3.1. The Leslie Matrix 49
Age-class
2
3
43
4
1
2
0
1
(i)
Age (x)
Figure 3.1. The relation between the continuous age variable x,usedinthelife
table functions m(x) and l(x), and the discrete age classes i, used in the projection
matrix parameters P
i
and F
i
.
nonhuman species. Humans too have many important attributes in addi-
tion to age. Attempts to fit these processes into the life table framework
have a strained and unsatisfactory feel, but matrix methods are equally
applicable to all of them (see Chapters 7 and 17, and AMD).
Matrix population models were introduced in the 1940s by Bernardelli
(1941), Lewis (1942), and especially Leslie (1945, 1948), whose name is
associated with them still. Leslie was an ecologist particularly interested in
populations of small mammals (Crowcroft 1991). In addition to his work
on matrix population models, he made the first life table calculation of the
intrinsic rate of increase for any nonhuman species and made significant
contributions to stochastic models and mark-recapture estimation. Matrix
models were largely neglected until the mid-1960s, when both ecologists
(Lefkovitch 1965) and human demographers (Lopez 1961, Keyfitz 1964,
Rogers 1966) rediscovered them.
3.1 The Leslie Matrix
We begin with a model for an age-classified population. We divide the
continuous variable age, which starts at 0, into a discrete set of age classes,
which start at 1. The scheme is shown in Figure 3.1. Age class i corresponds
to ages i − 1 ≤ x ≤ i. According to this convention, the first age class is
number 1. Some authors number the first age class as 0.
Our goal is to project the population from time t to time t+1. We assume
that the unit of time is the same as the age class width. We call this unit
the projection interval; its choice is one of the first steps in constructing a
matrix model. Not surprisingly, a model that projects from year to year will
differ from one that projects from month to month or decade to decade.
Suppose that the projection interval is one year, and that individuals
are classified into three age classes (0–1, 1–2, and 2–3 years). The state of
the population is described by a vector n(t), whose entries n
i
(t)givethe
numbers of individuals in each age class.
The individuals in age classes 2 and 3 at time t + 1 are the survivors of
the previous age classes at time t.Thatis,
n
2
(t +1)=P
1
n
1
(t)
n
3
(t +1)=P
2
n
2
(t),