Biology 103

A Method of Population Estimation: Mark & Recapture

!"ecti#es:
1. Learn one method used by wildlife biologists to estimate population size of wild animals.
2. Learn how sampling size effects accuracy and standard error of estimates.
3. Use the mark-recapture method to estimate the number of coins in a jar.
$% $ntroduction
o understand how different physical or biological factors influence the distribution or abundance of species!
we usually need to measure changes in population abundances o"er space or time. #cologists interested in
$uestions about populations and communities fre$uently face a "ery basic $uestion% how best to estimate the
size of a population of organisms in the field& 'owe"er! it usually is not possible to obtain a complete count or
census of a natural population of animals (and it is often difficult e"en for plants)*. +or this reason! ecologists
generally ha"e to rely on some kind of estimate of abundance or density. , "ariety of methods e-ist to do the
job. the utility of each "aries depending on such factors as the type of organism! the habitat in which the
population occurs! and the time and e$uipment a"ailable.
he methods ecologists use "ary in their le"el of precision and difficulty of application. ,t one e-treme is a
complete census of indi"idually identifiable organisms. his approach might in"ol"e tagging all trees in a
forest plot! or marking all the animals of a species li"ing in an area at a particular time. echni$ues for
indi"idually identifying animals include%
/ompiling a catalog of photographs or drawings that show uni$ue markings of indi"iduals (e.g.!
fin shape and markings whales. spot patterns on ,frican wild dogs. whisker patterns in lions. facial
e-pressions in primates*.
agging indi"iduals with markers "isible from a distance or in the hand (e.g.! numbered metal leg bands
on birds. colored plastic leg bands in uni$ue patterns on birds. strings of beads around the necks of
small primates. wing tags on bats*.
,t the other e-treme are 0$uick 1 dirty2 methods that gi"e crude estimates of abundance. #-amples include
counting all birds sighted along a roadside route one tra"els by car. counting the number of monarch
butterflies passing a point such as /ape 3ay during migration. counting the number of bird songs one hears
during a morning in the spring. scanning aerial photographs of wildebeest herds on the ,frican plains. or
counting animal tracks along a trail.
4ne may count the number of singing males of a species of bird in a gi"en community or defined area and then
estimate the size of the breeding population by assuming that for e"ery male there is one female. 4r one may
count the indi"iduals in a sample area and then e-trapolate to the larger area in which the whole population is
assumed to li"e. 5ecause each method has different assumptions and hence! different strengths and
weaknesses! it is recommended (though rarely done* that at least two estimation methods be used and
compared in any population study.
, "ariety of methods for estimating population size fall between the two e-tremes outlined abo"e. hese
methods in"ol"e marking a subset of a population! followed by later counts of the relati"e numbers of marked
and unmarked indi"iduals. his type of method is more precise than a crude census in which no organisms are
marked at all! but takes less time and e-pense than does an e-hausti"e marking program.
&he Mark and Recapture &echni'ue
5y far the most popular way to measure the size of a population is called the 3ark and 6ecapture
echni$ue. his techni$ue is commonly used by fish and wildlife managers to estimate population sizes before
fishing or hunting seasons. he mark and recapture method in"ol"es marking a number of indi"iduals in a
natural population! returning them to that population! and subse$uently recapturing some of them as a basis for
estimating the size of the population at the time of marking and release. his procedure was first used by /.7.8.
9etersen in studies of marine fishes and +./. Lincoln in studies of waterfowl populations! and is often referred
to as the Lincoln :nde- or the 9etersen :nde-. :t is based on the principle that if a proportion of the population
was marked in some way! returned to the original population and then! after complete mi-ing! a second sample
was taken! the proportion of marked indi"iduals in the second sample would be the same as was marked
initially in the total population. hat is!
R (marked recaptures* M (marked initially*
;;;;;;;;;;; < ;;;;;;;;;;;;
& (total in second sample* ( (total pop. size*
5y rearrangement we can estimate the population size! =! to be!
M)&
( < ;;;
R
+or e-ample! suppose you took 2>> mice out of a forest ha"ing an unknown number of mice! put leg bands on
them! return them to the forest and let them mi- thoroughly. :f you then take 2?> mice from the forest and find
?> of them to be ha"e leg bands! then M * +00! & * +,0! R * ,0! and the unknown total number of mice (=*
could be estimated as%
( * M)&-R * .+00/.+,0/-,0 * 1000 mice
@ample problem%
, pest control technician captures and applies ear tags to 23 brown rats! which he then releases. , week later
he traps 2A brown rats! 11 of which ha"e ear tags. Bhat is the estimate of the total population of brown rats&
Assumptions% he accuracy of this mark-recapture method rests on a number of assumptions being met.
