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MIDLANDS
STATE UNIVERSITY
DEPARTMENT OF APPLIED BIOSCIENCES AND BIOTECHNOLOGY
LEVEL 1.2
HABB 103: PRINCIPLES OF ECOLOGY
Lecturer: J. Bare
Module description
The
module
includes
four
one-‐hour
lectures
and
two
three-‐hour
practicals
per
week,
but
attendance
may
be
required
at
additional
meetings
for
extra
lectures
or
tests.
The
duration
of
the
modules
will
be
twelve
weeks.
Major
Topics
are:
What
is
Ecology?
Ecology
and
levels
of
organization
in
biological
systems:
individuals,
populations,
communities,
ecosystems,
and
the
biosphere;
distribution
and
abundance
of
organisms
in
space
and
time:
causes
and
consequences;
populations
growth;
estimation
of
population
size
and
density;
concepts
of
habitat
and
niche;
competition;
intra-‐
and
inter-‐specific
competition;
relations
among
species:
commensalisms,
mutualism,
parasitism
and
predation;
population
regulation.
Grading Criteria
In
this
module
the
final
examination
will
constitute
70%
of
the
final
mark.
The
remaining
30%
will
come
from
continuous
assessment
marks
as
follows:
20%
from
continuous
assessment
of
practicals
and
10%
from
continuous
assessment
of
tests
and
assignments.
For
your
theory
continuous
assessment,
you
will
write
two
in-‐class
tests
(one
midway
through
the
module
and
the
other
one
towards
the
end
of
the
module)
and
two
assignments.
References
STILING, P. D Ecology and Applications (2nd Edition) Prentice-‐Hall.
BEGON,
M.,
HARPER,
J.
H.
and
TOWNSEND,
C.
R.
(1990).
Ecology:
Individuals
,
Populations
and
Communities
Blackwell
Scientific
Publications,
Oxford.
BEGON,
M
and
MORTIMER,
M.
(1986).
Population
Ecology:
A
Unified
study
of
Animals
and
Plants
Blackwell
Scientific
Publications,
Oxford.
VARLEY,
G.
C.,
GRADWELL,
G.
R.
AND
HASSEL,
M.
P.
(1975).
Insect
Population
Ecology
Blackwell
Scientific
Publications,
Oxford.
OSBORNE,
P.
L.
(2000).
Tropical
Ecosystems
and
Ecological
Concepts.
Cambridge
University
Press.
BARBOUR,
M.
G.,
BULK,
J.
H
and
PITTS,
W.
D.
(1987).
Terrestrial
Plant
Ecology
(2nd
Edition)
Benjamin
Cummings,
Menlo
Park,
California.
CAMPBELL, N. A. (1993). Biology (3rd edition). Benjamin Cummings, Menlo Park, California.
CHAPMAN,
J.
L.
and
REISS,
M.
J.
(1992).
Ecology:
Principles
and
Applications.
Cambridge
University
Press.
COLLINVAUX, P. A. (1973). Introduction to Ecology. John Wiley and Sons, New York.
ENRICH, P. R. and ROUGHGARDEN, J. (1987). The Science of Ecology, MacMillan, New York.
KIKAWA,
J.
and
ANDERSON,
D.
J.
(1986).
Community
ecology:
Pattern
and
Process
Blackwell
Scientific
Publications,
Oxford
KNOX,
B.,
LADGES,
P.
and
EVANS,
B.
(1994).
Biology
McGraw-‐Hill
Book
Company,
Sydney,
New
York,
San
Francisco,
Auckland,
Bogota,
Caracas,
Lisbon,
London
,
Madrid,
Mexico
City,
Milan,
Montreal,
New
Dehli,
San
Juan,
Singapore,
Tokyo,
Toronto.
KREBS, C. J. (1978). Ecology (2nd Edition). Harper and Row, New York.
MACKENZIE,
A.,
BALL,
A.
S.
and
VIRDEE,
S.
R.
(1999).
Instant
Notes
in
Ecology.
Viva
Books
Private
Limited,
New
Dehli,
Mumbai,
Chenai.
MCINTOSH,
R.
P.
(1985).
The
Background
of
Ecology:
Concept
and
Ecology,
Cambridge
University
Press,
Cambridge.
ODUM, E. P. (1971). Fundamentals of Ecology (3rd Edition) W. B. Saunders, Philadelphia
ODUM,
E.
P.
(1983).
Basic
Ecology
and
our
Endangered
Life
Support
System.
Sinauer
Associates
Sunderland,
Massachusettes
Lecture
1:
INTRODUCTION
Module:
Principles
of
Ecology
Dept.
of
Applied
Biosciences
and
Biotechnology,
Midlands
State
University
1.1. What is Ecology?
1.2. Ecology and levels of organisation
1.3. Relations of ecology to other fields of biology
1.4. Doing ecology
___________________________________________________________________________
INTRODUCTION
What
is
Ecology?
The
term
“ecology”
was
coined
by
the
German
zoologist
Ernest
Haeckel
in
1866
to
describe
a
then
emerging
speciality
area
of
biology.
Ernest
Haeckel
derived
the
term
ecology
from
the
Greek
word
Oikos
which
originally
referred
(in
Greek)
to
a
family
household
and
its
daily
operations
and
maintenance;
the
same
root
(Oikos)
also
gave
rise
to
the
word
“economy”
which
referred
to
organisms
linked
together
in
conflict
as
well
as
in
mutual
aid:
Haeckel
elaborated
on
this
in
1869:
“By
ecology,
we
mean
the
body
of
knowledge
concerning
the
economy
of
nature,
the
investigation
of
the
total
relations
of
the
organism
(i.e.
animal,
plant,
protistan,
fungus,
bacterium,
etc.)
both
to
its
organic
and
inorganic
environment;
including
above
all,
its
friendly
and
inimical
relations
with
those
animals
and
plants
(or
any
other
organisms)
with
which
it
comes
directly
or
indirectly
into
contact.
In
a
word,
ecology
is
the
study
of
all
the
complex
interrelationships
referred
to
by
Darwin
as
the
conditions
of
the
struggle
for
existence”.
From then on ecology has been various defined:
1. Ecology
=
the
study
of
the
interaction
of
organisms
with
their
environments,
both
abiotic
(physical-‐chemical)
and
abiotic
(other
organisms).
The
problem
of
this
definition
is
that
it
is
very
broad
and
inclusive,
and
it
leaves
very
little
that
is
NOT
ecology,
and
there
is
need
to
narrow
it
down
somewhat.
2. Ecology= scientific natural history (Charles Elton, 1927, Animal Ecology).
Where:
Natural
history
=
the
study
of
organisms
in
nature
with
respect
to
all
aspects
of
their
biology,
from
mating
and
reproduction
to
predator
thwarting,
diet
and
finding
food,
etc.
Natural
history
gave
rise
not
only
to
ecology
but
also
to
systematics,
ethology
(behaviour),
comparative
biology
and
evolutionary
biology.
Advantage:
points
out
the
origin
of
many
ecological
problems
in
the
ancient
field
of
natural
history.
Problem: uncomfortably vague.
3. Ecology
=
the
study
of
structure
and
function
of
nature
(Eugene
Odum,
1963).
Advantage:
emphasises
the
form
and
function
idea
that
permeates
biology.
Problem: still not completely clear.
4. Ecology
=
the
scientific
study
of
the
distribution
and
abundance
of
organisms
(H.
G.
Andrewartha,
1961,
Introduction
to
the
study
of
Animal
populations).
Advantage: clear and restrictive definition.
Problem: this definition is static and it leaves out the idea of relationships.
5. Ecology
=
the
scientific
study
of
the
interactions
that
determine
the
abundance
and
distribution
of
organisms
(Charles
J.
Krebs,
1972,
Ecology).
Where:
Distribution
=
the
geographical
extent
of
a
population
or
any
other
ecological
unit.
Abundance= the number or quantity of organisms in a given area (density).
This
definition
brings
out
the
idea
that
ecologists
are
interested
in
knowing
where
organisms
are
found,
how
many
occur
there,
and
why?
For the purposes of this module we are going to use definition number 5.
Ecology and levels of organisation in biological systems
We have to recognise 9 different levels of organisation within biological systems:
Molecule
Organelle
Cell
Individual
Population
Community
Ecosystem
Biome
Biosphere
However,
ecology
is
that
branch
of
biology
that
deals
with
levels
of
organisation
from
the
individual
level
and
above.
That
is:
Individual
Population
Community
Ecosystem
Biome
Biosphere
Individual
organism=
this
is
the
basic
biotic
unit
that
has
to
be
studied
in
relation
to
the
environment.
It
is
easy
to
define
individual
in
some
species
(e.g.
in
humans),
but
more
difficult
in
colonial
(asexual
reproducing)
species,
especially
if
polyps
maintain
physiological
contact.)
Population=
a
group
of
individuals
of
the
same
species
inhabiting
the
same
area
at
the
same
time.
Community=
an
association
of
interacting
populations,
usually
defined
by
the
nature
of
their
interactions
or
the
place
in
which
they
live
(e.g.
herbivore
community,
plankton
community).
Ecosystem=
a
self-‐sufficient
habitat
where
living
organisms
and
the
non-‐living
environment
interact
to
exchange
energy
and
matter
in
a
continuing
cycle
(e.g.
forest
ecosystem,
ocean
ecosystem,
etc).
Biome=
large
areas
of
approximately
uniform
habitat,
consisting
of
distinctive
combinations
of
plant
and
animal
species.
Biosphere= the part of the planet containing living organisms, the living world.
Reductionism=
a
philosophy
which
states
that
the
higher
levels
of
organisation
of
complex
systems
can
be
fully
explained
through
knowledge
of
the
smallest
components.
In
other
words,
reductionism
is
the
idea
that
breaking
a
system
down
into
increasingly
smaller
parts
will
enable
one
to
understand
it
completely.
However,
biological
systems
are
extremely
complex,
especially
at
the
higher
levels
of
organisation
(populations,
communities
and
ecosystems),
and
are
subject
to
stochastic
(chance)
events
that
make
such
reductionism
impossible.
Emergence=
the
occurrence
of
characters
at
the
higher
levels
of
organisation
which
could
not
have
been
predicted
from
a
knowledge
of
the
lower
level
components.
Ecology
of
the
individual
-‐
deals
with
how
organisms
are
affected
by
(and
how
they
affect)
their
environment.
In
other
words,
we
are
not
concerned
so
much
with
the
interaction
between
individuals
and
their
environment,
but
rather
with
the
numbers
of
individuals
and
the
process
leading
to
changes
in
the
number
of
individuals.
Autecology=study
of
the
individual
in
relation
to
the
environmental
conditions
(e.g.
streamlined
bodies
of
animals
from
flowing
rivers;
floating
seeds
in
island-‐dwelling
palm
trees).
In other words, Autecology is the ecology of individual organisms or species.
Ecology
of
populations
(Population
Ecology)
-‐
deals
with
the
presence
or
absence
of
particular
species,
with
their
abundance
or
rarity,
and
with
the
trends
or
fluctuations
in
their
numbers.
There are two approaches at the population level:
(i)-‐Deals
first
with
the
attributes
of
individual
organisms,
and
then
considers
the
way
in
which
these
combine
to
determine
the
characteristics
of
the
population.
This
is
a
reductionist
approach.
(ii)-‐deals
directly
with
the
characteristics
of
populations,
and
tries
to
relate
these
to
aspects
of
the
environment.
This
is
a
systems
approach
which
allows
for
emergent
properties.
*historically tied to animal studies.
Community ecology – often considered together with ecosystem ecology.
It
deals
with
the
structure
of
communities
(e.g.
how
many
species,
which
species)
and
with
the
functions
of
communities/ecosystems
(i.e.
pathways
followed
by
energy,
materials,
nutrients
and
other
chemicals
through
them).
There are two approaches at the community level:
(i)-‐study
the
component
populations
to
gain
an
understanding
of
these
patterns
and
processes.
This
is
a
reductionist
approach.
(ii)-‐look
directly
at
the
properties
of
populations/ecosystems
themselves
(e.g.
species
diversity,
rate
of
biomass
production).
This
is
a
systems
approach
which
allows
for
emergent
properties.
*historically tied to plant studies.
Ecosystem ecology – as for community ecology.
Synecology = studies groups of organisms in relation to their environment, includes
population, community, and ecosystem ecology, i. e. the ecology of plant and
animal communities.
Relations of Ecology to other fields of biology
Ecology tends to overlap with:
(i)Physiology
(ii)Behaviour (Ethology)
(iii)Genetics
(iv)Evolution
Ecology and Physiology
How
do
physiological
mechanisms
act
to
affect
survival
and
reproduction
at
the
population
level?
Physiology
(a
morass
of
within-‐organism
characteristics
which
specify
how
an
individual’s
life
processes
respond
to
the
environment)
is
very
much
ignored
at
the
population
and
community
levels
but
provides
a
means
of
coupling
organism’s
response
to
the
environment.
–
effects
of
abiotic
factors
(e.g.
temperature,
light,
pressure,
pH,
O2
concentration,
etc.)
on
organisms,
act
through
physiological
changes.
–
effects
of
biotic
factors
(e.g.
food,
allelopathic
chemicals,
etc.)
on
organisms,
act
through
physiological
changes.
This can lead to a subject called physiological ecology or environmental physiology.
Ecology and Behaviour
How
do
behavioural
mechanisms
act
to
influence
survival
and
reproduction
at
the
population
level?
Behavioural
responses
of
organisms
to
the
environment
(e.g.
temperature
preferences,
prey
location
and
selection,
mate
choice,
sociality,
decisions
to
forage
or
to
look
for
mates,
etc.)
can
determine
survival
and
reproduction
at
the
population
level.
–
decisions
on
how
to
spend
your
time
and
energy,
e.g.
to
foraging
for
food
or
looking
also
affect
survival
and
reproduction
at
the
population
level.
This can lead to a subject called behavioural ecology.
Ecology and Genetics
– population level, population genetics
–
most
characteristics
of
organisms
that
affect
their
interaction
with
the
environment
have
a
genetic
component
(morphology,
physiology
and
behaviour).
–
Individuals
in
a
population
differ
genetically
and
this
can
result
in
differences
in
their
survival
and
reproduction
under
a
given
set
of
environmental
conditions.
Changes
in
the
genetic
make-‐up
of
a
population
over
time
results
in
EVOLUTION.
Ecology and Evolution
Ecological interactions can result in changes in populations over time (i.e. evolution).
This can lead to a subject called evolutionary biology or evolutionary ecology.
According
to
Evelyn
G.
Hutchinson,
there
are
two
causes
for
observable
patterns
or
relationships,
e.g.
for
the
observable
geographical
range
of
blue
Jays
in
North
America:
Proximal
(ecological)
cause
may
involve
limits
of
the
bird’s
tolerance
of
the
physical
environment
(e.g.
temperature,
scarcity
of
food,
abundance
of
parasites
or
predators,
etc.).
In
other
words,
the
proximal
explanation
looks
at
functions
causing
what
is
happening
here
and
now.
Ultimate
(evolutionary)
cause
addresses
how
the
bird
came
to
have
these
properties
in
the
first
place;
this
is
usually
a
result
of
past
evolutionary
changes.
In
other
words,
the
ultimate
explanation
of
the
present
distribution
and
abundance
of
this
bird
lies
in
the
ecological
experiences
of
its
ancestors,
i.e.
the
ultimate
explanation
looks
at
how
the
current
situation
was
influenced
by
past
events
including
evolution.
Adaptation
=
a
genetically
determined
characteristic
that
enhances
the
ability
of
an
individual
to
cope
with
its
environment;
or
an
evolutionary
process
by
which
organisms
become
better
suited
to
their
environment.
Doing
Ecology
What
do
ecologists
do?
Discover and describe patterns in nature (WHAT AND WHERE).
Explain observed patterns in nature (HOW AND WHY).
(a)Proximate
explanation
=
functions
causing
what
is
happening
here
and
now.
(b)Ultimate
explanation
=
how
the
current
situation
was
influenced
by
past
events
including
evolution.
Predict
or
Control
patterns
(APPLIED
ECOLOGY
=
use
of
ecological
principles
and
knowledge
for
human
benefit)
There are therefore three basic approaches to ecological problems:
Descriptive approach
This
is
where
the
investigator
asks
“what”
questions,
e.g.
what’s
there?
The
investigator
can
make
an
inventory
and
describes
what
he/she
finds,
e.g.
they
can
describe
the
major
vegetation
groups
(Tundra,
temperate,
deciduous
forests,
grasslands,
etc.).
This
is
mainly
natural
history.
–
Darwin
admonished,
“one
might
as
well
go
into
a
gravel
pit,
count
the
pebbles,
and
describe
the
colours”.
Functional approach
In this approach the investigator asks “how” questions, e.g. how does the system operate?
This
approach
is
oriented
towards
dynamics
and
relationships
studies
and
tries
to
identify
the
proximate
causes
of
the
dynamic
responses
of
populations
and
communities
to
immediate
factors
in
the
environment.
Evolutionary approach
The
investigator
asks
“why”
questions,
e.g.
why
does
natural
selection
favour
this
particular
ecological
solution?
–
studies
“ultimate”
causes
of
certain
patterns
and
considers
organisms
and
their
relationships
as
historical
products
of
evolution.
Methods in Ecology
The
principal
method
used
in
ecology
and
other
sciences
is
the
hypothetico-‐deductive
approach.
You
start
with
a
series
of
observations
which
appear
to
show
a
constant
pattern
or
relationship,
e.g.
you
note
that
in
Chimanimani,
pine
seedlings
do
not
grow
under
a
canopy
of
mature
deciduous
trees.
Based
on
these
observations
you
generate
a
theory,
e.g.
pine
seedlings
cannot
tolerate
low
light
levels
found
under
such
a
canopy.
Using
your
theory
as
a
guide
you
can
generate
hypotheses
which
predict
the
outcome
of
various
experiments
if
your
theory
is
correct,
e.g.:
●
Over
a
range
of
sites
there
should
be
a
positive
correlation
between
light
intensity
and
pine
seedling
density.
●
Pine
seedlings
should
show
inability
to
survive
at
low
light
levels
in
controlled
laboratory
experiments.
If
you
artificially
increase
light
intensity
under
a
forest
canopy
in
the
field,
you
should
be
able
to
see
an
increase
in
pine
seedling
abundance.
If
your
experiments
all
produced
the
results
you
predicted,
you
can
conclude
that
you
have
successfully
explained
the
observed
pattern;
if
not,
you
must
change
your
theory,
generate
new
hypotheses
and
run
new
experiments
to
test
them.
Deduction
(logic)
=
the
process
of
reasoning
in
which
a
conclusion
follows
necessarily
from
the
stated
premises;
inference
by
reasoning
from
the
general
to
the
specific.
Kinds of evidence used by ecologists
Hypotheses are tested by collecting data in two ways:
Observation
and
Monitoring
of
natural
systems,
e.g.
measuring
the
density
of
Blue
Jays
nests
in
areas
with
and
without
dogwoods
to
test
the
hypothesis
that
dogwoods
influence
Blue
Jays
nest
density.
Advantage = very close to natural conditions – results may be real.
Disadvantage
=
causes
of
results
may
be
difficult
to
identify
because
of
confounding
factors
(e.g.
areas
studied
may
differ
more
than
in
just
dogwood
density).
Experiments
Types
of
experiments
in
Ecology:
Field
Experiments
–
manipulation
of
natural
systems
to
test
hypotheses,
e.g.
remove
dogwoods
(independent
variable)
from
one
area
but
not
from
a
control
area,
then
compare
Blue
Jay
nest
densities
(dependent
variable)
between
the
two
areas.
An
effort
must
be
made
to
try
and
vary
only
the
independent
variable
of
interest
and
keep
all
other
independent
variables
the
same
in
all
treated
areas.
Advantage = can isolate and better identify causes.
Disadvantage = treatment may have unintended effects (e.g. dogwood
removal may stimulate production of other food items that
the Blue Jays may like).
*Therefore,
there
is
great
realism
(natural
system)
but
low
confidence
(complex
system
with
many
uncontrolled
variables).
Laboratory experiments – manipulation of artificial systems to test hypotheses.
Advantage = highly controlled system.
Disadvantage
=
even
more
unrealistic
than
field
experiments;
we
should
seek
simplicity,
but
distrust
it.
*Therefore,
there
is
low
realism
(unnaturally
simple
system)
but
high
confidence
(most
other
variables
tightly
uncontrolled).
Theoretical/mathematical Models – an abstract representation of ecological interactions.
Advantage = less costly, can answer many questions because it is flexible.
Disadvantage = difficult to account for complexity.
*Models
are
only
as
good
as
the
data
and
relationships
used
to
construct
them.
Used
primarily
when
field
experiments
are
impossible
(too
large
or
too
long,
e.g.
global
warming).
Lecture
2:
POPULATION
STRUCTURES
Module:
Principles
of
Ecology
Dept.
of
Applied
Biosciences
and
Biotechnology,
Midlands
State
University
2.1. Distributions of Populations
2.2. Dispersion – spacing between individuals
2.3. Temporal changes in population size
___________________________________________________________________________
POPULATION
STRUCTURES
The
term
population
is
defined
differently
in
various
sciences:
-‐ In
human
demography
a
population
is
a
set
of
humans
in
a
given
area.
-‐ In
genetics
a
population
is
a
group
of
inter-‐breeding
individuals
of
the
same
species,
which
is
isolated
from
other
groups.
-‐ In
ecology
a
population
is
a
group
of
individuals
of
the
same
species
inhabiting
the
same
area
at
the
same
time.
A
population
can
be
a
discrete
“natural”
unit,
e.g.
the
Blue
gill
sunfish
of
Conesus
Lake
in
the
USA,
or
a
population
maybe
arbitrarily
chosen
by
the
investigator,
e.g.
the
elephant
population
of
Hwange
National
Park.
Characteristics of Populations
(i) Distribution -‐ geographical range, or areal extent
(ii) Dispersion – spacing between individuals
(iii) Abundance – density in a given area or total number in the population
(iv) Age structure – relative or absolute numbers in different age classes
(v) Genetic Composition -‐
We
might
also
want
to
know
the
following
about
the
dynamics
of
the
population
(i.e.
its
changes
over
time):
(i) Dispersal and migration – movement through space
(ii) Population dynamics – change in number (or density) over time due to births,
deaths, immigration or emigration.
(iii) Changing age structure (demographics)
(iv) Changing genetic composition – evolution
Distribution of populations
-‐ Distribution
is
the
geographical
extent
of
a
population
(or
any
other
ecological
unit
such
as
a
species
or
community).
-‐ Biogeography
is
the
scientific
study
of
the
distribution
of
organisms.
With respect to distribution we can ask questions such as:
“Why are individuals of a given species present in some species but absent from others?”
“What determines (or limits) the geographical range of a population or species?”
The
distribution
of
a
population
is
determined
by
a
number
of
factors:
1. Dispersal
–
if
a
species
is
not
found
in
a
particular
area
we
can
ask
the
question;
is
the
species
absent
because
its
“propagules”
(seeds,
spores,
larvae,
adults,
etc.)
did
not
or
could
not
reach
the
area?
–
due
to
some
geographical
barrier
and/or
limited
dispersal
ability.
The
question
can
be
answered
by
transplant
experiments.
These
involve
the
experimenter
moving
some
individuals
to
an
area
where
they
are
not
found
to
see
if
they
can
survive,
grow
and
reproduce
there,
e.g.
the
sugar
maple
was
transplanted
to
Europe
and
succeeded
(a
good
habitat),
further
north,
it
failed
(because
it
was
too
cold),
further
south,
it
failed
(because
it
was
too
hot),
and
further
west,
it
also
failed
(because
it
was
too
dry).
Such
experiments
can
help
to
determine
the
potential
range
(ecological
range,
i.e.
all
sites
within
a
species’
fundamental
niche)
of
a
species.
2. Behaviour
(Habitat
selection)
–
in
some
instances
an
organism
or
species
can
survive
in
an
area
if
introduced
and
restrained
there,
but
does
not
occur
there
naturally
(even
if
its
propagules
reach
the
area).
This
may
be
due
to
the
fact
that
the
organism
might
be
limited
by
an
inflexible
behavioural
habitat
preference
–
i.e.
individuals
do
not
see
any
suitable
habitat,
and
they
continue
to
move
on.
This
behaviour
may
have
evolved
under
circumstances,
where
it
was
advantageous
(i.e.
had
adaptive
value)
in
the
old
habitat,
but
is
maladaptive
here.
