Otis Etal 1978

Statistical Inference from Capture Data on Closed Animal Populations
Author(s): David L. Otis, Kenneth P. Burnham, Gary C. White, David R. Anderson

Source: Wildlife Monographs, No. 62, Statistical Inference from Capture Data on Closed
Animal Populations (Oct., 1978), pp. 3-135
Published by: Allen Press
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WILDLIFE MONOGRAPH
A Publicationof The WildlifeSociety
3 9WYS1
I k 0s
| ° k 1
L l
STATISTICAL
INFERENCE
FROM
CAPTUREDATAON CLOSED
ANIMALPOPULATIONS
by
DAVIDL. OTIS,KENNETH
P. BURNHAM,
GARYC. WHITE,ANDDAVIDR. ANDERSON
OCTOBER 1978 No. 62

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mammalpopulationssuch
studiesare frequentlyconductedon small
FRONTISPIECE. Capture-recapture by LetaBurnham.)
as snowshoeharesLepus
(Photograph
americarlus.
I.
STATISTICAL INFERENCE FROM

CAPTURE DATA ON CLOSED
ANIMAL POPULATIONS
David L. Otis,l KennethP. Burnham,2Gary C. White,3

and David R. Anderson
UtahCooperativeWildlifeResearchUnit,UtahStateUniversity,Logan,Utah84322
CONTENTS
INTRODUCTION 7 Example --- ---------------- 35

Objectives 8 Example - 36
Assumptions 9 Discussion - -- 37
Unequal Sclpture Probczbilities 11 MODEL Mtb: CAPTUREPROBABILITIES VARY
Perspectives 12 BY TIME AND BEHAVIORALRESPONSE
Commerztson the Use of this Monogralph 12 TO CAPTURE 37
ACKNOWLEDGMENTS 13 Structure (lnd Use of the Model 37
FUNDAMENTALCONCEPTS 14 Discu.s.sion 38
Data und Parumeters 14 MODEL Mth: CAPTUREPROBABILITIES VARY
Statistics and Notation 15 BY TIME AND INDIVIDUALANIMAL 38
PczrumeterEstimation 16 Structure (lnd Use of the llSodel 38
Interval Estimation 17 Simulfltion Result.s 39
HISTORICALOVERVIEW 17 Di.scu.s.sion 39
MODEL MO:CAPTUREPROBABILITIES
ARE MODEL Mhh: CAPTURE PROBABILITIESVARY
CONSTANT 21 BY INDIVIDUALANIMALAND BY BE-
Structure and Use of the Model 21 HAVIORALRESPONSETO CAPTURE 40
StmulatzonResults 22 Structure clnd U-se of the lXloclel 40
txample 23 Simulsltion Re.sult.s 41
EJcample 24 Example ---------------------- 42
t_ . .
vlscusston 24 Discu.s.sion 42
MODEL Mt: CAPTUREPROBABILITIES
VARY MODEL XItbh: CAPTUREPROBABILITIES
VARY
WITHTIME 24 BY BEHAVIORALRESPONSETO CAPTURE,
Structure flnd Use of the Model 24 TIME, AND INDIVIDUALANIMAL 43
SimulcltionResults 25 Di.scu.ssion 43
Example ---------------------------- 28 REMOVALMODELS 44
Discussion 28 Introdueticsn 44
MODEL Mb: CAPTUREPROBABILITIES
VARY Structure and Use of the Getleralizecl Re-
BY BEHAVIORALRESPONSETO CAPTURE 28 rnorcll Moclel 44
Structure and Use of the Model 28 Simul(ltion Result.s 46
Simulation Results 30 Exclmple 46
Exfample 31 Example - ---------- 48
Lxample 32 Example 49
Discussion 32 Discu.ssion 49
MODLL Mh: CAPTUREPROBABILITIES
VARY TESTS OF MODEL ASSUMPTIONS 50
BY INDIVIDUALANIMAL 33 Philo.sophy of the A,D7vrotlch 50
Structure and Use of the Model 33 Summury vf Models clnd Estimltors 51
Simulution Results 34 Speczfic Te.st.s to Perform 53
1Present address: U.S. Fish and Wildlife Service, Denver Wildlife Rea.search Center7 Denver,
Colorado80225.
2Presentaddress:U.S. Fish and WildlifeServiceSOffice of BiologicalServices,Fort Collins, Colo-
rado80521.
3 Presentaddress:Los AlamosScientificLaboratory,
Los Alamos,New Mexico87545.
6 WILDLIFE MONOGRAPHS
On the Needfor an Objective Selection APPENDIXB 105

Procedure Estimation in Model Mo 105
56
An Objective Model Selection Procedure APPENDIXC 106
57
Estimation in Alternative Models Estimation in Model Mt ------------------ 106
61
Additional Examples of Model Selection APPENDIXD 107
62
A Test for Closure Estimation in Model Mb ------------------ 107
66 108
DENSITY ESTIMATION APPENDIXE
Introduction 67
Estimation in Model Ma ------------------ 108
67
Problem Formulation APPENDIXF 110
Statistical Treatment 68 Discussion of Model M,b ------------------ 110
69
Discussion APPENDIXG 111
72
STUDY DESIGN Discussion of Model Mth ------------------ 111
74
LivetrappirlgVersus Removal Methods APPENDIXH 112
75
Closure --- ---- Estimation in Model Mbh------------------ 112
75
Eliminating Variation due to Time, APPENDIXI 113
Behavior, and Heterogeneity Discussion of Model Mtbh ---------------- 113
75
Sample Size APPENDIXJ 114
77
Recording Data Estimation in Removal Models 114
80
Data Anomalies APPENDIXK 115
80
COMPREHENSIVE
EXAMPLES Tests of Model Assumptions 115
81
A Taxicab Example APPENDIXL 121
81
A Penned Rabbit Study Density Estimation Based on Subgrids 121
84
An Example of Trap Response APPENDIXM 123
87
An Example There No Model Fits General Simulation Methods 123
93
COMPREHENSIVECOMPUTERALGORITHM APPENDIXN 123
96
SUMMARY Simulation Results 123
96 133
LITERATURECITED APPENDIXO
97
APPENDIXA Interval Estimation 133
102
Notes on Estimation
102
STATISTICAL INFERENCE FROM CAPTURE DATAtis et al. 7
INTRODUCTION studies (a removalstudy is, of course,

slightlydifferent).A typicalfieldexperi-
The estimationof animalabundance is lnentis the following:a numberof traps
an important problemin both the theo- arepositionedin the areato be studied,
reticaland appliedbiologicalsciences. say144trapsin a 12 x 12grid,7 m apart.
Seriousworkto developestimation meth- At the beginningof the study(j= 1) a
ods began during the 1950s, with a saluplesize of nl is takenfrolnthe pop-
few attemptsbeforethattime.The liter- ulation,theanimalsaremarkedortagged
ature on estimationlnethods has infor future identification,and then re-
creasedtremendously duringthe past25 turnedto the population,usuallyat the
years(Corlnack 1968,Seber1973). samepointwheretheyweretrapped.Af-
However,in largepart,theproblemre- terallowingtimeforthe markedan(lun-
mainsunsolved.Pasteffortstowardcom- luarkecl animalsto mix,a secondsalllple
prehensiveandsysteluaticestimationof (1= 2, oftenthe followingday)of 112 ani-
density(D) or populationsize (N) have lllalsis then taken.The secondsaluple
been inadequate, in general.Whilemore norluallycontainsboth Inarkedand un-
than200 papershavebeen publishedon luarkedaniluals.The unmarked anilllals
the subject,one is generallyleft without areluarkedandall capturedanilnalsare
a unifiedapproachto the estimationof releasedbackinto the population.This
abundanceof an animalpopulation. procedurecontinuesfort periodswhere
Thissituationis unfortunate becausea t ¢ 2. The animalsshouldbe Inarkedin
numberof pressingresearchproblems such a way that the capture-recapture
requiresuch information. In addition,a historyof eachanilualcaughtduringthe
wide arrayof environlnental assessment studyis known.Inpractice,toesareoften
studies and biological inventorypro- clippedto uniquelyidentifyindividual
gramsrequirethe estiluationof animal animals(Taberand Cowan1969)or se-
abundance.Theseneedshavebeen fur- riallynumberedtagsaresolnetilnesused
thereluphasizedby the requirement for on largeranimals.
the preparation of Environmental Impact Suchcapturestudiesare classifiedby
Statements iluposedby the NationalEn- 2 schelnesthat are directlyrelatedto
vironmental Protection Actin 1970. whatclassof luodelsareappropriate and
This publicationtreatsinferencepro- whatparameters can be estiluated.The
ceduresforcertaintypesof capturedata firstclassification addressesthesubjectof
on closed animalpopulations.This in- closure.Closureusuallymeansthe size
cludes lnultiplecapture-recapture stud- of the populationis constantoverthe pe-
ies (variouslycalled capture-lmark-re-riodof investigation, i.e., no recruitlnent
capture,luark-recapture, ortag-recapture(birthor ilumigration) or losses(deathor
studies) involving livetrappingtech- emigration). This is a strongassuluption
niquesandreluovalstudiesinvolvingkill and,of course,nevercompletelytruein
trapsoratleasttemporary removalofcap- a naturalbiologicalpopulation. Forgreat-
turedindividualsduringthe study.Ani- er generality,we defineclosureto lnean
malsdo notnecessarilyneedto be phys- thereareno unknownchangesto the ini-
icallytrapped;visualsightingsof marked tial population.In practice,this lneans
animalsand electrofishingstudies also knownlosses (trapdeath,or deliberate
producedata suitablefor the methods removals)do notviolateourdefinitionof
describedin this monograph. closure.Ifthestudyis properlydesigned,
To providea frameof referencefor closurecanbe lnetat leastapproximate-
whatfollows,we give an exampleof a ly. Open or nonclosedpopulationsex-
capture-recaptureexperimentto esti- plicitlyallowforoneorluoretypesof re-
luatepopulationsize of smallanimalsus- cruitlnentor lossesto operateduringthe
ing live traps.The generalfield experi- courseof the experiment (Jolly1965,Se-
lnentis similarforall capture-recaptureber 1965, Robson1969, Pollock1975).
Onlyclosedpopulationswill be consid- and analysisInethodswe do not cover

eredin this monograph. here. We do not treatsequentialsaIn-
The secondclassification dependson pling studies(e.g., Samuel1968),strati-
the type of datacollectedwith 2 possi- fied populations(e.g., Darroch1961,Ar-
bilities occurring(Pollock1974,unpub- nason 1973), Bayesianschemes (e.g.,
lisheddoctoraldissertation, CornellUni- Gaskelland George1972),or changein
versity)Ithaca,New York): ratioestimation(e.g.,PaulikandRobson
1969).The subjectof stratifying the data
(1) only information on the recoveryof afterthe fact on such variablesas sex,
narkedanimalsis availableforeach age,or speciesis notdiscussedprilmarily
samplingoccasion,j, j - 1, 2, . . ., t. because thererarelyareenoughdatafor
(2) information on bothmarkedandun- sucha stratification. The contingencyta-
markedanimalsis availableforeach ble approach to estimation frommultiple
samplingoccasion,j, j = 1, 2, . . ., t. capturestudiesis a promisingnew de-
In case (1), populationsize (N) is not velopment(see Fienberg1972),but cur-
identifiable,however,otherparametersrentlyit is relativelyunexploredor de-
can be estimated(Brownieet al. 1978). veloped;we donotdiscussit. Finally,we
In case (2), N can be estimatedusinga do not treatstudiesor analysisInethods
wide varietyof approachesdepending forwhichthe goalis to computeonlyan
uponwhatwe wishto assume.Onlycase index to abundance(e.g., capturesper
(2)will be dealtwithhere. 100trapnights);standard statistical
tech-
niques are adequatefor those types of
studies.
Objectives Althoughour objectiveis to present
comprehensive methodsof analysis,the
The objectivesof this publicationare scientistInustrealizethatno amountof
twofold: sophisticatedstatisticalanalysiscancom-
pensatefor poor study design or field
(1) to give a thoroughtreatmentof the technique(suchas hightraplosses).The
estimationof populationsize given experimenter can do farmoreto ensure
multiple capture occasions (t > 2) valid estimates by having a properly
assuming
plannedandconductedstudythanhe can
a. populationclosure, by sophisticated analysisafterthe exper-
b. there mayexist 3 lnajortypes of ilnent.Wehavethereforeincludeda sec-
variationin captureprobabilities; tionon statistical aspectsof studydesign.
(2) to extendand makeavailablea pro- Thatsectionincludescommentson how
cedurefor estiluatingdensity(num- dealwithanomalies
berof animalsperunitarea)fromgrid toThis publication is
suchastraplosses.
intended foruse by
trappingstudies. biologists.Sucha goalis difficultto attain
This monograph is specificallyorient- due to the generallytechnicalandInath-
ed to the commonlydone gridtrapping elnaticallycomplexnatureof the subject
and removalstudieswhereclosurecan matter.We have developeda compre-
reasonablybe assumed.Specifically,we hensive coluputerprogramto compute
do nottreatthe caseof 2 livetrapping oc- estimatesand test statisticsfor the var-
casions(t = 2). Thissubject(i.e.,the Pe- iousmethodscoveredin subsequentsec-
tersenor Lincolnestimatorsand variations (programCAPTURE).Biologists
tions thereoflis adequatelycoveredin whowishto analyzedataareurgedto use
the literature(see Seber1973).In fact,to the computerprograIn ratherthanto try
use the methodspresentedhereforanal- to coluputethevariousestimatesandtest
ysis of gridtrappingdatawe suggestthe statisticsby hand.Also mostof theInath-
studyhave5 ormoretrappingoccasions. ematicaland statisticaldetailsare con-
Therearesometypesof studydesigns tainedin appendixesto this monograph.
STATISTICAL FROMCAPTURE DATA4)tis
INFERENCE et al. 9
We hope this publicationand the asso- Overtonand Davis (1969:404):"Com-

ciatedcomputerprogramwill be useful puterswill soonproveofverygreatvalue
withinthe framework of the assumptions in the routineprocessingof censusand
considered. surveydata.Whenthey becomegener-
Weundertook the theorydevelopment ally availableit will be desirableto ad-
andthe writingof thisreportfora variety vanceto evenmorerealisticandcolaplex
of reasons.Severalimportantadvances solutionsto the problems;therewill be
havebeen Inadebutareavailableonlyas no premiulnon siluplicityso longas the
unpublished dissertations (Burnha1nusersunderstand the principlesand are
1972,unpublisheddoctoraldissertation,able to comprehend the constraints ancl
OregonStateUniversity,Corvallis,Ore- limitationsof the modelson whichthe
gon;Pollock).NewInethods haveempha- coluputersolutionsarebasecl."
sized nonpara1netric approaches thatare
robustto the failureof certainassump-
tions.Further,the use of a sequenceof Assumptions
statisticalluodelsseelusappropriate. It is estimationlnethodis basedon
unreasonable to expecta singlemethod a Every set of assumptions.The generalas-
to performwell on studiesof variousspe- sumptions the capture-recapture
cies in differenthabitats,orthesamespe- lnethodswefor present here are listedancl
cies at differenttilnes.Pollock(unpub- cliscussedbelow. The assumptiollslor
lisheddissertation) treated4 models,each the relllovalexperilnent aregivenin the
basedon specificassumptions, and sug- sectiononremovalstuclies. assump-
gesteda statistical testingsequence.That tionsare necessaryfortheFour luost restric-
generalstrategy,followedin this publi- tive experilnental situations:
cation,allowsmodels(assumptions) that
are inadequateto be rejectedfor a par- (1) the populationis closed
ticulardataset. A methodinappropriate(2) animalsdo not lose theirmarksdur-
forfieldmicePeromyscus spp.Inaywork ing the experiment
well forvoles LHicrotus spp- (3) all marksarecorrectlynotedandre-
Thereexistsa largebody of standard cordedateachtrapping occasion j and
statisticaltheorythatis directlyrelevant (4) eachanimalhasa constantandequal
andapplicableto theestiluation probleln probabilityof captureon each trap-
ill capture-recapture and reluovalstud- ping occasion.Thisalso impliesthat
ies. Biologistsneed not, however,learn captureandmarking do notaffectthe
the theoryto be ableto use the resultsof catchability of the animal.
these advancedmethocls.The methods Beforediscussingthe alsove we lilUSt
employedhereareoftenbeyondthe for- eluphasizethat the focal point of our
lualtrainingof luostbiologists,although workhas beell to relaxAssuluption4.
they shouldbe able to Inakeproperuse That assuluptioll is not Inetin Inostcap-
of the results.Westressthatwe haveex- ture-recapture studies an(la largeper-
aluined the estimationand inference centageof pasteffortshave beelldirectecl
problemsin a rigorousstatisticalfralne- at relaxingit. Assumptions 1-3 IllUSt lge
workas opposedto variousad hoc pro- luadeforall luodelsconsidered here.We
cedures. discussthe first3 andthen elal-
Anotherobjectiveof thisluonograph is briefly
orate on the lastill the followingsectioll.
to bring to the biologists'and statisti-
cians' attentionthe computerprograln (1) Populatiotl closure. This aSSUIllp-
writtento implelnentthe complexanal- tionarisesbecausepopulation estilllation
yses describedhere.AAIithout the aid of Inodelswere initiallyconceptualized as
a computer to do the calculations, devel- extensionsof urn models(Feller 1950).
opmelltof sophisticated analysesis just Such tnodelsare basicallyintendedto
an academicexercise.Ourphilosophyin providea "snapshot" of the population
this matterhas been sumlnedup by size at a givenpointin spaceandtilne.
In thatcontext,openandclosedmodels catchable,or nearlyso, is indistinguish-

becomeessentiallynoncompeting, since ablefromonethatdiesoremigrates.Pol-
open modelsare morefrequentlyused lock (1972,unpublishedmaster'sthesis,
for purposesof monitoringpopulations CornellUniversity,Ithaca,New York)
overa longerperiodof time andobtain- discusseda testformortality in somede-
ing information concerningsuchproper- tail. The tests for recruitment are more
ties as survivaland recruitment rates.If difficult.Thus,the biologistis forcedto
estimatesof populationsize at a given considercarefullythe design of such
time are also desired,however,compe- studiesin an effortto assurethatthe clo-
titionbetweenthe2 typesof modelsdoes sureassumption is met.Finally,we note
arise. In general,open modelsrequire that the tests for closureimplicitlyas-
moredatathanclosedmodelsdueto the sumeequalcaptureprobabilities; there-
factthatassumptionsare morerigorous fore,such tests can rejectclosurewhen
and more parametersare involved. in factclosureis truebut equalcapture
Therefore,feasibilityoftenprohibitsthe probabilityis false.This greatlylessens
use of verygeneralstochasticmodelsfor the value and powerof such tests. We
estimatingpopulationsize of open pop- believe closurewill haveto be assessed
ulations(Jolly1965;Seber1965;Robson largelyfroma biologicalbasisratherthan
1969;Arnason1972a,1972b,1973;Pol- fromanydefinitivestatisticaltests.
lock 1975).If, forexample,a 10-dayex- Theclosureassumption canbe relaxed
perimentis considered,17 basicparam- in somecases.Seber(1973:70-71) showed
eters wouldhaveto be estimatedusing thatnaturalmortality will notbias some
Jolly's(1965)model.Hence, datafrom estimatorsif it acts equallyon marked
onanypopulationestimationexperiments and unmarkedsegmentsof the popula-
are inadequatefor obtainingestimates tion. In such cases,the populationesti-
withacceptableprecisionandsmallbias matethenrelatesto the size of the pop-
using models for open populations. ulationat the beginningof the study.
Moreover,unlike the models treated However,if recruitmentand luortality
here, none of those open population occurduringthe experiment,the esti-
modelsallowsforunequalcaptureprob- mateof N will be too high,on the aver-
abilitiesof individualanimals.Let it be age, forbothinitialandfinalpopulation
clear, we believe that well-developed, size (RobsonandRegier1968).
general models for capturedata from
open populationsare essentialin some (2) Permanency of marks. Lossof luarks
studies.However,we also believe that (tags)violatestheclosureassumption and
formanypopulations of interest,the clo- will resultin anoverestimate of N. If the
sureassumption canbe metapproximate-studyis of shortduration(tohelp assure
ly and the models discussed in this the closureassumption),it seems that
monograph will be useful.Forexample, loss of markswill generallybe a minor
closuremightbe assumedfor an 8-day problem.Someexceptions,suchas radio-
studyof cottontailsSylvilagus spp. dur- activeisotopeswitha veryshorthalf-life,
ing a nonbreedingperiodin a well-de- undoubtedlyoccur(cf. Seber 1973:93-
fined(sampled)area. 100).
A numberoftestsforclosurehavebeen
derived(RobsonandFlick1965,Robson (3) Reporting and recording marks
andRegier1968,Pollocket al. 1974),but (tags).-This assumptioncan be easily
they generallyhave little chanceof re- assuredby workingcarefully.Field re-
jectingclosureunlessthe sampleis large portsand keypunchedcardsshouldbe
andthereis a markeddeparture fromclo- editedandverified.Often,a pilot study
sure.In addition,closuretests areoften maybe beneficialto trainpersonneland
confoundedwithbehavioralresponseto identifyanyproblemswith the marking
capture,e.g.,ananimalthatbecomesun- lnethod.
STATISTICAL FROMCAPTUREDATAtiS
INFERENCE et al. ll
UnequalCaptureProbabilities casion).This situationmaybe common

The fourthassumptionis particularlyeven thoughthe numberof trapsmight
iluportantand,forthis reason,we focus be fixedduringthe courseof the study.
on it here. It is now widely recognized Forexample,a coldrainydayduringthe
thatthisassumption is commonly notInet studymightreduceactivityof the ani-
(e.g.,Younget al. 1952,Geis 1955a,Hub- malsand reducethe probabilityof cap-
er 1962,Swinebroad 1964).Edwardsand ture.Also, if differentcapturemethods
Eberhardt (1967),Nixonet al.(1967),and are used on each occasion,this model
Carothers(1973a)providedclear evi- couldbe appropriate.
dence that accuratepopulationestima- ModelMb allowscaptureprobabilities
tionusuallywill requiremodelsthatpro- to varyby behavioralresponseor "cap-
vide forunequalprobabilities of capture. turehistory,"anddealswithsituationsin
The effects of unequalcaptureproba- whichanimalsbecolnetraphappyortrap
bilitiesonestimatesderivedfrolnmodels shy.Carothers (1973a)referredto thisas
that assulne equal catchabilitieshave a contagion of catchability.
This iluplies
been studiedby computersimulationby that an anilnal's behavior tends to be al-
Burnhamand Overton(1969), Manly tered afterits initialcapture(e.g., per-
(1970), Gilbert (1973), and Carothers hapsthe anilnalwas frightenedor hurt
(1973b).Estiluators studiedwere gener- duringinitialcaptureand markingand
ally found to be significantlybiased thereafter it will notlikelyenteranother
whenthis assumption wasviolated. trap).
This luonograph presentsa nuluberof Model Mh allowscaptureprobabilities
modelsandestimators developedto relax to varyby individualanimal.This situa-
the criticalassumptionof equal catch- tion has been modeledonly with great
ability.We havedrawnheavilyfrolnthe difficultyandrequiresthatadditional dis-
workof Pollock(unpublisheddisserta- tributionalassumptionsbe lnade.Indi-
tion,pers.comm.)andBurnham (unpub- vidualheterogeneity of capturemayarise
lished dissertation).FollowingPollock in many ways. Perhaps accessibilityto
(unpublisheddissertation), we consider traps(as influencedby individualholne
a sequenceof modelseach allowingfor ranges),socialdoluinance, or differences
differentcolubinations of up to 3 typesof in age or sex cancausesuchan unequal
unequalcaptureprobabilities: probabilitystructure.This is an ilnpor-
tanttype of variationand has been rig-
(1) captureprobabilitiesvarywith time orously treated by Burnhaln(unpub-
ortrappingoccasion ModelMt, lisheddissertation), whosenonparametric
(2) captureprobabilities varydue to be- approach is presented in a latersection.
havioralresponses ModelMb, In additionto these 3 simplemodels,
(3) captureprobabilities varyby individ- we considerallpossibilecombinations of
ual animal Model Mh(h = hetero- the 3 typesof unequalcaptureprobabil-
geneityamonganimals). ities (i.e ., ModeIs Mtb, Mth, Mbh, and
Mtbh). We also treatthe "null"case in
Theassumptionsregarding unequalcap- which captureprobabilityis constant
tureprobabilities areto be explicitlyem- with respectto all factors(ModelMo)
bodied in probabilitymodels that de- ModelMocorresponds to the 4 assulnp-
scribecapturestudies. tionslistedearlier.Forsimplicity,we de-
We agreewith Carothers(1973b:146) note estilnatorsof populationsize for a
thatequalcatchability is an unattainable specificlnodelusingthe samesubscript
ideal in naturalpopulations(cf. Seber notation.ForexampleNodenotesthe es-
1973:81-84).We discussthe 3 simplest timatorderivedfromModelMo;Nt de-
waysto relaxthisassumption. notesthe estimatorderivedfromModel
ModelMtallowscaptureprobabilities Mt;Nbhdenotesthe estimatorderived
to varyby time (e.g., each trappingoc- frolnModelMbh, andso on.
MONOGRAPHS
WILDLIFE
12
Webelieverigorous probabilit,vmodels
Perspectives
incorporating various tentative
Wewish to emphasizethata specific explicitly assumptions representthe best approach
of assumptions
set is the basisfor a spe-
andmodel toward estimating populationsize N, or
model.The assumptions
cific D. Thetentativenatureof the as-
thenrepresenta tentative hypothesis density and the generaluncertainty
when analyzingtheresultsof a particular suluptionsbiologicalprocessesInaketesting
capture experimentconductedto esti- about Seber(1973)pointed
matepopulation sizeordensity.Cormack akeyconcern.As models shouldbe used
(1968:456)stated,"Inall caseseveryiota statistical
out,
caution,due to lack of controlover
information,
of bothbiologicalandstatis- with populations. All models depend
tical,lnust be gatheredto check and natural the validity of various underlying as-
countercheck the unavoidableassulup- on that are often difficult to
tions."Statisticaltestingwithinandbe- suluptions rigorously.
tweenInodels(assumptions) is empha- evaluate
here.In spiteof this,more work in Finally, we believethattheoryandap-
sized must be integrated.Eitherin
thisdirectionis clearlyindicated.Our plication absenceof the otherwill stifleprog-
approach is to derivemodelsforan array the Forthisreasonwe havetriedto in-
oftypesof unequalprobabilities of cap- ress.
ture.Weconductedstatisticalteststo en- tegrate the statisticaltheorywiththe bi-
modelfor ological application. We havevhowever,
ableselectionofanappropriate triedto separate the luorecomplexsub-
theanalysisof a particular dataset (cf.
Pollock unpublisheddissertation). Some jects andincludethenaasa seriesof tech-
Inodels areverysensitiveto smalldepar- nical appendixes.We urgebiologiststo
turesfrolnthe underlyingassumptions; tryto considerand understandthe ap-
andwe askstatisticians to con-
therefore, testing between luodelsand pendixes, to be concerned with the biological
investigating the robustnessof each es- tinue and realities before at-
timator areessential. complications
of such testingis re- tempting additional theorydeveloplnent.
The ilnportance Through an integrated team approach
inade-
flectedin the factthatuse of an highly on this
wecan expectfurther progress
quatemodelwill oftenlead to a problems.
biasedestilnateof populationsize. This seriesof estimation
isperhapsto be expected,if notobvious. Comments on the Useof
Moresubtleis thatestimatesof the sam- This Honogra ph
plingvariance(a measureof precision) Wecoverseveraltopicshere,andpre-
arequitedependenton the correctmod- as well as appliedre-
el. Biasofthe estiluator onaybe small,but sentmatheluatical includedataanal-
Topics covered
the estilllateof variancemay be very sults. of short-terlnlivetrapping and
poor,even with largesamples.This can ysis effortreluovalstudies,designof
cause,forinstance,associatedconfidence constant
such live trappingstudies,and simula-
intervalsto have very poor properties. Nu-
Theiluportance of assuluptions andtheir tionresultson inferenceprocedures. A vari-
Pau- merousexamplesarealso given.
testingcannotbe overeInphasized. ety of uses of this luonographare
lik (1963)noted that an approxiluately
correctestiInatewithlow precisionis al- anticipated by: (1) biologistswho must
waysbetterthana highlypreciseincor- analyze faced actual data,(2) biologists(and
rectestimate.Testsof assuInptions con- statisticians) withdesigningcapture
cerniIlgequal captureprobabilitiesare studies, (3) personsinterestedin perfor-
becauseestimators uanceof estiluatorspresented here, (4)
especiallyiluportant are statisticians interested in developing
basedon given sets of assuluptions advancedmodels,and(5)educators
usually not robustto departuresfrom more seekto teachcourseson the subject
those assumptions(Seber1970,Gilbert who
1973). of populationsize estimation.
FROMCAPTUREDATA-4OtiSet al.
STATISTICALINFERENCE 13
Biologistswho have datafromclosed will need to readmostsections,excep-

population livetrapping studieswill have tionsbeing HISTORICAL OVERVIEW, RE-
to readquitea bit of thismonograph be- MOVALMODELS,and materialfollowing
fore they can understandthe methods. the STUDYDESIGNsection.
Theydo notneedto readtheappendixes. If you are interestedin obtainingin-
Theywouldhaveto understand all sec- sightsintotheperforluance of variouses-
tionsthroughTESTS OF MODELASSUMP- timators,you shouldput special effort
TIONS, except REMOVAL MODELS. We into studyingthe numeroussilnulation
believethiscanbe doneby anyonehav- resultspresentedhere. This would re-
ing had a solid coursein college level quirereadingalluostallthetextandcare-
algebraandbeginningstatistics.In order ful studyof AppendixN
to understandthe essence of what we Personsinterestedin doingfurtherre-
present,the readerdoes not haveto fol- searchalongthe lines of the modelsand
low all the mathematical descriptionsof approach of this monograph will haveto
luodelsnor discussionsof Inodelprop- carefullystudyalmosteverythinghere,
erties.We have includednulnerousex- especiallythe appendixes.
alnples.In particular, the readershould Finally,this Inonograph and prograln
benefitgreatly frolnthe section on Co- CAPTURE have value for teaching and
PREHENSIVEEXAMPLES. learningaboutpopulationsize estilna-
If the readerintendsto do, say, data tion. The simulationfeature of CAP-
analysisaccordingto thesemethods,it is TURE can be especially valuable in
virtuallynecessaryto use progralnCAP- teachingthe conceptsof salnplingvaria-
TURE(see COMPREHENSIVECOMPUTER tionandproperties of estimators.Persons
ALGORITHM).This prograln is available interestedin performing suchsimulation
and thereis a user'snlanualfor it. Per- of the lnethodspresentedhere(eitherfor
sons with many data sets to be ana- design of studies, evaluationof esti-
lyzed should get the program.Con- lnators? or teachingpurposes)will need
versely,we do not recommend tryingto to iluplelnentthe progralll fortheirown
implementthiscomputerprogram if one use.
has only a few (orone) datasets to ana-
lyze.In thislattercase,it is betterto have ACKNOWLEDGMENTS
thedatarunforyou.Theauthorsarewill-
ing to assistin runningsuchdataprovid- Dr.K.H. Pollock,Ulliversityof Rea(l-
ed the userarrangeshis own keypunch- ing,providedseveralideasandcriticislns
ing of the datain the necessaryformat throughcorrespondellce; in addition,we
(we cansupplythisformat). havedrawnon the resultsof his doctoral
If one'sgoalis to analyzesomereluoval prograln. Drs.M.H. SmithandJ. B. Gen-
data,the relevantsectionsare those on try,Savannah RiverEcologyLaboratory,
ModelsMb,Mbh, andthereluovalmodels contributed to this workthrough(liscus-
(plus the introductory sections).Again, sion and providedsallaplesof their re-
the authorswouldtryto help usersana- searchto be used for progralntesting.
lyzerelnovaldata;withinreasonable lilll- Datato be used as exalnpleswere pro-
its we Inaybe ableto runthe dataanal- vided by A. D. Carothers,H. N. Cou-
yses or assistin settingthelnup. lolnbe,C. T. Cushwa,W.R. Eclwards, S.
Manyreaderswill sometimesbe faced W.Hoffinan,E. C. Larsen,R. F. Raleigh
withdesigninga capturestudy.The sec- andV. H. Reid.
tionOI1 STUDY DESIGN coverssolllefun- We appreciateextensive comments
damentaldesignaspectsof livetrapping nladeby A. N. Arnasonv A. D. Carothers,
studies for closed populations.If your W. R. Clark,S. W. Hoffinan,and R. P.
goalis to designa study,readthatsection Davisonon an earlierdraftof the manu-
at a minilnum;to get full advantageof script.The colulnentsby A. N. Arnason
this Inonograph in termsof design,you andG.A. F. Seberon the finalversionof
14 WILDLIFE NIONOGRAPHS
the manuscriptare also muchappreci- X1 1 X12 , . . Xlt

ated. [Xij]= X21 X22 ... X2t
Dr.R. G. Streeter,U.S.FishandWild- * . .
life Service, providedencouragement _ XN1 XN2 XNt _

duringthe investigations.
This workwas performed underCon- where
tract 14-16-0008-1224 of the Coal Pro- 1 if the ithanimalis caughton the
gram,Officeof BiologicalServices,U.S. xij = jth occasion
Fish and WildlifeService,to the Utah Ootherwise.
CooperativeWildlife Research Unit.
Fundsforthisworkweremadeavailable TheX Inatrix is a simplewayto record
to the Fish andWildlifeServiceas part the captureornoncapture of eachanimal
of the FederalInteragencyEnergy/En- in the populationon eachtrappingocca-
vironmentResearchand Development sion.Rowi givesthe trappingresultsfor
Program, Officeof Researchand Devel- individuali, whilecolumnj givesthe re-
oponent,U.S. Environmental Protection sults of the jth trappingoccasion.Note
Agency (IAG-EPA-D5-E385). Publica- thatthe luatrix X maynotbe observedin
tioncostswerepaidby theOilShalePro- its entiretybecausesolne animalsmay
gram,Officeof BiologicalServices,U.S. neverbe captured; therefore,thoserows
FishandWildlifeService,contract14-16- ofX areall zeros.
0008-1197.Finaldebuggingof the com- A seriesof specificmodelsforcapture
puterprograln, andpreparation of exam- datacanbe derivedif we definethe fol-
ples wereperformed underthe auspices lowinggeneralstructural model:
of the U.S.EnergyResearchandDevel- Pij= the captureprobabilityof the ith
opmentAdministration. individualin the populationon
the jth trappingoccasion,wherei
FUNDAMENTALCONCEPTS = 1,2, . . ., N, andj = 1,2, . . ., t.
Thissectionpresentsnotationanddis- For example,if we assumethe restric-
cussesthe statisticaltechniquesused in tionsPii= p forall i andj we get Nlodel
this monograph.The subjectmatteris Mo)the simplestpossibleInodel.All oth-
basicallytechnicalin nature,butwe have er Inodelswe introducemaybe thought
triedto keepthepresentation simpleand of as generalizations of Nlodel M0.
referthe readerto appendixesfor more Hence,in the followingsections,models
coluplexdetails.Webelieve it is impor- aredevelopedbaseduponcaptureprob-
tantforusersof themethodsdescribedin abilitiesbeingtimespecific,behaviorally
this publicationto understand the luate- related,ordifferingamongindividualan-
rialpresentedin this section. imals. Therefore,captureprobabilities
are the crucialelementof the series of
Data and Parameters modelswe discuss.
The abovestructureand assumptions
All the modelsdiscussedhereassulne sufficeto specifythe marginaldistribu-
populationclosure(exceptforknownre- tion of eachindividualXij(i.e., they are
lnovals).Therefore,the parameterwe Bernoullirandomvariables);however
wish to estimateis populationsize N theydo notspecifythejointdistribution
whichis constant.Nloreover, becausethe of all Xij.Therefore,we have assumed
salne individualanilualscomposethe joint independenceof the variablesin
population oneachtrappingoccasion, j = orderto havea completelyspecifiedgen-
1, 2, . . ., t, we canconceiveoftheindivid- erallmodelstructure. Specifically, we as-
uals as being numberedi = 1,2, . . ., N. sumethatgiventhe correctmodel(i.e.,
The basiccapturedataareconvenient- the correctspecification of captureprob-
ly expressedin Inatrixforlnas abilitiesPij),thenthe elelnentsof Xijare
FROMCAPTUREDATAtiS
INFERENCE
STATISTICAL et al. 15
mutuallyindependentrandomvariables. 2, ...,t.Notethatuj=nj-mjand
Thisassumptionis nottestableunlessone thatm1= 0,
first knows what the correctmodel is t
(whichwe neverwill forrealdata).It is m. = sumof the mj= E mj.

our opinionthis is not a restrictiveas- J=1
sumptionandit need not be a sourceof

concern. ThestatisticsUj,fj,Mj,andmjmayalso
be computeddirectlyfromtheX matrix.
Statistics and Notation However, the computationis not as
straightforward asthatofnjandis notgiv-
Probabilitymodels fromwhich esti- en here.WedenoteX<,as the numberof
matorsof populationsize N maybe de- animalswitha specificcapturehistoryco.
velopedare discussedin followingsec- For exampleXl00llrepresentsthose in-
tions.A few simplestatisticsareneeded dividualscaughton trappingoccasions1,
for these models.They are definedand 4, and5.Thesetofallthepossiblecapture
discussedbelow. historieswill be symbolizedas {X(O}. In
general,witht captureoccasionsthereare
nj= the numberof animalscaptured 2tpossiblecapturehistories.
in thejth sample,j = 1, 2, . . ., t, Twoothertermsusedfrequently in this
N
monograph are:
=xiix Robustness (of an estimator).A robust
i=l
n. = the totalnumberof capturesdur- estimator is onethatis notsensitivetothe

t breakdown of a particularassumption. A
ingthe study= E nj, specificmeasureof robustness is difficult
j=l to define.Therefore, a somewhatsubjec-
u;= the numberof new (unmarked) tive determination regardingthe robust-
animalscapturedin the jth sam- ness of an estimatoris maderelativeto
ple,j = 1, 2, . . ., t, thegeneralperformance of theestimator.
fj= the captures frequencies= the Performance is evaluatedwithrespectto
numberof individualscaptured the essentialcriteriaof bias, precision,
exactlyj times in t daysof trap- andconfidenceintervalcoverage.Forex-
ping,j = 1, 2, . . ., t. fOwillbe used ample, the estimatordevelopedunder
for the numberof individuals ModelMtperformsverypoorlywithre-
never captured(obviously,fOis spectto all criteriaif individualhetero-
notobservable). geneityto captureis presentin the pop-
Mt+l= thenumberofdistinctindividuals ulation.We saythatthis estimatoris not
caughtduringtheexperiment (re- robustto a particular assumption. In con-
call that t is fixed for a given trast,thejackknife estimator forModelMh
experiment), appearsto be fairlyrobustfora nuluber
t t of specificassumptions.
=fj = Euj,
j=l j=l Bias (ofanestimator).Biasis the differ-
Mj= the numberof markedanimals ence betweenthe expectedvalueof an
in the populationat the time of estimatorand the trueparameter being
thejth sample,j = 2, 3, . . ., t. (Note estimated, e.g., B = E(N) - N. Percent
thatM1--°) relativebias, 100{[E(N)- N]/N} is de-
M.= sumof the Mj[doesnot include notedas RB. Overtonand Davis(1969)
t gavea gooddiscussionof theseandother
Mt+l]= E Mj, relatedterms.
j=l A final note concernsthe differences
mj= the numberof markedanimals betweenparameters (truevalues)andes-
capturedin the jth sample,j= timates.We are concernedchieflywith
MONOGRAPHS
WILDLIFE
16
estimatesoftheparameters
making N and usingthe likelihoodfunctionde-
culus,
size and density,respec- fromtheprobability
rived model.In some
population
D, the estimatormaytakea simple,
cases,
We denoteoursampleestimators
tively. to use form.For example,the ML
parameters
ofthese asN andD. Biologists easy
referredto
are Kendall and Buckland of N forModelMtfor2 sample
estimator
for
(1971) definitionsof standard statisti- (t = 2) is thePetersenestimator
occasions
terms.
cal Nt = nl n2
m2
Parameter Estimation
orre- where nl, n2,andm2havealreadybeen
Thedatafromcapture-recapture defined.
moval studiesaresamples. Thisimposes However,in capture-recapture models
theneed fora probabilistic treatmentof werarelyfindthatthe exactMLestima-
thedatato derivecorrectestimationand tors existasa simpleformula asabove.To
inference procedures.The models we illustrate this,consider the model devel-
considerhere are termed stochastic opedby Darroch (1958) when 4 sampling
models. Unlikethemodelsforopenpop- occasions areconsidered(t= 4) andthe
ulations,the only stochastic component probabilities areassumedto vary
formodelsunderpopulationclosure recapture P2, p3, and p+).
onlyby time (i.e., P1,
tothesamplingprocess:i.e.,thecap- Theapproximate
lates ML estimatorof N for
Modelformulation in
tureprobabilities.
with a set of explicit thismodel (see Darroch1958) is the
thiscontext begins solutionof the equation
assumptions. A probability modelforthe unique
sampling distributionof theX matrix(the
basicdata)is derivedto quantitatively (1_ Ms )= (1_ nt)(l_n2)
expressthe assumptions.A probability rep-
function is a formof mathematical
resentation of the observed data under a
It provides a ( N) ( N)
specificset of assumptions.
basisforquantitatively andexplicitlyin-
corporating the specific assumptions Ingeneral,for ModelMtthe ML esti-
aboutcaptureprobabilitiesand for de- matoris the solutionof the equation
velopingthepointandintervalestiInators
by rigorousstatisticalestimationtech- N ) [I ( N)
niques. (
Mostparameter estimators in this pub-
licationwere derivedusingthe method For t greaterthan2, thisequationcannot
be solved algebraically for N. In other
of lnaximumlikelihood(ML). Several esti- words, it is not possible to arrangethe
models and their corresponding algebraically in such a waythat
lnatorsweretakenfromexistingliterature symbols N appears on one side ofthe equation
(e.g., Zippin1956,Darroch1958),often only all other terms appear on the other
with some modification.Othermodels and side. The equation can be solved, butonly
and estimatorswere derivedduring the
on a caseby casebasis using a numerical
courseof this study. are procedure. Wesaythe equationdoesnot
Estimators found by the ML method
a simple, "closedform"solution.
optimal,at leastforlargesamples.(Fora have Complex probability modelsoftendo not
discussionof optimality,referto Appen-
dixA).Thisis a generallyacceptedtenet have simple estimatorsand tests of as-
of statisticalestimationtheory(Moodet sumptions; nonetheless,complexmodels
of appear necessary to describeluanycap-
al. 1974).In general,ML estimators found ture-recapture studies adequately.
unknownparameters (e.g., N) are thatseveralof the
by application of resultsfrom siInple cal- Our work has shown
FROMCAPTUREDATAtiS
INFERENCE
approximations of N suggestedfor this IntervalEstimation

model in the past are fairlypoor.Fur-
thermore,someof the iterativesolutions One of the severaladvantagesof the
given (e.g., Darroch1958)produceonly probabilitymodel/MLapproachis that
approximate MLestimates.Wehaveob- estimatesof saluplingvarianceand co-
tained exact ML estimatorsfor all the variancescanbe computedas partof the
modelsinthispublication (exceptModels MLmethod.Thesemeasuresofprecision
Mh,Mtb,Mth,andMbth) by employingnu- are essentialin makinginferencesfroln
mericalprocedures on a digitalcomputer thesampleresultsoftheexperiment. The
(in factno estimators canbe derivedfor varianceand covarianceestimatorsare
the latter3 models).We havefoundthe derivedfrom"largesample''theoryand
uaxilnumof the likelihoodfunctionin usuallyareofunknown valueasmeasures
sucha wayas to obtainexactintegerval- of precisionin ''smaller''samples.We
ued MLestimatorsof N. The disadvan- haveperformed a largenuluberof Monte
tage here, of course,is that we cannot Carlo silnulation experiments (Appendix-
showsimpleclosedformestimators. This es M and N) to examine the sluall sample
subjectis discussedfurtherin Appendix properties ofsuchvariance estimators and
A. the confidence intervals that depend on
We findthatthe estimatorof N under theln. Intervalestimation is an old subject
eachmodelinvolvesonlysimplestatistics in the statisticalliterature,andwe refer
coluputedfromthe X luatrix.Individual the interestedreaderto thetextby Mood
capturesarenotemployed onlyvarious et al. (1974)fordetails.Seber(1973)also
sums(linearcombinations) derivedfrom gavenulnerousexamples.
the X matrix.Those sumsare statistics Typically,the MLestimator of N is not
suchas nj,n., u;,andMt+,.Foranymodel norlually distributed unlesslargesaluples
we consider,thereexistsa set of simple are taken.Becauseconfidenceintervals
statistics,calledminimalsufficientstatis- commonly useddependuponanassump-
tics(MSS). Estimators shouldbe basedon tion of we exploredalternative
norluality,
only MSS. The use of the ML method intervalestimation techniques(Appendix
resultsin estimators thatarealwaysfunc- O).The alternative procedures were not
tionsof the MSS. Thisis a desiredprop- totally satisfactory and we will use the
ertybecauseit canbe shownthattheMSS standard procedure asfollows to construct
containsall the inforluationavailable anapproximate 95 percentconfidencein-
frolnthe experilnentfor estiluatingthe tervalon N:
paralneter(s) of interest(in ourcase N).
Anestimator basedonstatisticsotherthan N + 1.96jVar(N).
MSSis notusingallavailableinforlnation
andis, therefore,not optimal.Solnesta- Thisprocedure hasits limitations but,all
tisticaltests of assumptions will depend thingsconsidered,appearsto be best at
on information otherthanMSS. present.
The numberof parameters thatcanbe
identified(estimated) is lessthanorequal HISTORICALOVERVIEW
to the numberof elementsin the MSS
(regardlessof the estimationlnethod Althoughthe basicconceptof obtain-
used).The subjectof"identifiability" of ing inforluation aboutan animalpopula-
parameters is importantin the material tionby Inarking someof its melnbersmay
thatfollowsand,therefore, we placesome be tracedas farbackas the 17thcentury
emphasison MSS.For example,under (Chapman 1948)andto Petersen's(1896)
ModelMoxtheMSS is n. andMt+l,where- expressionof the fundamental principle,
as underModelMh,the MSSis fj,j = 1, one mayarguethatthe practicalbegin-
2, . . ., t. We makefrequentuse of the nings of the literatureconcerningthe
MSS in the followingsections. Inarkingmethodcan be associatedwith
MONOGRAPHS
WILDLIFE sampling,
18 as random
such
use of band returns to assumptions,the validityof the method.
(1930)
Lincoln's influence
that
the size of the North
estimate
American
50 considerations
Such wereat leasthinted
population. In the nearly in Schnabel's(1938) paper
waterfowl (1930) initial work, however,
at, to the situation
sinceLincoln's
years method
that the of the population
extended
avoluminous literature hasresultedfrom inwhich members
directedtowardderiving and re- andreleasedbackintothe
efforts the capture- marked
were
techniques
fining based on
decade, 2 populationon more than 1 occasion. rela-
recapturemethod. In the past is madeof the percentage
the exist- Mention an experi-
attemptsto summarize
notable made. The first biasof the estimatorsin size is
tive
literature
ing have been in which the population
ment
provided a sur- that
isCormack
by (1968) who
proposed for andthe readeris cautioned
known, expected to
veyofmathematical models of the solutions
"none can be
experiments. ofthe gen-
in capture-recapture
use text by Seber morethananestimatetotalpop-
provide
second
The is an extensive
to bring together all orderof magnitudeof the Presum-
eral
thatattempted for estimating
(1973) ulation"(Schnabel1938:352). by the
proposedtechniques
the and relatedpa- ably, cautionis generated
some of randomsam-
abundance
population as a subsetof those that "assumptions
fact
Included and constantpopulation are only
rameters. concerned with the pling to the actual situ-
techniques arethose roughapproximations Moreconsid-
method. In the pre- (Schnabel 1938:352).
capture-recapture substan- ation" of the cap-
sentation of thosetechniques,adetail is wasgivento the uses
eration sequence
a1nountof mathematical technique in a
tial numerical examples. ture-recapture (1933, 1937, 1939,
provided, as are mustbe ofpapers by Jackson with
assumptions
Furthermore, ofa particular that whowas concerned not only
tech- 1940), also with
metto ensurevalidity for testing estimating populationsize but
nique arepresented; methods anddeath-emigration
validity of some of the assumptions birth-immigration Contained in the considera-
the parameters. is the concept
aregiven. and qualityof tionof those parameters "closed,"i.e.,
Because of the existence thatthe population is not
is made here
thecitedworks,no attempt associated population size is not constantthrough-
lnethods
topresentspecific methods the samplingperiod.Those
of capture-recapture and out applied mainly to fish
with the theory we present were then being
related experiments. Rather,
and insectpopulations andnot terres-
to
areviewthatfollows the chronological
trialwildlifepopulations, althoughsome
development of conceptual approaches didexist(e.g., Green andEv-
exceptions the worth of
inthe literature. of the artis well char- ans1940).Scepticismas to wildlife pop-
Theinitialstate statement themethod withrespectto
acterized by Lincoln's (1930:2)
estima- ulations was expressed by Dice
ofthe solutionto his population
accurate (1941:402), whostatedthat"theapplica-
tionproblem: "Given a fairly
tion of the proportional snethodof cal-
showingthe number of wild may
statement in any one culating luammalianpopulations as the cozn-
America
duckskilledin North numberof ducks oftenrequireas mucheffort whole
season,then the total forthatseason pletetrappingor countingof the effort
continent
presenton the by a percentagecom- samplepopulation." Nonetheless,
theory.
luaybe estimated therelationthatthe continuedin developmentof the (1943) pro-
putation, basedupon killed dur- Schumacher and Eschmeyer that of
banded ducks solution to
totalnumberof band carriers vided an alternative
of regression
ing their first season as banded."As Schnabel(1938)by the use their work was
bearsto the total number of techniques. Evidently,
no mention is made desireto develop an es-
onemightexpect, such an esti- spurredby the robust to de-
the statistical properties timator that wouldbe more
underlying
matormightpossessorof the
FROMCAPTUREDATAtiS
INFERENCE
parturesfrolnthe underlyingassump- theoryforestimatingsuchparalneters as