Assumption 1. Curing the inter"al between the preliminary marking period and the subse$uent recapture
period! nothing has happened to upset the proportions of marked to unmarked animals (that is! no new
indi"iduals were born or immigrated into the population! and none died or emigrated*. ,nother way to state
this assumption is that the population is assumed to be DclosedD. ,n e-ception is when there are e$ual
proportions of deaths or emigrants for marked and unmarked indi"iduals during the sampling period. :f
deaths occur in this manner the proportions of marked to unmarked will not change and thus we can still
estimate the population. =otice the same cannot hold true for births since a proportionate number of
indi"iduals will not be born with marks on them. 5irths can still occur and an accurate estimate can still be
made only if an e$ual number of unmarked indi"iduals lea"e (or die* and are born. his means that no
marked indi"iduals can lea"e! which would be highly unlikely.
=otice that inherent in this assumption is that the marking of the indi"iduals has no effect on their sur"i"al.
his is often the most difficult assumption to meet since the mark may make the indi"idual more noticeable
to predators or may slow them down! such that more marked indi"iduals die than unmarked between the
sampling periods.
Eou should also be aware that the length of the sampling period is "ery important. he shorter the time
inter"al between marking and recapturing indi"iduals reduces the likelihood that this assumption will be
"iolated. he longer the time inter"al the greater the chance that some indi"iduals will die! emigrate!
immigrate or e"en be born and hence become an DopenD population.
Assumption +. he chance for each indi"idual in the population to be caught are e$ual and constant for both the
initial marking period and the recapture period. hat is! marked indi"iduals must not become either easier or
more difficult to catch. 3arking an indi"idual does not make them more or less likely to be recaptured
relati"e to unmarked indi"iduals. +urthermore! there cannot be something uni$ue about those animals that
were initially marked that make them more conspicuous. :f only males are captured in the traps because
females are busy incubating eggs or feeding! then males may be more likely to be recaptured. 'ence the
population estimate would only be appropriate for those animals with that uni$ue beha"ior that make them
easier to capture (i.e. males*.
,nother problem deals with Ftrap shyF or Ftrap happyF indi"iduals. @ome techni$ues for capturing and
marking animals can ha"e a dramatic effect on the animal. 0rap shy2 animals $uickly learn to a"oid those
traps or that area where the capture occurred making them less like to be recaptured. he opposite can also
occur! especially if the trapping method uses bait. , 0trap happy2 indi"idual is lured into a trap with food
and is then marked. Later during the recapture period the animal may recall that these traps had rewards
(food* and that other than a little decoration (the mark* it made out pretty good. herefore it becomes 0trap
happy2 and is more likely to be captured in the trap compared to unmarked indi"iduals who ha"e not
e-perienced the rewards.
,lso notice that there cannot be any changes in beha"ior among indi"iduals that may affect capture
probabilities. +or e-ample! in some species there are different times of peak acti"ity for different classes in
the population. ,dult males may be most acti"e in the morning. 7u"enile males and females may be most
acti"e midday! and adult female may be most acti"e in the afternoon. :f you mark indi"iduals in the
morning you will capture and mark mostly males. 5ut if you perform your recapture sample at midday or
the afternoon! you will capture mostly unmarked ju"eniles or females! respecti"ely. ,gain! you should
realize that the length of time between the sampling periods plays an important role in whether this
assumption is "iolated.
Assumption 3 . @ufficient time must be allowed between the initial marking period and the recapture period for
all marked indi"iduals to be randomly dispersed throughout the population (so that assumption 2 abo"e
holds*. 'owe"er! the time period must not be so long that ,ssumption 1 breaks down. :f marked animals
failed to disperse from the sampling area! or if they tend to aggregate in groups! there is a higher probability
that they will be recaptured relati"e to unmarked indi"iduals (or not depending on where they aggregate*.
Assumption 0% ,nimals do not lose their marks. his is an important factor for animals that shed or molt as
they grow or as they respond to seasonal factors.