3. Interactions with other species – parasites, diseases, prey, competitors, predators,
etc.
A
species
may
suffer
severely
in
increased
mortality
and/or
reduced
fecundity
due
to
intense
parasitism,
disease
outbreaks,
food
shortages,
competition
or
predation.
In
order
to
find
out
whether
the
absence
of
a
particular
species
in
a
particular
area
is
due
to
interactions
with
other
species,
an
investigator
could
perform
transplant
experiments
that
also
exclude
suspected
interacting
species.
4. Physical
and
chemical
factors
–
abiotic
factors
(conditions,
abiotic
resources)
might
be
outside
the
range
tolerated
by
the
species.
For example, temperatures may be too high or too low for the species in question.
Two important considerations to make regarding geographical limitation:
(1) Geographical barriers – areas of inhospitable habitat, in which a species cannot
survive, separating areas of hospitable habitat (some of
which are not occupied), e.g. oceans for terrestrial
species, land for aquatic species, deep water for
shallow water species, mountains for low-‐altitude
species, valleys for high-‐altitude species.
(2)
Dispersal
ability
–
some
species
are
extremely
mobile
(e.g.
birds,
such
as
the
arctic
tern),
while
others
are
stationary,
at
least
as
adults
(e.g.
trees).
The
offspring/dormant
stages
(e.g.
seeds,
spores,
larvae,
etc.)
are
often
the
stage
that
disperses.
The
spread
of
a
species
accidentally
introduced
by
man
into
a
region
where
they
never
previously
occurred
(i.e.
“exotic”
or
“introduced”
species)
illustrates
the
dispersal
ability
of
a
species,
e.g.:
(i) Gypsy
moth
–
introduced
to
America
by
a
French
astronomer
working
at
Harvad
University
near
Boston
in
1850.
(ii) African
honey
bee
–
an
aggressive
subspecies
introduced
to
Brazil
in
1956
to
improve
honey
productivity.
(iii) Colorado
beetle
–
rare
case
of
spread
from
the
“New
World”
to
the
“Old
World”.
Dispersion – Spacing of individuals (spatial distribution of
organisms)
Dispersion = spatial pattern of distribution of individuals within populations.
The
spatial
distribution
of
all
living
organisms
depends
on
a
variety
of
factors
such
as
soil
type,
moisture
conditions,
temperature
variations,
the
presence
of
neighbours
or
competitors
and
so
on.
Distributions
may
change
with
time
as
they
are
affected
by
such
things
as
the
movements,
births
and
deaths
of
organisms.
Furthermore,
different
life
stages
of
the
same
species
may
have
different
distributional
patterns.
The individuals of a population can follow three basic types of spatial distribution:
(1) Random
distribution,
in
which
there
is
an
equal
chance
of
an
individual
of
occupying
any
point
in
an
area
irrespective
of
the
position
of
other
individuals.
A
striking
feature
of
random
distribution
is
the
lack
of
any
system,
e.g.
some
individuals
occur
in
groups
and
some
are
equally
spaced,
some
individuals
are
close
together
and
others
are
wide
apart.
Mechanism:
chance
events
(e.g.
passive
seed
dispersal
by
wind);
no
interaction
between/among
individuals.
(2)
Regular
(or
uniform,
even,
spaced,
or
over
dispersion)
distribution,
which
occurs
when
distances
between
individuals
are
approximately
equal
–
more
equally
spaced
than
is
expected
by
chance.
Mechanism:
either
individuals
avoid
each
other
(e.g.
when
animals
are
relatively
crowded
they
move
away
from
each
other
and
tend
to
be
the
same
distances
apart)
or
competition
eliminates
individuals
which
are
too
close
together.
(3)
Clumped
(aggregated,
contagious,
or
under
dispersion)
distribution,
in
which
individuals
are
closer
together
than
is
expected
by
chance.
Random
or
regular
distributions
seldom
occur
in
nature
and
most
populations
occur
in
definite
clumps
or
patches.
Mechanism:
this
comes
about
because
most
environmental
factors
either
are
unevenly
distributed,
i.e.
individuals
are
attracted
to
(or
survive
better
in)
a
given
area
(i.e.
the
habitat
is
“patchy”),
individuals
are
attracted
to
each
other
(gregarious
–
for
protection
against
predation),
or
offspring fail to disperse far from parents.
Temporal Changes in Population Size
Very few, if any, populations grow exponentially for any length of time in nature. We can
only see this in the case of population recovery after a disturbance (storm, drought, volcanic
eruption, etc.) or introduction of “exotic” species.
The
size
(or
density)
of
a
population
can
vary
temporally
(i.e.
over
time).
For
example
the
number
of
breeding
pairs
of
the
Grey
Heron
was
seen
in
the
Thames
Valley
to
fluctuate
during
the
period
1928
–
1970
(fig
1).
Fig.
1:
Population
fluctuations
of
the
Grey
Heron
in
the
Thames
individuals
are
attracted
to
Valley
The
decrease
in
population
size
occurred
in
the
cold
winters.
The
Grey
Heron
feeds
on
fish
and
in
cold
winters,
with
the
water
bodies
capped
with
ice,
food
supply
was
limited
and
this
lowered
the
population
size.
Populations
can
remain
stable
over
a
period
of
time,
they
can
fluctuate
cyclically
or
they
can
continue
to
decline
until
they
become
extinct.
Population fluctuations are caused by four processes (called demographic processes):
1. Mortality – death of individuals (decreases population sizes).
2. Natality (or fecundity) – production of new individuals (increases population sizes).
3.
Emigration
–
movement
of
individuals
out
of
the
population
(by
migration,
dispersal,
etc.)
(decreases
population
sizes).
4. Immigration – movement of individuals into a population (by migration, dispersal,
etc.) (increases population sizes).
We
can
summarise
how
these
four
processes
affect
population
size
over
time
in
the
following
equation:
Nnow
=
Nthen
+
B
–
D
+
I
–
E
Where:
Nnow = number (or density) at present time
Nthen = number (or density) at some time in the past
B = number of births over that time
D = number of deaths over that time
I = number of immigrations over that time
E = number of emigrations over that time
Similarly, we can predict the future population size (or density) as
Nfuture
=
Nnow
+
B
–
D
+
I
–
E
or
Nt+1 = Nt + B – D + I -‐ E
Here we will consider only the first two of these processes – birth and death.
Natality
(or
fecundity)
–
production
of
new
individuals
by
birth,
hatching,
germination
or
fission.
Mortality
–
loss
of
individuals
due
to
their
death
–
can
be
due
to
interactions
with
other
organisms
(e.g.
diseases,
parasitism,
predation,
starvation,
etc.)
or
loss
of
homeostasis
in
the
face
of
environmental
challenge
(change
in
conditions
=
physiological
death).
“Control”
of
population
numbers
can
result
from
the
density
dependence
of
mortality
and/or
fecundity.
A
processes
is
said
to
be
density
dependent
if
its
and
outcome
depend
on
the
number
of
individuals
in
the
population.
Density-‐dependence
reflects
intra-‐specific
competition
in
which
individuals
of
the
same
species
compete
amongst
each
other
for
food,
space,
light
or
other
resources.
The
tendency
for
N
to
increase
when
it
is
low,
and
to
decrease
when
it
is
high
results
in
logistic
growth.
Possible
patterns:
-‐ Both birth and death rates are density-‐dependent
-‐ Only birth rate is density-‐dependent
-‐ Only death rate is density-‐dependent
Stability and fluctuations of Populations
Population
fluctuations
(up
and
down
movements
on
the
graph)
are
due
to
environmental
changes,
predators,
parasites,
genotype,
mutations,
etc.
Equilibrium is a state of a system which does not change.
Any
equilibrium
may
be
stable
or
unstable.
For
example,
the
equilibrium
of
a
pencil
standing
on
its
tip
is
unstable;
that
of
a
picture
frame
on
the
wall
is
(usually)
stable.
An
equilibrium
is
considered
stable
if
the
system
returns
to
it
after
small
disturbances.
If
the
system
moves
away
from
the
equilibrium
after
small
disturbances,
then
the
equilibrium
is
unstable.
Types of population fluctuations
Three kinds of changes in population size may occur according to the discrete-‐time logistic
model (Ricker’s model). The discrete-‐time logistic model
Nt+1 Nt R [(K -‐ N)/K] can be written as Nt+1 = Ntexp[r0(1-‐Nt/K)]
This
version
of
the
discrete-‐time
logistic
model
(Ricker’s
model)
can
be
used
to
generate
simulations
of
the
three
types
of
population
dynamics,
which
are
as
follows:
1. Equilibral
This
is
where
we
can
get:
(a) Monotonous
increase
in
numbers
=sigmoidal,
described
by
the
logistic
equation,
(b) Damping
oscillations.
Fig.
2.1:
Monotonous
increase
in
numbers
under
the
discrete-‐time
logistic
model
Fig.
2.2:
Damping
oscillations
under
the
discrete-‐time
logistic
model
In
figs
2.1
and
2.2
the
model
has
stable
equilibrium,
only
the
patterns
of
approaching
the
equilibrium
are
different.
In
fig.
2.2
(Damped
oscillations)
N
increases
when
it
is
below
the
carrying
capacity,
K
and
then
decreases
after
overshooting
K,
and
then
stabilises
when
N=K.
In
this
case
the
variations
in
population
size
are
marked
at
first
and
then
decrease
with
time.
The
population
is
going
towards
stability,
e.g.
when
pests
are
disturbed
by
pesticides
and
then
develop
pesticide
resistance.
In
other
words,
the
magnitude
of
the
oscillations
decreases
over
time
until
eventually
N
stabilises
at
K.
Ricker’s
model
is
stable
if
0
<
r
<
2.
The
question
is
what
happens
to
model
populations
if
stability
is
lost?
Non-‐equilibrium
dynamics
may
be
of
two
types:
limit
cycle
–
when
the
trajectory
repeats
itself,
and
chaotic
when
the
trajectory
does
not
repeat
itself.
2. Cyclic (or periodic or limit cycle)
Populations that cycle can do so with various periods. A population cycling with a
period 2 repeats itself every two-‐time steps (Fig. 2.3).
Fig.2.3: Limit cycle with period=2 under the discrete time logistic model
A population cycling with a period 4 repeats itself every four-‐time steps (Fig. 2.4).
Fig.2.4: Limit cycle with period = 4 under the discrete time logistic model
3. Chaotic
(fluctuations,
but
without
a
regular
period).
Chaotic
populations
never
exactly
repeat
themselves
(fig.2.5).
Fig.2.5:
Limit
cycle
with
period
=
4
under
the
discrete
time
logistic
model
Natural populations appear to show all of these different kinds of dynamics.
Lecture
3:
POPULATION
GROWTH
Module:
Principles
of
Ecology
Dept.
of
Applied
Biosciences
and
Biotechnology,
Midlands
State
University
3.1. Population Growth
3.2. Population Growth forms
3.3. Isolated populations, unlimited environments and the exponential model
3.4. Isolated populations, in limited environments and the logistic model
___________________________________________________________________________
POPULATION
GROWTH
Self-‐reproduction
is
the
main
feature
of
living
organisms.
This
is
what
distinguishes
them
from
non-‐living
things.
Population
growth
is
essentially
a
multiplicative
process
(i.e.
populations
grow
by
multiplication
not
by
addition)
and
is
continuous
where
there
is
a
complete
overlap
of
generations.
Populations
grow
by
birth
and
immigration
and
decline
by
death
and
emigration.
If
birth
+
immigration
exceed
death
+
emigration,
the
population
grows
and
vice
versa.
The
rate
at
which
individuals
are
added
to
or
removed
from
the
population
by
birth
and
death
is
a
function
of
the
size
of
that
population.
Population Growth Forms
Population have characteristic patterns of increase which are called population growth
forms. These are the Exponential growth form for isolated populations in unlimited
environments and the logistic growth form for isolated populations in limited
environments.
The
shapes
of
the
characteristic
curves
of
these
two
population
growth
forms
are
the
J-‐
shaped
curve
for
exponential
growth
and
S-‐shaped
or
sigmoid
curve
for
logistic
growth:
Exponential
and
logistic
growth
models
help
to
solve
different
kinds
of
problems
in
ecology,
e.g.:
(i) How
long
will
it
take
for
a
population
to
grow
to
a
specific
size?
(ii) What
will
be
the
population
size
after
n
years
(or
generations)?
(iii) How
long
can
a
population
survive
at
non-‐favourable
conditions?
Isolated Populations in Unlimited Environments (Density-‐
independent Population Growth)
We will start by considering an isolated population (i.e. one not interacting with other
populations through predation, parasitism, competition, etc.) in an unlimited environment
(where growth is unlimited because resources are not limiting).
We can consider two possible situations:
Discrete Generations (and the Discrete-‐Time Exponential model)
Here the species has a discrete (short) breeding season and a life span of less than a year,
i.e. generations do not overlap, e.g. annual plants and “univoltine” insects.
Changes
in
population
size
in
this
case
can
be
described
by
the
Discrete-‐Time
Exponential
Model:
Nt+1
=
RoNt
Where:
t is time measured in generations
Ro is the net reproduction rate
Nt is the population size (females) at generation t
Nt+1 is the population size (females) at generation t+1
For monovoltine organisms, R is the average number of offspring per one parent. For
example, in univoltine insects with a sex ratio of 1:1, R = Fecundity/2.
If
all
the
offspring
survive
to
breed
the
following
season
the
population
size
after
one
time
step
will
be
given
by:
Nt+1
=
RoNt
Ro
here
can
be
called
the
multiplication
constant.
Here
we
are
taking
discrete
time
steps
which
can
be
described
by
a
difference
equation:
Nt+1 /Nt = Ro
The
discrete-‐time
exponential
model,
Nt+1
=
RoNt
is
a
difference
equation
and
it
gives
the
number
of
individuals
that
will
be
in
the
population
after
a
single
time
step
(e.g.
after
one
generation).
If
we
assume
Ro
is
constant
over
time,
we
can
calculate
the
size
of
future
generations
as
in
the
table
below:
Starting with an initial population of 10 individuals (N0 = 10) and R0 of 1.5:
GENERATION POPULATION SIZE (Nt)
0 10
1 15=(1.5)(10)
2 22.5=(1.5)(15)
3 33.75=(1.5)(22.5)
If
we
plot
this
sort
of
data
we
get
population
growth
curves
with
discrete
time
steps
but
they
will
be
J
–
shaped.
Note:
Different
R
values
give
different
curves.
Higher
R
means
the
population
grows
faster.
Different
N0
values
give
curves
of
the
same
shape.
*By
analogy,
if
you
put
$10
in
the
bank
at
5%
interest
you
w ill
get
a
lot
less
m oney
in
20
years
than
if
you
put
% 100
in
the
bank,
although
in
both
cases
your
balance
grows
in
the
same
way.
The
table
above
also
shows
that
the
size
of
the
increment
in
the
population
in
one
time
step
depends
on
the
size
of
the
population
at
that
time.
Populations
therefore
grow
by
multiplication
(proportional
or
geometric
increase)
rather
than
by
addition
(absolute
or
arithmetic
increase).
Overlapping Generations (and the Continuous-‐Time Exponential
model)
While the discrete-‐time exponential model is used to describe the growth of isolated
populations with discrete generations in unlimited environments, the Continuous-‐Time
Exponential model describes the growth of isolated populations, with overlapping
generations, living in unlimited environments.
Here
we
species
that
have
prolonged
or
continuous
breeding
(i.e.
no
specific
breeding
season)
and
overlapping
generations
and
there
is
need
for
a
continuous
model
(no
discrete
time
steps)
which
produces
a
smooth
curve.
This
kind
of
population
growth
is
best
described
by
the
continuous-‐time
exponential
equation:
dN/dt = rN = (b-‐d)N
Here dN/dt is the rate at which the population is growing at each instant.
In
the
continuous-‐time
exponential
model,
we
have
modelled
an
instantaneous
rate
of
growth,
not
the
number
that
will
be
there
after
a
single
time
step
(or
next
year).
The
continuous-‐time
exponential
model
means
that
changes
in
the
population
size
instantly
feedback
on
the
population’s
growth
rate.
If
we
add
a
single
individual
right
now,
the
population’s
growth
rate
will
instantly
change
from
rN
to
r(N+1).
The
discrete-‐time
exponential
model
means
that
the
number,
say
of
insects
hatching
in
the
spring
of
next
year
depends
on
the
number
of
insects
hatching
in
the
spring
of
this
year.
There
is
no
instantaneous
adjustment
of
the
population
growth
rate.
In
the
continuous-‐time
exponential
model,
rather
than
take
discrete
(relatively
large)
time
steps
as
in
the
discrete-‐time
exponential
model,
we
take
infinitesimally
small
time
steps
and
population
growth
can
be
described
by
a
differential
equation:
dN/dt = rN = (b-‐d)N
Where:
N = population size
t = time
r =intrinsic rate of increase (or per capita rate of population growth, etc.)
b = instantaneous birth rate
d = instantaneous death rate
Plotting the output of this differential equation yields an exponential growth curve:
The
integral
form
of
this
equation
is
Nt/No
=
ert
We
can
rearrange
this
to
give
N
at
any
given
time
as:
Nt
=
Noert
There
are
three
possible
model
outcomes:
(i) Population exponentially declines (r< 0)
(ii) Population exponentially increases (r> 0)
(iii) Population does not change (r = 0)
Both
the
continuous-‐time
and
discrete-‐time
models
of
density-‐independent
growth
result
in
exponential
growth.
When
a
population
is
increasing
exponential,
growth
continues
at
a
constant
rate
without
limit.
The
continuous-‐time
and
discrete-‐time
exponential
models
sometimes
do
have
similar
model
outcomes,
but
not
always:
→ Population exponentially declines if:
R < 0 (in the discrete-‐time model)
r < 0 (in the continuous-‐time model)
→ Population exponentially declines if
R = 1 (in the discrete-‐time model)
r = 0 (in the continuous-‐time model)
Parameter r is called:
-‐ Malthusian parameter,
-‐ Intrinsic rate of increase,
-‐ Instantaneous rate of natural increase, or
-‐ Population growth rate.
Assumptions of the Continuous-‐time Exponential Model
(i) Continuous reproduction (e.g. no seasonality),
(ii) All organisms are identical (e.g. no age structure), and
(iii) Environment is constant in space and time (e.g. resources are unlimited).
However,
the
exponential
model
is
robust,
it
gives
reasonable
precision
even
if
these
conditions
do
not
hold.
Organisms
may
differ
in
age,
survival,
and
mortality,
but
the
population
consist
of
a
large
number
of
organisms
and
their
birth
and
death
rates
tend
to
average.
Note
that
in
the
exponential
growth
form,
population
density
increases
rapidly
in
exponential
fashion
and
then
stops
abruptly
as
environmental
resistance
or
another
limiting
factor
becomes
effective
more
or
less
suddenly.
Applications
of
the
Exponential
Models
These
models
accurately
describe
the
initial
rate
of
growth
of
some
populations
and
therefore
find
application
in:
-‐ Microbiology
(e.g.
the
growth
of
bacteria
in
petri
dishes)
-‐ Conservation
biology
(restoration
of
disturbed
populations)
-‐ Fishery
(prediction
of
fish
dynamics)
-‐ Plant
and
insect
quarantine
(species
invading
new
habitats
often
grow
exponentially)
Isolated Populations in limited Environments (Density-‐dependent
Population Growth)
The Logistic Model
We
very
seldom
see
the
kind
of
explosive
population
growth
described
by
the
exponential
model.
Most
environments
are
limited
in
terms
of
resources,
and
growth
rate
depends
on
the
population’s
current
size
(N).
Even
if
an
organisms
is
released
into
a
good
habitat
with
a
large
quantity
of
available
resources,
as
its
population
size
grows,
the
amount
available
for
each
individual
becomes
limiting.
The
growth
of
the
population
slows
(by
increasing
mortality
or
decreasing
fecundity
or
both)
as
density
increases.
In
this
case
population
growth
is
density-‐dependent
and
is
best
described
by
the
logistic
model
which
produces
an
S-‐shaped
growth
curve.
The
population
begins
to
grow
exponentially,
but
as
population
size
increases,
the
rate
of
growth
declines.
Population
size
then
slowly
approaches
a
maximum
value,
K,
the
carrying
capacity
of
the
population.
K
is
the
maximum
number
(or
density)
the
habitat
can
sustain,
and
is
the
upper
asymptote
of
the
sigmoid
curve.
Each
individual
in
our
population
I
competing
with
other
members
of
the
population
for
limiting
resources,
i.e.
there
is
intra-‐specific
competition.
The
logistic
model
was
developed
by
Belgian
Mathematician
Pierre-‐Verhulst
(1838)
who
suggested
that
the
rate
of
population
increase
may
depend
on
population
density.
Density-‐dependent
population
growth
can
best
be
described
by
the
continuous-‐time
logistic
equation
for
populations
with
overlapping
generations,
and
the
discrete-‐time
logistic
equation
Nt+1 =R0Nt[(K – N)/K]
for populations with discrete generations. In both models, when the population is very
small, nearly 0 in size (K -‐ N)/K is nearly 1, so the population undergoes density-‐independent
growth. Similarly, when the population is at carrying capacity, N = K and so (K -‐ N)/K = 0 and
the population does not grow at all.
Thus, the population has density-‐independent growth at low density, and density-‐
dependent growth at high density.
Discrete Generations (and the Discrete-‐Time logistic model)
Here our species with a discrete (short) breeding season and a life span of less than a year
(i.e. generations do not overlap) is now living in a limited environment in which resources
will become limiting when a certain population size is reached.
The discrete-‐time logistic equation
Nt+1 =R0Nt[(K – N)/K]
is identical to the discrete-‐time exponential equation for population growth in unlimited
environments except that R0Nt is now multiplied by a new term (K – N)/K, which can be
viewed as a proportion of available “space” (or resources) left for any given N.
(K
–
N)
is
the
amount
of
available
space
(total
amount
of
space
minus
amount
already
occupied),
while
K
is
the
total
amount
of
space.
Parameter
K
can
also
be
interpreted
as
the
amount
of
resources
expressed
in
the
number
of
organisms
that
can
be
supported
by
these
resources.
The discrete-‐time logistic equation
Nt+1 =R0Nt[(K – N)/K]
is also a difference equation and it gives the number of individuals that will be in the
population after a single time step. In other words, the discrete-‐time logistic model treats
time as occurring in clear steps, lets say, of a year and describes the number of organisms at
the next time step.
Plotting
the
output
of
this
difference
equation
yields
an
S-‐shaped
or
sigmoid
curve:
Overlapping Generations (and the Continuous-‐Time logistic model)
Here we have species that have prolonged or continuous breeding and overlapping
generations living in limited environments.
Because
generations
overlap,
population
growth
is
best
described
by
a
differential
equation
which
produces
a
smooth
curve:
dN/dt = rN[(K – N)/K)]
This
is
a
rate
equation,
and
dN/dt
is
the
rate
at
which
the
population
is
growing
at
each
instant.
We
have
also
modelled
an
instantaneous
rate
of
growth,
not
the
number
that
will
be
there
after
a
single
time
step
(or
next
year).
This
equation
is
identical
to
the
continuous-‐time
exponential
equation
for
population
growth
in
unlimited
environments
except
that
rN
is
now
multiplied
by
a
new
term
(K
–
N)/K,
which
takes
into
account
the
effect
of
intra-‐specific
competition
between
the
individuals.
Just
like
in
the
continuous-‐time
exponential
model,
changes
in
the
population
size
instantly
feedback
on
the
population’s
growth
rate.
If
we
add
a
single
individual
right
now,
the
population’s
growth
rate
will
instantly
change
from
rN
to
r(N+1),
orif
you
added
a
lot
of
individuals
to
the
population,
so
that
there
were
more
than
K,
the
population
would
reduce
to
about
K
very
quickly.
Thus,
in
the
continuous-‐time
logistic
model
the
population’s
growth
rate
adjusts
itself
instantaneously,
so
that
there
is
a
gradual
slowing
of
growth
as
N
increases.
Consequently,
the
population
can
never
overshoot
its
carrying
capacity.
Under
the
discrete-‐time
logistic
model,
it
is
possible
for
the
population
to
overshoot
its
carrying
capacity.
There
is
no
instantaneous
adjustment
of
the
population
growth
rate.
The
discrete-‐time
logistic
model
tells
us
something
about
what
happens
when
the
effects
of
density-‐dependence
are
not
instantaneous,
but
lag
behind
the
population’s
growth
in
time.
