REMOVAL SAMPLING/REMOVAL METHOD

The removal method of estimating populations has been widely used

in studies of small mammals. The method involves the removal of

individuals from the population either permanently or for the dura-

tion of the study.

The basis of these methods is the expectation that the number

caught and removed from the population at a given time of trapping

will be greater than the number caught at later trapping using the

same sampling effort.

That is, as one reduces the population size, the size of the catch will

decrease.

Necessary assumptions of removal methods are listed below.

1. Each individual in the population has an equal and independent

chance of being captured. That is, the sampling must be random.

2. Except for the effects of the trapping, the population is not

increasing or decreasing in size (through the combined effects of

births, deaths, immigration, and emigration).

3. The probability of capturing an individual is the same for each

period of sampling.

Violation of any of these assumptions will result in inaccurate

population estimates.

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The first assumption relates to the collecting technique. Capture

should not favor one sex, one age class, or one individual over

another. Each sample taken should represent a random collection

from the entire population. For example, if some members of the

population consistently avoid capture, then the size of the population

will be underestimated.

In capture-recapture theory, births or immigration may not occur

during the study, although random mortality and emigration are

acceptable. When removal methods are used, a population may have

births and immigration, but (according to assumption 2 above) they

must balance with normal deaths and emigration so that population

size remains constant. An increase in the population size between

sampling periods would give an overestimate of population size.

According to the third assumption, if the chance of capturing

animals in the population changes from the first sampling period to

the second sampling period, then the population estimate will be

biased. Therefore, the sampling effort must be the same for each

sampling period.

If the probability of capture increases between sampling periods,

then the population would be overestimated. In capture-recapture

sampling, the probability of capture need not be constant; what is

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required instead is that the ratio of marked to unmarked individuals

remains the same.

Trap-shyness or trap-proneness can affect the probability of

capture. If trap-prone animals are more likely caught the first

sampling period, the probability of their capture will be high.

Therefore, during the second sampling period, the probability of

capture will be lower, and fewer animals will be caught due to a

higher proportion of trap-shy animals.

Other factors, such as bait acceptance, weather conditions, and

differential activity of ages and sexes, affect the probability of

capture. Thus one must ensure that sampling and environmental

conditions are as identical as possible during all sampling periods and

that the sampling effort remains constant.

Finally, it should be realized that the error in estimating population

size by these methods is smallest when large proportions of the

population have been sampled. If only a very small proportion of the

total population has been captured, then the confidence in the

population size estimate is very low.

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Procedures

The procedure requires at least two periods of sampling. Sampling

requires setting the same number of traps for several days or

nights. For small mammals these traps may be snap traps or live

traps. If live traps are used, all mammals caught in each sampling

period must be marked and not counted if trapped again. The

sampling may involve successive removals of animals from the

population, the Hayne (1949) method; or it may involve only two

periods of sampling, the Zippin (1958) method.

A field procedure is as follows:

1. Set the traps in a grid system, 3 traps to a station; or in two

parallel lines 50 meters apart, 20 stations to a line, 3 traps at

each station. Space the stations 25 or 50 meters apart, depending

upon the nature of the vegetation or study site.

2. Prebait for best success.

3. Trap for several successive periods, or for two periods on each of

several dates. Depending upon the species, trapping periods may

be 24 hours or nighttime only.

A problem with trapping over a number of successive periods is the

possibility of attracting new animals into the sampling area as the

population is reduced, particularly if the sampling design is a grid.

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Traps on the outer stations usually capture more animals than those

in the center of the grid, because animals in the border zone react

to the sudden removal of animals in the center.

These immigrants, picked up by the border traps, contribute

significantly to the catch on the outer grid lines and influence

population estimates.

Regression Method

In the Hayne method the daily catch is plotted against the

number of animals previously caught. A line can be drawn through

the data points to cut the horizontal axis. The point at which the

horizontal axis is cut represents the population estimate. A more

accurate method is to calculate a simple regression line from the

catch data. The slope of the line represents the average proportion

of the population during each sampling period.

250

200
Num ber Caught, Y

y = -0.5x + 200
150

100

50

0
0 100 200 300 400
Total Num ber Previously Caught, X

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Figure 1. The number of captures each sampling period as a function
of the accumulated prior catch. The data are form Table 1.

Table 1. Numbers of animals caught in four successive sampling

periods. These data are plotted in Figure 1.

Sampling period (i) 1 2 3 4

Number of animals 200 100 50 25
caught (Yi = ni)
Accumulated prior 0 200 300 350
catch (Xi)

One useful method for graphical estimation of population size, N, is

based on successive removals of animals from the population; such

procedures have developed since the early part of this century.