tionsof the Schnabelmethod.Moreover, deathrateand populationsize, and de-
unlikeSchnabel,they providedan esti- votedmucheffortto the examination of
uatorfor the standarderrorof the esti- assumptions.
uate.Siluilarregression techniqueswere The appearance of suchmathematical
also investigatedby Hayne(1949a)and treatments generatedInostof the ilupor-
DeLury(1958). tantimlnediatelysucceedingworkonthe
A significant changein boththequality developlnentof the theory.As an exam-
andquantityof workin the fieldof cap- ple, one canconsiderthe workof Zippin
ture-recapture theoryoccurredwith the (1956),who provideda morecomplete
appearanceof severalimportantpapers statisticaltreatment of the removalmeth-
of the early 1950s. Those paperssig- od firstsuggestedby Moran(1951).An
naledthe beginningof a morerigorous ilnportant exampleis theworkof Darroch
Inathematical treatmentof the theoryin (1958),who was responsibleforthe cor-
termsof both estimationand testingof rect derivationof the probabilitymodel
assumptions. Bailey(1951),forexample, forthe multiplecapture-recapture exper-
proposeda binomialInodelforthe single iment firsttreatedby Schnabel(1938).
mark-release situation.He usedthe ML Moreover,Darrochpresentedexpres-
theoryto developanestimator of thepre- sionsforthe asymptotic biasandapprox-
cision of the populationsize estimator, imatevariancefor his approxiluate ML
the latterestimatorbeing the salne as estimatoranda methodforconstructing
Lincoln's(1930).Chapman(1952)con- confidenceintervals. Darroch's(1958,
sidereda hypergeometric modelforthe 1959)workontheclosedmodel,thebirth
multiplecapture-recapture experiment only and deathonly modelsstandsas a
and derivedan approximate expression cornerstonein the developmentof the
for the resultingMLestimator.He also theory.
gave a test fordeterminingwhetherthe Given the methodsavailable,it was
probabilityof captureis independentof nowpossibleforresearchers to directef-
tagging.Alternativesamplingschemes forttowardthe development of statistical
wereproposedby suchauthorsas Chap- tests of assumptionsunderlying the
uan (1952,1954)and Goodman(1953). Inethodsof estimation.One of the most
Such schemeswere designedto avoid generallyinvokedassumptionsof pro-
undesirablestatisticalpropertiesassoci- posed estimationtechniqueswas (and
ated with the direct samplingmethod stillis) thatall anilnalsin the population,
that considersthe total numberof ani- regardlessof capturehistoryand other
malscaughton each occasionas a fixed individualcharacteristics, are equallyat
paralneter. Forinstance,Chapman (1952) riskto captureon eachtrapping occasion.
pointedout thatan estimatorof popula- Leslie (1958)deviseda test directedto-
tion size obtainedvia inversesampling wardthathypothesis,whichwaslaterex-
(i.e., consideringthe numberof marked tended by Carothers(1971). Corluack
anilualscaughton eachoccasionas fixed) (1966)Inadethe iluportant pointthatfail-
is unbiased,whereasthe estimatorasso- ure of the above assumptionInaybe
ciatedwithdirectsamplingis biased.In causedeitherby eachanimalin the pop-
addition,removaldata,similartothetype ulationpossessingan "innatecatchabil-
of datausedin marking experiments,was ity"whichvariesamongindividualsover
usedin alternative methodsproposedby the population,or by an individual's
Moran(1951)andDeLury(1951).Alsoat probabilityof capturebeing affectedby
thattime,progresswasmadein the the- its capturehistory,or both. Cormack
ory of estimationin open populations (1966)provideda test forthe formeras-
througha sequenceof papersby Leslie sumingthe latteris false. Seber(1962,
andChitty(1951),Leslie(1952),andLes- 1965)andRobsonandYoungs(1971)con-
lie et al. (1953).Thoseauthorsused ML sideredthe problemof testingwhether
markingan animalaffectsits probability tiorzclosurewas reasonable,

of captureon subsequenttrappingocca- homogeneous and non-
sions,andManly(1971)provideda meth- bilities were caused by individual capture proba-
od forestimatingtheeffectof marking the sampling
on schemesused.Various'4Schnabel type?'
survivalof the animal.Duringthatperi- estimators7 that assume equal capture
od Seber(1965)and Jolly (1965)inde- probabilities, were reportedas having
pendentlydevelopedwhatis nowknown substantial bias. In addition,2 regression
as the Jolly-Sebermethodof estimating typeestimators proposed
open populationparameters by Tanakaand
frommulti- Kanamori (1967) and Marten (1970),
ple capture-recapture experiments. That of whichassumeda certainformofeach un-
lmodel,aspectsof whichwere latergen- equalcapture
eralizedby Robson(1969)and Pollock duce significantly probabilities, failed to re-
(1975),allowsthe populationto be ex- biasof the "Schnabel" the magnitudeof the
periencingdeath,recruitment, estiluators.
immigra- The appearance in 1965 of the Jolly-
tion,ancbpermanent emigration. Arnason Sebermethodof estimatingparameters
and Baniuk(1977)provideda compre- ofopenpopulations didnotprecludethe
hensivecomputeralgorithmto compute developtnentof additional
estimatesfor variousmodels for open techniquesin the literature,estimation in spite of
populations. Existenceof suchopenpop- the fact that Corluack(1968:487)
ulationmodelspointsout the need for lievedthe methodto be be-
testsforclosureof the populationunder powerful "anextremely
study.Unfortunately, generalforInulation" ofthecap-
good tests of that ture-recapture experiment. Although the
assuluption arestill notavailable. method is general in the sense
The importance of developingandus- lowsfor such processesas that it al-
ingvalidtestsof modelassumptions recruitment
was andmortality,it is restrictedby the as-
furtheremphasizedhy resultsappearing sumption thatall animalshavethe same
simultaneously in the literatureconcern- probability of captureona giventrapping
ingthe operatingcharacteristics of exist- occasion.In manyexperimental
ingestimationtechniques.Edwardsand tions,the assumptions situa-
Eberhardt's of populationclo-
(1967)studyon a confined sureand unequalcaptureprobabilities
rabbitpopulation of knownsize revealed
largebiasesin boththe Schnabel(1938) constitute a morerealisticset of assump-
tionsthanthe set requiredforthe Jolly-
and Schumacher-Eschmeyer(1943) Sebermodel. Hence,
methods of estimation.The authorscon- timators parameteres-
jectured that those biases were due to different derived from luodels basedon
sets of assumptions than
'<individualanimalshavingdifferentor ly-Seberrnodelscontinuedto betheJol-
changingprobabilitiesof capture.'7A opedby researchers devel-
silnulationstudyby Braaten(1969)indi- Kanamori suchas Tanakaand
catedseriousbias in the estimatorsde- Marten (1967),Eberhardt (1969a),and
(197()).
rivedfromDeLury's(1947)catch-effort forbias reduction The jackknife technique
modelif the assumptionof "constant ouille proposedby Quen-
(1949,1956)wasusedby Burnham
catchability'is violated.A similarlackof (unpublished dissertation) to derive an
robustnessto unequalcaptureprobabili- estimator for the situation
tiesamonganimalswasexhibitedby es- onember in whicheach
timators of the population has an <'in-
examinedin a computersimu- nate'> probabilityof capturethatvaries
lationstudy by Burnhamand Overton among individuals. Thateffort
(1969),
who generated"populations" us- auniqueattemptto developarepresents robustes-
ing
the familyof betadistributions. lMore timator of
recently,Carothers(1973b)sampleda parametric7 population size that is non-
populationof knownsize, the members assume i.e.7onethatdoesnotneedto
of
whichwerethe taxicabsof the city of tributed aowcaptureprobabilities aredis-
Edinburgh. over the population. Such non-
The assumptionof popula- parametric approachesare appealing
INFERENCE et al. 21
because they are robust to specific Unfortunately, it has been shown that
assumptionsregardingthe experiment Inisinformation resultsif, fora givenex-
andtendnotto sufferfrombreakdown of periment,assumptionsare not valid or
specificassumptions used to parameter-statisticalestimatorsare not appropriate
ize the model.Pollock(unpublished dis- or both.Thus,it shouldbe obviousthat
sertation)also consideredestimationof a rigorousapproachto paralneteresti-
populationsize underthe assumption of mationin capture-recapture experilnents
heterogeneityof captureprobabilities, will includea statisticaltesting algorithln
but withthe addedcomplication thatan thatallowsthe datato aid in selectionof
animal'sprobabilityof capturemay be the "best"set of assumptions forthe ex-
alteredby its capturehistory.However, perilnent.Althoughsolnetestsof specific
no specificestimationprocedureforthat assumptions havebeen introduced, uni-
lnodel had been proposed in the litera- fied approaches to the problem have not,
tureuntilthe appearance of the general- for the mostpart,receivedattentionin
ized reluovalmethoddescribedin this theliterature (anexceptionis theworkof
nonograph. Pollock,unpublisheddissertation). The
Thisoverviewwouldnotbe complete conceptof a unifiedapproach is thebasis
withoutluakingsome observations con- forthe developmentof this Inonograph.
cerningmethodsof densityestimationin Webelievean approach basedon sucha
capture-recapture experilnents. The no- conceptis a step in the directionof iln-
tionthatthe effectiveareaof trappingis provedanalysesof datafromcapture-re-
greaterthanthe actualareaof the trap- captureexperilnents.Furtherlmore, we
ping grid(i.e., the so-callededge effect) hopethatfutureresearchwillbe directed
has long been recognized.Dice (1938, to thatsalneobjective.
1941)correctedforthe effectby adding
to the grid areaa stripof one-halfthe MODEL MO:CAPrURE
holne rangeof the animal,and thatre- PROBABILITIES ARE CONSTANT
uainsthe Inostcommonpracticeat pre-
sent. Otherauthors(Stickel1954,Mohr Structt4reand Use of the Model
andStumpf1966,Smithet al. 1975)have Assumptions andParameters
used recaptureradiito correctfor edge
effect bias. Assessmentlines have also The simplestof all Inodelsundercon-
been used to estimatedensity(Kaufmansiderationresultsfrolnthe assulnption
et al. 1971,Smithet al. 1971).Morere- that all membersof the populationare
cently, Burnhamand Cushwa (pers. equallyat riskto captureon everytrap-
comm.) have forlualizedMacLulich's ping occasion.Moreover,the occasions
(1951)techniquefor estimatingdensity themselvesdo not affectcaptureproba-
thatinvolvesusingconcentrictrapgrids bilities.We thushavea Inodelin which
to allowsilnultaneous estimationof den- thereis noheterogeneity ofcaptureprob-
sityandedge width. ability,no behavioralresponseto cap-
An underlyingthemeof this historical ture,andnovariation in theexperilnental
overviewis that any capture-recapturesituationovertime.Thismodelis desig-
experimentrequiresthatthe researcher natedModelMov andinvolvesonly2 pa-
make specific assumptionsconcerning rameters:N, the populationsize, andp,
the manyfactorsthataffectthe resultsof theprobability thatananimalis captured
theexperiment. Theassuluptions thatare on anygiventrappingoccasion.
chosendeterminewhich statisticalesti-
mationproceduresshouldproducethe Statistical Treatlnent
bestresultsavailablefromthedata.Many
estimationprocedureshave been pro- The probability distributionof the set
posedbecauseseveraldifferentassump- ofpossiblecapturehistories{X(O} is given
tionscanoftenbe Inadefora givenfactor. by (cf.Darroch1958):
22 WILDLIFE IUONOGRAPHS
forModelMoaregivenin TableN.1.bof
P[{Xco}]= N! AppendixN.
[tI Xc,,!
] (N - Mt+l)
!
* pn- ( 1 - p)tN-n- ConfidenceIntervals
Achievedconfidencecoefficients ofthe
wheren. = E nj= total numberof cap- confidenceintervalproceduresimulated
=1 tures in the experi- were consistentlyat or above the 0.90
ment,and level and hence were close to the
Mt+1 = numberof differentanimals claimed 0.95 coverage.However,the
capturedin the experiment.widthof an averageintervalis so large
forsmallvaluesof p thatnotmuchinfor-
An algorithmfor producingML esti- luationconcerningtrue populationsize
matorsof N andp is derivedin Appendix is provided.Forinstance,forN = 400,t =
B. (Whent= 2, a closed formML esti- 5, andp= 0.05 (Trial3) expectedwidth
matorof N existsandis givenby No= (n1 [= 2 1.96 AveVVar(N)] is 628.2;andfor
+ n2)2/4m2, wherem2is the numberof re- N = 400,t= 5, p = 0.10(Trial2) thisval-
capturesin the secondsample.)These ue is 217.7. However, with p= 0.30
estimatorsare necessarilyfunctionsof (Trial1) expectedwidthdropsto an av-
the minimalsufficientstatistic{n., Mt+1}.erageof 46.9, indicatingthatthe model
Thus,all the information relevantfores- providesuseful information concerning
timationpurposesis containedin the N whenp is reasonably large.The num-
nuluberof differentanimalscaptured and ber of replicationsforthese 3 examples
thetotalnumberofanimalscaptured dur- were 500, 200, and 200, respectively.
ing the courseof the experiment. Appen- Oneshouldkeepin mindthatextremely
dix B also gives an estimatorfor the wide confidenceintervalstendto reveal
asymptotic varianceof Nothatwe usedin poor experimentalconditions,i.e., low
the construction of confidenceintervals values of p, and thus can be of use in
forN. providingthe experimenterwith infor-
mationconcerningthe successor failure
Simulation Results of the experiment.See Table N.1.b of
AppendixN forfurtherdetailsof the sim-
Bias ulationresults.
A computerwas used to simulateex-
perimentsfrompopulationssatisfying Robustness
theassumptionsofModelMo Byvarying
the population parameters N andp, some BecauseModelMois builtfromthe as-
insightinto the smallsamplebias of N suluptionthatno factorsthataffectcap-
was obtained.Resultsindicatethat the tureprobabilitiesarepresentin the ex-
bias of N is negligibleforvaluesof p at periment, it is not surprising that
leastas largeas 0.10andt ¢ 5. Forsmall- simulationresultsreveal that the esti-
er probabilitiesof capture,however,pos- matorderivedfromthis modelis not ro-
itive relativebiasesof 15-20percentare bustto anytypeof variability in the cap-
realized.Forexample,fromAppendixN, tureprobabilities. In particular2
if capture
TableN.l.b, fora population of size N = probabilities varyby animalNoexhibits
400,onesimulation consistingof200rep- significantnegativebias. This property
lications with p = 0.10 and t = 5 pro- has been documentedin the literature
duced an averagevalue of No of 406.0 (Robsonand Regier1964,Gilbert1973,
(Trial2), while another,based on 500 Carothers1973b).Commonsense and
replications,producedan averagevalue somereflectionon the natureof the ex-
of No of 456.9 with p = 0.05 (Trial3). perimentshouldtell us notonlythatbe-
Completeresultsof the simulationof No havioralresponsewill causebias in the
STATISTICAL FROMCAPTUREDATA C)tiSet al.
INFERENCE 23
NUS3ER OF TRAPPING OCCAS

IOIQSWAS 5
estimator,but will also indicatethe di- NUSER OF ANIMALSCAPTO.
OF CAPTS,
MXT+I) ( US
N., WAS
w
2M
rectionof thatbias.Thatis, animalsbe- TOTALR
comingtrapshy will causeNoto overes- ESTIMATEDPABILITY OF CAPTWE, P-T - .4672
timateN, and vice versawhen animals

becolnetrapaddicted.These assertions
are supportedby the simulationresults POPULATIOtlESTIMATEIS IOZ WITHSTANA E 2.389
given in TablesN.3.band N.4.bof Ap- APPROXIMATE CONFIDE:NCE

95 PE:RCENT INTERVAL 97 TO 107
pendixN. Resultsalsorevealthatthe es- FIG. 1. Exampleof populationestimatiollwithcon-

timatoris somewhatrobustto changesin stantprobabilityof captureunderModelMowith
captureprobabilitiesover time. How- simulated data based on N= 100, t= 5, alld
ever,Seber(1973)recommended, on the p = 0.5.
basisof Darroch's(1959)work,thatthe
estimatorassociatedwith ModelMonot theassumptions of ModelMoarefulfilled
be usedeven if the captureprobabilitiesforthiscylinderInodel.Thatis, the pop-
are suspectedof not varyingwith time. ulationis closed becausemarblesmay
This is goodadviceif largenumbersof not enter or leave the container,and
anilnalsarebeingcaughtbut sucha rule everyindividualhasthesameprobability
couldresultin someloss of efficiencyfor of captureon everytrappingoccasionbe-
smallsamplesizes. cause (1) all Inarblesare the samesize
and thus are not"heterogeneous,"(2)
Example whiteandblackmarbleshave the same
captureprobability andthusthereis no
A capture-recapture experimentthat "behavioral responseto capture," and(3)
satisfiesthe conditionsof ModelMOcan the salne''salnpling cylinder"is used in
be analogousto an urn experiment,a the sameInanneron all t occasionsand
samplingexperiment conductedin order thusthereis no "timevariation."
to estimatethe numberof marblesin a The factthatan analogycanbe drawn
cylinderwhen all marblesare the same betweena capture-recapture experiment
size. We may visualizea cylindercon- modeledby ModelMoand the simple
tainingN white marbles(individuals),urnexperimentillustratesthe pointthat
each of whichhas an equalprobabilityit is not reasonableto expectthatmany
(p) of beingpickedfromthe cylinderon capture-recapture studies can be ade-
any given occasion.On each of t occa- quately represented by Model Mo
sions,the followingsamplingschemeis Therefore,to presentan exampleof the
carriedout.A "samplingcylinder,"with estimationprocedureof ModelMo we
a dialneterthatis 100percentof the di- simulatedcapture-recapture salupling
ameterof the cylindercontainingthe for5 occasionson a population of 100in-
luarbles,is insertedinto the container dividuals,eachof whichhada 0.5 prob-
and a randomsample of marblesre- abilityof capture.As Fig. 1 shows,the
moved.The numbersof white (individ- value of the minimalsufficientstatistic
uals not previously "captured")and {n., Mt+1} is {238,98}.Thesevalues,and
black("recaptures") marblesin the sam- the value of t, are used to producethe
ple are recorded.All white marblesare population estimateof 102.BecauseN =
paintedblackand returnedto the con- 100,thisestimateis only2 percentgreat-
taineralongwiththe blackmarbles,and er thanthetruevalueof N. Notealsothat
all the marblesarerandomly mixed.The the lower limit of the large salnple9S
nuluberofwhiteandblackmarblesin the percentconfidenceintervalextendsbe-
salupleis recorded.Usingthe datafrom low the numberof differentmarbles
theset samples,theestimation procedure seen. This undesirableoperatingchar-
associatedwith ModelMoprovidesthe acteristicis revealedthroughout the re-
appropriate estimatorof N, the number sults of this study,and is discussedin
of marblesin the cylinder.Noticethatall AppendixO. When this happens,the
NUSBEROF TRAPPt NG OCCA51ON5 14AS

NUSER Ofo ANIMALSCSTURED, M(T+I 1, B5
5
106 The estimationprocedureassociated
TOTALNUMBER Of CAPTURE5, N., WAS 149
withthistnodelproducedtheresultspre-
ESTIMATEDPR%ABILITY y CAPTWE. P-HAT s .1726
sentedin Fig.2. Noticethatthe calculat-
ed 95 percentconfidenceintervalis rel-
ativelynarrow,probablydue to the fact
POPUI AT I ON EST I MATE I S 173 WI TH STANOARD ESRRf:BR 17.9t18 thatthe estimateof captureprobability
p
1S nearly u.z.
1 ,\ n
APPROX
I MTE 95 PERC:ENT
CONFICENCEl NTERVAL 137 T9 209
FIG. 2.Exampleof populationestimationwithcon-

stantprobabilityof captureunderModel Mowith Discussion
meadowvole data fromE. Larsen(pers. comm.).
Model Morepresentswhat mightbe
calledthe "best"of all possibleexperi-
lower limit shouldbe increasedto the mentalsituationsconsideredherein that
numberof distinctindividualsseen. Fi- a mlnlmumnumDerot nulsance pa- * * 1 r cs . X
nally,we mentionthatone wouldnotex- rametersis involved(one)if one is con-

pect the estimatorNoto be robustto de- cernedonly with estimationof popula-
parturesfromthe assumptions of the urn tion size N. This lack of nuisance
experiment. For instance, if black parameters resultsof coursefromthe re-
(marked) marbleswerelargerthanwhite strictiveassumptions onwhichthemodel
(unmarked) ones,andthushada higher is based.We believethatthoseassump-
probability of selection,we couldexpect tionsare in mostcases unrealistic,and,
Noto exhibitsignificant negativebias. therefore,the estimatorbased on the
modelis, in general,of limiteduse. The
Example caseagainstthemodelis strengthened by
the fact thatits associatedestimatorNo
E. Larsen(pers. comm.)used live- appearsextremelynonrobust to variation
trappingto estimatethe populationsize in captureprobabilities causedby behav-
of meadowvoles Microtus ochrogaster ioral responseor heterogeneity.More-
on a gridnearthe Flint Hills of Kansas over,it appearstruein generalthatlittle
in June1974.A 10 x 10gridof live traps, is gainedby usingModelMoinsteadof
spaced40 feet(12.2m)apart,waslaidout Model Mt when only time specific
in a tall-grassprairiethathad been un- changes in probabilitiesare present.
burnedandungrazedfor3 years.Onthe Therefore,the greatestutilityof Model
first 2 nights of trapping,traps were Molies in providinga "null"modeluse-
placedon top of the deep, dense litter ful in testingforsourcesof variation, and
thatuniformly coveredthe substrate,and in providing a basic model that can be
as a resultalmostno animalswere cap- generalizedin a numberof different
tured.Onthethirdnight,holesweredug ways.Suchgeneralizations are the sub-
in the litterandthe trapswereplacedin ject of concernin the following7 sec-
theholes.Thattrapping occasionyielded tions.
only12animalscaptured, perhapsdueto
the adverseeffectthatdisturbance of the
environmentmayhave had on the ani- MODEL Mt:CAPTURE PROBABILITIES
mals. On the last S nightsof trapping, VARY WITH TIME
however,relativelylargenumbersof an- Structureand Use of the Model
imalswere capturedconsistently.Thus Assumptions andParameters
we havechosento analyzethe datafron]
only those occasions.Whenappliedto The set of assumptions usedas a basis
thosedata,the discrimination procedure for ModelMtis the sameset associated
describedin the sectionentitledTESTS with the classicalmultiplecapture-re-
OF MODEL ASSUMPTIONSchose Model capture experiment. It is assumedthatall
Moas the appropriate modelforthe data. membersof thepopulation areequallyat
FROMCAPTUREDATAtiS
INFERENCE
riskto captureon the jth trappingocca- algorithmnecessaryto producethe true

sion. Thus, all animalshave ie same ML estimatefora glven set of dataare
probabilityof captureon any particulargivenin AppendixC. The estimatewill
trappingoccasion,but that probability dependonlyon the valueof the minimal
canchangefromoneoccasionto thenext. sufficientstatisticforthe model,namely
The set of parameters involvedin this {n1,n2, . . ., nt,Mt+1}.Notethatthemodel
model containsN, the populationsize, involvest + 1 parametersand that the
and the pj, j = 1, . . ., t, wherepj is the dimensionof the MSSis alsot + 1. This
probability of captureon thejtn occasion. assuresidentifiability of all the parame-
tersofthe model.Darroch (1958)alsode-
Treatment
Statistical velopedan estimatorof the asyluptotic
varianceof the MLestimator thatcanbe
ModelMthasreceivedmorestatistical used in the construction of a confidence
attentionthananyotherin the literature intervalforN (see AppendixC).
(see Cormack 1968).Schnabel(1938)first
usedthe aboveset of assumptions to de- Simulation Results
velop a model from which the well-
knownSchnabelestimatorwas derived. Bias
Her model,however,assumedthat the If the experimentalsituationis well
valuesof the Mj,the numberof marked represented by ModelMt,it is important
animalsin the populationat timej, are to know what biases luay be expected
known a priori,for j = 1, . . ., t. It re- fromthe estimatorof N discussedin the
mainedfor Darroch(1958)to derivethe precedingsection.Computer simulations
correctmodelforthe situation.Usinghis of experiments on populationssatisfying
results,we maywritetheprobability dis- theassumptions of ModelMtproducethe
tributionof the set of possiblecapture samegeneralconclusionsconcerning the
histories{X,O} as: biasof Ntasthoseproducedin ModelM
P[{X}]= N! Thatis, if the probabilities of capturep
are,on the average,closeto 0.1 orlarger,
] (N-Mt+l)!
[tl Xfi,! the bias of Nt is not significant.Again,
however,if the pj'Sbecomesmallerthan
pj)N-n3
0.1, significantbias results.Someexaln-
ples aregivenin Table1 forexperiments
* 11 pJni(1 -
j=l
conductedfor5 and7 trappingoccasions
where (additional resultsarepresentedin Table
nJ= numberof animalscaughton the N.2.b of Appendix N). In Table1, as in
jth occasion,and all tablesin this publicationAveE] rep-
Mt+1 = numberof differentanimalscap- resentsthe averagevalueof the quantity
turedin the experiment. in bracketsover all silnulatedreplica-
tions.R representsthe numberof repli-
Whent = 2, a closedformexpressionfor cations performedand RB represents
the maximumlikelihoodestimatorof N percentrelativebias.
exists and is given by Nt= nln2/ln2,
wherem2is the numberof recaptures in ConfidenceIntervals
the secondsample.This is the familiar
LincolnIndex. Darroch(1958)derived Confidenceintervalswereconstructed
an expressionthatmaybe solveditera- fromsimulatedexperimentsto compare
tivelyto give an estimatorof population achievedconfidencecoeffiicients to the
size fort > 2. One is led to believe that stated value of 0.95. Those achieved
this estimatorproducesestimateswithin levels dependon a numberof factorsof
unityof the trueMLestimateof N, but whichthe mostimportant are the accu-
this is notin factthe case.Detailsof the racyof the varianceestimatorof Nt?the
26 WILDLIFE NIONOGRAPHS
TABLE1. RESULTSOF COMPUTER STUDIESOF THE BL&SOF Nt (ALSOSEE APPENDIX

SIMULATION
TABLEN.2.b)
Percent Averageof True Model Mt Number Number
relative bias estimates populat}on Probabilitiesof capture of reps of occasions
RB Ave[NJ N pj j = 1, 2, . ., t R t Trial
22.0 1,015.6 800 0.01, 0.01, 0.02, 0.03, 0.03 200 5 11

10.7 442.7 400 0.03, 0.04, 0.05, 0.06, 0.07 200 5 5
2.2 408.6 400 0.10, 0.10 0.10, 0.10, 0.01 200 5 3
-0.7 198.7 200 0.30, 0.4070.10, 0.40, 0.30 100 5 11
-0.4 398.6 400 0.50, 0.20, 0.10, 0.10, 0.10, 0.10, 0.10 100 7 7
-0.2 399.4 400 0.20, 0.40, 0.30, 0.10, 0.20, 0.30, 0.20 100 7 9
degreeto whichNt is normallydistrib- estimatestend to be prohibitivelylarge

uted, and the accuracyof Nt itself. Re- when probabilitiesare small. In cases
sults indicatedthat the achievedconfi- where capture probabilities are ex-
dence level was in most cases at least tremelylow, lower confidenceinterval
0.90. Thoseresultsare encouraging be- limitsarenegative,thusprovidingno in-
causein manyof the experimental aboutN. As pointedoutin the
situ- formation
ationsinvestigatedthe assumptions nec- discussionof ModelNlohowever,such
essary for the strict validity of the experimentalresultsdo informthe re-
confidenceintervalused arenot met(cf. searcherthathis datahavebeen able to
AppendixO for a discussionof the as- tell him essentiallynothingaboutpopu-
sumptions).However,the amountand lationsize. In thatrespect,therefore,the
the kindof information containedin such calculatedconfidenceintervalis provid-
an intervalvariesfromone situationto ing relevantinformation, not aboutpop-
the nextdependingon the valuesof the ulationsize but aboutthe failureof the
captureprobabilitiespj, j= 1,2j...,t. If experimentitself.
thoseprobabilitiesareon the averageas To illustratethese points,Table2 has
largeas 0.20or 0.30,confidenceinterval been constructed, givingselectedsimu-
coverageis good and intervalwidth is lationresultstakenfromTableN.2.bof
small.Thus,veryusefulandreliablein- AppendixN regardingconfidenceinter-
forluationconcerningN canbe obtained val coverage(the proportionof replica-
undersuch experimental conditions.As tionsR in whichthe confidenceinterval
the capture probabilitespj decrease, containedN),andaverageconfidencein-
however so does the information about tervalwidth(Ave[C.I.width]).
populationsize providedby the confi-
dence interval.That is, intervalwidth Robustness
tendsto be so largeas to notprovideany
useful informationwith respect to N. Becausethe maximum likelihoodesti-
ThisresultagreeswithChapman's (1951) mator(orapproximations thereoflof pop-
argument, forthecaset = 2>thatvariance ulationsize N has been so frequently
CONFIDENCE
TABLE2. SIMULATED INTERVAL USINGNt (ALSOSEEAPPENDIX
WIDTHSANDCOVERAGE
TABLEN.2.b)
Model M
Population Probabilitiesof capture Number Numberof
Averasze size of reps occasions
[C.I. wiAth] Coverage N Pl P2 P3 P4 P5 R t Trial
,348.3 0.89 400 0.01 0.01 0.02 0.03 0.03 1,000 5

40.2 0.94 100 0.05 0.05 0.10 0.15 0.15 1,000 5 12
31.3 0.88 200 0 40 0.30 0.10 0.30 0.40 100 5 10
15.1 0.96 400 0.45 0.45 0.50 0.55 0.55 200 5 1
STATISTICAL INFERENCE
FROM CAPTURE DATAtis et al. 27
used in practiceover the last 40 years, OCCAS I ON

AN I MALS CAUGHT
J.
N 1J ) t
1 2
l5
3
16 29
t 5
l9
6
7
questionsof robustnessof those esti- 7
matorswith respectto departuresfrom TOTAL AN I HALS CAPTD 45
the assumptions of ModelMtarepartic- P-HAT(J)s .14 .30 .32 .48 .38 .14
ularlyrelevant.Simulationresultsindi-
catethatNtunderModelMtis nonrobust Pa>ULATION ESTIMTE IS 50 W I TH STAA EMW Z. 4
to failureof the assumptionthatall ani- APPROX I MTE 95 PERCENT CONF I DENCE I NTERVAL 44 TO 56
mals, regardlessof capturehistory,are

equallycatchableon the jth trappingoc- HISTOGRAH
OF N( J )
casion.Aspreviouslyemphasized, depar- FREQUENCY

_ _ _ _ _ _ _ _ _ _ _ _ _ _ _
7
_ _ _ _
15
_ _ _ _ _ _
16
_ _ _ _
24
_ _ _ _ _
19
_ _ _ _ _ _
7
_ _ _ _ _
turesfromthatassumption maybe dueto EACH* EQUALS 3 POINTS
captureprobabilities varyingwiththean- 2
21 .
§
imal,or the animal'sprobabilityof cap- 18

15 * *
*
*
*
turebeing alteredafterfirstcapture,or 12 * * * &

*
both. Althoughboth departuresmay 9

6
3
*
*
*
*
.
* *
. . .
createsignificantbiasin Nt,the natureof _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _

*
_ _ _ _ _
*
_ _ _ _ _
*
_ _ _ _ _
*
_ _ _ _ _
v
_ _ _ _ _
the bias differs.Thatis, departurefrom FIG. 3. Example of populationestimationwith

equal catchabilitycausedby heteroge- time specific changes in probabilityof capture
neity results in Nt being negatively under Model Mt with least chipmunkdata fro
biased,with the magnitudeof the bias V. Reid (pers.comlll.).
proportional to the amountof heteroge-
neitypresentin the population. Thisob- havioralresponseto first capture,and
servationcorresponds withthe resultsof Model Mh,in which heterogeneityof
the study by Edwardsand Eberhardt captureprobabilities occurs.In Table3,
(1967)on a rabbitpopulationof known 2 examplesof ModelMbareconsidered:
size. Biasof Ntthatis a manifestationof Trial8, whereanimalsthathavenotbeen
animalsexhibitinga behavorial response capturedpreviouslyhavea probability of
to capture,however,maybe eitherposi- captureof 0.2 on everytrappingoccasion
tiveornegative,according towhetherthe and a probabilityof capture0.05 after
animalsbecometrapshyortrapaddicted, firstcapture,andTrial9, wheretheprob-
respectively.Thisis the sameresultnot- abilityof captureforpreviouslycaptured
ed forthe estimatorof N associatedwith animalschangesfrom0.2 to 0.5. Three
ModelMoeThemagnitude of thebiasde- examplesof ModelMhare considered;
pends on the degreeto whichthe ani- forexampleTrial1, in which200animals
mal'sbehavioris changed.Table3 illus- have a 0.05 probabilityof capture,100
tratesthe bias of Nt when it is used in animalshave a 0.15 probabilityof cap-
simulated experimentsconducted on ture,and 100animalshavea 0.50proba-
populations satisfyingtheassumptions of bilityof capture.Whenthe estimatorNt
ModelMb,in whichanimalsexhibitbe- was computedfromthe datagenerated
TABLE 3.-COMPUTER SIMULATIONRESULTSILLUSTRATINGTHE NONROBUSTNESSOF THE DARROCH

ESTIMATOR Nt USINGDATAGENERATEDUNDEROTHERMODELS.DATA MODELSAREDESCRIBEDIN TABLES
N.3.a AND N.4.a OF APPENDIXN (ALSOSEE APPENDIXTABLES N.3.b AND N.4.b)
Percent Population Number Number of

relative bias size of reps occasions
RB Ave[N,] N R t Data model
199.3 299.3 100 100 5 Mb,Trial 8

-28.3 71.8 100 100 5 Mb,Trial 9
-43.0 228.0 400 100 5 Mh,Trial 1
-13.2 173.6 200 100 10 Mh,Trial6
-12.1 87.9 100 100 5 Mh,Trial 12
t For example, data generated under Model Mb were used to estimate N using estimator N.
WILDLIFE VIONOGRAPHS
28
underthis trialof ModelMh,an average spect to populationsize. As
percentbiasof -43.0 resulted(Table3). (1968)noted, Cormack
"thehigherthe proportion
of the population markedthe morestatis-
Example tically precise will be the estimateof
In the summerof 1975,V. Reid(pers. population size."If the captureprobabil-
ities are small,the varianceestimateof
comm.)laid out a 9 x 11 livetrapping Nt,andhencethe confidence
grid with trapsspaced50 feet (15.2m) N, tends intervalfor
to be quitelarge,tellingthe ex-
apartat a Coloradolocationin a bottom perimenterthatthe estimate
areadominatedby sagebrushandsnow- tion size N is unreliable. of popula-
berryand peripherallyby gambeloak, apparentpositive correlation Becauseof the
serviceberry,and juniper.Least chip- average capture probabilitybetween and the
munkEutamias minimus were trapped amountof useful
for6 consecutivedays (t= 6). The dis- containedin theexperiment, information about N
crimination proceduredescribedin the the experimenter it is wise for
TESTS OF MODEL
to calculatethe maxi-
ASSUMPTIONSsection mumlikelihoodestimates of thepj,j = 1,
choseModelMtas the mostappropriate2,. . .,t, fromthe dataat hand.
lmodelforthe data.Althoughthatchoice forthese MLestimators (Formulas
is subjectto some suspicion(the good- pendixC.)Such aregivenin Ap-
nessof fit test of ModelMtcouldnotbe videmorefeel for calculations shouldpro-
performed both the quantity and
becauseof insufficientdata), qualityof information contained in the
thedata were analyzedusing the esti- data.
mation procedureassociatedwithModel
Mtfor purposesof illustration.Results
(Fig.3) indicatethatestimatesof the pj MODEL \/t b CAPrURE
arelargeenoughforone to expectvalid PROBABILITIES VARYBY
andusefulconfidenceintervals,andsuch BEHAVIORALRESPONSETO
anintervaldoes appearto result.The CAPrURE
pointestimateof 50 animalsalso seems Structureand Useof the l\Jodel
tobe in line withthe observeddata,i.e.,
thefact thatu6 was 0 (no new animals Assumptions andParameters
caught on day6) leadsus to believethat Thismodeldealswith
thevalueof Ntshouldnotbe muchlarger assumption thefailureofthe
than that
the numberof differentanimalscap- affectthe probability initial capture does not
tured.A histogram of capture
of the njvaluesis in- sequentoccasions.Thatis, the on sub-
cluded to help the researcher to visually lowsananimalto exhibita behavioral modelal-
examine the data. re-
sponseto captureand become either
"trap addicted"or"trapshy."Overton
Discussion andDavis(1969)pointedout that"it is
Given the simulationresults of this well imals
knownthatso-calledtrap-happy
are often
an-
section,one mightconjecturethat fre- theexistenceof trap encountered." Moreover,
response been
quentuse of the"Schnabel method,"i.e., welldocumented(Geis 1955a,has 1955un-
Model in practicehas been unfortu-
nateunless tests of the assumptionsof published
lut,
doctoraldissertation,Michi-
ganState University, East Lansing,
Model lut haveindicated the modelmay Michigan; Tanaka1956, 1963; Flyger
beappropriate. Moreover,if Slodel Mt 1959; Bailey
appears to be anadequaterepresentation ly,Modellqb 1968; Pucek1969).Formal-
assumes
ofthe experimental situation,relatively trapping thaton anygiven
large
values(atleast0.2 on the average) have occasion, all unmarked animals
of
theparameters one probability of capture, and all
pjusuallyarenecessary marked animalshaveanotherprobability
toproduceuseful information with re- ofcapture.It is assumedthatthereis no
FROMCAPTUREDATA C)tis et al. 29
differencebetween trappingoccasions, Mt+1 = numberof differentanimals

i.e., thatcaptureprobabilities do notvary caughtduringthe entireex-
with time. An implicationof those as- periment.
sumptionsis thatall membersofthepop-
ulationhavethe sameprobability of cap-
tureat the beginningof the experiment. A MSSfor this probabilitydistribution
Note also that the assumptionis made is given by {Mt+l,m., M.}. Noticethat
thatan animal'scaptureprobability is al- this is a 3-dimensional vectorof statis-
tered only once, afterfirstcapture.Ai- tics, and because the model involves
thoughone mightthinkit morerealistic 3 parameters, each of those parameters
to allow the captureprobabilityto be is identifiable. Whatis mostnoteworthy
changedmorethanonce(e.g.,afterboth concerning the abovedistribution is the
firstand secondcapture)this moregen- factthatthe estimationof c is indepen-
eralassumptionturnsout to haveno ef- dentof the estimationof the parameters
fect on the estimationof populationsize N and p. That is, in Model Mb,once
N. Therefore,forsimplicityof presenta- ananimalhasbeencaptured, subsequent
tion the assumptionis made that all recapturesof the animalprovideno in-
Inarkedanimals,regardlessof the num- formation withrespectto the estimation
ber of timesthey mighthavebeen cap- of N and p. Therecapture information of
tured,havethe samecaptureprobability.theexperiment is usedonlyin theestima-
Theassumptions of ModelMbresultin tionofthenuisanceparameter c,theprob-
3 modelparameters: N, populationsize; ability of recapture. The reader is referred
p, the probabilityof captureof an un- to Appendix D for details
of the abovear-
markedanimalon any trappingocca- gument and for the derivation of the maxi-
sion;andc, theprobability thatananimal mumlikelihoodestimatorsof N and p.
is capturedon any trappingoccasion Thoseestimators are essentiallyequiva-
subsequentto the occasionon whichit lent tothose given byZippin(1956,1958),
wasfirstcaptured. who provided a basic statisticalanalysis
of the removalexperiment firstproposed
Statistical Treatment by Moran(1951).In Zippin'sdiscussion,
The probability distribution however,animalsusuallyarephysically
of the set removed
of possiblecapturehistories{Xs,,} froma ing, whereas by killtrappingor electrofish-
multiplecapture-recapture experiment "removed" in ModelMh,animalsare
on a populationsatisfyingthe assump- marked;in both,the from population by being
tionsof ModelMbmaybe writtenas(Pol- dependonlyuponfirstcaptures. estimators and N andp
lockunpublisheddissertation): Since estimationof populationsize
under the conditionsof Model Mh is
P[{X@}] = I| X!(N - Mt+l)! equivalentto estimationin Zippin'sre-
co
movalmodel,the "failurecriterion"as-
sociated with the removal method
pMt+I (
1 - p)N-M,+l-M-
applies.SeberandWhale(1970)showed
* Cm (1 - C)M-m. thatvalidMLestimators forN andp are
obtainedfromthe data when the cri-
whereM. = E Mj= sum(overj) of the terion
=1 t
numberof markedanimals E (t + 1 - 2j)(nj- mj)> 0
in thepopulation atthetime j=l
of the jth trappingoccasion, is satisfied.If thatcondition,whichtests

m. = E mj= totalnumberof whetherthe populationis being suffi-
j=l ciently"depleted"by the "removal" of
markedanimals,caught, new animals,is satisfied,a confidence
and intervalforN maybe constructed using
TABLE 4. RESULTS OF COMPUTERSIMULATIONSTUDIES OF THE BIAS OF Nb (ALSO SEE APPENDIX N,

TABLE N.3.b)
Percent Average of True Probability Number Number of

relative bias estimastes population of capture of reps occasions
RB Ave[Nb] N P R t Trial
-18.0 82.0 100 0.10 161 5 10

15.4 461.5 400 0.10 196 5 4
9.8 109.8 100 0.20 199 5 8
-0.7 198.6 200 0.25 100 7 6
-1. 1 197.9 200 0.30 200 5 5
the asymptoticvarianceestimatorof Nb frequentlyin cases whereN is small,a

givenby Zippin(AppendixD). relativelylarge numberof potentially
largeestimatesof N are"lost."Thisphe-
Simulatiorl Results
nomenonthereforeeffectsa significant
reductionin the valueof Ave[N].
Bias
ConfidenceIntervals
Becausebehavioralresponseto cap-
tureis commonin practice,an estimator Confidenceintervalswere simulated
of populationsize thatexhibitsrelatively forvariouspopulations satisfyingthe as-
smallbiases in populationsthat satisfy sumptionsof ModelMb(Table5).A high
the assumptionsof Model Mbwould correlation betweenthe valuesof p and
proveuseful in practice.Simulationre- adequateperformance of confidencein-
sults of experimentsconductedon such tervalsis indicatedby Table5. Evidence
populationsindicatethatNb,the MLes- indicatesthathighconfidencelevelsand
timatorof N, is a good estimatorif the small(henceinformative) intervalwidths
probabilityof capturefor an unmarkedcan be expected from experimentsin
animal(p) is at least 0.2. Frequencyof whichtheprobability of firstcaptureis at
"failure" of the experiment is substantialleast 0.30. On the otherhand,in an ex-
(20%)withp values60.1, andwhenthe perimentwithN = 100andp = 0.10low-
experimentdoes succeedin these cases, er limits of confidenceintervalswere
Nbtendsto be significantly biased.A few negativein morethan40 percentofthose
examplestakenfromTableN.3.bof Ap- simulatedexperimentsthat succeeded.
pendixN of the simulatedbiasof Nbare Suchresultsillustratethe pointthat,in
given in Table4 which illustratesthat general,confidenceintervalsconstructed
biaseson the orderof 15-20percentoc- fromexperimentswith an insufficiently
cur when p = 0.10, but that such bias largevalueof p serveonlyto informthe
graduallydecreasesas probability of first researcherthathis datacannotprovide
captureincreasesto reasonablelevels anyrealinformation withrespectto pop-
greaterthan 0.20. Althoughthe first 2 ulationsize.
simulations includedin Table4 havethe
sameprobability of firstcapture,the case Robustness
forN = 100is negativelybiased,whereas
the caseforN = 400 is positivelybiased. If oneexaminesthefailurecriterionfor
These seeminglycontradictory results estimationin ModelMb,it becomesclear
arecausedby the estimatorfailingmore thatthe numberof newanimalscaptured
oftenforthe N = 100case.Whenthe es- (removed)shoulddecreaseforeachsuc-
timatoris closeto failing(i.e.,the failure cessivetrappingoccasion.Simulation re-
criterionis close to zero),the estimates sultsseemto indicatethatsteepdeclines
tend to be biasedhigh. Becauseactual in new capturesovertimeproducegood
failureof the methodoccursmuchmore estimatesof and informative confidence
STATISTICAL
INFERENCE
FROMCAPTUREDATAtiS et al. 31
TABLE 5.-SIMULATED CONFIDENCEINTERVALWIDTHSAND COVERAGE

USINGNb (ALSOSEE APPENDIXN,
TABLE N.3.b)
Population Probability Number Numberof

Average size of capture of reps occasions
[C.I. wiath] Coverage N p R t Trial
253.9 0.71 100 0.10 161 5 10

145.0 0.95 20() 0.20 98 5
51.5 0.90 200 0.30 100 5 5
78.0 0.92 400 0.30 100 5 1
intervalsfor the parameterN. (Thisre- ulationof size 400 with200anilualsthat

latesdirectlyto the previousobservation have a 0.05 probabilityof capture,100
thatlargervaluesof theparameter p pro- with a 0.15 probability,
and 100 with a
duce estimatorswith betterproperties.) 0.50 probability (Triall), an experilnent
One can see thatif captureprobabilities with t= 5 trappingoccasionsproduced
varyfromone trappingoccasionto the an averageestimateof 258 animals.
next,the vectorof''reluovals'' represent-
ed by (u1,u2,...,ut)lnaybe perturbedin Example
such a way as to preventthe desired
monotonedecreasein the removalsover E. Larsen(pers.comm.)reportedthe
time.Thisresultsin Nbexhibitinga large resultsof a livetrapping experilnentcon-
bias and confidenceintervalsforN hav- ductedin the summerof 1976as partof
ing largeexpectedwidth.These obser- a studyon comlnunity succession.A total
vationssupportthe conjecturethatesti- of 55 live traps,spaced15 In apartand
mationof N basedon NlodelMhwill be covering0.81ha, were set out in a field
sensitiveto significantchangesin cap- in whichvegetationwasdominatedby a
tureprobabilities overtime.Moreover, if mixtureof sagebrushArtemesia triden-
captureprobability variesamongindivid- tata and rabbitbrushC1irysothamnus
uals,independentlyof the animal'scap- spp.Trappingwasconductedfor10con-
ture history,Nbtends to underestimate secutive nights.Data collected on the
N. The magnitudeof the bias depends deer mouse Peromyscus maniculat?4s
directlyon the numberof animalsin the fromthose10occasions(Fig.4) wereana-
populationthatareessentiallyuntrappa- lyzed by the discrimination procedure
ble, i.e., thosethathavesmall(<0.1)CCin-describedin the sectionentitledTESTS
nate"probabilities of capture.This non- OF NIODELASSUMPrIONSandit wasde-
robustnessto heterogeneityof capture terminedthatlModelMbwouldbe anap-
probabilityis of the samenatureas that propriatemodelforthe data(thesignifi-
exhibitedby Noand Nt. These remarks cance level of a goodnessof fit test of
arebasedon resultsobtainedfromsim- Model Mbwas approximately0.47).
ulationexperimentson populationsthat Thereforethe estimationprocedureas-
satisfythe assumptions of Modell4lh (see sociatedwithNlodelMb wasusedto pro-
followingsectionfora completedeserip- duce pointandintervalestimatesfor N.
tion of this model).A few examplesfol- Two aspectsof those estimates(Fig. 4)
low concerningthe biasof Nbin experi- areworthnoting:animalstendtobecome
ments on Model Mhpopulations.In a traphappysincerecapture probability(c)
populationof size 100,with 40 animals is morethan twice the value of initial
thathavea 0.05probability of capture,40 captureprobability(p)andthe sluallval-
animalswith a 0.10 probability,and 20 ue ofp = 0.09is themajorreasonwhythe
animalswitha 0.30probability (Trial8), 95 percentconfidenceintervalfor N is
an experimentwith t= 10 trappingoc- wideandthelowerlimitofthecoluputed
casionswassimulatedresultingin an av- intervalextendsfarbelowthe numberof
erageNbestimateof 78 forN. In a pop- animalsactuallyseen (as mentionedbe-
OCCASI ON J- 1 2 3 q 5 6 7 8 9 10 OCCAS
ION Js 1 2 3 4 5 6
TOTAL CAUGHT "{J)" 0 14 19 30 37 42 W8 51 57 6] SS TOTALCAUGHT HzJ ) s 0 15 23 29 32 35 38
NEwr CAUGHT UIJ)x 19 5 11 7 5 6 3 6 W 8 NEWLYCAUGHT U( J ) s 15 a 6 3 3 3
ESTIMATED PROBABILITY OF CAPTORE, P-HAT s .091 oe6
ESTIMATEDPROILITY OF CAPTURE,P-HAT w .342w13
E5TIMATED PROBABlLITY OF RECAPTURE, C-HAT - .2339e3
ESTIMATEDPAB IL I TY w KCAPTUK t C-HAT s . 61 1g0
ffRtATI ESTI>TE IS 112 WITH STA<AZ E 31.1
50 TO a 74
KPULATION £5T IMTE 15 WI WI TH STAZA ERROR 3 * 0518
APPROXIthATE95 PERCEN7 CONFIDENCE INTERVALS
AffltOX I MATE 95 PERCENT CONF I DENCE I NTERVALS 35 TO W7
HISTOG9v1 OF UtJ)
HISTOGRWt OF U{J}
FREauENcY 14 5 11 7 5 6 3 6 n 8
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ .
EACH * EOUALS 2 POI NTS FREQ(£M;Y IS 8 6 3 3 3

_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
EACH * EaJ^LS 2 PO I NTS
* * *
16
***** * .
14 §
14 ********v
lZ *
12 ****.***§
- - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - X
10
10
8
8 * *
FIG. 4. Example of populationestimationwith 6 6

4
§
*
*
*
.
* * * *
constantprobabilityremovalestimatorunderMod- _ _ _ _ _ _ _
Z
_ _ _ _ _ _ _ _
*
_ _ _ _ _
*
_ _ _ _ _
*
_ _ _ _ _
*
_ _ _ _ _
*
_ _ _ _ _
.
_ _ _ _ _
el Mb with deer mouse data from E. Larsen

(pers.comm.). FIG. 5.Exampleof populationestimationwithcon-
stant probabilityremovalestimatorunder Model
Mb with deer mouse data from V. Reid (pers.
comm.).
fore,the lowerlimitshouldbe takenas
69, not50, in this situation).
Example Discussion
V. Reid(pers.comm.)reportedthe re- Simulation resultsconcerningthe per-
sultsof livetrapping deermicePeromys- formance of Nbin populations thatsatisfy
cus maniculatus in a drainagebottomof the assumptions of ModelMbseem en-
sagebrush,gambeloak,andserviceberrycouraging in thatthe estimator andits as-
withpinyonpine andjuniperon the up- sociated confidenceintervalsprovide
lands.The area,in Rio BlancoCounty, usefulinformation in the presenceof be-
Colorado,wastrappedfor6 consecutive havioralresponseto firstcapture,if the
nightsin the summerof 1975.Trapswere probability of firstcaptureis sufficiently
arrangedin a 9 x 11 gridandspaced50 large(>0.1). (Hence,calculationof the
feet (15.2 m) apart.Analysisof the re- MLestimateof p shouldassistin assess-
sultingdataby the modelselectionpro- ingtheamountofinformation concerning
cedureindicatedthat ModelMbwould N thatis containedin thedata).However,
be appropriate for use in estimatingN the estimatorof N appearsnonrobustto
(e.g.,the significance level of a goodness otherfactorsthatmayhave an effecton
of fittestof ModelMbwasapproximatelyan animal'sprobability of capture.More-
0.43).The dataand the resultsof using over,theestimation procedure associated
Nb to producepoint and intervalesti- withModelMbis a specialcaseofa more
matesforN aregivenin Fig.5. Notethat generalestimationprocedurewhichin-
the mice tend to becometraphappy,as volvesfewerassumptions thanthoseas-
in the previousexample.In thatstudy, sociatedwithModelMb.Thisprocedure
however,the estimateof first capture is describedin the sectionon removal
probability is p = 0.34,a factthatlargely models.Forthesereasons,the estimator
accountsforthe narrowwidthof the 95 of populationsize N associatedwith
percentconfidenceinterval.Again,the Model Mbis useful only in those in-
lower limit of the confidenceinterval stanceswhereallunmarked animalshave
couldbe takenas 38, the numberof dif- the samecaptureprobability on all trap-
ferentanimalscaptured. plng occaslons.
* .
FROMCAPTUREDATA C)tiSet al.
INFERENCE
STATISTICAL 33
MODELMh:CAPrUREPROBABILITIES Statistical Treatment

