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Biology 206

LAB 8

Objectives

1. To become familiar with standard procedures and calculations used in estimating

population sizes of organisms.

2. To understand the assumptions associated with mark-recapture censusing techniques.

3. To explore how mark-recapture methods can be used to estimate the number of

organisms in a population.

Some text in this lab modified from: Kingsolver, R.W. 2006. Ecology on Campus. Pearson Education, Inc.

One of the first questions an ecologist asks about a population is, How many individuals are

here? This question is trickier than it appears. First, defining an individual is easier for some

organisms than others. A head count of Canada geese captured on the ground during their

summer molt gives a clear indication of adult numbers, but should eggs be counted as members

of the population or not? In plants, reproduction may occur sexually by seed, or asexually by

offshoots that can remain connected to the parent plant. This reproductive strategy, called clonal

reproduction, makes it difficult to say where one individual stops and the next one begins.

Population ecologists must make decisions about how to define an individual when counting

organisms.

Once the individual is defined, ecologists working with stationary organisms such as trees or

corals can use spatial samples, called quadrats, to estimate the number of individuals in a larger

area. Mobile animals are usually simpler to define as individuals, but harder to count, because

they tend to move around, mix together, and hide. Quadrats are not a good approach with mobile

animals because immigration and emigration in and out of the study site make it hard to know

what area the entire population occupies. For largemouth bass in a pond, you could easily draw

a line around a map of the population, but how would you define the edges of a population of

house sparrows in your community? Although house sparrows tend to be more concentrated in

towns and urban areas, they do not stop and turn back at the city limit sign. For zoologists, a

fuzzy definition of the space occupied by the population often forces an arbitrary designation of

the survey group, such as the population of robins nesting on campus in the spring. Knowing

the number of animals in a designated area is interesting, but we must bear in mind that the

ecological population is defined in terms of interactions among organisms of the same species

and not by convenience.

After defining the individual and establishing the limits of the population you wish to count, your

next task is to choose a counting method. Arctic and prairie habitats such as the tundra lend

themselves to accurate survey by aerial counts. This approach works poorly in forests, at night,

underwater, or in soil habitats. If animals can be collected or observed in a standard time or

collecting effort, you can get an idea of relative abundance, but not absolute numbers. For

1

Biology 206

LAB 8

example, the number of grasshoppers collected in 50 swings with an insect net through an old

field community produces data that could be used to compare relative abundance in different

fields, but would not tell you how many grasshoppers were in the population.

For estimates of absolute numbers, mark-recapture methods can be very effective. The first

step is to capture and mark a sample of individuals. Marking methods depend on the species:

birds can be banded with a small aluminum ankle bracelet, snails can be marked with waterproof

paint on their shells, butterflies can have labels taped to their wings, large mammals can be fitted

with collars, fish fins can be notched, and amphibians can have nontoxic dyes injected under the

skin. Marked animals are immediately released as close as possible to the collection site. After

giving the animals time to recover and to mix randomly with the whole population, the ecologist

goes out on a second collecting trip and gathers a second sample of the organisms. The size of

the population can then be estimated from the number of marked individuals recaptured on the

second day.

The assumption behind mark-recapture methods is that the proportion of marked individuals

recaptured in the second sample represents the proportion of marked individuals in the

population as a whole. In algebraic terms,

=

N = estimated population size

R = number of animals recaptured at a second time

S = size of the sample on the second capture

Lets consider an example. Suppose you want to know how many box turtles are in a wooded

park. On the first day, you hunt through the woods and capture 24 turtles. You place a spot of

paint on each turtles shell and release all turtles back where you found them. A week later you

return, and with an extraordinary effort, you catch 60 turtles. Of these, 15 are marked and 45 are

unmarked. Since you know how many turtles you marked, sampled, and recaptured, you can

figure out the size of the whole population. By the definitions above, M = 24 marked and

released, S = 60 in the second sample, and R = 15 recaptures. If the second sample is

representative of the whole population, then:

15 24

24 (60)

=

This can be rearranged to: =

= 96 turtles.

60

15

Lincoln-Peterson Index:

population size. (See box to the right.)

M = marked individuals released

S = size of second sample

R = marked animals recaptured

Biology 206

LAB 8

In the rearranged version of the general formula, notice that the smaller the number of

recaptures, the larger the estimate of population size. This makes good biological sense,

because if the population is very large, the marked animals you release into the wild will be

mixing with a greater number of unmarked animals, so you will recapture a lower percentage of

them in your second sample.

The Lincoln-Peterson method is fairly simple, and its calculations are straightforward, but it does

depend on several assumptions. Violating the conditions of the Lincoln-Peterson model can

seriously affect the accuracy of your estimate, so it is very important to bear these assumptions in

mind as you interpret your results:

1.

2.

3.

4.

5.

6.

7.

Individuals with marks have the same probability of survival as other members of the

population. It is important to choose a marking method that does not harm your animal.

If a predator used your paint marks to locate and capture marked turtles at a higher rate

than other turtles, your number of recaptures would be lower, and the population estimate

would therefore be too high.

Births and deaths do not occur in significant numbers between the time of release and the

time of recapture. If marked individuals die and are replaced with newborns, then you

will recapture few or no marked individuals, and your estimate will be too high. This is

not a large concern in studies of box turtles, but can significantly affect estimates for

rapidly breeding organisms.

Immigration and emigration do not occur in significant numbers between the time of

release and the time of recapture. If marked individuals leave the study area and new

individuals come in to replace them, you will get fewer recaptures than the equilibrium

population size would lead you to expect. To think about this another way, the real

population covers a much larger area than the habitat you thought you were studying.

