along
with satisfying their thirst. The majority of food consumed by bats
includes insects, fruits and flower nectar, vertebrates (fish, frogs, lizards,
birds, and sometimes other bats) and blood. Almost three-fourths of the
world’s bats are insect eaters. Insects consumed by bats include both
aerial insects, and ground-dwelling insects. Each bat is typically able
to consume one third of its body weight in insects each night, and
several hundred insects in a few hours. This means that a group of
one thousand bats could eat four tons of insects each year. If bats
were to become extinct, the insect population is calculated to reach an
alarmingly high number.
136
vii) Pathogens and role in the transmission of zoonoses

Among ectoparasites, bats occasionally carry fleas, but are one of the
few mammalian orders that cannot host lice (most of the others are water
animals).
Bats are natural reservoir for a large number of zoonotic pathogen

including rabies, severe acute respiratory syndrome (SARS),
Henipavirus (i.e. Nipah virus and Hendra virus) and possibly ebola virus.
Their high mobility, broad distribution, and social behaviour (communal
roosting, fission-fusion social structure) make bats favourable hosts and
vectors of disease. Many species also appear to have a high tolerance for
harbouring pathogens and often do not develop disease while infected.
If a bat is found in a house and the possibility of exposure cannot be

ruled out, the bat should be sequestered and an animal control officer
called immediately, so that the bat can be analysed. This also applies
if the bat is found dead. If it is certain that nobody has been exposed to
the bat, it should be removed from the house. The best way to do
this is to close all the doors and windows to the room except one that
opens to the outside. The bat should soon leave.
Due to the risk of rabies and also due to health problems related to
their faecal droppings (guano), bats should be excluded from inhabited
parts of houses. The Center for Disease Control and Prevention provides
full detailed information on all aspects of bat management, including
how to capture a bat, what to do in case of exposure, and how to bat-
proof a house humanely. In certain countries, it is illegal to handle bats
without a license.List the two suborders of bats?
1. What has made Bat the most successful groups of
mammals?
2. What is Bat Echolocation?
4.4 Summary
Bats are flying mammals in the order Chiroptera. There are about 1,100
bat species worldwide, which represent about twenty percent of all
classified mammal species. About seventy percent of bats are
insectivores. Most of the rest are frugivores, or fruit eaters. Flight has
137
enabled bats to become one of the most widely distributed groups of

mammals.
Baerwald, E.F., G.H. D’Amours, B.J. Klug, R.M. Barclay. 2008.

Barotrauma is a significant cause of bat fatalities at wind turbines.
Current Biology 18:R695-R696.
Barbour, R.W, and W.H. Davis. 1969 Bats of America. The University
Press of Kentucky.Lexinton, KY.
Bat Conservation International [BCI]. “White Nose Syndrome.” Bat

Conservation International. 2009. (accessed Dec 2009).
Bat Conservation International [BCI]. “Bat Species Profiles: Myotis

lucifugus.” Bat Conservation International, 2012. (accessed Sept
2012).
Bergeson S.M. 2012. Examining the suitability of the little brown bat
(Myotis lucifugus) as a surrogate for the endangered Indiana bat
(Myotis sodalis). Masters thesis, Ball State University,
Munci IN. Boyles J.G., P.M. Cryan, G.F. McCracken, T.H. Kunz. 2011.
Economic importance of bats in agriculture. Science 332:41-42.
Brittingham, M.C. and L.M. Williams. 2000. Bat boxes as alternative

roosts for displaced bat maternity colonies. Wildlife Society
Bulletin 28:197-207
Hunter, P. (September 2007). "The nature of flight. The molecules and