ANIMAL
VARYBYINDIVIDUAL Recall that the randomvariableX
Structureand Use of the Model takes on the value1 whenthe ithanimal
Assumptions andParameters is caught on thejth trappingoccasionand
is zero otherwise.If it is assumedthat
The assertionthateachmemberof the {Pi}resultsfroma randomsamplefrom
population hasits ownprobability of cap- F(p)thenthe probability distributionfor
tureindependentof all othermembersof the set of variables{Xij}revealsthata
the populationis the basisof ModelMh. sufficientstatisticforModelMhis given
The assumptions are madethatthereis by the capturefrequencies{fl,f2,...,ft},
no differencebetween trappingocca- wherefj= the numberof animalscaught
sionsandno behavioral responseto cap- exactlyj timesin the experiment. Hence,
ture, but that there is heterogeneity alltheinformation forestimatingN is con-
amongthe captureprobabilities of indi- tainedin the frequencyof capturesta-
viduals.Cormack (1968)statedthata test tistics. Estimatorsbased on frequency
for the assumptionof"heterogeneityof of capturestatisticshavebeen proposed
individuals"is impossibleunlessan in- (Craig 1953, Tanton 1965, Eberhardt
dependentexperimentis conductedon 1969a),but are the resultof essentially
a populationof knownsize thatis "rep- ad hoc approaches.If the particular
resentative" of thepopulation of interest. familyof distributions of whichF(p)is a
Becauseof thatapparentdifficulty(and memberis specified,a luaxilnulnlikeli-
others),modelssimilarto ModelMhhave hood estimatorfor N may be derived.
been largelyignored,althoughthe as- For example, Burnham(unpublished
sumptionsbehindthe modelfitmoreex- dissertation)assumedthat {Pi} results
perimentalsituationsthan do most of froma rando1n samplefrolna 2-parame-
those commonlyused. As Eberhardt ter beta distributionand investigated
(1969a)pointedout,"varioussetsof data the propertiesof the resultingMLesti-
indicate. . . thatthe equal-probability-of- matorof N. Theoreticaland simulation
captureassumption is notfulfilled." resultsindicatedthatthis estimatorhas
Conceptually, ModelMhinvolvesN + quite unsatisfactoryoperatingcharac-
1 parameters: the populationsize N and teristics,and this inspiredBurnhamto
the set of captureprobabilities{Pi},i = developa nonparametric estimationpro-
1,2,...,N, wherePi is the probabilityof cedurethat would not requirespecifi-
captureof the ithanin1alon anytrapping cationof F(p). Such an estimatorwas
occasion.This luoregeneralforlnulationdevelopedusinganextensionofthejack-
of ModelMhdoesnotallowestimation of knifelnethodof biasreductionfirstpro-
populationsize N becauseof the pres- posed by Quenouille(1949,1956).The
ence of too largea numberof nuisance resulting form of the estimatorNh
parameters.Therefore,for most of the canbe writtenas
developlnentin this chapter,it is luore
usefulto thinkof {Pi}asa randomsample Nh = E ajfj.
of size N frolnsome probabilitydistri- i=l
butionF(p)definedon the interval[0,1]. Eachset of constantsal,a2,...,at,gener-

Unfortunately, thereis no colupletely atedby usinga different"order"of the
satisfactory estimatorwhen ModelMhis jackknifeprocedure, corresponds to a dif-
true. Therefore,one shoulddesign the ferent "jackknife"estimator.Burnham
studyto luinimizeheterogeneity. In fact, (unpublisheddissertation)suggesteda
it is worthrepeatingthatit is very im- sequenceof statisticaltests designedto
portantto designanycapturestudycare- choosethe best one of the estimators for
fullywith the goalof havingas few fac- any given dataset. Fora moredetailed
tors as possible affecting capture presentationof that estimationproce-
probabilities (see STUDYDESIGN). dure,consultAppendixE whichalsopre-
TABLE 6. RESULTS OF COMPUTERSIMULATIONSTUDIES OF THE BIAS OF Nh (ALSO SEE APPENDIX,

TABLE N.4.b)
Percent Average of True Number Number of

relative bias estimaXtes population Probabilities of capture of reps occasions
RB Ave[Nhl N pi i = 1, 2, . . ., N R t Trial
-17.2 331.1 400 Pi = 0 05, i = 1,200; Pi = 0.15, i = 201,300; 200 5 1

Pi = O.S0, i = 301,400.
0.2 100.2 100 Pi = 0.05,i = 1,40; Pi= O.lO,i = 41,80; 200 10 8
Pi = 0.30, i = 81,100
15.0 460.1 400 Pi= 0.10, i = 1,100; Pi = 0.20, i = 101,200; 200 5 3
Pi = 0 25, i = 201,300; Pi = 0.30, i = 301,400.
4.3 417.1 400 Pi = O.Ol,i = 1,50; Pi = 0.15, i = 51,200; 100 5 4
Pi = 0-25, i = 201,300; Pi = 0.30, i = 301,400.
3.7 207.0 200 Pi= 0.05,i = 1,50; Pi= 0.15,i = 51,150; 100 10 6
Pi = 0-25, i = 151,200.
11.0 443.9 400 Pi = 020, i = 1,100; Pi = 0.30, i = 101,200; 100 5 5
Pi = 0.40, i = 201,300; Pi = 0.50, i = 301,400.
13.4 226.8 200 Pi= 0.15,i = l,S0; Pi = 0.20,i = 51,100; 100 7 7
Pi = 0-25, i = 101,150; Pi = 0.30, i = 151,200.
sents a formulafor a varianceestimator range;and (2) even thoughsignificant

of Nh bias may sometimesbe presentin the
jackknifeestimator,the magnitudeof
Simulation Res?4lts suchbiaswill, in general,be substantial-
ly less thanthe biasthatwouldbe asso-
Bias ciatedwith any otherestimatorthusfar
Cormack(1968:497)noted that when discussed.Evidenceof thisassertioncan
animalsexhibit heterogeneouscapture be seenin TablesN.l.b, N.2.b,andN.3.b
probabilities"anyavailableestimateof of AppendixN thatpresentthe resultsof
populationsize [is] markedlybiased." using estimatorsotherthan Nh on data
We performedsimulationexperiments simulatedfromModelMh
withModelMhpopulations to determine
whether,in a givenpopulation, "marked ConfidenceIntervals
bias"is associatedwithNh.Table6 lists
someexamplesof suchresults.Thecom- Constructionof confidenceintervals
plete detailsof the simulationaregiven fromsimulatedexperimentaldata gen-
in TablesN.4.aandN.4.bofAppendixN. eratedfrompopulationsof Model lth
Resultsof thosesimulations andof those show a largevariancein the achieved
conductedby Burnham(unpublished confidencecoefficients(Table7). Cover-
dissertaion)seem to indicate 2 major ageis poorandrangesfroman estimated
pointsconcerningthe bias of the jack- O percent(Trials2, 11) to 87 percent
knife estimator:(1) generalstatements (Trial6) (AppendixN, TableN.4.b).Be-
concerningthe magnitudeanddirection cause resultsindicatethatthe distribu-
of the biasarenot easilymadesincethe tion of thejackknifeestimatoris approx-
estimatorexhibitsboth smalland large imatelynormal,poorcoverageis caused
andpositiveandnegativebias,depend- eitherby significant negativebiasof the
ing on the valuesof N andthe set {Pi} varianceestimator,a significantbias of
Onemayconiecture, however,thatif the the estimatorNh,or both.Unlessan ex-
numberof trappingoccasionsis suffi- perimenterhas data froman adequate
cientlylarge(saygreaterthanS) andif a numberof trappingoccasionsand feels
negligiblenumberof animalsis for all thatveryfew,if any,membersofthepop-
practicalpurposesuntrappable, thenthe ulationunderstudyareuntrappable, the
bias of Nh will be within a tolerable confidence intervalconstructed by the
STATISTICAL FROMCAPTURE DATAtis
INFERENCE et al. 35
TABLE 7. SIMULATEDCONFIDENCEINTERVALWIDTHSAND COVERAGE ESTIMATOR

USINGTHEJACKKNIFE
Nh (ALSOSEE APPENDIXN, TABLE N.4.b)
Population Number Number of

Average size Probabilities of capture of reps occasions
[C.I. widthl Coverage N ps i = 1,2, . . ., N R t Trial
89.9 0.18 400 ps = 0.05, i = 1,200; Pi = 0.15, i = 201,300; 200 5 1

Pi = 0.50, i = 301,400.
104.9 0.40 400 Pi = 0.10, i = 1,100; Pi = 0.20, i = 101,200; 200 5 3
Pi = 0.25, i = 201,300; Pi = 0.30, i = 301,40().
45.4 0.87 200 Pi= 0.05, i = 1,50; Pi = 0.15, i = 51,150; 100 10 6
Pi = 0.25, i = 151,200.
45.3 0.68 100 Pi = 0.05,i = 1,40; Pi= 0.10,i = 41,80; 200 5 8
Pi = 0.30, i = 81,100.
methods associated with Model Mh jackknifeestimatorwill providean ade-

should be consideredunreliable.The quateestimateof populationsize in an
readermayreferto TableN.4.b of Ap- experilnentin whichmanyanimalsare
pendixN forcompleteresultsconcerning caughta relativelylargenuluberoftimes.
simulatedcoverageof these confidence Someexamplesof the bias of the jack-
intervals. knifeestimatorwhendataaresimulated
frommodelsotherthanMh aregiven in
Robustness Table 8. Moreinformation maybe ob-
tained froln Tables N.2.b, N.3.b, and
Thejackknifeestimatorwas construct- N.5.b of AppendixN. In the table,the
ed with the objectiveof havingan esti- informationunder the heading Data
luatorthatis robustto heterogeneityof modelrefersto the populationon which
capture probabilities.The degree to the simulatedexperimentswere per-
whichthe objectiveis satisfiedhasbeen formed.Fordetailsof thesepopulations,
discussedpreviously.The questionre- see AppendixN, TablesN.2.a,N.3.a,and
luains:is the jackknifeestimatorrobust N.5.a.
to otherformsof departurefromthe as-
sumptionof equalcatachability? Results Example
indicatethatof the estimatorsdiscussed
in this monographthe jackknifeis the Carothers(1973a)conducteda cap-
Inostrobust.This does not meanthatit ture-recapture experilnenton the "pop-
is a good estimatorno matterwhatas- ulation"of taxicabsin Edinburgh,Scot-
sumptionsapplyto the populationunder land.Thepopulation, knowntobe of size
study.In general,it maybe saidthatthe 420 andassulnedto be closed,was sam-
TABLE 8. COMPUTER SIMULATIONRESULTS ILLUSTRATINGTHE ROBUSTNESSOF THE JACKKNIFE

ESTIMATORNh TO DATAGENERATEDUNDEROTHERMODELS(ALSOSEE APPENDIXN, TABLESN.2.b, N.3.b
AND N.S.b)

RB Ave[Nh] N R t Data nodel
-12.5 349.2 400 200 5 Mt, Trial 31

- 1.3 98.7 100 1,000 5 Mt, Trial 12
-17.6 82.3 100 100 5 Mb, Trial 9
-13.1 347.7 400 200 5 Mb, Trial 4
0.7 402.9 400 100 5 Mth,Trial 1
19.7 478.7 400 100 s luth, Trial 2
' For example, data generated under Model Mtwere used to estimate N using the estimatorNh.
36 WILDLIFEMONOGRAPHS
NU%ER OF TQ^ IK OCCASI BS 6

tet R OF YRAF'PINC OCCAS t ONS WAS lO
ER OF ANlrlALS CAPTD. Mt r I ) . WAS 68
N9ffR OF ANIMALS CAPTUREO. M(T+I , WAS 283 rorz NUSBER OF CAPTURES. N . . WAS 145
*o r^ tAJ"3t R OF CAPTJRES, N ., WAS 50 0
FRtQUENCIES Of CAPT,Ff II
FREOUENCI £ 5 0F CAPTURE, F ( I )
Is I 2 3 4 5 6
It I Z 3 4 5 6 7 8 9 10 Ff I 8s 25 ee 13 5 1 2
Ftl)- zqe 81 W9 7 3 1 0 ° O °
COff>UTED JACKKNI FE COEFFI C I ENtS
COMPUTEOJACKKNI FE COEFFI C I ENTS
Nf I ) Nte) Nt3) Nt4) Nt5)
hit l ) Nt2) Nt3} Nl4) NiS)
| a .833 2.500 3. 000 3. 333 3.500
1 1.900 2.700 3.40Q 4.000 b.500
e X. ooo . 61 - .233 - .833 - I . 167
2 1 . ODO .289 - .878 -2 .278 -3 .722
3 1 . 000 1 . 000 1 .225 1 .542 I .750
3 1 . 000 1 .000 1 . W76 2.535 q . 042
4 1.000 1 . OoO I . OoO .956 .9 1q
W I . 000 1 . 000 l . 000 .79 3 . 077
1 . 103 5 1 .000 1 .000 1 .000 l .000
5 1 .000 1 .000 l .000 1 .000 I .001
THE RESULTS OF THE JACKKN[FE COMPUTATIONS THE RESULTS OF THE JACKKNIFE COMPUTATIONS
I Nt I ) SE( I ) .95 CONF. LlMlTS TESt OF N( 1+1 ) VS. N( I ) I N( I } SE ( I ) .95 CONF. L IMIT5 TEST OF Ns i + 1 ) VS. N( I )
0 283 CHI-SQUARE( I D.F. 0 68 CHI-SOUAREXI O.F, )
1 4 10.8 15.58 380. 3 44 1 .3 25.877 1 88.8 6. Ia 76.7 lOO.9 1.4]0
2 466.8 Z5.20 4 17.4 5i6.2 4.225 2 93.8 9.40 75.4 1 i2.Z .000
3 495.0 36.42 423.7 566.4 t.807 3 93.8 12.65 69.0 t 18.6 .078
4 516.9 49.98 418.9 614.9 1.735 4 92.8 15.95 62.5 123.1 I 18
5 540.4 65.29 412.4 668.4 0.000 5 92.Z 17.06 58.7 125.6 0.000
AVERAGEP-hAT s . 1062 AVERAGEP-HAT = .2778
INTERPOLATEDPOPULATiON ESTIMATE IS 47t WITH STANDARDERROR 26.8266

INTERPOLATEDPOPULATlON ESTIMATE IS 81 WlTH STAZARD ERRZ 5.8Z20
APPROXIMATE95 PERCENT CONFIDENCE INTERVAL 418 ro 5Z4
APPROXIMATE95 tRCENT CONFIDENCE INTERVAL 75 TO 99
HISTOGRAMOF F t I ) HI STOGRAMOF F t I )
FREOuENCY 142 81 49 o
FREQUENCY 25 22 l 3 5 l 2
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
EACH Y EOVALS 15 POINTS

EACH * EOUALS 3 POI NTS
135 29
xeo 21 *
105 i8
90
15
75 12 * *
60
9 * *
45 * *
6 * *
30
3 * * * *
15 _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
FIG. 6. Example of populationestimationwith FIG. 7. Example of populationestimationwith

variableprobabilityof captureby animal under variableprobabilityof captureby animal under
NIodel Mh with the colnplete set of Scheme A Model Mh with snowshoehare data from Burn-
taxicabdatafromCarothers(1973a). hamand Cushwa(pers.colnm.).
pled for 10 consecutivedays.The sam- intervalconstructedfor N does in fact

plingwasdoneby observingtaxicabsthat containthe truepopulation valueof 420.
passedpreselectedpointsandrecording These resultsare as satisfactoryas any
"capture"or "recapture"accordingto producedby variousestimationtech-
whetheror not the particularcab had niquesusedby Carothers.
been observedpreviously.In the au-
thor'ssamplingscheme"A,"a different Example
set of samplingpointswasselectedeach
day,andthe timeof sampling(i.e.,morn- In 1972,Burnhamand Cushwa(pers.
ing,afternoon,night)wasalsovaried.We comm.)laid out a livetrappinggridin a
appliedthe model selectionprocedure black spruceforest30 miles (48.3 kln)
(describedin TESTSOF MODELASSUMP- northof Fairbanks, Alaska.Thebasicgrid
TIONS)to the data collected fromthis was 10 x 10, with trapsspaced200 feet
scheme.Theprocedure indicatedthatan (61 m) apart.Trappingfor snowshoe
appropriate modelforthe datawouldbe haresLepus americanus was carriedout
ModelMh.(Forexample,a goodnessof for 9 consecutivedays in earlywinter.
fit test of ModelMhhas a significance Trapswerenotbaitedforthefirst3 days,
level greaterthan0.99).Thus,the esti- andthereforewe havechosento analyze
mationprocedureassociatedwithModel the datafromthe last6 daysof trapping.
Mhwas usedto producepointandinter- Themodelselectionprocedure designed
val estimatesforN (Fig.6). AlthoughNh to identifya propermodelforestilnating
overestimates the truevalueof N by ap- N choseModelMhas the Inostappropri-
proximately12 percent,the confidence ate modelforthe data.A goodnessof fit
FROMCAPTUREDATAtis et al. 37
testof ModelMhhada significance level MODEL Mtb: CAPTUREPROBABILITIES

of 0.06.The data,andthe estimatespro- VARYBY TIME AND BEHAVIORAL
duced by the estimationprocedureof RESPONSETO CAPrURE
ModelMharegivenin Fig.7. Theresults
of the entire9 daysof trappingshow74 Stractureand Useof theModel
differentanimalscaughtwhilethe lower Assumptions andParameters
95 percentconfidencelimit for N given
in Fig.7 is 75.Althoughthisresultis sat- If it is assumedthatan animal'sprob-
isfying, we reemphasizethat the true ability of capturechangesafter initial
confidencecoefficientof the confidence captureand thattemporalchangesalso
intervalsassociatedwithModelMhis, for have an effecton captureprobabilities,
most populations,much less than the ModelMtbresults.This modelhasbeen
statedlevel of 0.95. conceptualizedin the literature(see
Eberhardt et al. 1963,Seber1973)butit
hasnotreceivedmathematical treatment
Discussion
with respect to estimation of population
parameters. Althoughmanyvariationsof
Theoretically, 2 maincomplicating fac- the basic Inodel exist(see AppendixF),
torsare associatedwith the existenceof forpurposesof this chapterwe shallde-
heterogeneity of individualcaptureprob- finethe parameters of ModelMtbas:
abilities.First,a parameter mustbe in- N = populationsize,
troducedinto the modelfor everyindi- pj= the probability an unluarked animal
vidualin the population, andthismakes is capturedon thejth trappingocca-
straightforward statisticalestimationof sion,j = 1,2,. . .,t,
populationsize impossible.Second,the c; = probabilitythat a marked(previ-
assuluptionof heterogeneitycan render ously captured)anilnalis captured
solneindividualsnearly"invisible" with on the jth trappingoccasion,j=
respect to any estimation procedure 2,3,. . .,t.
basedon Inarking lnethodsbecausesuch
individualshavenearlyzerocatchability. Notethatagain,as in ModelMh,an ani-
Thejackknifeestimation procedureis the malexhibitsonly 1 behavioralresponse
mostrobustprocedurethusfarproposed to captureafterits initialcapture.
for dealing with problelnsassociated
with heterogeneity.Moreover,the esti- Statistical Treatlnent
lnatorseems robustto otherfactorsthat
mayhavean effecton captureprobabili- Thejointprobability distribution ofthe
ties. It shouldbe pointedout,however, set of possiblecapturehistories{X,,}can
thatin solne cases the bias of the esti- be writtenas
matoris notnegligible.In general,if the
jackknifeprocedureis to be considered P[{Xco}] = N! II
a good nonparametric and robusttech- [tI X! ](N - Mt+l)! j=l
nique, the trappingmustbe performed
on a largenumberof occasions,andthe * (1 - pj)N Mj+t(Cj)mi(l - Cj)Mj-mi
nulnberof recaptureson each occasion where
nustbe substantial.
We relnarkthatthe observations con- u;= nulnberof unmarked anilualscaught
tained in this section reflect Gilbert's on thejth occasion,j = 1,2,. . .,t,
(1973:524)opinionthat"thepresenceof Mj= numberof markedanimalsin the
heterogeneity per se is not as important populationthe the time of the jth
as the particular patternof heterogeneity sample,j = 1,2,. . .,t,and
and whetherthe heterogeneityis near m.j= numberoflnarkedanimalscaptured
zeroor one. ,,
on thejth occasion,j = 1,2,. . .,t.

A minimalsufficientstatisticforthis dis- MODEL Mth: CAPrURE PROBABILITIES

tributionis given by {ul, u2,...,ut, m2, VARYBY TIME AND INDIVIDUALANIMAL
m3,...,mt}whichhas dimension2t- 1.
Sincethe modelinvolves2t parameters, Structureand Useof the Model
not all parameters canbe estimatedand Assumptions andParameters
maximumlikelihood estimationof N
provesto be impossible.The structure of If, on the jth trappingoccasion,the ith
P[{X,O}] revealsthatestimationof the pa- animalhas a captureprobability of Pii =
rametersN, P1, P2, . . , Pt dependsonly PiPithatis independent of its capturehis-
upon the vectorof removals{ul,u2,.... tory(i.e.,thereis no behavioral response
ut}.Thus,estimationof N in ModelMtb to capture),thenModelMthis the appro-
wouldbe equivalentto the estimationin priateprobability modelfora capture-re-
Zippin's(1956)removalmodelgeneral- captureexperiment on sucha population.
ized to allowthe probability of removing Notice that the structureof Pijimplies
an animal,pj, to be differentfor every thatvariation in captureprobabilities due
trappingoccasion.Thisinvolvesestimat- to time is independentof the variation
ing t + 1 parameterswith t statistics. causedby individualheterogeneity.In
Hence,in orderto makeN anidentifiable effect,thismeansthatthe factorsrespon-
parameter, one has to makethe assump- siblefortimevariation, e.g.,environmen-
tion that at least 2 of the pj are equal. tal conditions,affectall membersof the
Becausethereappearstobe nobiological population similarly.Forpurposesof this
justificationformakingsuchan assump- sectionit shallbe assumedthatthePi, i =
tion, only ad hoc estimationprocedures 1,2,. . .,N,area randomsampleof size N
areavailable.Forexample,Tanaka(1951, fromsomeprobability distribution-func-
1952) has proposeda regressiontech- tion F(p;8) thatis parameterized by the
nique that involves plottingyj= mj/n vectoro anddefinedforp in the interval
versusMjon a logarithmicscale;how- [O,1]. Notethatit is necessarythateach
ever,Seber(1973)pointedout thereare of the parameters,pj, j= 1,2,...,t, be
serious problemsof interpretationin- subjectto the constraint thatO> PiPi> 1
volvedin usingthis technique.It is true fori = 1,2,...,N. Conceptually, the form
that graphicaltechniquessuch as this of the distributionfunctionF(p; @)and
lmaypossesssomeutilityin thattheyen- the parameters@,Pl,P2,. ., Pt are un-
couragethe researcherto examinethe knownelementsof the model,as is the
data carefully. However, regression parameter N.
methodsarenotcontainedin the classof
estimationmethodsconsideredin this Statistical
Treatment
monograph.Therefore,Tanaka's(1951,
1952)methodandthosesimilarto it will Estimationof populationsize under
notbe consideredhere. the assumptionsof NIodelNIthhas not
been consideredin the literature.If one
is willingto completelyspecifythe dis-
Discussion tributionF(p; 8), then maximumlikeli-
hoodestimationof N becomespossible
In view of the previousdiscussion,we (cf.AppendixG).In general,suchan as-
believe thatModellutb iS not usefulfor sumptionis not realistic.Furthermore,
estimationpurposes.However,the mod- simulationresultsof Burnham(unpub-
el does have utilitywith respectto the lished dissertation) indicatethatan ML
problemof choosinga "best"estimation estimatorresultingfrolnsuchas assu1np-
proceduregiven the dataat hand.For tion luayhavepooroperatingcharacter-
detailsof howthe modelis usedin such istics.
a testingproceduresee the sectionon If no assumptionsconcerningF(p; @)
modelselection. aremade,MLestimationof N is notpos-
STATISTICAL
INFERENCE
TABLE 9. COMPUTERSIMULATIONOF THE:BLASASSOCIATEDWITH THE ESTIMATORSNh AND Nt FOR

DATAGENERATEDUNDER MODEL Mth (ALSOSEE APPENDIXN, TABLE N.5.b)
Percent Percent Population Number Numberof

relative bias relative bias size of reps occasions
RB Ave[Nh] RB Ave[NJ N R t Trial
0.7 402.9 -24. 1 303.6 400 100 5 1

19.7 478.7 - 7.S 369.9 400 100 5 2
- 1 1.5 353.9 -22.8 309.0 400 100 5 3
- 1 1.8 352.8 -31.8 272.8 400 100 5 4
8.9 217.7 - 6.1 187.9 200 100 7 5
- 10.9 178.2 - 17.9 164.2 200 100 5 6
' The probability of capture for data generated under Model Nlth iS complex; therefore, the specific values are given in Appendix N,
Table N.5.a.
sible.Atpresent,we arenotawareof any matorsof ModelMhand ModelMtwas

rigorousestimationtechniquethatis ap- poor(cf.TableN.5.b,AppendixN). The
propriateforestimationin ModelMth lackof coverageis due,in general,to the
significantbiaspresentin the estimates.
Simulation Results Therefore,we recolumendthat confi-
Bias dence intervalsconstructedfroln data
thatapparently fit ModelMthbe consid-
To gain some insightinto the perfor- ered only as a very crudeindicationof
mance of estimatorsassociated with thepossiblevaluesof N andnotasa valid
modelscloselyrelatedto ModelMth,we statementof inference.
simulated capture-recaptureexperi-
mentson populationsthatsatisfythe as-
sumptionsof ModelMthand calculated Robustness
the estimatorsassociatedwith ModelMt A discussionof the robustnessof the
and ModelMhfromthe resultingdata. estimatorassociatedwith ModelMthis
Examplesare givenin Table9. See Ta- somewhatinappropriate becauseno sin-
bles N.S.aand N.5.bof AppendixN for gle estimatoris exclusivelyassociated
a descriptionof the populationsusedand withthemodel.Therobustness ofNhand
forresultsof othersimulations. A choice Nthasbeen discussedpreviously.Those
betweenthe 2 estimators withrespectto discussionsindicatedthatthe formeres-
performanceis obviouslynot clearcut. timatoris relatively robustand thatthe
Themagnitude of thebiasof Nhseemsin latteris not. Suchassertions seem to be
generalto be less thatthatof Nt. How- supportedby the resultsof this section
ever,directionof biasof Nhis notconsis-
tent,whereasthe estimatorNtappearsto concerningbias.
consistentlyexhibitnegativebias.Thus,
if the dataseem to fit ModelMth,com- Discussion
putingboth estimatesof N shouldgive ModelMthis one of the mostdifficult
the experimenter somesense of the size models dealt with in this monograph.
of the population.However,it shouldbe Thatdifficultyis reflectedin the factthat
emphasizedthat the model does not no estimation procedure can proposed
seem particularly useful in providinga thatis specificallysuitedtobe
all assump-
single satisfactoryestimateof the value tionsof the model.Estimators associated
of N. with2 previousmodelshavebeeninves-
ConfidenceIntexvals tigatedforpossibleusebutneitherseems
completelyadequate.Thus, at present,
Coverageof confidenceintervalscon- satisfactorytechniquesfor constructing
structedboth on the basis of the esti- pointandintervalestimatesof N arenot
available.Hopefully,furtherresearchin- butionfunctionG(p,c; @)thatis param-

volvingModelMthwill be forthcomingeterizedby the vectorof unknowncon-
so thatthe modelcanbe dealtwith in a stants@.If one assumesthatthe formof
moreadequatestatisticalmanner.Until the distribution G(p,c; @)is knownand
then, one mustbe satisfiedwith obtain- if the dimensionof @ is no largerthan
ing onlyveryroughestimatesof popula- [t(t+ 1)/2]- 1, wheret is the numberof
tion size when ModelMthis the appro- trappingoccasions,then theoretically
priaterepresentation of the experiment. maximum likelihoodestimationcouldbe
usedto providean estimatorof N. How-
ever,a "nonparametric" approach is also
MODEL Mbh: CA7ruRE PROBABILITIES possible.Wediscussthisapproach in the
VARY BY INDIVIDUAL ANIMAL ANDBY contextof the assumption thatthe bivar-
BEHAVIORAL RESPONSE TO CAPruRE iate densityfunctionG(p, c; @)can be
factoredas G1(p;Ol)G2(c;492)Thatis, an
Structureand Use of the Model individual'sinitial captureprobability
and its probabilityof recaptureare in-
Assulnptions andParameters dependent.With that assumption,Pol-
ModelMbh assumesthateverymember that lock (unpublisheddissertation) showed
the probability distribution
of the populationhas a specificpairof of possiblecapturehistories{X(O} of the set
captureprobabilities: Pi, the probabilitywrittenas can be
thatthe ithanimalis caughton anytrap-
ping occasiongiventhatit has not been
previouslycaptured;and ci, the proba- PE{X@}] =U1tU2! . . . UtT(N - Mt+l)!
bilitythattheithanimalis recaptured giv-
en thatit hasbeencapturedat leastonce 87TtUt(l - Erj)N-M,+1
previously.Thus,the modelallowsboth 1 1T2 2 * * *
behavioralresponseto firstcaptureand *P*[{X@} I Ul,U2, * * *,Ut],

individualheterogeneity of captureprob-
abilities. An importantand appealing
characteristic of the modelis thatit al- where
lows the behavioralresponseto capture
to varywiththe animal,i.e., all members uj= number of unmarkedanimals
of the populationdo notexhibitan iden- caughtat timej, j = 1,2,. . .,t,
ticalresponseto firstcapture. r
The most generalformulation of the 7rj= E[(1 _ p)j-lp]= J (1 _ p)j-lpdG
modelinvolves2N + 1 totalparameters o
consistingof N (populationsize), and 2 j = 1,2,.. .,t,and

captureprobabilitesfor each of the N P*[{XCt)}lUlX U2, * * ., Ut] =
membersof the population.Obviously, a conditionalprobability distribu-
no real experimentwill be able to pro- tion that does not dependupon
vide enoughinformation forthe explicit the parameter N or the distribu-
estimationof this numberof parameters. tionGl(p;@l).
Furtherassumptionsare necessarybe-
foreestimationof populationsize is pos- (Referto AppendixH forfurtherdetails).
sible. Thus, the "removals"{ul,u2,...,ut}are
Statistical
Treatment the relevantstatisticsforthe purposeof
estimatingN and the parameters 81 in
Pollock(unpublisheddissertation) in- Gl(p;81),andthesestatisticshavea mul-
troducedModelMbhand assumedthat tinominaldistribution withparameters N
the pairs(Pi,ci) are a randomsampleof and T1,X2, . . ., 7Tt (Seber 1973:316gave
size N froma bivariateprobability distri- thismodelforthe specialcaset = 3). We
STATISTICAL FROMCAPTUREDATA{)tiS et al.
INFERENCE 41
can transformthe rj by writing rj= 62(c; 82) be possiblein orderto use the
(1 - p1)(l- P2) (l - Pi-1)Pi, j = 1,. . .,t, generalizedremovalmethodto estimate
wherepjis the averageconditional prob- populationsize. It is important to realize
abilityof captureon the jth occasionfor that this methodis a generalization of
those animalsnot previouslycaptured. Zippin's(1956, 1968)reluovalmethod
The distributionof the t-dimensional whichassumesno variationin firstcap-
vectorof removalsnow dependsupon tureprobabilities. Therefore, thefactthat
the t + 1 parameters N, P-1,P-2,- .,P-t,and the generalized relmovalmethodhasutil-
thusall the parameters of the modelare ityin removalexperilnents aswellascap-
notidentifiable. Thisis similarto the sit- ture-recapture experiments is notsurpris-
uationthatresultedin ModelMtbwhere ing (cf.REMOVALMODELS).
it was conlcudedthatMLestimationof Finally,we mentionthatthe lnethod
N is notpossible.If individualprobabil- can "fail"if a mathematical criterionin-
ities of firstcapturevaryoverthe popu- volvingthe removalsul,u2,...,utis not
lation,however,it is logicalto assume satisfied.Thisfailurecriterion,siluilarto
that P1> P2> P3 > * * *) P-t,becausethe the one involved in Zippin'sremoval
individualswithhighfirstcaptureprob- method,ensuresthata sufficientdecline
abilitywill tendto be removedfirst,the in the nuluberof newlycapturedanimals
animalswith slightlylowerfirstcapture is beingeffectedby successivetrapping
probabilities removedsecond,andso on. occasions.A formulaexpressingthis cri-
Moreover, theassumption that(P1- P2)> terionis givenin AppendixH.
(P2- P3) > * *) (Pt-l - Pt), i.e., the big-
ger differencesin the conditionalproba- Simulation Results
bilities of removaloccurin the initial
stagesoftheexperiment, doesnotseemil- Bias
logical.Theseassumptions arethe basis Experilnents weresilnulatedon popu-
forthedevelopment of anestimation pro- lations behavingaccordingto the as-
cedureappropriate forNIodelMbh. Thees- sumptionsof ModelMbh. The estiluator
timatorNbhderivedunderModell!wlbh iS Nbhwascalculatedto developsomeidea
obtainedby sequentially testing(through ofthe biasinvolved.Resultsindicatethat
goodness of fit tests) for differences relativebiasrangesfroln3 to 38 percent
amongthe pj. The processbegins with forpopulationsused in the simulations.
testingwhetherornotallthepjareequal. Thatrangeis somewhatlmisleading how-
If not, we let P1 be differentand test ever,becauseforall populationsexcept
whetheror not P2= P3= *-- = Pt. This one (in which half the populationwas
testing continuesuntil it is concluded essentially untrappable),the range of
thatthe last t- k + 1 captureprobabili- biaswas3 to 15percent.Someexaluples
ties arenotsignificantly different,where- aregiven in Table10. The readeris re-
as the firstk captureprobabilities do dif- ferredto TablesN.6.aand N.6.bof Ap-
fer.In the case k= 1, we aresayingthe pendixN forfurtherresultsand for de-
simpleModel/Ib (allpjequal)adequate- scriptionsof the populations. In general,
ly fitstheremovals.Fork > 1 we arecon- it seemsthereis no seriousbiasin Nbhif
cludingNlodelMb does not fit the data, relativelyfewmelubersofthepopulation
due to the presenceof heterogeneity (or are essentiallyuncatchable(i.e., proba-
tiInevariationin captureprobabilities).bilityof firstcaptureless than0.05)and
The estimatorNbhis the ML estimator the numberof trappingoccasionsis ad-
underthe selectedmodel. equate.(Recallthatsincethe estiluation
The abovetechnique,calledthe gen- techniquedepends on removalsonly,
eralizedremovalmethod,is furtherde- probabilities of recapturehaveno effect
scribedin AppendixH. The appendix on the perforlmance of Nbh ) Considering
alsopointsoutthatit is notnecessarythat the complicated modelstructure andthe
the factorizationG(p, c; 0)= Gl(p; 81) assumptionsrequiredto produce the
TABLE 10.-COMPUTER SIMULATIONOF THE BIAS vide reasonableestimatesof population

ASSOCIATEDWITH THE ESTIMATORNbh FOR DATA
GENERATEDUNDER MODEL Mbh (ALSO SEE AP-
size if the numberof unmarked animals
PENDIXN, TABLE N.6.b) capturedon each trappingoccasionex-
hibitsa definitedecreaseovertime.Such
Percent
relative
Popula- Number Number
tion of of
a trendindicatesthatconditionalproba-
bias
RB
^ size
Ave[Nbhl N
reps occasions
R t Trials'
bilitiesof captureon thejth occasionare
reasonablylarge,which is the primary
-38.4 246.6 400 100 5 1 requirementfor adequateestimationof
- 14.8
- 5.9
340.8
94.1
400
100
100
100
5
5
2
7
populationsize.
- 12.3 175.5 200 100 10 5
- 3.2 193.7 200 100 7 6 Example
- 4.3 383.0 400 100 5 4
l The probability E. Larsen(pers.comm.)providedthe
of capturefordatageneratedunderModelMbh
resultsof livetrapping
is complex;therefore,the specificvaluesare given in Appendix
N, TableN.6.a.
GreatBasinpocket
mice Perognatht4s parsus in a desert
community,CurlewValley, Utah.The
generalizedremovalestimator,these re- areawastrappedfor7 consecutivenights
sults are encouragingwith respect to duringJune 1977.Trapswere arranged
practicaluse of the estimator. in a 12 x 12 gridandspaced15 m apart.
Analysisof theresultingdataby themod-
ConfidenceIntervals el selection procedure(described in
TESTS OF MODEL AssuM7rIoNs) indicat-
Simulationresultsare not so encour- ed thateitherModelMtbh or ModelMbh
aging,however,whenit comestoplacing wouldbe appropriate foruse in estimat-
a confidenceintervalon N. Thevariance ing N. The dataon firstcaptureandthe
estimatorassociatedwithNbhappearsill resultsof usingNbhto producepointand
behavedand the distributionof Nbhis intervalestimatesforN aregivenin Fig.
nonnormal. Thosefactorsresultedin sim- 8. Note that the estimatorrejectsthe
ulatedconfidencecoefficientsaveraging modelfork= 1, but acceptsk= 2. Also
approximately 0.50.Furtherworkis nec- of interestis thatthe estimationmethod
essaryto developusefulconfidencein- failsfork = 3, butestimatesfork = 4 and
tervalsfor N when Model Mbh seems k= 5 are producedthatappearreason-
appropriate because normal theory able. The estimatedvarianceof Nbhis
confidenceintervalsapparentlyare in- quitelargefork = 2. Possiblytimevari-
appropriate forpracticaluse. ationis quiteimportant and causesthis
imprecision. Theprobabilities of capture
Robustness decline for k= 2, as they should under
ModelMbh. A histogram of the Ujvalues
Thegeneralizedremovalestimator can is includedto help the researcher visu-
be expectedto be robustto ModelsMh allyexaminehis data.
and Mb,since both are specialcases of
ModelMbh.If captureprobabilitiesex- Discussion
hibit significantvariationovertime,the
utilityof the methodis undoubtedly de- ModelMbhis one of the mostrealistic
creased.Thatis particularly trueif prob- and useful modelsfor a capture-recap-
abilitiestend to increasewith time, al- tureexperiment consideredin thismono-
thoughthe failurecriterionshouldhelp graph. Realismresults from allowing
to providea safeguard againstuse of the every animalto possessits own proba-
methodwhenit is inappropriate. In gen- bility of firstcaptureand fromnot con-
eral,it is expectedthatthe generalized strainingthe membersof the population
removalmethodused in the contextof a so thateveryindividual'sbehavioralre-
capture-recapture experimentwill pro- sponseto firstcaptureis identical.The
FROMCAPTUREDATAtiS
INFERENCE
OCCASI Jw 1 2 3 4 5 6 7
TOTALCAUGHT MIJ ) s 0 23 32 35 40 46 52 55
CAlJGHT U1 ) w
NEl^LY 23 9 3 5 6 6 3
K N-HAT i(N} CHI -sa . Psos . ESTI MATEDP-3AR ( J ), Jw I, . . ., 7
60 . 6 1 W * 563 - 9. 965 . 0762 .2802 .2802 .2802 .2802 . 280Z .2802 .2802
2 76.73 2W.66909 3,F .4B01 .2996 .1377 . 1377 . 1377 . 1377 . 1377 . 1377
3 FAILURE CRITERIOEJ- -2 N0 EST I MATESFOR THI S srEp .
67.W 18. 19 I .308 .S201 .3410 .2025 .0846 .2067 .2067 .2067 .2067
5 58 . 65 5 +8967E15 .823 .3643 . 3922 .2525 .1 126 .2114 .3953 .3953 .3953
POPUIAT1 EST I HATE I S 77 W1TH STAZA E9 24 . 6691
APPR()XI MTE 95 PERCENTCONFI DEN(DEI NTERVAL 28 TO 126
HI STOGRAMOF U ( J )
FREQUENCY 23 9 3 5 6 6 3
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
EACH * EWKS 3 w I NTS
21 *
18
15
12 *
9 t *
6 * * * * *
3 * * * * * * v
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
FIG. 8. Exampleof populationestimationwith variableprobabilityremovalestimatorwInderModel

Mbhwith pocketmousedatafromE. Larsen(pers.colllm.).
luodel is useful because an estimation in ModelMtbh, everyothermodelconsid-

procedurebased on the assumptionsof ered here is of coursea specialcase of
the model is availableforestimatingpop- thismodel.Thusit maybe usefulto think
ulationsize. Althoughthe operatingchar- of ModelMtbh as the mostrealisticof all
acteristicsof the estimatorare not com- modelsthatassumepopulationclosure.
pletely satisfactory,the estimatorwould Unfortunately, this extremerealismpre-
seem superiorto any otherthat mightbe cludesthe modelfromhavinganyprac-
used if, in factthe assumptionsof Model ticaluse in the estimationof population
Mbhare satisfied.Moreover,the method size.If the experimenter is led to believe
is nonparametricin the sense that one thatalltheassumptions of ModelMtbh are
requiredfor the capture-recapture
does not have to specifya probabilitydis- ex-
tributionfor the pairs(Pi,ci). perimentat hand,none of the models
consideredin this publicationare capa-
MODEL CAPTURE PROBABILITIES
ble of producingvalid statisticalinfor-
Mtbh:
mation concerning thepopulation param-
VARYBYBEHAVIORAL RESPONSE etersof interest.If sucha failureoccurs,
TO CA7ruRE,TIME,AND the researcher is forcedto reevaluatethe
INDIVIDUAL ANIMAL experimental designto discernhow the
Discussion experiment mightbe conductedto obtain
useful information.This reevaluation
For purposesof this monographMod- maycausethe capture-recapture tnethod
el NItbhis useful only conceptualiy.Be- to be discardedin favorof solne other
cause all 3 factorsthatmayaffectcapture completelydifferenttechnique,e.g.,line
probabilitiesareassulnedto be operating transectmethods.Whateverthe conclu-
44 MONOGRAPHS
WILDLIFE
sions,such an approachis much pre- condition that units of effortact inde-

ferredover one that ignoresnecessary pendently, a numberof estimationtech-
assumptions associatedwith the appro- niques have been proposed.Although
capture-recapture
priate modelandpro- maximum likelihoodestimationof N is
ceedsto use some simpler but inappro- possible (cf. Seber 1973:297),the best
priate model. This latter approach known estimation techniquesare the
requiresthe researcher to give credence regression techniquesproposedby Les-
tostatisticallyinvalidinformation about lieandDavis(1939),DeLury(1947),and
thepopulation, andtherefore ba-
violates Ricker (1975).All thosemethodsinvolve
sictenetsof scientificresearch. regressing catchper unit effortagainst
somefunctionof the "catchability coef-
ficient" and the efforts expended, and
REMOVALMODELS squares analysisto pro-
thenusingleast
Introduction videan estimatorof N and its variance.
In a capture-recapture experiment, all Suchcatch-efforttechniqueswill not
ourapproach toremoval
lnembers of the populationcapturedon beconsideredin for a number of reasons.
agiventrapping occasionaremarkedand experimentation
and most important, we restrictour
releasedbackinto the population.In a First, consideration of removal studiesto those
removalexperiment,however, as the inwhicheffortis deliberatelykeptcon-
nalneimplies,membersof the popula- stant onalloccasions.In suchstudies,the
tionare permanentlyremovedand are concept of efforthas no utility.Second,
notreintroduced intothepopulation after
firstcapture.Theremovalmaybe accom- because the general approachin this
monograph has been to void paraInetric
plishedby killtrapping,electrofishing, assumptions concerningcaptureproba-
trawling, or merelylivetrapping the ani-
malsand physicallydisplacingthem to bilities, the assumptionthat pj= 1-
exp(-kej) is inappropriate for our pur-
another area. Third, it is necessary forpurposes
poses.
Removalmethodshave traditionally of leastsquaresanalysisto makecertain
beenassociatedwith catch-effort exper-
iments,whichhave been used oftenin assumptions that involve the variance
of the observations. Becauseto
practice(Omand1951, Ketchen1953, structure each postulated variance structure there
Fischler 1965). Such experimentsare corresponds a formulaby which the es-
basedontheideathatthe sizeofthepop- timateof N is calculated,such proce-
ulationwill be decreasedgraduallyas duresareto someextentarbitrary in the
will the catchperuniteffortby applying absence of valid tests for determining
a knownamountof removalefforton a propervariancestructure.Finally,Braa-
nuluberof occasions.The basicassump- ten (1969)investigated the robustnessof
tion behind catch-efforttechniquesis the DeLury(1958)estimatorto depar-
thatthenuluberofanimalsremovedfrom turesfromsoIneof the assumptions and
thepopulation directlydependsuponthe exhibit sig-
amountof effortexpendedin removing found thatthe estiluatormay
bias.
theIn.By usingthe size of the successive nificantnegative
decreasesandthe known efforts that ef-
fectedthosedecreases,initialpopulation Structureand Use ofModel the Generalized
size N canbe estimated.Theexperiment Removal
usuallyis modeledby assumingthatall
animalshaveprobability of capturepj = If it is assumedthatsaInplingeffortis
1- exp(-kej) on the jth trappingocca- constant over trappingoccasions,and
sion,wherek is oftencalledthe "Poisson that all animals in the populationhave
catchability and
coefficient" ej represents the same probability of removal,thenthe
the knowneffortexpendedon thejth OC- removalmodel and estimationmethod
casion.Undersuch conditionsand the considered by Moran (1951)results.That
STATISTICAL INFERENCE FROM CAPTURE DATAtis et al. 45
method was discussed in conjunction models.Such a procedurerepresentsa

with estimationin ModelMb.The con- generalization (1951)andZip-
of Moran's
texthere,however,is differentsincean- pin's(1956,1958)constantprobability re-
imalsarenot reintroduced intothe pop- movalmodelto the case in whichheter-
ulationafterinitialcapture.Zippin(1956) ogeneousprobability of removalexists.If
showed that the joint distributionof the set of removalprobabilitiesPi, i=
. . .,ut}, where u; representsthe 1,2,...,N, is assumedto be a random
{u1,u2,
numberof animalsremovedon thejthoc- sampleof size N fromsomeprobability
casion,canbe writtenas distributionG(p;@)parameterized by o
P[{ul,u2,...,ut}] =
and definedon [O,1], then the distri-
bution of the vector of removals
N! pM,+, {u1,u2,...,ut}canbe writtenas
[II uj!](N- Mt+l)! P[{UI,U2,@@,Ut}] =
tl!J- E (t - j + l)uj
NE
*(1-p) 1=1 t { E[p]}U.
{E[(1 - p)p]}u2
[Iluj!](N- Mt+l)!
wherep is the probability of removalof i=l
any animalon any trappingoccasion. *. . {E[(1_ p)t _ p)t}N-Mt+,
lp]}Ut {E[(1
Becausethe parameterspace is 2 di-

mensional(N andp arethe onlyparam- where
eters)andthe minimalsufficientstatistic rl
E[(1 _ p)j-lp]= | (1 _ p)j-1pdG(p;o),
J()
{ t+l,JEj i}
j= l,...,t,
is 2 dimensional,both parametersare E[(1_ p)t] = 1 - E E[(1_ p)j-lp].
identifiableand may be estimatedby j=l
maximum likelihood.Suchestimators are
derivedin AppendixD, andarethe same This distribution is identicalto the dis-
as those used to estimateN and p in tributionof the removals{ul,u2,...,ut}
ModelMb underthe conditionsdiscussedin the
An estimatorof populationsize N sectionon ModelMbh.Thus,it is easily
basedon the abovemodelwill clearlybe argued(cf.AppendixJ) thatthe general-
unsatisfactory if all animalspresentdo izedremovallnethoddevelopedforMod-
nothavean equalprobability of removal el Mt,his alsoappropriate forestimation
on a giventrappingoccasion.In removal in removallnodelsin whichit is assumed
experilnents, unequalcaptureprobabili- thatheterogeneity of firstcaptureproba-
ties can be causedby tiInevariationor bility exists. Details of this estiluation
heterogeneityor both.Behavioralvaria- technique are given in AppendixH,
tionis nonexistentsincelnelnbersof the alongwithanestiluator of theasylnptotic
populationare removedafterfirstcap- varianceof the populationestilnatorNR
ture.We maintainthatproperplanning (theR standsfor"removal," butwe note
and design of the reluovalexperiment that NR is luathematically the same as
can be used to controlor reducetilne Nbh)and a "failure"criterionthatInust
variation(see STUDY DESIGN). Thus,the not hold if parameters areto be validly
mainproblelnlies in dealingwith het- estimated.This criterionensuresthata
erogeneityof capture(removal) probabil- sufficientdecline in the populationis
ities. In the followingdevelopment,it is beingeffectedby the successiveremov-
arguedthattheestimation procedure pre- als.
viouslyproposedfor ModelMbhis also As explainedin the materialon esti-
appropriatefor estimationin removal uationof N in the sectionon ModelMhh,
ourprocedureis to lookat a sequenceof thatthissectiondealswithstrictremoval

removalmodels.These correspondto a data.
sequenceof assumptions as follows:all Robustness
pj are equal, or P17&P2 but P2= P3
= = Pt, or P17&P2 + p3, but p3 = p4 In the contextof removalexperiments,
= = Pt,andsoforth.Themostgeneral the generalizedremovalmethodrepre-
modelallowsall pj to be different,but sents a significantstep forwardwith re-
this mostgeneralmodeldoes not allow spectto robustestimationof population
estimationof N. We definethe specific size. It shouldbe emphasized, however,
removal model MRkas the model in thatthe procedureis designedto be ro-
whichthe last t - k + 1 valuesof pj are bustto failureof the assumption thatall
the same,andfork > 1, the firstP1to Pk animalshavethe sameprobability of first
aredifferent. capture,andnotto failureof the assulup-
The estimationprocedureis to testthe tionthatsamplingeffortis uniformover
goodnessof fitof the removalmodelsse- trappingoccasions.Thereis some indi-
quentiallyfromMR1to MRt_2. Thus,we cationin the simulationresultsthatthe
are first testing whetherthe constant methodperforms adequatelywhenthere
probability modelfits.If it does,we use is noheterogeneity of captureprobability
the corresponding maximumlikelihood but there is nonuniformity in sampling
estimatorof N. If this simplemodel is rates over time. The methodperforms
rejectedby the chi-squaretest (at the verypoorly,however,when bothheter-
20%significancelevel in programCAP- ogeneityandnonuniform samplingrates
TURE),we thenexaminethe case(mod- are present.First,the percentageof ex-
el MR2) of P17&P2, but P2 = P3 = = Pt periments which"fail,"as determined by
The estimatorof N, NR, usedin thisgen- the failurecriterion,can be very high.
eralizedremovalapproach is takenas the For instance,in the populationdefined
ML estimatorof N for the selectedre- by Trial1 of ModelMth, the experiment
movalmodel. failed in every one of 100 simulations.
Secondly, when the experimentdoes
Simulation Results succeed,biasis usuallysignificant. Some
BiasandConfidenceIntervals simulated examples are given in Table 11
whichhelpto substantiate theseremarks.
A discussionofthebiasoftheestimator Completesimulationresultsaregivenin
of N associatedwith the generalizedre- TableN.6.bof AppendixN.
movalmethodandof theusefulnessof its
associatedconfidenceintervalswas pre- Example
sented in the materialon Model Mbh.
Briefly,the simulationstudy revealed Andrzejewski and Jezierski(1966)re-
thatthe biasof the estimatorof N wasin portedthe resultsof a studydesignedto
most cases not serious,but that confi- estimatepopulationdensityof European
denceintervalsachievedonly50 percent hareLepus europaeus on experimental
coverageon the average.Althoughthe huntinggroundsin Poland.Hareswere
results and discussionof that section capturedand removedby drivingthem
were presentedin the contextof a cap- intonetssurrounding the area.Resultsof
ture-recapture experiment, all the mate- the applicationof thegeneralizedremov-
rialis directlyapplicablehere. Because al methodto the dataresultingfromthe
the2 estimation procedures areidentical, 4 drives(removaloccasions)aregivenin
the relevantdatain both cases involve Fig. 9. Whenk= 1 (i.e., all animalsare
only the removalstatisticsfrompopula- assumedto havethe salneprobability of
tionswithheterogeneous probabilities of removal),a poor fit to the dataresults
firstcapture.Denotingthe estimator of N (chi-square goodnessof fit valueis 13.5
as NRis intendedto remindthe reader 2 df, and p = 0.0014).For k = 2, how-
STATISTICAL
INFERENCE
OCCASI ON 1 2 3 4
TOTAL CAUGHT 0 722 913 982 1018
NEWLY CAUGHT 722 19 1 69 36
K N-HAT i (N) CHI -SQ . PROB. ESTIMATEDP-BAR( J 3, J= 1, . . ., 4
1 1028.21 3*789703 13.150 .001W .6806 .6806 .6806 .6806

