Population Models using Leslie Matrices

Goal: To build a discrete time age structured model for the female
population of organisms using only birth and survival rates. These
models were introduced by Patrick Leslie in 1945.
An individual in the study has a specific life span, say L years (this
could be days or some other unit of time depending on the individ-
ual). The life span [0, L] is divided into n equal intervals [0, L/n),
[L/n, 2L/n), [2L/n, 3L/n), , [(n 1)L/n, L]. We use age i to
refer to the interval [iL/n, (i + 1)L/n). For each age i interval where
0 i n 1, we let bi denote the average number of female
offspring from each female in age i and si+1 denote the percent of
females in age i that survive to age i + 1. It follows that the bis are
nonnegative numbers and sis are positive numbers.
In many species, reproduction is highly age dependent. An indi-
vidual has to reach a certain age before it begins reproducing. For
example, some plants this can be years and for a cicada it takes 13-
17 years.
Since only females produce offspring we need only follow the fe-
male population. The plan is to count the female population size in
age groups every L/n years (a generation) in order to study the total
female population.
Let t be the variable for the generation number and Ni(t) represent
the number of females in the population of age i at generation t.

1
Define
N0(t)
N (t)
1
N2(t)

N (t) =
N3(t)

..
.
Nn1(t)
as the vector valued function that describes the number of females in
each age class at time t.
What do we have to take into account for population change? For
this model we will consider birth rates and survival rates.
Example 1:
We divide the female population into 4 age groups - age 0, age 1, age
2, and age 3.
Survival rates: Assume that 40% of the females age 0 live to age 1,
30% of the females of age 1 live to age 2 and 10% of the females of
age 2 live to age 3. Assume no female lives beyond age three.
Birth rates:
Assume females age 0 have no offspring, age 1 females have 2 fe-
male offspring on average, age 2 females have 1.5 female offspring
on average and females age 3 have no offspring.
Census at generation t + 1.
N0(t + 1) = 0 N0(t) + 2N1(t) + 1.5N2(t) + 0 N3(t)
N1(t + 1) = .4N0(t)
N2(t + 1) = .3N1(t)
N3(t + 1) = .1N2(t)

2
Translate this to a matrix system:

N0(t + 1) 0 2 1.5 0 N0(t)
N1(t + 1) .4 0 0 0 N1(t)

N (t + 1) = =
N2(t + 1) 0 .3 0 0 N2(t)

N3(t + 1) 0 0 .1 0 N3(t)

0 2 1.5 0
.4 0 0 0

This matrix L = is called the Leslie Matrix.
0 .3 0 0
0 0 .1 0

1000
200

Suppose the initial population vector is N (0) = . Then
100
10

0 2 1.5 0 1000 550
.4 0 0 0 200 400

N (1) = =
0 .3 0 0 100 60

0 0 .1 0 10 10
What is the population distribution 4 years later?
N (4) = LN (3) = L(LN (2)) = L(L(LN (1))) = L(L(L(LN (0)))) =
L4N (0).

811 1000 702
248 200 269

N (4) = . 10 years later? N (10) = L10 = .
107 100 87
7 10 8

Example 2: Now, if given a Leslie matrix for a specific population

3
you should be able to interpret the matrix:
Given
5 7 1.5
L = .2 0 0
0 .4 0
What do we know?

1. The population is divided into three age classes: 0, 1 , 2.

2. The top row represents fecundity ratesbirth rates for each age
group. Thus each individual in the age 0 group has 5 female
offspring on the average, in the age 1 group has an average of
7 female offspring and in the age 2 group has an average of 1.5
female offspring.
3. The subdiagonal elements represent the survival rates for each
age group. 20% of the age 0 group survive to age 1, and 40% of
the age 1 group survive to age 2.

Example 3: Suppose we have an organism with an age structured

population whose Leslie Matrix is
 
1.5 2
L=
.08 0

We would like to find a growth pattern where the percentage of or-

ganisms in each age class remains the same over successive genera-
tions. (i.e. we want the ratio to remain the same) In other words we
want to find an age distribution vector x such that Lx = x. We can
solve this by finding the eigenvectors. We call this the stable range
distribution.

4
  
 1.5 2 
det(I A) =   = ( 1.5)() .16 = ( 1.6)( + .1)
.08 
1 = 1.6 and 2 = .1
Theorem: A Leslie matrix L has an unique positive eigenvalue 1
(called the dominant eigenvalue) and 1 has multiplicity one in the
characteristic equation of L. A corresponding eigenvector of 1 is of
the form
1

s1/1

2
(s1s2)/(1)

x1 =

(s1s2s3)/(1)3



...

(s1s2 sn1)/(1)n1
Then N (k) = Lk N (0) c(1)k x1. That is, for large k, N (k)
c(1)k x1.
1 is called the growth rate and x1 is called the stable age distribu-
tion.
 
1 20
Back to the example : rref (1.6I L) =
0 0
 
1
Thus eigenvectors corresponding to 1 = 1.6 are t where t 6= 0
.05
 
1
and x1 = .
.05
This tells us that a stable range distribution has a ratio of 20 to 1 for
the age 0 group to the age 1 group. What happens over the long term
if the population begins with this ratio?
Start with 2000 age 0 and 100 age 1 individuals, (The ratio is 20: 1.)
What happens in 5 years? 10 years? Check the ratio:
5
        
2000 20, 971 2000 219, 902
L5 = and L10 = .
100 1, 048 100 10, 995
However, by the above Theorem, for large k, N (k) c(1)k x1. That
is, for large k, N (k) is approximately a scalar (i.e., c(1)k ) times the
vector x1. That is, as k gets large, the population N (k) converges to a
scalar multiple of x1. The scalar c depends on the  initial
 population.
100
In this last example, lets start with N (0) = and compute
100
10
c.
 We will  use k = 10as a large value. N (10) = L N (0) =
23, 283 1
c(1.6)10 or 23, 282 = c(1.6)10 1. Thus, c 211.76.
1, 764 .05
The conclusion is that for large k,
 
1
N (k) 211.76(1.6)k .
.05

Check it out for k = 15:

     
244, 148 1 244, 143
N (15) = and 211.76(1.6)15 = .
12, 207 .05 12, 207
 
3 5
Practice Problem for Final: Let L = .
.8 0
(a) Find the eigenvalues for L.
(b) Find the growth rate 1.
(c) Find the stable age distribution vector x1.
 
100
(d) For N (0) = , find a scalar c such that for large k, N (k)
100
c (1)k x1.