In
both
models
of
logistic
growth,
population
growth
rate
declines
with
population
size,
N,
and
reaches
0
when
N
=
K.
if
the
population
size
exceeds
K,
then
the
population
growth
rate
becomes
negative
and
population
size
declines.
The
logistic
equations,
therefore
describe
changes
in
rates
of
growth
of
populations.
Growth
is
slow
at
first,
becomes
faster
and
faster
until
a
maximum
is
reached
and
then
falls
away
finally
to
zero
as
the
system
reaches
equilibrium
(where
birth
rate
=
death
rate).
The logistic model has two equilibria: N = 0 and N = K.
The
first
equilibrium
is
unstable
because
any
small
deviation
from
this
equilibrium
will
lead
to
population
growth.
For
example,
immigration
of
a
small
number
of
organisms
results
in
growth
of
a
population.
The
population
never
returns
to
its
equilibrium.
Instead
population
numbers
increase
until
they
reach
the
stable
equilibrium
N
=
K.
N
=
K
is
stable
because
after
a
small
disturbance
the
population
returns
to
its
equilibrium.
The
dynamics
of
the
discrete-‐time
logistic
model
are
similar
to
those
of
the
continuous-‐time
logistic
model
if
the
population
growth
rate
is
small
(0
<
R
<
0.5).
However,
if
the
population
growth
rate
is
high,
then
the
model
may
exhibit
more
complex
dynamics
such
as
damping
oscillations,
cycles
or
chaos
because
of
a
time
delay
in
feedback
mechanisms.
There
are
no
intermediate
steps
between
time
t
and
time
t+1.
Thus,
over
compensation
may
occur
if
the
population
grows
or
declines
too
fast
passing
the
equilibrium
point
(K).
Assumptions of the Continuous-‐Time logistic model
(i) The feedback mechanism is instantaneous.
(ii) There is complete overlap of generations
(iii) Constant carrying capacity
(iv) All individuals equal in reproductive potential.
The
provisors
of
the
logistic
growth
are
met
in
the
growth
of
yeast
and
floating
pond
weed
in
culture.
In both cases the sigmoid growth pattern is a close approximation to the logistic function.
The
feedback
mechanisms
are
the
production
of
alcohol,
which
limits
the
growth
of
the
thin-‐walled
daughter
yeast
cells
after
budding,
and
mutual
shading
of
the
pond
weed
fronds,
which
limits
photosynthesis.
Both
mechanisms
regulate
the
populations
below
levels
where
nutrients
in
the
culture
are
exhausted.
However,
not
all
populations
show
the
logistic
growth.
In
many
cases,
the
population
will
briefly
overshoot
its
carrying
capacity,
then
falls
back
down
below
K,
etc,
resulting
in
oscillations.
This
overshoot
of
K
and
resultant
oscillations
is
typically
due
to
a
time
lag
between
the
acquisition
of
resources
and
the
production
of
offspring,
i.e.
the
number
of
offspring
produced
when
the
population
reaches
K
may
actually
have
been
set
some
time
when
N
was
still
below
K,
etc.
Applications of the logistic Model
-‐ Microbiology (growth of yeast),
-‐ Biological control (introduction of the natural enemy),
-‐ In managing game reserves,
-‐ Cattle ranging and
-‐ In the study of almost all populations with a strong interaction among individuals.
Lecture
4:
ESTIMATION
OF
POPULATION
SIZE
AND
DENSITY
Module:
Principles
of
Ecology
Dept.
of
Applied
Biosciences
and
Biotechnology,
Midlands
State
University
4.1. Censuring a whole population
4.2. Quadrat Sampling
4.3. Simple Random or Systematic Sampling
4.4. Stratified Sampling
4.5. Capture-‐Mark-‐Recapture Method
___________________________________________________________________________
Estimation
of
Population
Size
and
Density
Recall
we
define
ecology
as
the
scientific
study
of
interactions
that
determine
the
abundance
and
distribution
of
organisms
–
here
we
concentrate
on
the
abundance
of
populations.
Two different measures of abundance can be considered:
(i) Total
number
of
individual
in
the
entire
population
–
seldom
measurable
(ii) Density
=
the
number
of
individuals
in
a
population
per
unit
area
(e.g.
number/m2
or
number/ha)
or
volume
(e.g.
number/m3
or
number/litter).
Censuring a whole population (total counts)
In cases where individuals are readily detected (often by sight), population size can be
determined by direct counts of all individuals in an area, especially if their numbers are
small and the area is well bounded and not too large.
Example: trees in a small isolated forest.
Migrating
populations
can
be
censured
using
aerial
photography.
This
method
is
often
used
when
the
population
has
seasonal
migration.
When
numbers
are
too
large
or
effort
to
obtain
total
counts
would
be
too
great
(
especially
when
organisms
are
cryptic),
the
experimenter
can
count
only
a
small
but
representative
portion
of
the
population
and
use
this
sample
to
estimate
the
population
size.
Quadrat
Sampling
-‐
good
for
sessile
organisms
(e.g.
plants,
sessile
animals
like
adult
barnacles
and
mussels).
-‐
the
site
is
marked
off
in
grid,
and
a
few
quadrats
are
randomly
selected
for
intensive
study.
-‐
the
results
are
used
to
estimate
the
density
of
the
entire
population
(i.e.
mean)
as
well
as
reliability
of
the
sampling
estimate
(i.e.
variance).
Simple random or Systematic Sampling
-‐ Two possible objectives for sampling:
(i) to estimate average population density, and
(ii) to make a map of population density.
Traditionally,
random
sampling
was
preferred
to
systematic
sampling
because
random
sampling
helped
to
avoid
subjective
selection
of
sampling
locations.
However,
systematic
sampling
has
no
elements
of
subjectivity
if
the
sample
location
is
selected
prior
to
examining
the
area.
For
example,
there
are
no
subjective
decisions
if
we
sample
every
tenth
potato
plant
and
count
the
number
Colorado
potato
beetles
on
each
plant.
Moreover,
systematic
sampling
has
an
advantage
over
random
sampling
if
the
number
of
samples
is
large
because
of
more
uniform
coverage
of
the
entire
area.
It
is
especially
important
for
making
population
maps.
Random
sampling
can
be
used
if
the
objective
is
to
estimate
the
mean
population
density
and
the
number
of
samples
is
not
large.
Traditional
statistical
methods
include
estimation
of
the
mean
population
density
(M),
standard
deviation
(S.D),
and
standard
error
(S.E),
which
is
the
standard
deviation
of
the
sample
mean.
S.D = S.D or S_ = Sx
√N X √N
The equation for standard deviation is derived assuming that all samples are independent.
This is a very strong assumption which is unrealistic in many situations. Samples separated
by a small distance are often positively correlated. Before using standard statistics it is
important to test if the samples are correlated. This is however, beyond the scope of this
module.
Stratified
sampling
Stratified
sampling
is
used
if
the
sampled
area
(or
volume)
is
heterogeneous.
In
stratified
sampling,
the
area
is
subdivided
into
two
or
more
portions
which
are
sampled
separately.
For
example,
pine
sawflies
prefer
to
spin
their
cocoons
close
to
the
tree,
thus
the
area
adjacent
to
trees
(within
1m
radius)
can
be
sampled
separately
from
the
rest
of
the
area.
The
mean
population
density,
M,
is
estimated
as
a
weighted
mean
of
the
mean
densities
Mi,
in
each
stratum,
i,
with
weights,
wi
equal
to
the
area
covered
by
statum
i:
M
=
∑MiWi
The
standard
error,
S.E,
of
the
mean
is
equal
to:
SE
=
√∑SEi2.Wi2
Where:
SEi
is
the
standard
error
for
the
mean
in
stratum
i.
Capture-‐Mark-‐Recapture
Sampling
This
method
is
good
for
mobile
organisms
(e.g.
most
animals).
In
the
first
step
of
this
method,
a
sample
of
individuals
is
captured
from
an
animal
population,
marked
in
some
way
(e.g.
by
painting,
tagging,
clipping
ears
or
toes,
etc.)
and
released.
The marked animals are given sufficient time to mix back into the population.
A
second
sample
is
then
taken
from
the
population.
By
assuming
that
the
ratio
of
the
marked
animals
to
the
total
number
of
animals
in
the
recaptured
sample
is
the
same
as
the
ratio
of
the
individuals
in
the
first
sample
to
the
total
number
of
individuals
in
the
whole
population,
the
size
of
the
population
can
be
estimated.
Suppose
the
population
is
of
size
N,
such
that
N
is
the
number
we
wish
to
estimate.
Suppose
M
organisms
were
captured,
marked
and
released
back
into
the
population.
After
some
time
which
should
be
sufficient
for
the
organisms
to
mix,
n
organisms
were
captured,
and
m
of
these
were
marked.
The
proportion
of
the
recaptured
organisms
is
assumed
to
be
the
same
as
the
proportion
of
the
marked
organisms:
m/n
=
M/N
Population
size
can
be
found
as:
N = (nM)/m (the Lincoln Index)
Where:
N = population size you wish to estimate
M = number of animals captured, marked, and released
n = total number of individuals in the recapture sample
m = number of marked individuals in the recapture sample
For
example,
you
capture
and
mark
100
animals,
you
recapture
200
animals
and
find
that
50
of
them
(i.e.
¼)
were
marked,
your
estimate
of
the
population
size
will
be:
N
=
nM
=
200
x
100
m
50
=
4/1
x
100
= 400 animals
The following conditions should be met:
(i) No
immigration,
emigration,
births
or
deaths
between
the
release
and
capture
times.
(ii) The
probabilities
of
being
caught
are
equal
for
all
individuals
(including
the
marked
ones),
i.e.
the
marks
should
not
affect
the
behaviour
of
the
marked
animals
in
any
way.
(iii) Marks
(or
tags)
are
not
lost
and
are
always
recognizable.
Lecture
5:
CONCEPTS
OF
HABITAT
AND
NICHE
Module:
Principles
of
Ecology
Dept.
of
Applied
Biosciences
and
Biotechnology,
Midlands
State
University
5.1. The Niche Concept
5.2. Niche Segregation and Competition
5.3. Overlapping Niches and Competition
5.4. Character displacement and Competition
___________________________________________________________________________
The
Niche
concept
The
habitat
of
an
organism
is
the
place
where
it
leaves.
The
ecological
niche,
on
the
other
hand
is
a
more
inclusive
term
that
includes
not
only
the
physical
space
occupied
by
an
organism
but
also
its
functional
role
in
the
community
(e.g.
its
trophic
position)
and
its
position
in
environmental
gradients
of
temperature,
pH,
soil
and
other
conditions
of
existence.
Organisms
of
any
given
species
can
survive,
grow,
reproduce
and
maintain
a
viable
population
only
within
certain
temperature
limits.
Their
range
of
temperature
is
the
species’
ecological
niche
in
one
dimension.
Organisms
of
the
species
in
question
may
also
be
able
to
survive,
grow,
reproduce
and
maintain
a
viable
population
only
within
certain
limits
of
relative
humidity.
Taking
temperature
and
relative
humidity
together,
the
niche
becomes
two-‐dimensional
and
can
be
visualised
as
an
area
and
so
on.
The
niche
has
been
variously
defined
as
either
the
predominant
trophic
role
of
an
organism
within
a
community
(by
Charles
Elton,
1927)
or
the
range
of
physical
environments
in
which
a
species
can
live
(by
Joseph
Grinnell,
1917).
The
first
definition
emphasised
the
“profession”
of
the
species
and
the
second
one
emphasised
its
“address”.
Evelyn
G.
Hutchinson
(1957)
defined
a
niche
as
a
multi-‐dimensional
space
or
hyper
volume
within
which
the
environment
permits
an
individual
or
species
to
survive
indefinitely.
The
definition
is
more
close
to
Grinnell’s
definition.
It
became
popular
because
the
range
of
tolerance
to
ecological
factors
can
be
easily
measured,
whereas
species’
“profession”
cannot
be
measured
easily.
Hutchinson
also
defined
the
Fundamental
Niche
of
a
species
as
the
extent
of
its
total
environment
which
it
could
potentially
exploit
in
the
absence
of
other
species
and
its
Realized
Niche
as
the
resources
which
it
actually
utilizes
when
other
species
are
present.
Niche segregation and competition
Within the same species competition is often reduced when different stages in the
organism’s life history occupy different niches, e.g. the tadpole functions as a herbivore and
the adult frog as a carnivore in the same pond. Niche segregation may even occur between
the sexes, e.g. in the woodpecker of the genus Dendrocarpus the male and the female differ
in bill size and foraging behaviour.
In
Anopheline
mosquitoes,
females
are
blood
suckers
whilst
males
feed
on
plant
juices.
This
means
there
is
no
competition
for
resources
between
the
sexes
in
these
species.
Overlapping
Niches
and
competition
Fig
1:
Distribution
of
Platyhelminth
flat
worms
Planaria
gonocephala
and
P.
montengrina
along
temperature
gradients
in
streams
where
they
occur
together
(sympatry)
and
alone
(allopatry)
The
above
example
illustrates
that,
if
two
competing
species
coexist
in
a
stable
environment,
then
they
do
so
as
a
result
of
niche
differentiation,
i.e.
differentiation
of
their
realized
niches.
If,
however,
there
is
no
differentiation,
one
of
the
competing
species
will
dominate
or
exclude
the
other.
Character displacemet: Sympatry and Allopatry
Differences in closely related species are often accentuated in sympatry (when species occur
in the same area) and weakened in allopatry (when species occur in different geographical
areas) by an evolutionary process called character displacement.
Character displacement has two adaptive values:
(i) It enhances niche displacement thus reducing competition, and
(ii) It enhances genetic segregation by maintaining species distinctiveness (i.e.
preventing hybridization) and thereby maintaining greater species diversity in
the community.
Specialists and Generalists
A
species
which
exploits
a
narrow
niche
is
called
a
specialist
while
one
which
exploits
a
broad
niche
is
called
a
generalist.
Lecture 6: COMPETITION
Module: Principles of Ecology
Dept. of Applied Biosciences and Biotechnology, Midlands State University
6.1. Introduction
6.2. Intra-‐specific
competition
6.2.1. Scramble
competition
6.2.2. Contest
competition
___________________________________________________________________________
COMPETITION
Introduction
Competition
is
the
use
or
defence
of
a
resource
by
one
individual
that
reduces
the
availability
of
that
resource
to
other
individuals,
of
the
same
species
or
of
other
species.
In
other
words,
competition
is
an
attempt
to
gain
exclusive
or
prior
access
to
a
reserve
which
is
in
limited
supply
and
it
leads
to
a
reduction
in
the
survivorship,
growth
and/or
reproduction
of
the
competing
individuals.
Resource
=
a
substance
or
object
required
by
an
organism
for
normal
maintenance,
growth
and
reproduction.
What
constitutes
a
resource
is
very
much
species-‐
specific,
e.g.:
o Most
photoautotrophs
require
sunlight,
CO2,
H2O
and
a
number
of
inorganic
nutrients
(including
a
source
of
nitrogen,
phosphorus,
potassium,
etc.);
some
also
require
various
organic
molecules
(e.g.
certain
vitamins)
as
well.
o Most
heterotrophs,
on
the
other
hand,
require
organic
molecules
as
an
energy
source
and
various
inorganic
substances
to
maintain
the
composition
of
their
body
fluids
(e.g.
NaCl),
but
may
also
require
specific
organic
molecules
they
cannot
synthesize
themselves
(e.g.
various
vitamins
and
essential
amino
acids).
“Space”
can
also
be
viewed
as
a
resource,
e.g.
nesting
and
roosting
sites
in
birds
and
attachment
sites
in
barnacles
and
mussels.
If
the
supply
of
a
resource
is
scarce
relative
to
its
demand
by
an
organism,
it
is
referred
to
as
a
limiting
resource
for
that
organism.
Competition
is
among
the
most
important
factors
in
population
dynamics
of
many
species.
It
often
determines
the
upper
limit
of
fluctuations
of
population
numbers.
Intra-‐specific
competition
Intra-‐specific
competition
is
competition
between
individuals
of
the
same
species.
Main features of intra-‐specific competition
Intra-‐specific competition leads to decreased rates of resource intake per individual,
perhaps to decreased rates of individual growth, or to decreases in amounts of stored
reserves. They may, in turn, lead to decreases in survivorship and/or decreases in fecundity.
The first common feature of intra-‐specific competition is that its ultimate effect is a
decreased contribution of the individual to the next generation.
The
second
common
feature
of
intra-‐specific
competition
is
that
the
resources
for
which
individuals
compete
must
be
in
limited
supply.
The
third
common
feature
of
intra-‐specific
competition
is
that
the
competing
individuals
are,
in
essence,
equivalent
–
but
in
practice
very
much
less
so.
The
fact
that
they
have
been
classified
as
the
same
species
implies
that
they
have
many
fundamental
features
in
common,
and
they
may
be
expected
to
use
similar
resources
and
react
in
much
the
same
way
to
conditions.
However,
it
should
be
noted
that
the
competing
individuals
are
not
reciprocal.
There
are
many
occasions
when
intra-‐specific
competition
is
very
much
one-‐sided:
a
strong,
early
seedling
will
probably
shade
a
stunted,
late
one;
and
tall
genotypes
of
maize,
for
instance,
will
usually
shade
and
suppress
short
genotypes
of
the
same
species.
Thus,
competing
individuals
of
the
same
species
are
not
entirely
equivalent.
This
lack
of
exact
equivalence
means
that
the
ultimate
effect
of
competition
is
far
from
being
the
same
on
different
individuals.
Weak
competitors
may
make
only
a
small
contribution
to
the
next
generation
or
no
contribution
at
all.
Strong
competitors
may
have
their
contribution
only
negligibly
affected.
Indeed,
a
strong
competitor
may
actually
make
a
larger
proportional
contribution
when
there
is
intense
competition
than
when
there
is
no
competition
at
all.
Thus
competition
can
increase
fitness
in
a
species.
The
fourth
and
final
common
feature
of
intra-‐specific
competition
is
that
the
effect
of
competition
on
any
individual
is
greater,
the
more
competitors
there
are.
The
effects
of
intra-‐specific
competition
are
therefore
said
to
be
density-‐dependent.
Anything
that
has
a
relationship
with
the
density
of
the
population
is
said
to
be
density
dependent.
For example, the continuous-‐time logistic equation:
dN/dt = rN[(K -‐ N)/N]
shows
that
the
rate
of
population
increase
depends
on
population
density.
At
first
population
growth
rate
increases
with
population
density
and
then
decreases
with
density
as
intra-‐specific
competition
creeps
in.
we can talk of the density dependence of a number of parameters, e.g.:
-‐ the dependence of mortality on density
-‐ the dependence of birth rate on density
-‐ the dependence of growth rate on density
With
density
dependent
birth/death
rate,
there
is
a
tendency
of
birth/death
rare
to
increase
with
density.
With
density
dependent
growth
rate,
there
is
a
tendency
of
growth
rate
to
decrease
with
density.
Mortality,
however,
can
be
density
independent.
Mortalities
caused
by
changes
in
weather
conditions
(e.g.
frost,
heat
waves,
etc.);
volcanic
eruptions,
earthquakes,
etc.,
are
density
independent
(in
contrast
with,
lets
say,
mortalities
caused
by
competition
or
food)
and
were
.
Mortalities
caused
by
predation
are
also
classified
as
density
independent
because
they
are
not
determined
by
the
density
of
the
prey
population.
Mortality
of
insects
due
to
pesticide
application
is
also
density-‐independent.
Scramble and contest
-‐ Intra-‐specific competition can be conveniently divided into scramble and contest
competition
Scramble
Competition
Scramble
competition
occurs
where
there
is
exactly
equal
partitioning
of
resources
with
no
clear
losers
or
winners.
All
individuals
within
a
species
are
affected
to
the
same
extent.
One
individual
is
unlikely
to
gain
the
whole
resource
and
under
high
population
levels
this
form
of
interaction
may
result
in
insufficient
of
the
resource
for
all.
Interactions
between
individuals
are
usually
indirect,
i.e.,
competing
individuals
do
not
interact
with
each
other
directly.
Instead,
individuals
respond
to
the
levels
of
a
resource
which
has
been
depressed
by
the
presence
and
activity
of
other
individuals.
Thus,
grasshoppers
competing
for
food
are
not
directly
affected
by
other
grasshoppers,
but
by
the
reduction
in
food
level
and
the
increased
difficult
of
finding
good
food
that
has
been
left
by
the
others.
In
such
cases,
competition
may
be
described
as
exploitation
in
that
each
individual
is
affected
by
the
amount
of
resources
that
remains
after
it
has
been
exploited
by
the
others.
Main features of scramble competition
(i) Interactions between individuals are usually indirect.
(ii) Competition is usually for food, nesting sites or mates.
(iii) Resources must be limiting
(iv) Scramble involves rapid, efficient location and prior use of resources
(v) The resources are rapidly depleted.
(vi) Competitors are approximately equivalent. Each individual does less well than if
it had exclusive access to the resource.
(vii) There is a threshold density above which the population rapidly declines
Fig.1: Plot of mortality against population density
Fig. 1 shows a generalised relationship between population density and percentage
mortality resulting from scramble competition.
Contest
Competition
Contest
Competition
occurs
when
there
is
unequal
partitioning
of
resources
so
that
there
are
clear
winners
and
clear
losers.
Under
the
conditions
of
contest
competition,
the
competitors
interact
in
such
a
way
that
some
individual
retreat
or
are
eliminated
so
that
the
entire
resource
can
be
utilised
by
the
victor.
When
individuals
interact
directly
with
each
other,
one
individual
will
actually
prevent
another
from
occupying
a
portion
of
the
habitat
and
so
from
exploiting
the
resources
in
it.
In
such
cases
competition
may
be
described
as
interference.
This
is
seen,
for
instance,
in
motile
animals
that
defend
territories.
The
result
is
often
that
the
territory
itself
often
becomes
a
resource.
For
example,
Lion
prides
in
Savuti
national
Park
in
Botswana
mark
their
territories
with
urine
and
fiercely
guard
them
against
invasion
by
other
lion
prides.
In
most
cases
fierce
fighting
occurs
and
may
result
in
death
if
the
loser
cannot
escape.
In
many
other
vertebrate
populations,
however,
the
contest
may
become
ritualised
so
that
the
dispute
can
usually
be
settled
without
substantial
damage
to
the
contestants.
A
classical
example
of
this
form
of
interaction
is
the
study
of
woodland
owl
populations
by
Southern
(1970).
The
woodland
was
divided
up
into
a
number
of
territories
through
intense
vocal
conflict
between
the
owls.
Only
pairs
of
birds
holding
a
territory
were
able
to
breed.
The
young
birds
were
expelled
from
the
woodland
and
formed
a
reserve
population
of
animals
which
only
breed
following
the
death
or
successful
challenge
of
the
resident
pair.
Main features of contest competition
(i) Interactions between individuals are direct.
(ii) Interactions are usually for space incorporating critical resources.
(iii) Resources potentially or occasionally limiting.
(iv) Contest involves active exclusion of competitors (i.e., interference).
(v) Competitors are not equivalent, there being clear winners and losers.
(vi) There is a threshold density above which a fixed number of individuals survive.
Fig.2:
Plot
of
k-‐value
against
log
population
density
in
(a)
scramble
competition,
and
(b)
contest
competition.
Fig.
2(a)
shows
that
at
the
threshold
density
(T),
mortality
caused
by
scramble
competition
rises
sharply
from
0%
to
100%.
In
contrast,
the
k-‐value
for
contest
competition
(Fig
2
(b))
rises
with
a
slope
=
1,
i.e.,
a
constant
number
of
individuals
survive.
However,
real
slopes
are
less
than
this
because
of
phenotypic
and
genotypic
variations
between
organisms
(see
plots
with
broken
lines
in
figs
2(a)
and
2(b)).
Figs.
2(a)
and
2(b)
show
that
ain
both
scramble
and
contest
competition
there
is
no
competition
at
all
at
low
densities:
all
individuals
have
as
much
resource
as
they
need,
and
all
individuals
need
and
get
the
same
amount
(%
mortality
=
0).
Above
a
threshold
density
of
T
individuals,
in
scramble
competition,
all
the
individuals
still
get
an
equal
share,
but
this
is
now
less
than
what
they
require,
and
as
consequence
they
all
die
(as
is
found
in
dung
insects,
and
insect
forest
defoliators).
That
is
the
slope
of
the
ideal
slope
in
fig
2
(a)
suddenly
changes
from
0
to
infinite
as
the
threshold
T,
is
passed.