2 1039.10 7.658751 1.528 .2164 .6948 .5916 .5916 .5916
POPULATI ON EST I MATE 15 1039 WI TH STANDARO ERROR 7.6588
APPROX
IMATE95 PERCENTCONFI DEKE INTERVAL 1023 TO 1055
HISTAM OF U(J)
FREt)UENCY 722 191 69 36

___________________________________
EACH ' EQlJALS 73 POINTS
730
657
58q
511
438
365
2g2
219
146
73
FIG. 9. Exampleof populationestimationunderthe variableprobabilityremovalInodelwith clataon
EuropeanharefromAndrzejewski andJezierski(1966).
ever,anacceptable fitresults(p = 0.2164) NR= 1,039+ 7.66 seems muchprefera-

and,therefore,the pointandintervales- ble to the estimateof 1,010thatresults
timatescorresponding to ModelMR2 are from the (commonlyused) regression
chosen.In particular,the pointestimate methodused by Andrzejewski and Je-
TABLE 1 1 .-COMPUTER SIMULATIONRESULTSILLUSTR TING THE ROBUSTNESSOF THE GENERALIZED

REMOVALESTIMATORNR USING DATA GENERATEDUNDER OTHER MODELS (ALSO SEE APPENDIX N,
TABLES N.2.b, AND N.5.b)

RB Ave[NR] N R t Data model
- 3.2 387.1 400 100 10 Mt,Trial 6

- 9.0 364.1 400 100 7 Mt,Trial 7
2.6 410.3 400 100 7 Mt,Trial 9
- 0.2 199.7 200 100 5 Mt,Trial 10
- 1.9 272.6 400 100 5 Mth,Trial 2
22.3 489.1 400 100 5 Mth,Trial 4
-10.3 179.4 200 100 7 Mth,Trial 5
I For example, data generated under Model Mt were used to estimate N, using the generalized removalmethod NR.
OCCASIObJ Js 1 2 3 4 5
TOTAL CAUGHT M(J)s 0 181 192 196 201 204
NElPlLYCAUGHT U{J)s 181 11 4 5 3
K N-HAT i (N) CHI -SQ . PROB. EST I MATEDP-2AR ( J ), J= 1,, . ., 5
1 204 .00 .2094765 70 .820 0 . 0000 .8160 .8160 .8160 .8160 .8160
2 206.77 3.697937 I .555 .4596 .8754 . 4029 . 4029 .4029 . 4029
3 208 . 43 8 . 365668 1.021 .3124 .8684 .40 10 . 3306 . 3306 . 3306
I ON EST I MATE I S
POF>ULAT 207 W1TH STANDARDERROR 3.6979
IMATE95 KRCENTCOS IDEKE I NTERVAL

APPROX 199 T0 215
HISTOGRAMOf U(J)
FREQUENCY 18 1 11 4 5 3
________________________________________
EACH ' EXALS 19 w I NTS
190 *
171 *
152 *
133 *
1 14 *
95 *
76
57 *
38 *
19 * *
________________________________________
FIG. 10. Exampleof populationestimationunderthe variableprobabilityremovalmodel with aquatic

insectdatafromR. F. Raleigh(pers.comm.).
zierski,becausethenumberofharesactu- generalizedremovalestimationproce-
allyremoved was1,018.Theestimated cap- durewas used to producepointand in-
tureprobabilitiesforModelMR2 areP1= tervalestimatesforN (Fig.10).As in the
0.6948,and P-2=p3= p4= 0.5916,a re- previousexample,thereis a poorfit for
sult that supportsthe idea that hetero- the simplemodelwith constantcapture
geneityis operating. probability(k= 1). When k= 2, how-
ever,a goodfitto the dataresultsandan
Example estimate NR= 207, correspondingto
ModelMR2 is produced.Notethe differ-
R. F. Raleigh(pers.comm.)provided ence between P1 = 0.8754 and p=
someresultsof a removalexperimentin- 0.4029,indicatingthata significanthet-
volvinga speciesof mayflyEphemerop- erogeneityin removalprobabilities may
weresampledfrom10ran-
tera. Nlayflies exist.The computedconfidenceinterval
domlyplaced0.25-m2areasin a section is somewhatunsatisfactory becauseits
of the PoudreRiverstreambed nearFort lowerlimitis less thanthe numberof an-
Collins,Colorado,with S removalocca- imalsactuallyseen,andremindsonethat
sions at each site. A special benthic in removalandlivetrapping experiments,
aquaticsamplerwas used in the study. conditionsnecessaryfor construction of
Becausethesearetrueremovaldata,the normaltheoryconfidenceintervalsare
INFERENCE et al. 49
OCCASION Jw 1 2 3 q 5 6 7
TOTALCAU()HT M( J ) z 0 25 51 66 79 91 104 109
NEWLYCAUGHT U(J)- 25 ^ 15 13 12 13 5
K N-HAT SE (N) CHI -sa ^ PROB. EST I MATEDP-BAR ( J ), J= 1, * . ., 7
1 138.07 14.67g3 3.a79 .5670 . 19B0 . 1980 . 1980 . 1980 . 1980 .1980 . IgEO
2 132.46 13.6W 3.623 .b594 I 887 .221B .2Z18 .2218 .2Z18 .2218 .2218
3 141.63 25.85191 2.754 .q31 1 . 1765 .2229 . 1829 . 1829 . 16Z9 . 18Zg . 1829
4 130 .96 20 a I W541 2 . 688 .2608 1909 .2454 . 1876 .2339 2339 t 2339 .2339
5 1 IB.54 10.21073 2. 133 . 1Wb2 .2109 . Z779 . 2Z2 1 .2474 3675 .3675 .3675
POPUIATI ON EST I HATE I S 138 W I TH STAZARO ERROR 1b . 6799
APPROXI MATE95 PERCENTC I DENCE I NTERVAL l o9 To 167
HISTOGRAJ1
OF U(J)
FREaJENCY 25 26 15 13 12 13 5
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
EACH * EWALS 3 POINTS
27
*
29
*
21
* *
18
* * *
15
* + * * * *
12
* * * * * §
9
* * * * * * *
6
* v * * * * *
3
_ _ - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - -
_ _ _ _ _ _ _ _ _ _ _ _
FIG. 1l. Exampleof populationestilnationunderthe variableprobabilityreluval z1l>delwith whitefish

dataioln Ricker(1958.150).
notsatisfied.Inthiscase,oneshouldtrun- sion methodassumingequaleffort.(We

catethe lowerliInitto the actualnuluberwish to reemphasizethaton theoretical
removedratherthanusinga lowerliluit groundsthe MLestimationprocedureis
less thanMt+1. superiorto the variousregressiontech-
niquesoftenusedwithremovalstudies.)
Example The 95 percentconfidenceintervalesti-
xuateof [109, 167] is very informative.
Ricker(1958:15())removedwhitefish NIoreover7 the true confidencelevel of
Coregonus clupeafornis frozmShake- theintervalprobably is closetothestated
speareIslandLakein OntarioS Canada, level of 0.95,sincethe estiluatecl proba-
on 7 successiveoccasionsby meansof bilityof removalis nearly0.20.
gillnetting. NleInbersof whitefish re-
uovedin the 13-to 14-inch(3.3-3.6cln) Discussion
lengthclass are shownin Fig. 11, with
the resultsof the generalizedremoval Conductinga relnovalexperilnentfor
snethod.Noticethata goodfit to the re- purposesof estimatingpopulationsize
movalsis achievedfork = 1, so thatZip- maysozuetirlles provemorefeasiblethan
pins (1956,1958)constantprobabilityre- a capture-recapture approach(referto
mova model seems appropriatefor STUDY DESIGNforfurtherdiscussion) . In
estimatingN. Thepopulationestimateof such cases, the experilnenter has avail-
NR= 138 is in close agreementwith able 2 classesof estiluation proceclures,
Ricker7s estimateof 136 whichwas ob- the catch-effort techniquesusuallyasso-
tainedby usingDeLury's(1947)regres- ciatedwith LeslieandDavis(1939)and
o WILDLIFE MONOGRAPHS
DeLury(1947)or the "removal"tech- (i.e., assumptions) possibleforthe given

niquesfirstintroducedby Moran(1951) data.Thena seriesof specificalternative
refinedby Zippin(195671958),andgen- modelsshouldbe developedbased on
eralizedhere. It is felt thatcatch-effort specificassumptions thatcoverthe var-
techniques are often not appropriate, iouscasesof realisticortheoreticalinter-
either because of the assumptionsin- est. Those alternativemodelsformthe
volvedor becausethe conceptof effort basisforbothtestingof assumptions and
uaybe meaninglessin manyexperimen- estimatingparameters.Under FUNDA-
tal situations.In those frequentsitua- MENTALCONCEPrS,we gavethe general
tions,we believethe generalizedremov- modelframework consideredhere:pop-
al methodprovidesthe best approach to ulationclosureis assumedand captures
estimatingpopulationsize. The operat- are assumedto be independentevents
ing characteristicsof this methodareby withcaptureprobabilities
no means completely satisfactory,in PiJ,j= l,...,t,i= 1,...,N.
view of the results concerningconfi-
dence intervalcoverageand the failure Sucha luodelstructureis usefulonlyas
of the experimentin some situations. a conceptualstartingpointbecausenone
However,the factthatthe methodfails of theparameters areestimable.Next,we
in a givenexperimentdoes at leastpre- recognized3 differentsourcesof varia-
vent the use of wildly inaccurateesti- tion actingon these captureprobabili-
matesin practiceandhelpsto informthe ties:(1)variation overtirne,(2)behavior-
experimenterthat the assumptionsof al variation asa resultoffirstcapture(trap
the methodare not met and the quality response)and(3)variationoverindivid-
of the experilnentneedsto be upgraded uals(heterogeneity). Wealsorecognized
or the designaltered,orboth.Neverthe- the various possible combinationsof
less, the methodis the mostgeneralnow thesesourcesof variation.
availablein the literaturethatis capable An almostinfinitevarietyof veryspe-
of providingusefulresultsand withfur- cific modelscould be constructed.We
therresearch,improvements in themeth- examined8 modelslikelyto be generally
od shouldbe forthcozning. usefillforestimationortestingpurposes.
Those modelscan all be expressedin
TESTS OF MODELASSUMPTIONS termsof the assumptions aboutvariation
In precedingsections,we haverecog- in the capture probabilities.If eachof the
nized 3 distinctsourcesof variationin 8 luodelsintroduced hadits ownunique
captureprobabilities, and have given 8 estimationprocedure,then selectionof
modelscorresponding to presenceorab- an estimatorwouldbe synonymous with
sence of specifictypesof variability.No selection of a model. Suchis not the case
estimatorof population size hasbeende- with only 5 estimators and8 models. Our
rivedfrom3 of those models(Mth)Mtb, strategyis to presenta testingsequence
Mtbh). Consequently, we haveintroduced designedto leadto selectionof the most
only 5 estimatorsthatare identifiedbe- appropriate model.Wedefine"appropri-
low, alongwith the modelswherethey ate model" as the siluplestmodel that
shouldbe used.Themainpurposeof this "fits'the data.Whenthatselectiongives
sectionis to give a strategyforselecting a modelsuchas Mthforwhichthereis no
one of the estiluationproceduresbased theoreticallyappropriateestimator,we
on tests of the variousunderlyingas- recommend (insomeinstances)usingthe
sumptions. nextbest fittingmodelforwhichan es-
timatorexists.It will occasionally be nec-
Philosophyof the A pproach essary to conclude thatno estimate of N
can legitimatelybe calculatedfroznthe
Ourphilosophyis thatone shouldfirst data.
presentthe mostgeneralmodelstructure It is important to recognizethatwe do
INFERENCE et al. 51
notclaimthismodelselectionprocedure below,in termsof assumptions andcor-

resultsin the "true"model.Bytruemod- respondingcaptureprobabilities.
el fora capturestudywe meanthatthe MODELMo Pu-P
statedmodelis anexactrepresentation of
the truecaptureprobability structurefor Colument:This is the tnostrestrictive
thatstudy.Anymodelis a simplification uodelbecausecaptureprobabilities
thatcan realisticallyrepresentonly the are assumedto be constant.
morepredominant featuresof the study. Estimation: We recommendthe NIL
Also, we must recognize that the luost estiluator(see AppendixB); it does
appropriate modelwill dependuponthe notexistin closedformexceptfort -
amountof datawe have.Withgooddata 2.
(largegrids5manyoccasions)one might MODELi\lt Pii--pj5 t
be ableto show,forexampleS thatModel j = 1> * * *
Mbh wasnecessary.Yet if the samestudy Comment: Capture probabilities vary

used a much smallergrid, and fewer only with time. This is the luodel
days it mightbe impossibleto delnon- mostfrequentlyassumed(oftenim-
strateheterogeneity fromthe data them- plicitly)in the literature.
selvesandModelMbmightbe indicated Estimation:We recommendthe ML
as appropriate. estimator(see AppendixC), it does;
The conceptualgoal of ourluodelse- not exist in closedform,exceptfor
lection procedureis to achieve an ac- t- 2 (in which case it is the Lin-
ceptabletrade-offbetween precisionand coln estimator).The Schnabelesti-
bias.If one uses too simplea model the luator,or variations thereof,is often
estimatedpopulationsize is likelyto be used with this model, but is onlyan
severelybiasedyet have a deceptively approximation to the ML estiluator.
smallsamplingvariance.If the modelis
comples,but still the wrongone, then MODELMb:Pi -4 [p forfirstcapture
estimatorsareagainof uncertainbias.If J (Cforrecapture.
a modelis chosenthatis toogeneral,the ComInent: This is the simplestmodel
riskof bias is muchreduced,or negligi- of behavioral(trap)responseand it
ble but the samplingvarianceis larger hasbeenrecognizedin theliterature
thanit needs to be (i.e., we lose preci- (Tanaka1956).
sion).Of those2 casesythe saferone in Estimation: The MLestimator,which
ouropinionis to choosethemoregeneral is recommended (AppendixD)>is a
modelin hopesof minimizing biases.We special case of the generalizedre-
statethisas ourconceptualgoalbecause7 moval estimator.Zippin'sreluoval
notknowingthetruemodel we cannev- modelis equivalentto ModelMb for
er be certainwhen analyzingreal data purposesof estimatingN but his
thatwe have madethe best choiceof a suggestedestimatoris only a close
model. approximation to the true NILesti-
This model selection procedureis mator.
basedon testsof assumptions of 2 types:
(1)testsbetweenspecificmodelsand(2) MODELMh:Pii-Pi i - 1, . >N
generalgoodnessof fit tests to specific Comment:This modelallowshetero-
models.Beforediscussingthesetestswe geneityof captureprobabilities over
sutumarizethe primarymodelsand as- animalsS but allowsno variationin
sociatedestiluators. captureprobabilities overtime.
Summary of L1Wodels and Estimators Estimation: The jackknife estimator
(Appendix E) is recommended when
Capture-Recapture Models Mhis the underlyingmodel5thates-
Becausethese modelshave been de- timatorwas derivedspecificallyfor
scribedin detail,onlya summary is given thismodel.Note however,thatthere
is no way to derivean optimalesti- TABLE 12.-MODELS AND SUGGESTEDESTIMATED

matorfor Model Mhif nothingis PROCEDURES
knownabouthow the captureprob- Theoretically
abilitiesvary. Model appropriateesffmator
Mo Null (No)
epjforanyfirstcap- Mt Darroch (Nt)
MODEL Mtb:Pii-< ture j-1,. . .,tS Mb Zippin (Nb)
ci for any recapture Mh Jackknife (Nh)
> j = 2, . . .,t. Mtb (none)
Mth (none)
CommentThismodelallowsvariation Mbh Generalized removal (Nbh)
in captureprobabilities due to both Mtbh (none)
time and behavior(trapresponse).
Estimation:Populationsize N is not
estimableunderthis model.There- case in whichall 3 sourcesof varia-
fore thereis no satisfactory estima- tionoperate.
tion procedureassociatedwith this Estimation:Populationsize N is not
luodel. estimableunder this model using
straightforward methods.Therefore,
MODEL u Mth: PiJ-p@j j - 1>.. .,tSi - there is no satisfactory estimation
, ,>. procedureassociatedwiththis mod-
Cotnment: This is usefulas a concep- el.
tual modelof how time andhetero-
geneity mightoperateas a simple Relationships BetweenModels
product.Noestimator canbe derived
fromthismodel. Table 12 summarizessome of the
Estimation.Populationsize N is not above information. It is important to un-
estimable under this model if derstand the relationship of the models
straightforward methodsare used. to one anotherin orderto understand
Thereforethereis no satisfactory es- testingbetween models.One can only
timationprocedureassociatedwith truly test one model against another
this model whenone modelis a specialcase of the
other.ForexampleModelMois a special
MODELMbh: Pifor
Pii-v c, forall
first capture case
recaptures
of eachof the 3 modelsMt Mb,and
Mh.Butthose3 luodelsarenotrelatedto
i= 1S...SN. eachotherin anysimplemannerthatal-
Golument.This modelassumesheter- lows one to chooseortestbetweenthem
ogeneitywith trapresponseS but no with any simplestatisticaltests.
timevariationas suchin the capture The relationships amongthesemodels
probabilities. is shown in Fig. 12 usingan arrowbe-
Estimation:The generalizedremoval tween 2 models to indicate thatone is a
estimatorshouldperformwell here special case of the other. For exaInpley
(AppendixH). However if none of M) Mt meansModelMois a special
the generalizedremovalmodelsfits caseof ModelMtS becauseif ModelNIt iS
no estimatecan be producedusing restricted by the assumptionPl = Pz
this model. = = Pt= p ModelMoresults.
The selectionof an appropriate cap-
M°DEL Mtbh- p- 3 Pii forfirstcaptureS ture-recapture model is neither simple
* 1 <cij forall recaptures, norstraightforward andthereis, infact,no
j= l,...,ty optimalor rigorousstatisticaltheoryto
i- 1>...>N. guide this selection.The jumbleof ar-
Comment.This is a conceptually use- rowsin Fig. 12 is indicativeof the dif:fi-
ful modelbecauseit representsthe culty.
n .r rn r 21e t
*At O nOe1 iS imml 5pe1a1
STATISTICAL
INFERENCE
FROMCAPTUREDATAA)tiS et 1. rYn
5 pectpc 1 ests to reryorm Mtbh
Betorewe can glve our strategyfor

r
/
tnodelShence estimatorS selectionlit is \
necessaryto disoussexactlysvhathypoth- \
esis can be testedconcerningvariations \K
in captureprobabilities.Two types of
testsaremade. ih Mbh
Mtb XMt
(1) Specifictests of on@ tnodelversusa
luore generalInodel.For exampleS
testingModelMovs. ModelMttexts
\/
for time variationin averagedaily /\
captureprobabilities; X Mh
(2) Generalgoodnessof fittestsof a giv- Mt M
en modelto the data.ForexampleS
testing whetherModel Mt fits the
dataamountsto testingwhethertime
rariationis the only sourceof varia-
tionin captureprobabilities.
M
Thereis a basicdifferencein the nature FIG. 12. RlM10nSh1PS AINQNg loetS. ff()wS i-
of the hypothesesassociatedwith those tase nf an>ter In>det
2 differentclassesof tests.The objective
of the firstclass of tests is to establish
whetheror not the moregeneralInodel
producesa slgnificantly betterfit to the
datathanthe Inorerestrictivemodel.On basisto believe thereis anyheterogeneity
the otherhandSthe goodnessof fit tests of captureprota zi ities.
ofthesecondclasstestwhetherornotthe the siluple tests for behav-
SiluilarlyS
datamightreasonablyhave arisenfroln ioral variation(Model Ml)vs. Model MhS

the specifiedlnodel. test 2) and for tizuevariatigxn (Model M>
Table 13 gives the 7 specifictests we vs. Moc e >Mty test 3) are not significant
in
haveused in the nlodelselectionproce- .iog t *
rn nloSeS * *n
tt!itS gIat. Slglllhtnte
dure.The readershouldstudyTable13 levels of 0.98056and().31935> respective-
beforeproceedirlg.ProgralnCAPTURE ly. Basedon only these results we would
hasbeen used to producean exalupleof (correctlySin this case) conclude the ap-
thetests.iResultsarepresentedin Fig 143>propriatemodel is Model M(J. When a
that is basetl on simulateddataunder InodelInorecoznplexthan M>is the ap-
ModelMOwithparameters N -lOOS p- propriateone7additiorlaltestsareneeded.
0.50>and t- 5. The first3 tests of the In the second partof the Inodelselec-
Inodelselectionprocedure separately test tion procedureS4 luore tests are pre-
each of the silupletnodelsMh MhS Mt sented. The first 3 are simple goodness
against Model M. These are all chi- of fit tests to lnodelsMhSM)>and Mt.The
squaretests(see AppendixKfordetails). seventh is a test for behavioralresponse
Thesignificance levelofthetestsis given in the presence of heterogeneityshenee
by the prograIn.ForexampleS in Fig. 13 this test helps makea choice between Mb
underthe testof ModelMl}vs. ModelMh and Mbh.That last test is included here
(test1) we havea chi-square test statistic because we lave an estilnatorfor Model
of 5.007(3 degreesof freedom)and the lvlbh
significancelevel (labeledPROBABILITY The goodness of fit test of Model Mh

OF LARGER VALUE) is 0.17127. ThusSthat (test 4) in Fig. 13 results in a chi-square
test is not significant7and we have no value of 4.S56(4 dfl andhasa significance
OCCASION Js 1 2 3 4 5
ANI MAL5 CAUGHT N{ J ) s 55 47 54 60 W8
TOTAL CAUGHT M( J ) w 0 55 75 88 96 98
NERLY CAUGHT U( J ) w 55 20 13 8 2
FREOUENCI ES F( J )- IO 3a 26 20 4
I . TEST FOR HETEROGENEI TY OF TRAPPI NG PROBA8I L I T I ES I N POPULATI ON.

NULL HYPOTHESIS OF MOOCL n(o ) vs . ALTERNATEHYPOTHESI S OF MODEL M ( H)
CHI-SOUARE VALUE w 5.007 DEGREES OF FREEDOM- 3 PROBABILITY OF LARGER VALUE = . 17127
2. TEST F9R BEHAV19RAL RES AFTER INITIAL CAPTURE.

NULL HYWTHESI S OF L M( O) VS . ALTERNATEHYPOTHESI S OF MODEL M( 8 )
CHI -SaJARE VALUE - . 001 DEGREES OF FREED()M- 1 PROBA2I L I TY OF LARGER VALUE = . 98056
3. TEST FOR TIME SPECIFIC VARIATI IN TRWPIK ASILITIE5.

NtLL HYPOTHESIS OF MOOEL Ml9) VS. ALTERMTE HYWTHESIS OF MODEL M(T)
CHI -SQUARE YALW - w . 791 DEZEES OF FREENM w 4 PROBABI L I TY OF LARGER VALUE = . 31935
4. GOaDJESS 6 FIT TEST 9F MOOEL MIH)

IULL HYPOTHESIS 9F MOOEL MIH) VS. ALTETE HY THE5IS 9F NOT EL M(H)
CHI -SQUARE VALX - 4 .59 DEES OF FREE - 4 PROBABI L I TY OF LARGER VALUE = . 3360 1
TEST OF MOOELM( H) BY FREKY OF CAPTURE

(FRE£NCIES LE55 THAN 2T ARE NOT CALCULATED.)
ER OF CAPTURES CHI -SWARE D. F . PR%AB I L I TY

_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
1 5. 000 4 .28730
2 1.29f3 4 .B6167
3 I.949 4 .74519
4 5.500 4 .23973
5 * GDCX)NESSOF F I T TEST OF MODEL M( B )

NULL HYPOTHESIS OF MODEL M(B) VS . ALTERNATEHYPOTHESIS OF NOT MODEL M(B)
CHI-SOUARE YALUE - 5^ IS3 DEGREES OF FREEDOMw 6 PROBABILITY OF LARGER VALUE = .5Z053
5A . CONTRI BUTI ON OF TE5T OF HOMOGENE

I TY OF F I RST CAPTURE PROBA8I L I TY ACROSS T I ME
CHI-SQUARE VALW w 2.459 X9EES OF FREEDOM- 3 PROBA8JLITY OF LARGER VALUE = .48276
58 . CATR I WT I ON OF TE5T OF H< I TY OF RECAPTUREPRO8ABI L I T I ES ACROSS T I ME
CHl-SQUARE VALW s 2.724 DEES OF FREEDOMw 3 PRO8ABILITY OF LARGER VALUE = .43610
6. (:iOOONESSOF FIT TEST OF MOOEL M(T3

MJLL HYPOTHES15 OF L M( T ) VS . ALTERNATEHY THES I S OF NOT M%EL M( T )
CHI -S(;IUARE VALUE - 59.465 OEZEES w FKE s 54 PROBABI L I TY OF LARGER VALUE = . Z8338
7. TEST FOR BEHAVIWAL REE IN WESEKE w HETEENEITY.

N1L HYPOTHE5I S OF PlODEL M( H1 VS . ALTERNATEHYPOTHESI S OF M:L M ( BH)
CHI-SaJARE VALUE - 7.471 DE9EES w FREE - 10 PR%ABILITY OF LARGER VALUE = .68036
SOEL 5ELECTION CRITERIA. MOOELS£LECTEO HAS SXI VAL.
EL HlO) M(H) M(S) Ht - ) M(T) M(TH) M(T8) M(TSH)

CRITERIA 1.00 .96 .2r; .51 0.00 .44 .33 .64
FROMCAPTUREDATA{)tiS et tzl.
INFERENCE
STATISTICAL 55
USEDIN THEMODEL
TABLE13.-SPECIFICTESTSOFASSUMPTIONS PROCEDURE
SELECTION
Test Source of variation

number tested for Null hypothesis Alternativehypothesis Comments
1. Heterogeneity Model Mofits Model Mhfits This test examinesthe capture

the data the data frequenciesto see if there is
evidence of variabilityamong
individualcaptureprobabilities
2. Trap response after Model Mnfits Model Mbfits This is a test for grossbehavior
first capture the data the data effects on captureprobabilities
3. Tlme varlatlon ln Model Mofits Model Mt fits This tests for variationin aver-
capture probabilities the data the data age daily captureprobabilities.
Unfortunately the actualtest is
also sensitiveto ModelMb
4. Trap response and/or Model Mhfits Model Mhfails to If Mhis the true model we ex-
time variation given the data fit the data pect this test not to reject.We
neterogeneity would also expect test 1 to re-
ject ModelM, in favorof Mh
4a. As above As above As above These tests are a specificform
of test 4, where for each sig-
nificantly large capture fre-
quencyan individualtestcanbe
made for trapresponseor time
variationor both
5. Heterogeneity andlor Model Mbfits ModelMbfailsto If Mbis the true model we ex-
time variation given the data fit the data pect this test not to reject.We
trap response also would expect test 2 to re-
ject Model Moin favorof Mb
This goodnessof fit test can be
partitioned into two specific
tests, useful in their own right
(5a,5b)
5a. Heterogeneity andlor First capture prob- Firstcaptureprob- Thistestis identicalto the good-
time variation using abilities are abilitiesvaryby ness of fit test forthe simplere-
first capture only constant time and/or movelmodel
animals
5b. Heterogeneity and/or Recapture prob- Recaptureprob- If Mbis true then this null hy-
. . ..
.:lme varlatlon uslng

.
abilities are all abilitiesvaryby pothesisshouldnot be rejected

only recaptures constant time and/or
animals
6. Trap response and/or Model Mt fits ModelMtfailsto If Mtis the model,then we ex-
heterogeneity given the data fit the data pect this test to fail to reJect,
time variation and we expect ModelMoto be
rejectedin favorof Mt
7. Trap response given Model Mhfits ModelMbhfits This test is usefulbecauseif we
heterogeneity the data the data reject Mh in favor of Mbhthe
estimatorto use is the gen-
eralizedremovalmethod
with
13. Exampleof the model selectionproceduretests with simulateddata.True model is ^X1,>
FI(;.
modelis M,,.Suggestedestimatoris Null.
parametersN = 100,t = 5, andp = 0.5. Appropriate
1 * . . n
W.
ILDLIFE \ ONOG>PHS
level of 0.33601.Thus,in this example goodnessof fit tests to ModelMbexarn-

ModelMhfitsthe data.This is expected ine whetherany othersourceof varia-
becauseModelMois a specialcase of tlOn 1S OperatlNg.
* . .
NIodelMh,andhencethetestsshouldnot The 7 tests have been siInulatedto

reject. studytheirsize and power,and the re-
Whenindividualcapturefrequencies sults are presentedin AppendixN. In
arelargeenough(we haveused the cri- generalall thetestshavegoodsizeprop-
teriafi > 2t) we have computeda good- erties,i.e., they tend not to rejectwhen
ness of fittest of ModellbvIh basedon the t le nu zypotzesis is true. However,
dataforanimalscapturedexactlyfitimes. mostof the testslackpower,i.e., theydo
Theoretically, thiscanbe doneforallfre- notalwaysrejectwhentheyshould.This
quenciesfl, . . .,ft-lthatarelargeenough. is particularly truefortests of heteroge-
The goodnessof fit test of ModelMb neity(X1,,vs. Mhandthe goodnessof fit
(test5) canalsobe partitioned into2 com- test of Mh)@
ponents.Test5a is actuallythe goodness
of fittestto Zippinsremovalmodel,i.e., On the Needfor an Objective
ittestswhethertheprobability offirstcap- SeZectionProcedure
tureis constantovertime.In Fig. 13,the
test statisticvalue is 2.459 with signifi- The abovediscussionand Fig. 13 il-
cance level 0.48276.Thus,in this case lustratethe generalmodelselectionap-
thereis no evidenceof variationin first proach.Conceptually,we examinethe
captureprobability overtime.Thesecond resultsof the 7 testsof assumptions, and
componentof the test of ModelNIb (test on the basisof the resultschoosethe ap-
5b)examinesfortimevariationin recap- propriatemodel. In the exampleused
tureproDa z11tles. n t ig. , t ze teststa- (Fig. 13),the selectionwas not difficult.
tisticvalueis 2.724.The goodnessof fit However,forothermodels,andformuch
test statisticfor NIodelMbis the sumof realdatawe haveseen,thejudgemental
those2 chi-square values.In Fig. 13 the selectionof an appropriate estimatorcan
valueis 5.183(6dflwithsignificance level be verydifficultbecause(1)the testsare
0.52053.Thus,we concludeNlodelMbfits not independent,in particular if behav-
the data.Again,we pointoutthatModel ioralvariationin captureprobabilities is
Mois a specialcase of Mb,so the result presentthis can stronglyaffectthe test
1S no surpr1s1ng. for tilne and heterogeneityeffects;and
. . .
Thefinalgoodnessoffittestis forMod- (2)forrealdataall3 sourcesofvariability

el Mt(test6). In Fig. 13,the test statistic probablyarepresentin varyingdegrees,
valueof 59.465(54 dfl is not significant so all we canhopeto dois selectthemost
(P- 0.28338). appropriate model(theone thatbest de-
An alternateway of thinkingof these scribesthe data).Withrespectto the first
testsis as follows.The test of ModelMo point,considerwhathappensif Model
vs. Mhtests whetherthereis heteroge- Mb iS true.Forexample,assume7 trap-
neity of captureprobabilitieswhile the pingoccasionswith]\Ib asthetrueluodel,
goodnessof fit test of Model Mhtests and p = 0.40, c= 0.20. Thatis, priorto
whetherthereis anyadditionalsourceof beingcapturedanimalshavea 0.40 cap-
variationin captureprobabilitiesdue ture probability7 but due to behavioral
eitherto behavioror timeor both.If we response(trapavoidancein this case)
rejectedModelMoin favorof Mhandthe their captureprobabilitiesdropto only
goodnessof fittestto Mhindicatesalh fits 0.20afterfirstcapture.Theexpecteddai-
the data,we couldconcludeNIodelMh ly captureprobability is givenby
wasappropriate.
Similarly,the testsof ModelMovs. Mb pj= E(n.)
NJ, j- 1J**X7
test forwhetherthereis behavioral vari-
ationin captureprobabilities, while the These are constantonly if the expected
. n
STATISTICAL FROMCAPTUREDATAtis
INFERENCE et al.
57
dailycapturesare constant.Butas trap- cal versionof this probleln,one has a
pingprogresses theaveragedailycapture sample of rneasurementsfrom each
probabilitydecreasesas moreand more knownpopulationv andfroznthosedataa
animals"shift"theircaptureprobability uathematical ruleis constructed forclas-
froln0.40 to 0.20 as a resultof trapre- sifyingfuturecasesbasedon theirlnea-
sponse.A formu]aforpjin this caseis surelnents.In our case, the '<lneasure-
ments"are the significancelevels froln
_ (1-(1-pVi-l) C + (1-p)i-lpS the 7 tests.The 4'populations" arethe 8
Pi 1 - (1 - p)i Inodels.Whatis unknownis whatmodel
j = 2, . . ., t best fitsanycapturedataat hand.Given
this conceptualization, we chose to use
(P1-p). For the case above, we have discriminantfunctionanalysesto con-
struct the classificationfunction (see
J Pi Cooley andLohnes1962).The objective
1 0.400 of the discrilminant analysisis to weight
2 O.275 and linearly coznbinethe significance
3 0.237 levels of the 7 tests in solnefashionso
4 0.220 that the modelsareforcedto be as statis-
5 0.211 tically distinctas possible.
6 0.207 The modelclassification functionwas
7 0.204 derivedfromsimulateddatawherethe
WhenModelMb iS true,the testfortilne trueluodelis known.The captureprob-
variationin captureprobabilities (Model abilitystructureof these simulateddata
Movs. Mt)will tendto rejectModelMoZ 1S presentec1n Wa zi e Z. 1 aeseparame-
becausethebehavioral responsedoesin- tervalueswerechosenbecausetheyare,
deed"cause"timevariation tobe present in our opinion,representative of luany
also. real capture studies. Foreach populationt
The samesortof argumentshowsthat: 100 replicationswere generated,giving
behavioralresponse"causes'heteroge- a total of 1,600 cases. The probability
neity on any given day. Consequently,levels fromeachof the 7 testswerethen
whenModelMb iS trueSthereis a strong used to constructthe classification func-
tendencyforall of the simpleteststo re- tion.In additionto the 7 probabilities, 9
ject ModelMoSmakingselectionof the additionalvariableswereconstructed by
correctmodeldifficult. taking all possible productsbetween
In thefinalanalysis,selectionofa Inod- probability levelsfrolntests1-3 andtests
ew. lnvoves examlnlnga polntln a t-c1- 4-6>i.e. by takingthe 9 pairwiseprod-
. . . . . os .
lnensionalspaceandtryingto classifyit ucts of the probabilities fro1nthe 3 spe-

into 1 of 8 categories.We havenotbeen cific tests againstModelM, anelthe 3
successfulourselvesin providinga set of goodnessof fit tests. This providesa
simplerulesforthis choiceandbelieve totalof 16variables thatweretransformed
thatfieldbiologistswithoutrigoroussta- byX = log(P+ 0.01).Thattransforluation
tisticaltrainingmighthavegreatdifficul- tends to weightthe smallerprobability
ty arrivingat a properchoice. levelsmoreheavilythanthelargervalues
close to 1. The classification ruleresult-
An Objective Xlodel ing from that seriesoftransformations was
Selection Procedure one of many rules examine(l; this par-
ticularrule was chosenbecauseit had
Theproblelnidentifiedin theprevious the best erformance
sectionfallsin the realmof multivariate The transformed variableswere then
statistics,specificallyclassification of an usedto computea classification function
unknownentityinto one of severaldif- using the SPSSdiscrilminant procedure
ferentknownpopulations.In the classi- (Nieet al. 1975). Theclassification func-
TABLE 14.-CAPrURE PROBABILITYSTRUCTUREFOR THE SIMULATEDDATA USED TO GENERATETHE

CLASSIFICATIONFUNCTION. ONE HUNDRED REPLICATIONSOF EACH TYPE WERE GENERATED,GIVING
A TOTALSAMPLESIZE OF 1,600
Model ProbabilityStructure
LARGE POPULATION (N = 400, t= 7)

MO P = 0 30
Mh Pj = 0.15, i = 1,100; PX= 0.25, i = 101,200; Pj = 0.35, i = 201,300; Pj = 0.45, i = 301,400.
Mb P = 0.2; C= 0.5.
Mbh p; = 0.15, Cj= 0.375, i = 1,100; Pj = 0.25, CZ = 0.625, i = 101,300; PX= 0.35, Cj= 0.875,
i = 301,400.
Mt Pl = 0.15; P2 = 0.45; p3 = 0.35; p4 = 0.25; p5 = 0.30; P6 = 0.20; p7 = 0.40.
Mth Pij = Pi-P;; Pi = 0.45, i = 1,100; Pi = 0.55, i = 101,200; Pi = 0.65, i = 201,300; Pi = 0.75,
i = 301,400; P1 = 0.65; P2 = 0 75; P3 = 0.55; p4 = 0.35; p5 = 0.60; P6 = 0.50; p7 = 0.80.
Mtb Pi; = P;-C; C = 2.50; P1 = 0 05; P2 = 0.35; p3 = 0.25; p4 = 0.15; p5 = 0.20; P6 = 0.10; p7 = 0.30.
MtBh Pi; = Pi Pj C; Pi = 0.15, i = 1,100; Pi = 0.25, i = 101,200; Pi = 0.35, i = 201,300; Pi = 0.45,
i = 301,400; C = 2.50; P1 = 0.65; P2 = 0 75; P3 = 0.55; p4 = 0.45; p5 = 0.60; P6 = 0 50;
P7 = 0.70.
SMALL POPULATION (N = 100, t= 5)

MO P = 0.10.
Mh Pi = 0.05, 1 = 1,25; Pi= 0.15, i = 26,50; Pi= 0.25, i = 51,75; Pi = 0.35, i = 76,100.
Mb P = 0.40; C = 0.20.
Mbh PI = 0.15, Ci = 0.075, i = 1,33; Pi = 0.25, Ci = 0.125, i = 34,66; Pi = 0.35, Ci = 0.175, i = 67,100.
Mt P1 = 0.15; P2 = 0.45; p3 = 0.35; p4= 0.25; p5 = 0.30.
Mth Pi; = Pi P;; Pi = 0.35, i = 1,25; Pi = 0.45, i = 26,50; Pi = 0.55, i = 51,75; Pi = 0.65, i = 76,100;
P1 = 0.65; P2 = 0 75; P3 = 0.55; p4 = 0.35; p5 = 0.60.
Mtb Pij = P; C; C = 0.5; P1 = 0 35; P2 = 0.25; p3 = 0.15; p4 = 0.20; p5 = 0.30.
Mtbh Pi; = Pi Pj*C; Pi = 0.15, i = 1,25; Pi = 0.25, i = 26,50; Pi = 0.35, i = 51,75; Pi = 0.45, i = 76,100;
C = 0.75; P1 = 0.65; P2 = 0 75; P3 - 0.55; p4 = 0.45; p5 = 0.60.
tionconsistsof 8 setsofcoefficients(1 set centroidsforthe transformed probability

foreachof the 8 models)thatareusedto levels.The ithfunctionhasthe form
forlnlinearcombinations of the 16 vari-
ablesplusanintercept.Theclassification Ci- ciO+ cil Xl + ci2X2
equationsare derivedfromthe pooled + + ci16X16
within-nodelcovariancematrixand the The 136 classificationcoefficientsare
TABLE 15. CLASSIFICATIONFUNCTIONCOEFFICIENTSUSED IN THE MODEL SELECTIONPROCEDURE
COeffiCient M(O) M(H) M(B) M(BH) M(T) M(TH) M(TB) M(TBH)

C0 -8.09154 - 14.33446 -25.69392 - 17.15981 -31.90722 -23.20432 -26.11839 - 15.07368
C1 2.44865 1.55835 2.41361 2.09104 4.85352 4.52842 4.58145 1.01685
C2 0.01513 0.76894 0.34328 -0.30063 4.19868 3.36253 - 1.18146 -0.78482
C3 5.60627 7.61750 4.45155 2.65005 -3.83028 -4.33794 3.81010 1.97090
C4 2.31654 2.43157 - 1.90418 1.85727 4.11214 3.84041 0.07484 5.00658
C5 0.39009 0.24251 2.57294 2.95937 -0.37644 -0.30277 -0.73170 - 1.36695
C6 1.59847 1.50362 5.40943 0.82379 3.29315 2.76447 4.77294 0.91642
C7 - 1.07536 -0.75833 - 1.39342 -0.92010 - 1.87794 0.47319 - 1.46921 -0.87191
C8 -0.95870 -2.99247 -0.29576 0.83512 - 1.39938 - 1.28509 - 1.44532 -2.06608
C9 -4.23372 -5.76230 - 1.30645 -3.97856 -4.93318 -4.85310 -3.75607 - 7.44977
C10 0.73672 2.16608 1.71422 -0.11956 1.32575 2.28866 1.74703 2.68257
C11 -0.90843 -0.51321 0.06944 -2.67855 -2.69316 -2.01668 - 1.73071 1.08225
C12 1.54069 2.72345 -4.56388 -2.53049 -5.55330 -2.64194 0.53878 2.03048
C 13 -5.49376 -6.30792 -2.38615 -2.14175 4.38634 -0.03381 -2.06993 -5.17029
C14 -3.30107 -2.40404 -5.13204 -2.38473 -3.79996 -4.33330 -4.76823 -2.41632
C15 -0.19891 - 1.42895 -2.26381 -0.48135 0.35309 -2.42112 - 1.92578 -0.09665
C16 -2.04687 -3.46579 -4.06512 - 1.73548 -4.64956 - 1.59132 -3.89432 - 1.80314
STATISTICAL
INFERENCE
FROMCAPTUREDATA{:)tiS et al. 59
TABLE 16. CAPTUREPROBABILITY

STRUCTUREUSED TO SIMULATEGOOD,MEDIUM,AND POORDATATO
STUDYTHE OPERATINGCHARACTERISTICS
OF THE MODELSELECTIONPROCEDURE.A POPULATION
OF 400
WASTRAPPED7 OCCASIONSFOR 200 CASES
Model ProbabilityStructure
GOOD DATA (Average p 0.35)

Mo P =0 35-
Mh Pi = 0.5, i = 1,100; Pi = 0.4, i = 101,200; Pi = 0.3, i = 201,300; Pi = 0.2, i = 301,400.
Mb p= 0.5; c = 0.2.
Mbh Pi = 0.65, ci = 0.10, i = 1,100; Pi = 0.55, ci = 0.15, i = 101,200; Pi = 0.45, ci = 0.10, i = 201,300;
pj = 0.35, ci = 0.50, i = 301,400.
Mt P1 = 0 35, P2 = 0.45, p3 = 0.25, p4 = 0.35, p5 = 0 20, P6 = 0 50, p7 = 0.35.
Mth Pii = Pi-Pi; Pi = 0.9, i = 1,100; Pi = 0.8, i = 101,200; Pi = 0.7, i = 201,300; Pi = 0.6, i = 301,400;
P1 = 0 5; P2 = 0-6; p3 = 0.4; P4 = 0.5; P5 = 0-35; P6 = 0.65; p7 = 0.5.
Mtb Pii = pj-C; c = 1.5; P1 = 0-3; P2 = 0.45; p3 = 0-15; p4 = 0.3; p5 = 0-2; P6= 04; P7 = 0-3
Mtbh Pij = Pi *pj c; Pi = 0.5, i = 1,100; Pi = 0.4, i = 101,200; Pi = 0.3, i = 201,300; Pi = 0.2, i = 301,400;
c = 1.5; P1 = 0 7; P2 = 0 7; P3 = 0 4; P4 = 0-4; P5 = 0-7; P6 = 0-9; p7 = 0.87.
MEDIUM DATA (Average p 0.20)
Mo p= 0.20.
Mh Pi = 0.3, i = 1,100; Pi = 0.25, i = 101,200; Pi = 0.15, i = 201,300; Pi = 0.10, i = 301,400.
Mb p = 0.15; c= 0.03.
Mbh Pi = 0.35, ci = 0.05, i = 1,100; Pi = 0.4, ci = 0.1, i = 101,200; Pi = 0.2, ci = 0.05, i = 201,300;
Pi = 0-25, ci = 0.2, i = 301,400.
Mt P1 = 0-02; P2 = 0-35; p3= O.OS; p4 = 0-2; p5 = 0-1; P6= 0-3; p7 = 0-2
Mth Pij = Pi Pj; Pi = 0.7, i = 1,100; Pi = 0.55, i = 101,200; Pi = 0.45, i = 201,300; Pi = 0.3, i = 301,400;
P1 = 0-4; P2 = 0.55; p3= 0.25; p4 = 0-4; P5 = 0.3; P6= 0-5; P7 = 0.4.
Mtb Pij=pj*c; c=0.33; P1=0-30; P2=0.45; p3=0. 15; p4= 0.30; p5= 0-20; P6=0-40; P7= 030-
Mtbh Pii = Pi-Pi-e; Pi = 0.3, i = 1,100; Pi = 0.25, i = 101,200; Pi = 0.15, i = 201,300; Pi = 0.10,
i = 301,400; c= 2.00; P1 = 0-50; P2 = 0.65; p3 = 0.35; p4= 0-50; P5= 0-40; P6= 0-70;
P7 = 0.50.
POOR DATA (Average p 0.05)

Mo P = 0-05-
Mh Pi = 0.09, i = 1,100; Pi = 0.05, i = 101,300; Pi = 0.10, i = 301,400.
Mb p = 0.075; c= 0.01.
Mbh Pi = 0.07, ci = 0.03, i = 1,100; Pi = 0.03, ci = 0.08, i = 101,300; Pi = 0.01, ci = 0.09, i = 301,400.
Mt P1 = 0-05; P2 = 0.01; p3 = 0.09; p4 = 0-05; p5 = 0-07; P6 = 0 03; p7 = 0.05
Mth Pij = Pi-Pi; Pi = 0-20, i = 1,100; Pi = 0.25; i = 101,200; Pi = 0.05, i = 201,300; Pi = 0.10,
i = 301,400; P1 = 0-33; P2 = 0.10; p3= 0.10; p4 = 0.333; P5 = 0-50; P6= 0-50; P7 = 0.33-
Mtb Pii = pj-c; c = 0.10; P1 = 0.10; P2 = 0.15; p3 = 0.05; p4 = 0.10; p5 = 0.19; P6 = 0.01; p7 = 0.10.
Mtbh Pii = Pi-Pi-c; Pi = 0-20, i = 1,100; Pi = 0.25, i = 101,200; Pi = 0.05, i = 201,300; Pi = 0.10,
i = 301,400; c= 2.00; P1 = 0.20; P2 = 0.15; p3 = 0.20; p4= 0-05; P5= 0-05; P6= 0.15;
P7= 0.37.
given in Table 15. The 8 Ci valuesare N = 400 andt = 7. Thegooddatahadan

then standardized overthe interval0 to approximateoverall average capture
1 to give the model selectioncriteria. probability p of 0.35,mediump wasap-
Thesestandardized "modelselectioncri- proximately 0.20,andpoorp wasapprox-
teria"areprintedby program CAPTURE imately 0.05. The captureprobability
rightafterthe 7 tests(cf.Fig. 13). structureof these datais givenin Table
Given the classification functions,an 16.
evaluationof their usefulnessmust be Weanalyzedeachdatasetandselected
made.Again,dataarerequiredwherethe a modelbasedon the classification func-
underlying modelsareknown;thisis only tion describedabove. This allows an
possiblewithsimulateddata.Hence,we evaluationof the selection procedure,
simulated3 additional datasets:200repli- i.e., how oftena wrongmodelis chosen,
cationseachof good,medium,andpoor and whichmodelstend to be confused.
dataforeachmodel.In all cases,we used Theseresultsaregivenin Tables17-19
TABLE 17. PERFORMANCE OF THE MODELSELECTIONPROCEDUREWITHGOODDATA.THE TRUEMODEL

FROMWHICHTHE DATAWEREGENERATED IS AT THE TOP,AND THE MODELSELECTEDBYTHE CLASSIFICA-
TION FUNCTIONON THE SIDE. THE ENTRIESARETHE PERCENTAGE OF TIMESTHE MODELWASSELECTED,
THE AVE(N), AND THE 95g0 CONFIDENCEINTERVALCOVERAGE
Data generated frommodel

Model All
selected Mo Mh Mb Mbh M, Mth Mtb Mtbh data
70.0 2.5 0.0 0.0 0.0 0.0 0.0 0.0 9.1

Mo 399.7 382.2 - - - - - - 399.1
0.957 0.200 - - - - - - 0.931
21.5 93.5 0.0 0.0 0.0 0.0 0.0 0.0 14.4
Mh 414.0 427.4 - - - - - - 424.9
0.349 0.059 - - - - - - 0.113
0.0 0.0 90.0 0.0 0.0 0.0 0.0 0.0 11.3
Mb - - 399.7 - - - - - 399.7
- - 0.922 - - - - - 0.922
6.5 1.5 0.5 93.0 0.0 0.0 0.0 0.0 12.7
Mbh 394.2 396.3 399.0 396.6 - - - - 396.4
0.692 1.00 1.00 0.575 - - - - 0.591
1.0 0.0 0.0 0.0 82.0 58.5 2.0 0.0 17.9
Mt 404.0 - - - 399.8 396.3 372.0 - 398.0
1.00 - - - 0.951 0.829 0.0 - 0.888
0.0 1.0 0.0 0.0 6.5 39.5 0.5 0.0 5.9
Mth _ _ _ _ _ _
0.0 0.5 9.5 0.0 11.0 2.0 95.0 37.0 19.4

Mtb - - - - - - _ _ _
1.0 1.0 0.0 7.0 0.5 0.0 2.5 63.0 9.4

Mtbh - - - - - - _ _ _
Mean population
estimate 402.5 425.8 399.7 396.6 399.8 396.3 372.0
Coverage 0.808 0.077 0.923 0.575 0.951 0.829 0.0
forthe 3 typesof data.However,ourin- tion,thecorrectmodelis muchlesslikely

terestin these simulateddatadoes not to be selectedforpoordata.Notethatin
stopwiththe resultsof the modelselec- Table19 (thepoordatacase),ModelM0
tion.Evenmoreinformative is howwell is selected 34.7 percent of the time
the estimatorperformed whenthe wrong (whereasit is the truemodelonly 12.5No
modelwas selected(we knowthatesti- of the time),andthatAve(NO) is 745.5.
matorperformance usuallyis acceptable For good data,such a discriluination
when the correctmodelis used).Infor- procedurewill do quitewell in selecting
mationon the Ave(N)andconfidencein- the appropriate model.We suggestthat
tervalcoverageis also given in Tables usersknowledgeable in statisticsandex-
17-l9 forall estimators.The tablesalso periencedin analysisof capturedatamay
providesummariesby model selection sometimesbe ableto renderanimproved
andby datatype. judgementaboutthe approrpriate model
Thosetablesagainemphasizethe im- forrealdata.However,thefieldbiologist
portanceof high captureprobabilities. shouldprobablyacceptthe model rec-
The estimatesand coveragecoefficients ommendedby the modelselectionalgo-
drasticallydecline in usefulnessas the rithm.The primaryreasonforthis is the
captureprobabilitiesdecline. In addi- problemswith dependenceamongtests
MO
Mbh 409.3
400 0.00() 382.0
8 361.1
471.8
0.399 390.1 234.0
0.Q00 - - 467.1 410.6
- 388.4Q.31Q
STATISTICAL
INFERENCE
FROMCAPTUREDATAtiS et 1. 61
TABLE 18. PERFORMANCEOF THE MOI)EL SELECTIONPROCEDUREWITH MEDIUMDATA. THE TRUE

MODELFROMWHICHTHE r)ATAWEREGENERATEDIS AT THE TOP AND THE MODELSELECTEDBY THE
CLASSIFICATIONFUNCTIONON THE SIDE. THE ENTRIESARE THE PERCENTAGEOF TIMES THE MODEL
WAS SELECTED,THE AVE(N),,AND THE 95% CONFIDENCEINTERVALCOVERAGE
Data generated frommodel

Model All
selected Mo Mh Mb Mbh Me M,h Mtb Mthh data
68.0 28.0 0.0 0.0 0.0 0.0 Q.0 Q.5 12.1
0.971 0.089 - - - - - 0.000 0.710

18.5 66.5 0.5 0.0 0.0 0.0 0.0 0.0 10.7
Mh 484.7 449.7 890.4 - - - - - 459.9
1.0 0.() 81.0 58.0 Q.() O.Q 0.0 1.5 17 7

427.0 - 397.2 388.4 - - - 783.0 397.9
0.50() - 0.951 0.681 - - - 1.00 0.837
7.5 3.0 9.0 32.0 0.0 0.0 0.0 3.5 6.9
0.667 0.333 0.949 0.688 - - - 1.000 0.727

1.0 0.0 0.0 0.0 57.5 67.S 0.0 1.5 15.9
'l[t 405.0 - - - 400.3 377.9 - 251.3 386.7
1.000 - - - 0.974 0.593 - 0.000 0.761
3.0 2.0 O.Q ().() 1().5 25.5 0.() 1 1.5 6.6
Mth _ _ _ _ _ _
Q.0 0.() 8.5 8.5 31.5 6.5 98.0 65.0 27.S

Mtb _ _ _ _ _ _
1.0 0.5 1.(3 1.5 0.5 0.5 2.0 16.5 2.9

Mtbh - - - - -
Mears population
estimate 418.0 422.2 407.4 389.0 400.3 377.9 - 4'tl.9
Coverage 0.755 0.308 0.945 0.683 0.974 0.593 - 0.714
arldthefactthatwithrealdatait is highly ingarlestimator.If sucha mode]is founll,

unlikelythatanyof these 8 luodelswill and if the relevanttests of lmodelas-
be exactly"true.'> suluptionssuggestthatthe gooclnessof
fit of the modelis adequatethe corre-
Estnation in Alternattte Models spondirlgestiluatorcan be used with
some confidencXe. We would cautionn
Whenthe luodelselectionprocedure however,againstusingmodelswith se-
describedabovehasclassifieda modelas lectionvalllesless tharl0 75 to produce
the best one for a given set of data,2 estimatesof N especiallyif there is a
problemsluaystillarise.First the model poorfit of the modelto the clata.In the
Inaynot have an associatedestimation secondcase,noneof theInodelswithcor-
procedureforestinlatingN. Second,the respondingestimationproceclures seelus
model with the largestselectiorlvalue to fitthe datawell;wherefit is judgedby
(always1.0) luaynot reallyfit the data, the luodel selectionprocedureand irl-
even thoughit is the "best"modelavail- spectionof individualtests,thenwe can
able. In the firstcase, the investigator see nojustification
forgrantingstatistical
shollldsean for a large seleetionvalue validityto anycalculatedpoplllationes-
(say0.90)corresponding to a Inodelhav- tiluate.
selected Mo Mh Mb Mbh Mt M h Mtb Mtbh data
TABLE 19. PERFORMANCE OF THE MODELSELECTIONPROCEDUREWITH POORDATA.THE TRUE MODEL

FROMWHICHTHE DATAWEREGENERATEDIS AT THE TOP, ANDTHE MODELSELECTEDBY THE CLASSIFICA-
TION FUNCTIONON THE SIDE. THE ENTRIESARE THE PERCENTAGEOF TIMESTHE MODELWASSELECTED,
THE AVE(N), AND THE 95g0 CONFIDENCEINTERVALCOVERGE
Datageneratedfrommodel
Model All
79.0 74.0 39.5 83.0 0.0 1.0 0.0 1.0 34.7

Mo 417.5 316.3 3,229.7 270.2 - 342.5 - 146.5 745.5
0.949 0.547 0.557 0.386 - 0.500 - 0.000 0.615
13.0 18.5 4.5 11.0 1.0 9.0 0.0 5.0 7.8
Mh 377.7 336.5 612.2 255.6 374.6 347.1 - 210.5 342.8
0.923 0.514 0.000 0.000 1.000 0.611 - 0.000 0.452
0.5 1.0 14.0 0.0 0.0 0.0 0.0 0.0 1.9
Mb 158.0 156.0 272.8 - - - - - 261.6
0.000 0.000 0.250 - - - - - 0.226
3.0 4.0 33.0 2.0 0.0 0.0 0.0 0.0 5.3
Mbh 174.7 172.1 359.5 123.3 _ _ _ _ 317.2
0.000 0.000 0.924 0.000 - - - - 0.726
1.0 0.5 0.0 1.5 83.5 24.5 2.0 68.0 22.6
Mt 366.5 246.0 - 216.7 406.7 325.9 2,983.0 164.5 331.0
1.000 0.000 - 0.000 0.940 0.653 0.000 0.059 0.550
1.5 0.5 0.5 1.5 5.0 5.0 0.0 0.5 1.8
Mth - - - - - _ _ _ _
0.5 0.0 4.5 0.0 1.0 0.0 92.5 0.0 12.3

Mtb - - - - - - _ _ _
1.5 1.5 4.0 1.0 9.5 60.5 5.5 25.5 13.6

Mtbh - - - - - - - _ _
Mean population
estimate 402.7 312.2 1,604.5 264.7 406.3 331.9 2,983.0 167.4
Coverage 0.912 0.510 0.615 0.328 0.941 0.638 0.000 0.054
Additional Examples of
Model Selection was donetwlce dally,mornlngandeve-
ning,forS days.Thusthereare 10 trap-
As partof a populationecologystudy ping occasions,but we can expecttime
on salt marshrodents,Coulombe(1965, variationin captureprobabilitiesbe-
unpublishedmaster'sthesis, University tweenmorningandnightoccasions.The
of California,Los Angeles,California), entiredataset of 173distinctindividuals
conducteda livetrapping
studyonanout- capturedincludedyoungandadult,and
breakof feralhousemice musculus male and female.Thus, we mightalso
L7dUs
in asaltmarshinmid-December 1962,at expect some heterogeneityof capture

BallanaCreek,LosAngelesCounty,Cal- probabilities.
ifornia.A square10 x 10 gridwas used Fromthe model selectionprocedure
withtrapsspaced3 m apartandtrapping (Fig. 14),thereis clearevidenceof time
FIG. 14. Exampleof the model selection procedureapplied to Coulombe's(unpublishedthesis) full

dataset. Appropriate
modelprobablyis Mth.No estimatorresultsfromthe model.
STATISTICAL
INFERENCE
OCCASI ON J= 1 2 3 4 5 6 7 8 9 10
ANI MALS CAUGHT N( J ) = 68 61 62 52 74 41 76 36 76 39
TOTAL CAUGHT M(J)= 0 68 102 128 140 156 159 171 171 173 173
NEWLYCAUGHT UIJ ) = 68 39 26 12 16 3 12 0 Z 0
FREQUENCI ES F (J ) = 2 64 40 31 16 13 5 1 0 1
I . TEST FOR HETEROGENEI TY OF TRAPP1NG PROBABI L I T I ES I N POPULATI ON.