Presented here is the method of Hayne (1949), which is a

modification of Leslie’s method (Leslie and Davis, 1939).

In this procedure, one obtains a series of collections, capturing ani-

mals at different times and removing them from the population. The

amount of collecting effort must be the same each time. So, for

example, one may tabulate the number fish caught in an 8-hour

period, or the numbers of mammals trapped in 24-hour period, on

each of several dates.

The numbers of animals caught are then plotted against the total

numbers previously caught, as shown in Figure 1 and for the data of

Table 1.

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In this example, 200 animals were caught, with a sampling effort, in

sample 1; 100 were caught with same sampling effort in sample 2;

and so on. A total of 0 animals were accumulated prior to sampling

period 1; 200 animals were accumulated before sample 2 was taken;

200 + 100 = 300 animals were removed from the population prior to

the third sample; and so on.

If the probability of capture remains constant, the points on the

graph should fall along a straight line. In this example, probability of

capture is 0.50 (50% of the remaining population is removed with

each sampling). If this line is extrapolated to the horizontal axis

(the line in Figure 1), the total accumulated catch of 400 would then

be the total original population size, N. This point of extrapolation

represents the theoretical condition of a total census when all

animals have been removed and counted.

For the extrapolation to be dependable, however, one must count a

very large proportion of the population and obtain enough samples to

draw a reliable line through the several data points. The line might

be drawn by eye if the points are obviously along a straight line

(although the linearity of points in Figure 1 is unlikely to be encoun-

tered with real data). But in general, Microsoft Excel should

calculate a regression line automatically. In which case the computed

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slope of the line indicates the average proportion of the population

removed with each sampling.

To determine the statistics a and b in the equation;

y = mx t c = formula for linear regression

Yi = a + bXi (1)

where Xi is the accumulated catch at period i, and Yi is the number

caught at period i (Yi = ni, the number caught in sample i). The slope,
b, of the regression line will be a negative value, for Yi decreases as
Xi increases. Once a and b have been calculated by regression
analysis, one can calculate N, which is the value of Xi when Yi = 0.
That is, by substituting in Equation 1,

0 = a + bN (2)

and it follows that

N = —a/b (3)

For example, if a = 200 and b = -0.50, as in Figure l, then

N = —(200)/(—0.50) = 400.

4. The Moran-Zippin Method

This procedure for estimating population size (Moran. 1951; Zippin,

1956; 1958) requires fewer sampling periods than Hayne’s

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extrapolation method, although the two population estimates should

be the same or very similar.

The basis of the method is as follows.

Let N be the population size, n1 be the number of animals caught and

removed during the first sampling period and n2 be the number

caught and removed on the second sampling period. Thus, the

proportion of the original population captured in the first group of n1

animals is removed from the population, N – n1 animals remain. The

proportion of this remaining number of animals captured in the

second sample is n2 /(N – n1). If we can assume that the two

trappings caught the same proportion, p, of animals (which is the

same as saying that p is the probability of animal being captured),

then

p = n1 /N (4)

and

p = n2 /(N – n1) (5)

Therefore,

n1 = n2 (6)
N N - n1

Solving this equation for N, the population size, we find that

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N= n12 (7)
n1 – n2

Applying this equation for estimating N to the first two samples in

Table 1, we compute

N= (200)2
(200 – 100)

= 40000/100 = 400,

the same answer obtained using Hayne’s regression method

The standard error of this population estimate is

SE = (n1)( n2) n1 + n2 (8)

(n1 – n2)2

For the above example,

SE = (200) (100) 300

(200 – 100)2

= 346410/10000 = 34.6

An approximate confidence interval for our estimate of N may be

calculated as

N + (t)(SE) (9)

Where t is Student’s t for DF = ∞. (Thus, for 95% confidence

interval use t = 1.96 and for a 99% confidence interval use t = 2.58).

Therefore, we may conclude, with 95% confidence, that the size of

the population we sampled is

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400 + (1.96)(34.6) = 400 + 68;
that is, the true population size is estimated to be between 332 and
468.

Further considerations

Often one does not want to remove animals from a study area

because this may affect the behavior or vigor of the population. In

these situations, you may use removal methods by marking each

captured animal and releasing such animals back into the population,

but not counting any marked animals subsequently caught.

If 30 animals are captured, marked, and released during the first

trapping, and 20 unmarked and 10 marked individuals are caught

during the second, then, using Zippin’s equation, N = (30)2/(30 — 20)

= 90.

Using the Lincoln-Petersen method, you will find N = (30)(30)/10 =

90. Although the two estimates of N are identical, their standard

errors are 42 and 19 respectively. In general, populations that can

be sampled by either method are sampled more reliably and

efficiently using capture-recapture procedures (Zippin, 1958).

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