In
contest
competition,
on
the
other
hand,
individuals
fall
into
two
classes
when
the
threshold
is
exceeded.
T
individuals
still
get
an
equal
and
adequate
share
of
the
resource,
and
survive
(these
are
the
winners);
all
other
individuals
(the
losers)
get
no
resource
at
all,
and
therefore
die.
Thus,
in
Fig
2
(b)
the
ideal
slope
changes
at
threshold
from
0
to
1.
Whilst
there
can
be
no
survivors
in
scramble
competition,
there
are
always
just
T
survivors
in
contest
competition
irrespective
of
the
initial
density,
because
mortality
compensates
exactly
for
the
excess
number
of
individuals.
Scramble
and
contest
can
also
be
viewed
in
terms
of
fecundity.
Below
the
threshold
there
is
no
competition,
all
individuals
produce
the
maximum
number
of
offspring.
Above
the
threshold,
scramble
leads
to
production
of
offspring
whatsoever,
while
contest
leads
to
T
individuals
producing
the
maximum
number
of
offspring
and
the
rest
producing
none
at
all.
In conclusion we need to note that intra-‐specific competition results in:
(i) Evolution of efficiency in use of resources.
(ii) Broaden niche breath.
(iii) Regulation
of
size
through
contest
competition
and
the
exclusion
of
some
individuals
from
access
to
critical
resources.
Lecture
7:
INTERSPECIFIC
COMPETITION
Module:
Principles
of
Ecology
Dept.
of
Applied
Biosciences
and
Biotechnology,
Midlands
State
University
7.1. Inter-‐specific competition
7.2. Exploitative competition
7.3. Interference competition
7.4. Lotka – Volterra Model
7.5. Competitive Exclusion Principle
7.6. Coexistence
___________________________________________________________________________
INTER-‐SPECIFIC
COMPETITION
Inter-‐specific
competition
is
competition
between
individuals
of
different
species,
e.g.,
between
lions
and
hyenas,
cattle
and
donkeys,
etc.
The
essence
of
inter-‐specific
competition
is
that
individuals
of
one
species
suffer
a
reduction
in
fecundity,
survivorship
or
growth
as
a
result
of
resource
exploitation
or
interference
by
individuals
of
another
species.
Inter-‐specific completion occurs between similar species with similar ecological
requirements and usually results, if the populations are resource limited and emigration is
prevented, in the decline and ultimate extinction of one of the species.
If
competing
species
are
ecologically
identical
(i.e.
use
the
same
resources),
then
inter-‐
specific
competition
is
equivalent
to
intra-‐specific
competition.
Each
individual
competes
with
all
organisms
of
both
populations.
As
a
result,
the
population
growth
rate
of
each
species
is
determined
by
the
sum
of
numbers
of
both
populations:
dN1 = r1N1(K1 – (N1+N2))
dt K1
dN1 = r1N1(K1 – N1-‐N2)) equation 1
dt
K1
for species 1, and
dN2 = r2N2(K2 – (N2+N1))
dt K2
dN2 = r2N2(K2 – N2-‐N1)) equation 2
dt
K2
for
species
2.
The
species
with
a
higher
carrying
capacity
(K)
always
wins.
Higher
carrying
capacity
means
the
species
can
endure
more
crowding
than
the
other
species
(e.g.,
due
to
more
effective
search
for
resources).
If
the
competing
species
are
sufficiently
different
then
intra-‐specific
competition
is
stronger
than
inter-‐specific
competition.
Individuals
of
another
species
are
not
considered
as
“full”
competitors.
As
a
result,
the
number
of
inter-‐specific
competitors
is
multiplied
by
a
weight
αi<1:
dN1 = r1N1(K1 – N1-‐α12N2)) equation 3
dt
K1
dN2 = r2N2(K2 – N2-‐α21N1)) equation 4
dt
K2
Suppose
that,
together,
10
individuals
of
species
2
have
the
same
competitive
inhibitory
effect
on
species
1
as
does
a
single
species
1
individual.
The
total
competitive
inhibitory
effect
on
species
1
(both
intra-‐
and
inter-‐specific)
will
then
be
equivalent
to
(N1
+
N2/10))
species
1
individuals.
We
call
the
constant
–
1/10
in
this
case,
the
coefficient
of
competition
and
denote
it
by
α12
since
it
measures
the
competitive
inhibitory
effect
on
species
1
of
species
2.
In
other
words,
multiplying
N2
by
α12
converts
it
to
a
number
of
N1
–
equivalents.
Note
that
α12<1
means
that
species
2
has
less
inhibitory
effect
on
species
1
than
species
1
has
on
itself,
α12>1
means
that
species
2
has
a
greater
inhibitory
effect
on
species
1
than
species
1
has
on
itself.
α21 measures the competitive inhibitory effect on species 2 of species 1.
Species
coexistence
is
possible
if
intra-‐specific
competition
is
stronger
than
inter-‐specific
competition.
This
occurs
if
competing
species
have
different
preferences
in
resource
utilisation.
Inter-‐specific competition is conveniently divided into exploitative and interference
competition.
Exploitative
Competition
In
Exploitative
Competition
individuals
interact
with
each
other
indirectly,
responding
to
a
resource
level
which
has
been
depressed
by
the
activities
of
competitors.
When
inter-‐specific
competition
involves
resource
exploitation,
both
species
consume
the
resource
reducing
its
availability.
This
depletion
may
move
either
or
both
species
from
their
“saturated
range”
down
into
their
“limiting
range”
or
from
the
limiting
range
to
below
the
“threshold”
(where
the
individual
dies
or
the
population
goes
extinct).
Examples of exploitative competition include the following:
-‐ American Bison and domestic cattle compete for grass
-‐ Goats and sheep compete for grass
-‐ Tsessebe and Wildebeest compete for grass
-‐ Various fresh water algal species compete for inorganic nitrogen
Interference
Competition
In
Interference
Competition
interaction
between
individuals
is
direct.
An
individual
of
one
species
may
prevent
individuals
of
other
species
from
occupying
a
good
habitat,
from
using
resources,
or
may
actually
kill
individuals
of
the
other
species,
i.e.
the
behaviour
of
one
species
limits
others’
access
to
or
use
of
resources.
Thus interference competition:
-‐ may be aggressive (antagonistic) behaviour, e.g., scavengers at a Zebra kill on the
Serengeti plains in Tanzania or fierce fighting between lions and hyenas at a Zebra
kill in Savuti National Park,
-‐
may
be
“unintended”
–
e.g.
turbulence
generated
by
filter-‐feeding
water
fleas
(Cladocera:
like
Daphnia)
can
disrupt
filter-‐feeding
by
the
much
smaller
rotifers
(note
Cladocerans
may
also
kill
the
smallest
rotifers),
or
-‐
may
be
due
to
chemicals
(e.g.
toxins)
–
like
“allelopathic”
chemicals
produced
by
several
plant
species
that
inhibit
the
germination
of
other
(potentially
competing)
species.
Field
Experiments
We
can
ask
various
questions
about
competition
in
the
field:
-‐ If two species coexist naturally do they compete?
*
To
answer
such
a
question
you
can
perform
a
removal
experiment
–
you
can
remove
one
species
and
observe
the
effect
on
the
other
species
–
if
the
population
of
the
other
species
increases,
it
means
they
may
have
been
competitors.
-‐ If two species do not coexist, why not, is it competition?
*
You
can
perform
an
addition
experiment
and
determine
whether
the
addition
of
one
species
affects
the
density
of
the
other
–
does
competition
determine/limit
distribution.
Examples if interference competition from field experiments
1. Bedstraws – Tansley (1917)’s experiment with two species of bedstraws (Galium
spp) in England provides a clear example of both interference competition and the
competitive exclusion principle.
-‐
Bedstraws
of
the
genus
Galium
are
small
plants
which
can
grow
on
soil
of
any
pH.
Galium
hercynicum
(Gh),
however,
occurs
on
acidic
soils,
while
Gallium
pumilum
(Gp)
occurs
on
calcareous,
basic
soils
in
England,
but
when
alone
either
species
can
grow
on
soils
of
any
pH.
-‐ The fundamental niche for both Gh and Gp are soils of all pH’s.
-‐ When grown together, Gh wins on acidic soils and Gp wins on basic soils.
-‐
The
realized
niche
for
Gh
is
made
up
of
acidic
soils
only,
while
the
basic
soils
constitute
the
realised
niche
for
Gp.
-‐
Gh
drives
Gp
to
extinction
on
acidic
soils
while
Gp
drives
Gh
to
extinction
on
basic
soils.
This
shows
that,
which
species
is
the
winner
depends
on
the
habitat
and
that
unless
competing
species
partition
the
soil
pH
dimension,
one
will
be
driven
into
extinction.
-‐
The
mechanism
of
exclusion
in
this
example
involves
overgrowth
and
shading
(i.e.
shoot
competition).
2.
Barnacles
–
Connell
(1961)’s
work
with
barnacles
in
Scotland
also
provides
another
clear
example
of
both
interference
competition
and
the
competitive
exclusion
principle.
-‐
Barnacles
are
marine
crustaceans
found
in
the
rocky
intertidal
zone.
The
adult
barnacles
are
sessile
animals
attached
to
rocks.
When
covered
with
water
they
filter-‐
feed
on
suspended
particles
such
as
zooplankton
and
detritus.
-‐ The larvae, which are planktonic, disperse, then settle and metamorphose.
-‐
The
adults
of
two
species
of
barnacles,
Balanus
balanoides
and
Chthamalus
stellatus
co-‐occur
in
the
same
areas
in
the
field,
but
do
not
overlap
in
tidal
level
occupied.
-‐
Balanus
(B)
is
restricted
to
the
lower
levels
(near
the
low
tide
mark),
Chthamalus
(Ch)
is
restricted
to
higher
levels
(near
the
high
tide
mark).
Desiccation
is
frequent
at
the
high
tide
level
but
infrequent
at
the
low
tide
level.
-‐
The
larvae
of
both
species
settle
in
all
zones
but
the
subsequent
disappearance
of
Ch
from
the
B
zone
suggests
either
that
B
excludes
Ch
from
the
zone
of
their
potential
overlap
or
that
Ch
is
simply
unable
to
live
there.
-‐
Connell
carried
out
a
removal
experiment
in
order
to
distinguish
between
the
two
alternatives.
He
removed
B
from
some
plots
and
compared
the
results
with
those
from
the
controls
(unaltered).
Results:
Ch
survives
at
all
levels
where
B
is
removed
–
so
Ch’s
fundamental
niche
is
the
entire
intertidal
zone,
but
its
realized
niche
is
only
near
the
high
tide
line.
-‐
In
regions
where
B
eliminates
Ch,
it
does
so
by
interference
competition:
B
smoothers,
undercuts
or
crushes
Ch.
-‐ Connell concluded that Ch’s distribution was limited by competition.
In
the
opposite
experiment,
Connell
removed
Ch
from
some
plots
and
noted
the
effect
on
B.
Results:
removal
of
Ch
had
no
effect
on
b
whose
distribution
was
still
limited
to
the
lower
reaches.
-‐
Connell
concluded
that
the
distribution
of
B
is
restricted
by
a
physical
factor
(probably
inability
to
resist
desiccation),
not
by
competition.
-‐
Although
the
competition
between
B
and
Ch
is
markedly
one-‐sided,
these
two
barnacles
partition
the
niche
on
the
tidal
level
dimension.
Laboratory
Experiments
A
laboratory
is
a
controlled
environment
–
you
can
force
two
species
to
compete
for
one
resource
(e.g.
you
can
put
them
in
the
same
test
tube)
–
the
work
of
Gause.
Example of interference competition from laboratory experiments
G. F. Gause was a Russian ecologist who studied competition in laboratory experiments
using three species of Paramecium in 193 and1935.
Paramecium is a ciliated protozoan which can be maintained in test tubes feeding on
bacteria (and yeast) growing on oatmeal.
(i) P. aurelia (Pa) vs P. caudatum (Pc):
Result: Pa wins; Pc goes into extinction – complete exclusion
(ii) P. bursaria (Pb) vs P. caudatum (Pc):
Result:
with
these
two
species
we
get
stable
coexistence
(no
competitive
exclusion),
But
careful
examination
of
the
tubes
indicates
that
the
two
species
do
not
overlap
in
space:
Pc
tended
to
live
and
feed
on
the
bacteria
suspended
in
the
medium,
whilst
Pb
was
concentrated
on
the
yeast
cells
at
the
bottom
of
the
tubes.
Pb
has
symbiotic
green
algae
and
algal
photosynthesis
provides
02
to
the
paramecium
even
in
the
hypoxic
zone
of
the
test
tubes.
So
here
the
species
partition
the
niche
even
though
they
are
confined
to
a
single
test
tube.
The Lotka -‐ Volterra Model
While the logistic model is a population growth model with takes into account intra-‐specific
competition, the Lotka – Volterra model is a competition model including the effect of both
intra-‐specific and inter-‐specific competition on population dynamics.
Remember:
-‐ differential equations:
dN/dt = slope of live individuals at a particular instant in time
-‐ exponential equation:
dN/dt =rN – no intraspecific competition (unlimited environment)
-‐ logistic equation:
dN/dt = rN[(K -‐ N)/K] – includes intraspecific competition.
As N approaches K, dN/dt declines
If N>K, dN/dt becomes negative.
Now the Lotka – Volterra Model is like the logistic model, but it now includes the density of
the other species.
Considering
the
population
dynamics
of
species
1
in
the
presence
of
species
2,
the
logistic
equation
becomes:
dN/dt1 = r1N1[(K1 – N1 – α12N2)/K1]
so in the final term (previously (K1 – N1)/N1 we now add N2 multiplied by α12 to N1.
Where:
α12
is
the
competition
coefficient
and
measures
the
competitive
effect
of
species
2
on
species
1
When
α12
has
a
large
value
(i.e.
approaches
1.0)
it
means
species
2
is
a
strong
competitor
and
exerts
a
large
inhibitor
effect
on
species
1.
α12
defines
the
inhibitory
effect
of
species
2
on
species
1
in
terms
of
equivalent
numbers
of
species
1
individuals.
For example:
If
species
2
is
very
similar
to
species
1,
α12
approaches
1.0
such
that
an
individual
of
species
2
has
nearly
as
much
effect
on
dN/dt1
as
an
individual
of
species
1
would,
i.e.
there
is
near
complete
niche
overlap:
(K1 – N1 -‐ α12 N2)/K1 approaches (K1 – N1 –N2)/K1
If
species
2
is
quite
dissimilar
to
species
1,
there
is
only
slight
niche
overlap
-‐
α12
approaches
zero
(N2
has
little
effect
on
dN/dt1):
(K1 – N1 -‐ α12 N2)/K1 approaches (K1 – N1)/K1
Similarly, the population dynamics of species 2 can be described by the equation:
dN/dt2 = r2N2[(K2 – N2 – α21N1)/K2]
Note:
some
authors
use
α
(instead
of
α12)
for
the
effects
of
species
2
on
species
1
and
β
(instead
of
α21)
for
the
effects
of
species
1
on
species
2
or
even
W1
and
W2
for
α12
and
α21,
respectively.
Possible
outcomes
of
the
Lotka
-‐
Volterra
Model
We
can
employ
a
simple
graphical
model
to
predict
the
outcome
of
competition
between
two
species
if
we
know
the
initial
densities
of
each
(i.e.
N1
and
N2)
and
the
respective
competition
coefficients
(i.e.
α12
and
α21).
We
can
start
by
graphing
“zero
net
growth
isocline”
(ZNGI)
for
species
1
on
a
plot
of
N1
vs
N2.
This
line
shows
all
the
combinations
of
N1
and
N2
that
will
give
no
net
growth
(or
decline)
in
the
population
size/density
of
species
1
over
time,
i.e.
where:
dN/dt = 0
In order to draw the N1 isocline we will use the fact that on it dN/dt = 0
i.e. r1N1[(K1 – N1 – α12N2)/K1] = 0
This is true, less importantly, when r1 or N1 are zero, and more importantly when:
K1 – N1 – α12N2 = 0 or N1 = K1– α12N2
Since
our
isocline
is
a
straight
line
we
can
draw
it
by
finding
two
points
on
it
and
joining
them.
Along
the
x-‐axis
(i.e.
along
N1,
where
N2
=
0),
dN/dt
=
0
when
N1
=K1,
as
there
are
no
individuals
of
species
2.
(N2
=
0;
N1
=K1)
is
point
A
in
fig
1.
Along
the
y-‐axis
(i.e.
along
N2,
where
N1
=
0),
dN/dt
=
0
when
N2
=K1/α12
where
there
are
enough
individuals
of
species
2
to
completely
fill
up
all
the
spaces
for
species
1,
e.g.
if
K1
is
100
and
each
species
2
individual
fills
up
half
of
a
species
1
individual
space,
α12=0.5
and
so
you
need
K1/
α12
=
100/0.5
=
200
N2
individuals
to
reach
species
1’s
ZNGI.
(N1
=
0;
N2
=K1/α12
)
is
point
B
Fig.
1:
N1
isocline
Anywhere
“below”
(to
the
left
of)
the
ZNGI
there
is
“available
space”
for
species
1
(i.e.
it
is
below
its
carrying
capacity),
so
N1
will
increase
towards
the
line.
Anywhere
“above”
(to
the
right
of)
the
ZNGI
there
are
more
individuals
of
species
1
than
there
is
available
space
for
them
(i.e.
it
is
above
its
carrying
capacity),
so
N1
decreases
towards
the
line.
Fig.2: (a) N2 – isocline (b) N1-‐ and N2-‐ isoclines
Similarly,
we
can
graph
the
ZNGI
for
species
2
on
the
same
axis
(Fig
2(b))
Fig
2(b)
shows
that
species
1
drives
species
2
into
extinction.
When
both
isoclines
are
parallel,
the
species
with
a
higher
carrying
capacity
will
win.
Fig.3: Isoclines for N1-‐ and N2-‐ showing outcomes of competition
Fig. 3 (a) shows that:
K2 > K1/ α12 and K1 > K2/ α21
i.e. K1 < K2 α12 and K2 < K1α21
This
means
that
interspecific
effects
are
more
important
than
intraspecific
effects:
both
species
are
strong
interspecific
competitors.
There
are
two
stable
points
(N1
=
K1;
N2
=
0
and
N2
=
K2;
N1
=
0)
and
an
unstable
equilibrium
combination
of
N1
and
N2.
This
means
that
one
species
always
drives
the
other
into
extinction,
but
what
species
will
be
excluded
depends
on
initial
densities.
In
other
words,
if
α
>
1
and
isoclines
intersect,
one
species
will
exclude
the
other,
but
which
species
excludes
the
other
depends
on
initial
densities
of
both
populations.
Fig. 3 (b) shows that:
K1/ α12 > K2 and K2/ α21 > K1
i.e. K1 > K2 α12 and K2 > K1α21
This
means
that
intraspecific
effects
are
now
more
important
than
interspecific
effects:
both
species
are
weak
interspecific
competitors,
and
the
species
can
coexist
at
a
particular
equilibrium
combination
of
N1
and
N2.
Species
coexistence
is
possible
if
intraspecific
competition
is
stronger
interspecific
competition.
This
occurs
if
competing
species
have
different
preferences
in
resource
usage.
Using
the
Lotka
–
Volterra
model,
we
can
therefore
envision
4
possible
relationships
between
the
two
species
each
with
a
different
outcome:
1
Species
1
strong
competitor,
species
2
weak
competitor:
-‐ Species 1 wins, species 2 is excluded….no matter what the initial densities are.
2 Species 2 strong competitor, species 1 weak competitor:
-‐ Species 2 wins, species 1 is excluded….no matter what the initial densities are.
3 Both species strong competitors:
-‐
Which
species
wins
depends
on
initial
densities.
Coexistence
is
possible,
but
the
equilibrium
will
be
unstable
–
diverges
at
slightest
disturbance.
4 Both species weak competitors:
-‐ Result is stable coexistence, stable equilibrium.
-‐ Get same equilibrium densities, irrespective of initial densities.
Problems with the Lotka -‐ Volterra Model
-‐ Simplest possible model; biological unrealistic.
-‐ Linear effect of interspecific competition.
-‐ How to determine α values in the field or lab
-‐
Does
not
explicitly
define
resources
or
nature
of
interaction
(e.g.
exploitation
vs
interference)
–
underlying
explanation
may
be
very
complex.
Theory: The Competitive Exclusion Principle
-‐ Complete competitors cannot coexist at equilibrium in a stable environment.
-‐
If
two
competing
species
coexist
in
a
stable
environment,
and
there
are
no
differences
in
their
realized
niches,
then
one
will
be
competitively
excluded.
-‐ If they do coexist in the field, there must be some niche differentiation.
-‐ Can we disprove the theory?
-‐
If
we
find
coexistence,
we
look
for
niche
differentiation
–
if
we
do
not
find
any,
it
means
we
did
not
look
hard
enough.
Examples of competitive exclusion from laboratory experiments
1. Gause’s work with Paramecium
P. aurelia vs P. caudatum
P. aurelia excludes P. caudatum from the test tube.
2. Park (1954)’s work with flour beetles Tribolium confusum and T. castenium
In
a
series
of
simple,
sterilised
cultures,
Park
held
most
environmental
variables
constant
but
varied
the
climate.
In
all
conditions,
both
species
were
able
to
survive
in
monospecific
cultures:
the
fundamental
niches
of
both
species
spanned
the
whole
climatic
range.
In
mixed
cultures,
however,
T.
castenium
completely
excluded
T.
confusum
under
hot
–
moist
conditions
and
T.
confusum
excluded
T.
castenium
completely
under
cold-‐dry
conditions.
Examples of competitive exclusion in the field
1. Connell’s barnacles in Scotland
-‐ Balanus excluded Chthamalus from the zone of their potential overlap.
2. Bedstraws
-‐
Galium
hercynicum
excluded
Galium
pumilum
on
acidic
soils
while
G.
pumilum
excluded
G.
hercynicum
on
basic
soils.
-‐ Competitive exclusion, in this case, was very localised.
Coexistence
When
competitive
exclusion
does
not
occur
(i.e.
when
coexistence
occurs),
we
must
have:
1. Niche differentiation (i.e. competition not complete, niche overlap not complete).
2. Unstable environment (not at equilibrium).
3. Spatial patchiness (not at equilibrium)
We can now consider these in greater detail:
(a) Niche differentiation
Species can coexist when:
-‐ they are not complete competitors, and
-‐ when differences exist in their realized niches.
For example, Paramecium in the Lab exhibited niche differentiation with one species
restricted at the bottom of the test tube and the other at the top.
Field
example:
coexistence
of
three
species
of
birds
(tits)
in
English
broadleaved
woodlands
–
work
of
Lack
(1971):
Lack
concluded
that
the
three
species
of
birds
are
separated
from
each
other
at
most
times
of
the
year
by
their
feeding
station
and
the
size
of
their
insect
prey:
-‐ Blue tit (Parus caeruleus): feeds in oak trees on insects <2mm in length.
-‐ Marsh tit (P. palustris): feeds in shrubs on insects 3 -‐4mm in length.
-‐ Great tit (P. major): feeds on the ground on insects >6mm in length.
So, we have two-‐dimensional niche differentiation.
The
niche
differences
are
also
reflected
in
differences
in
body
size,
and
size
and
shapes
of
the
bills.
Other
examples
of
niche
differential
include
mixed
herds
of
grazers
that
graze
together
in
the
savannah
grasslands.
For
instance
mixed
herds
of
zebras
and
wildebeest
graze
together
on
the
vast
plains
of
the
Serengeti
but
do
not
compete.
Wildebeests
eat
tender
grass
shoots,
while
zebras
crop
tall,
tough
grass
that
the
wildebeest
cannot
bite
off.
Why do niche differences exist?
Niche differences exist either because of current competition, competition in the past or
some other reasons with nothing to do with competition.
(i) Current competition
The
species
can
restrict
each
other
to
smaller
realized
niches
through
completion.
Lack
did
not
do
field
experiments
to
show
competition.
He
could
have
removed
one
species
to
show
increased
growth,
survivorship
and/or
fecundity
of
the
other
species
–
or
show
expansion
of
niches
as
these
differences
may
reflect
choice
or
behavioural
limits.
(ii) Competition in the past
The species might have evolved niche differences to avoid competition.
It
could
be
that
natural
selection
favoured
those
individuals
with
less
overlap,
and
so
with
less
inter-‐specific
competition.
Individuals
of
both
species
in
the
area
of
overlap
could
have
been
negatively
impacted
resulting
in
lower
fitness.