NULL HYPOTHESI5 OF MODEL M(O) VS. ALTERNATEHYPOTHESI5 OF MODEL M(H)
CHI -SQUARE YALUE = 98. 576 DEGREES OF FREEDOMa 6 PROBABI L I TY OF LARGER VALUE = 0 . 00000
2 . TEST FOR BEHAVI ORAL R'ESPON5:EAFTER I N I T I AL CAPTURE,

NULL HYPOTHESIS OF MODEL M(O) VS. ALTERNATEHYPOTHESIS OF MODEL M(B)
CHI-SQUARE VALUE = 1.848 DEGRE£S OF FREEDOMs 1 PROBABILITY OF LARGER VALUE = . 17400
3. TEST FOR TIME SPECIFIC VARIATION IN TRAPPING MWABILITIES.

NULL HYPOTHESIS OF MODEL M(O) YS. ALTERNATEHYPOTHESIS OF MODEL Ml T )
CHI-SaJARE VALUE 5 62.246 DEGREES OF FREEDOMs 9 PROBABILI TY OF LARGER VALUE = O.00000
4 . GOESS OF F I T TEST OF MOOEL M( H)

MJLL HYPOTHESIS OF MOOEL MIH) VS, ALTERNATEHYPOTHESIS Of NOT MODEL M(H)
CHl -SOUAR.E VALUE = 57.151 D£GREES OF FREEDOMx 9 PROBABILI TY OF LARGER VALUE = 0. 00000
TEST OF MbEL HIH) BY FR£WENCY Of CAPTURE

lFREQUENCIES LESS THAN 2T ARE NOT CALCULATED.)
NU1BER OF CAPTURES CHI -SOUAR£ D. F . PROBABI L I TY

_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
2 1W.027 9 .12136
3 6.857 9 .65199
4 27.387 9 .00121
5 . Gi>)ONESS OF F I T TEST OF MODEL M( B 1

NULL HYPOTHE5I S OF MODEL M( B ) VS . ALTERNATEHYPOTHESI S OF NOT MODEL M( B )
CHI -SQUARE YALUE = 6a . 087 DEGREES OF FREEDOM= 15 PROBABI L I TY OF LARGER VALUE = 0 . 00000
5A . CONTRI BUTI ON OF TEST OF HOMOGENE

I TY OF F I RST CAPTURE PROBABI L I TY ACROSS T I ME
CHI -SOUARE YALUE = 2W.780 DEGREES OF FREEDOM= 7 PROBABI L I TY OF LARGER VALUE = . 00083
5B . CONTRI BUTI ON OF TEST OF HOMOGENE

I TY OF RECAPTUREPROBABI L I T I ES ACROSS T I ME
CHI -SQUARE VALUE = 93. 307 DEGREE5 OF FREEDOM= 8 PROBABI L I TY OF LARGER VALUE = . 00000
6 (£55 OF F I T TEST OF MODEL M( T 1

NULL HYPOTHESI S OE MOt)EL M{ T ) VS . ALTERNATEHYPOTHESI S OF NOT MODEL M( T )
CHI -SGIUAREVALUE = 162 . 329 DEGREES OF FREEDOMs 125 PROBABI L I TY OF LARGER VALUE = . 01 386
7 . TEST FOR BEHAYI ORAL RESPE I N PRESEKE OF HETEROGENE

I TY.
NULL HYPOTHESIS OF MODEL M(HI V5. ALTERNATEHYPOTHESI5 OF MODEL MIBH)
CHI -S(kJARE VALUE = 31 .939 DEGREES OF FREEDOMz 24 PROBABILI TY OF LARGER YALUE = . 14153
MODELSELECTI OEJCRI TERI A . KDEL 5ELECTED HAS MAXI MUMVALUE.
M(X)ICL M1O) MIHI M(B) M(BH) M(T) M(TH) M(TB) M(TBH)

CRITERIA . J5 .29 .03 .22 .26 1.00 0.00 .66
andheterogeneity variation,butlittlein- all 5 possibleestimatorsto be in close

dication of behavioralvariation.Note agreementwiththe valueof 173.In fact
alsothatnoneof the simplemodels(Mh, the resultsfor Coulombesfull dataset
Mb,Mt)fit the data.The modelselection erCt1 w .
criteriastronglysuggestModelMthas ap-
propriate, but thereis no estimatorasso- tstzmator va ue Standard
_ .
error s T
ciatedwith this model.To obtainan es- No 176 1.8

timator,we can look for the next most Nt 175 1.6
likelymodel,whichis Mtbh (selectioncri- Nb 174 1.6
teria= 0.66);however,thereis no esti- Nh 175 1.8
matorforthatmodeleither.In fact,to get Nbh 173 0.2
a modelallowingestimationwe would
haveto use eitherMt,Mh,or h. Unfor- These are all very siluilararldprecise.
tunately,the value of the selectioncri- Notethatin generalit is notappropriate
teriacorresponding tothosemodelsis too to computeall estimates.Werecomlnend
low to allowlegitimatechoiceof one of it onlywhenthereis strongevidenceall
the models for estimation purposes. animalswerecaptured,in whichcase it
Therefore,the investigator mustrealize providesan additionalcheck.
thatnone of the modelscan be used to Asa furtherexample,Fig. 15showsthe
estimatepopulationsize withthe datain luodel selectionresultsusing only the
the presentform. morningcapturedatafromCoulombes
One alternativeis to simplytake the (unpublished thesis)study(i.e.>pretend-
totalnumberseen (lVlt+l) as the best es- ing the evening captures never oc-
tiluate.Given good data, Mt+1will be curred).Presumably,this would elimi-
nearlyequalto N. Also with good data nate mostof the time variation,but not
one will tend to reject the simpler neterogene1ty. l ne selectloncr1terza ver-
1 * 1 1 * * *
luodels.The real measure,however,of ify thisconjecture.

gooddatacomesfromsuchthingsas t, n./ Model selection
Mt+l,thepatternof thenewcaptures(the Model crlterla
uj's)and the apparentaveragecapture Mo 0.99
probability(averagep is 0.3 here using 1.00
N= 173= NIll).In good data,very few Mh
Mb ().58
new animalswill be caughtby the last 0.74
fewoccasions;in Coulombe's data,only2 Mt
Mbh
0.00
new anirnas were caugzt attert re sev- 0.46
enthoccasion.Similarly,the capturefre- Mth
0.53
quencydatashouldshowmanyanimals Mtb
0.80
M tbh
caught2, 3, 4, or moretimesand there
shouldnot be a strongspikeat fi (cap-
turedonce only).By all these measures, Whenthe criteriavaluefor2 or more
Coulorllbe's datasuggestthatalmostall modelsexceed 0.95, the programdoes
the populationwas caught.Thus,here not just suggest I, but names the 2
we wouldsuggesttakirlg173as the esti- models that have the highest criteria.
uatedpopulationsize. Thus in this case the choice between
Underthosecircumstances, we would modelsMoand Mhis not clearcut.As a
also expectthe pointestimateof N from standardoperatingE>rocedure we rec-
FIG.15. Example of the model selection procedureapplied to Coulombe's(unpublishedthesis)

morningtrappingoccasions (Occasions1, 3, 5 7> 9). Appropriatemodel probablyis Mh or M1).
Suggestedestimatoris jackknife.
STATISTICALINFERENCEFROMCAPTUREDATAtiS et al. 65
OCCASI ON Jx 1 2 3 4 5
ANI MALS CAUGHT N{ J ) w 68 62 79 76 76
TOTAL CAUGHT Mt J ) a 0 68 103 127 153 160
NEWLYCACHT U( J ) w 68 35 24 26 7
FREQUENCI ES F(J ) 5 W5 59 36 15 5
I . TESt FOR HETEROGENEI TY OF TRAPPI NG PRO6ABI L I T I £5 I N POPULATI ON.

NULL HYPOTHESIS OF MOOEL M(O) VS. ALtERNATE HYPOtHESIS OF MODEL M(H)
CHI-SQUARE VALUE s 5.051 DEGREES OF FREEDOM- 3 PROBABILITY Of LARGER VALUE = * 16813
2. TEST FOR BEHAYIORALRESPONSE AFTER INITIAL CAPTURE.

NlJLL HYP9THESI S OF MOOEL Mf O) VS . ALTERNATEHYPOtHESI S OF MODEL Ml E )
CHI-SOUARE VALUE s 2.271 DEGREES (X FREEDOM- 1 PROBABILITY OF LARGER VALUE = . 13184
3 . TEST FOR T I ME SPEC IF I C VAR}ATI NQ IN TRAPRI NC MOBAB I L I T I ES .

NULL HYPOTHESI S OF MOOEL M1O) VS . ALTERNATEHYPOTHESI S OF MODEL M( T }
CHI -SUARE VALUE = 3 . 667 DEGRE:ESOF FREEDOMw 4 PRO8ASI L I TY OF LARGER VALUE = . 45295
4 . ESS OF F I T TEST OF MOXL M( H}

NULL HYPOTHE515 OF MOOiEL M( H) VS . ALrERNAtE HYPOtHESI S OF NOT MODEL Mt H)
CHI-S(BUARE VALUE s 3.674 DEGREES OF FREEDOMs 4 PROBABILITY OF LARGER VALUE = .45191
TEST OF MODEL" I H) BY FREKY OF CAPtURE

(fREaUENCIES LESS THAN 2T ARE NOT CALCULATED.)
NUMBEROF CAPTURES CHI -SQUARE D. f . PROBABI L STY

_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
1 3. 333 4 .50367
2 4 . 362 4 . 359Z8
3 3. 074 4 .5455 1
4 2.667 4 .61506
5 . GOOONESSw F I T TEST y MODEL M( B )
NULL HYPOTHESI S OF MODEL M( B ) VS . ALTERNATEHYPOTHESI S OF NOT MODEL M1B )
CHI-SQUARE VALUE = 10.240 DEGREES OF FREEDOM= 6 PRO8ABILITY OF LARGER VALUE = . 1 1492

I TY OF F I R5t CAPTURE PROBAdI L I TY ACROSS T I ME
CHI -SQUARE VALUE = 7 . 735 DEGREES OF FREEDOM= 3 PROBABI L I TY OF LARGER VALUE = . 05183

1tY OF RECAPTUREPR0RA8 I L I T I ES ACROSS T l ME
CHI -SQUARE VALUE = 2.505 DEGREES OF FREEDOM= 3 PROBABI L I TY OF LARGER VALUE = .47440
6. NESS OF f I T TEST OF MODEL M( T 1

WLL HYPOTHESI S w MODEL M( T ) YS . ALTERNATEHYPOtHESI S OF NOT MODEL M( r )
CHI -SOUAkE YALUE = 115.230 DEGREES OF FREEDOM= 10 1 PRO8ABI L I TY OF LARGER VALUE = .15766
7. TEST FOR BEHAVIORALRESPONSE IN PRESENCE OF HETEROGENEITY.

NULL HYPOTHESIS OF MO()EL M(H) VS. ALtERNATE HYPOTHESIS OF MODEL M(BH)
CHI -SWARE VALUE = 16. 497 DEGREES y FREEDOMw 10 PR0BABI L I TY 0f LARGER VALUE = . 08626
MOOELSELECTlaQ CRITERIA. MOXL SELECTEB HAS MAXIMUMVALUE.
MODEL Mf01 MlHl M(B) M(BH1 M(T) M(TH) M(TB) M(T8H)

CRITERIA .539 1*00 .58 .74 0.00 .46 .53 .BO
w n X n rW r n r rn t
OVERALLTEST RESVLTS - - test statisticvalue (z-value)is -0.403,

Z-VALUE - . 4C
PR0{3ASI L I TY OF A SMALLERVALUE
and the significancelevel of the test is . 393E
.oot>. aus,torcou ome s cata,c Fo-
sureis notrejected.
TEST OF CLOSVREBY FREQUENCYOF CAPTURE.
(FREWENCIES LESS THAN 10 ARE NOT COMPUTED. )
If individualcapturefrequenciesare
10 or more, programCAPTUREalso
NUtwlBEROF CAPTURES Z-VALW PROBABI L I TY computesandprintsa partitioned version
________________________________________
of the closuretest forthosefrequencies.

.25477
For example,fromFig. 16, for animals - 660
- . 322 .37360
3 caughttwicethe testvalueis -0.660 and
-.066 .47356
4 is not significant.Thesepartitionedtest
.694
-1 .085
.75607
5
. 13906
6
cases are of interest,but we emphasize
the overalltest is the one to use forthe
FIG. 16. ExalnpIeof the test forclosureprocedure finaljudgement on closure.
with bral house mouse datafiom Coulombe(un- Failure of closure meansthat during
publishedthesis).
the studyanimalsare eitherenteringor
leavingthe.population at risk1ofr capture,
ommencluslngtneJacnlte estlmatorln or 30t. r a1S
1 . 1.
COU C. S)e causee,torexam-
. 1 1 .r n
suchcasesbecauseit is robustto heter- ple, by death,emigration, orthetrapgrid

ogeneityandwill tendto do well even if itselfattracting animalsfromsurrounding
a competlngmocle1S true. qort ze morn- areas(especiallylikely in removaltrap-
. 1 . _
ing data)the estimateof N fromthejack- ping).In anyof thosecases,the animals

knifeestimator w as 194+ 7.6,whilethat that enter or leave have zero capture
fromthe null estimatorwas 171+ 4.1. probabilities duringthetimetheyarenot
Hence bothchoicesgivenby the model partof the trappedpopulation.The cor-
selectionprocedureprovidedsimilares- respondingprobabilitymodelhas simi-
timates. laritiesto ourbehavioral ModelsMb Mbh,
As a finalcheckon whethertimevari- MtbandMtbh. Indeed,as previouslystat-
ationexistsin the formof morningand ed, behavioralresponsesare extrernely
evenlng( ltterences,notet zattaaerewere difficultto untanglefromtruefailuresof
. rr
norecapturesin theInornings (occasions closure. Our simulationresults have

1, 3 5, 7, 9) thanin the evening(occa- shown the closuretest rejectsstrongly
sions2, 4, 6, 8>10). whenModelMbis true,andis nota truly
unbiasedtest wheneverthere is strong
A Test Jor Closure behavioral variationin captureprobabil-
r 7
ities. sowever,t ae test cWoes notseemto

Throughoutthis luonographour phi- be affectedby heterogeneityor random
losophy has been that assumptions timevariations. We havenot used other
shouldbe tested.Oneof the mostcritical tests fromthe literature(e.g., Pollocket
assumptionsbehindthis entireworkis al. 1974) because, to our knowledge,
thatof populationclosure.Althoughit is those tests all are implicitlydeveloped
c esira) e to testc xosureS t zereareno tru- underthe assumptionthatModelMois
ly suitabletestsforthisassumption. Clo- t ae true modei under t ae c osure as-
sureis difficultto test for becausesozue sumption Becausewe feel Model MO
types of variationsin captureprobabili- probablyis neveracceptable,thosetests
ties (especiallybehavioral) aredifficultto will be even more untrustworthy than
distinguishfrolna failureof closure. ourcurrentclosuretest.
Burnhamand Overton(pers.comm.) The biologistis advisedto look care-
suggesta closuretestbasedon ModelMh fullyattheclosuretest.If theteststatistic
ct. Appenctlx X . r lg. lt glves an exam- is notsignificant,
r . 1 . Tr
thistendsto supportthe
- r ,} .
ple producedby programCAPTURE of validityof the closureassuluption. If that

this closuretest appliedto Coulombes testrejectsclosure,beforeacceptingthat
(unpublishedthesis) full data set. The result,it is necessaryto lookat the other
STATISTICAL
INFERENCE
FROMCAPTUREDATA C)tiSet al. 67
tests and the indicatedmodel.If strong problem.Two of theln rely on Dice's

behavioral variationis indicatedSthe clo- (l938) conceptof a boundarystripabout
suretest is notvalid. the gridsuchthatthe effectivetrapping
Additionalevidenceregardingclosure areais the gridareaplus this boundary
is obtainedfroma test of averagecap- striparea.Dice assumedthe boundary
turesper trapas a functionof trapdis- stripto be one-halfthe averagediameter
tancefromthe edge of the grid.Thattest of the home range of the population
is discussedin the sectionon DENSITY e1ng trappee. ze trst z approac
, 1 r n
zes at-
f)
ESTIMATION;it testsforwhetherthe grid tempt to estiluatethis strip width, W,

attractsanimals. from the capture data. These 3 ap-
Fina y, we mentiont lat t le c osure proachesaregivenbelow:
test used hereis orientedtowarddetect- (1) Holne rangesize is estimatedfroln
lng Dreatc£owns
* t 1 1
ln closureon y durlng
. 1 1 .
the locationsof differentcapturesfor

the initialandfinalstagesof the experi- the sameanimaland is used to esti-
ment.The test is not appropriateS forin- matethestripwidthW.Avarietyofap-
stance,foridentifyingsituationsin which proaches havebeen used;all areba-
animalsemigrateduringthe tniddleof sicallyad hoc andsubjecttonumerous
the study period and then ilumigrate problems, e.g., resultsdependupon
backto the studyareain the latterstages trap spacing and numbersof recap-
of an experiment. tures (Hayne 1949b Stickel 1954>
anaia 1Y {Y . .i le asic idea can De
r n s rert r n 1
developedintoanelaborate statistical
DENSITY ESTIMATION estimationproblem (Jennrichand
Introduction Turner1969)>but as faras we know
The luodelsdiscussedto thispointin- estimationof W l)asedon luovement
volve only populationsize N as the pa- (2) data The
remainsunsatisfactory.
rameterof interest.Often,interestmay parameter W is directlyestimat-
lie in populationdensity,the numberof ed basedon datadrawnfromselected
animalsperunitarea(e.g.,squirrels/hec- subgrids 19697
(MacLulich19517Hansson
tare).One couldnaivelytakeN divided Seber 1973:51 Smith et al.
by the areaenclosedby thetrappinggrid 1975). As suggested by Burnham and
as an estimateof density.Thatapproach Cushwa(pers.comm.)thatideacanbe
however,leadsto severeoverestimation developedintoa procedureallowing
as a resultof whathasbeen called'Cedge jointestimationof D andWfromdata
effect, i.e., notall animalshavetheiren- on 1 suffiiciently largegrid.We dis-
tirehozuerangewithinthe trappinggride cussthisapproach in detailbelow.
but tnay still be caughtbecause some (3) The use of 44assessment>' lines is the
trapsnearthe gridboundaryare within most complex approach to densityes-
their home range.Althoughbiologists tirnation. It involves designing the
have recognizedthis problemfor de- study to specifically estimate the ef-
cades (Dice 1938 1941; Stickel 1954)> fective trappingareaas well as the
statisticians concernedwithestiluation of size of the populationat riskof cap-
populationabundance havetendedto ig- ture. Therearenumerousvariantson
noreor lave failedto recognizetheprob- thisapproach (Smithet al. 1971?19727
leln. This probablyis due in partto the 1975, Swift and Steinhorst1976
factthatabstract modelsforcapturestud- O'Farrell et al. 1977).We have not
ies suchas ball-and-urn modelsS haveno pursued this approach here because
spatialcoluponent) hencedo notinclude the proper data analysis depends upon
any conceptof densityas distinctfrozn the study design.
nuln Dersot an1ona
1l nree
r
s.
. approac
as1c zes are g1ven1n Ofthose3 approaches, onlythesecond
the biological literatureto solve this seemsto be formulated in a rigoroussta-
.h X w X X
, ,e, X r X---
fi X X
wS X ,&, X X w ^ X st. I §
I
> . 0 .. , t r s, , o, i |
68 WILDLIFE MONOG>PHS
X X X X X X X X 8 X X X ^- s As
X X X X X X X X X X X X X X X
} 4 X
X X X X X X X K X X X X X-K X { l 3 1 1
X X X X X X X X X X X X X X X
X X X X X X X X X: X v X X X
t t 1 2 t | |
X X X w X X X X X X X ^ X X X
X w X v X X -w- X H K X X X w X I i I r n l I I
t l I I I I 0 t
X X X X X X X X X s X w X w X
X x X ^ X - X X X ^ X X X X X
X X X X X X X X X X X X X w X lrYrA_ I I I I
| I I
t _J | |
I I
:
X X X X X X X X X X X ^ X X X
t t _______________J
l |l lt
X w X X X X H- X X X X x X X X
;
X X X X X X X X X X X X )( X X } 1 1 1
l \_______________________ l
X X X X X X X-- X X X -K-- X Pi- x X t I
X X X X X X X X X X X X >S X X
l ,
!
- X- X K X X X X- x x x -K X }(--- X zX
F1G 1X. series vf 4 neswd subgridswith a
A
FIG. 17* A 17 x 17 trapplngd VY1dK
4 nested bourldarystrip of wldth W aroundeach subvid.
subgrids Note the torFwerof the boundaryseip is a quarter
cireIeof radlesW.
tisticalfashionwithoutarbit assump- in the calculationsT>ps were3 m apart
tions. IoweverS even the second ap- and the areaenclosedby the grid was
proachcannotbe tnadeworliablewlthout ()*073ha We can delineatesubvids of
sozue arSwaryad hee ieares.
sluallerslzeby examlnlng smallergroups
ProblemFormulon or t>ps ln pco ar we can coneew
tuallzea seriesof nestedsubvids(seefor
We will illustratetlle proljlemwth exampe filg JW . _t
sotnedatafroma liveWapping sdy of The size of the populationat risk of

feral house mice Mtks musculus in a canturefor each noustb stilnated
southernCallfornia coastalsaltmarshn The choice of a population estimator
ffrid
DecelnSerl962 by Coulolmle(unpub- snou d e ased on t e mockel select:ion 8 r % s s
ltshedthesis)5thatwere also tesell to il roceduresusin the dataforthe entLre

lustratethe tnodelselectionprocedure. griclFori - 1>2>** **kSassumethatapop-
0nly the basicspeelficationsof the sdy ulationestimateN hasbeenmadeforthe
will be neededhere. 100 Shernlal;w live ithgrid by using dataobtainedonly fr:>m
aps were laid out in a 1(1x 10 square that gid The naive estimatorof densit
gridXMorningandevenlngeapp1ngses- of the Lthgr3Wd 1s gven by
sionswereconductedforS daysandpro
vided 10 trappingoccasions7although YL Ni/AT
only the morningsesssonswill be used i iS2>.*.>kS
TABLESO.THE INFOAoN NEEDEX3

TO ES4E DENSITYUSEX T W NESD SCX9S AND
TH:EDATAFROMCOULOMBE
(PUBLISHED THESIS)
2 . - t
No. Grid AreajAi (individllals) ( indiviZoals ) Be(ts)
t 4x4 innergrid 0.r81 47 5770.9 607.8

2 (i X 6 mlddlenner ld O*V225 tO7 47S0*7 452.6
3 8 X8 mldElleouSrvid Q+N41 lM 3768.S S43.4
4 IO X IO nrE 1 0t0729 IN S69.4 IN*9
INFERENCE et al. 69
whereAiis the size (area)of the ithtrap- of trapsin the shapeof a squareor rec-
pinggrid.The griddataaregivenin Ta- tangle.Thatis, by connectingthe outer-
ble 20. Notethatthe standard errorof Y mosttraps,a squareor rectangleresults.
is Se(Ni)/Ai Let P be the rneasured perimeterof the
The fourYivaluesin Table20 cannot grid LetA be the areawithinthisperim-
reasonablybe consideredeach an un- eter,andletA(W)equaltheareaobtained
biased estimateof a single pararneter.by addinga boundarystripof constant
Thebiasis attributed to thephenomenon W1Ct 1 w. ilen t le tuntamentalre atlon-
. sxY r n fb 1
of edge effect,whereinthe areausedby shlpS 1S

1 . .
individualsat risk of captureis not re- A(W)=A+PW/c+7rW2/c,

strictedto the areacontainedwithinthe
trappinggrid;rather,thereareareasout- wherec is a conversionfactorto express
side the trappinggridthatarepartof the PWor w2 in unitsof A. Forexaluple,to
homerangesof individualsat riskof cap- convertm2to ha,c = 1,000InSha.
ture. Let there be k such gridsidentified.
Weassumethatthepopulationdensity These maybe subgridsof one gricl,or
is constantin the areaof trapping.That luaybe physicallyseparatestudygrids.
is) thereis not a trendin densityacross Assumea constantdensity applies for
thegrid.Undertheseconditionsit is rea- eachgrid.Then,forgridi we wouldex-
sonableto thinkthatthereis an areaof pectto have
constantwidth aboutthe trappinggrid Ni = D Ai(W)- D[Ai+ PiW/c
suchthattheactualareausedby thepop- + X W2/c],
ulationat riskof captureis the totalarea
of the trappinggrid plus that of the whereNi is the population at riskof cap-
boundary strip.LetWequalthe widthof turewith respectto the ithgriclof traps
this strip.This is illustratedforthe four only.Conceptually, we shouldlet theac-
gridsin Fig. 18.Theconceptof a bound- tualnulubersof individuals atriskof cap-
arystripgoesbackat leastas faras Dice tureon gridi be a randolnvariablewith
(1938).We need not interpretW, but expectationDAi(W).Then we woulel
fromthe inceptionof the boundarystrip write E(Ni)= DAi(W),and it is really
concept,biologistshaveconsideredthat E(Ni)we are estiluatingforthe ithgrid.
W is relatedto home rangesize (Dice Lettingi= 1) . . ., k we oltain structural
1938, 1941) In fact they arbitrarily as- equationsrelatingthe parameters D) Wt
sumeW is one-halfthe Inaximum linear andthe induce(:lparallleters Ni.Thearea
dimensionof the averageholllerangeof Ai, and the grid perilneterPi tnust l)e
the species. known.Nextwe redefinethe basicstrue-
turalequationsas
Statistical Treatment
y _ Ni
i - D[1 + aiW + biW2]
The approachto density estimation
thatwe advocateforuse with gridtrap- i = 1, 2, . . . k,
ping is to formulatethe problemas one
of jointestiluationof D andW, withthe where
paralneterN being a functionof D and
W. Then,by having2 or Inoregridsof ai = Pi andhi =
Aic Aic
differentsizes, we can estiInatethose
parameterswith a weightednonlinear Assumethatfrolnthe trappingclataof
leastsquaresprocedure. each grid we have estimatesof the Ni,
In whatfollows,we assuluethatden- expressedas Ni, and estiluatesof their
sity (D) is expressedin individualsper variancesVar(Ni).ASSUme furtherthat
unit areaand stripwidth(W)in linear the Ni are good estimatesin the sense
units,suchas meters.Considera layout thattheyhavesluallbias,so we canwrite
innergridlargeenoughso thatN1is well

Ni = yi = D[1 + aiW+ biW2]+ e} estimated.Otherpossibilities(not rec-
Ai
i- 1>2>...k, ommended)areto includesubgridscon-

sistingof halvesandquartersof the total
wheresi is a randozn errorwithE(e)- 0, gridSorboth.
and covariancematrixE(e st)-}. The The basicidea behindthis procedure
smallbias of the Ni is a big assumption wasintroduced by MacLulich (1951)>
but
thatrelies heavilyon previoussections. until recentlyno methodof estimating
Note that Yi- Ni/Aiis what we would thevariancesof D andWhadbeengiven
call the naive estimatorof density,D, (Smithet al. 1975).
fromthe ithtrappinggridSclearlyit can In Fig. 19 a completeanalysisis pre-
be significantly biasedunlessAlis large sentedforthe California saltmarshferal
relativeto (PiWlc+ xW2/c),the areathat tnouse study. First note that we have
is addedby assuminga stripwidthof W. computedthe matrixof capturespertrap
Notethatfora sequenceof k subgridsof station.Thatmatrixincludesthe datafor
increasingsizeSthe mostbiasedestimate both the morningand eveningtrapping
of D is obtainedfromYl becauseAl is sessionsS orforall 10occasions.Multiple
smallestrelativeto (PlW/c+ 7rW2lc). On capturesper trapcause some entriesto
the otherhand,Ykis the best estimateof be greaterthan 10. Visualstudyof this
D becauseAkis largestrelativeto (PkW/ matrix(Fig. 19a)does not discloseany
c + vW2/c).If we had a verylargegrid, grosstrendsin mousedensityacrossthe
saythestateofWyomingthecontribution grid.Fig. l9b showsthe resultsof 3 chi-
ofareaduetotheboundary stripWwould squaregoodnessof fit testsywhere the
be negligible,and we could ignorethe capturematrixis firstcollapsedby rowsS
problem.Becausethisis notthecaseewe then by columns5and finallyby rings.
use ourbiasedestimatesof Yito findan Thosetests generallytend to rejectthe
unbiasedestimateof D. nullhypothesisof uniformprobability of
WeassumeE e fr) -X becausein gen- captureby rows or by columns.HQW-
eral the naivedensityestimates,YiSare ever5we are moreconcernedin identi-
not incbependentfrom one anotherif fying grossirregularities in mouseden-
thesesulogrids derivefrom1 overallgrid. sity suchas no capturesin one cornerof
Hence,we mustestimatethe covariance the gridSor a strongtrendof decreasing
Inatrix Aiof thek estimatesof Yi.A simple probabilityof capturefromrightto left.
andintuitiveprocedureis to assumethe The tests identifya problemof higher
correlation betweenYiandYJis just the trapsuccessat t ze gridedges.Thisis in-
proportionof overlappingareabetweem dicatedby the testof rowsSwherea large
gridi andgridj includingtheirboundary portionof thechi-square valueis contrib-
strip.Thenthe covariance of YiandYJis utedby the ninthandtenthrowsyandby
Se(Yi)*Se(YJ)COrr(YiS YJ).With the re- the ringtestSwherethe outsideringof
sulting covariancematrixSgeneralized trapshad muchbettersuccessthanex-
nonlinearleastsquarescanbe performed pectedSanc>t ze innerrsngpoorersuccess
to estimatedensityandstripwidthas D thanexpected.ForlargetrappinggridsS
and WSrespectively.Note that$ is a the problemcouldbe correctedby not
functionof WSbecause the amountof usingthe dataforthe outerringof traps.
overlapbetween grid i and gridj is a This is very wastefulof dataShoweverS
functionof the estimatedstripwidthW. andwe will ignorethe problemhere.
In the mostcommonsituationS thereis ze next4 pagesof outut (Figs.19e-f)
one gridsuchas ourexamplein the Cal- consistof the populationestiluatesfor
iforniasalt marshS and one mustchoose thewholegridandeachofthe4 subgrids
a smallnumberof subgridsto use in this coluputedwith the jackknifeestimator
approach. Wereeommend 4 ormorenest- derivedfromModelMh.Thatmodelwas
ed grids.Themainconeernis to keepthe selectedbasedon the ouWutforthetests
tt r .
STATISTICALINFERENCEFRQMCAPTUREDATA{)tTS et al. 71
MATRI X OF CAPTURES PER TRAP STAT I ON.

luatesof D andW.The columusheaded
uts
_ _
2
_ _ _ _
3
_ _ _ _
q
_ _ _ _
5
_ _ _ _
6
_ _ _ _
7
_ _ _ _
8
_ _ _ _
9
_ _ _ _
10
_
NAIVE DENSITY PEEMETEWAREAs and ^r 1
to YiS aiy and iS re-

- - - - - - - +
l - PI/AREA corresponc *
RW 1 1 9 q 6 6 6 7 5 5 9 6
spectivez -Y1and the COVASANCE MATEUX
RSl 2 1 12 6 4 7 2 W 4 2 4 8
corresponds to $ forthe initialvalueof
R)l 3 1 W 8 9 7 4 8 3 3 3 7 W. 1 ze rlumzerot lteratlonsrequlrec . to
R(IU 4 1 B 9 8 Z 4 10 6 7 6 3 + the paral
estllnate ute ers 1S slVen1
' aldfr
l O
Grid
R1EJ 5 1 7
this exampleit is 189. The maximum
9 4 4 6 4 7 7 6 5 1
nutuberallowedin prograzn CAPTURE 2

ROU 6 1 5
-4 is 300. The approximate
8 3
nuluberof sig- 4 3 4 9 5 4 5 3
ROW7 7 1
nificantdigitsof D andWis alsoprinted.
6 I 8 3 3 5 2 3 6
Rt);J 8 5 1 A tablelistingthegridnuluberS
5 1 thenaive 5 7 2 4 6 5 3
RS 9 1 5 density1andthe predictedvalueis then

9 6 6 10 5 4 7 B 14
RO#J10 1 IO giver.Thisallowsthe investigator

9 6 to get 8 8 6 5 8 11 9
a feel forhowwell thezuodel fitshis data.

FIG. 19a. Nested subgridsused ill the deuxity Notethatall the residuals in theexaluple
estinlationprotedurewii iraI house mousedata arenegative;this is becausetheyareall
iom Coulolube(unpublishedthesis).In the matrixS highlycorrelatedS and hence we expect
trapcoonlinate§are roundedto it nearestwhole sign.Thelnultiple
integer. In goodnessof St testsytrapcoordinates theln to be of the salne
that are not integersand noarectangular trappiLngcorrelation coefficientS RSis also printed
gridswill cause spuriousreswtlts. to help assessthe fit of the luodel.The
valueof R2is interpreted as the propor-
describedin the section on model selection of the variance in the data thatis ex-
tion(E;ig.15). plainedby the lnodel(0 S R2S 1). The
The final page of output(Fig. 19g) computedvaluesof 1) andW arehighly
gives the generalizedleast squaresesti- negatively correlatedSindicating the
.
CHI -SQUARE TEST OF UNIFORMDENSI TY BY ROWS
z 4 6 8 9 10
ROW
53 56 63 59 50 44 43 74 80
ORSERVEO 63 58.500 58.500 58.500 58.500 58.500 58.500 58.500 58.500 58.500
EXPECTEO 58.500 .517 . 107 .346 .00# I .235 3.594 4. 107 4. 1 07 7.902
CH I -SaJARE . 3W6
22 . 26 W1TH . PRO2ASI L I TY OF LARGERVALUE=

9 DEGREESOF FREEDOM . 0081
TOTALCHI -SWAK a
.
CHI -SaJARE TEST OF UNIFORMDENSI TY LY COLUMNS
4 6 8 9 lo
COLUt 2
OBSERVED 72 73 48 57 53 53 52 52 59 66
EXPECTED 58.500 58.500 58.500 58.500 58.500 ;Z8 . 500 58.500 58.500 58.500 59.500
CH I -SQUARE 3. 1 15 3. 59 I . 885 .038 .517 .517 .722 .722 .004 .962
l 2 . 08 W I TH 9 DEGREESOF FREE . PROBA2I L I TY OF LARGERVALUE= . 2090

TOTALCH1- =
CHI-SaJARE TEST OF UNIFORtl DENSITY SY RIKS (NTER RlK IS ER I).
2 3 4 5
RING
OBSERVED 2q7 160 96 65 17
EXPECTED 210.600 163.800 1 17.000 70.200 23.400
CH X-SQl JARE 6.291 .088 3.769 . 385 I .750
12.28 WI TH q . PROSASIL I TY OF LARGERVALUE=

DEGREESOF FREEDOM . 0154
TOTALCHI -SOUARE s
FIG. l9b. Chi-squaretests of uniformdensitywithferalhousemousedatafromCoulombe(unpublished

thesis).
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ of Fig. 19a(X - Y - 3 8).
hUMBER QF tRAPP I NG OCCAS1 WAS 5 NU"BER OF TRAPPING OCCASIONS US 5

NU"3ER OF ANIS CSTURED. MfTv I ), WAS 33 NUMBEROF ANIMALS CAPTUREO. H(TsI }, WAS 67
rorK NU"BER OF CAPTURES. N., UAS 50 TOTAL NU"BER OF CAPTURES. N ., WAS 110
FREWENCIES OF CAPTURE8tl I } FREOt,JENC

IES OF CAPTUREF ( I )
ww X e 3 X 5 1^ 1 2 3 # 5
Ft 11* 18 13 2 O O Ft 11. 37 21 5 # 0
I ENTS
CUTED JACKKNI FE CWFF I C COMPUtEOJACKKNIFE COEFFIC1ENT5
NX3) NlQl Nf5) Nl I ) N(Z) Nt3) N(W} Nt5

Nt I )
Nt2 e . S00 3.000 3. DOt 1 1.800 2.90Q 2.800 3.000 3. OQ0
X X. eoo 2. 00 .050 - .250 - . 25Q e l . OoO . 55C .050 - .250 - .250
2 1 .000 .550 I . 133 f vU50 1 wE50 3 1 . Ooo 1 . QQo a . 133 I, ZS0 1 .250
3 1 . OOQ I .000 l I .000 1 .000 1 . ooo .992 . S9e
l . 000 .992 . g9E!
o I .000 I .000
I .000 1 OQ0 l . OOQ 5 1 .000 1 .000 1 .000 I .000 l . Q00
5 l . QOO I .000
THE RE5ULTS OF THE JACKKNIFE CUTATI THE RESULTS OF THE JACKKNIFE COMPUTATIONS
Nl I 1 SEX 11 .95 CONF. LIMIT5 TESTr OF Nt 1+1 ) VS. NX11 Nt t ) SEI I ) .95 CONF. LIMITS TEST
r OF N( 1+1 ) VS, Nl I )
o rHI-513UAKt l D.F. 1
33 Cl o 61 vHI -SQUAREf I D . f . }
Cl
I 2.839
47.4 5.09 37.4 S7.4 l 96*6 7.30 82.3 1 10.9 10.573
e . 198 e 109.#
1 10.91 88.0 130.7 e.23l
52.4 7.57 37.5 6v.e
3 . ooe 124.3 13.65 87.E t4} . I .789
53.3 9.51 34.7 12.0 3 1
53.3 10.62 3e.4 79.1 0. QOO 4 1116.0 15.17 86.2 lW5.7 O.OOD
5 O .000 1116.0 15.17 86.2 1q5.7
53.3 10.62 32.W 74. } 5 O .000
AWRAX P >-HAT
AVERAGEP . Z056
'-HAT - .2128 a
INTERPOLATEOPOPU.ATlON ESTIMATE IS 47 Ul TH STAARC1 ERROR # .> INTERPOLATEOPOPlATION ESTIMATE IS 107 W} TH STANDARDERROR 10 . l 827
APPROXIMATE95 PERCENT CQNFIDENCE INTERVAL 37 ro 57 APPROXIMATE 95 PERCENT C 1DENCE I NTERVAL 86 TO IzV
HlStO0RAH w Ft 11 HISTOGRAMOF f ( 1 }
FRECOUENCY 18 13 2 0 0 FREOUENC:Y 37 Z1 5 4
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ .
EACH * EOUALS 2 POINTS EACH ' EQUALS 4 POI NTS
8 36
32 *
14 * 28
12 * . e4
10 * X eo * *
8 * X 16
6 * * le
4 * . 8
e 4 * * *
FIG. 19C. Exampleof populationestimatiQu wiffi FIG. 19d. Exampleof populationestilnationwith

variableprobabilityof capturehy allimal under variableprobabilityof captureby animal under
Modellqh w1thiral house mousedata(Coulotnbe ModelMh with bral house mousedata(Coulonlbe
(ullpublishedthesis) frolninner innergrid of Fig. unpublishedthesis) from the middle inrwergrid
19a(X = 4-7 Y- 7).
stronginverserelationbetweendensity animalsythus zCinducing>> a higherden-

andstripwidth.Thisnegativecorrelation sitythanwouldotherwisebe found.The
makesit difficultto estimateeitherpa- probem ofthegridattracting animalscan
-
ralneterwith a smallstandarderrorS because severe zlas. t attractionoccurs . rW
cause other combinationsof D and W duringtrapping,then the assumptionof

also resultin almostas gooda fit as the populationclosurewill be violated.With
valuesselected Finallya testofwhether removaltrappingS over a long enough
W is significantlydifferentfromzero is timethe problemis sureto develop.The
printed.In thatexamE)lethe highlysig- problemmaystill occurir}livetrapping
nificantdifferenceof W fromzerv indi- studies.One possibletest forthis effect
catesthatthe estimateof D is muchbet- is the CCring>>testgivenin Fig. l9b. Even
ter thanthe naiveestimatebasedon tlle withsomeapproach likeprebaiting to al-
actualgridarea. low this attractioneffectto stabilizebe-
fore trappingboth methods(recapture
DTSG8SSXOB movenaentsand directestimationof W
arldD) will be invalidif the griditself
ThedensityestiInation problemresults attractsanimals.In thatcase it appears
entityin assesslnentlines wouldhaveto be used.
becausethe gridis an artiIScial
the environmentand animalstrapped If the abovemethodproducesa poor
use areasboth iIlside and outside the resultSan alternative approach is to base
grid. A practicalproblemillustratedln an estimateof W on animalmovements
the exampleis thatthe grid can attract as determinedfromrecapturelocations.
FROMCAPTUREDATAtiS
INFERENCE
NUMBEROF TRAPPI Nt; OCCASI ON5 WAS 5

NUMBEROF ANI MAL5 CAPTUREO , Mt T+ 11, tWAS 160
NU"BER OF TRAPPlhG OCCASIONS WAS 5
TOTAL NUMBEROF CAPTURES, N ., WA5 356
NUMBEROF ANIMALS CAPTUREDs M(T+I }, WAS I 16
TOTAL hW1BER OF CAPTURES. N., WAS 202
FREQUENC I E5 OF CAPTURE,F t I )
1- I Z 3 4 5
FREQUEhClES OF CAPTURE.FZI 1
FX I }s 45 59 36 15 5
1- 1 2 3 4 5
F I 1- 55 43 11 7 0
CO+UTED JACKKNI FE COEFFI C IENT5
CO"PUTED JACKKNI FE COEFFl C IENTS
NX I N(2 N(3) N(4) N(5)
N(3) N(41 Nf 5 I I .800 2.800 3.000 3.000
Nl I ) Nt2) Z .400
1 1.800 2.400 2.800 3.000 3.000 2 1 .000 .550 . 050 - .250 - .250
2 1 . 000 .550 . 050 - .250 - .250 3 1 .000 I .000 I . 133 1 .250 1 .250
3 1.000 1.000 1. t33 1.250 l .250 4 1.000 I .000 I . 000 .992 .992
4 1 .000 | .000 1 .000 .992 .992 5 1.000 I .000 1 . OOG I .000 1 .000
5 1 . ooa 1 . ooo 1. ooo ] . ooo i .000
THE RESULTS OF THE JACKKNIFE CO"PUTATION5

