Discrete time

mathematical
models in ecology
Andrew Whittle
University of Tennessee
Department of Mathematics
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Outline

Introduction - Why use discrete-time models?

Single species models

Geometric model, Hassell equation, Beverton-Holt, Ricker

Age structure models

Leslie matrices
Non-linear multi species models

Competition, Predator-Prey, Host-Parasitiod, SIR

Control and optimal control of discrete models

Application for single species harvesting problem

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Why use discrete

time models?

Discrete time
When are discrete time models appropriate ?

Populations with discrete non-overlapping

generations (many insects and plants)
Reproduce at specific time intervals or times
of the year
Populations censused at intervals (metered
models)
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Single species
models

Simple population
model
Consider a continuously breading population

Let Nt be the population level at census time t

Let d be the probability that an individual
dies between censuses
Let b be the average number of births per
individual between censuses

Then

Suppose at the initial time t = 0, N0 = 1 and = 2, then

We can solve the difference equation to give the

population level at time t, Nt in terms of the initial
population level, N0
Malthus population, when unchecked, increases in a
geometric ratio
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Geometric growth

Intraspecific
competition
No competition - Population grows unchecked
i.e. geometric growth
Contest competition - Capitalist competition
all individuals compete for resources, the ones
that get them survive, the others die!
Scramble competition - Socialist competition
individuals divide resources equally among
themselves, so all survive or all die!
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Hassell equation
The Hassell equation takes into account intraspecific
competition

Under-compensation (0<b<1)
Exact compensation (b=1)
Over-compensation (1<b)
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Population growth for

the Hassell equation

Special case:
Beverton-Holt model
Beverton-Holt stock recruitment model
(1957) is a special case of the Hassell
equation (b=1)

Used, originally, in fishery modeling

Cobweb diagrams
Steady State

Stability

Cobweb diagrams
Sterile insect
release
Adding an
Allee effect
Extinction is
now a stable
steady state
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Ricker growth
Another model arising from the fisheries
literature is the Ricker stock recruitment
model (1954, 1958)

This is an over-compensatory model which

can lead to complicated behavior
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Nt

a
Period doubling to chaos in the
Ricker growth model

richer behavior
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Age structured
models

Age structured models

A population may be divided up into separate discrete

age classes

At each time step a certain proportion of the population

may survive and enter the next age class

Individuals in the first age class originate by

reproduction from individuals from other age classes

Individuals in the last age class may survive and remain

in that age class

N1t

N2t+1

N3t+2
1

N4t+3

N5t+4

Leslie matrices

Leslie matrix (1945, 1948)

Leslie matrices are linear so the population level of

the species, as a whole, will either grow or decay

Often, not always, populations tend to a stable age

distribution
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Multi-species
models

Multi-species models
Single species models can be extended to multi-species

Competition: Two or more species compete against

each other for resources.

Predator-Prey: Where one population depends on

the other for survival (usually for food).

Host-Pathogen: Modeling a pathogen that is specific

to a particular host.

SIR (Compartment model): Modeling the number

of individuals in a particular class (or compartment).
For example, susceptibles, infecteds, removed.
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multi species models

Growth

Growth

Nn

Pn

die

die

Competition model
Discrete time version of the Lokta-Volterra
competition model is the Leslie-Gower model
(1958)

Used to model flour beetle species

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Predator-Prey models
Analogous discrete time predator-prey
model (with mass action term)

Displays similar cycles to the

continuous version
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Host-Pathogen models
An example of a host-pathogen model is the
Nicholson and Bailey model (extended)

Many forest insects often display cyclic populations

similar to the cycles displayed by these equations

SIR models
Susceptibles

Infectives

Removed

Often used to model with-in season

Extended to include other categories such as
Latent or Immune

Control in discrete
time models

Control methods
Controls that add/remove a portion of the
population

Cutting, harvesting, perscribed burns,

insectides etc

Adding control to our

models
Controls that change the population system

Introducing a new species for control, sterile

insect release etc

How do we decided what is

the best control strategy?
We could test lots of different scenarios and see
which is the best.
However, this may be teadius and time
consuming work.
Is there a better way?
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Optimal control
theory

Optimal control
We first add a control to the population
model
Restrict the control to the control set
Form a objective function that we wish to
either minimize or maximize
The state equations (with control), control set
and the objective function form what is called
the bioeconomic model
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Example
We consider a population of a crop which has
economic importance
We assume that the population of the crop
grows with Beverton-Holt growth dynamics
There is a cost associated to harvesting the
crop
We wish to harvest the crop, maximizing
profit
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Single species control

State equations

Control set
Objective functional

how do
we find the
Pontryagins
discrete
maximum
best control
princple
strategy?

Method to find the

optimal control
We first form the following expression

By differentiating this expression, it will provide

us with a set of necessary conditions
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adjoint equations

Set
Then re-arranging the equation above gives the adjoint
equation

Controls

Set
Then re-arranging the equation above gives the adjoint
equation

Optimality system
Forward
in time
Backward
in time
Control
equation
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One step away!

Found conditions that the optimal control must
satisfy

For the last step, we try to solve using a

numerical method
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numerical method
Starting guess for control values
State equations
forward

Update
controls

Adjoint equations
backward
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Results

B large

B small
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Summary
Introduced discrete time population models
Single species models, age-structured models
Multi species models
Adding control to discrete time models
Forming an optimal control problem using a
bioeconomic model
Analyzed a model for crop harvesting
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