If
niche
dimension
is
a
heritable
trait,
those
in
overlap
will
decrease
over
time.
Character displacement – species can evolve morphological adaptations which
allow them (largely if not totally) to avoid competition.
Niche
differentiation
due
to
natural
selection
could
have
arisen
in
order
to
avoid
competition.
In
Connell
(1980)’s
words
we
can
explain
niche
differences
in
the
present
by
invoking
the
“ghost
of
competition
past”.
(iii) Evolved niche differences for some other reasons
During
the
course
of
their
evolution,
species
might
have
adapted
to
their
in
environment
in
different
ways,
but
in
ways
with
nothing
to
do
with
interspecific
competition.
Its
quite
possible
that
the
species
did
not
compete
at
any
time
in
the
past,
and
do
not
compete
at
present.
We
cannot
distinguish
between
these
hypotheses
based
on
field
observations
alone.
We
need
to
carry
out
experiments
to
determine
whether
the
species
are
currently
competing.
It
is
impossible
to
demonstrate
competition
in
the
past
because
observed
niche
differences
may
be
consistent
with,
but
not
sufficient
to
demonstrate
current
or
past
competition.
(b) Unstable Environment
E.G. Hutchinson’s “Paradox of the Plankton”
When
the
principle
of
competitive
exclusion
became
widely
known
among
ecologists,
it
seemed
to
contradict
with
some
well-‐known
facts
and
this
contradiction
was
formulated
as
“paradoxes”
for
example
the
“paradox
of
the
plankton”
focused
on
the
variability
of
plankton
organisms
which
all
seemed
to
use
the
same
resources.
All
phytoplankton
species
use
the
same
essential,
limiting
resources:
light,
P,
N,
and
trace
elements.
There
are
not
so
many
mineral
components
dissolved
in
water
as
compared
to
the
large
variability
in
the
phytoplankton
species.
This
means
that
there
is
a
broad
overlap
in
niches.
In
some
systems
spatial
heterogeneity
(or
patchiness)
allows
species
to
coexist,
but
no
such
spatial
patchiness
(no
physical
niche
differentiation)
exists
in
the
case
of
the
lake
as
all
the
mineral
components
occur
throughout
the
mixed
zone
of
lakes.
Apparent
paradox
–
many
species
coexist,
but
how?,
with
no
competitive
exclusion.
The
answer
is
that
the
environment
changes
before
exclusion
can
take
place
(i.e.
the
environment
is
unstable).
As
the
environment
changes,
different
species
become
favoured
and
a
state
of
equilibrium
will
never
be
reached
–
rate
of
environmental
change
is
faster
than
the
rate
of
exclusion.
(c) Spatial patchiness
1. Ephemeral habitats – environment only exists temporarily – e.g. vernal ponds
(which dry up), rotting corpses, fruits and dung.
The
inferior
competitor
may
be
able
to
coexist
with
a
superior
competitor
by
being
a
better
colonizer
(which
finds
new
habitats
faster)
and/or
being
able
to
complete
its
life
cycle
before
the
habitat
vanishes
(early
age
at
first
reproduction;
matures
at
smaller
size,
etc.)
-‐ the system not at equilibrium
2.
Unpredictable
gaps
–
e.g.
light
gaps
in
tropical
rainforests.
Light
intensity
is
very
low
under
the
canopy
(due
to
interception
by
the
canopy)
–
seedlings
cannot
establish
under
a
canopy,
but
only
grow
to
adults
in
“light
gaps”.
Light
gaps
are
caused
by
tree
falls,
e.g.
with
high
winds,
the
tallest
canopy
trees
can
be
blown
down.
One
possible
strategy
then
is
to
be
a
“fugitive”
species
which
colonises
light
gaps
(or
newly
formed
ephemeral
ponds,
etc.)
first
and
the
produce
mobile
or
long-‐dispersed
seed.
E.g.
fast
growing
vines
can
coexist
with
superior
competitors
if
enough
light
gaps
are
formed.
In
light
gaps,
species
are
not
at
equilibrium
–
eventually,
the
gap
closes
in
and
dominant
canopy
trees
win.
An
inferior
competitor
can
“pre-‐empt”
space
(by
getting
an
earlier
start,
the
inferior
competitor
can
persist
longer
than
it
would
if
it
started
to
grow
at
the
same
time
as
the
superior
competitor).
General conclusions about competition
-‐ the interference/exploitation dichotomy in interspecific competition is somewhat
similar to that between scramble and contest (in intraspecific competition) in that
contest always involves interference, while scramble usually involves exploitation.
Interference and exploitation, however, have none of the extreme threshold
characteristics associated with scramble and contest competition.
-‐ competitive exclusion happens:
● winner often depends on environment.
-‐ coexistence is possible if:
● there is niche differentiation, and
● the system is not at equilibrium.
Lecture
8:
RELATIONS
AMONG
SPECIES
Module:
Principles
of
Ecology
Dept.
of
Applied
Biosciences
and
Biotechnology,
Midlands
State
University
8.1. Introduction
8.2. Commensalism
8.3. Mutualism
___________________________________________________________________________
RELATIONS
AMONG
SPECIES
Introduction
We
define
ecology
as
the
scientific
study
of
the
interactions
that
determine
the
abundance
and
distribution
of
organisms.
These interactions are with:
-‐ the abiotic environment (e.g. temperature, pressure, pH, pO2), and
-‐ the biotic environment (i.e. other organisms).
Theoretically,
populations
of
two
species
may
interact
in
basic
ways
that
correspond
to
combinations
of
0,
+
and
–
as
follows:
00, ++, +0, -‐0, and +-‐
Where:
+ stands for benefits
-‐stands for suffers, and
0 stands for unaffected.
The five major types of interactions are as follows:
COMPETITION -‐-‐ BOTH COMPETITORS SUFFER
PREDATION +-‐ PREDATOR BENEFITS, PREY SUFFERS
PARASITISM +-‐ PARASITE BENEFITS, HOST SUFFERS
COMMENSALISM +0 ONE BENEFITS, THE OTHER UNAFFECTED
MUTUALISM ++ BOTH MUTUALISTS BENEFIT
Three
of
these
combinations
(++,
-‐-‐
and
+-‐)
are
commonly
subdivided,
resulting
in
9
types
of
interactions.
These
are
as
follows:
(i) Neutralism
(00)
=
in
which
neither
population
is
affected
by
association
with
the
other.
(ii) Competition:
Direct
interference
type
(-‐-‐)
=
in
which
both
populations
actively
inhibit
each
other.
(iii) Competition:
Resource
use
type
(-‐-‐)
=
in
which
both
populations
suffer
when
a
common
resource
is
in
short
supply.
(iv)
Ammensalism
(-‐0)
=
in
which
one
population
is
inhibited
and
the
other
is
not
affected.
(iv) Predation
and
Parasitism
(+-‐)
=
in
which
one
population
adversely
affects
the
other
by
direct
attack
but
nevertheless
dependent.
(v) Commensalism
(+0)
=
in
which
the
commensal
benefits
while
the
host
is
not
affected.
(vi) Protocoorperation
(++)
=
in
which
both
populations
benefit
by
the
association
but
relations
are
not
obligatory.
(vii) Mutualism
(++)
=
in
which
the
interaction
is
favourable
to
both
populations
and
the
association
is
obligatory.
COMMENSALISM
Commensalism
is
an
interaction
in
which
one
organism
(or
species)
beneficially
affects
the
second
organism
(or
species),
but
the
second
has
no
effect
on
the
first.
Commensalism
often
involve
one
organism
that
is
already
dead,
which
is
then
consumed
by
another
(decomposition,
detritivory),
or
the
consumption
of
the
waste
products
of
one
species
by
another
(e.g.
coprophagy).
Decomposition
is
the
breakdown
of
complex
energy
rich
organic
molecules
to
simple
inorganic
constituents
–
also
called
“mineralisation”
and
is
an
important
step
in
material
recycling
(nutrient
regeneration).
Decomposers
are
typically
bacteria
and
fungi.
The
later
may
dominate
in
acidic
habitats,
while
the
former
may
dominate
in
more
basic
sites.
Bacteria
dominate
in
the
decomposition
of
the
more
easily
degradable
“parts”
(or
molecules)
of
dead
organisms
(e.g.
simple
carbohydrates,
lipids,
proteins
and
nucleic
acids);
while
fungi
dominate
in
the
decomposition
of
hard
to
degrade
molecules
(e.g.
celluloses,
hemicelluloses
and
lignins
of
woody
plants).
Forest
mushrooms
are
fruiting
bodies
of
the
fungi
that
breakdown
wood
and
leaves.
Detritivory
is
the
consumption
of
dead
organic
matter
(detritus)
usually
together
with
the
associated
decomposers
(bacteria
and
fungi).
Detritivory
can
also
include
carrion
feeders.
These
range
from
scavengers
(like
vultures)
most
of
which
can
also
act
as
predators
if
the
opportunity
arises,
to
carrion
feeding
insects
like
blowflies
in
which
the
adults
find
carcasses
by
smell,
lay
their
eggs
on
or
in
the
carcasses
after
which
the
maggots
(larvae)
consume
the
flesh,
pupate
and
emerge
as
adults.
Burying
beetles
of
the
genus
Necrophorus
are
also
carrion
feeders.
They
will
burry
small
mammal
carcasses
they
will
find
(to
reduce
competition
with
blowflies)
and
then
tear
off
and
consume
pieces
of
flesh
and/or
eat
any
blowfly
larvae
they
encounter.
Most
streams
and
some
lakes
derive
most
of
their
organic
matter
from
the
surrounding
watershed,
especially
dead
plant
parts
(mostly
leaves)
of
terrestrial
plants
that
fall
into
streams.
These
support
a
diverse
assemblage
of
aquatic
insect
larvae
and
other
invertebrates.
Coprophagy
While
the
guts
of
herbivores
are
longer
than
those
of
carnivores,
plant
matter
is
typically
more
resistant
to
digestion
than
are
animal
tissues
and
the
faeces
of
herbivorous
vertebrates
still
contain
significant
quantities
of
organic
matter.
Some
animals
have
specialised
on
feeding
on
the
faeces
of
other
animals,
e.g.
the
large
African
dung
beetle
(Heliocopris
dilloni)
feeds
on
the
dung
of
African
elephants.
The
winged
adults
of
the
beetle
smell
fresh
dung,
fly
to
the
site,
eat
some
of
the
dung
immediately,
and
cut
off
pieces
that
they
roll
away
and
burry
in
the
ground.
The
adult
female
lays
a
single
egg
in
each
of
the
dung
balls.
The
larva
then
hatches
out
and
consumes
the
dung
ball,
pupates
and
then
emerges
as
an
adult
to
look
for
fresh
dung
to
lay
its
eggs.
MUTUALISM
Mutualistic
relationships
are
mutually
beneficial
relationships,
in
which
individuals
of
both
species
have
higher
fitness
in
the
presence
of
the
other
species.
Fitness
is
the
relative
contribution
that
an
individual
makes
to
the
gene
pool
of
the
next
generation.
Fitness
is
very
difficult
to
determine
in
most
cases
–
rather,
we
take
short-‐cut
measures
to
estimate
fitness,
such
as:
-‐ growth,
-‐ survivorship, and
-‐ fecundity (reproductive output).
Mutualisms
generally
involve
one
species
gaining
access
to
resources
(food
or
inorganic
nutrients)
while
the
other
is
provided
with
protection
from
enemies,
a
favourable
environment,
or
a
service
(e.g.
pollination,
seed
dispersal,
rid
of
parasites,
etc.).
Coevolution of mutualistic relationships
-‐ Interacting species often show reciprocal changes through time.
-‐
Coevolution
–
each
species
must
change
in
response
to
the
changes
in
the
other
species.
-‐
Symbiosis
(=same
lives)
is
the
ultimate
expression
of
coevolution
–
two
species
live
as
one,
often
one
within
the
cells
or
tissues
of
the
other.
-‐
a
symbiosis
is
an
intimate,
long-‐lived
relationship
and
often
obligatory
association
of
two
species.
Mutualisms
can
be
divided
into
two
broad
categories:
Non-‐symbiotic
and
symbiotic
mutualisms.
Facultative vs Obligate Mutualisms
-‐ in facultative mutualisms, each species gains a benefit from the association but can
live on its own.
-‐
in
obligate
mutualisms,
on
the
other
hand,
each
species
can
only
live
in
the
presence
of
the
other.
Nonsymbiotic
Mutualisms
Protection
(or
Defensive)
Mutualisms
In
this
case,
one
species
is
defended
against
predators,
parasites
or
competitors
by
the
other,
which
may
in
turn,
receive
shelter,
etc.
Ant/Plant Protection Mutualisms
-‐
Seen
in
many
taxa
of
plants
but
especially
well
developed
in
the
South
American
Acacia
and
its
associated
ant
species.
-‐
Ants
protect
Acacia
trees
from
herbivorous
insects
by
actively
attacking
any
that
land
on
the
plant.
Ants
also
protect
the
Acacia
from
competitors
by
cutting
off
any
leaves
and
branches
of
nearby
plants.
-‐ In return, Acacia trees provide the ants with:
(i) nesting/brooding sites in the form of hollow thorns into which the queen
ant bores
(ii)
protein-‐rich
food
in
the
form
of
“beltian
bodies”
produced
at
the
tips
of
the
leaves,
and
(iii)
carbohydrate-‐rich
food
in
the
form
of
“nectar”
secreted
by
nectaries
at
the
leaf
bases.
-‐
South
American
Acacias
lack
more
traditional
plant
defences
such
as
spines
and
secondary
metabolites
to
deter
herbivores
and
so
they
depend
on
ants.
Farming of Fungi by Leafcutter Ants
-‐
Leafcutter
ants
defoliate
trees,
carry
the
leaf
pieces
back
to
underground
burrows
they
have
dug,
chew
the
leaves
and
hang
the
chewed
matter
from
the
roofs
of
their
galleries
where
they
become
infected
with
a
fungus.
The
fungus
“digests”
the
tough
leaves
and
is
harvested
for
food
for
larval
and
adult
ants.
-‐
The
fungus
gains
a
constant
supply
of
“food”
and
an
agent
for
dispersal.
When
the
ants
move
to
the
new
burrow,
the
queen
takes
a
supply
of
the
fungus
with
her
to
start
new
“gardens”.
The
fungus
is
never
found
in
the
absence
of
the
ants.
-‐
Harvester
termites
which
lack
gut
symbionts
found
in
other
termites
to
break
down
cellulose,
produce
subterranean
fungal
gardens
in
a
similar
manner.
Farming of Fungi by “Ambrosia” beetles
-‐ One group of beetles exhibits mutualisms with fungi similar to that of leafcutter ants,
however, here the food for the fungus is the wood of the dead and dying trees. The
beetles burrow into the wood forming a network of tunnels, which it infects with
fungi carried in its gut or attached to specialised “hairs”. The beetles visit the tunnels
periodically to harvest the ”ambrosia” for food.
-‐ Again the fungus gains a supply of food and a means of dispersal.
-‐ The larvae of wood wasps live similarly.
Farming
of
Aphids,
Leafhoppers
Scale
insects
and
other
bugs
(Homoptera)
by
Ants
-‐
Numerous
species
of
true
buds
(Homoptera),
such
as
aphids,
treehoppers,
leafhoppers,
and
scale
insects,
feed
on
phloem
sap
of
vascular
plants.
However,
since
they
must
pierce
the
plant
with
tubular
mouthparts
which
cannot
be
rapidly
withdrawn,
they
are
vulnerable
to
predation
by
other
insects
when
feeding.
-‐
Many
of
these
homopterans
have
mutualistic
relationships
with
ants.
The
ants
protect
the
homopterans
from
predators.
-‐
In
return,
the
ants
drink
the
“honeydew”
(dilute
sugary
faeces)
constantly
produced
by
the
bugs.
The
ants
often
stimulate
the
bugs
to
release
the
honeydew
by
stroking
them
with
their
antennae.
The
bugs
must
process
large
quantities
of
sap
so
as
to
ensure
that
they
obtain
enough
protein
and
are
able
to
leave
some
sugars
undigested
for
the
ants.
Farming of Caterpillars (Lepidoptera) by Ants
-‐
Caterpillars
of
the
butterfly
family
Lycaenidae
have
mutualistic
relationships
with
ants.
These
relationships
are
most
complex
in
the
blue
butterflies
where
the
relationships
are
between
single
species
pairs
of
butterflies
and
ants.
-‐
The
caterpillars
feed
on
plants
up
to
their
third
instar,
then
fall
to
the
ground.
The
caterpillars
mimic
ant
larvae
chemically
and
behaviourally
and
they
are
picked
up
by
ants
and
carried
back
to
their
nest.
-‐
The
ants
protect
the
caterpillars
from
predators
and
parasitoids,
as
well
as
feeding
them
with
plant
material.
-‐
In
return,
the
caterpillars
supply
the
ants
with
sugary
secretions
produced
by
the
“honey
glands”
when
they
are
stroked
by
the
ant’s
antennae.
The
caterpillar
secretions
may
also
contain
an
addictive
chemical.
-‐
The
relationship
between
the
butterfly,
ant,
and
the
plant
on
which
they
feed
may
be
so
restrictive
that
loss
of
any
one
species
may
cause
extinction
of
all
the
three,
e.g.,
loss
of
grasslands
in
England
has
dramatically
reduced
the
amount
of
wild
thyme,
the
plant
on
which
the
blue
butterfly
and
its
two
species
of
attendant
ants
depend.
The
butterfly
is
now
extinct
in
England.
Goby fish/Burrowing Shrimp Mutualism
-‐
Marine
shrimp
of
the
genus
Alpheus
have
mutualisms
with
goby
fish
of
the
genus
Cryptocentrus.
-‐ The shrimp digs burrows which the fish use for shelter.
-‐
The
shrimp,
which
is
blind,
keeps
one
antennae
constantly
in
contact
with
the
fish
when
outside
the
burrow.
This
provides
the
shrimp
with
early
warning
of
danger.
Mixed Species Aggregations
-‐ Mixed species aggregations such as the herds seen on the Serengeti Plains (with
Zebra, Wildebeest and Gazelle, for example), achieve increased vigilance against
predators as the different species combined will have better vision, hearing, smell,
etc.
Sea Anemone/Clown fish Mutualisms
Many
species
of
clown
fish
associate
with
sea
anemones.
The
clown
fish
gains
protection
by
retreating
into
the
sea
anemone’s
tentacles
when
threatened.
They
accumulate
slime
from
the
sea
anemone
to
prevent
triggering
the
nematocysts
themselves.
The
sea
anemones
also
gain
protection,
as
the
clown
fish
attacks
fish
species
that
normally
eat
sea
anemones.
Cleaning Mutualisms
-‐
A
number
of
coral
reef
fish
species
function
as
“cleaners”
of
other
often
larger
fish.
They
set
up
“cleaning
stations”
on
the
reef
to
which
they
attract
“customers”
with
their
body
posture.
-‐
The
customers
allow
the
cleaners
to
move
over
their
body
surface
and
even
into
their
mouth
and
gill
chambers
to
pick
out
and
eat
parasites
attached
to
them.
-‐ The cleaner gains a source of food while the customer is cleaned of parasites.
-‐
In
Africa
,
birds
known
as
the
oxpeckers
remove
ticks
and
other
parasites
from
large
mammals.
Recently,
oxpeckers
have
disappeared
from
certain
areas
as
they
have
been
poisoned
by
feeding
on
ticks
attached
to
cattle
that
were
dipped
to
kill
parasites.
The
wild
animals
have
suffered
tremendously
as
ticks
have
since
increased
dramatically.
Transport (or Dispersive) Mutualisms
Plants have developed mutualistic and parasitic relationships with animals for the
movement of their gametes (especially sperm in pollen) and propagules (especially seeds).
Pollination
Mutualisms
-‐
Wind
pollination
is
very
wasteful.
Most
of
the
pollen
never
reaches
another
flower.
Inducing
an
animal
(insect,
bird,
bat,
rodent,
etc.)
to
visit
flowers
by
providing
a
food
reward
in
the
form
of
nectar
(a
sugary
solution)
or
excess
pollen
(high
protein)
provides
a
more
efficient
way
of
delivering
pollen
from
one
flower
to
the
next.
-‐
Pollinators
and
flowers
may
be
generalists
(each
interacting
with
many
species
of
the
other)
or
specialists.
If
pollinators
and
flowers
are
specialists,
efficiency
in
pollen
delivery
is
even
greater
as
none
is
wasted
on
flowers
of
the
“wrong”
species.
However,
specialisation
which
can
even
lead
to
obligate
mutualisms
(e.g.
between
the
figs
and
the
fig
wasps),
makes
each
species
prone
to
extinction
should
the
other
be
wiped
out.
Seed Dispersal Mutualisms
-‐ Many plants produce fleshy fruits as an inducement for birds and/or mammals to
consume their seeds. The seeds are then carried for some time or distance before
being regurgitated or excreted, hence dispersing the seeds. The seeds, however,
must be digestion resistant.
-‐
Most
plants
have
their
seeds
consumed/dispersed
by
a
great
variety
of
species.
A
few
cases
of
specialisation
are
known,
however.
For
example,
some
Australian
plants
produce
seeds
with
edible
appendages
(called
elaiosomes)
which
are
eaten
only
by
ants.
Many
seeds
are
carried
some
distance
by
the
ants
before
elaiosomes
are
eaten.
A Miscellaneous Nonsymbiotic Mutualism
Honey Guide/Honey badger Mutualism
When
the
African
bird
known
as
the
honey
guide
(Indicator
indicator)
finds
a
bee’s
nest,
it
flies
around
looking
for
a
honey
badger
(also
known
as
the
ratel,
Melliovora
capensis),
or
a
human
being,
and
guides
it
back
to
the
nest.
The
powerful
mammal
rips
open
the
nest
and
eats
the
honey
and
larvae.
When
it
is
done,
the
bird
then
comes
in
and
eats
the
beeswax
and
the
remaining
larvae.
The
bird
has
gut
mutualists
capable
of
digesting
wax.
It
was
first
described
stealing
candles
from
mission
churches.
Symbiotic
Mutualisms
Digestive
Symbioses
(Gut
Mutualisms)
While
technically
outside
the
tissues
of
their
hosts,
gut
symbionts
or
mutualists
aid
the
digestion
of
complex
molecules
in
a
number
of
animals
and
synthesize
important
organic
molecules
like
vitamins
needed
by
their
host
using
the
host’s
food
as
raw
materials.
Ruminant
Mammals/Ciliates/Bacteria
-‐ Ruminants
(dear,
cattle,
antelope,
etc.)
which
feed
primarily
on
hard-‐to-‐digests
grasses,
have
four-‐chambered
stomachs.
The
rumen
contains
a
complex
community
of
cellulolytic
bacteria
and
protozoans,
most
of
which
are
specialists
that
cannot
live
elsewhere.
The
ruminants
use
the
fermentation
products
of
their
mutualists
as
food.
Termites/Flagellates/bacteria
-‐
While
75%
of
termite
species
produce
their
own
cellulose
to
digest
their
food
(wood),
and
some
species
cultivate
fungal
gardens,
other
species
possess
a
diverse
assemblage
of
mutualists
within
out-‐pockets
of
their
guts.
-‐
Anaerobic
flagellate
protozoans
ingest
wood
particles
and
ferment
them,
releasing
acetic
acid
and
other
organics
used
by
the
termite.
-‐
The
flagellates
are
covered
by
two
types
of
bacteria:
spirochaetes,
which
provide
locomotion
for
the
flagellate
in
exchange
for
“food”
(there
are
so
many
on
the
surface
of
the
protozoans
that
they
were
initially
classified
as
ciliated)
and
nitrogen-‐
fixing
bacteria
which
fix
gaseous
nitrogen
in
exchange
for
food.
General Gut flora
-‐ Bacteria in the guts of animals may synthesize vitamins and other molecules which
their hosts may need but cannot make on their own.
Photosynthetic
Symbioses
In
some
“marginal”
habitats,
inorganic
nutrients
needed
by
photoautotrophs
are
scarce
as
are
“prey”
for
heterotrophs.
Mutualisms
between
these
groups
provide
a
close
coupling
with
inorganic
nitrogen,
phosphorus,
etc.
passing
from
animals
(which
secrete
them
as
wastes)
to
“plants”
(where
they
are
nutrients)
and
some
photosynthate
passing
from
“plants”
to
animals.
These
mutualisms
typically
involve
algae
living
within
tissues
or
cells
of
animals.