,ll these assumptions must be considered "ery carefully and play a strong role in the techni$ues used to
perform the study. , certain method of trapping may "iolate one assumption while another trapping method
may "iolate another assumption. 'a"ing too short of a time inter"al between sampling may lead to the
"iolations of one assumption (which one(s*&* while an inter"al that is too long will "iolate others (which
one(s*&*.
/learly the decisions the researcher must make during such a study re$uire an e-tensi"e knowledge of the
natural history of the organism. Co beha"ioral patterns change o"er the day& Bhen do they breed& Bhen does
most of their mortality occur& Bhat are their predators& 'ow $uickly can they mo"e& ,re they territorial&
Guestions like these must be answered before the e-periment is designed and performed so that a minimum of
"iolations to the assumptions are made.
$$% Methods 11 Eour goal is to estimate a population size using the mark-recapture method. Eou will use an
unknown amount of beans as your model population.
#ach pair of students needs these materials%
1-H>> ml 5eaker which represents the habitat to be sampled
1-small 5eaker (sampling beaker*
1-I>> ml 5eaker with a known number of white beans 1-
I>> ml 5eaker with colored beans (marked indi"iduals*
1-I>> ml beaker (empty! to hold beans remo"ed from the population*
Effort 1:
1. 4btain a population of white beans! a large habitat beaker! colored beans! and one small beaker. he white
beans will represent the population to be sampled.
2. Marking% Using the small beaker as a JtrapK! push through the white bean population once! filling the trap
with beans to the 1> ml line. /ount the number of white beans caught and for each white bean caught!
substitute a colored bean and record that number as% (um!er marked * M% (#.g.! if you JtrappedK 23 white
beans with your sampling beaker you would put 23 colored beans into the original population*.
3. horoughly mi- the marked and unmarked bean population. (Bhich assumption does this fulfill&*
I. Recapturing% =ow sweep through the mi-ed bean population with the trap once! filling the trap to the 1>
ml line. :f you reco"ered at least one colored bean with this sweep! record the number of marked beans
(recaptures* as R. 6ecord the total number of beans caught as &! ( i.e. marked 2 unmarked*.
?. =ow calculate the bean population size using the formula ( * M)&-R
Effort +:
6epeat steps 2! 3! I! and ?! but this time fill the trap to the 2> ml line at L2 and LI. hat is! you will be
doubling your sampling effort.
Effort 3%
6epeat steps 2! 3! I! and ?! but this time fill the trap to the 3> ml line at L2 and LI. ,gain! you are increasing
your sampling effort.
$$$% Results
:n a graph! compare the entire classFs mean estimates for population size for each of the three sampling efforts.
Bhat happens to each of the parameters listed below when sampling effort is first doubled (1> ml "s. 2> ml*
and then increased again by ?>M (2> ml "s. 3> ml*&
Mean of the estimates at each 3Effort3
4tandard error of the mean of the estimates
Range of estimates
,ssuming cost is an issue! which of the three sample sizes would be the best &
(:n the sampling of actual wildlife! cost usually is an issue. :t can be e-pensi"e and time-consuming and labor-
intensi"e to sample wild populations. 8etting accurate results without consuming to many resources Ntime!
money! possible harassment of animal populationO is a worthy goal. *
$5% 6onclusion
1. #"aluate how well you were able to satisfy each of the assumptions of the mark-recapture techni$ue.
2. :f migration occurred in a natural population being studied! how would this influence the reliability of
your estimate of population size determined using the mark-and-recapture techni$ue& Bould your
population estimate be too high or too low! or would you not be able to predict how your estimate would
be biased& (6emember% migration consists of both immigration and emigration.
3. Ciscuss the kind of bias you would e-pect to be generated in the estimate of population size (andPor the
standard error* if each of the remaining assumptions (2! 3! and I* were "iolated. (+or e-ample% :f
,ssumption 2 is not met! would it cause the size of the population to be o"erestimated& underestimated& 4r
ha"e no effect& * =ote that it is possible that "iolating se"eral different indi"idual assumptions may ha"e
the same effects.
ne final statement% here are a number of other models (designs and e$uations used to estimate population size. #ach has their own set of
assumptions. hey are often grouped into open and closed models depending on whether the population is assumed to be open or closed.
here are many modifications of this model that allow one to rela- some of the assumptions but typically this re$uires
more sampling effort. 5ut as you can imagine the comple-ity of the mathematics can greatly increase.