AGE STRUCTURE AND SURVIVORSHIP

Populations, whether animal or plant, vary in their proportions of

young and old individuals. Time units such as weeks, months, or years
can describe ages. Or individuals can be assigned to qualitative age
classes such as nestling, juvenile, subadult, and adult, or egg, larva,
pupa, and adult. The proportions of individuals belonging to the
various age groups are collectively referred to as the age structure
or age distribution of the population.

Three different procedures may be used for obtaining the age

structure of a population.

The vertical approach follows a particular cohort. A cohort is a group

of individuals born within the same time interval. Thus, by knowing
the age of cohort members, you can follow their survival until all have
died.

The horizontal approach uses data on all ages within a given

population at one time; that is all cohorts in the population are
examined at the same time. In the latter method, one assumes a
stable age structure and constant birth and death rates.

A third approach involves knowing the age at death for members of a

population. Such data are commonly obtained for a game species.

Knowledge of age structure is important, for the age distribution of

a population affects its growth and dynamics. From a knowledge of
age structure, a table of age-specific mortality, survivorship, and life
expectancy can be constructed – a life table. In addition, population
growth rates may be estimated from data on births per female in the
population.

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 LIFE TABLE

To construct a life table we must be able to determine the age of

the organisms in questions and distribute the population members
into age classes or age intervals. Age intervals can vary according to
the longevity of the organism. For small rodents or lagomorphs the
age interval may be one month, for deer one year, for humans five
years. For insects age categories may be instars or life history
stages. We need information on survival, mortality, or rate of
mortality by age classes for a given population. Data on survivorship
in each age class provide the information needed for survivorship
column, lx.
Table 1. A Life Table. The data in the x and Lx columns were obtained from a
population of animals. Then, all other columns of data were derived from them,
as described in the text.

Age Cohort Number Number Number Probability Probability Animal- Live to

(yr) (age in Living Dying of Dying of Years Expectancy,
interval) Cohort, at Start, during Life Surviving Live, ex (yr)
x Lx lx x, dx during x, Interval x, Tx
qx sx
0-1 0 33 46 26 0.57 0.43 63 1.37
1-2 1 16 20 8 0.40 0.60 30 1.50
2-3 2 9 12 6 0.50 0.50 14 1.17
3-4 3 4 6 4 0.67 0.33 5 0.83
4-5 4 1 2 2 1.00 0.00 1 0.50
5-6 5 0 0 Σdx = 46

Life Tables
In a life table various statistics are compiled for each age class, or
cohort (designated x). Data are commonly collected as numbers of
individuals in each age class. Lx is the number of individuals in age
class x. It is assumed that Lx is the number alive at the middle of
age class x (for example, in above table, 33 individuals assumed to be
0.5 year old, even though the true ages of the 33 might range
between 0 and 1 year old).

We designate lx as the number of individuals alive at e beginning of

age class x. Thus, Lx may be defined as

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 Lx = (lx + lx+1) (1)
2
(i.e., Lx is the number alive at the midpoint of age class x), and

lx = 2Lx – lx+1 (2)

For example, in Table 1.

Lo = 33
L1 = 16
L2 = 9
L3 = 4
L4 = 1
L5 = 0

Since L5 = 0, we can set l5 = 0. Then by applying Equation 2,

l5 = 0
l4 = 2(1) – 0 = 2
l3 = 2(4) – 2 = 8 – 2 = 6
l2 = 2(9) – 6 = 18 – 6 = 12
l1 = 2(16) – 12 = 32 – 12 = 20
l0 = 2(33) – 20 = 66 – 20 = 46

The number of individuals in the population that die during interval x

is

dx = lx – lx+1 (3)

Therefore,

d0 = 46 –20 = 26
d1 = 20 – 12 = 8
d2 = 12 – 6 = 6
d3 = 6 – 2 = 4
d4 = 2 – 0 = 2

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Note that the sum of the dx values must equal l0; that is,
Σdx = l0 (4)

And in the example,

Σdx = l0 = 46

The age-specific mortality rate (qx) is the proportion of individuals at

the start of age interval x who die during the age interval:

qx = d x (5)
lx

also expressed as the probability of an individual dying during that

interval. For our data, the age-specific mortality rates are

q0 = 26/46 = 0.57
q1 = 8/20 = 0.40
q2 = 6/12 = 0.50
q3 = 4/6 = 0.67
q4 = 2/2 = 1.00

The age-specific survival rate (sx) for age interval x is the proportion
of individuals alive at the start of the interval who do not die during
that interval period. In other words, sx is the probability of surviving
age interval x:
sx = 1 – qx (6)