Reef-‐building
Corals/”Zooxanthellae”
-‐
Corals
(coelenterate)
polyps
contain
symbiotic
algae
in
their
tissues
to
“harvest
sunlight”.
The
symbiotic
algae
of
most
cnidarians
are
non-‐motile
dinoflagellates,
called
Zooxanthellae.
-‐ The zooxanthellae also provide their hosts with excess photosynthate.
Sea Anemones& Jellyfish/Zooxathellae & Zoochlorellae”
-‐ A number of species of sea anemones and jellyfish (coelenterate or cnidarians)
contain symbiotic dinoflagellates (“Zooxanthella”) or green algae (“Zoochlorella”). In
cnidarians, these are contained in the gastrodermal cells and they pass on to their
host, excess photosynthate, in addition to harvesting sunlight.
“Green Hydra” (Hydra viridis) / Chlorella sp.
-‐ The freshwater, solitary coelenterate Hydra viridis contains symbiotic green algae of
the genus Chlorella within its cells. The algae can provide their hosts with oxygen as
well as photosynthate.
Paramecium bursaria/Chlorella sp.
-‐ The single-‐celled ciliate protozoan Paramecium bursaria contains numerous
intracellular symbiotic cells of green algae of the genus Chlorella. Not only do the
algae provide their hosts with photosynthate, they also supply them with oxygen in
hypoxic/anoxic environments.
“Lichens”: Fungi/Algae
-‐
About
25%
of
fungal
species
can
be
lichenized,
i.e.
infected
with
algae
to
form
lichens.
These
can
occupy
extremely
poor
habitats
such
as
bare
rock
and
tree
trunks.
-‐
The
algae
provide
photosynthate
(and
possibly
fixed
nitrogen)
to
the
fungus.
The
fungus
might
serve
the
alga
by
efficiently
taking
up
inorganic
nutrients
from
the
environment,
even
from
dilute
rainwater.
Nutrient Uptake Symbioses
“Lichens”: Fungi/Algae
Mycorrhizae: Plants/Fungi
The
roots
of
many
plants
form
intimate
relationships
with
various
fungi.
The
fungi
typically
penetrate
into
the
tissues
of
the
root
or
even
into
individual
cells.
Fungal
hyphae
extend
into
the
organic
upper
layers
of
the
soil
and
take
up
inorganic
nutrients
from
the
soil,
passing
them
on
to
the
plant.
In
exchange,
the
plant
passes
some
photosynthate
to
the
fungus.
Mycorrhizal fungi can extend between many plants, even trees of different species.
Chemosynthetic
Symbioses
Red
tube
worm
(Phylum
Pogonophora)/chemoautotrophic
Bacteria
-‐
Deep
sea
hydrothermal
vents
release
sulphides
which
support
unique
communities
of
pogonophorans
(red
tube
worms),
polychaete
worms,
crabs
and
large
molluscs.
-‐
The
red
tube
worms,
which
lack
a
mouth
and
a
digestive
tract,
have
a
mass
of
tissue,
called
the
trophosome
that
is
packed
with
symbiotic
bacteria
(chemosynthetic
bacteria).
The
bacteria
oxidize
sulphides,
producing
carbon-‐containing
compounds
which
they
share
with
their
pogonophoran
host.
-‐
The
vascular
haemoglobin,
which
accounts
for
the
red
colour
of
these
large
worms,
is
important
in
delivering
the
large
amounts
of
oxygen
required
by
the
bacteria.
-‐
The
worm
thus
appears
to
provide
a
safe
habitat
with
appropriate
conditions
for
the
bacteria.
-‐ Molluscs have their own chemosynthetic bacteria in their gills.
Nitrogen-‐fixation
Symbioses
In
many
habitats
photoautotrophs
are
limited
by
nitrogen
(or
phosphorus).
While
eukaryotes
lack
the
enzymes
to
‘fix”
the
abundant
atmospheric
nitrogen
(N2)
into
an
organic
form,
this
ability
is
widespread
in
the
prokaryotic
bacteria.
Mutualism
between
nitrogen-‐
fixing
bacteria
and
some
plants
ensures
that
the
later
has
an
adequate
supply
of
nitrogen.
Legume/Rhizobia
-‐
A
number
of
bacterial
genera
collectively
known
as
Rhizobia
form
complex
relationships
with
legumes.
-‐
The
presence
of
the
bacteria
triggers
the
root
cells
to
produce
root
nodules,
in
which
the
nitrogen-‐fixing
bacteria
will
be
contained.
The
nodules
are
kept
hypoxic,
as
the
nitrogenase
enzyme
only
functions
at
very
low
oxygen
concentrations.
-‐
“fixed”
nitrogen
(mainly
asparagine)
is
passed
on
to
the
plant.
In
return
the
plant
passes
photosynthate
on
to
the
bacteria.
Actinomycetes/non-‐legume symbioses
-‐ Actinomycets
of
the
genus
Frankia
form
nodules
on
the
roots
of
non-‐legumes
such
as
Alder
(Alnus),
which
is
an
important
early
colonist
in
some
habitats.
Cyanobacteria/Plant
Symbioses
-‐ A
number
of
plant
species
from
different
groups
form
symbioses
with
“blue-‐green
algae”
(prokaryotic
cyanobacteria),
some
of
which
can
fix
N2.
The
cyanobacteria
are
housed
in
special
structures
on
the
plant
thallus,
leaves
or
roots.
Cyanobacteria/Heterotrophic Bacteria Symbioses
-‐ Some free-‐living, filamentous cyanobacteria like Aphanizomenon contain special,
thick-‐walled cells called heterocyst which are the sites of nitrogen fixation. To ensure
low oxygen concentration in the heterocysts, they leak photosynthate which attracts
heterotrophic bacteria which consume most of the oxygen in the surrounding water.
Luminescence
Symbioses
Luminescence
Bacteria/Animals
-‐
A
large
number
of
marine
fish
and
invertebrate
species
are
capable
of
producing
light
(bioluminescence).
In
many
cases
the
fish
or
invertebrate
itself
produces
the
chemicals,
but
in
other
cases,
light
is
produced
by
luminescent
bacteria
housed
in
special
regions
of
the
animal.
-‐
The
light
produced
serves
the
anima
in
“counter-‐shading”,
finding
food
(e.g.
the
lantern
fish),
attracting
food
(e.g.
deep-‐sea
Anglerfish),
attracting
mates,
etc.
-‐ The animal provides the bacteria with appropriate conditions and food.
Serial Endosymbiosis Theory of Eukaryotic Evolution
-‐ In 1975, Lynn Margulis proposed that eukaryotic cells evolved by intracellular
symbiosis between/among prokaryotes.
-‐ Fermentative anaerobes could become infected by Krebs cycle-‐containing
eubacteria. The later becoming mitochondria.
-‐ Motile bacteria (such as spirochaetes) could infect the cells to become cilia and
flagella (like in the flagellate in the termite gut).
-‐ “ingestion” of photosynthetic cyanobacteria could give rise to chloroplasts, etc.
-‐
The
fact
that
both
mitochondria
and
chloroplasts
have
their
own
DNA
is
support
for
their
origin
as
intracellular
symbionts.
-‐
Given
the
success
of
eukaryotes,
these
symbiotic
mutualisms
(if
indeed
that
is
what
they
are)
can
be
viewed
as
the
most
important
of
all.
Lecture
9:
PARASITISM
Module:
Principles
of
Ecology
Dept.
of
Applied
Biosciences
and
Biotechnology,
Midlands
State
University
9.1. Introduction
9.2. Micro-‐parasites
vs
Macro-‐parasites
9.3. Endo-‐parasites
vs
Ecto-‐parasites
___________________________________________________________________________
PARASITISM
AND
INFECTIOUS
DISEASE
In
parasitism
and
infectious
disease,
one
organism
(the
parasite
or
pathogen)
benefits
at
the
expense
of
its
host.
Taken
together
parasitism
and
infectious
disease
have
devastating
effects
on
the
health
of
individuals
and
size
and
dynamics
of
most
(if
not
all)
wild
populations.
A
parasite
is
an
organism
that
consumes
part
of
the
host’s
tissues,
usually
without
killing
the
host
(at
least
immediately).
As
such,
parasitism
can
be
regarded
as
partial
predation.
The
term
disease
refers
to
an
unhealthy
condition
of
an
individual
that
impairs
a
vital
function.
An
infectious
disease
is
thus,
one
caused
by
an
infection
by
a
foreign
organism,
and
is
different
from
autoimmune,
metabolic
or
genetic
diseases.
A
disease
organism
is
an
organism,
usually
endoparasitic,
that
causes
a
disease
condition
in
another
organism.
Microparasites vs Macroparasites
-‐ Microparasites (including most disease organisms) are very small and typically can
complete their entire life cycle within a single host. They may rely on a
vector”organisms, however, to transport their propagules to infect new hosts.
-‐
One
example
of
a
microparasite
is
Plasmodium
spp.,
a
protozoan
which
causes
malaria
in
humans.
The
vector
organisms
of
malaria
are
the
females
of
a
number
of
species
of
mosquitoes.
-‐
Macroparasites
are
large
and
typically
live
on
the
surface
of
their
host
or
in
the
host’s
gut
cavity.
However,
the
blood
flukes
(Schistosoma
spp.),
which
are
classified
as
microparasites,
live
in
the
blood
vessels
of
their
hosts
and
cause
a
devastating
disease
in
humans
called
schistosomiasis.
–e.g.
the
Lamprey
and
leeches
on
fish
and
ticks
and
fleas
on
mammals.
-‐
Macroparasites
often
cannot
complete
their
entire
life
cycle
on
or
within
a
single
host.
They
often
produce
eggs
or
dispersive
larvae
and
frequently
involve
an
intermediate
host
in
their
life
cycle.
For
example,
schistosomes
infect
humans
as
adults
and
fresh
water
snails
(the
intermediate
host)
as
one
larval
form.
The
adults
(infecting
humans)
release
eggs
in
ponds
and
streams,
which
develop
into
ciliated
miracidium
larvae,
which
in
turn,
seek
out
and
penetrate
snails,
transforming
into
sporocysts
which
reproduce
asexually
to
produce
rediae
and
cercariae,
the
later
of
which
leave
the
snail
to
infect
humans
(either
actively
or
passively
by
encysting
and
being
eaten
by
the
host).
-‐
While
the
hosts
can
be
viewed
as
habitats
for
the
parasites
which
supply
them
with
controlled
conditions
and
a
constant
supply
of
resources,
the
host
possess
defence
mechanisms
in
the
form
of
resistant
surfaces,
immune
systems,
macrophages,
etc.
-‐
These
defences
co-‐evolve
with
“escape”
systems
in
the
parasite
in
a
continual
“arms
race”.
For
example,
immune
systems
of
hosts
produce
antibiotics
in
response
to
“foreign”
proteins
on
the
surface
of
the
parasites.
These
then
bind
to
the
surface
of
the
parasites
and
target
them
for
destruction
by
macrophages.
However,
parasites
can
escape
this
control
by
coating
themselves
with
host
proteins,
living
inside
cells
where
the
immune
system
cannot
reach
them,
or
constantly
changing
their
surface
proteins
so
that
the
host
cannot
‘keep
up”.
-‐
To
illustrate
the
normal
effect
host
defences
have
we
need
only
consider
the
cases
where
disease
and
parasites
have
reached
new
host
populations
that
evolved
in
their
absence.
The
diseases
may
devastate
these
‘naïve”
populations
(e.g.
European
humans
bringing
small
pox
and
other
diseases
to
the
native
Americans).
Endoparasites vs Ectoparasites
-‐ Endoparasites live within the tissues or cells of their host –e.g. most infectious
disease organisms including Plasmodium and Schistosoma.
-‐
Ectoparasites
live
on
the
surface
of
their
host
–e.g.
the
Lamprey
and
leeches
on
fish
and
ticks
and
fleas
on
mammals.
Lecture
10:
PREDATION
Module:
Principles
of
Ecology
Dept.
of
Applied
Biosciences
and
Biotechnology,
Midlands
State
University
10.1. Introduction
10.2. Predation
10.3. Types
of
predators
10.4. Predator/Prey
coevolution
10.5. Adaptations
of
predators
and
prey
___________________________________________________________________________
PREDATION
Introduction
Predation
and
parasitism
are
examples
of
antagonistic
interactions
between
two
species
which
result
in
negative
effects
on
the
growth
and
survival
of
one
of
the
populations.
Predators
use
their
prey
as
a
source
of
food
only,
whereas
parasites
use
their
hosts
both
as
a
food
source
and
a
habitat.
Importance of the study of predation and parasitism:
-‐
In
many
species
predation
and
parasitism
are
dominating
among
ecological
processes.
The
dynamics
of
the
se
populations
cannot
be
predicted
and
understood
without
considering
natural
enemies.
-‐
Pest
species
of
insects
and
weeds
can
be
suppressed
by
introduction
of
natural
enemies
(biological
control).
-‐
Natural
enemies
may
cause
side
effects
in
pesticide
applications.
The
numbers
of
arthropod
natural
enemies
may
be
reduced
due
to
pesticide
treatment
which
may
result
in
increasing
pest
populations.
Predation
Predation
is
the
consumption
of
one
organism
(the
prey),
in
whole
or
in
part,
by
another
organism
(the
predator),
in
which
the
prey
is
alive
when
the
predator
first
attacks
it
(i.e.
not
detritivory).
Predation
benefits
the
predator
(which
gains
a
resource
for
survival,
growth
and
reproduction)
but
is
costly
to
the
prey
(which
suffers
mortality
or
injury).
A
predator
will
require
more
than
one
prey
individual
in
order
to
grow
so
that
its
search
for
prey
is
continuous.
Types of Predators
a “functional” classification
-‐
True
Predators
–
are
animals
that
consume
other
animals.
They
kill
prey
more
or
less
immediately
after
attack;
eat
many
prey
individuals
per
life
time;
usually
consume
the
entire
prey
–
e.g.
lions,
cheetahs,
eagles,
snakes,
etc.
-‐
Herbivores
or
grazers
–
are
animals
that
eat
plants;
rarely
kill
prey;
ear
only
parts
of
many
prey
individuals
per
life
time,
called
browsers
if
they
eat
woody
parts
of
woody
plants,
e.g.
sheep,
cattle,
goats,
etc.
-‐
Parasites
–
are
organisms
(animals
or
plants)
that
live
in
an
obligatory,
close
association
with
a
host
individual;
rarely
killing
prey
(hosts)
outright;
eat
only
part
of
one
or
very
few
prey
individuals
per
life
time,
e.g.
tapeworms,
liver
flukes,
leaf-‐
mining
and
leaf-‐galling
insects,
various
bacterial
and
fungal
diseases.
-‐
Parasitoids
–
are
insects,
mostly
wasps
(Hymenoptera)
and
flies
(Diptera),
which
are
free-‐living
in
their
adult
stage,
but
lay
eggs
on
or
within
their
host.
The
larval
parasitoid
begins
eating
the
host,
initially
doing
little
harm,
but
eventually
almost
totally
consuming,
and
therefore
killing
the
host.
Parasitoids
eat
one
prey
individual
per
life
time.
a
taxonomic
classification
-‐
Carnivores
–
consume
animals
-‐ Herbivores – consume plants
-‐ Omnivores – consume both plants and animals
Predator/Prey
Coevolution
The
act
of
predation
affects
the
survival
and
ultimately
fecundity
of
both
predators
and
prey.
The
negative
effects,
however,
tend
to
be
quantitatively
small
where
the
interacting
populations
have
had
a
common
evolutionary
history
in
a
relatively
stable
ecosystem.
Natural
selection
tends
to
lead
to
a
reduction
in
detrimental
effects
or
to
the
elimination
of
the
interaction
altogether,
since
continued
severe
depression
of
the
prey
population
by
the
predator
can
only
lead
to
the
extinction
of
one
or
both
populations.
Consequently,
severe
interactions
are
most
frequently
observed
when
the
interaction
is
of
a
recent
origin
or
when
there
has
been
large-‐scale
or
sudden
changes
in
the
ecosystem.
Predators
and
their
prey
are
likely
to
have
co-‐evolved
and
there
is
a
continuous
selection
pressure
on
prey
to
avoid
death
at
the
hands
of
their
predators,
and
a
reciprocal,
continuous
selection
pressure
on
predators
to
increase
their
predation
efficiency.
Adaptations of Predators and Prey
Since the act of predation affects the survival and fecundity of both predators and prey (i.e.
affects their fitness), we see strong selection pressure for adaptations that increase:
-‐ the efficiency of predation in predator populations, and
-‐ the changes of escape or evasion in the prey populations.
Predation
can
be
viewed
as
a
multi-‐step
(or
multi-‐stage)
process
and
the
adaptations
to
predation
of
predators
and
prey
are
stage-‐specific
(co-‐evolution):
STAGE
PREDATOR
PREY
COUNTER-‐
ADAPTATIONS
ADAPTATIONS
Encounter
Encounter
behaviours
Rarity;
apparent
rarity
(ambush
or
search);
search
(hiding
and
refuge
use,
or
set
up
an
ambush
in
a
being
active
when
limited
area
where
prey
is
predators
are
not);
abundant
Movement,
e.g.
seasonal
migration
away
from
centres
of
predator
density);
Aggregation
Detection
Sensory
acuity;
search
image
Crypsis;
Immobility;
Polymorphism;
detect
predator
first
and
move
away;
Dispersion
Recognition/Selection
Image
of
“food:;
assess
Polymorphism;
profitability
(discrimination)
masquerading
as
an
learning
inedible
object;
Warning
colourations
(Aposematism);
Batesian
mimicry,
Mullerian
mimicry
Attack/Capture
Motor
skills
(speed,
agility);
Motor
skills,
escape
offensive
weapons
(see
behaviours
below)
(unpredictable
movements;
flee
to
cover;
startle,
bluff
or
threaten
predator);
Aggregation
(predator
saturation)
Handling/Subduing
Subduing
skills
and
offensive
Strength
to
escape;
weapons
(e.g.
teeth,
claws,
Active
defence
filter
sieves);
strength
to
(offensive
weapons);
detain;
killing
skills;
venom
Group
defence;
Spines;
Tough
integument;
Resistance
to
venom
Consumption
Tools
to
dismember
prey
Protection
for
safe
(teeth,
jaws
and
claws);
passage
through
the
detoxification
ability;
gut;
Toxins;
Spines;
digestive
tract
Emetics
Encounter
stage
-‐
In
order
to
encounter
prey
predators
may
lie
in
wait
or
move
around
in
search
of
prey.
In
other
words
Ambush
predators
may
set
up
in
a
localised
area
where
prey
is
abundant,
e.g.
at
water
holes
while
others
have
to
move
around
in
search
of
prey.
Which
mode
is
used,
however,
depends,
in
part,
on
the
movement
(or
lack
of
movement)
of
the
prey
(i.e.
it
makes
no
sense
for
a
herbivore
to
be
an
ambush
predator).
-‐
If
prey
are
rare
or
simply
“appear”
to
be
rare
(due
to
hiding
or
by
being
active
when
predators
are
not),
the
chances
of
an
encounter
are
reduced.
-‐
Therefore
prey
species
can
reduce
encounters
with
predators
through
refuge
use.
For
example:
-‐ Small organisms may hide under rocks or in holes.
-‐
Zooplankton
may
remain
in
deep
waters
during
the
day
to
avoid
detection
(at
least
when
predators
are
most
active).
-‐ Some herbivores may take cover in the dense bushes.
-‐
Some
species,
however,
avoid
cover
so
that
they
can
keep
a
good
lookout
for
predators,
but
others
need
it,
paradoxically,
for
the
purpose
of
hiding
from
predators.
Thus,
while
the
Thompson
Gazelles,
Wildebeest
and
Zebras
like
the
open
plains
and
may
well
be
less
vulnerable
there,
others
such
as
the
waterbucks,
reedbucks
and
duikers,
take
shelter
in
places
with
denser
vegetation.
-‐
Refuge
use,
however,
may
be
costly
to
the
prey
in
that
they
may
be
unable
to
search
for
their
own
food
when
in
hiding.
-‐
Prey
may
also
reduce
encounters
with
predators
by
being
active
when
predators
are
not,
e.g.
prey
species
can
adopt
nocturnal
feeding
habits
in
order
to
reduce
encounters
with
predators
which
hunt
during
the
day.
-‐
The
movement
of
the
prey
may
“be
designed”
(i.e.
shaped
by
natural
selection)
to
limit
encounters
with
predators,
e.g.
seasonal
migration
away
from
centres
of
predator
density.
In
some
cases,
herbivores
migrate
and
their
predators
do
not.
This
means
that
the
predators
only
live
in
plenty
when
the
game
passes
through.
Detection
stage
-‐
Predators
often
have
acute
senses
(sight,
smell,
hearing,
etc.)
to
help
them
detect
prey;
they
also
develop
“search
images”
(e.g.
symmetry,
contrast,
etc.)
to
help
them
detect
prey
“signals”
against
the
environmental
“background”.
-‐
Conversely,
prey
often
have
adaptations
to
reduce
their
chances
of
being
detected
by
predators.
These
include:
-‐
Cryptic
colouration
or
camouflage
–
this
is
a
passive
but
essential
defence
strategy
that
makes
potential
prey
difficult
to
spot
against
its
“background”.
A
camouflaged
animal
needs
only
remain
still
on
an
appropriate
substratum
to
avoid
detection.
Camouflage
may
also
include
transparency
and
counter-‐
shading
in
pelagic
species.
-‐
Immobility
(or
Akinesis)
makes
prey
less
detectable
by
predators
whose
vision
responds
best
to
motion.
-‐
Polymorphism
–
if
prey
are
polymorphic
(i.e.
the
population
or
species
consists
of
individuals
with
many
different
forms
or
colours)
this
can
confound
the
predator’s
search
image.
-‐
Sensory
acuity
–
if
prey
have
acute
senses
as
well,
they
may
detect
the
predators
first
and
move
away.
-‐
Interspecific
co-‐operation
(or
mixed
species
aggregations)
e.g.
Impala
and
Baboon
vs
leopard
and
the
mixed
species
aggregations
of
Zebras,
Wildebeests
and
Gazelles
(in
the
Serengeti)
ensures
increased
vigilance
against
predators
as
the
species
combined
will
have
better
vision,
hearing,
smell,
etc.
for
example,
the
Impala/baboon
combination
relies
on
the
impala’s
good
sense
of
smell
and
the
baboon’s
good
eye
sight
to
escape
from
the
leopard,
i.e.
they
are
able
to
detect
the
leopard
first
and
move
away.
-‐
Dispersion
–
if
the
predators
require
more
than
one
individual
for
detection,
dispersion
by
the
prey
may
be
advantageous.
-‐
Distraction
displays
-‐
distraction
displays
direct
the
attention
of
the
predator
away
from
a
vulnerable
prey,
such
as
a
bird
chick,
to
another
potential
prey
that
is
more
likely
to
escape
such
as
the
chick’s
parent.
Recognition (selection) stage
-‐ Typically predators do not “attack” all of the “potential” prey items they encounter.
Most are selective predators to some extent (some extremely so) and they asses the
profitability of the detected prey item before they attack – what constitutes “good
food” can be shaped by learning.
-‐
Again
polymorphism
(in
the
prey
population)
may
foil
the
predator’s
image
of
“good
food”
as
can
masquerading
as
an
inedible
object.
-‐
In
some
cases
prey
that
have
chemical
defences
that
affect
latter
stages
(e.g.
are
distasteful
or
toxic
and
hence
rejected
as
they
are
being
consumed)
advertise
this
fact,
often
by
being
brightly
coloured,
a
warning
to
predators
known
as
aposematic
colouration
(or
aposematism).
This
is
prevalent
in
several
groups,
e.g.
“poison
arrow”
frogs,
butterflies,
etc.
the
fire
Salamander,
which
can
squit
a
nerve
poison
from
glands
on
its
back,
is
brightly
coloured
in
black
and
yellow
to
deter
would-‐be
predators.
-‐
Mimicry
–
since
predators
learn
to
associate
warning
colourations
with
distastefulness/toxicity
and
the
refrain
from
attacking,
some
other
species
have
evolved
to
mimic
the
colourations
of
these
protected
species
and
hence
gain
protection.
-‐
Mimicry
is
a
phenomenon
in
which
the
mimic
bears
a
superficial
resemblance
to
another
species,
the
model.
Defensive
mimicry
in
prey
often
involves
aposematic
models.
-‐
In
Batesian
mimicry,
a
palatable
or
harmless
species
mimics
an
unpalatable
or
harmful
model.