mechanics of flight in animals". Science and Society 8 (9): 811–
813. doi:10.1038/sj.embor.7401050. PMC 1973956. PMID
17767190. http://ukpmc.ac.uk/articlerender.cgi?artid=1635883.
Retrieved 2009-07-17.
https://www.cbd.int/financial/values/g-ecobats.pdf
https://dnr.wi.gov/files/pdf/pubs/er/er0705.pdf
https://ec.europa.eu/environment/nature/conservation/species/action_pl
ans/pdf/EU%20Bats%20Action%20Plan.pdf
http://www.arkive.org/golden-capped-fruit-bat/acerodon-jubatus/
138
http://animals.jrank.org/pages/2834/Kitti-s-Hog-Nosed-Bat-
Craseonycteridae-PHYSICAL- CHARACTERISTICS.html
https://ncep.amnh.org/index.php/Detail/objects/1054
https://www.nature.com/articles/d41586-018-05648-2
https://www.youtube.com/watch?v=pmsEzGqHApg
https://www.wsl.ch/en/biodiversity/conservation-biology-and-nature-
reserves/bats.html
139
Answers to SAEs 1
1. Flight has enabled bats to become one of the most widely
distributed groups of mammals. Apart from the Arctic, the
Antarctic and a few isolated oceanic islands, bats exists all over
the world. Bats are found in almost every habitat available on
Earth. Different species select different habitat during different
seasons — ranging from sea-sides to mountains and even
deserts.
2. Bat echolocation is a perceptual system where ultrasonic

sounds are emitted specifically to produce echoes. By
comparing the outgoing pulse with the returning echoes the
brain and auditory nervous system can produce detailed images
of the bat's surroundings. This allows bats to detect, localize and
even classify their prey in complete darkness. At 130 decibels in
intensity, bat calls are some of the most intense airborne animal
sounds
140
Contents
5.1 Introduction
5.3 Proximate causation and Optimization theory
5.3.1 Evolutionarily stable strategies (ESS)
5.3.2 Tinbergen's Four Questions
5.4 Summary
5.1 Introduction
Behavioural ecology, or ethoecology, is the study of the ecological and

evolutionary basis for animal behavior, and the roles of behaviour in
enabling an animal to adapt to its environment (both intrinsic and
extrinsic). Behavioural ecology emerged from ethology after Niko
Tinbergen (a seminal figure in the study of animal behaviour) outlined
the four causes of behaviour.
If an organism has a trait which provides them with a selective advantage

(i.e. has an adaptive significance) in a new environment natural selection
will likely favour it. Adaptive significance therefore refers to the
beneficial qualities, in terms of increased survival and reproduction, a
trait conveys.
For example, the behaviour of flight has evolved numerous times in

reptiles (Pterosaur), birds, many insects and mammals (bats) due to its
adaptive significance—for many species, flight has the potential to
increase an animal's ability to escape from predators and move swiftly
between habitat areas, among other things, thereby increasing the
organism's chances of survival and reproduction. In all instances, the
organism adapting to flight had to have "pre-adaptions" to these
behavioural and anatomical changes. Feathers in birds initially evolving
for thermoregulation then turned to flight due to the benefits conveyed
(see Origin of avian flight); insect wings evolving from enlarged gill
plates used to efficiently "sail" across the water, becoming larger until
capable of flight are two good examples of this. At every stage slight
improvements mean higher energy acquisition, lower energy
expenditure or increased mating opportunities causing the genes that
convey these traits to increase within the population. If these organisms
did not have the required variation for natural selection to act upon either
141
due to phylogenetic or genetic constraints, these behaviours would not

be able to evolve.
However, it is not sufficient to apply these explanations where they
seem convenient. Viewing traits and creating unsubstantiated theories or
"Just So Stories" as to their adaptive nature have been deeply criticized.
Stephen Jay Gould and Richard Lewontin (1979) described this as the
"adaptationist programme". To be rigorous, hypotheses regarding
adaptations must be theoretically or experimentally tested as with any
scientific theory.
The hypothesis of the evolution of insect flight for example has been
tested through wing manipulation experiments. Empirical observations
which adhere to the conditions prosed also provide evidence. For
instance, one can suppose that when birds are not at risk of being eaten
they might lose the ability to fly as the construction of functional wings
are costly to produce and take away energy which could be used to
increase offspring production or survival, a trend many island flightless
birds such as the Kakapo, the Penguin and the now extinct Dodo
demonstrate in the absence of natural predators.
At the end of this unit, students should be able to:
 Understand behavioural ecology and the importance of the