ION5
THE RE5ULTS OF THE JACKKNIFE C0MPUTAT
N(ll SE1I} .95 CONF. LIHITS TEST OF N(lsl] v5. N{l)
Nt I ) SEX I ) .95 CONF. LIMXTS TEST OF Nl l+l ) V5. Nt o 160 CHI -5QUARE( I D . F . )
o
1 16 CHI-5OUAREI l O . F . ) l 196.0 8.05 180.2 211.8 .007
160.0 8.90 142.6 177.4 6.866 2 196.5 1 1.69 173.5 219.4 2.000
2 173.7 13.20 147.8 199.5 .195 3 189.8 15.15 160.1 219.4 2.847
3 175.6 16.64 143.0 208.2 .073 185.1 17.31 151 ve 219.0 0.000
4 t8.62 138.4 211.4 0.000
174.9 5 185.1 17.31 151.2 219.0 0.000
5 l74.9 18.62 138.4 Zl t .4 0.000
AVERAGEP-HAT = 243 AVERAGEP-HAT w .3670
I NTERPOLATEOPOPULATI ON EST I MATE I S 166 WITH STANDARDERROR 10.7335 INTERPOLATEDPOATION ESTlMATE IS 194 WITH STA<ARD ERROR 7.6455
APPROXIMATE95 PERCENT CONFIDENCE INTERVAL 1q5 TO i 88 AOX I MATE 95 PEtENT C I DEKE I NTERVAL 178 TO 209
HISTOGRAMOF F(l) HISTOGRAMOF Ft I )
FREOUENCY 55 43 11 7 O FREOUENCY 4S 59 36 15 5
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
EACH * EOUALS 6 POtNTS EACH v EQUALS 6 PO I NTS
54
48 60
42 * * 5q
36 ' 48
30 42 * +
2W * 36 *
18 * 30 * .
12 * * 29 * *
6 * * 18 * * X
12 * * * *
6
FIG. Exalnpleof populationestimationwith
19e.
with
variableprobabilityof captureby animal under FIG. l9f. Exal}ple of populatin es;tilllatiear}
ModelMhwith feralhouse mousedata(Coulombe variableprohabilityof capttlre1)v allinal under
wlnpublishedthesis) from the middle outer grid Mdel Mhwith iral hovIsellluse clrlta(Coulombe
of Fig. 19a(X - 2A91Y = B9). unpublishediesis) iom the entire grid of Fig
19a (X = 1-lOs Y - 1-1()).
Let W be suchan estimate,forexample andthe standard errorof D is simply

one-halfof the averagemaximumdis-
tance Detweentrappedlocationsfor all Se(D)- +/tar(D).
animalscapturedatleasttwice.Thestan-
darderrorof W,Se(W),canbe computed Thesevarianceformulasarevalidwhat-
fromthe datathemselves.The estimator everthe techniqueforestimating Wfrom
of densityis then recapture locations.
D- N In the exalupleof theferalhousemice
A(W) W is calculatedas 3.63 m with Se(W)-
0.149 and comparesfavorablywith the
whereN is basedon all the data(i.e.,the estimatedvalue of W= 4.65 frolnFig.
entiregrid)and l9g.
A(W)= Ag[l+ aW+ bW2] The methodsoutlinedin this section
require largeamountsof datato achieve
fora andb as definedpreviouslyin this satisfactory results.Botha largetrapping
section. An estiluatorof the salupling gridanda largenuluberof capturesare
varianceof D is givenby required.A grid size of 9 x 9 probably
Var(D)- Var(N) can be consideredthe minilnuln;how-
[A(W)]2 ever a largergrid such as 15 x 15 is
lnuchbetter.Goodtrappingsuceessto
+ (D)2(1+ aW+ bW2)r(W) achieve a large numberof capturesis
necessaryto providea usefulpopulation
74 MONOGRAPHS
WILDLIFE
STARTING VALUES FOR DENSITY ESTIMATION-- movals.However, proper planning,

andfieldconductof suchstudies
4
NUMBER 0f GR I DS
TRAP I NTERVAL
3 °°
10 00 0 . 00 designS
UN I TS COtSVERSI ON
I N I T I AL DENS I TY ES T I MATE
I N I T I AL STR I P W I DTH E S T I t1ATE
. 5089
882 . 6797
necessaryto obtainmeaningful
is datafor
GRID
I
NAIVE DENSITY
Yl I B
PERIMETtR/AREA
At I 1
Pl/^REA
8( 1 )
STARTING COVARIANCE MATRIX
Many
analysis. factorsmlust be consid-
1
2
5770.927
4750.710
.4444444
.2^^&i67
. 3879E-0 I
. 1396E-01
. 369E-06
.273E+06 .205E+06
.592E+05 whenplanninga capture-recapture
ered
orremoval study to estimate animal
.7124E-02 . 146E-06 . 109E+06
3 3768.521 .1904762 .1 10Es05
.Ei25E-05 .468E+05 .254E+05
.1481481 .4309E-02
4 2.554.402
abundance.We considerthese as falling

2 broadcategories:statisticaldesign
0f ITERATIONS
REStJLTS 189
FUNCT I ON E VALUAT I ONS REQUI RED
ESTI"ATED SIGNIFICANT DIGITS OF PARAt1ETER
VALUES 8
into
anddata recordingconsiderations, and
F I TTED ItOOEL CCt1PARED TO THE DATA
procedures,
field althoughthe distinction
GRIDt I )
1
2
Yt I )
5770.927
4750.7 10
Ft I )
5505.869
3583. 134
may sometimesseema bit arbitrary. Ex-
3
4
3768. Y
2654.402
1 2875 . 007
25 1 1 .652
amples of such considerations include
arenotlimitedto)the following:
COMELATION COfFICIENT IS .93181
MULTIPLE
OENSITY-
ESTltRATEO
STRIP WIDTH-
ESTltbtATEO
1408.934
4.653
172.1260
1.0576
-
-
ITS STANOARD ERROR
ITS STANARO ERROR
(but
-.9471
Statisticalcorlsiderations
CORRELATION Of ESTlt1ATORS
ZERO.
OF ESTIMATED STRIP WIOTH GREATER THAN
TEST
numberof captureoccasions
.0000
Z-VALUE
s 4.3998 PRO8A8ILITY OF LARGER VALUE -
F I NAL COVARI ANCE ÂTR I X
timebetweenoccasions
size andshapeof trappinggrid
. 3694Es06
. 1560Ev06 .2048E-06
.5398E+05 .7085E-05 .5924E+05
.2127Ev05 .1779E+05 . 1100E+05
spacingof traps
.1621E+05
Fl6.lGg Exampleof joillt estimationof (lensity numberof trapsat a point

stripwidth with bral hollse 1OUSt
and bt}llndary
dataSom Coulombe(unpublishedthesis). numbering of traps
Field procedurecoresideratior7s
estimatefor each grid.This becomesa >captures
aSlvevs. removai
.
problemespecially with the smaller choiceof traptype

grids,whereonly a fractionof the data c lolce ot e ectronsnlnggear . o f}
areused to estimatethe population.Be- methodof markingortagging

sidesrequiringa good populationesti- methodof recapture
mate,a goodvarianceestimatefor N is use of bait
required becausethatquantityis usedin timeof dayto checktraps
constructingthe estimateof the weight- handlingthe animals
ingmatrix}-1 Poordataresultin a poor The purposeof this sectionis to present
weightingluatrixS whichthen resultsin
poorestimatesof D andW. somesuggestionsandcautionsregarding
Possible methods to increase the the aspectsof studydesign.The general
amountof dataare(1)to placetrapsclose themespresentedareto conductthe ex-
perimentso that(1) assumptions canbe
togetherto increaserecaptures(2) to tested,(2) the closure assurnptionis met,
place gridsin a uniformhabitatso that (3) the simplest possible modelis appro-
discontinuitiesin densitydo not occur, priate7and (4) the numberof animals
and(3) to increasethe numberof traps. capturedis maximized(includingrecap-
r
o repeat t le meta
n
zOCIrequlres<arge . 1
atnollntsof data.A carefullydesigned tures). We begin by mentioning

studyis reguiredto reliable
obtain values livetrappingversus removalmethods,
of D and W andonly can
rarely a typical then discuss closure proceedthrough
studybe madetoyield waysof eliminating variationdueto tilne
capture-recapture behavior,and heterogeneitySand then
reasonableestimates. c Wiscuss sample size considerations such
as grid size andnumberof traps.
STUDY DESIGN For acciitionaldiscussionof design
The objectiveof this monographis to consideration in gridtrappingtlle reader
is referred to Overton and Davis(1969),
presentrnethodsforthe thoroughanaly- Smithet al. (1969,1971,1975),Tanaka
sis of capturedatawhenthe targetpop- andHansson(1974).
ulationsareclosedexceptforknownre- (1970)7
FROMCAPTUREDATAtiS
STATISTICALINFERE1NTCE Ct 1.
75
It is beyondthe intendedscopeof this riods of recuitlnent(e.g., juvenilesbe-

uonograp ] to provideguidanceon the comingtrappable) or iluluigration.Also
operational aspectsof capturestudies,al- keep the durationof the experilnentas
thoughtheyareimportant. Forexaluple, shortas possible.If it is necessaryto
if the methodof luarking(ortagging)is studythe populationat suchtimes,open
suchthatluarksarelost,thena basicas- populationlmodelsshouldbe usecl for
sumptionneededforlneaningfulresults dataanalysis(Seber1973,Pollock1975,
is violated.Whenmakingdecisionsabout ArnasonandBaniuk1977).
a fieldstudy,the scientistshouldconsid- Gridsare oftenthoughtto attractani-
er the probableeffectsof the experiInen- uals. For example,when kill trapsare
tal designon assuluptions necessarylor used, anilualsfromthe areaaroundthe
dataanalysis.For additionaldiscussion gridwill luoveontothe gridas localan-
of fieldprocedures the readeris referred ilualsareremoved(cf.Gentryet al. 1968,
to Davis (1956),Southwood(1966),and Sluithet al. 1975:38).This violatesclo-
TaberandCowan(1969). sure,ancltheonlygoodsolutionseelusto
be to keep the lengthof the study(in
Livetrcipping VersusRemovalHethods days)veryshortso thatthestudywillend
beforesignificant canoccllr.
imluigratioll
As discussedin the previoussections,
reluovalInethodsare a specialcase of ElimitlatingVarifltiotlDue to Tiz7le,
livetrapping methods.Thatis, theremov- Behavior,atld Heterogeneitzy
al estimators canbe usedon livetrapping
data.Hence, we recolnmendthat live- Given that closure is satisfactorilv
trappingInethodsshouldbe usedif pos- achieved,the next luostimportantCO1;-
sible becauseof the widerarrayof op- siderationis twofold:(1) achievinga
tions availablefor the dataanalysis.A largeenoughnumberof capturesto ob-
hazardof removalstudiesis thattheydis- tainreliableresults,and(2) achievinga
ruptthe population,and as substantial studyforwhichthebestmodelis thesim-
aniluals are removed) ilumigrationluay plestpossibleone (e.g.,ModelNl0rather
occur;thisviolatesclosure. than/ttbh, or ModelMb ratherthanNItb)*
For livetrappingstudies,all possible In this section,we discusslnethodsof
precautionsshouldbe takento prevent eliIninating variationof captureprobabil-
deathsof the anilualswhile in the traps, ities due to time,behavioralresponseto
e.g., shadingthe traps in sulumer,or firstcapture,and heterogeneityof indi-
avoidingperiodsof extremecold. Simi- viduals.
larly,it is assumed(iluplicitly)thatthe Ofthe3 factorsthataffectcaptureprob-
lnethodof markingwill not induceluor- abilities,timeis the one mosteasilycon-
tality.If substantialmortalitycannotbe trolledby thebiologist.He canselectthe
avoided,thenone mustanalyzethe study seasonof the yearthe studiesare to be
as a removalexperiInent. conducted,the lengthof the trappingpe-
riod,andthe tilneof daywhentrapping
Closure is to be done.In all thosedecisions,the
objectiveis to reducevariationin capture
For the analysislnethodspresented probabilitiesover time. Amongother
here,the singlemostimportant assump- things, this means that equal effort
tiOllis closure.Closureis verydifficultto shouldbe expendedon each occasion.
test for,yet anyviolationof thisassump- For exaluple use the same nuluberof
tionbiasesthe testsandpopulationesti- trapsthoughout, trapat the sallletimeof
uatorspresentedhere. day,andif baitis used,use thesalnetype
Considerations to help assureclosure andamounton all occasions.The study
include,forexample,tiluingthetrapping shouldbe donewhenweatherconditions
to avoidknownluigration tilnes,andpe- will be as constantas possible,because
variableweatheris likely to causetime eachtrappingoccasion.Weareunsureof

variationin captureprobabilities(Getz the usefulness of the technique, and
1961). would like to see furtherstudies con-
Behavioralresponse is commonin ductedto see if significantreductionsin
small mammals,and it is doubtfulif heterogeneityresult.For logisticalrea-
muchcanbe doneto avoidit. Apparentlysons,the randomization of traplocations
for this reason, many biologists have on eachoccasionprobablyis notfeasible.
studiedandused strictremovallnethods A valuablecheckon the livetrapping
forsmallmammal population estimation. methodspresentedin this monograph
If thereis any choice,a methodof cap- canbe luadeby usinga secondmethod
ture (for livetrapping)shouldbe used of estimatingthe proportionof luarked
that will not resultin a trapresponse. animalsin thepopulation. A columonex-
Oneapproach is to use differentmethods ampleis killtrapping orhuntingafterthe
of captureon eachoccasion(cf. Overton capture-recapture experimenthas been
andDavis1969).Thisis anexcellentpro- completed.Then, using the numberof
cedure,but its applicationto multiple markedanilualsin the population,Mt+1,
captureoccasionsis severlylimitedbe- andthe ratioof markedto unmarked an-
cause one cannotusuallyfind 5 or 10 imalsobtainedfromthe killtrapping or
quite different capture lnethods. We hunting,a secondand somewhatinde-
note, however,thatthe use of different pendent populationestiluate is made
trappingmethodsprobablywill resultin with Chapman's(1951)versionof the
timevariation(thedifferentmethodsoc- Lincolnestimate,
curof necessityat differenttimes).Con-
sequently,there is a trade-offhere: re- (Mt+l + 1)
ducedbehavioral variationmayresultin xT= (allanimalskilltrapped + 1) _ 1
increasedtilnevariation. (marked animalskilltrapped + 1)
A columonsourceof heterogeneityis
the lackof equalaccessto trapsif traps (cf.Seber1973).Ofcoursetrapsaretraps,
are far apart relative to home range andkilltrapsstillpresenta strangeobject
(Eberhardt 1969a).The abovephenolne- in the anilual'senvironment.Another
na are partof the reasoningbehindour possibilityforsmallmammals markedby
recomlnendationof 4 trapsper home toe clippingis to use tracksto obtaina
rangewhenwe discusssamplesize con- ratioof markedto unmarked animals.Bi-
siderationsin the nextsection. ologistshaveusedsmokedplates(Justice
Othersourcesof heterogeneity aredif- 1961)or dustcoveredtiles to obtainan
ferencesin activityorcatchability related estimateof the proportion of markedan-
to measurablecharacteristicssuch as iluals,and thusa secondpopulationes-
size, age,or sex. Suchsourcescanbe re- tiluateto comparewiththatobtainedby
luovedby stratification if the attributeslivetrappingmethods.The use of tracks
are recordedand samplesize permits. hasthedistinctadvantage ofbeinga truly
Unfortunately, samplesizes are seldonl differentmethodof saluplingthe popu-
adequateto allow stratification. Hetero- lation.Muchconfidencein the popula-
geneityluayalsobe due to someunrec- tionestiluateis obtainedwhellthe 2 es-
ognized attribute,and thus cannotbe timatesaresimilar.
eliminatedby stratification.Different In additionto design considerations,
methodsof captureluay increase the poststratification of the datacanbe used
numberof aninlalscapturedin thiscase, to createsubsetsof datawhicharemore
butwill noteliminatethe basicproblem holuogeneouswith respect to capture
of heterogeneity of captureprobabilities.probabilities.
Thisis nothingmorecoln-
One possible Inethodof eliminating plex thanpartitioning the datainto sub-
heterogeneity, and possible trap re- sets on variablessuch as species, sex,
sponse, is to locate trapsrandomlyon age, weight,etc. If there are sufficient
FROMCAPTUREDATAtiS
INFERENCE
is a valuable
data,suchpoststratification estimatestripwidthWas well as N. This
device.The subsetsso createdare then requiresthatthetrapsbe laidoutto eover
analyzedseparatelyby the methodsde- anarea,ratherthanin a singleline.Nlore-
scribedin this monograph.The onlyad- over,we mustbe able to associateeach
ditionaltestingone mightdo is for ho- trapwithanarbitrary X-Yeoordinate sys-
mogeneityaluongthese strata(subsets) tem. For praetiealreasons,this implies
(see White1975). solne sort of regulargrid layout(often
squareorreetangular) withequalspaeing
Sample Size between traps.This latteraspeetof trap
layoutis notneeessary justforestiInation
To obtainreliableestimatesof popu- of N: for example,if trappingwas on a
lation size, a sufficientlylarge saluple sInallisland,or in an enelosure,knowl-
mustbe taken.Typicalsamplesize con- edge of traploeationin a eoordinate sys-
siderationsarenot applicablehere(e.g., temwouldnotbe needed.Finally,forthe
deterluining the numbersof plotsto saln- densityestimateto be Ineaningful, the
ple). Rather,"saTnple size"relatesto the gridshouldbe plaeedin a homogeneous
nuluberof animalscaptured.Fora live- habitattype, to assureuniforlndensity
trappingstudy, one lnust have both a overthe grid.
largeenoughnumberof distinctanimals Fora ehoieeof r ande whentheobjec-
capturedand a sufficientnumberof re- tiveis densityestiluation, we suggestthat
captures(exceptforModelNlb andMbh) bothvaluesbe greaterthanorequalto 5;
The factorsthat controlexpectednum- asa minimumwe recommend r + C ¢ 20.
bersof capturesare(1)gridsize,in terms Examplesarea squaregrid10 x 10 or a
of area coveredand numbersof traps rectangle5 x 15.Wenotemuchworkin
used, (2) captureprobabilities,and (3) the literaturerelies on 16 x 16 gricls
nulaberof trappingoccasions.We dis- (Gentryet al. 1968,Smithet al. 1971),
cussthese3 factorsin relationto the size and we suggestthatgridsshoulclbe at
of the experimentnecessaryto achieve least that largefor attemptsto uxe the
precisepopulationestimates. methodbasedonnest-
densityestimatioll
Thesize of the gridis the firstdecision ed subgrids.We base this recolulllenda-
to be luade.Gridsize is a functionof trap tion on the factthata largenumberof
spacing,s, andthenumberof rows,r,and capturesis requiredin each subgrid;
colusuns,c, of traps.Equaltrapspacing hence,the largerthe size of the subgrids
anda systematicgridlayoutaresuggest- the betterthe chancethatlargenullllers
ed. Ateachgridstationtherewill be one of captureswill be achieved.
or luoretraps.If densitiesareveryhigh, The next decisionto be llladeis the
we recolumend2 or moretrapsper sta- spacingof traps(i.e.,thevalueof s). NIost
tion to avoiclcolnpetitionfortraps.Few workwithsmallluamnlals uses 15-lntrap
studieshavebeen madecomparing 2 or spatingor less (Barbehenn1974,himith
luoretrapsperstationwith1 trapstation, et al. 1975).The rationale,when one is
and we suggestfurtherresearchis re- given,relatesto the size of holneraIlge.
quiredto see wheremultipletrapsper Let s be the spacingbetweentraps,and
stationincreasesthe probabilityof cap- let 2Wbe the averagelinearholnerange
tureforindividualanimals. size.HomerangesInayrarelybe eircular,
The olejectiveof gridtrappingover a but assuluingfor design purposesthat
shorttimeperiodis to estimatethe pop- theyluaybe circular,thentheirra(liusis
ulatiollsize andusuallyalso the density W.We suggestat least4 trapsperholne
at the gridsite. Becauseof"edgeeffect" range.This ilupliess S (N/2)W. Forlest
(as discussedin the sectionon density resultswe suggests S W/2.Clearly,this
estimation),it is necessaryto estiluate iInpliessomeknowledgeof holnerange
effectivetrappingareaas well as N. In sizes hefore a gooclstuclyean be (le-
practice,this lneanswe mustbe able to signed.This is olotunreasonable to ask;
the biologistshouldhavesomebehavior- subgridswill otherwisebe too sparsefor

al knowledgeof the speciesbeingstud- reliableresults.Webelievethatreliable
ied so that samplingdecisionscan be densityestimatesusingthe subgridap-
Inadeintelligently.In fact,in anystatis- proachrequirea gridof atleasta 10 x 10
ticalsamplingproblemS a goodstudycan- andas a minimum75 to 100differentan-
not be planned without soIne prior imalscaught.
knowledgeof the populationparalneters We now consideran approachto de-
to be estimated. termininga minimumgrid areaon the
1 e anayses presentec
r .
v ln t llS mono- basisof the abovecriteria.

.
We startwith
graphfor estiInatingpopulationsize re- the relationshipN= D A(W),where
quire sufficientnulubersof capturesto A(W)is the effectivetrappingarea.For
produce satisfactoryresults. Defining a rectangular grid,thisis
"sufficientnumbers"is an extrelnely N = D[LrLe+ 2(Lr+ L)W + qrW21,
complicatedtask.Basedon our experi-
ence with bothrealand simulateddata, whereLris the lengthof a row [Lr=s
however,somecrudeguidelinesmaybe tr- F anc -cr c- s c- J 1S tSae engtR
, \ .
stated. For instance, experimentsin ot a co,umnot traps.. iausyt ae areacov-

o rn
which Mt+1is on the orderof 10 or 20 eredby the gridis LrL.

animalssiluplydo not provideenough As mentionedabove, without soIne
inforluationforthe procedures discussed knowledgeof D andW,a suitablestudy
hereto performwell. Thenllmberof dif- cannotbe designed.AssumeDf,andWo
ferentanilualscapturedneedsto be sev- arethe best guessesof the valuesof the
eraltilneslarger,andwill dependheavi- paralneters. To determineif a gridstudy
ly Ollthe probabilities of captureof the is at all feasible,set N - 50, substitute
populationInenabers beingstudied.That D, andWOin the aboveequation,set Lr
is, a populationin whichmelubershave = Lc- L (a squaregrid),andsolveforL
an 'average'captureprobability of 0.40
or 0.50mightonlyhaveto be as largeas L - +/0.8584(Wo)2
+ NlDo- 2Wo
50 beforetheestimation andtestingtech-
niquesbecolneuseful,whereasa popu- Thenthe actualareathe gridmustcover
lationsize of 200 or so mightrequirean is L2.
averagecaptureprobability of only0.20. This procedureis not difficult;how-
ForInoststudies,a relativelylargenum- ever one mustbe carefulto use the same
berof recaptures mustbe realizedbefore basicunitsforD L, andW.Forexample,
the experilnenthas a chanceto produce let W()be 100feet (30.5m)andassulnea
useful results and this againrelatesto densityof 1 animalperacre(1/0.4ha).For
the magnitude of theprobabilities of cap- compatibility
of unitsput Doin termsof
tureinvolved.In general,the probabili- squarefeet then Do- (1/43,560)feet2.
ties lnust be largerfor stnallerpopula- SolvingforL gives
tions butin no instanceshouldN be less
than25 or averagecaptureprobabilities L - /(0.8584)(100)2 + (50)(43,560)
- 200
less than0.10whentrappingsmallluam- - 10279feet.
malsforonlya fewoccasions(sayt S 10).
These recommendations do not guaran- Thistranslates backinto37.5acres(15.2
tee thatthe datacanbe satisfactorily ana- ha) as an absoluteminimulngrid size
lyzedSbutwe haveseen enoughrealand [37.5= (1,279)V43>56()].
simulateddatato saythatif the datafail This is clearlyconservativebecause
these criteriait is iluprobable thata pre- notall animalswill be caught.Iluproved
cise estiluatewill be achieved. planningrequiresus to deterlmine grid
Estiluationof densityby the neste size so thata given nu1nberof aniluals
subgridapproachrequireseven larger Mt+1will be caught.But the expected
saluple sizes; the data on the smaller numberof distinctanilualscaughtde-
FROMCAPTUREDATA{)tiS Xt al.
INFERENCE
STATISTICAL
79
pendsuponthe trueunderlyingcapture Thus, to get reliableresultsin such a
probabilities, whicharenot known.The studywe wouldsay the truepopulation
only practicalapproachis to luakethe densityshouldequal or exceed21 ani-
best guess at the overallaverageproba- mals/ha.If the biologisthas goodreason
bilityof firstcapture,p, applicableduring to believe truedensityis only 10 or 12
the studyandthenuse the formula animalslha, the studyis not even worth
Mt+l= N[1- (1 _ p)t]. doing.
In additionto controlling the salnpling
Becausethe valueof t (nuluberof occa- effortthroughthe size of the gridandthe
sions)has been introduced, the relevant nuluberof traps,the biologistcan also
colnputations for several realistic values selectthe nuluberof trappingoccasions.
of t cannowbe performed. In theory,the moretrappingtimesthere
For example, assulne p = 0.30, set are the better,but this ignoresthe fact
Mt+1 = 50,andsolveforN forseveralval- thattheclosureassuluption becolnesless
ues of t. Giventhese valuesof N, solve realisticas Inoretilne passes.We rec-
forthe valueof L, andhence luinimuln omInenda minimutnof 5 trappingocca-
gricbsize: sions,but 7 to 10 is better.The interval
*
grlc . slze
between occasionsshouldbe short.In
in acres is eitheronceatrapping
practice, most of sluallanimals
t N L (ha)
day (Inorning) ortwicea
4
rw
1,498 51.S (20.8)

day (morning and evening). Trapping
6 S7 1,378 43.6 (17.6)
only once a day is far less likely to intro-
8 53 1S323 40.0 (16.2)
duce tilne variation. With Inorning and
eveningtrappingthereis very likely to
In practice,this examplemeansif trap- be a differencein captureprobabilities
ping were only for 4 days, one would betweentimes.If variationof behavior
needa 16 x 16grid,trapsspaced100feet and heterogeneityshouldalso be pre-
(30.5 m) apart.For an 8-dayperiodof sent,the correcttnodelends up as Mtbh
trapping,the same(expected)datacould forwhichno suitableestiluatoris avail-
be obtainedwitha 14 x 14 gridof traps able.Morning andeveningtrappingluay,
spacedlO0feet (30.5m)apart. however,be aiinedat differentspecies.
This processcan be reversed.Let us Thena workabledesignwouldbe 5 (or
saya studyis plannedwitha squaregrid 7) daysof trappingin bothmorningand
of 16 traps,spaced15 m apart(W= 7.5 evening,butwithseparateanalysisofthe
m).ThenLr= Lc= 225m.Substitution in mornlngancleveningdata.
* 1 1
the basicequationgives In removalstudies,an absolutemini-

N= D 57,552 (In2) lnuln is t = 3 occasions(not 2, as is often
done)becauseit is impossibleto test for
or equal captureprobabilitieswhen t= 2.
N= D 5.76 (ha). We recoznmend thatat least4 reluovals
be done.
Assumefurtherthe studyis to last5 days Anothervaluablemethodof testingthe
and the averagecaptureprobabilityis adequacyof the designbeforegoingto
aboutp = 0.30.Thenwe have the fieldis to simulatethe experiment on
M6= N(0.83). a computer. Approximate parameter val-
ues can be chosen,and the experiment
We need to get at least 100animalsbe- replicatedas manytimes as necessary.
causethe smallestsubgridrequires50,so Amongthe criteriathatcanbe observed
the densityshouldbe largeenoughthat are selectionof the appropriate model,
N > 120,or the bias of selectedestimators,and the
D ¢ 120 _ 20.8animals/ha. achieved confidencelevel. Obviously,
5.76 thevalidityof the simulations to thefield
. v v
(1,1) (lt2)
In recapture studies, each animal
f (1,3)
shouldbe markeduniquely,otherwise
substantialinformation maybe lost and
it will be impossibleto computesomeof
the testsforsourcesof variation. Though
st
(2,1) (2,2) (2,3)
it should be obvious,
x we do mention
* that
it is crucialto correctlyrecordall data
(e.g., animalnumberand trapnumber).
To knowthe traplocation)trapsmust
be numbereduniquely.Moreover,the
biologistmustknowthe relationship be-
(3,1) (3,2) (3,3) tweenthetrapnumberandits coordinate
on some (arbitrary) rectangular X-Y co-
ordinatesystem.Fromthe standpointof
data analysis,the best approachis to
x X
identifythe trapsin the fieldby theseX-
x x
(4, 1) (4w2) (4,3) Y coordinates.We stronglyrecommend
FIG. 20. Exampleof trap numberingfor a 4 x 4 use of thissystemandwe stressthatden-
gridwith a standardcoordinatesystem. sityestimation usingprogram CAPTURE
requiresdatato be collectedin the con-
study will dependon the similarityof text of a coordinatesystem.We recom-
parameters selectedto the actualparam- mendthata cornertrapbe numbered(1,
etervalues.However,insightsaboutthe 1)thentherowsbecomethe "Xaxis>' and
experimentcan be gainedthroughsim- columns the '<Yaxis."For example, a4
ulationsthatcannotbe achievedin any x 4 grid wouldlook like Fig. 20. This
otherway. We note thatprogramCAP- systemcanbe extendedto coveranyreg-
TUREhasthecapability to conductthese ularrectangular gridof r rowsandc col-
slmuatlonstucbles. . *
umns.
Whentrapsarecheckedbothmorning
RecordingData and evening,it is necessaryto recordnot
onlythedayof capture,butalsothetime.
If densityis to be estimatedon the ba- Data recordingwill be facilitatedby
sis of grid trappingdata,the minimum using standardfield formsand standard
information thatmustbe takenwhen a conventionsfortrapnumbering andani-
captureoccursincludes:(1)animaliden- malidentification. Oneexampleofa stan-
tificationcode, (2) traplocation,and (3) dardizedmethodis presentedin Brotz-
trappingoccasion.Usually,the species, manandGiles(1966).
sex, and age are also recordedfor each
animal,thoughforanalysesgivenhereit Data Anomalies
has no purposeexceptto partitionthe
databy species,sex,andage.Whileana- Various anomalies and unplanned
lyzingthe dataseparatelyby species is eventsmayoccurin trapping.Forexam-
recommended, thereoftenarenotenough ple, (1) severalanimalsmaybe foundin
datato furtherpartition by sexandageof 1 trap,(2)animalsmaybe founddeadin
animal. traps,(3) releasedanimalsmaybe found
Fortrueremovalstudies(e.g.,electro- furtherdownthe gridtrappedagainon
fishing),thereis no animalidentificationthe sameoccasion,and(4) a trappedan-
code. As suggestedby Raleigh (pers. imal mayescapewhen one attemptsto
comm.),it is very important in removal removeit fromthetrap.WeInakethe fol-
studies to record the individual by lowingsuggestionsregarding these hap-
speciesandto analyzethedataby atleast penings:(1) morethan1 animalpertrap
presentsnoproblem.Recordeachanimal
. . .
a 1 f\t tfX V fAlA f+ 1 fW ffUl 1 YS Ct

lllC;zlUl tCXAtJll\Jlll1t_ ;1VH
FROMCAPTUREDATA4)tiS et al.
INFERENCE
STATISTICAL
81
separately. Thistypeof datadoesnotin- OVERALL

2-VALUE
TEST R£5UlTS --
- 1 . 08Lt
validatethe analysespresenthere;(2)an f3AB IL ITY A SMALLERVALX . 13925
w
animaldeadin the trapin a livetrapping
studyis a moreseriousproblem.If it is TEST OF CLOSUREBY FR1DQUE<YOF CAPT?E.
t ze asttrappingoccasionit cBoes notmat- lFR£CkUlEKI£5 LESS THAN 10 ARE N0T COWUTED. )
ter.Otherwisethe dataanalysismustbe
luodified.Solneofthemethodsdescribed ER OF CAPTURE5 SZ-YALU£ PROBABt L I TY
here can allowforknownremovalsin a
________________________________________
true livetrapping study (specifically 2 . l> .57306

ModelsMoandMt).Becausenotall tests 3 t}64 .6786t
and estitnatorscan be so modified,we 9 X.208 .88650
have not dealthere with such modified tionFIG. 21a. Exampleof test procedurefor popula-
Inodels.Werecommend the following:if closurewith the SchemeB taxicabdatafrom

Carotherstl973b).
trapdeathsareless than5 percentof total
captures,removethosedatafromthe to-
tal results,runthe analysesandaddthat
numberof deadanimalsto N, and then densityestimationmayalso be desired.
Inultiplythe densityestimateDvby (1 + Finally,in the courseof these analyses
proportion dead).If suchdeathsareluore varioussummarystatisticscan be com-
than20 percentof totalcaptures,use the puted.Below,we givenseveralcomplete
generalizedremovalmethodof analysis examplesofthe entireanalysisprocessof
on firstcaptures.For 5-20 percenttrap livetrapping dataforpurposesof estiluat-
deaths,the onlysafeanalysismaybe the ing populationsize. We do not include
removalInethod.These modifications densityestimationforall theseexaluples
give N and D relativeto the firstdayof becauseits essentialfeaturesarealways
the study.If one desirestheseestimators the same.Also,no furtherexaluplesare
to apply to the populationremaining givenhereoftheanalysisof reluovaldata
alive afterthe study,then simplydelete becausethatsubjectis substantially Silil-
all trapdeathsfromthe dataset priorto pler thanthe analysisof capture-recap-
analysis;(3) the sameanirnalis caught turedata.
norethanonce on a givenoccasion,the
only addedinformation providedis on A Taxtcs Example
* 7
movement.We recommendthat both

capturesbe recorded, butonlytheresults Carothers(1973b)conductedan inge-
of the firstcapturebe usedfordataanal- nious capture-recapture experilnenton
ysis; and (4) an animal escapes during the taxicab population of Edinburgh,
handlingbeforeit is tagged,orbeforethe Scotland.Such a studyhas the advan-
markis read.Do nothingbut recordthe tagesof knownpopulationsize yet the
tact. Jo not attelnptany sortof anaysis populationis a realone (thoughnot in-
o _
In essence,it does not volvinganiluals)as opposedto a coln-

of this C<record.''
b colnep artofthe aa. putersimulation experiment. Twodiffer-
ent saluplingmethodswere used; we
COMPREHENSIVE EXAMPLES havealreadyused Carothers' SchemeA
in the sectionon ModelMhto illustrate
Precedingsectionshavegiventhe de- the jackknifeestimator(see Fig. 6), the
tails of the specificInodels,estimators, entire 10 days of observations("trap-
andtests thatarethe basisforouranal- pings")frolnSchetneB are used as an
ysis of capturedata.Theanalysisof a set examplehere.In thatscheme,observers
of livetrappingdata by these methods hadfixedstationsin the city.Thiscorre-
will involvethe Inodelselectionproce- spondsto a trappingstudywith 10 days
dure,followedby estimationof N under of trappingat fixedtraplocations.The
the selected(ormostappropriate model); truepopulation size was420,andwe can
OCCASION JX I 2 3 4 5 6 7 8 9 10
ANIMALS
CAUGHTN1J ) m 48 52 47 44 48 45 48 43 47 53
TOTALCAUGHT M1J)X 0 48 90 122 146 173 188 203 213 Z25 241
NEWLYCAUGHT U(J)= 48 42 32 24 27 15 15 10 12 16
FRENENC
IES F (J ) - I04 67 51 12 6 1 0 0 0 0
I . TESTFORHIETEROGENE
ITYOFTRAPPINGPROBAB
ILITIES IN POPULAT
ION.
NULLHYPOTHES
IS OFMOOELM(O) VS. ALTERNATEHYPOTHESIS OFMODEL
M(H)
CHI-SQUARE
VALUE
s 7.913 DEGREES
OFFREEDOM
= 4 PROBABILITY
OFLARGER
VALUE
= .09482
2 . TESTFORBEHAV
IORALRESPONSE
AFTERINITIALCAPTURE
.
NULLHYPOTHESIS
OFMODELM(O) VS ALTERNATE
HYPOTHESIS
OFMODEL
M(B)
CHI-SQUARE
YALUIE
s .095 DEGREES
OFFREEDOM
s 1 PROBABILITY
OFLARGER
VALUE
= .75743
3. TESTFORTIMESP£CIFICVARIATIONIN TRAPPING
PROBABILITIES.
WLLHYPOTKSIS OFMODEL M( O) VS. ALTERNATE
HYPOTHES
IS OFMODEL
Ml T)
CHI_SaJAR£VALUIE
s 2 . 247 DEGREES
OFFREED()M
= 9 PROBAB
ILITYOFLARGER
VALUE
= . 98693
4 GOOONESS
OFFIT TESTQFMODELM(H)
NULLHYPOTHESIS
OFMOOELM{H)VS. ALT£RNATE
HYPOTHESIS
OFNOTMODEL
M(H)
CHI-SaJAREVALUE
s Z. 300 DEGREES
OFFREEDOM
s 9 PROBAB
ILITYOFLARGER
VALUE
= . 9857B
TESTOFMODEL
M(H) BYFREQUENCY
OFCAPTURE
(FREQUENCIES
LESSTHAN 2T ARENOTCALCULATED.
)
NUF8ER
OFCAPTURES
CHI_SaJARED. F. PROBAB
ILITY
1 6.769 9 .66113
2 6.246 9 .71504
3 4.714 9 .85847
5. GONNESSOFFIT TESTOFMODELM(B1
NULLHYPOTHESIS
OFMODELM(B) VS. ALTERNATE
HYPOTHESIS
OFNOTMODEL
M(B)
CHI-SQUARE
VALUE
= a.957 DEGREES
OFFREEDOM
= 16 PROBAB
ILITYOFLARGER
VALUE
= . 91516
5A. CONTR
IBUT
IONOFTESTOFHOMOGENE
ITYOFFIRSTCAPTURE
PROBAB
ILITYACROSS
TIME
CHI-SQUARE
VALUE
= 7 . 598 DEGREES
OFFREEDOM
= 8 PROBAB
I LITYOFLARGER
VALUE
= . 47369
58 CONTR
IBUT
IONOF TESTOFHOMOGENE
ITYOFRECAPTURE
PROBAB
ILITIES ACROSS
TIME
CHI-SQUARE
VALUE
= 1. 359 DEGREES
OFFREEDOM
= 8 PROBAB
ILITYOFLARGER
VALUE
= . 99480
6 . GOOOKSS
OFFIT TESTOFMODEL M( T)
WLLHYPOTHES
IS OFtlODELHIT) VS. ALTERNATE
HYPOTHES
IS OFNOTMODEL
M(T)
CHI-SQUARE
VALUE
= 188. 341 DEGREES
OFFREEDOM
= 168 PROBABILITY
OFLARGER
YALUE
= . 13483
7 . TESTFORBEHAV
IORALRESPONSE
IN PRESENCE
OFHETEROGENE
ITY.
NULLHYPOTHESIS
OFMODELM(H) VS. ALTERNATE
HYPOTHESIS
OFMODEL
M(BH)
CHI-SaJAREVALUE
= 15.148 DEGREE5
OFFREEDOM
= 26 PROBAB
ILITYOFLARGER
VALUE
= . 95458
MBELSELECT
IONCRITER
IA. MOOEL
SELECTED
HASMAX
IMUM
VALUE
.
MODEL M(O) M(H) M(B) MlBtl1 M(T) M(TH) M(TB) M(TBH)
CRITER
IA . 93 1. 00 . 37 . 52 0 . 00 . 46 . 36 59
FIG. 21b. Exampleof model selection procedurebased on Scheme B taxicabdata from Carothers
(1973b).Appropriate
modelprobablyis Mh.Suggestedestimatoris jackknife.
APPROXIMATE 95 ERCENT COSIX<E INTERVAL TO 406
FROMCAPTUREDATA C)tiSet al.

INFERENCE
STATISTICAL 83
NU"8ER OF TRAPPING OCCASI ONS UAS

NUt18ER OF ANIMALS CAPTUREO. M{T+I ), WAS
10
241
lowedby the computedselectioncriteria
TOTAL NU"BER OF CAPTURES, N., UAS 475
anda suggestedappropriatemodel.The
FREOUENCI ES OF CAPTURE,F { I )
1s 1 2 3 4 5 6 7 a 9 lo first3 testscompareModelMoto Models
Ft I t 104 67 51 12 6 1 0 0 0 0
Mh,Mb,and Mt,respectively.FromFig.
COPPUTED JACKKNI FE COEFFI C I ENTS
Nt I ( N(2) N(3} N(4) N(5)

21b results are sumluarizedbelow for
I
2
I . gO0
1 .000
2 700
.289
3.400
- .878
4 .000
-2 .278
4. 500
-3 .722 those3 tests:
3 1 .000 1 .000 1 .476 2.535 4.042
4
5
1 .000
1.000
1 .000
1 .000
1 .000
1 .000
.743
1 .000
.077
1 . 103 Chi-squclre Significczzlce
THE RESULTS OF THE JACKKNIFE CO"PUTATIONS Test of Moclel value level
I
0
N( I }
241
MoversusMh 7.913 SE t I )0.09482 . 95 CONF. L I MI TS TES T OF N ( I * I } YS . N( I )
CHI -SIZUARE( I D . F . )
1
2
334 .6
370 e
MoversusMb 0.095 0.75740
13. 34
21.53
308 .5
328.0
360 . 7
412.4
13. Ee5
2.344
3
4
389. 1
408.6 MoversusMt 2.247 0.98693
3 1 . 54
44.09
327 . 3
322.1
450.9
495.0
1 .680
2.137
5 433.3 58.67 318.3 548.3 0.000
AVERAGEP-HAT = The onlyindicatiollof variability

.1301 in cap-
INTERPOLATEDPOPULATION ESTIMATE IS
ture probabilitiesis heterogeneity(sig- 55 UITH STAAM E 20.30?4
nificantat the lOSo level).The studywas

designedto achieve constantnulubers
H 15TOGRAMOF F ( I ) captureclevery day; thereforeit is not
FREOuENCr surprisingthereis no indicationof tilne
104 67 51 12 6 1 0 0 0 0
EACH * EQUAL5
variabilityin captureprobabilities.
1 I POINTS
Also,
99
88 we do not findit surprising
.
* thatthereis
77
66
55
no indication,fromtest 2, of behavioral
*
*
*
'
'
response.One wouldnot expect"trap"
§
44 * *
33 *
22
I l
responsesfromtaxicabs.
* * * * .
Tests4, 5, and6 examinethe goodness

_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
FIG. 21C. Exampleof populationestimationwith

variableprobabilityof captureby animal under of fit of ModelsMh,Mb,and Mt,respec-
Model Mh with Scheme B taxicab data from tively.Whenanyof thosetestsreject,we
Carothers(1973b). aresayingthatmodeldoesnotappearap-
propriateforthe givenstudy.Test 4 in-
dicates no departurefrom MoclelN1h
be reasonablycertainof the closureas- (overallchi-squareof 2.3 with9 df).Nor
suluption. doestest 5 rejectModelMb(overallchi-
In thisexample,the conceptof density squareof 8.957with16df).Thegoodness
is not applicable,noris it meaningfulto of fit test to ModelMtdoes notcauseus
identify"trap"locations.Consequently,to rejectMt,butdoesluakeus suspicious
the firstpage of computeroutputis the of thatmodel(probability of a largerval-
test of closure(Fig.21a).FromFig. 21a, ue = 0.13483).Finally,test 7 givesus no
we have the overall test value z= causeto suspectModelMhshouldbe re-
-1.084. This is not significant,conse- jectedin favorof ModelMbh(chi-square
quently we would not reject closure. of 16.148with27 dfl.
Therewereenoughresightings2, 3, and By itself,noneof these7 testsis defin-
4 timesof the sametaxicabto allowspe- itive in suggestingthe appropriate luod-
cific tests of closureon just those cabs e ; ratherit is necessaryto considerthe
seenthatluanytimes.Noneof the3 tests resultsof all thetests.In thisexaluplewe
aresignificant. see someevidenceof heterogeneity, but
Thenextsectionof outputis theluodel no evidenceof timeor behavioralvaria-
selectionprocedure(Fig.21b).Afterthe tionsin captureprobabilities. Bearingin
headings,some sum1nary statisticsare luindthe robustnessof the jackknifees-
presented(dailycapturesnj,cumulative timatorof ModelMh we are willingto
markedMj,new animalsUj,andcapture concludetheappropriate modelforthese
frequenciesfj).The restof the modelse- datais Mh.The C<model selectioncrite-
lectionprocedureoutput(Fig.21b)is de- ria"computedby the programsuggests
voted to the 7 tests of assumptionsfol- Mhas appropriate (withModelMl,as the
W.
84 ILDLIFE ONOGPHS
VM
best secondchoicel.GiventheseresultsS OERALL TEST ZSWT5 -- -2. 132

the usershouldlooknextatthejackknife PROBAB
Z- VALtJE
IL I TY OF A SHALLERVALW .01650
estimatefor these dataand shouldcon-
siderall otherestimatorsas being inap-
.
proprlate. TESTOFCLRE BYFRE<Y OFCAPT.

(FR£NCIES LESSTHAN
10 ARENOtCOMPUTED.
)
Fig. 21c gives the resultsof estima-
tionof N fromCarothers' SchemeB data. Wt1B£R OF CAPTURES Z-VALl"JE PWBABIL I TY
Thispageof outputwill alwaysgive the ____________________________________ ___
summarystatisticsMt+l,n., andthe cap- 2 3 .691 . 99989

ture frequencies.Then the jackknife FIG. 22a. Exampleof test for populationclosure
coefficientsare given forthe numberof procedurewith cottontaildatafromEdwardsand
trappingoccasionsused.Next,theresults Eberhardt(1967).
of computingthe firstS jackknifeesti-
mates are shown. For example,Nh=
334.6 with a standarderrorof 13.34 edge there have been few other con-
while Nh5- 433.3,and SE(Nh5) - 58.67. trolledstudieslike thatdone, which is
In thatexample,the selectedestimateof unfortunatebecause it would be very
N is 365 withstandard errorof 20.3.The valuableto have moredatasets on real
approximate 95 percentconfidenceinter- populations whereN is known.
val on N is 325 to 406. Thatconfidence It thatstudy,135wildcottontails were
intervalfails to coverthe trueN - 420, capturedandplacedin a 40-acre(16.2ha)
reflectingthe previouslydiscussedpoor rabbit-proof enclosure.Afterallowing4
coverageof the confidenceintervalsas- daysforthe rabbitsto adjustto theirnew
sociatedwiththejackknifeestimator, but surroundings, livetrapping was conduct-
the relativeerrorof the estimatoris only ed for 18 consecutivenights.Whenpro-
13.1 percent(whichis consistentwith gramCAPTURE wasusedto analyzethe
oursimulationresultson Nh).In termsof resultantdata,the resultswere disap-
real capture-recapturestudies this pointing
amountof biasis, in ouropinion,accept- Fig. 22a showsthe resultsof the clo-
,
able. suretest appliedto Edwardsand Eber-

Finally we note thatjackknifeesti- hardt's(1967)data.Becausez=-2.132
matoris better(i.e.,nearerto N = 420)for (P= 0.0165)one would normallyques-
CarothersSchemeB datathanthe com- tionwhetherclosurewastrue.In thatex-
tnonlyusedestimators as Petersen(1896) ample,we attributethe resultto a time
or Schnabel(1938). variationin daily captureprobabilities,
As a furtheraidin judgingthe validity specificallytherewerefewercapturesto-
of t ze study,the estimatedaveragecap- wardthe end of the 18 daysthanat the
tureprobability is given.FromFig. 21e, start.As mentionedbefore,this 4'closure
AVERAGE P-HAT = 0.1301. Our studies test"candetectonlycertain typesoftime
have indicatedthata valueof less than variations of individualcaptureprobabil-
0.10suggeststhe captureresultsmaynot ities.It cannotof itself'<knowi7 thecause
Detrustedto producegoodresults.A 0.13 of the variations. Therefore,all the evi-
averageprobability is notveryhigh,but dencein the dataor otherwiseavailable
it is acceptablewhenthe truepopulation mustbe used to reachfinalconclusions
size is as highas 420. a Doutc.osure,or aboutotherquestions
such as the presenceof behavioralre-
A PennedRabbitStudy sponse(whichalsogetsconfounded with
closure).
EdwardsandEberhardt (1967)report- A briefdatasummaryandthe tests of
ed the resultsof a livetrapping studyon assumptions aregivenby the modelse-
a pennedpopulationof 135wild cotton- ection procedure(see Fig. 22b). From
tailsSylvilagusfloridanus.Toourknowl- tests1 25 and3, we see thatModelMois
NULL HTHES I S (lf MO4EL ( H 1 VS . ALTETE HT<S I S w L Ml > )
STATISTICAL
INFERENCE
FROMCAPTUREDATA{)tiS et 1. SS5
(3CCASI Js l 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18
AN{>LS cSGHr N(J)s 9 8 9 14 8 5 18 11 4 3 16 5 2 7 9 0 4 10
TOTK CAT M( J ) - Q 9 15 18 29 33 34 44 51 52 53 62 62 63 68 74 74 74 76
ffiLY CAT U( J ) w 9 6 3 11 4 1 10 7 1 1 9 0 1 5 6 0 0 2
FRE:QtJENC
IES F (J ) s 43 16 8 6 0 2 1 0 0 o 0 0 0 0 0 0 0 0
I . TEST FOR HETERO(3EFE

I TY Of TRAPPING A8 I L I T I ES IN POPWAT1N .
KLL HT<SIS w L M(O} YS. ALTEATE HYPOT<SIS w ML MSH)
CHI -fi4lJ YALUEw I 1 I 10 XBREES 8f FREEDtBtlw 3 PR(:IBAB

1L I Tr w LARBERYALK x . 01 1 15
2. TEST FW KEY 1wK KtiDONS£ AFTER I N1T1AL OWT .

NlLL HYMTHE515 OF P100EL MfO1 VSt KTETE HY T<SIS w ML M(B1
CHI -54UARE VALUEs . 062 X9EES 4B FREEDI:t1w I M%AB IL I TY w LARBERVALW s . 80367
3. TEST F0R TIME SPE:CIFICYARIATIN IN TRIK ILITIES.