The
larva
of
the
hawkmoth,
for
example,
puffs
up
its
head
and
thorax
when
disturbed,
looking
like
the
head
of
a
small
poisonous
snake
complete
with
eyes.
The
mimicry
even
involves
behaviour,
the
larva
weaves
its
head
back
and
forth
and
hisses
like
a
snake.
-‐
Additional
examples
of
Batesian
mimicry
are
the
many
harmless
snakes
that
mimic
the
conspicuous
red,
white
and
black
markings
of
the
poisonous
coral
snake.
-‐
For
Batesian
mimicry
to
be
effective,
however,
the
models
must
generally
outnumber
the
mimics,
otherwise
predators
would
learn
that
animals
with
a
particular
colouration
are
good
rather
than
bad
to
eat.
-‐
Batesian
mimicry
was
named
after
its
discoverer,
Henry
Bates,
a
19th
century
English
naturalist
and
explorer.
-‐
In
Mullerian
mimicry,
which
was
named
after
another
19th
century
explorer,
the
German
Fritz
Muller,
two
or
more
unpalatable
aposematically
coloured
potential
prey
species
resemble
each
other.
Each
acts
as
both
a
model
and
a
mimic.
Each
species
gains
an
additional
advantage,
because
the
pooling
of
numbers
causes
predators
to
learn
more
quickly
to
avoid
any
prey
with
a
particular
appearance.
-‐
Prey
may
employ
crypsis
and
akinesis
at
this
(and
the
following)
stage
too.
Once
detected,
the
prey
may
stop
moving,
for
example,
if
the
predator’s
vision
relies
on
motion
for
detection,
the
predator
may
lose
track
of
the
prey.
Attack (capture) stage
-‐ Once the predator encounters a prey item and deems it acceptable, it will “attack”.
This may range anywhere from a lion chasing a Gazelle to a squirrel picking up an
acorn.
-‐
Predators
may
exhibit
some
rapid
locomotion
and/or
offensive
weapons
(teeth,
claws,
filtering
sieves,
etc.)
to
help
them
approach
and
capture
prey.
-‐
Predators
may
also
use
mimicry
to
capture
prey.
For
example,
some
snapping
turtles
have
tongues
resembling
a
wriggling
worm,
thus
luring
small
fish,
any
fish
that
tries
to
eat
the
bait
is
itself
quickly
consumed
as
the
turtle’s
strong
jaws
snap
shut.
-‐
Motile
prey
may
attempt
to
escape.
Fleeing
is
perhaps
the
most
direct
antipredator
response,
although
it
can
be
energetically
very
expensive.
During
the
course
of
evolution
there
has
been
selection
for
fast
locomotion
and
some
prey
species
can
exhibit
evasive
behaviours,
e.g.
unpredictable
movements.
-‐ Aggregation by prey can “saturate” predators, i.e. diluting risk to prey.
-‐
Prey
may
attempt
to
startle,
bluff,
or
threaten
the
predator,
e.g.
impalas,
the
ink
in
the
Cephalopod
molluscs
can
confuse
predators.
Handling (subduing) stage
-‐ Predators show adaptations to increase ability to subdue (or even kill) prey, e.g.
physical strength, claws, pointed teeth, etc. lions, for instance, will use their physical
strength, claws and pointed teeth to subdue a Wildebeest and then go for the neck
to suffocate it.
-‐
Some
predators
(e.g.
many
snakes)
even
employ
venoms
(often
neurotoxins)
to
immobilise
prey.
-‐
Prey
may
use
their
strength
or
“defensive
weapons”
(e.g.
some
prey
can
attack
predators
using
horns,
spines,
gelatinous
sheaths,
etc.)
in
an
attempt
to
escape.
-‐
Prey
may
employ
group
defence
–
because
as
a
group
they
are
more
likely
to
be
able
to
fend
off
predators.
-‐
Some
prey
show
adaptations
to
make
them
hard
to
handle
(manipulate
in
preparation
for
consumption)
by
predators,
e.g.
spines,
gelatinous
sheaths,
etc.
-‐
Some
potential
prey
escape
simply
by
being
too
large
for
predators
to
handle
or
consume,
e.g.
whales
and
elephants.
This
is
called
“escape
in
size
from
size
selective
predation”.
Consumption
stage
-‐
Predators
may
have
to
dismember
prey
(mechanical
preparation,
i.e.
convert
it
into
smaller
pieces)
before
they
can
ingest
it
–
they
may
use
teeth,
jaws
and/or
claws.
-‐
Since
prey
may
be
toxic,
the
predator
may
have
to
be
able
to
detoxify
the
prey,
e.g.
some
pythons,
such
as
the
yellow
python,
can
eat
rattle
snakes
because
they
can
detoxify
rattle
snake
venom.
-‐
The
digestive
tract
of
the
predator
must
be
able
to
chemically
digest
(hydrolyse)
the
food
prior
to
absorption.
-‐
Even
when
captured,
prey
will
show
the
last
line
of
defence
against
being
eaten.
This
can
be
in
the
form
of
armour
or
spines
or
bad
taste
(including
toxics
or
emetics).
Predators
may
learn
from
bad
experience
with
one
individual
not
to
prey
on
similar
items
in
future.
-‐
But
how
does
this
help
the
dead
organism
(i.e.
how
can
such
a
trait
be
selected
for
if
the
organism
has
to
die
for
the
defence
of
other
individuals?)
–
by
kin
selection.
This
could
feedback
on
the
predator’s
selection
of
prey
to
attack.
Lecture
11:
PREDATION
(CONTINUED)
Module:
Principles
of
Ecology
Dept.
of
Applied
Biosciences
and
Biotechnology,
Midlands
State
University
11.1. Effects of Predation on Prey Populations
11.2. Regulation of Prey Populations by Predation
11.3. Effects of Predation on Predator Populations
11.4. Numerical Response
11.5. Functional Response
___________________________________________________________________________
EFFECTS
OF
PREDATION
ON
PREY
POPULATIONS
We
can
ask
two
questions
on
the
effects
of
predation
on
prey
populations:
(i) can predators regulate prey populations?
(ii) Why don’t predators typically drive prey populations to extinction?
Regulation of Prey Populations by Predation
A number of field studies have indicated that predator populations can regulate (limit) prey
species abundance:
Huffaker and Kenett (1956):
-‐ Their work involved two species of mites found in association with strawberry plants.
The Cyclamen mite Tasonemus pallidus is herbivorous on strawberries. The other
species Typhlodramus occidentalis is carnivorous and predatory upon the Tasonemus
pallidus.
-‐
In
the
presence
of
its
predator
(Typhlodramus),
the
cyclamen
mite
maintained
low
population
levels,
but
in
the
absence
of
its
predator
there
was
a
25-‐fold
increase
in
its
population
size
(treatment
with
a
chemical
insecticide
killed
the
“predator”
but
not
the
“prey”).
-‐
This
was
a
removal
experiment
and
it
illustrated
that
predators
normally
keep
prey
densities
low.
Cactus
“prickly
pear”
(Opuntia
sp.)
and
the
cactus
moth
(Cactoblastis
cactorum)
-‐
Caterpillars
of
the
cactus
moth
(Cactoblastis
cactorum
prey
upon
the
cactus
(and
also
carry
pathogens
and
rot
causing
organisms).
-‐
The
cactus
was
introduced
in
Australia
from
South
America.
It
spread
rapidly
and
dramatically
increased
in
abundance,
i.e.
the
cactus
expanded
its
distribution
and
abundance
in
the
absence
of
its
natural
enemy
(or
predator).
-‐
The
introduction
of
the
cactus’
natural
enemy,
the
cactus
moth
(from
South
America)
to
control
the
cactus
saw
a
dramatic
decline
in
abundance
and
distribution
and
restrictions
in
the
range
of
the
cactus.
-‐
This
an
example
of
an
addition
experiment
which
shows
that
predators
can
regulate
or
limit
prey
populations.
-‐
The
cactus
now
persists
as
a
“fugitive”
species
as
it
can
disperse
more
quickly
than
the
moths
(some
seeds
reach
areas
where
the
moth
is
not
established,
and
can
grow
and
reproduce
before
the
moth
catches
up
with
them).
Why don’t predators typically drive prey populations to extinction?
As a prey species becomes rarer, it becomes harder for predators to find individuals to
consume. The predator will, therefore, spend a longer time searching for that prey type.
This means that the “costs” of predation will go up and the “returns” will go down. Some
predators will then switch their search image to a more abundant (and hence more
profitable) prey type. This prey switching by the predator reduces the predation pressure on
the rare prey species. The rare prey species is thus, not driven into extinction.
On
the
effect
of
predation
on
prey
populations,
we
need
to
note:
Prey
populations
reduced
by
their
predators
will
experience
a
compensatory
decline
in
the
depressant
effects
of
intraspecific
competition,
while
those
that
grow
large
through
the
rarity
of
predators
will
suffer
the
consequences
of
intraspecific
competition
all
the
more
intensely.
Effects of Predation on Predator Populations
Since prey are an obvious resource for their predators, we can ask:
-‐ are predators limited by prey density?
Or
-‐ how do predator populations respond to prey density?
Holling
(1959)
studied
the
predation
of
small
mammals
on
pine
sawflies
and
he
found
that
predation
rates
increased
with
increasing
prey
density.
This
resulted
from
two
effects:
(1)
each
predator
increased
its
consumption
rate
when
exposed
to
a
higher
prey
density,
and
(2)
predator
density
increased
with
increasing
prey
density.
Holling
considered
these
effects
as
two
kinds
of
responses
of
predator
population
to
prey
density,
with
the
increase
in
consumption
rate
being
a
functional
response
and
an
increase
in
predator
density
being
a
numerical
response.
Numerical
response
Numerical
response
means
that
predators
become
more
abundant
as
prey
density
increases.
This
may
result
from
two
different
mechanisms:
(i) increased
rate
of
predator
reproduction
when
prey
are
abundant
(numerical
response
per
se).
(ii) Attraction
of
predator
to
prey
aggregations
(“aggregational
response”).
The
reproduction
rate
of
predators
naturally
depends
on
their
predation
rate.
The
more
prey
consumed,
the
more
the
energy
the
predator
can
allocate
for
reproduction.
Mortality
rate
also
decreases
with
increased
prey
consumption.
The
most
simple
model
of
predators’
numerical
response
is
based
on
the
assumption
that
the
reproduction
rate
of
predators
is
proportional
to
the
number
of
prey
consumed.
This
is
like
the
conversion
of
prey
into
new
predators.
Lets
say,
for
example,
as
10
prey
are
consumed,
a
new
predator
is
born.
We can define the following:
-‐
Threshold
–
same
as
threshold
level
or
light
compensation
point
–
is
the
minimum
density
needed
to
sustain
a
predator
population.
-‐
Limited
region
–
increases
in
prey
density
over
this
range
cause
increases
in
predator
fitness.
-‐
Saturated
region
(“unlimited
region”)
–
at
prey
densities
above
the
“saturation
point”,
further
increases
in
prey
density
do
not
further
increase
predator
fitness.
-‐
Satiation
–
predators
can
only
consume
prey
at
a
limited
rate
(in
the
zone
“saturation”
above).
Satiation
leads
to
the
prey
strategy
of
explosive
reproduction,
in
which
the
prey
try
to
swamp
predators.
Food quality
Another
factor
complicating
the
relationship
between
predation
rate
and
predator
fitness
is
food
quality.
It
may
take
more
of
a
lower
quality
prey
to
support
the
same-‐sized
predator
population.
In
particular,
herbivores
are
greatly
affected
by
the
quality
of
their
food,
especially
nitrogen
content.
Sinclair
(1975)
monitored
the
protein
content
of
the
food
available
to
the
Wildebeest
in
Serengeti
during
1971.
He
also
monitored
the
fat
reserves
in
the
bone
marrow
of
the
live
males,
and
of
males
that
had
died
from
natural
causes
(these
reserves
being
the
last
to
be
utilised).
It
can
be
concluded
from
Sinclair’s
work
that
the
wildebeest
consumed
food
that
was
below
the
level
necessary
even
for
maintenance
(5
–
6%crude
protein)
in
the
dry
season
and
judging
by
the
depleted
fat
reserves
of
the
dead
Wildebeest,
poor
food
quality
was
an
important
cause
of
mortality.
McNeil
(1978)
also
showed
that
seasonal
peaks
in
the
densities
of
insects
feeding
on
the
grass
Holcus
mollis
are
related
to
peaks
in
food
quality
(measured
as
soluble
nitrogen)
in
the
leaves
and
stems.
Aggregational
Response
Aggregation
of
predators
to
prey
density
is
often
called
“aggregational
response”.
Aggregational
response
is
very
important
for
several
predator-‐prey
systems.
Predators
selected
for
biological
control
of
insect
pests
should
have
a
strong
aggregational
response,
otherwise
they
would
not
be
able
to
suppress
prey
populations.
Also
aggregational
response
increases
the
stability
of
spatially-‐distributed
predator
–
prey
systems.
Functional
Response
The
density
of
prey
is
of
crucial
importance
to
the
predator.
As
prey
density
increases,
in
general
the
consumer
eats
more
prey
(because
the
prey
becomes
easier
to
find,
etc.).
The
relationship
between
an
individual’s
consumption
rate
and
food
density
is
known
as
its
functional
response.
In
other
words,
functional
response
is
the
way
in
which
the
predation
rate
of
predators
is
influences
by
prey
availability.
Holling
(1959)’s
model
of
functional
response
illustrates
the
principle
of
time
budget
behavioural
ecology.
It
assumes
that
the
predator
spends
its
time
on
two
kinds
of
activities:
(i) Searching for prey
(ii) Prey handling (which includes chasing, killing, eating and digestion).
Consumption
of
the
predator
is
limited
in
Holling’s
model
because
even
if
prey
are
so
abundant
that
no
time
is
needed
for
search,
a
predator
still
needs
to
spend
time
on
prey
handling.
Holling (1959 identified three basic types of functional responses:
“Type 2” functional response
-‐ Most frequently observed and most typical
Fig.11.1 Type 2 functional response
-‐
In
this
type
of
functional
response
the
response
is
curvilinear.
Consumption
rate
initially
rises
with
increasing
prey
density
–
the
rate
of
increase
then
slows,
and
finally,
a
plateau
is
reached
at
maximum
ingestion
rate.
Holling
attributed
the
form
taken
by
the
curve
to
the
existence
of
the
predator’s
handling
time
(the
time
the
predator
spends
persuing,
subduing
and
consuming
each
prey
item
it
finds,
and
then
preparing
itself
for
further
search).
-‐ Examples: dragonfly nymphs eating waterfleas (Daphnia), slugs eating grass, etc.
Explanation:
-‐
A
predator
has
to
devote
a
certain
amount
of
handling
time
to
each
prey
item
it
consumes,
during
this
time
it
cannot
search
for
additional
items.
-‐
As
prey
density
increases,
finding
prey
becomes
increasingly
easy
(search
time
becomes
trivial),
but
handling
individual
prey
still
takes
the
same
amount
of
time,
and
handling
takes
up
an
increasing
proportion
of
the
predator’s
time.
-‐
Thus,
at
high
prey
densities,
the
predator
effectively
spends
all
its
time
handling
prey,
and
the
predation
rate
reaches
a
maximum
determined
by
the
maximum
number
of
handling
times
that
can
be
fitted
into
the
total
time
available.
Therefore,
even
when
prey
is
superabundant
and
search
time
is
nil,
predation
rate
is
limited
by
the
finite
handling
time
→plateau.
Implications of “type 2” functional response
-‐ The
plateau
represents
predator
saturation.
Prey
mortality
declines
with
prey
density.
Predators
of
this
type
cause
maximum
prey
mortality
at
low
prey
density.
For
example,
small
mammals
destroy
most
of
the
gypsy
moth
pupae
in
sparse
populations
of
the
gypsy
moth.
However,
in
high
density
defoliating
populations,
small
mammals
kill
a
negligible
proportion
of
the
pupae.
“Type 1” functional response
-‐ much less common
Fig.
11.2
Type
1
functional
response
In
this
type,
there
is
a
linear
increase
in
predation
rate
with
increasing
prey
density
until
the
maximum
predation
rate
is
reached,
then
abrupt
plateau
at
the
maximum.
The type 1 functional response is found in passive predators like spiders and filter feeders.
Examples:
-‐ Spiders – the number of flies caught in the net (web) is proportional to fly density.
-‐
Daphnia
magma
(waterflea)
filter-‐feeding
on
yeast
cells
(Saccharomyces
cerevisiae)
–
work
of
Rigler
(1961)
-‐
Rigler
(1961)
studied
the
feeding
rate
of
Daphnia
magma
with
the
yeast
Saccharomyces
cerevisiae
as
its
prey.
Daphnia
magma
is
a
filter-‐feeder
and
it
can
extract
yeast
cells
from
a
constant
volume
of
water
washed
over
its
filtering
apparatus.
-‐
At
low
prey
density
(below
105
yeast
cells
ml-‐1)
the
predation
rate
is
directly
proportional
to
food
concentration
because
the
Daphnia
can
filter
and
swallow
yeast
cells
simultaneously.
-‐
Above
105
yeast
cells
ml-‐1,
however,
Daphnia
are
unable
to
swallow
(i.e.
handle)
all
the
food
they
filter.
At
such
concentrations,
therefore,
they
ingest
food
at
a
maximal
rate,
limited
by
their
handling
time.
Fig.
11.3.
Functional
response
of
D.
magma
to
different
concentrations
of
yeast
cells
Explanation
-‐
Passive
predators
(e.g.
spiders
which
use
webs,
and
filter
feeders)
can
search
for
and
handle
prey
simultaneously.
-‐
Below
the
plateau,
search
time
(filtering
rate
in
this
case)
is
limiting.
Daphnia,
for
example,
filter
water
at
the
maximal
rate
–
ingestion
rate
increases
with
increasing
prey
density
(as
all
that
are
found
are
captured
and
consumed
rapidly).
-‐
When
the
plateau
is
reached,
ingestion
rate
is
at
its
maximum
and
“suddenly”
this
is
the
limiting
factor.
“Type 3” Functional Response
-‐ In this type, the response of predation rate to prey density is sigmoidal. The
predation rate initial increases exponentially with increasing prey density, but with
further increases in prey density, ingestion rate declines until it plateaus at a
maximum (i.e. slope increases, then decreases)(Fig 11.4).
Fig
11.4.
Type
3
functional
response
-‐ The
type
3
functional
response
occurs
in
predators
which
increase
their
search
activity.
In
other
words,
type
3
functional
response
occurs
whenever
attack
rake
increases,
or
handling
time
decreases,
with
increasing
prey
density.
Exapmles:
-‐ Blue bottle flies feeding on sugar droplets,
-‐ Shrews feeding on pine sawflies,
-‐ Ichneumonid wasps parasitizing their insect hosts.
Explanation:
-‐
Seen
whenever
an
increase
in
prey
density
increases
the
predator’s
searching
efficiency
and/or
decreases
its
handling
time.
Either
or
both
will
increase
ingestion
rate
“faster”
than
linear.
-‐
These
generally
involves
learning
on
the
part
of
the
predator-‐i.e.
with
practice
the
predator
will
become
better
able
to
find
and
handle
prey.
-‐
Also,
type
3
functional
responses
result
from
switching
preferences.
When
the
predator
switches
to
alternative
prey,
the
mortality
of
the
initial
prey
declines.
Polyphagus
vertebrate
predators
(e.g.
birds)
can
switch
to
the
most
abundant
prey
species
by
learning
o
recognise
it
visually.
Mortality
first
increases
with
increasing
prey
density,
and
then
declines.
Functional responses and the regulation of Prey densities
-‐ Can predators regulate the population sizes of their prey?
Or
-‐ Are prey population dynamics stable?
-‐
In
order
to
have
regulation,
the
mortality
of
the
prey
must
increase
with
increasing
density
N,
i.e.
only
positive
density
dependent
mortality
can
regulate
the
dynamics
of
prey
populations.
-‐
If
predator
density
is
constant
(e.g.
birds,
small
mammals),
then
they
can
prey
density
only
if
they
have
a
type
3
functional
response
because
this
is
the
only
type
of
functional
response
for
which
mortality
can
increase
with
increasing
prey
density.
-‐
However,
the
regulating
effects
of
predators
are
limited
to
the
initial
portion
of
the
type
3
functional
response
where
there
is
a
positive
slope
in
the
prey
mortality-‐vs-‐
prey
density
curve,
i.e.
the
interval
of
the
prey
density
where
mortality
increases.
Increases
in
prey
density
at
this
interval
lead
to
increases
in
predation
pressure
on
the
prey
population.
This
process
is
density
dependent
and
has
a
potentially
important
stabilising
effect
on
the
interaction.
If
prey
density
exceeds
the
upper
limit
of
this
interval,
then
mortality
due
to
predation
starts
declining,
and
predation
will
cause
a
positive
feedback.
As
a
result
the
number
of
prey
will
get
out
of
control.
They
will
increase
in
numbers
until
some
other
factors
(intraspecific
competition,
for
example)
will
stop
their
reproduction.
This
phenomenon
is
known
as
“escape
from
natural
enemies”
and
was
first
discovered
by
Takahashi.
-‐
Wildlife
managers
are
familiar
with
type
3
functional
responses:
e.g.
in
the
presence
of
wolves
,
whitetail
deer
densities
are
relatively
stable.
When
wolves
are
exterminated,
whitetail
population
densities
may
fluctuate
wildly.
This
can
lead
to
the
collapse
of
the
population
if
it
seriously
overgrazes
its
range.
-‐
Type
3
functional
responses,
which
are
capable
of
stabilizing
prey
population
densities,
have
traditionally
been
referred
to
as
“vertebrate
functional
responses”.
It
was
believed
that
because
they
involve
learning
only
vertebrates
would
exhibit
them.
However,
insects
can
also
learn
hence
blue
bottle
flies
and
ichneumonid
wasps
also
exhibit
type
3
functional
responses.
-‐
Type
2
functional
responses
have
traditionally
been
referred
to
as
“invertebrate
functional
responses”.
-‐
Hunting
can
be
viewed
as
another
“type
3
functional
response”
because
you
can
become
a
better
hunter
with
experience
–
more
efficient
at
locating
and
capturing
prey.
Lecture
12:
PREDATION
Module:
Principles
of
Ecology
Dept.
of
Applied
Biosciences
and
Biotechnology,
Midlands
State
University
12.1. Predator Behaviour
12.2. Diet Composition -‐ Preferences
12.3. Diet Width – The Optimum Foraging Theory
12.4. Foraging Behaviour and Predation Risk
___________________________________________________________________________
PREDATOR
BEHAVIOUR
-‐
Organisms
face
a
multitude
of
problems
through
their
lives,
but
dominated
by
three
things:
eating,
avoiding
being
eaten
and
reproducing.
Those
organisms
that
do
any
of
these
better
than
their
fellows
will,
on
average,
leave
more
progeny.
If
the
trait
which
makes
them
better
has
a
genetic
component,
it
will
become
more
common,
i.e.
it
will
be
“selected”
by
“natural
selection”.
So,
over
time
we
would
expect
that
organisms
become
as
good
as
possible
at
doing
the
jobs
they
have
evolved
to
do
(finding
food,
mates,
and
avoiding
predators).
-‐
Using
simple
arguments
based
on
some
basic
biological
assumptions
and
the
premise
that
natural
selection
happens
we
can
make
predictions
about
hoe
organisms
should
behave.
For
the
process
of
food
gathering
and
eating
(collectively
known
as
“foraging”)
we
can
predict
that
organisms
will
have
been
selected
to
maximise
the
profitability
of
their
feeding
(energy
per
unit
time)
whilst
minimising
the
risks
of
being
eaten.
-‐
The
field
of
behavioural
ecology
is
concerned
with
looking
at
the
behaviour
of
organisms
and
seeing
if
their
behaviour
makes
“sense”
in
the
light
of
evolutionary
biology.
By
looking
at
foraging
behaviour,
we
are
also
trying
to
understand
how
natural
selection
has
favoured
particular
patterns
of
behaviour
in
particular
circumstances.
-‐ Predator behaviour:
-‐ is a product of evolution (natural selection),
-‐ coevolves with prey defences
-‐ affects predator-‐prey population dynamics.
Diet Composition – Preferences
-‐ We can start by asking the question:
-‐ What determines the diet composition for a particular predator?
-‐
Predators
are
classified
as
either
monophagus
(feeding
on
a
single
prey
type),
oligophagus
(feeding
on
few
prey
types)
or
polyphagus
(feeding
on
many
prey
types).