Tinbergen's Four Questions.
 Explain optimization theory and evolutionarily stable strategies
(ESS)
5.3 Proximate causation and Optimization theory
Proximate causation is divided into two factors which are ontogenetic

and mechanistic. Ontogenetic factors are the entire sum of experience
throughout the lifetime of an individual from embryo to death. Hence,
factors included are learning the genetic factors giving rise to behaviour
in individuals. Mechanistic factors, as the name implies, are the
processes of the body that give rise to behaviour such as the effects of
hormones on behaviour and neuronal basis of behaviour.
Behavioural ecology, along with other areas of evolutionary biology, has

incorporated a number of techniques which have been borrowed from
optimization theory. Optimization is a concept that stipulates strategies
that offer the highest return to an animal given all the different factors
142
and constraints facing the animal. One of the simplest ways to arrive at
an optimal solution is to do a cost/benefit analysis. By considering the
advantages of a behaviour and the costs of a behaviour, it can be seen
that if the costs outweigh the benefits then a behaviour will not evolve
and vice versa. This is also where the concept of the trade-off becomes
important. This is because it rarely pays an animal to invest maximally
in any one behaviour. For example, the amount of time an ectothermic
animal such as a lizard spends foraging is constrained by its body
temperature. The digestive efficiency of the lizard also increases with
increases in body temperature. Lizards increase their body temperature
by basking in the sun. However, the time spent basking decreases the
amount of time available for foraging. Basking also increases the risk of
being discovered by a predator. Therefore, the optimal basking time is
the outcome of the time necessary to sufficiently warm itself to carry out
its activities such as foraging. This example shows how foraging is
constrained by the need to bask (intrinsic constraint) and predation
pressure (extrinsic constraint).
An often quoted behavioural ecology hypothesis is known as Lack's

brood reduction hypothesis (named after David Lack). Lack's hypothesis
posits an evolutionary and ecological explanation as to why birds lay a
series of eggs with an asynchronous delay leading to nestlings of mixed
age and weights. According to Lack, this brood behaviour is an
ecological insurance that allows the larger birds to survive in poor years
and all birds to survive when food is plentiful.
Ultimately, behaviour is subject to natural selection just as with any other

trait. Therefore animals that employ optimal behavioural strategies
specific to their environment will generally leave greater numbers of
offspring than their suboptimal conspecifics. Animals that leave a greater
number of offspring than others of their own species are said to have
greater fitness. However, environments change over time. What might
be good behaviour today might not be the best behaviour in 10,000 years’
time or even 10 years’ time. The behaviour of animals has and will
continue to change in response to the environment. Behavioural ecology
is one of the best ways to study these changes. As geneticist Theodosius
Dobzhansky famously wrote, "nothing in biology makes sense except in
the light of evolution."
5.3.1 Evolutionarily stable strategies (ESS)
The value of a social behaviour depends in part on the social behaviour

of an animal's neighbors. For example, the more likely a rival male is to
back down from a threat, the more value a male gets out of making the
threat. The more likely, however, that a rival will attack if threatened, the
less useful it is to threaten other males. When a population exhibits a
143
number of interacting social behaviours such as this, it can evolve a

stable pattern of behaviours known as an evolutionarily stable strategy
(or ESS). This term, derived from economic game theory, became
prominent after John Maynard Smith (1982), recognized the possible
application of the concept of a Nash equilibrium to model the evolution
of behavioural strategies.
In short, evolutionary game theory asserts that only strategies that, when
common in the population, cannot be "invaded" by any alternative
(mutant) strategy will be an ESSs, and thus maintained in the population.
In other words, at equilibrium every player should play the best strategic
response to each other. When the game is two player and symmetric each
player should play the strategy which is the best response to itself.
Therefore, the ESS is considered to be the evolutionary end point