NtLL HYPOTEESISOF t"ODEL M(O1 VS. ALTERNATE
HYPOTHESISOF M9OIELMlT1
CHI -SaJARE VALUEw 96. 9 XES y FZE w 17 PR%AB1L I rY y LARGERVALW s . 00012
s . ESS w F I T tEST OF MbEL " I H)

NILL APOTHESIS OF HODEL H(H) VS. ALTErE HTHESIS OF KT ML M(H)
CHI-SU VALUEw 55.502 DEGREESOF FZE " 17 4IL I TY OF LARXR VALW - .00001
TEST OF FlOOlELMlHl BY FREKY OF CWTWE
IFREaJEt<:IES LE55 tHAN 2T A*E NOT CALCUAtED.l
NUwBERX CAPTURES O11-5;au D. F . f>Rt8AEl

IL I TY
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
I w .023 17 . C0002
5. (iOOOKSS CF F I T TEST 1:F L M( 51

NULLHYPOTHESIS9F MODEL M(B) VS. ALTERNATE
HYPOTHESISOF NOT MODEL M(S)
CHI-At VALW X 102. 913 DE9EES OF FREEDOM

w 31 PROOAB
IL I TY w LARGERVALUEs 0 . 00000
5A . CONTR
l WT 1 w TEST w < I Tr OF F IRST CAPTWE PRZAB IL I TY ACROSSTI z
CHI -SQIJAK YALW ' 7 . 065 X9EES w FKEDt w 15 PR%ABIL I TY w LARGERVALUE- * 00004
5B . Ct:3NTR
IOUTI ON OF JEST OF HOt£NE I TY OF RECAPT PRWACI L I T IES ACROSST1 S
CHI-SQUAREVALW w 55.-9 X(3REES w FREEDOM

w 16 WOBABILITYw LARGERVALUEw .00000
OF F I T TEST OF M()OEL Mf r)
6 . (ilMOONESS
NULLHTHESIS w L M(T} VS. KTErE HYWTHESISX KT EL MIT1
EXPECTEDVAL<S TW SMLL. TEST KT KRFO
7 . TEST FOR BEHAV

IORAiLRESPONSEIN PRESENt:Ew F£TEROGENE
t TY.
CHI-EiaJ YALW w 9.023 X9EES w RREEWHw 17 PROBA8tLlTY w LARGERVALW w 00002
tSOOELSELECTION CRI TER1A. L WLECTEDHAS MX 1 vALV .
HODEL M(Q) H(H) HIB) M{>) MtT} MfTH) HxT8) H8T-

CRITERIA 15 O.00 .N .Ol 1.00 .64 .22 .M
FIG. 22b. Excllllpleof model selection pr()cedurebased on cottolltail datclS0111Edwclrclscllel Elerharelt

(1967). Appropriatemodel probably is XIt. Suggestecl estilllatr is Darr)ch.
notacceptablemoreover, the testsshow (tests3 and 1). Althoughtest 2 fails to

thatthereis clearlytimevariationin av- suggest that Model Mbis better than
eragedailycaptureprobabilitiesandthat ModelM,, we cannotSfromthatalone,
there probablyis some heterogeneity conclude there are no behavioralre-
1 1 1 1. _ 1 1 , 1
WILDLIFE MONOGRAPHS
86
(:)CCASI 1' 1 2 3 4 S 6 7 8 9 10 11 12 13 14 15 16 17 18
ANIS CAT N(J)- 9 8 9 1W 8 5 18 11 4 3 16 5 Z 7 9 0 4 10
TOTAL AN I MAL5 CAPTD 76
P-HAT {J ) - . 12 . II . 12 . 18 Î I . 07 . 24 ffi19 . 05 . 04 .21 . 07 . 03 . 09 . 12 0 . 00 . 05 . 13
TU
POPULAT EST I>TE IS 76 WI TH STAZARD E
.0165
OX IMTE 9fi PERCENTCONFICEKE INTERVAL 7S TO 77
HISTF w N(J)
FREQUENCY 9 8 9 14 8 5 18 11 4 3 16 5 2 7 9 0 4 10
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
EACH * EaJ^LS Z POI NTS
18 -
16 * *
14 * * *
lZ { . . *
10 * * * v * * * *
8 * * * * v . . . . * *
6 * * * * * * * * v * * . *
4 * * * * * * * * * * * * * * * *
Z * v * * * * * § * * * * * * . § §
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
FIG. 22C. ExaIliple of populationestimationwith time specifie changes in probabilityof capture

underModelMtwith cottontaildatafromEdwardsand Eberhardt(1967).
sponses In fact, test 7 (p = 0.00002) mate of populationis 76 which is the

shows thatthere are time variationsin sameas the numberof distinctanimals
captureprobabilitieswhichmaybe be- captured(i.e., M19=76). As furtherevi-
havioral. dencefordetermining whetherNtis a re-
Only 2 of the goodness of fit tests (4 liable estimate (andmodel),one should
and5) couldbe computed.Thegoodness compute the overall average capture
of fit to NIodelMtcouldnotbe donebe- probability:
causeofthesmallnumbersof individuals n.
caughtmostdays.Theother2 tests,how- tN
ever,resultedin rejection,indicatingthat
neitherModelMhnorMbis a satisfactory ForEawarcls ana EDernarats aata:
modelforthe data.
Whenthegoodnessof St testcannotbe
computedforModelMt,the modelselec- P (18)(76)
tion criteriaare computedbasedon the The advantageof usingthis formulain-
assumption thattest 6 wouldgive a sig- steadof
nificancelevel of 0.50.Thistendsto give t
undueweightto ModelMt,but thereis P - E Pj /(t)

reallyno goodsolutionto this problem. i=l
ForEdwardsandEberhardt's data,Mtis is thatwe knowthe truepopulationsize

the indicatedbest fittingmodel, with is greaterthanor equalto 76. Thus we
ModelMththe nextbest. This is consis- knowthis valueof p is an upperbound
tent with the observationthat capture onthetrueexpectedvalueofp, andhence
probabilities do appearaffectedby both averagecaptureprobability is lessthanor
timeandheterogeneity. equal to 0.10. This shouldraise a red
The estimationof N basedon Model f ag to t ze investigatoraveragecapture
-
Mtis given in Fig. 22c. The pointesti- probabilitymaywell be less than0.10,

B
STATISTICAL
INFERENCE
FRQMCAPTUREDATA{)tiS et 1. 87
ANIMAL t^1. MAXI MUM AYERAl:iE STANOAF?S) ANI MAL NUM. MAXI MUM AVERAGE STANARO ANI MAL NUM. MAXIMt AVERAGE STARAM
I.De CAP. D15T. DIST, ERRW l.D. CAP. D15T. DIST. ERROR l.D. CAP. DIST. 015T. ER
268 6 4.5 I .7 .69 269 3 1 ,4 .7 .71 6 z.o .8 .37

272
273 1.4 .3 .2a
27b s I ,0 .3 .25
c:
1.4 .7 29
276
277 6 2.2 I .5 .30 279 5 8.5 b.0 I .75 4 2.Z .8 .42
280
281 3 5.0 3 .E I .79 282 J I .4 I .0 O .00 5 7. 1 3.3 I .45
285
w 6 2.4 .7 29
2B7 5 2.0 5 .29 6 2.:2 .8 .20
288
299 5 I .4 .W
.35 29 3t0 I .s l .DQ 3 2.0 2.a 0 F 00
300
163 4 I .s .7 .33 165 4. I 3.5 .63 2 a .o 0. o O .00
166
167 2.0 .7 .67 169 2.2 I .2 .65 5 Z.O .5 .50
170
t71 5 4. I I .8 .61 172 2 I .4 I .4 O .00 3 5 S0 4. I .92
173
175 l O .0 O .o O .00
176 3 O .0 0.0 3 I .0 l .0 o.on
O 4 00 177
1- I 4.0 0.0 0.00
IB7 2.2 .7 .75 3 0.0 C .0 o .oo
188
189 2 3.4 3.0 191 I .0 .3 3 0.0 O .0 o .oo
0.00 .33 192
193 3 4.0 O .0 0.00
196 3 I .4 I .2 .21 l O .C O .0 o .oo
198
199 2 5.4 5 0 O .00 200 2 2.2 2.2 5.00 z O .0 O .0 O.OQ
86
89 l 0.0 O .o 90 I O .0 0.0 l o.a O .0 O.QO
0.00 O .00 91
2 I .4 o ,00
93 2 O .0 0.0 0.00 2 4. I 4. I 3.00
95
g8 I o.0 0.0 100 I Q.0 O .0 I 0.0 a .o 0.00
O .00 Q.OO 360
NOTE TAT AERAX DlST£ Y ZFERS TO 0157ANC£ KTZEN CESSIVE CAPTURES,
WHILE MXIMUt1 DISTAKE REFEF T4 TZ ATEST OISTAKE Kt<EN ANY TZ CAPt POINTS.
ALSO DISTANCE IS IN ITS 6 TRW INTERVALS, I.Ew, IF TZ INTER-TRAP OISTAKE IS 5 KTERS,
AN) TS: HAX 015TE IS 1.4, T<N TZ HAX DISTANCE 1N nETERs IS 1.4*5 w 7 STERS.
-RYBY FKKY w CT w nAXI OISTAKE KTZEN CAPt POINTS.
ER 5M1PLE F£ANOF STANDARI)

CAPT5 S l ZE HAXOIST. £
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
z 9 .91
I .e6
3 10
l .58 .6 14
4 8 Z .23 .360
5 8 3.45 I *O 18
6 7 392
2 wiM
TOTt 42 .575
_ _ _ _ _ _ _ _ _ _ _ _ Z .23
_(.____
FIG. 23a. Exampleof anilnalby animalsulnmaryof deer mouseeapturedatafromV. Reid(pers.comm.).
in whichcase this analysisis not trust- An Example of Trap Response

worthywhen (apparent) populationsize
is around 100. Manydatasets we haveseen on Pero-
In fact if we use the truevalueof N - myscus 1naniculatus have fit Model/Ib
135,then averagecaptureprobability in (trapresponse).The followingis an ex-
thisexperiment was0.056.Wealsopoint ample supplied by V. Reid (pers.
out that even after18 days of trapping comm.).The datawere takenin a 6-day
only 53 percentof the populationhad livetrapping study near Wet Swizer
been captured.This exampleillustrates Creek,RioBlancoCountyColorado, Au-
ourcontentionthatit is veryimportant to gust 1975.A rectangular grid of 9 x 11
have averagecaptureprobabilitieswell trapswasused with50-foot(15.2-m)trap
above O.OSor 0.10 for the population spacirlg. OneSherluan live trap(forsInall
sizes typicallyencounteredin capture- mammals) wasplacedat eachgridpoint
recapturestudies(50 to 150).Consider- andtrapping wasdonetwicedaily(morn-
ing thatN - 135andthe populationwas ing andnight) we haveusedonlyInorn-
penned we suggestthatthe '4true'' situ- ingcapturesforthisexample.Thereader
ationwas as follows.Therewas signifi- should study Figs. 23a-e which give
canttiInevariation andheterogeneity but the basicresults,beforeproceeding.
little real behavioralresponse.The av- A summaryof luovernentinformation
erage daily captureprobabilitiesde- fromrecaptures is givenin Fig. 23a.We
clinedin the last 9 (of the 18) daysand reluindthe readerthesedistancesarein
this causedrejectionof boththe closure unitsof 1 trapspacing[i.e.,50 feet (15.2
testandtest7. No goodestiluator of pop- m rere. .. zus t ile meanmaxlmum diS-
r n . 1
ulationsize is availablefor dataof this tanceof 2.23 impliesthe averagemaxi-

typeandquality. mulnmovementwas 111.5feet (34 m).
OCCASI ON J= 1 2 3 4 5 6
ANI MALS CAUGHT Nt J ) - 16 29 27 29 32 38
TOTAL CAUGHT M( J ) w 0 16 31 3B 44 48 51
NEWLYCACHT U(J) w 16 15 7 6 4 3
FREQUENCIES F(J)s 9 9 10 8 8 7
1 TEST FOR HETERNE I TY OF TRAPPI NG PROBABI L I T I ES I N POPULATI ON.

WLL HYPOTHESIS OF MODEL M(O) VS. ALTERNATEHYPOTHESIS 9F MODEL MtH)
CHI -SOIJARE VALUE w 37 . 489 DEGREES OF FREEDOMs 4 PROBABI L I TY OF LARGER VALUE = . 00000
2 TEST FOR BEHAVI ORAL RESE AFTER I N I T I AL CAPTWE .

Nl)LL HYPOTHESIS OF MOOEL M(O) VS ALTERNATEHYPOTHESIS OF MODEL M(B)
CHI -SaJARE VALUE X 24 . 342 DEGREES OF FREEDOMw I PROBA8I L I TY OF LARBER VALUE = . 00000
3. TEST FW TIXE SPECIFIC VARIATION IN TRA ING ABILITIES.

NULL HYPOTHESIS OF MOOEL M10) YS. ALTERMTE HYPOTHESIS OF MODEL M(T)
CHI -SWSE VALUE s 24 . 773 DE9EES OF FREEDOMw 5 PROBABI L I TY OF LARGER VALUE = . 00015
4. ZOONESS OF f IT TEST OF MOOEL M(H)

NULL HYPOTKS I S OF MOOEL M( H) VS . ALTERNATEHYPOTHESI S OF NOT M(3DEL M( H1
CHI-SOUAffE VALUE - 25.225 DEGREES OF FREEDOMw 5 PROBASILITY OF LARGER VALUE = .00013
5. GOOOKSS OF F I T TEST OF MODEL M( B )

NJLL HYPOTHESIS OF EL M(B) YS. ALTERNATEHYPOTKSIS w NOT M%EL M{8)
CHI-SaJARE VALUE s 5.978 DEGREES OF FREEDOM = 8 PROBABILITY OF LARGER VALUE = .6468
5A . CONTR I BUT I ON OF TEST OF HOMOGENE I TY OF F I RST CAPTURE PROBAS I L I TY ACROSS T I ME
CHI -SaJE VALUE = 1.135 DEGREES OF FREEDOM = It PROBAB I L I TY OF LARGER VALUE = . 88866
5B . CONTR I BUT I ON OF TEST OF HOM%ENE I TY OF RECAPTURE PROBAB I L I T I ES ACROSS T I ME
SH I -SQUARE VALUE s 4 . 843 DEGREES OF FREEDOM = q PROBAB I L I TY OF LARGER YALUE = . 30379
6 . GXONESS OF F IT TEST OF HOOEL M( T )

NJLL HYPOTHES IS OF MOOEL M( T ) YS . ALTERNATE HYPOTHES IS OF NOT MODEL M( T )
EXPECTED VALUES TOO SMALL. TEST NOT PERFO - ED.
7 . TEST FOR B£HAV I ORAL RESSE IN PRESENCE OF HE TEROGENE I TY .

WLL HYPOTHESIS OF HOOEL H(H) VS. ALTERNATE HYPOTHESIS OF MODEL M(BH)
CHI-SaJA*E VALUE s 22.217 DEGREES OF FREEDOM s 7 PROBABILITY OF LARGER YALUE = .00233
HOD£L SELECTION CRITERIA. EL SELECTED HAS MAXIMUM VALUE.
HOOEL HIO1 H(H) H{B) H(BH) M(T) M(TH) M(T8) M(TBH)

CRITERIA 26 .17 1.00 .65 0.00 .28 .55 .27
FIG. 23b. Exampleof modelselectionprocedurebasedon deer mousedatafromV. Reid(pers.comm.).

Appropriatemodelprobablyis Mb.Suggestedestimatoris Zippin.
One estimateof W (stripwidth)is pro- ing 106 feet, 32.3 m, here).Eitherap-

ducedby takinghalfthis averagemaxi- proachprobablywouldbe an iluprove-
mumdistance(56 feet, 17 m, here),or ment over using W- 0, but these are
halfof it plusone intertrap
distance(giv- essentiallyad hocapproaches.
,
FROMCAPTUREDATAtiS
INFERENCE
OCCASION J- 1 2 3 4 5 6 MATRIXw CAPTUR£5 P£R TRAP STATION.

TOTAL CAUCHT H(J)- 0 16 31 38 94 98 51
NEZY CAUGHT U(J)- 16 15 7 6 4 3
CtUwNS 1 2 3 q 5 6 7 8 9
_ _ _ _ _ _ _ + _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
ESTI>TED PRA3ABILITY OF CAPTURE, P-HAT X 3191 19

l
EST l MATED PROBAB I L I TY w RECAPTURE, C-HAT - . 67766 Rz3U | X a o 1 3 7 3 5 4 4
l
F 2 1 3 3 9 4 1 5 0 1 2
PLAT ION ESTIMATEIS 56 W I TH STAZAZ ER 4 .2859 l
f 3 i 2 2 2 0 0 4 2 q 2
APMOXt MATE9! i PERCENTC IDEKE INTERVALS 97 TO 65
l
F?04q1 3 2 3 e ° I l 1 2
Gri d
H I STOGRAIn1OF U ( J ) F?O#J 5 1 l O 0 I O O I q 5 1
2
61 O 2 2 e 2 0 q O 0
FR£0tJENCY 16 15 7 6 4 3 3
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
2 PO I NTS RS 7 1 6 I 2 2 3 3 I O O 4
EACH * EtZUALS
16 v ROIi 8 1 2 Z q 0 4 0 3 2 3
1 *
12 *
10 *
ROW 9 1 W O 5 0 2 0 D 0 O
8 * * . I
6 * * * * R)l 10 1 O 3 0 1 ° ° ° ° i 0
4 * * * * *
I
2 * . . . .
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ RC)W 11 1 0 4 e o 1 2 0 1 1
FIG. 23C. Exampleof populationestimationwith FIG. 23d. lNestedsubgridsused in the dellsity

constantprobabilityremovalestimatorunderMod- estimationprocedurewithdeermousedatafrolllV.
el Mhwith deer mouse data fromV. Reid (pers. Reid (pers.colllm.).In the Inatrix,trapcoordiIlates
comln.}. are roundedto the nearestwhole illteger.II) the
following goodlless of fit tests, trap coorclillates
In this example,the test for closure that are not integersand 1lonreetangular trappillg
gave z = - 1.586 (P= 0.05633).But be- gridswill eausespuriousrestllts
cause the closuretest is knownto be
biasedwhenthereis trapresponse,in the The suggestedmodelis Mb, and the
finalanalysiswe concludeModelMbad- correspondingestilnatorof population
equatelyfits the dataand closureis ac- size is, essentially,the Zippinreluoval
ceptable.A secondbitof evidenceonclo- estimator.(Wehave used the maximum
sureis givenby the "ring"test (see Fig. likelihoodestimatorof N underthe Zip-
23e).Thattestexaminesforanyapparent pin removalInodel.Recallthatthis dif-
attractionof animalsby the grid;it is not fers slightlyfrolnthe estiluatorusually
significantin this example,indicating computed.)Fig. 23c shows the esti-
thatthereis nobasistothinkimmigration mate of populationsize under Model
occurred. Mbto be N = 56, witha standard errorof
Fromthe modelselectionresults(Fig. 4.3. The approximate 95 percentconfi-
23b)we findthattests 1, 2, and3 all re- denceintervalon N is 51 to 65. Notethat
ject. This typicallyhappenswhen there the progralndoes not truncatethe coln-
is strongtrapresponse.Tests4, 5, and6 putedlowerliluit of 47 backto 51 (the
examine the goodness of fit of the 3 nuluberactuallyseen),but thiscouldbe
Models(]!w/th, ]!w/tb, and lut, respectively). donein reportingthe results.The prob-
We see thatModelMh does not fit, but abilityof initialcaptureis p = 0.32 and
ModelNlbdoes fit the dataadequately. the probabilityof recaptureis c= 0.68.
Unfortunately, the goodnessof fit of Mt Thesearesignificantly differentbecause
is the luostdifficulttest to coluputeand ModelMowasrejectedin favorof Model
withthesmallnumbersofanimalscaught Mb.Thissortof dramatic increasein cap-
it couldnotbe reliablycomputed. afterinitialcaptureis en-
tureprobability
Test7 examinesforpossiblebehavior- tirely consistentwith the propertiesof
al variationsin captureprobabilitiesal- the data,for example,the observedin-
lowingforanyheterogeneity whichmay creaseovertilne in the nj's(dailynuln-
be present(Fig.23b).Thattest strongly berscaptured).
suggests some forln of behavioralre- The areacoveredby the trappinggrid
sponseis present. was 4.59 acres(1.86ha);the naiveesti-
5
9o WILDLIFE MONOGRAPHS
-SQUARE:
I .
TEST OF UNIFfR1 DENSI TY BY ROWS
CH
9 o
7 8
I 3
l t 4
ROW
IB 4 5 12 18 20
(W;ERYED 27 23 :12 15.545 15.545
15.545 15 15.545 545
15 15.545
EXPECTED 15.545 15.545 15.545 I.329 8575
.388 15.545 .809 4 38a I276
9
I -AK
CH 8.4W0 3.575 .809
4 019
ROW l l
OBE;ERVED l l
EXf>ECTED 15.545
I -SQUARE
CH 1329
.
FK£. %ABILITY y LARGERYALUE= .0027

2fi,99 W1TH 10 XES w
CH1-
TOTAL $
IS .
I -SUARI: TEST OF UNI FORM[)ENSI TY BY COlU
CH
8 9
3 6
COL 17 17 19
2
25 15 20 18
OBSER\fED 2l l9 t9.000 1*9000
. oao 19.000 19.000 t9.000 19.000
EXPECTED i9.000 .211 .211 O.000
ooo
1. B95 . 8t2 * 053 .053
I -S(BUARE
CH .211 o * ooo
1L I TY w LARGERYALZ w ^9012
3.47 W I TH B XE5 w FZE . AB
CH I -SQUARE
TOTAL s
IF X%ITY BY RIKS {XTER RlK 15 R I) .

CHl- TEST6
l 2 3
RlF*
73 2 36 zo
35ERVED 25*go9
EXPECTED 62.182 8*5 3 .545
I . 88Z .837 w06 1 I .348
CHI -9UAK
%ABILlTY w LARGERYALW = *2479
4.13 WITH 3 XE5 w FK£.
TOTALCHI -A s
V. Reid(pers.comlll.).
FIG. 23e. Chi-squaretests of uniforlndensitywith deer Illousedatafrom
luateof densityis thus 12.2 mice/acre choicethe of subgridsas showntendsto Inin-

(30.71ha).Because of edge effect we imize problem.
knowthis is an overestimate.As dis- For each subgridan estimateof the
cussedabove,smovementdatafromre- population at risk of captureon that
capturesis one basis for estimatingthe As subgridwas obtainedusing ModelMb.
stripwidthWto get an estimateof effec- section, explainedin the densityestimation
tive trappingarea.A conceptually better we canthenestimateW.The re-
approach is to estimateWbasedon trap- sultsare shown in Fig. 23f.The 4 naive
pinggridsof differentsizes. Figs. 23e-f densities are shown withthe necessary
give results relevantto this approach computed constants (basedon gridsizes),
and the initialcovariance matrixof the
forReidisWetSwizerCreekdata.Four naivedensitites.
nestedsubgridswere chosen(as shown Thevalueof Wis 105witha (typically
in Fig.23d)asthebasisofthisprocedure standard errorof 58. NonethelessS
fromthe numberof capturespertrapsta- large) value of W is significantlygreater
tion thereis no apparentevidenceof a this
densityoverthe grid.Figure thanzerogso we acceptas our density
nonuniform (13.7/ha),
23e gives somelogicaltests of this uni- estimateD - 5.54animalslacre
iormdensityassumption.HoweverS our (SE(D)-1.92).
experience with the tests is that they Whenthis subgridapproachto esti-
oftenrejectwhenthereisSin fact, no real mation of D fails one mustuse
densitygradient(i.e.>systematicchanges D = NIA(W)
in density over the grid). Thus even
thoughthere maybe some variationin A(W) = Ag[l + aW + bW2]>
densityby '4rows>> it is minor.Also the
STATISTICALINFERENCEFROMCAPTUREDATAtlS et al. 91
STARTItIG YAL<5 FOR DENSITY ESTIMATION--

NUR OF GRI DS 4
TRAP I NTERYAL 50 . 00
UtJIT5 CObIVER51ON 43560?00
INITIAL DENSITY ESJIMATE 30.6995

INITIAL STRIP WIDTH E5tIMATE .5228
GRI D NAI VE XN5 I TY PERl K TER/ AREA P I / A*EA 5 TART I:NG COVARI ANCE MATRI X
I Y( 11 A( I ) Bt I )
t 32.67000 .3000000E-01 * 1571E-03 150.

2 61.71000 .1666667E-01 .5236£-04 507. .152£+05
3 17.qA00 .1 I?E-O1 ?261K-04 7.74 232. 14.1
W 12.10 .9000OOOE-02 .:i571E-09 1 * 16 34+7 2.11 .871
KiLT5 OF I TERATI
FlJKT I9J EVALUAT10N5 KWIKD 87
E5T1MTED SI;IFiCANT D:IGIJ5 6 PbRAKT£R VALW5 8
F I TTED MOD£L CN"PARED TO THE DATA

GRIDXI ) YXI ) F 11)
1 38*6tO 32.651
2 61.710 18.460
3 17.424t 13.941
4 12.197 11.745
MU T I PLE CELAT I 0b1 C6F I C IENT I 5 . 337 I B
EST I MATEDDEN5 I TYs S . 538 1 . 9241 w I TS STANDARDEf?ROR

ESTIMATED STRIP WIOTH- 105*2^ 57.w03 w ITS STAZAW ERRW
CORRELAT I ON w E5T I STORS - .}
TEST w EST I MATED5TR IP W1DTH GR£ATER STHANZERO.

Z-VA:LUE = 1 . 8145 PMBAB I L I TY OF LAR VALUE w . 034B
F INAL COYAR1ANCE MATRI X
150 0
91B.6 . :1521E+O5
18.80 311.4 14. lO
3+359 55.6Z Z.518 .87I4
FIG. datafr(:
23f: Exalnpleof jOillt estimationof density alld boundarystripwidth with deer III()EISU
V. Reid (pers.colnm.).
whereAgisthe gridarea,Wderives from 55.75 (half the averageluaximumdis-

eithermovelnentdataoran independent tanceof 111.5)we get
source anda andb areconstantswhich A(W)= 4.59 x 1.5506= 7.117
can be computed.In fact,the program
printsthese constantson the densityes- andhence
tiluationpage. FroznFig. 23f, we have 56
(for the total gricl)>a= 0.009 and b= 7117
0.00001571.
As an illustrationonly,if we use W= (Forpreviouslygivenreasons,however,
92 MONOGRAPHS
WILDLIFE
OCCASI 4N Jw 1 2 3 4 5
ANI MALS CAUGHT N( J ) s 37 5W 58 65 69
TOTAL CAUGHT MtJ)s 0 37 68 77 98 1 10
NEWLYCARHT U(J)s 37 31 9 21 12
FREQUENCIES F(J)s 34 20 28 15 13
I . TEST FOR HIETEROGENEI TY OF TRAPPI bK; PROBABI L I T I ES I N POPULATI ON.

NULL HYPOTHESI S OF MODEL M( O) YS . ALTERNATEHYPOTHESI S OF MODEL Mt H)
CHI-SQUAR£ VALUE = 49.016 DEGR£ES OF FREEDOMs 3 PROBABILITY OF LARGER VALUE = 0.00D00
2. TEST FOR BEHAVIWAL REE AFTER INITIAL CAPTURE.

NULL HYPOTHESIS OF KOiEL M(O) VS. ALTERNATEHYPOTHESIS OF MOOEL M(B)
CHI-SaJARE VALUE a 35.B65 DEZEES OF FREEDOM- I PROBABILITY OF LARGER VALUE = 0.00000
3 . TEST FOR T I ME SPEC I F I C VARI ATI ON I N TRAPPI NG PROBABI L I T I ES .

FaLL HYPOTHESIS OF MODEL M(O) VS. ALTERNATEHYPOTHESIS OF MODEL M(T)
CHI-SQUARE VALUE - 24.071 DEES 4F FR:EENM - 4 PROBASILITY OF LARGER VALUE = .00008
4 . GOOONESSOF F I T TEST 4F MaBEL M( H)

NULL HYP()THESI S OF MODEL M1H) YS . ALT£RNATE HYPOTHESI S OF NOT MXEL M( H)
CHI-SQUARE VALUE w 25.504 DEGRE£5 OF FR£EDOM s 4 PRO8ABILITY OF LARGER VALUE = .00004
TEST OF MODELMt H) 8Y FREWENCY OF CAPTWE

(FREaJENCIES LESS THAN 2T RE NOT CALCULAT£D.)
NUMBER4F CAPTURES CHI -SQUARE D . F . PROf3A8I L I TY

_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
1 18.647 4 .0009Z
Z 18.333 4 .00106
3 6. 048 4 . I 9562
4 6.000 4 .19915
5 . NESS OF F I T T£5T OF MODEL M ( B )

NVLL HYPOTHESI S OF MODEL Mz B ) VS . ALTERNATEHYPOTHESI S OF N9T SDEL M( B )
CHI -SGIUAREVALUE s 13 . 51 0 DEGREES OF FREEDOMs 6 PROBABI L I TY OF LARGER VALU£ = . 0 3562
I TY OF F I RST CAPTURE PROBABI L I TY ACROSS T I ME

CHI -SQUARE YALU"EX 9 . 220 DEGR£ES OF FREEDOM- 3 PROBABI L I TY OF LARGER VALUE = . 02650
I TY OF RECAPTUREPROBABI L I T I ES ACROSS T I ME
CHI -SOUARE VALUE w 4 .289 DEGREES OF FR£EDOM w 3 PROBASI L I TY OF LARGER VALUE = . 23187
6. ESS OF F I T TEST OF EL M ( T3
NlLL HYPOTHE5I S OF MODEL M( T ) VS . ALTERNATEHYPOTHESI S OF NOT MODEL M( T )
CHI -SaJARE VALUE - 84 .12 1 DEGREES OF FREEDOMw 66 PROBABI L I TY OF LARGER VALUE = . 0656Z
7 . TEST FOR BEHAVI ORAL RESE I N PRES,EKE OF HETER%EK I TY.

NULL HYPOTHESIS OF MOOEL MIH) YS. ALTERNATEHYPOTH"ESISOF MODEL M(BH)
CHI-SaJAR£ VALUE - 47 . 1 DE9EES w FRE£NM w 10 PROBABI L I TY OF LARGER VALUE = . 00000
MOOELSELECTI ON CRI TERI A . MXEL SELECTED HAS MAXI MUMVALUE.
MOOEL M(O) M1H) M(B) M(BH) M(T) M(TH) MSTB) M(TBH)

CRITERIA .55 .q2 . .6 0.00 .55 .8B 1.00
STATISTICAL
INFERENCE
we suspectthat W= 55.75is an under- OCCAS

ION
TOTALCAUGHT
Ja
M(J) s
I
0
Z
37
3
68
4
77
5
98 t 10
estimateandsuggestW = 105is a better NERLYCAUGHT U(J)- 37 31 9 21 12
valueto use.) ESTIMATEO

PR08ABI LI TY OF CAPTURE,P-HAT t . 255222
The samplingvarianceof D is comput- ESTIMATEDPR08ABILI TY OF RECAPTURE,C-HAT - . 617857
ableby the formulagivenin the density

estimationsection. The only missing POF^JLATI ON EST I MATE I S 142 W I TH STANDARD ERROR l 6 . 4Z 17
terlnis Var(W)which is obtainedfroln APPROX I FTE 95 PERCENT CONF I DENCE I NTERVALS I 09 TO 175
the standarderrorof Ineanmaximum dis-

tance(Fig.23a) HISTOGRA11
OF U( J )
FREaJENcY 37 31 9 ZI IZ
Var(W)
= ( 50 0 575 ) = 20664 _ _ _
EACH* EWALS
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
4 POINTS
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
ThenusingtheforlnulaforVar(D)we get 36
32
v
* v
Var(D)= 0.98,or SE(D)= 0.99. Z8

2q
ZO
*
*
*
-
+
16 ^ *
12 * *
An ExampleWhereNo LModel
Fits 8
4
*
*
*
*
In previousexamples,the luodelselec- FIG. 24b. Exampleof populationestirnationwith

tionprocedureusuallyindicateda model constantprobabilityremovalestimatorunderMod-
thatfit the data,andforwhichtherewas el Mbwithdeer datafrolllS. Hoffman(pers. IllOUSt
an estimator.However,with lmanyreal comm.).
datasets we have foundthatno luodel

fits the data.In those cases, the proce-
duresgiven in this monograph serveto lationremainsuntrapped. Phraseddiffer-
warnthat no reliableestimatorcan be ently,thereis no evidencehere to sug-
computedfromthe data(unlessit canbe gest Mt+1 is a reliableestimateof N.
judged the entire populationhas been FromFig.24a,it canbe seenthattests
caught). 1, 2, and3 rejectModelNIO overwhelln-
In this example(Figs.24a-e),we use ingly. We thus concludethere is some
livetrappingdatacollectedby S. Hoff- type of variabilityin captureprobabili-
man(pers.comm.)in mid-July1974on ties, and are suspiciousthat Inorethan
deermice.The studysite was in a sage- one sourceis operating.Test 4 also re-
brush-greasewood comlnunityat 4,500 jects the null hypothesisthatModel Mh
feet (1,372m) elevationin CurlewVal- fits.Atthispointwe canbe fairlycertaill

ley, Idaho.A 12 x 12 grid of Sherman thereis solneformof tilnevariabilityin
live traps,spaced15 m apart,was used. dailycaptureprobabilities, butit maybe
Trapping wason5 consecutivemornings, the oftenencountered resultof behavior-
110individualswerecaught(i.e.,Mt+l= al variability.
110)andtherewere283 capturesin all. The goodnessof fit test to Model,,
Fig. 24a shows some summarysta- indicatesthatthisInodelis nota gooclfit
tisticsandthe Inodelselectionprocedure to the data(P = 0.03562).Test6 alsosug-
results.Thedailycapturesincreased(37, gests ModelMtis not an adequatefit to
54,58,65,69)>andalthoughthenewcap- the data.Test7 againconfirlusthatsolne
tures tended to decrease,they variecl formof behavioral responseis presentso
substantially(37, 31, 9, 21, 12). Froln thatModelMhis inadequate. Thisleaves
thesesummary statistics,we wouldjudge us uncertainas to howtimeandbehavior
a substantialpartof the catchablepopu- variability areoperating. Tests5aand5b
FIG. 24a. Exalnpleof model selection procedurebased on deer mouse data from S. Hoffillan(pers.
coIllm.).Appropriate
modelprobablyis Mtbhor Mh. Suggestedestimatoris Zippin.
I
MONOGRAPHS
WILDLIFE
94
.
PER TRAPSTATION
IX OF CAPTURES
MATR
9 10 11 12
5 6 7 8
1 2 3 4
COLUMNS
_ _ _
_
_
_
_
__
_
_______________________________________
_ _
_ _
1
+
_ _
0 3 4 3
5 5 3 2
3 5
lI 4 3 0
ROW
I 4 3 0 1
0
2I 5 54 0 1
ROW 1 3 3 1
l 1 0 2 2
31 40 23
ROI*I Z 4 2 3
l 1 0 2 5
I 13 54
ROR 0 1 5 2
I 0 1 3 0
51 O4 1 3
RO;J 0 0 q 5
I 1 0 3 2
61 30 5 1
ROW 2
I O O l 3
2 0 2
71 2 0
ROW
I 1 0 0 3
0 0
81 44 1 2
4 4 2 4
RV l 4 0 0 2
9 l 1 4 2 2
RO>J I 1 0 2 4 4
2 0 0
l0 1 2 50 2
RO;J I 4 2 2
O O 0 3
1 3 O
11 l 2
ROWl 1 4 4
0 0 0
3 0 2
4 4
12 1
ROW
OLE INTEGER .
THE NEAREST
I
CO()RO NATES ARE ROUNOEDTO THAT ARE NOT INTEGER5
I X, TRAP COORDINAT£S
THE MATR
ABOVE
I N T TESTS, TRAP KSULTS .
GOOOFESSOf F I CAUK SPURI OUS
N THE ING
FOLLOiJ
TR ING GRI DS WI LL
NON-RECTALAR
AND ER 1) .
(tTER RlK IS
OF UNIF DENSITY SY RINGS
CHI-SQUARE TEST
6
4 5
2 3 4
32 11
76 53 7.861
RI^JG 107 39.306 23.583
70.750 028
55. 1.B96
OBSE:RVED 472
86, 1358
. 6.714
EXECTED .390 .075 VALULz . 0091
-AK
I
4.873 ILITY OF LARGER
. PROBAB
CH OF FREEDOM
15 31 WITH from S.
5 DEGREES HoSman (pers. comln.).
CHI-SQUAREs
TOTAL deer mouse daLta
tests of vIniformdensity with
24C. Chi-square
FIG.
model selection criteriaarea sig-
Test5a The data.In this
to clarifymatters somewhat. aid
nificant to interpreting models
help
goodness of fit test to Zippin's example, the criteriavaluesfor0.99, 0.89,
isthe probability removalmod- Mb, Mbh, Mtb,and Mtbhare Recallthat
constant that
(1956)
el,i.e., the null hypothesishere is Test 0.88,and 1.00,respectively. a
probability is constant. theluost likely modelis alwaysgiven values
firstcapture thatre- criteria
5bexalmines the null hypothesis This valueof 1, andthe other hence, we cannot
constant.
probabilitiesare suggestingthat arescaledaccordingly, data.It is
capture 0.23187 Mtbh actuallyfitsthe
testhas P = sayModel allaccount
latter
probabilities maywell be con- 5a significantthat these4 modelscorroborat-
recapture By contrastwithTest thus
stantovertime. that firstcap- forbehavioralresponse,that there is a strong
(P= 0.02650)we conclude These 7 ing ourcontention in captureprobabil-
probablyvary. strongbehavioral variation most
tureprobabilities
that behavior is the Because NlodelMbis the next
tests suggest
capture probabilities likelymodel,one mightselect bethe
ities. it as
est factor affecting
is the next most significant for estimation. Buttherecan no
andthattilne basis
factor.
STATISTICAL FROMCAPTUREDATA+)tiS et al.
INFERENCE 95
SU1s1ARY BY FREQUENCYOF CAPTURE OF MAXI MUMD I STANCE 8ETWEEN CAPTURE PO I NTS . OVERALLTEST RE:SULTS--
NUMBER SAMPLE MEAN OF STANDARD Z-VALW -Z. 142
CAPTURES S l ZE MAX D I ST . ERROR
IL I TY OF A SMALLERYALUE
PF?0f3AB , Ol 6 10
2 20 I .W6
.279
3 28 2.93
.410
q 15 2.26
. 388 TEST OF CL95 9F CAPTUR£.
BY FREX31JENCY
5 13 2 .#
TOTAL 76 2. 14
.338
.W01
tFREaINCIES LES5 THAN 10 ARE NOT CUT£D. )
FIG. 24d. Sumlnaryby frequencyof captureof NR w CAPTS Z-YALW PR%ABIL I TY

(listancebetweeneapturepoints.
InSililUIll
________________________________________
2
- I . I 18 .13178
strongrelianceon Nbor any otheresti- 3 -1 .708 .04383
. 1959
matorhere,becausenoneof the models -I .05W
fit the data.FromFig. 24b,the firstcap- FIG. 24e. with
Exalllple (f test fEar
deer mouse
poplllati>ll
(lata iolll S.
cl)sure
Hoffilla
ture probability(p) and the recapture procedtlre
(persi.t0lilln.).
probability underModelMbareestiluat-
ed to be
p = 0.26andc = 0.62. units.Convertingthis to sneters(2.14x
Theseareknownto be significantly dif- 15) and dividingby 2 we have 16 ln
ferentbecausetest2 rejectedModelMb. (52.7feet) as a conservative estimateof
FromModelMbtheestimateofN is Nb= W.Thisis typicalof suchvaluesseen for
142,with an estimatedstandarderrorof deerluiceonlivetrapping gridswith15-n
16.4.The approxiluate 95 percentconfi- trapspacing.It is alsoknownto be anun-
dence intervalis 109-175.Otheralter- derestimate. UsingW = 16m wouldgive
nativesare also basicallyunsatisfactory.a lessbiaseddensityestimatethanW= 0,
NlodelMbhcould be consideredbut it but it would still probablyresultin an
doesnotfitthe dataforanyof its specific overestimate of ID.The matteris further
cases.This maybe becausefirstcapture complicatedif animalswere in fact at-
probabilitiesvaryover tilne. ModelMh tractedto the grid. This phenomenon
does not fit, andthereare strongindica- cannotbe adequatelydealtwith,except
tionsthatbehavioris thedominant source by assessmentlines(ordesigninga study
of variation.Basedon these factors,we so that no attractionoccurs).Dropping
cannotrecomlnenda valid estiluation the outerring of trapsand reanalyzing
procedureforthesedata. the datais anotherpossibility,but then
If densityestimationis desiredhere, we are treatingthis outerringas an as-
furtherproblemsarise. FrolnFig. 24c, sessmentline.
thereis novisualevidenceofnonuniforln Norlually,we shouldhave lookedat
densityoverthe 12 x 12 grid.However, the closuretestearlyon in the modelas-
the ringtest suggestssignificantly more sessmentprocedure(see Fig. 24e); the
anilualswere caughtin the outertraps, resultsare z= -2.142, P = 0.0161.Be-
whichcouldbe evidenceof the grid"at- causethereis strongbehavioral variation,
tracting"animals. This phenozuenon we mustdiscountthis test;it cannotbe
does affectdensityestimationbasedon reliedupon.Thus,the onlyevidencewe
nested subgrids;in fact,this procedure have of a closurefailureis the ringtest
failedforthese dataas the estiluatedW of Fig. 24c.
valuewasnotsignificantly differentfrom We proposethe followingas a plausi-
zero.The onlyrecourseis to estiluateW ble explanation of thesedata.FromHoff-
frommovelnentdata,orto use a valueof man(pers.colum.)we knowtherewasno
W knownto be representative for deer prebaitingof thesetraps,no timewasal-
lnice. lowedforthe animalsto becomeusedto
Fig. 24d showsthe sulumaryof max- the traps.Fromthe analysiswe know
imumdistancesmovedforrecaptured an- therewasbehavioral response,andprob-
imals.The overallaverageis 2.14 trap ablytimevariationin firstcaptureprob-
. n n
abilities(butnot in recaptureprobabili- and54 subroutines. In addition,thereare

tzes
ae. - ac c ot preDa1t1ngotten 7 commonblocks.Includedin the source
Tr . . r
recommended in theliterature)mayhave code are amplecomlLnent statelnentsto

resulted in animalseither increasing followprogram flow.The totalcodecon-
theirfirstcaptureprobabilities overtime sists of about6,000cards.Corerequire-
as the studyprogressed,or in an influx lnentson anIBM360areabout200Kfor
occurringduring(ratherthanbefore)the the code withoutan overlaystructure,
studystarted.Prebaitingmay solve the but this can be reducedconsiderably
firstproblem.It will notsolvethe second withan overlaystructure.
problemof the grid attractinganimals The dilnensionsof the programare
andtherebycreatingartificially highden- presentlyset to allowup to 2,000indi-
.
slnes.
. .
vidualanimalsand31trapping occasions.
The productof the numberof captured
COMPREHENSIVE COMPUTER animalsandthe numberoftrapping occa-
ALGORITHM sionsmustbe less than4,000.This will
allow,forexample,30 trappingoccasions
The computations necessaryto calcu- and 80 aniinals,or 120 animalsand 20
late luanyof the estimatesdescribedin trappingoccasions.Thesevaluescan be
the precedingpages are quite lengthy, changed by changing DIMENSIO1!;
and in most cases, nearly ilupossible statementsin the program.
Wltout a computer.. aeretore,to pro-
vide methodsof populationestimation S UMMARY
usefulto the biologist,a comprehensive
FORTRANcomputerprogram,CAP- Theresultsof thisstudyprovidemeth-
TURE,has been written.The inputto ods forthe estimationof animalpopula-
the programhas been writtenin a *ee- tion size N and densityD frolmcapture
forlnandnaturalstyleto makeit easyto experiments. Bothcapture-recapture and
use. reInovalstudiesto estimatepopulation
The program is writtenin ANSIFOR- size aretreatedin detail.Theprimary fo-
TRANIV with severalsmallexceptions cus of the workhasbeen to relaxthe as-
so thatit will functionon mostbrandsof suluptionof equalprobability of capture.
digitalcomputers.Statementsknownto Threebasictypesof variationin proba-
causecompilationerrorson IBM,CDC, bilities of capturewere examined:(1)
Burroughs, Univac,Xerox,Honeywell,or captureprobabilitiesvary with tilne,
DECequip1nent havebeen flaggedwith ModelMt,(2) captureprobabilities vary
commentstatements,and the correct by behavioralresponse,ModellMb,and
statelnentforthe specificbrandof com- (3)captureprobabilities varyby individ-
puterincludedon a commentstatelnent. ual animal,ModelMh.Modelsallowing
A lmagnetictape with the FORTRAN these assumptions andvariouscolubina-
code andthe 13 sets of exampledatail- tions of assumptions(i.e., Modelslutb,
lustratedin thisluonograph areavailable Mth,Mbh,and Mtbh) are treated.Popula-
from tionclosureis assumedthroughout.
SHAREProgram LibraryAgency An integratedapproachwas followed
P. O. Box12076 and the result is a hlend of practical
ResearchTrianglePark,N.C.27709 lnethods,examplesof the analysisof real
data, statisticaltheory,and results of
computersilnulationstudies revealing
at a costof $40.00.Specifications forthe sozue small sample propertiesof the
tape(e.g.,7 or 9 track,800 or 1,600bpi, 1nethods.The estiluationand testing
etc.)shouldbe givenwhenorderingthe probleInhas been treatedin a standard
sourceprogram (No.360D-17.5.002). and usually rigorousstatisticalfraIne-
Theprogram consistsof a mainroutine work.Aboveall,the necessityof assump-
STATISTICAL
INFERENCE
FROMCAPTUREDATAtiS et 1.
97
tionsntheir importanceand statistical volving captureof animals.Biologists
tests of theirreasonableness have been haveall toooftengoneintothe fieldwith
emphasized.Assumptions lnustbe care- only a few trapshopingto obtainmean-
fullyevaluated,bothbiologicallyandsta- ingfuldata.Rarelywill this be possible7
tistically,before a particularestimator even if only an estiluateof population
canbe usedjustifiably. size is the goal.If densityestimatiorl is
A sequenceof 8 modelsthatincorpo- a goal, then filrtherdatarequirelllents
ratevariousassuluptions hasbeendevel- nustbe met.
oped.Pointandintervalestimators have As a referencepoint?it probablyis ap-
beenderivedformostofthelmodels. Fur- E)ropriate to thinkin terlusof 12 x 12)
therluorea coluprehensive andobjective 16 x 16,or2()x 20 squareorrectangular
(butnot optisnal)modelselectionstrate- gridsanda trappingperiodof 8-9 days.
gy is provided.This is crucialbecause Furtherguidelineson effortandsample
use of an incorrectluodelandiluproper size are given irl the text. Ourpriluary
assuluptions is aptto producebiasedes- contentionis that Inostcaptureexperi-
tiluatesand incorrectinferences.In ad- mentsconductedin the pastwere quite
dition,properinterpretatiors of the test inadequate in terlusof design,elifort, and
statisticsand their interrelationshipsis saluple size, which virtuallyprohibits
shownto be somesvhat difficult. justifiableil[ferencesto be drawnfro
Althoughthe practicalanalysisof data suchstudies.
frolncaptureexperimentshas been ex- Ourcomputersimulationexperilnents
tendedn additionalresearchneedsareap- have examinedthe sluallsampleprop-
parent.Statisticaltestingwithinand be- erties of the variouspoint arsdiIlterval
tween lmodelswill requireluorework. estilnators andthetestsof modelassump-
NIodelsalloving othersets of assumptions.The resultshavebeen inforlnative
tions need to be developed.Alternative andprovidesolnebasisforcautiousop-
estiluationschemes(e.g.,the generalized timisln.Capture-recapture and reInoval
jackknifeas opposedto thestandard max- metlods zavebeenoverrated in the past
imulnlikelihoodapproach)need atten- andthis probablyhascontributed to the
tion, particularlyfor solne of the luore lackof emphasison design,samplesize
coznplexluodels.Additionalworkwith andanalyticalmethods.Estilmation prob-
incompletecontingencY tablesmavprove lems relatedto D andN representdiffi-
fruitful(see Fienberg1972) Intervales- cult subjects.Ourresultsprovidehope
timatesin general,andparticularly inter- thata rigorousanalysismayvftenallow
val estilalates
forthe removalmodelsre- useful inferencesto be drarn if future
quireadditiorlal research.Bettertestsfor experilllentsare well (lesigneciandpro-
the iluportalltclosure assumptionare vide adequatecaptureand recaptureor
needed.Additiorwal controlledstudiesare removaldataforanalysis.
neededwherepopulationsize is known,
such as that reporteciby Edwardsand LITERATURECITED
Eberhardt(1967). Behavioralstudies
aimed at specific animal-trapinterac- ANDRZEJEWS}U7 R.SAND W. JEZIERSKI.1966. Stucl-
tions prollliseto provideinterestillgin- ies on the EtlropeaIlhare:XI. Estimationof
sights. populationdensity and attemptto plan the
yearlytakeof hares.ActaTheriol.11:433S48.
Researchresultsindicatethataccept- ANONYNIOUS. 1971. Mathematicalintrinsics:Infor-
able estiIllatescan le obtainedif ade- mationmanllal.Burroughs Corp.E1Nlonte)Ca.
quatedataare collectedproperly.How- 37 pp.
ever it is clearthatbiologistshave not ASSTASON,A. N. 197Sa. Predictionmethodsand
correctlyconceptualized captureexperi- varianceestilmates
forthe parameters of the tri-
ments over the past 4 clecades.These ple eatch-two populationmodel with migra-
tionanddeath.Univ.ManitohaSci.Rept.54 1-
lilmitationsmustbe correctedif progress 31.
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APPENDIX A
Notes on Estimation
Justificationfor the Use of MaximumLikelihoodEstimation
Our philosophy throughoutthis work has been to present a variety of

explicitmathematicalmodels for capture data based upon fully specified
assumptions.Given these modelsthatincorporatevarioustypes of variability
in capture probabilities(time, behavior, and heterogeneity),we are con-
cernedwith optimal estimationof populationsize, N, under each model.
Toachieve that optimality,we turnto the field of mathematicalstatistics.
A numberof very generalapproachesto optimalstatisticalinferencehave
been developed duringthe past 50 years. We feel that not all of them are
suitableforuse by biologistsforthe problemat hand(e.g., decisiontheoretic,
orBayesianapproacheswouldrequireinputsfromthe biologistthatwe doubt
theywould be willing, or able, to supply).We have chosen to use the method
of luaximumlikelihood (ML)estimationand inference;it is one of the best
developed,omnibustools of mathematicalstatistics.
The propertiesof ML estimationare well known (see Mood et al. 1974).
For many practicalmodels, ML estimatorsare optimal in many desirable
ways.Becauseof its generalapplicabilityandgoodproperties,MLestimation
has been the basis for most modernmethodsof estimatingparametersfrom
animalmarkingexperimentsof all kinds (Seber 1973).
The only requirementfor applicationof ML estimationis that one have
a well-definedparametricmodel with fewer parametersthan the dimension
of the vectorof minimalsufficientstatistics.Thus, we have been able to use
ML estimationfor Models Mo Mt, Mb,Mbh,and the removal models, but
not for Model Mh.Only for Model Mhdo we hesitate to claim there cannot
be significantimprovementsin the estimator.
It is not necessaryto understandthe mechanicsof how a ML estimateis
computed;it suffices to understandthis is not an ad hoc technique, but
rathera well-developedinferencemethod.
For any of the models dealt with here, there is a sampling probability
distributionfor the basic data (the {Xij}) that can be represented as a
mathematicalfunctionof appropriateparameters;symbolicallyit is
p{xij | N,p},
forN ¢ Mt+1,and with p representinga set of captureprobabilityparameters
(e.g., p = (p,c) under Model Mb).Given an actualsample, we can substitute
these data for the Xj variablesand treat this formulaas strictlya function
of the parametersN and p. This function (of N,p) is called the likelihood
function.The ML estimatorsN and p are those values of N and p which
maximizethe function
L(N,p) = P{Xij| N,p}
or, equivalently,which maximizethe log of L(N,p). In this way, the probleln
of deriving parameterestimators is reduced to the classic problem of
maxiluizinga given function over a set of possible values of specified
variables.
STATISTICALINFERENCEFROMCAPTUREDATA-4OtiS et 1. 103
The theoryof ML estimationgoes muchdeeperthanjust derivingpoint

estimatorsof parameters. It also gives methodsfor derivingapproximate
samplingvariancesforthe estimatorandprovidesjustification forthe usual
procedureforconfidenceintervalconstruction. Also,thereis a strongtie be-
tweenMLtheoryandthe theoryof minimalsufficientstatistics.
A sufficientstatisticis a condensation of the sampleintoa smallernumber
of statisticsthatstill containall the information thereis aboutN andp. For
example,Mt+1 andn. arethe minimalsufficientstatisticsforN andp under
ModelMo(representing quite a condensationof the entiresampleof Xij
values).The ML estimatoris alwaysa functionof the sufficientstatistic,
even if one has not explicitlydeterminedthe sufficientstatistic.Partlybe-
cause MLestimatorsare functionsof the sufficientstatistic,they have,for
largesamples,the smallestpossiblesamplingvariancesin the classof con-
sistentestimators. Stateddifferently,MLestimators are(asymptotically) the
most efficientestimatorspossible under a samplingtheoryapproachto
inference.Forall the abovereasons,we haveused likelihoodtheoryas the
mainbasisof estimationin thismonograph.
NumericalMethods
Exceptforthe specialcase of only 2 Brapping occasions(t= 2), the ML
estimatorsdescribedin the precedingchaptersdo not haveexplicitmathe-
maticalexpressions.Hence numericalmethodsmustbe used to maximize
thelikelihoodfunctionsandtherebyfindthepopulation estimatesona caseby
case basis.Conceptuallythis turnsout to be easy formodelsto whichwe
can apply ML estimation.Fromabove,the likelihoodfunctionis repre-
sentablesimplyas L(Np). In all applicablecases (excludeModelMhand
modelswhere estimationis not possible),we find there is a closed form
expressionfor the value of the captureprobabilitieswhichmaximizethe
functionL(N,p)foranyfixed N ¢ NIt+l Representthis as p(N).Thuswe can
write
maxmaxL(N,p)= maxL(p(N)|N)
N p N
= maxA(N),
N
whereA(N)is a function(analytically expressible)onlyof N. The maximiza-
tion of A(N)over N mustbe done numerically,but this is a simpleone-
dimensionalsearch.
Analgorithm by Brent(1973)(alsosee Fletcher1972)thatdoesnotrequire
analyticderivativesis used by programCAPTURE to maximizeA(N).The
algorithmsearchesbetween2 end points.Thoseend pointsarefirsttaken
as the numberof animalscaptured,Mt+l,forthe lowerboundanda linear
approximation to N plus Mt+1forthe upperbound.If the upperboundis
determinedto be the maximum forthe rangespecified,a new set of values
Iyingnextto the old valuesis selectedandthe searchoverN continuesin
thisfashionuntila truemaximum is found.Thealgorithm searchesalongthe
real line althoughonly integervaluesare appropriate. Becausethe search
is one dimensional,the algorithm is veryefficient.
The algorithmdoes not requirethe analyticfirstderivativesof A(N).This
meansit is notnecessaryto evaluatethederivativeofthelog-gamma function
withrespectto N(N!and(N - Mt+l)!appearin all the likelihoodfunctions).
IntegerN andConfidenceBounds
The domainof N includesonly integervalues. Therefore,the integer
thatproducesa maximum in the likelihoodfunctionis takenas the estimate.
The numericalsearchis conductedalongthe real numberline; then the
valuereturnedis truncatedto an integer,and a checkis madeto see that
it is the estimate,and not N + 1. A somewhatarbitrary roundingto the
nearestwhole integeris requiredfor the jackknifeestimatorto maintain
consistencywitR t ze otzerestimators.
Forall estimators, the varianceis computedon the basisof the valueof
N beforeit is roundedto an integer.We believe this procedureproducesa
slightlybetterestimatorof Var(N),and at the sametime avoidsnumerical
problemsthatwouldoccurwhenN equalsMt+1 (thissituationdoesoccur).
Confidenceintervalsof 95 percentarecomputedas N + 1.96gE(N).Con-
fidenceintervals(butnot standard errors)arecomputedaroundthe integer
valueof N. The lowerboundis thentruncatedto an integerandthe upper
boundroundedupwardto an integer.Those intervalend pointsare thus
technicallyoutsidethe true95 percentconfidenceinterval.Whenthe lower
boundis less thanthe numberof animalscaptured,the valuecouldbe reset
to Mt+1, a realisticprocedurebecausewe knowthatN is not less thanMt+1.
Thecoverageoftheintervalis notchangedbythisprocedure of usingMt+lfor
the lowerboundwhenthe calculatedlowerboundis less thanMt+1.
Admittedly, a confidenceintervalprocedurethatcanyield a lowerbound
less thanthe numberof distinctanimalscaptured(Mt+1) is not desirable.
We investigatedalternatives thatavoidthis problem,but we judgedthem
even less appropriate thanthe simpleapproachdescribedabove(see Ap-
pendixO).
Estimation of SamplingVariance
Maximum likelihoodtheoryincludesa generalmethodfor obtainingthe
largesample(i.e., asymptotic) formulaforthe truesamplingvarianceof N.
We have denotedthis simplyas Var(N),thus suppressingthe fact that it
is anapproximation to the "true'samplingvarianceof N. Theapproximation
is generallygoodforlargesamples;unfortunately, in the capture-recapture
contextwe -raveno goodmeasureof whena sampleis sufficientlylarge.A
furthercomplication entersbecausethe formulaforVar(N)is almostalways
a functionof the unknownparameters N andp; symbolically it is generally
of the form
Var(N)= Ng(p),
withthe formof the functiong known.ButbecauseN andp arenotknown
we mustestimateAlar(N) y
Var(N)=Ng(p)
The propertiesof Var(N)canbe different(sometimesquiteso) fromthose
of Var(N),which itself is only an approximation to the appropriate finite
samplingvarianceof N. One particular problemthathas been notedin the
literatureconcernsthe factthatN and gE(N)= war(N) - lNg(p) can be
substantially
correlated. Thisis notsurprising,
because,of course,N and/S
are highlycorrelated.The effect of this correlationis to cause an under-
estimateof SE(N)whenN is lowerthanthe trueN (whichit will be much
FROMCAPTUREDATAtiS
INFERENCE
of the time). This, in turn, causes the confidence intervalsin such cases to
be shorterthan they should be. An area suitable for furtherresearchon
capture-recapturestatistics(both closed and open models) is the question
of improvedestimationof samplingvariancesand confidenceintervals.
APPENDIXB
Estimation in Model Mo
In Model Mo parameterizedby the parametersN and p, the relevantpart
of the log-likelihoodfunctionis given by
1-
!) + (n.)ln(p)+ (tN - n VInX
( (N - Mt+1) )
where pe[OSl]and NeX= {Mt+l,Mt+1+ 1, Mt+1+ 2S...}. Given the value
of N, the ML estimatorp(N) of p is given as the solutionto
,98lnL(p| N,X) = O,
which reduces to
n. _ tN-n.
p(N) 1- p(N)
This results in the solution
p(N)= nX
Now, the ML estimatorNoof N satisfies
InL(p| N,X)]
lnL(Nop(No)| X) = mNa[ma[O
= NevY plnL(p(N) | N,X) 1
NevS Lln (N- M )s) + (n.)ln(n.)