For the present data,

S0 = 1.00 – 0.57 = 0.43

s1 = 1.00 – 0.40 = 0.60
s2 = 1.00 – 0.50 = 0.50
s3 = 1.00 – 0.67 = 0.33
s4 = 1.00 – 1.00 = 0.00

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We can calculate age-specific life expectancy (commonly done for
human populations) as follows. Let us define Tx, as the number of
time units left for all individuals to live from age x onward; this is
obtained by summing Lx values as follows:

last
Tx = Σ Li (7)
i=x

or Tx = Lx + Tx + 1 (8)

and expressing Tx in time units; so

T4 = L4 = 1 yr

T3 = 4 + 1 = 5 yr
or T3 = L3 + T4 = 4 + 1 = 5 yr

T2 = 9 + 4 + 1 = 14 yr
or T2 = L2 + T3 = 9 + 5 = 14 yr

T1 = 16 + 9 + 4 + 1 = 30 yr
or T1 = L1+ T2 = 16 + 14 = 30 yr

T0 = 33 + 16 + 9 + 4 + 1 = 63 yr
or T0 = L0 + T1 = 33 + 30 = 63 yr

Then, the life expectancy for an individual of age x is

ex = Tx / lx (9)

so
e0 = 63 animal-yr/46 animals = 1.37 yr
e1 = 30/20 = 1.50 yr

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 e2 = 14/12 = 1.17 yr
e3 = 5/6 = 0.83 yr
e4 = 1/2 = 0.50 yr

Life expectancy represents the average additional length of time

that an individual will live, once it has reached age x.

To compare different populations, the numbers dying (dx) or surviving

(lx) are often expressed as numbers per 100 or per 1000 individuals
entering the population at age 0; that is, 10 is set to 100 or 1000, and
all other life-table entries are expressed relative to this value. For
the example in our table, we have 10 = 46 and 11 = 20. If we set 10 =
100, then we would have an 11 value of (20/46) x (100) = 43.5. In
order words, for every 100 individuals born into the population, 43.5
survive to age 1. If we begin with 10 = 100, then of course Σdx = 100.

In the above example, the data collected were six values of Lx, from
which we computed all the values of lx, dx, qx, sx, Tx, and ex. If,
instead, the data collected were lx’s, then all of the Lx’s would have
been calculated by Equation 1, and the other quantities would have
been ascertained using Equations 4 through 9. If the original data
were values of dx, then we would have used Equation 4 to determine l0
and then, for x > 0,

lx = l x-1 – d x-1

which would have yielded

lx = 46
lx = 46-26 = 20
lx = 20-8 = 12
lx = 12-6 = 6
lx = 6-4 = 2
lx = 2-2 = 0

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The remaining life-table statistics would be computed using
Equations 5 through 9.

Although the construction of a life table is straightforward, given

data on survival of various age classes, the life table may be
inaccurate or invalid. Especially questionable are life tables based on
capture-recapture of marked or banded individuals. Life tables based
on such data involve two assumptions:

1. Annual survival rate is age-specific; it varies only by age and not

by year.
2.Recovery rates are constant over all ages and all years. Rarely are
these assumptions met. In fact, annual survival rates, especially
among birds, do vary by year, often influenced more by weather
than by age.

Survivorship Curves

Various types of graphs may be constructed from life-table data,

including mortality rate curves, life expectancy curves, and
survivorship curves. Most widely used among ecologists, the
survivorship curve is prepared by plotting (usually on semilogarithmic
graph paper) the logarithm of the number of survivors against age.
For comparative purposes use lx data bases on 100 or 1000. From this
graphical presentation, three basic types of curves are recognizable.

In the type 1 survivorship curve, there is a high survival rate of the

young and a low survival rate after a particular old age. In the type 2
curve, a constant rate of’ mortality occurs at all ages (a constant
percentage of population decreases each time period).

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The type 3 curve shows a high juvenile mortality and a relatively low
rate of mortality thereafter. Most populations exhibit survivorship
curves between types and 2 or between types 2 and 3. Some adult
birds approach type 2, and modern humans approach type 1. Including
data from very early life stages (even eggs) tends to introduce type
3 curve characteristics.

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 FECUNDITY TABLE

If we know the productivity of each age class of females, mx,

determined by litter counts, brood counts, placental scars, young
fledged, and so on, we can construçt a fecundity table.

The fecundity table includes the age categories, x; age-specific

survivorship from the female life table, lx; age-specific productivity,
mx; the mean number of female young produced by each female of
age x, lxmx; which is mx weighted by survivorship. The sum of the lxmx
column gives the net reproductive rate, Ro.

Added to the fecunditv table is another column xlxmx, which records

the values obtained by multiplying the lxmx by the appropriate age.
The sum of this column is used to compute the rate of increase.