subsequent to the interactions. As the fitness conveyed by a strategy is
influenced by what other individuals are doing (the relative frequency of
each strategy in the population), behaviour can be governed not only by
optimality but the frequencies of strategies adopted by others and are
therefore frequency dependent (frequency dependence).Behavioural
evolution is therefore influenced by both the physical environment and
interactions between other individuals.
5.3.2 Tinbergen's Four Questions
Most behavioural ecologists focus on answering one of the following

questions in their studies. They want to know either what is the cause of
the behaviour; how did the behaviour develop within the individual's
lifetime; what function, or functions, does the behaviour serve; or how
did the behaviour evolve? These questions are known as Tinbergen's
Four Questions and are named after the behaviourist Niko Tinbergen
who developed them.
Question number one deals with both environmental and internal factors
that may cause behaviours, such as hormones or infringing on another
animal's territory. The answers to question number two may include both
genetics and the animal's past experiences, similar to the "nature versus
nurture" problem in psychology. Question number three asks how the
behaviours will impact the animal immediately and its ability to adapt,
such as asking what will happen if an animal's territory is taken by
another. Question number four concentrates on how the behaviours
originated and how they have changed or will change, such as a horse's
flight instinct being exacerbated by modern horse-keeping practices.
There are several reasons to study behavioural ecology. Behavioural

ecology can be used as a way to interpret human social problems (several
144
ground-breaking psychological studies were based on observing animal

behaviour), applied to the study of neurobiology (neuroethology
combines animal behaviour and neuroscience) , find solutions to
environmental issues (animal behaviour is indicative of health of the
environment), help with animal welfare (behavioural observation is
necessary for deciding how to protect endangered species), and bring in
new interest in biological sciences (many students find animal behaviour
more interesting than other science disciplines). There is also a lot of
interest in animal behaviour from the public. Consider all of the recent
documentaries and books that have come out, and the popularity of safari
trips.
Dr. Jennifer Borgo, a visiting assistant professor at Coker College in

Hartsville, SC, is a wildlife behavioural ecologist who has studied flying
squirrels, ducks, woodpeckers, and grouse. She studies ultimate
behaviours as opposed to proximate behaviours. Proximate behaviours
refer to question one of Tinbergen's Four Questions, and ultimate
behaviours refer to question three. Dr. Borgo focuses on interspecific
interaction, which she describes as the interactions between members of
different species (such as one animal hunting another). What is
behavioural ecology? What is Ontogenetic factors?
1. State Tinbergen's Four Questions?
2. What is Optimization theory?
5.4 Summary
Behavioural ecology emerged from ethology after Niko Tinbergen (a

seminal figure in the study of animal behaviour) outlined the four causes
of behaviour.
145
Marden, J.H. & M.G. Kramer (1994) Surface-skimming stoneflies: a

possible intermediate stage in insect flight evolution. Science,
Vol. 266, 427-430
Amundsen, T.; Slagsvold, T. (1996). "Lack's Brood Reduction

Hypothesis and Avian Hatching Asynchrony: What's
Next?". Oikos 76 (3): 613–620.
doi:10.2307/3546359. http://jstor.org/stable/3546359.
Pijanowski, B. C. (1992). "A Revision of Lack's Brood Reduction

Hypothesis". The American Naturalist 139 (6): 1270–1292.
doi:10.1086/285386.
J.R. Krebs and Nicholas Davies, An Introduction to Behavioural

Ecology, ISBN 0-63203546-3
J.R. Krebs and Nicholas Davies, Behavioural Ecology: an evolutionary

approach, ISBN 0-86542-731-3
https://www.sciencedirect.com/topics/earth-and-planetary-
sciences/behavioral-ecology
https://behavioralecology.com/
https://behaviouralecologyconservation.weebly.com/an-introduction-to-
behavioural-ecology.html
https://www.bing.com/videos/search?q=What+is+behavioural+ecology
&&view=detail&mid=679859470CD19778B328679859470CD
19778B328&&FORM=VRDGAR&ru=%2Fvideos%2Fsearch%
3Fq%3DWhat%2Bis%2Bbehavioural%2Becology%26FORM%
3DHDRSC3
146
Answers to SAEs 1
1. Answer: Most behavioural ecologists focus on answering one of
the following questions in their studies. They want to know either
what is the cause of the behaviour; how did the behaviour develop
within the individual's lifetime; what function, or functions, does
the behaviour serve; or how did the behaviour evolve?
2. Optimization is a concept that stipulates strategies that offer the