-‐
An
equally
important
classification
is
between
specialists
(broadly,
monophagus
and
ologophagus)
and
generalists
(polyphagus).
-‐
It
should
be
noted
that
polyphagus
and
oligophagus
predators
are
not
indiscriminate
in
what
they
choose
from
their
acceptable
range.
Many
predators
are
selective
–
they
show
preferences
for
some
prey
items,
i.e.
they
eat
a
prey
out
of
proportion
of
its
abundance.
-‐Thus,
to
measure
food
preference
in
nature,
it
is
necessary
not
only
to
examine
the
animal’s
diet
(usually
by
analysis
of
gut
contents)
but
also
to
access
the
availability
of
different
food
items.
-‐
Examples:
1.
Bluegills
eating
Daphnia
of
different
sizes
(the
work
of
Werner):
PREY SIZE % IN THE ENVIRONMENT % IN THE GUT
1.4mm 25 10
1.9mm 25 20
2.5mm 25 31
3.6mm 25 39
This
is
an
example
of
a
RANKED
PREFERENCE
–
predators
prefer
prey
with
the
highest
energy
content
(but
these
also
require
greater
handling).
Prey
can
be
ranked
on
a
simple
scale:
Benefit/cost = Energy gain/handling time
Attacking, capturing, and ingesting prey cost time and energy.
2.
BALANCED
PREFERENCE
–
based
on
the
need
for
a
balanced
diet
of
complementary
prey
items:
-‐ Fitness higher with a mixed diet,
-‐
Complementary
resources,
or
avoid
eating
too
much
of
any
one
plant/animal
toxin.
3.
SWITCHING
PREFERENCE
–
predators
tend
to
prefer
the
most
common
prey.
If
the
most
common
prey
changes
(maybe
due
the
action
of
the
predator),
the
predator
switches
to
the
“new”
most
common
prey
type:
When
prey
is
abundant,
the
predator
“prefers”
it
and
consumes
it
in
greater
proportion
than
its
availability.
When
this
prey
becomes
scarce,
the
predator
switches
to
a
more
abundant
prey
item
and
consumes
it
in
lesser
proportion
than
its
availability.
Predator
switching
is
based
upon
a
learnt
ability
to
specialise.
Certain
predators
especially
vertebrates,
develop
specific
search
images
which
enable
them
to
search
more
effectively
(since
they
know
what
they
are
looking
for)
and
result
in
them
concentrating
on
their
“image”
prey
to
the
relative
exclusion
of
their
“non-‐image”
prey.
Predators exhibit RANKED PREFERENCES in discriminating between resource
types that are perfectly “substitutable” and exhibit BALANCED PREFERNCES
between resource types that are complementary.
Ranked
preferences
predominate
when
food
items,
can
be
classified
on
a
single
scale.
Many
predators
exhibit
a
combination
of
ranked
and
balanced
preferences.
They
select
food
that
is
generally
of
high
quality,
but
they
also
select
items
to
meet
specific
requirements.
Diet Width -‐ Optimal Foraging theory
-‐ Knowledge of the diet width among predators and of the ways in which predators
distribute their effort amongst prey items will give insights to the ways in which
predators tend to maximise their profits.
-‐
Natural
selection
has
shaped
foraging
behaviour
and
all
the
foraging
behaviours
tend
to
converge
on
the
optimum
(=the
best
possible)
–
“the
ghost
of
predation
past”.
-‐
The
optimal
foraging
theory
allows
us
to
predict
the
optimum
foraging
behaviour
for
a
predator
(e.g.,
actively
searching
for
prey,
or
ambush
predation).
In
other
words,
the
aim
of
the
optimal
foraging
theory
is
to
predict
the
foraging
strategy
to
be
expected
under
specified
conditions.
We
can
compare
data
from
the
field
to
the
theory’s
predictions
and
judge
whether
the
predator
is
truly
optimal.
Model of Diet Width
-‐ Once a prey is encountered, should it be eaten?
-‐ The model of diet width addresses which prey to include in the diet – preferences.
-‐ We rank prey items by profitability.
-‐
Then
ask
if
the
predator
should
expand
its
diet
by
including
the
next,
most
profitable
prey
item?
-‐
We
can
divide
the
time
spend
by
the
predator
(with
respect
to
foraging)
into
two
categories:
1.
Search
time
–
the
time
spent
trying
to
locate
prey
items
(S
=
mean
time
spent
searching
for
food).
2.
Handling
time
(hi
for
the
ith
item;
h
=mean
handling
time
for
all
items)
–
this
is
the
time
spent
between
the
location
and
the
final
consumption
for
all
prey
items,
and
includes:
(a) time spent pursuing the item until capture,
(b) time spent manipulating and consuming the item after capture.
-‐
If
a
predator
is
a
SPECIALIST,
it
will
pursue
and
consume
only
profitable
prey
items,
but
it
may
expend
a
great
deal
of
time
and
energy
searching
for
them.
-‐
If
a
predator
is
a
GENERALIST,
then
it
will
spend
relatively
little
time
searching
but
it
will
pursue
and
consume
both
profitable
and
unprofitable
prey
items.
-‐
An
OPTIMAL
FORAGER
“should
balance
the
pros
and
cons”
of
whether
or
not
to
include
a
given
item
so
as
to
maximize
its
overall
rate
of
energy
intake.
Calculating profitability:
For the ith species:
Profitability = Energy Content of Prey/handling time = Ei/hi
Profitability = E/h = Mean Energy Content/Mean handling time
If a predator eats prey type i its rate of energy gain is:
Ei/hi
If
a
predator
ignores
prey
type
i,
it
must
search
for
time
S
to
find
another
prey
item
during
which
its
rate
of
energy
gain
is:
E/(S + h)
We predict that the predator will include the item i in its diet only if:
Energy Gain if taken ≥ Energy Gain if ignored
or
Ei/hi ≥ E/(S + h)
So,
a
predator
should
add
item
i
to
its
diet
if
it
increases
its
average
rate
of
energy
gain
by
doing
so.
Predictions from the Optimal Foraging Theory
Some predictions of the Optimal Foraging Theory include:
1.
Predators
with
handling
times
that
are
typically
short
compared
to
their
search
times
should
be
generalists.
-‐ hi small → Ei/hi is large
-‐ while hi small, S is large, and Ei/hi remains unchanged
then,
for
many
prey
items,
Ei/hi
will
be
greater
than
E/(S
+
h),
so
they
will
be
included
in
the
diet,
e.g.,
birds
that
“glean”
stationary
insects
from
foliage
are
generalists.
2.
Predators
with
handling
times
that
are
long
compared
to
their
search
times
should
be
specialists.
-‐ S is small, so E/(S + h), is effectively = E/h
-‐
Therefore,
maximizing
E/(S
+
h),
is
effectively
equal
to
maximizing
E/h
which
is
done
simply
by
including
the
most
profitable
prey
items
(i.e.
those
with
the
largest
Ei/hi
)
in
the
diet,
lions
spend
much
time
pursuing,
capturing
and
consuming
their
prey.
As
a
result
they
select
the
old,
lame
and
weak
which
are
easier
to
catch
(hi
small
→
Ei/hi
large).
3.
In
unproductive
environments
(i.e.,
where
prey
are
scarce
and
hence
S
is
large),
predators
should
have
broader
diets
than
in
productive
environments
(where
S
is
small):
e.g.,
this
is
observed
in
both
bluegill
sunfish
and
the
Great
tit.
Within
each
species,
diet
width
depends
on
productivity.
4. Whether or not a predator should include an ith item in its diet should depend on:
(a) The profitability of that item (Ei/hi)
(b) The profitability of items already in the diet (E/h)
(c) The search times for items already in the diet.
…………………………….but it should not depend on:
the
search
time
of
the
ith
item
(Si)
because
it
is
being
encountered
during
the
“normal”
search
(S),
nor
therefore
should
it
depend
on
i’s
abundance
or
the
rate
at
which
the
predator
encounters
it.
………………………………therefore
(a)
Predators
should
specialise
when
profitable
prey
items
are
common
and/or
differences
in
profitability
are
great.
(b)
Predators
should
not
indiscriminate
when
profitable
prey
items
are
rare
and/or
differences
in
profitability
are
slight.
(c) Predators should ignore insufficiently profitable prey types
irrespective of their abundance.
What are the constraints on reaching optimality?
(i) Other
aspects
of
the
predator’s
behaviour
may
influence
fitness
more
than
foraging
behaviour,
such
as
avoidance
of
the
predator’s
predators.
(ii) For
many
predators
(especially
herbivores
and
omnivores),
energy
intake
may
not
be
the
crucial
factor.
Rather,
it
may
be
some
other
factor
(e.g.,
nitrogen
content)
o
obtaining
a
balanced
diet
by
consuming
a
mixture
of
prey.
(iii)
While
foraging
behaviour
evolved
under
one
set
of
circumstances
(e.g.,
to
limit
predator
or
to
supply
a
balanced
diet),
present
emphasis
may
be
on
energy
–
there
may
have
been
not
enough
time
to
evolve
the
optimal
strategy
(phylogenetic
constraint).
However,
if
energy
intake
rate
is
the
crucial
factor
and
other
aspects
like
predation
risk
are
taken
into
account,
the
optimal
foraging
theory
is
a
powerful
tool
–
lands
insights
into
the
foraging
decisions
a
predator
must
make.
Foraging Behaviour and Predation Risk
-‐ Organisms face a multitude of problems in their lives, chief among them eating,
reproducing and avoiding being eaten. Foraging strategies will not always be
strategies for simply maximizing feeding efficiency. Natural selection actually favours
foragers that maximize the profitability of their feeding whilst minimizing the risks of
being eaten.
-‐
The
modifying
influence
of
predators
on
foraging
behaviour
was
studied
by
Werner
et
al
(1983)
working
on
bluegill
sunfish.
-‐ Here we have three trophic levels:
Prey → our predator → our predator’s predator
Our predator is the forager and our predator’s predator is the predator
The forager must balance eating – vs – being eaten → conflicting demands
Example – Bluegills (work of Werner et al)
Predator = largemouth bass (fish)
Forager = bluegills
Prey = aquatic insects, zooplankton, etc
In
the
absence
of
the
predators,
the
bluegills
spent
much
of
their
time
for
aging
in
“open
water”
(few
weeds)
where
food
was
easy
to
spot
and
capture.
When
predators
(largemouth
bass)
were
present,
the
bluegills
shifted
to
increasingly
forage
in
more
heavily
vegetated
areas
where
there
is
more
cover
to
avoid
detection
by
the
bass
and/or
to
facilitate
escape.
In
the
absence
of
the
predators,
the
bluegills
move
amongst
the
available
sites
to
maximize
their
energy
intake
→
makes
optimal
foraging
predictions.
In
the
presence
of
predators,
the
bluegills
spent
much
more
time
in
prey
poor
“refuges”
–
energy
intake
much
less
(decreased
growth
and
reproduction),
but
overall
survival
is
increased.
When
we
looked
at
the
niche
concept
and
interspecific
competition
we
saw
that
realized
niches
can
be
highly
constrained
by
the
effects
of
competitors.
The
bluegills
example
also
illustrates
that
realized
niches
are
also
constrained
by
predators.
Lecture
13:
PREDATOR
BEHAVIOUR
Module:
Principles
of
Ecology
Dept.
of
Applied
Biosciences
and
Biotechnology,
Midlands
State
University
13.1. Lotka-‐Volterra Predator-‐Prey Model
13.2. Do Predator-‐Prey Cycles occur in nature?
13.3. Conclusions about Lotka-‐Volterra Predator-‐Prey Model
___________________________________________________________________________
THE
POPULATION
DYNAMICS
OF
PREDATION
We
can
look
at
the
effects
of
predation
on
the
population
dynamics
(i.e.,
changes
in
density
over
time)
of
both
the
predator
and
its
prey.
We
will
begin
by
considering
a
relatively
simple
model
based
on
differential
equations
considering
just
one
predator
and
one
prey
species.
Lotka-‐Volterra Predator-‐Prey Model
The Lotka-‐Volterra model is the simplest model of predator-‐prey interactions. The model
was developed independently by Lotka in 1925 and Volterra in 1926.
The model has two important variables:
C = number (or density) of the predator population
N = number (or density or biomass) of the prey or plant population
Here
we
have
two
simultaneous
differential
equations,
one
for
the
prey
and
the
other
one
for
the
predator.
1)Prey Population growth rate:
Prey Population growth rate = exponential birth rate – predator caused death rate
Assumption
#1:
in
the
absence
of
predators,
the
prey
population
increases
exponentially,
i.e.,
no
density-‐dependence:
dN/dt =rN
However,
in
the
presence
of
predators
prey
individuals
are
removed
(i.e.
killed)
by
predators.
The
rate
of
prey
removal
depends
on
the
rate
of
predator-‐prey
encounters
and
the
efficiency
of
the
predator.
Encounters
increase
as
(1)
the
number
of
predators
(C)
increase
and,
(2)
the
number
of
Prey
(N)
increases.
The predator’s efficiency can be designated a’ the “attack rate” of the predator.
a’ increases with (1) increasing searching efficiency and (2) increasing capture efficiency.
Assumption #2: a’ comes from a linear functional response.
The
rate
of
successful
encounters
(i.e.,
the
consumption
rate)
will
then
be:
a’CN
and the overall equation for the prey population dynamics becomes:
dN/dt = rN – a’CN equation 1.
2)Predator Population Growth Rate
Predator population growth rate = birth rate – background m ortality
In
the
absence
of
prey
(i.e.,
food),
predator
individuals
lose
weight
and
starve
to
death.
Therefore,
in
this
model,
predators
are
assumed
to
suffer
“background”
mortality.
Thus,
the
predator
population
declines
exponentially
through
starvation
in
the
absence
of
prey.
dC/dt = -‐ qC
where:
q = mortality rate
This loss of predators is counteracted by predator birth which depends on:
(i) the
rate
at
which
food
(prey)
is
consumed:
a’CN
(ii) the
predator’s
efficiency(f)
of
turning
this
food
into
predator
offspring.
There are two additional assumptions here:
Assumption #3: there is no mutual interference or cooperation between predators.
Assumption
#4:
the
conversion
of
prey
into
predator
offspring
has
a
linear
numerical
response
–
no
thresholds
or
saturation:
-‐ The predator birth rate then becomes:
fa’CN
and the overall equation for the predator population dynamics becomes:
dC/dt = fa’CN – qC equation 2
equations (1) and (2) above together constitute the Lotka-‐Volterra Predator-‐Prey Model.
To
begin
to
investigate
the
properties
of
this
model,
we
start
with
a
plot
of
prey
abundance
(x
–
axis)
vs
predator
abundance
(y
–
axis)
and
find
the
ZERO
NET
GROWTH
ISOCLINES
for
each
species
(similar
to
what
we
did
in
the
Lotka
–
Volterra
Model
of
Interspecific
competition).
ZNGI’s
are
lines
along
which
the
population
just
maintains
itself,
neither
increasing
nor
decreasing.
On
one
side
of
the
isocline,
the
population
will
increase,
on
the
other
sid,
it
will
decrease.
1) Prey Zero Net Growth Isocline – this is when:
dN/dt = 0, or dN/dt = rN –a’CN = 0
rearranging gives:
rN = a’CN
or
C =r/a’
Graphically:
2) Predator Zero Net Growth Isocline – this is when:
dC/dt = 0, or dC/dt = fa’CN – qC = 0
rearranging gives:
fa’CN = qC
or
N
=q/fa’
Graphically:
To
get
the
behaviour
of
both
species
(i.e.,
the
whole
system)
we
can
plot
both
isoclines
together
and
use
vector
addition
as
before.
We
end
up
with
a
number
of
cycles
=
coupled
oscillations:
These
coupled
oscillations
continue
indefinitely
(frictionless
pendulum)
i.e.
the
cycle
does
not
degrade
over
time.
For
a
single
predator
species
and
a
single
prey
species,
the
predator
tends
to
increase
in
abundance
when
there
are
large
numbers
of
prey.
However,
there
should
then
be
an
increase
in
predation
pressure
on
the
prey,
leading
to
a
decrease
in
prey
abundance.
This
will
ultimately
lead
to
food
shortage
for
predators,
a
decrease
in
predator
abundance,
a
concomitant
drop
in
predation
pressure,
an
increase
in
prey
abundance
and
so
on.
Thus
predators
and
their
prey
undergo
coupled
oscillations
in
abundance,
with
predator
numbers
“tracking”
those
of
the
prey.
“Neutral
stability”
–
populations
follow
precisely
the
same
cycles
indefinitely
unless
some
external
influence
(i.e.,
some
disturbance)
shifts
them
to
new
values,
after
which
they
follow
the
new
cycles
indefinitely,
e.g.
So,
if
a
given
cycle
is
disturbed,
there
is
no
tendency
to
return
to
the
original
cycle
–
(i.e.,
there
is
no
regulation).
The
Lotka
–
Volterra
predator
–
prey
model
is
useful
in
that
it
points
out
the
underlying
tendency
for
these
interactions
to
generate
cycles.
The
fact
that
we
do
not
generally
see
perfectly
regular
cycles
“in
nature”
could
then
be
explained
by
the
frequent
disturbances
which
set
up
new
cycles
(due
to
changes
in
climate,
food
availability
to
prey,
etc.).
The cycling results from a series of time lags:
(i) Between
“many
prey”
and
“many
predators”
–
time
needed
to
produce
new
predator
offspring
after
prey
abundance
increases.
(ii) Between
“many
predators”
and
“few
prey”
–
time
needed
for
predator
to
reduce
the
size
of
the
prey
population.
(iii) Between
“few
prey”
and
“few
predators”
–
time
needed
for
the
predator
to
starve
after
their
resource
has
declined.
(iv) Between
“few
predators”
and
“many
prey”
–
time
needed
for
the
prey
population
to
recover
following
reduction
in
predation
pressure.
Do Predator – Prey Cycles Occur in Nature?
Snowshoe hare (Lepas americanus) and the Lynx (Lynx canadensis)
The
snowshoe
hare
follows
a
roughly
10
year
cycle,
with
10
to
100
fold
changes
in
abundance.
The
cycle
is
indirectly
driven
bt
predation
by
the
lynx.
First
note
that
there
also
cycles
in
other
prey
species
taken
by
the
lynx,
such
as
a
bird
known
as
the
ruffed
grouse.
The
decline
in
hare
abundance
is
only
indirectly
due
to
predation
by
the
lynx,
but
more
directly
due
to
overgrazing
of
winter
food
(woody
browse).
When
the
hares
are
abundant,
they
overgraze
woody
browse,
leading
to
food
shortages,
low
birth
rates,
low
juvenile
survivorship,
high
weight
loss
and
low
growth
rates
in
the
hares
result.
In
addition,
when
the
woody
plants
are
heavily
overgrazed,
they
produce
high
levels
of
toxins
(takes
2
–
3
years
for
these
to
build
up).
This
results
in
a
decreased
quality
of
food.
These
two
factors
(low
food
quality
and
quantity)
together
lead
to
a
poor
“condition”
in
the
hares.
This
makes
them
more
susceptible
to
predation
by
predators.
So,
the
cycles
in
the
prey
population
(hares)
are
really
generated
by
a
time-‐lag
in
the
hare-‐
plant
interactions
and
have
little
to
do
with
the
lynx.
When
hare
numbers
decline,
woody
browse
can
increase
in
quantity
and
quality
–
woody
browse
is
released
from
predation.
Also,
the
cycles
in
the
predator
(lynx)
population
simply
track
the
cycles
in
their
prey
(the
hares),
they
do
not
generate
them:
When
the
hare
population
recovers,
the
predator
to
prey
ration
declines.
This
means
more
food
for
the
lynx,
higher
lynx
birth
rate
and
therefore
an
increase
in
the
abundance
of
the
lynx.
When
the
hare
population
declines,
the
predator
to
prey
ration
increases.
This
means
less
food
for
the
lynx,
lower
lynx
birth
rate
(and/or
death
rate
due
to
starvation),
and
therefore
a
decline
in
numbers
of
the
lynx.
But
additionally,
when
hares
become
scarce,
the
lynx
may
switch
to
the
ruffed
grouse
and
other
(alternate)
prey.
Thus,
the
lynx
can
drive
cycles
in
the
grouse
but
not
in
the
hare.
Conclusions about the Lotka -‐ Volterra Predator – Prey Model:
-‐ You have to be very careful when applying simplistic models to (often complex) field
situations.
-‐
Recall
that
the
Lotka
–
Volterra
Model
makes
some
very
basic
assumptions
that
are
rarely
expected
to
be
met,
e.g.:
(i)
Density
–
independent
population
growth
of
prey.
(ii) Simple linear functional response.
(iii) Linear numerical response.
-‐
Other
models
have
been
developed
(such
as
the
“Theta
logistic”)
which
allow
logistic
population
growth
of
prey,
type
1/type
2/type
3
functional
responses,
etc.
-‐
These
models
still
indicate
that
cycles
are
likely
under
most
conditions
(but
not
seen
if
there
is
type
3
functional
response).
-‐ But can get dumped oscillations, etc. as well.
Lecture
14:
POPULATION
REGULATION
Module:
Principles
of
Ecology
Dept.
of
Applied
Biosciences
and
Biotechnology,
Midlands
State
University
14.1 Density
dependence
and
Population
regulation
___________________________________________________________________________
POPULATION
REGULATION
Recall
that
the
size
(or
density)
of
a
population
can
vary
temporarily
due
to
four
demographic
processes:
1. Mortality – death of individuals (decreases population size).
2. Natality (or fecundity) – birth of individuals (increases population size).
3.
Emigration
–
movement
of
individuals
out
of
a
population
(by
migration,
dispersal,
etc.)
(decreases
population
size).
4. Immigration – movement of individuals into a population (by migration, dispersal,
etc.) (increases population size).
Begon
and
Mortimer
(1986)
define
regulation
as
the
ability
to
decrease
the
size
of
populations
which
are
above
a
particular
level
and
to
allow
an
increase
in
the
size
of
populations
which
are
below
a
particular
level.
This
particular
level
will
therefore
be
a
point
of
equilibrium.
Populations
below
it
increase,
those
above
it
decrease,
and
populations
actually
on
it
neither
increase
nor
decrease:
population
size
is
subject
to
negative
feedback.
In
the
case
of
the
effects
of
intraspecific
competition,
this
equilibrium
level
is
called
the
carrying
capacity
of
the
population.
In
other
words,
there
is
a
tendency
for
N
to
increase
when
it
is
low
and
to
decline
when
it
is
high.
This
results
in
logistic
growth.
Regulation
has
a
well-‐defined
meaning
and
by
definition
can
only
occur
as
a
result
of
a
density
dependent
process.
The
regulation
of
most
populations
results
from
density
dependence
of
mortality
and/or
fecundity.
There
are
three
possible
patterns:
(i) both birth and death rates are density-‐dependent.
(ii) only birth rate is density-‐dependent.
(iii) only death rate is density-‐dependent.
Examples of density-‐dependent mortality (i.e., mortality which increases as N increases):
-‐ Daphnia, in the Lab.
-‐ Daphnia, in the field.
-‐ Flour beetle, in the Lab.
-‐ Buffalo, in the field (Serengeti).
Examples of density-‐dependent fecundity (i.e., fecundity which increases as N increases):
-‐ Daphnia, in the Lab.
-‐ Annual plant, in the field.
-‐ Sparrow, in the field (Serengeti).
Unlike
regulation,
which
can
only
result
from
density
dependent
processes,
abundance
will
be
determined
by
the
combined
effects
of
all
the
factors
and
all
the
processes
that
impinge
on
a
population
(Begon
and
Mortimer
(1986).
In
conclusion,
it
should
be
noted
that
some
density-‐independent
factors
such
as
climate
can
drive
density
dependent
processes,
for
example,
Andrewartha
and
Davidson
felt
that
weather
acted
as
a
density-‐dependent
component
of
the
environment
during
winter,
by
killing
the
proportion
of
thrips
(Thrips
imaginis)
population
inhabiting
less
favourable
situations
(if
the
number
of
safe
sites
is
limited
and
remains
roughly
the
same
from
year
to
year,
then
the
number
of
individuals
outside
these
sites
killed
by
weather
will
increase
with
density.
POPULATION
REGULATION
Recall
that
the
size
(or
density)
of
a
population
can
vary
temporarily
Recall
that
the
size
(or
density)
of
a
population
can
vary
temporarily
due
to
four
demographic
processes:
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