In Table 2, age categories have been converted to mean age to

permit the correction calculation of xlxmx, starting with year class 0.
Some female gray squirrels may produce a litter late in the year of
their birth.

The value of R0 (1.169) is just above the replacement rate that would
be expected in an unhunted gray squirrel population. A hunted
population would have a higher reproductive rate and few, if any,
individuals in the 5 to 7 year age classes.

lx = age-specific survivorship from the female life table

mx = age-specific productivity
lxmx = the mean number of female young produced by each female of
age x
Ro = net reproductive rate

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Table 2. Fecundity table for the Gray Squirrel Population

x lx mx lxmx xlxmx
0-1(0.5) 1.000 0.05 0.050 0.025
1-2(1.5) 0.253 1.28 0.324 0.486
2-3(2.5) 0.116 2.28 0.264 0.660
3-4(3.5) 0.089 2.28 0.203 0.710
4-5(4.5) 0.053 2.28 0.132 0.594
5-6(5.5) 0.039 2.28 0.089 0.489
6-7(6.5) 0.025 2.28 0.057 0.370
7-8(7.5) 0.022 2.28 0.050 0.375
Σ lxmx = 1.169 Σ xlxmx =
3.709

Ro = net reproductive rate = 1.169

In keeping with our earliest interpretation of Ro, this value of Ro

indicates that the population is growing and that each individual
produces on the average 1.169 offspring. Remember that if Ro is less
than 1.0, then the population would be expected to decline, and if it
exactly 1.00, then the population should remain constant in size.

RATE OF INCREASE

Given a life table and a fecundity table, we can determine the rate of
increase, rm, for a population in a particular environment.
An approximation of r can be obtained by the equation

r = Σ lxmx logeΣ lxmx

Σ xlxmx

For the squirrel population an approximation is

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r = 1.169 (0.1561) = 0.049
3.709

Another method requires more calculations but gives the same result.
The first step is to determine mean generation time

Tc = Σ xlxmx
Ro

For the squirrels

Tc = 3.709/1.169 = 3.173

To find an approximate value of rm the following formula applies:

rm = loge Ro /Tc = loge1.169/3.173

= 0.1561/3.173
= 0.049

MEAN GENERATION TIME

Tc = Σ xlxmx
Ro

When generations are overlapping, the length of a generation is not

clear. One way to visualize the generation length in this case is as the
average age of an individual when offspring are born.

Where Tc is the generation length or mean generation time.

Tc = 3.173

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 REPRODUCTIVE VALUE

The life table and fecundity table provide the data needed to
calculate the reproductive values of females age x in the population.

The reproductive value is the relative number of female offspring

that remain to be born to each female age x.

To state it differently, it is the number of offspring that will be

produced by a female from age x until the end of her life. It can be
estimated relative to the reproductive value of the female at birth,
vo, which is 1 by the formula given by the geneticist R.A. Fisher.
∞
vx/vx = rrx / lx Σ e-ry lymy
y=x

Because the reproductive value of a female at birth is 1, vo can be

removed from the formula. The formula can also be written as

vx = erx Σ e-ry lymy

lx y=x

where y = all ages a female passes through from age x on up.

Essentialy the formula states that the number of female offspring
produced at any one moment of time by females aged x and over is
divided by the number of females of age x at any one moment.

To calculate reproductive values for females of each age class (Table

3).

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1. Add the e-rx lxmx column from the bottom up to obtain a value for
e-ry lymy for each age class

2. Divide this value by e-rx lx for each age class. The reproductive
values for the squirrel population are graphed in Figure 1. Note that
the reproductive values rise and fall by age.

Table 3. Determination of reproductive values for squirrel population

Let r = 0.050

x lx rx e-rx e-rx lx e-rx lxmx e-ry lymy vx

0.5 1.000 0.025 0.9753 0.9753 0.0478 0.9999 1.025
1.5 0.253 0.075 0.9277 0.2347 0.3005 0.9521 4.056
2.5 0.116 0.125 0.8825 0.1024 0.2329 0.6516 6.360
3.5 0.089 0.175 0.8394 0.0747 0.1703 0.4187 5.605
4.5 0.058 0.225 0.7958 0.0463 0.1054 0.2484 5.365
5.5 0.039 0.275 0.7595 0.0296 0.0675 0.1430 4.831
6.5 0.025 0.325 0.7225 0.0180 0.0412 0.0755 4.194
7.5 0.022 0.375 0.6873 0.0151 0.0343 0.0343 2.271

vx = e-ry lymy
e-rx lx

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 7

5
Reproductive value

0
0 1 2 3 4 5 6 7 8
Age in years

Figure 1. Reproductive value of the squirrel population described in

Table 3.

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