highest return to an animal given all the different factors and
constraints facing the animal.
Glossary
cm = centimeters
CO2 = Carbondioxide
CH4 = methane (CH4)
ft = feets
Hz = Hertz
in = inches
Ib = pounds
Kgs = kilograms.
km = kilometers
m = meters
mm = millimeters
O2 = Oxygen
oz = ounce
L = length
% = percentage
g = grams
spp = species
147
1. What is Apparent Competition?

2. What is Terrestrial animal?
3. What is population cycle?
4. List few characteristic features of African bat?
5. What is behavioural ecology?
148

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COURSE

Course Team O.A. Olajuyigbe -Adeyemi Colledge of

NATIONAL OPEN UNIVERSITY OF NIGERIA

© 2023 by NOUN Press

Printed 2022, Reprinted 2023

Animal Ecology (BIO 313) is a 2 credit-hours course for undergraduate

The Comprehensive Objectives of the Course are to;

1. Explain the historical background of animal ecology,

WORKING THROUGH THIS COURSE

Module 1 Introduction to Animal Ecology

Unit 1 Historical Background of Animal Ecology

Module 2 Ecology of Animal Population

Unit 1 Growth rate and Age structure of animal population

Module 3 Population Cycle in Animal Ecology

Unit 1 Competition in Animal Ecology

REFERENCES AND FURTHER READINGS

You would be required to read the recommended references and

HOW TO GET THE MOST FROM THE COURSE

This course exposes you to Animal Ecology, a sub-discipline and very

Do not delay to contact your facilitator by telephone or e-mail for

• You have difficulty with the self-assessment activities.

Dr. Esenowo, Imeh Kokoete is a Senior Lecturer of Ecology and

Module 1 Introduction to Animal Ecology ……………… 1

Unit 1 Historical Background of Animal Ecology ……… 1

Module 2 Ecology of Animal Population……………….. 46

Unit 1 Growth rate and Age structure of animal

Module 3 Population Cycle in Animal Ecology …… 107

Unit 1 Competition in Animal Ecology …………. 107

Module 1 Introduction to Animal Ecology

Unit 1 Historical Background of Animal Ecology

Unit 1 Historical Background of Animal Ecology

Ecology (from Greek: οἶκος,"house"; -λογία, "study of") is the study

integrate into a complex and regenerative spatial arrangement of types,

Ecology is a sub-discipline of biology, the study of life. The word

Ecology is not synonymous with environment, environmentalism,

1. life processes and adaptations

There are many practical applications of ecology in conservation

1.2 Intended Learning Outcomes (ILOs)

At the end of this unit, the student should be able to

 explain the historical background of animal ecology and the

1.3 Historical Background of Ecology

Studies of animal distribution began in the nineteenth century, but the

Although some of the early work in animal ecology, particularly in the

perspective also emerged in soil science, as in Sergei Winogradsky's

World War II proved to be a watershed for ecology. Although earlier

post-war emergence of a more strongly Darwinian perspective in

The post-war years also saw a shift toward quantitative aspects of

The devastation brought by World War II also contributed to greater post-

Who is the founder of British Ecological

You have been introduced to the field of animal ecology including

1.5 References/Further Readings/Web Sources

Donald Worster, Nature's Economy: A History of Ecological Ideas, 2d

Eugene Cittadino, Nature as the Laboratory: Darwinian Plant Ecology

Leslie A. Real and James H. Brown, eds., Foundations of Ecology

Gregg Mitman, The State of Nature: Ecology, Community, and

Michael Shortland, ed., Science and Nature: Essays in the History of

Robert P. McIntosh,The Background of Ecology (1985).

Sharon Kingsland, Modeling Nature: Episodes in the History of

Stephen Bocking, Ecologists and Environmental Politics: A History of