Marked individuals mix randomly with the population at large. If your marked turtles do

not move among unmarked turtles, and you recapture them near the place you released

them, then recaptured turtles may be overrepresented in your second sample, driving

down your population estimate.

Marked animals are neither easier, not harder, to capture a second time. If marking an

animal frightens it so that it hides from you a second time, then recaptures will be

underrepresented in a second sample. If animals become tame and are easier to

recapture, then the opposite error is introduced.

Marks do not come off your marked organisms. Invertebrates molt and shed marks,

mammals can wriggle out of their collars, and many things can happen to obscure your

marks. If this happens, recaptures will be undercounted, and your estimate will be too

high.

Recapture rates are high enough to support an accurate estimate. The Lincoln-Peterson

calculation tends to overestimate the population size, especially if the number of

recaptures is small.

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Biology 206

LAB 8

generate sufficient recaptures in large populations. To

correct for this source of error, ecologists often use a

slightly modified form of the Lincoln-Peterson index,

called the Bailey correction (see box at right). There are

other mark-recapture methods designed to account for

violations of several other assumptions of the LincolnPeterson method, but most of these require repeated

sampling and day-specific marks. For this lab, we will

simply use the Lincoln-Peterson method with the Bailey

correction to estimate the size of a population.

Lincoln-Peterson Index:

=

( + )

+

M = marked individuals released

S = size of second sample

R = marked animals recaptured

After you calculate a value for N, you should use the following formula to calculate the 95%

confidence intervals on your estimate of population size. Confidence intervals give you an idea

of the accuracy of your estimate. By adding a standard error term for the upper limit, and

subtracting the same amount for a lower limit, you know with 95% confidence that the true

population size falls within this range. In common language, we could say the true population

size is our estimated value of N, give or take the confidence interval. With a 95% confidence

limit we would be wrong only 5% of the time. The formula for 95% confidence intervals, using

the Bailey method is shown in the box below:

Confidence Interval for population size estimate:

( + )( )

!

% = .

( + ) ( + )

N = population size estimate

M = marked individuals released

S = size of second sample

R = marked animals recaptured

In summary, to estimate the size of a population: 1) define what you mean by an individual, 2)

determine the area you will study, 3) decide on a counting method, and then 4) calculate your

estimate and confidence intervals.

Biology 206

LAB 8

Procedure

*Please be sure youre using the right color pen and marking your crickets on the correct place

on the crickets bodies. Your TA will let you know the color/location of marks to use.*

1. One member of your group should take one cricket castle (portion of an egg carton) out of

the aquarium try not to disturb too many of the other cricket castles - and gently shake the

crickets into the small plastic tank. Make sure youve collected between 4-6 crickets for your

group. Replace the cricket castle (and extra crickets, if necessary) in the aquarium.

2. Take the crickets in your container to your desk and mark all of these crickets with a paint

pen. Be careful not to let them escape! Record the number of crickets you marked (Mi) in the

table below.

3. Gently return the marked crickets back to the aquarium.

4. After at least 15 minutes, take another sample of crickets from the aquarium. Remember, you

are trying to sample randomly, so dont look at the crickets themselves before selecting your

cricket castle. Continue until youve captured at least 5 crickets.

5. Bring your second sample of crickets back to your bench. Record the number of crickets in

your second sample (Si), and the number of them which are marked (Ri) (with the marks made by

YOUR section) in the table below.

6. Sum up the values from Table 1 below for all the groups in your section. Record the summed

values in Table 1 of the worksheet. Use those data for the worksheet calculations.

Table 1. Data on marked and recaptured crickets

from YOUR group. Do not use these data for

calculations! Share this information with your

section and use the combined values from all groups

in your section to do your calculations.

Paint-marked garden

snail (Cepea normalis).

Eastern bluebird

(Sialia sialis) marked

with color bands.

5

Mi

Si

Ri

elaphus).

Biology 206

LAB 8

Lab 8: WORKSHEET

1. Complete the table below with the summed values of all the groups in your section.

Table 1.

Variable

Value

M

S

R

2.

What was your true population size (the actual number of crickets in the aquarium your

TA will give you this information!)? _____

3. Use the values in Table 1 to calculate your estimated population size using the standard

formula for the Lincoln-Peterson index. Show your work!

N = ______________________

4. Use the values in Table 1 to calculate your estimated population size using the formula

for the Bailey Correction for the Lincoln-Peterson index. Show your work!

N = ______________________

Biology 206

LAB 8

5. Calculate the 95% confidence intervals for the population estimate you calculated in

number 4.

6. How close was your estimated population size (N) to the actual number of crickets in the

population? Which formula for calculating N resulted in a more accurate estimate? Was

the true population size within the 95% confidence interval you calculated?

7. Why was it important to randomly choose the cricket castle for your second sample,

rather than picking out one with clearly visible crickets? Can you imagine similar

sampling issues arising with the box turtle example from the Introduction?

8. Why is it important that we waited for a while between the first and second sample? If

we hadnt, how would that have affected your estimate of population size? Can you

imagine a comparable problem arising in a field study?

Biology 206

LAB 8

9. Looking at the formula for confidence intervals, how would you expect the 95% interval

to be affected by increasing the size of the second sample? How would it be affected by

marking a larger number of individuals?

10. Suppose some of the crickets escapes from the aquarium between the release of your

marked individuals and your second sample. Would your estimate of population size be

too high or too low? Explain why. How might a comparable problem arise in a field

study?

11. Suppose your TA added unmarked crickets to the aquarium between the release of your

marked individuals and your second sample. Would your estimate of population size be

too high or too low? Explain why. How might a comparable problem arise in a field

study?

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