+ (tN - n.)ln(tN- n.) - tNln(tN)]
For a given data set, a search over X is performedto locate the ML esti-
mate No This value is then used in the calculationof the 1NILestimate
p(No)- p via
n.
P = ^
tNo
The asymptoticvarianceof NOis (cf. Darroch1959)

Var(NO) = N[(1 _ p)-t - t(l - p)-1 + t- 1]-1.
An estimateof this varianceis
Var(NO)= No[(l- p)-t_ t(l _ p)-l + t l]-l.
APPENDIX C
Estimation in Model Mt
ModelMt is parameterized by the t + 1 parametersN, P1,P2,. , Pt. The
relevantlog-likelihood
functionforestimationof the parameters
is givenby
InL(N>P
IX) = ln( (N - Mt+1)!
) JE1
+ E (N - nj)ln(1 - pj),
3=1
where NeX= {Mt+tMt+1 + 1, Mt+l+ 2, }, P = {P1>
Ps * ,Pt}7pieLO,llfor
j = l, 2, . . ., t. Giventhe valueof N the MLestimators
pi(N)of PJaregiven
as the solutionsto the systemof equations
,98 lnL(p|NSX)
- 0 j- 1s27...,t.
Thejth one of theseequationsreducesto
nJ _ N - nj
pj(N) 1 - pj(N)'
whichresultsin the solution
pj(N)- n3
Now,the MLestimatorNtof N satisfies
L(N ^ (N) ^ (N) X) _ maxrmaxInL(pr
1 P2,. . ,Pt|N,X)
t>P1 t,**,Pt tl -e*NLpje[0ll
- Ne.hz[InL(p(N)?p2(N) ***,Pt(N)| N,X)]
NeX [ ((N - M )!)+ E niln(ni)
+ (N-n)ln(N-nJ)
j-l
- tNln(N)J.
A searchoverX is performedin orderto locatethe MLestimatorNt.It is

thenpossibletocalculatetheMLestimatesPj(St)- Piofthepjforj= 1r2,. . .t
via
^ ni
Pj = Wv
The asymptotic
varianceof st iS givenby Darroch(1958)as:
Var(St)-N t 1 + t-1 _ z (l _ p
Il(1-P;) j=l
-i-1
FROMCAPTUREDATAtiS
INFERENCE
Anestimateof thisvarianceis
- 1 t --1
Var(Nt) = Nt t 1 + t-1-E (1 _ pj)-
Il(l-pj) i=l
-i=l
APPENDIXD
Estitnation in .\lodel lVb

ModelMb iS ?arameterized by the parameters N,p,andc. The partof the
is givenby
necessaryforestimationof the parameters
log-likelihood
((N - Mt+l)!)+ Mt+lln(p) + (tN - N1- M )1 (1

+m.ln(c)+ (M.- m.)ln(1- c),
whereNeX= {Mt+ln Mt+1 + 1, Mt+l+ 2, . . .}, pe[O,l],ce[O,l].TheMLestilua-
torc of c is producedby the equation
adlnL(N,p,c|X) = O,
whichreducesto
m. _ M.- m.
A A
C 1-C
Solvingthis equationgives c = m./M..Thus,we see thatestimationof c is

of N andp. Now,the relevantpartof the log-
independentof the estiInation
likelihoodfunctionforpurposesof estimatingN andp is
+ (tN - M.- 5Flt+l)ln(l

) + Mt+tln(p)
rPI ) In((N- NIt+l)! _ p)
p(N)of p is providedby the equation
Giventhevalueof NSthe MLestimator
30 InL(p| N,X)= O,
whichreducesto
Mt+1= tN - M.- Mt+
p(N) 1- p(N)
The solutionto thisequationgives
P( ) tN- M.
Now,the MLestimatorNbof N satisfies
( b,P( b) IX) ,NTev&^[pe[0,l] ]
= NmaXX,[lnL(p(N)|N,X)]
(tN - M.)ln(tN- M)]
W,
108 ILDLIFE MONOGRAPHS
Nd[ ((N- Mt+l)t)+ Mt+1ln(Mt+l)

- Me- Mt+1)
+ (tN - M. - Mt+1)ln(tN
AsearchoverX is preformedto locatethe MLestimateNb.It is thenpossible

tocalculatethe MLestimatep(N) = p of p via
^_ ^ Mt+1
. )-
.' tNb-M.
varianceof Nbis givenby Zippin(1956)as:
The asymptotic
Var(Nb) = [1 _ ((1 _ ))t]2_ t2p2(1 _ p)t-l'
Anestimateof thisvarianceis givenby

Var(N) = Nb(l- p)t[1- (1 - p)S
p)
[ 1 - ( 1 - p)t]2 _ tSp2 ( 1 -
b
APPENDIXE
Estimation in Model Mh
The mathematical detailsof ModelMharecoveredin detailin Burnham
(unpublisheddissertation) and Burnhamand Overton(pers.comm.).The
followinggives a few of the basicresultsforthis modeland the jackknife
estimator.
UnderModelMhwe assume
PiJ= Pi i-1S...7N7
and we furtherassumethatthe Pl ..PN are a randomsamplefromsome
(unknown) probabilitydistributionF(p) pe[O,l].ForanyF(p)>the MSScon-
tains only the capture frequenciesfi...,ft, and their distributionis
multinomial:
] )
fi, . . .>ft
(N - Mt+lS H
where
7rj= j (] )pjXl-- p)t-idF(p) j - O,. . .>t.
If one assumesa parameteric formforF(p),thenstandard approaches are
(e.g.,
possible ML).Burnham (unpublished investigated
dissertation) the case
andfoundthatapproach
whereF(p)is theclassof betadistributions basically
unacceptable.
estimatorusedherewas developedby application
The <'jackknife'> of the
generalizedjackknife (Gray
statistic and Schucany 1972)to the naive esti^
matorMt+ly assumingthe bias of Mt+ as an estimatorof N is expressible
STATISTICALINFERENCEFROMCAPTUREDATAtiS et al. 109
TABLE E.1. THE JACKKNIFE Nhk OF POPULATIONSIZE, FOR K = 1 TO 5

ESTIMATORS
Nhl = Mt+l + ( t )fl
Nh2=Mt+l- ( t )fl- (t(t- )) f2
3t - 6 Vf _ (3t ( 15 1) )f2 + t(t - l)(t - 2)
( t ) ( t(t 1) ) ( t(t l)(t 2) ) t(t l)(t 2;(t 3)
h5 Mt+l+ ( t )fi- ( lOt ( 70t > 125)f2+ (lOt3-120t2 + 485t-660)f.
_ t (t- 4)5- (t- 5)5 j (t - 5)5 f

\ t(t - l)(t - 2)(t- 3) y 4 t(t - l)(t - 2)(t- 3)(t - 4) 5
in a power series in l/t. The end result of that applicationis that for the kth
orderjackknifethe estimatoris a linear functionof the capturefrequencies.
For example,we have
Nh1= Mt+1+ ( t )f1-(1 + t )f1 + E fi
These point estimatorsof N have been determinedfor up to the fifthorder,

and are given in Table E.1.
Because
Mt+1= E fj,
i=l
any Nhkis expressible as a linear combinationof the capturefrequencies,

say as
Nhk E ajkfj
i=l
Using the fact that the fj are multinomialrandomvariablesan approximate

varianceestimatorof Nhkis
Var(Nhk) = E ( aJk) fj Nhk,
i=l
and confidenceintervalscan be constructedon the basis of the asymptotic

normalityof Nhk
A procedurefor selecting one of the estimatorshas been suggested by
Burnhamand Overton(pers. comm.).It involves testing whether Nhk+1is
significantlydifferentfromNhkand stoppingwhen no significantdifference
is found.
APPENDIX F
Discussion of Model Mtb

The mostgeneralmodelone mightassumein the instancewhereboth
behavioralresponseto captureandvariability amongtrappingoccasionsare
foundinvolvesthe followingset of parameters:
N - populationsize
pj= probabilityof captureon trappingoccasionj of an animalnot pre-
viouslycaptured, j = 12 . . . t
C*kj= proa ility of captureon trappingoccasionj of ananimalfirstcaptured
onoccasionk,j=k+ l,k+2,...,t;k- l,...,t- 1.
The corresponding probability distribution
of {Xfi,}is givenby
P[{Xfi,}] - [ [I pjU,(l - pj)N-Mj+l]
[llXfi,!] (N - Mt+l)
! i=1
t-l t
. Il [| c kjRk( 1 - C*kj)Uk-E; S
k=l j=k+l
where
u;= numberof animalsfirstcaughton occasionj, j = 1,2,. . .,t,
Mj= numberof marked(previouslycaptured)animalsin the populationat
the timeof thejth sample,j = 2,3,. . .,t,
Mt+1= totalnumberof different animalscapturedin the experiment
(noticeMt+l- E uj),
j=l
Rkj-numberof animalscaughton occasionj thatwere firstcapturedon

occasionk,j = k + 1, k + 2, . . ., t; k = 1, . . ., t - 1.
A MSSforthisdistribution is {Ul, U2, * * *, Ut, R12, R13, * * *, Rlt, R23, * , R2t, * *,
Rt+lst}, that has dimensiont(t+ 1)/2. There are t(t+ 1)/2+ 1 parameters
involvedandstraightforward MLestimationof 19is notpossible.
If we restrictthe modelso thatc*kj= Cj,i.e., the probability of recapture
on occasionj does not dependon the occasionof firstcapture,then NIodel
Mtb(as previouslydefined)results.It is easilyverifiedthat,whenC*kj=c;,
P[{XX,,}]givenabovereducesto the probability distribution givenin the sec-
tionconcerningModelMtbwherea discussionof the nonidentifiability of N
is presented.Thus,forpurposesof estimatingN, the assumption thatC*kj =
c;hasno utility.
ML estimationof N is theoreticallypossibleif one is willingto further
restrictthemodelso thatthereis somerelationship betweenC*jandpj.There
is nouniquerestriction, andasanexamplewe considerC*j= Opj, j = 2, 3, . . ., t,
whereOspjs 1, j = 1, 2, ..., t and0S; Apjs 1, j-2, 3, ...> t. Withthis as-
sumption,the probability distribution of {X<,}becomes
llX<>,
!(N-M t+l)! [Ill Pi ( 1-p jV
n-Mj+t
]
0
* [I1 ( opj) m;(1 - opj) Mj-mj]

STATISTICAL
INFERENCE
FROMCAPTUREDATA<)tiS et al. lll
= Ilk!(N M )! plUl(l-pl)N-U(m)
co
*I1 ( 1 _ pj) N Mj+ ( 1 - op j) M;-mj p n

i=2
A sufficientstatisticforthis distributionis {u1,u2,. . .,ut,m2,m3,

. . .,mt},thathas
dimension 2t - 1. Since there are t+ 2 parametersinvolved, the model
allows all parameters,in particularN, to be identified if t ¢ 3. Estimation
under this model is not consideredhere, however, due to the questionable
assumptionthatrecaptureprobabilitiesbear a constantrelationshipto initial
captureprobabilitiesover all trappingoccasions, for all animals.Thus, at
present we are not able to present an estimatorof N that is appropriate
underthe assumptionsof Model Mtb.
APPENDIX G
Discussion of Model Mth

In the sectionconcernedwith ModelMh,the randomvariableXijwas stated
to be distributedaccordingto a Bernoulli distributionwith parameterPij,
that representsthe probabilitythat the ithanimal is capturedon the jth OC-
casion. Here, we assume that the elements of the set {Xij} constitute a
mutually independent collection of randomvariables, and that Pii= PiPi,
i= 1,2,...,N, j- 1,2,...,t. Furthermore,we let the Pi, i= 1,2,...,N be a
randomsample of size N fromsome probabilitydistributionfunctionF(p;o)
definedon [0, 1],andwe restrictthe pj,j = 1 2, . . ., t so that0 S PiPiS 1 forall
i,j. Under those conditions, the probabilitydistributionof the observed
sample {Xij},where i = 1, 2, . . ., Mt+1,
j = 1 2, . . ., t can be writtenas
P[{Xij}] = P[{Xij} | Mt+l]P[Mt+l],

where
P[{Xij}6Mt+l]= (l| pjni) ( [I; l o PYi [t| ( 1-ppj)l-X,j] dF(p;(J)),

Yi= the numberof times the ithanimalis captured,
P[Mt+1] = the probabilitydistributionof the number of different animals
captured in the experiment.This distributionwill involve the
parametersN, P1,P2,. . .,Pt,and the distributionF(p;o).
Obviously,if the formof F(p;o)is left unspecified,ML estimationof N will
not be possible. If the formof F(p;(1)is specified,but the vectorof parameters
o is not, likelihoodestimationwill be possible if the dimensionofthe minimal
sufficient statistic is sufficiently large to ensure identifiabilityof N. As
mentioned in the section on Model Mth,estimationof N will be possible
if F(p;o) is completely specified. In that case, a MSS of dimension t + 1
is {nl,n2,. . .,nt,Mt+l},and the numberof parametersto be estimatedis also
t + 1, i.e., Pl,P2,. . .,Pt,N.Therefore,unless the experimenteris willing to
make the doubtfulassumptionthat at least the formof F(p;o)is known,we
can present no satisfactoryestimation procedure appropriateunder the
assumptionsof Model Mth
_ , .
112 WILDLIFE sMONOGRAPEiS
APPENDIX H
Estimation in Model Mbh

Pollock(unpublisheddissertation) consideredModelMbhand foundthe
distributionof the setofpossiblecapturehistories{Xfi,} undertheassumption
that(pi,ci),whichrepresentsthe pairof initialand recaptureprobabilities,
respectively,of the ithanimal,is the ithmemberof a randomsampleof size
N from some bivariatedistributionfunctionG(p,c,@).Let us consider
rewritingthe distributionP[{X} |G(PEC;o)] . . .,ut}|G(P>C;0)]
as P[{ul,u2,
|{ul,u2,. . .,ut}, G(p,c;8)].The distributionof the removalsP[{u,,
P[{Xfi,}
u2,...,utElG(p,c;o)] will be multinomial, with parameterN and cell prob-
abilitiesthatare functions of the moments of G(p,c;o).The conditionaldis-
tribution P[{XXO}I{ul, u2, . . ., ut},G(p,c; @)]will alsodependorlthe moments
of G(p,c;o)but not on the parameterN. Therefore,the multinomialdis-
tribution
P[{Ul,U2, * * *,Ut} |6(p,c;8)] = t N! (Hri ')(Tt+l) ,
[n uj!] (N - Mt+l)! j I
whererj is a functionof the momentsof G(p,c;o)and

Xt+l - 1 - E rj
i=l
is the relevantdistributionfor purposesof estimatingN. Pollock(unpub-

lisheddissertation) showedthatif G(p,c;0)- Gl(p;01)G2(c;82), then7rj- E[p
(1 _ p)j-l]andthe conditional distribution P[{X,,,} | {ul,u2, . . C(p,c;o)]de-
.,ut},
pends only upon G(c;82). Regardless ofwhether p and c are independent, once
we agreeto baseestimationof N solelyon the firstcapture"removal" type
of data,then the problemcan be reformulated. The behavioral response is
then irrelevant and all we need to consider is the probability distribution
of firstcaptures.Thatis, we assumethatP1, . *,PNarea randomsamplefrom
somedistribution G1(p;o).
Considertransforming theparameters 71,X2 ,Vt intothesetofparameters
P1,P2, , Pt by using the relationship 7rj - (1 - pl)(l - p2)...(1 - Pi-l)Pi,
j = 1,2, . . ., t. Thus,pjis a conditional probability thatrepresents theaverage
firstcaptureprobabilityof those membersof the populationthathave not
yet been capturedat the timeof the jth trappingoccasion.Giventhis inter-
pretation,it is not unreasonable to assumethatP1> P2> > Pt.Further-
more,we assumethat(P1 - P2) > (P2 - P3)> > (Pt-1- Pt),so thatlarger
differencesin averagefirstcaptureprobability occurinitially.(Note:if G1(p;o)
is theclassofbetadistributions both assumptions areeasilyshowntobe true.)
The assumptionsand the distributionP[{ul,u2, . . .,Ut} |G(PEC;Ø)] which we
abbreviate P[u1,u2, . . form
.,ut], the basis of the generalized removal method
outlinedbelow.
Fork- 12 t-2
. . .,Pk-l differ.This
(i)AssumePk = Pkil = * * * - Pt- p and that P1,P2,
reducesthe numberof parameters involvedin the estimationof N
to k + 1.
(ii)EstimateN by the ML method.This task is greatlysimplifiedby
. .
rewrl :lng
STATISTICAL
INFERENCE
FROMCAPTUREDATA{)tiS et al. 113
P[ul,u2,. . .,Ut]as II P[uj| ul,u2,. . .,uj-l],where P[uj| ul,u2,. . .,uj_l]

j=l
is the conditionaldistributionof the jth removal,given the values of

the previollsremovals.
Then
(iii) Choosethe smallestvalue of k thatproducesa sufficientfit to the data,
and take as the estimate of N the estimateassociatedwith this value
of k. The fit of the data u1, u2, ...., ut can he measuredby the usual
size cechi-squaregoodness of fit test. We have used (x= 0.20 because
of the seriousness of Type II errors.If significancelevels of all the
tests (one for each value of k) are less than 0.20 we have chosen the
value of k correspondingto the largest achieved significancelevel.
For a given value of k7the asymptoticsamplingvarianceof Nbhis
t
Var(Nbh)= N / '=1 kel p 2 (t - k + 1)2p2 _l

l-Erii=,vj
.
Pi el-P)t(l-Pj)+Eri-ll
J-I - i=l j=l -
An estimateVar(Nbh) of Var(Nbh)is obtainedby replacingN,p,p,,p2,. ,Pk-lS

71,1J2, * * *,tt by their respective ML estimates.
Finally, we mention that for a given value of k, it is possible that the
experiment<'fails,"i.e., valid estimationof N is not possible. Recall that
Seber and Whale (1970) provided a failure criterion for the estimator
associated with Model Mb. (This model correspondsto the case k- 1.)
Following their method of proof,it is easily shown that the failurecriterion
for any value of k is
t
(t+ k- 2j)uj<0.
i=k
If the experimentfails for a value of k, the correspondingmodel is clearly

excluded from those eligible for selection as the appropriatemodel for
estimatingN.
APPENDIX I
Disc1lssion of Model Mtbh

In Model Mtbh,it is assumedthat the ithanimalin the populationhas, on
the jth trappingoccasion,both a unique probabilityof first capturePijand
a unique probability of recapture cij. The model therefore requires tN
parametersconcerning first capture, (t-1)N parametersconcerning re-
capture (because C11=C21=...=CN1=0),and the parameterN for its com-
plete specification.This totals (2t-l)N + 1 parametersand obviously all
parametersare not identifiablefor estimationpurposes.
The assumptioncan be made that the 2t- 1 dimensional vectors (Pll,
P 12, * * *,P 1t,C 12,C 13, * * *,C 1t), (P21,P22, * * *,P2t,C22,C23, * * *,C2t), * * *, (PN1,PN2, * * *,PNt,C N2,
...,cNt) are a randomsample of size N from some probabilitydistriblltion
function F(pl,p2,...,pt,c2,c3,...,ct;o)paralneterizedby the vector o and de-
finedon [0,1]2t-l.The probability

distribution
functionof {Xh,}can thenbe
writtenas
P[{X@]} = n[E[v@]]
[tt Xfi,!](N
- Mt+l)! @
ct)
wheregreO-
a scalarrandomvariablethatis a functionof the 2t- 1 dimen-
sional randomvariable(p1,p2,...7ptc2,c3,...,ct)
corresponding
to the capturehistory, and
E[X(O]
= t t 7r,,dF(p1,. . .,pt,c2,
. . .,ct;o)
o o
For instance,if t= 4 and ct)= {1,1,0,1}, then rfi,- plc2(1- C3)C4. If one is
willingto makecertainassumptions concerningthe familyof distributions
to which F( ;0) belongs,the dependencestructureamongthe variables
{Pl,P2,. . .,pt,c2,c3,
. . .,ct}andthe vectorof parameters@,thenMLestimation
of N will theoreticallybe possible.However,it is not unreasonableto
suspectthatthe amountof numerical computationandthenumberofassump-
tionsrequiredwill prohibitsuchestimatesfrombeingusefulin practice.
APPENDIX J
Estimation in Removal Models

For the removalexperiment,it is assumedtheremaybe heterogeneity
amongthe captureprobabilities of the N membersof the populationsubject
to removal.Moreover, the N capture(removal) probabilities
Pi,i= 1,2,. . .,N,
are a randomsamplefromsome probabilitydistributionfunctionG(p;@)
definedon [0,1]and parameterized by the vector@.Underthoseassump-
tions, the distributionof the vectorof removals{u1,u2,...,ut}is given by
P[ul U2, . . *> Ut] = N!t (nXi)Xt+l t+l
[tI uj!] (N - Mt+l)

! i=1
j-l
where
rl
7rj= E[p(1- p)j-l]- J p(1 - p)i-1dG(p;@), j = 1,2,. . .,t,
o
rl
Xt+l = E[(1_ p)t] = J (1 -
o
p)t dG(p;@)
=l-E[p(l-p)i-l].
3=1
Noticethatthisdistribution of {u1,u2,
. . .ut} is of the form,andthe param-
etersof the samenatureas the distribution of {ul,u21.. .,ut}in ModelMbh.
Therefore, the generalizedremovalmethoddevelopedforModelMbhis also
applicableto removalexperimentswhere estimationof N is desired.Al-
thoughthe experimental situationsassociatedwiththose2 modelsarequite
different,the factthatthe removalsare the only statisticsrelevantforpur-
posesof estimatingN, andthe natureof the parameters 7rjcombineto make
the generalizedremovalmethodappropriate in bothcases.
FROMCAPTURE DATA4)tis et al. 115
APPENDIXK
Tests of Model Assumptions

The detailsof the statisticaltests describedin the sectionentitledTESTS
OF NIODELASSUMPTIC)NS aregivenhere.Testshavebeen numerically iden-
tifiedand correspond to the identification
nulubersused in programCAP-
TURE.
Test 1
Since ModelMois a specialcase of ModelMh,a likelihoodratiotest of
Ho: Pi= p, i= 1,...,N versusHA: all Pi are not equal, seems plausible.
However,due to the nonidentifiability of parametersin NlodelMh a valid
likelihoodratioprocedureis notpossible.
An alternativeapproachis takenby examiningthe fit of the observed
frequenciesf1,. .,ft (recallthese are the elementsof the MSSfor Mh)to
theirexpectedvaluesunderModelMO. Theresultingteststatisticis intended
to be sensitiveto departures fromModelMOin the directionof ModelNlh.
If Ho is true, we would expectthe test statisticTl to be approximately
distributedas a chi-squarerandomvariablewith t - 2 degreesof freedozn,
where
T _I E
t (fj_ fj)2
A
i=l j
fj= No(J) pi(1 _ p)t i,
andNoandp arethe MLestimatesof N andp underModelM

Test2
Testingthe null hypothesisof ModelMoversusthe alternativeof Model
Mbis equivalentto testingthe nullhypothesisHo p = c versusthe alterna-
tive HA:P # C. If it is assumedthat the bivariaterandomvariable{p c}
is distributedas a bivariatenormalwithmeanvector{p c} andcovariance
matrix
rVar(p) O 1,
L0 Var(c)j
then, underHo the quantityT2'= (p-c)2/[Var(p)+ Var(c)]is distributed
as a chi-squarerandomvariablewith 1 degree of freedom(cf. Theorem
4.4.5in Graybill1976).Here,p andc arethe MLestimatesof p andc under
ModelMb(cf.AppendixD), andwe use Var(p)= p2q2(1 - qt)/N[q(1_ qt)2 _
p2t2qt],whereq= 1 - p (cf. Seber1973:312).Furthermore, we approximate
Var(c)by treatingc as a binomialvariablewith M. fixedso thatVar(c)=
c(l - c)/M..Obviously,bothVar(p)andVar(c)will haveto be estimatedby
substituting the MLestimatesof N, p, and c underModelMb,so thatthe
actualtest statisticbecomes
T2- (p- c)2
Var(p)+ Var(c)
It followsthatT2has an approximate chi-squaredistributionwith 1 degree
of freedomunderHo.
If Ho is false, T2has an approximate noncentralchi-squaredistribution

with1 degreeof freedom and parameter
noncentrality +
A= (p - c)2/2[Var(p)
(cf.
Var(c)] Theorem 4.4.5 in 1976).
Graybill Thus,theoreticalapproximations
tothe powerof the abovetest canbe foundforfixedvaluesof N, p, andc.
Theseapproximations, obtainedby the use of the noncentralchi-square
tablesof Haynamet al. (1970),are given for the alternativesinvolvedin
Table8, AppendixN. Finally,it shouldbe mentionedthat,due to the con-
ditionalnatureof Var(c),it wasnecessaryto substitute
t k-2
E qe
E[M] = NpE
e=o k=l
A.
parameter
forM.to enableevaluationof the noncentrality
Test3
Testingthenullhypothesisof ModelMoversusthealternative of ModelMt
is equivalentto testingHO:Pj= P, j- 1,2,...,t, againstHA: Not all the p
are equal. We assumethat the t-variaterandom variable(P1,P2, ,Pt)is
distributedas a normal
t-variate distributionwith mean vector(P>P2, ,Pt)
andcovariance matrix
Var(pl) O
, where
O Var(pt)
Pi= nJNt- MLestimatorof PiunderModelMt,and

= pj/N,
Var(pj) q = 1-pj, j-1,2,...,t.
Now,usingTheorem4.4.5in Graybill(1976),the randomvariable
- ( pi/Var(pi))
T3'= E pj2/Var(pi) / Var(pi)-l
hasa chi-squaredistributionwitht - 1 degreesof freedomunderHo As in
test2, the will
Var(pj)
quantities haveto be estimatedusingtheMLestimates
pj and Nt. Uponmakingthese substitutions the test statisticT3'reducesto
t / t \2 /
.
T3= z nj/qj-tE nj/pjqj)/ E nj/pj2q i=l
j=l i=l
UnderHoST3hasan approximate chi-squaredistributionwitht- 1 degrees

of freedom.
Anapproximationto thepowerofthistestgivenanyalternativeis provided
by the knowledgethat,underHA,T3 has an approximate noncentralchi-
witht - 1 degreesof freedomandnoncentrality
squaredistribution param-
eter
A= (N/2)[ pj/%-( 1/%)/ (pjqi)1].
j=l i=l j=l
were
Utilizingthe tables of Haynamet al. (1970),these approximations
calculatedfor the involved
alternatives in Table9 of N,
Appendix and the
resultsarecontainedtherein.
J 1 3 JAL NFERENCE
A _rrp n r s
FROM
CAPTURE
IJATAtiS et al. 117
Test4
Anoverallgoodnessof fittestof ModelMhcanbe thoughtof as equivalent
to testingHo:PU- Pi versusHA:not all pu= Pi,i-1,2, . . .7Mtl,j - t.
17 * * *
Thus7the null hypothesisassumesheterogeneous individualcaptureprob-

abilitiesthatdo not changeovertime,andthe alternativestatesthat,given
heterogeneitycaptureprobabilitiesalso changeovertime. NoticethatHA
does not statehow probabilitieschangeovertime,i.e., whetherthe change
is due to behavioralresponse,variationin trappingoccasionsor a com-
binationof both.Thussif Ho is rejected,it is not rejectedin favorof an
alternativeModel;ratherit is the goodnessof fit of ModelMhthatis being
rejecte .
Burnham(unpublisheddissertation) has shownthat underHo the test
statistic
t
2 (n-nlt)2 t 1
T4_ i=1
jft)(
JE t)
has an approximate chi-squaredistributionwith t- 1 degreesof freedom
andis appropriate fortestingH<,vs. HA.The test is conditionalon the fre-
quencyof capturestatisticsfi,f2,...,ft.Burnhamalso recommended thatif
tk is argeenough,a statisticappropriate fortestingHo:Pii= Pi versusHA:
-
notall PiJ- Pi forall i suchthatYi= k, is givenby

t k 2
= X ZkJ t fk t - 1
T4a= kf t-k'
t k
where z,; - numberof animalscaughton dayj thatwere capturedexactly
k times and
Yi= numberof timesthe ithanimalwascaptured.
UnderHO,T4ahasanapproximate chi-squaredistributionwitht - 1 degrees
of freedom,conditionalon the valueof fk.Noticethata test statisticof the
formof T4acanbe constructed foranyk = 1,21. . .7t 1 as longas fkis large
enough.We haveused the criterionthatfkmustbe largerthant beforethe
test is performed.
Test5
An overal1goodnessof fit test of ModelMbcan be constructed by com-
biningthe resultsof 2 independenttests.The firstof these testswas intro-
duced by Zippin(1956)fiortestingHQ:Pj = P versusHA. Not all PJ- P,
j- 1,2,...>t. In the contextof ModelM pj representsthe probabilityof
firstcaptureon the jth trappingoccasion,and thus ZippinXs test attempts
to determinethe constancyof firstcaptureprobabilityovertime.The test
statistic5whichhasanapproximate chi-square
distribution
witht - 2 degrees
otr treeclom
r 1
w zen .01S true 1S glvenas
T _ E (uj- Nbpqit)2+ (lqb - Mt+ - Nl,qt)2
j=l Nhpq N qt s
whereNband p are the ML estimatesof N and p underModelMb,and

q= 1 - pe(Note:we haveusedthe following4pooling>7 strategyforthischi-
squaretest. If Nbpqr-1 < 2 and NbpqS-l ¢ 2 for s = 1,2,. . .>r- 1, then the
corresponding
'cells>> to r,r+ 1,. . .,t arepooledinto 1 cell. Thiswill reduce
thedegreesof freedomassociatedwithT5ato r-2.)
A test (independent of the aboveprocedure) forthe constancyof recapture
probability overtime can be constructedusingthe so-calledvariancetest
forhomogeneityof binomialproportions (cf. Snedecorand Cochran1967:
240).Thatis a test statistic appropriatefor testingHo Cj- c versusHA:
notall Cj- c, j = .
2S3, . .,t, is givenby
T5b = E Mi(cj - c)2/c(l - c),
j_S
where
CJ - mj/Mj,
c - MLestimateof c underMb- m./M..
approximate distribution
chi-square witht - 2 degrees
The
rP
om . hasan
statisticTia
wnen 1Strue.
OStreec nQ
BecauseT5aand T5bare independentand have chi-squaredistributions

when ModelMbis true,an overalltest statisticfor the goodnessof fit of
ModelMbis givenby T5- T5a+ TsbThis test statistichas an approximate
chi-squaredistributionwith 2t- 4 degreesof freedomunderModelMb.
Test6
Testingthe goodnessof fit of Model Mt can be interpretedas being
equivalentto testingHO: PU- Pj versusHA: not all Pu= pj,i - 1>2>...,Mtel
and j= 1+2,...,t. Thus, the null hypothesisassumesvariationin capture
probabilities amongtrappingoccasionswhile asseflingthaton a given QC-
casionall animalshave the same probabilityof capture.The alternative
HAallowscaptureprobabilities to differamongmembersof the population
on a givenoccasion,as well as allowingtrappingoccasionsto affectcapture
probabilities.NoticethatHAdoesnotspecifywhycaptureprobabilities differ
amonganimals on a given occasion. That is, this difference may be due to
behavioral response individual heterogeneity or a combination of both.
Thus whenHois rejected,it is not in favorof a specificalternative model;
ratherit is the goodnessof fit of Mt that is being rejected.We have chosen
to adoptthe test procedure proposed by Leslie (1958) for the purpose of
testingHo versus HA The proper test statistic is given by
T _ E3 [t-l fk(fl(k - 1 _ y )2] I
J_1 k=ly._ R2j{/UJS
E
{=j+l
where
- numberof animalscapturedexactlyk timesthatwerefirstcaptured
fkfi)
on thejth occasion,
u, = numberof animalsfirstcapturedon thejth oCCaSiOn
Rjt- numberof animalsrecapturedon the {th occasionthat were first
caughton thejth occasion,
^
k=l
(k - l)/uj, and
= t-E+,lfk(j)
Ii _Jlifuj¢20
l 0 otherwise.
Notice that we have followed Leslie's recommendationsthat a "cohort>'of
newly identified animalsfirst capturedon the jth occasion not be included
in the overalltest unless it consistsof morethan 20 individuals,i.e., u; must
be greaterthan 20. Furthermore,a new cohortis not included unless it is
subject to at least 3 subsequent trappingoccasions, i.e., j= 1,2,...,t- 3.
Under the null hypothesisof"equicatchability,"T6has an approximatechi-
squaredistributionwith
t-3
E (Uj- l)Ij
i=l
degrees of freedom.
It shouldbe mentionedthatCarothers(1971)proposedan improvedversion
of Leslie's test. However, practicaluse of the procedure requires some
arbitrarytrimmingof the dataand thereforeis difficultto use for simulation
purposes.
Test 7
Pollock(unpublisheddissertation)developed a proceduredesigned to test
Ho Model Mhfits the dataversus HA:Model Mbhshould be used. The test
statisticdepends on the 2 vectorsof statistics
where fki) is the number of ani-
f* = {fl(1,fl<2,...,flit,...,ft-lel,ft-l2>,ftel)}
mals capturedexactly k times that were first capturedon the jth OC-
casion,
f= {f1,f2,...,ft}, where fj is the number of animals capturedexactly j
times.
This is a chi-squaregoodness of fit test formedby pooling t - 1 indepen-
dent chi-squaretests. The kthof these tests is conditionalon the value of fk
and has t - k degrees of freedom.The overalltest statisticis given by
t-1 t-k+l {fk [(t - k -g + l)/(k)] }

k=l i=l [(t k - J + 1)/(k) fk
Under Ho T7has an approximatechi-squaredistributionwith

t-l
E (t - k) = t(t - 1)/2
k=l
degrees of freedom.(Note: for each of the t - 1 distributionswe have used

the same pooling strategydescribedfor the test involvingT5a.In this case,
the quantitychecked for sufficientlylargeexpectationis
[(t k -J + l)/(k)] fk
Thereforeit maybe thatin a given case degreesof freedomwill not be

t(t - 1)/2,butratherthis quantityless the appropriate
numberof degreesof
freedomlostby pooling.)
A TestforClosure
An approachto a closuretest suggestedby Burnhamand Overton(pers.
comm.)can be conceptualized by firstconsideringthe null hypothesisHO:
Pij= Pi,j = 1, 2, . . ., t foronly thoseanimalscaptured2 or moretimes.Es-
sentially,HOmerelyassertsthatindividualcaptureprobabilites areinvariant
overtime.Now consider,however,an alternativehypothesisHAthatstates
thatfor at least some i, i.e., for some animalsthatwere capturedat least
twice,Pil- Pi2- = Pir= Oand/orPis- = Pit= 0, where1 S r < s S t.
HAstatesthatsomemembersof the population werenotpresentin the pop-
ulationforsomeinitialor terminalpartof the study,or both.If thatalterna-
tive is true,one mightexpectthatthe timebetweenfirstandlastcapturefor
animalsis, on the average,shorterthanone couldexpectunderHO.That
conjectureis the rationalefor the test proceduregiven below. Note that
the test is designed to detect birth-deathor immigration-emigration
phenomena,or both,thatoccuronly duringthe initialand latterstagesof
the study,not phenomenaoccurringtowardthe middleof the study.For
example,the test wouldnotbe appropriate whensomeanimalsarepresent
duringthe initialstagesof the study,leave the studyareafor a time, and
thenreturnpriorto the termination of the study.
Giventhatthe ithanimalwas capturedexactlyYitimes,andthatYi¢ 2,
let Qi= Wi- Vi, where Wi is the occasionof last captureand Vi is the
occasionof firstcapture.Thus,Qi is merelythe timebetweenfirstandlast
captureforthe ithanimal.Conditional uponthe valueof Yi,the expectation
andvarianceof Qiare:
E(Qi|yi= k) - (k - l)(t + l)/(k+ 1),
Var(Qi|yi
= k) = 2(t - k)(k- l)(t + l)/(k+ 2)(k+ 1)2.
UnderHoX
the statistic
t(Q Jk) - fl E QiI
k i=l
hasthe conditionalexpectationgivenabove
whereI _ {1 if the ithanimalwascapturedexactlyk times
O otherwise.
Iffkis largeSthe test statistic,

C _ E(Q|k)- (k - l)(t + l)/(k+ 1)
k p2(t- k)(k- l)(t + 1)11/2 ' k -_ 2,. . .,t - 1
L (k + 2)(k+ 1)2fk g
canbe assumedto be approximately distributedas a standardnormal.(We
haverequiredthatfk¢ 10.)An overalltest statisticappropriate for testing
HOversusHAis givenas
FROM CAPTURE DATAtis et al. 121
tE1[E(Q|k)- (k - l)(t + l)/(k+ 1)]

C -k=2
ptz 2(t - k)(k- l)(t + 1)11/2
Lk=2(k + 2)(k+ 1) fk 2
The test statisticC is also approximately distributedas a standardnormal
if HO is true and the numberof animalscapturedat least twice is large
(greaterthan10).Testsof HOversusHA calculatedby usingeitherCkor C
shouldbe one-sidedtests since the alternativespecifiesthat the E(Q|k)
shouldbe smallerthan E(Qi|yi= k). Thus,closureis rejectedonly if the
test statisticis small.Finally,we emphasizethatthe test involvingCkis
conditionalon the value of fk,and the test using C is conditionalon the
valuesof f2,f3,. . .,ft-1
APPENDIXL
Density EstimationBased on Subgrids

The density estimationprocedureusing nested subgridsis based on
Dice's(1938)boundarystripidea.The fundamental aspectof this approach
was proposedby MacLulich(1951).Let the trappinggridhaveareaAg and
a perimeterlengthof P.Thenforanyconvexgrid(thatincludesallrectangular
grids)the effectivetrappingareaA(W) is
A(W) = Ag + PW/c + rW2/c,
where c is a conversionfactorto expressPW and w2 in the units of A.
Fromthatequation,we derivethe expectedpopulationsize atriskof capture
as
E(N)= DA(W),
where D is the true densityof animals.Dividingthroughby the known
areaAg we derive
E(N)= D[1 + aW+ bW2],
Ag
where
P b= v
Ag(c) Ag(c)
Notethatthe unknownparameters areD andW andthatE(N)is estimable
fromthe trappingstudy.
Givenat least 2 gridsof differentsizes, we can estimatethe parameters
D and W. Assumethereare k differentgrids(thesemaybe subgridsof 1
overallstudy).The relevantequationscanbe writtenas
ANi
= D[1 + aiW+ biW2]+ si, i = 1, . . ., k.
g
By assumption,E(e)= O. Let the variancecovariancematrixof e be Qi.
Weareassumingthe samedensity(D) andstripwidth(W)applyto all grids.
Thisseemsespeciallyreasonablewhenthe gridsarenested(see sectionon
DENSITY ESTIMATION).
r 1 YY A Y§T * r .
GivenestimatesNi,the aboveequationsputestimationof D andWin the

frameworkof generalizednonlinearregression.All we need to carryoutthe
estimation a knowledgeof the variance-covariance
is matrix$. LettingYi=
NIAg)we have
Var(Yi) = Var(Ni)/Ag2,
whichis estimable.Alsowe knowthat

Cov(Yi,Yi) = rijSE(ti)SE(Yj),
whererijis the correlation is notknown.Burn-

of Si andNJ.Thiscorrelation
hamand Cushwa(pers.comm.) suggestedthe following way of approxi-
. e
matlng riJ
The correlation and Nj will dependin largeparton the overlapof
of lCTi
the2 populations of size NiandNjthatin turndependson the overlapof the
areasAi(W)andAj(W).Thuslet
= Ai(W)U AA(W)'
ru= corr(Ni,Nj)
thatis, rijis the areaof the intersectionof Ai(W)andAj(W)dividedby the
areaof theirunion.
Usingthisformula, we canarriveatanestimator of$ thatseemsreasonable;
hence,fromthe live trappingdata we can compute Y1,. . .,Ykand$.
We are now in a position to obtain weighted nonlinearleast squares
estlmators ot Lsane w as Ls,w satlstylng
min (y _ tt $-1(y _ f), where
fi = D[1 + aiW+ biW2], i - 1>.. .,k

An approximate variance-covariance matrixfor the estimatorsf) and W is
givenby the 2 x 2 matrix V't-1VwhereV is the Jacobianmatrix
V= F af af 1
LdD' AW]
The elementsof the columnsof the matrixV are,respectively,

aft = 1 + aW+ bW2
1 ,
AD 6
a fi = D[ai+ bi2W], i = 17 . . .t.
Becauset dependsuponW (butnot D) an iterativeprocedureis needed

whereinan initialvalueof WOis chosen,t is computedbasedon it, andthe
new f)1>911obtained.Iterationis continueduntilstableestimatesof D and
Wareobtained.
STATISTICAL FROMCAPTURE DATAtis
INFERENCE et al.
123
APPENDIX M
General Simulation Methods

The variousestimators andtests describedin this monographweresimu-
lated to studytheir smallsamplepropertiesand operatingcharacteristics.
The asymptoticpropertiesof the tests and estimatorsare knownin most
cases,butnotthe finitesampleproperties.MonteGarlosimulations provide
a methodby whichourestimationproceduresmaybe evaluatedunderthe
exactmodelfromwhichthey were derived.Becausewe wouldnot expect
realdatato fitanymodelexactly,by simulatingdata,we canstudythe prop-
ertiesof the procedureswithoutinterferencedue to the datanot fittingthe
model.
Pseudo-random uniform(0,1)variablesweregeneratedusingtheBurroughs
6700 FORTRANintrinsicfunctionRANDOM.The mixed congruential
methodis usedby thisfunction(Anonymous 1971).
Beta variablesfor the simulationof heterogeneousprobabilitiesof first
capturewere generatedusing subroutineGGBTAfromthe IMSL (1976)
package,withthe requireduniformvariablesfurnishedby RANDOM. That
routineuses a rejectionmethod(Fishman1973).
In the mostgeneralsimulationsn the trappingprocessis simulatedforeach
animalin the populationfor each trappingoccasion.The probabilityof
capturefor a particular animalon a particularday is comparedagainsta
uniform(0,1)variable.If the valueof the randomvariableis less thanthe
probabilityof capture,then the animalis assumedcaptured,and thatele-
mentof the X matrixis set to unity.Otherwise,the valuein the X matrix
is set to zero,indicatingno capture.Whenthe X matrixis completelyfilled,
the necessaryMSSare computedand the tests and estimationcompleted.
APPENDIX N
Simutation Results
Simulation ResultsRegardingEstimationProcedures
In developingthismaterial,we used6 differentmodelsto generatesimu-
lateddata:all capture-recapturemodelsexceptMtbandMtbh. Foreachof the
datasets variousestimation proceduresconsideredherewereappliedto gain
insightinto the operatingcharacteristicsof such procedures.The results
of this simulationstudyarepresentedherein tabularform.
Foreachmodelthereare2 corresponding tables.Obviously,foranygiven
modelone maychooseany numberof sets of parameter valuesneededto
completelyspecifythe model.Thus the firsttableforeach modellists all
the differentsets of parametervalues(eachof whichis calleda Trial)used
in the simulationstudy.Forexample,Trial1 in TableN.l.a indicatesthat
therewere400 animalsin the populationandthateveryanimalhada 0.30
probabilityof captureon eachtrappingoccasion.Thatinformation is all that
is requiredto specifyan exampleof ModelMo
The secondtableforeachmodelpresentsthe simulationresultsforeach
estimationprocedureused on datageneratedfromthe Trialsof thatmodel.
Columnheadingsin the tablesare:
WILDLIFEMONOGRAPHS
124
Estimatoridentifiesthe estimationprocedureused; e.g.,
thatthe estimationproceduredescribedin AppendixC "&b indicates
appliedto the data. forModelMbwas
R-the numberof replications(datasets).
t-the numberof trappingoccasions.
Ave(N)-the averagevalueof N takenoverall replications,
i.e.,
Ave(N)= R E EJk
a(s) the '<sample'>
standard
errorof ACJ
takenoverall replications,
i.e.,
(9) 4 (&k Ave(9))2/(R - 1).
k=l
Ave+/Var(N)-
-the averagevalueof +/Var(N)
takenoverall replications,
i.e.,
Ave/Var(N)- R E jVar(Nk) )
k=l
Coverage-theproportionof replicationsin which the

confidenceintervalcontainedthe truepopulationsize N.constructed95%
Trial identifieswhichpopulation(set of parameter
data. values)generatedthe
Twopointsconcerning(N) and Ave|Var(N)shouldbe
thevalue of s;r(N) estimatesthe true standarderrorof the mentioned.First
anygiven Trialand Modelso that a measureof the estimatorN in
estimatorA/Var(N) is givenby |Ave+ar(N)- (r(N)|/C(N). relative bias of the
confidence Second the
intervalwidthin a given Trialand Modelis givenby 2average
Ave/larN).
ls.
:e v
(1.96)
As an xamplesconsiderthe firstline of Table N.l.b.
datasets, eachconsistingof 5 trappingoccasions We see that200
toModel Mo Trial 1 and that the estimation were generatedaccording
Model Mowasappliedto eaehset.Forthose2Q0data procedureassociatedwith
unbiased setsSNowasessentially
- (Ave(N,)-N
- --0.7), as was the estimatorof the standard error
ofNo(Avewar(NO) - (N) = 0.05).Moreover, 95
confidence percent of
intervalscoveredthe truevalueof N. Thisachieved the constructed
coefficientof 0.95 corresponds confidence
exactlywith the statedlevel of the
namely 0.95. Such excellentperformance intervalS
cedureis due in largepartto the factthatofltothe confidenceintervalpro-
and war(N) appearto be
essentiallyunbiased.Finally,note thatthe averagewidth of the 200 con-
structed
confidenceintervalsis givenby 2(1.96)(11.96) = 46.88.
Simulation ResultsRegarding the Size andPowerof TestingProcedures
Becausethe testingof assumptions plays a vital role in the analysisof
capture-reeapture
datait is important
-
to havesomeinsightintotheoperating
characteristics
of such tests. Simulationstudiesof the tests T1
defined
in AppendixKwerecarriedouton throughT7
datasimulated from all
providesuchinsight,andthe resultsarepresentedin Tables 8 models
to
N.13.In addition,Tables N.8 and N.9 give resultsof an N.7 through
to
the powerof testsT2andT3basedon theoretical approximation
resultsin AppendixK.
STATISTICAL
INFERENCE
The columnheadingsof the tablesare:

ct indicatesthe nominalsize of the test.
DataModel indicatesthe Trialand Modelthatwereused to generatethe
data.
R indicatesnumberof replications (datasets).
t indicatesnumberof trappingoccasions.
The readerwill notethatin someinstancesthe numberof replications R
appearsto be strange.The explanation is thata few of the estimatorsand
tests simulatedare subjectto "failure,>'
i.e., a particular
dataset mayhave
characteristics
thatdo notallowcalculationof the desiredstatistics.In those
cases,the datasets areexcludedfromthe reportedsimulationresults.
Finally,in TableN.14we givea deseription oftheTrialsof ModelsMtband
Mtbh. ThoseTrialswere involvedin the simulationof someof the tests of
modelassumptions.
TABLE N. l.a. lDEscRIprIoN ON THE TR1ALS OF

MODEL MO
N p Trial
400 0.30 1
400 0.10 2
400 0.05 3
200 0.25 4
100 0.20 5
100 0.15 6
TABLE:N.l.b.-SIMULATION RESULTSOF ESTIMATIONPROCEDURESUSED ON r}ATAGENERATEDACCORD-

ING TO MODEL MO
Estimator N Ave(N) (r(N) Ave) Coverage R t Trial
No 400 399.30 11.91 11.96 0.950 200 5 1

No 400 456.93 219.31 160.25 0.922 500 5 3
No 400 405.97 52.84 55.54 0.930 200 5 2
No 100 101.70 19.43 - - 200 5 6
No 400 397.34 46.56 - - 200 5 2
No 400 453.82 203.75 - - 372 5 3
No 400 405.73 52.54 - - 200 5 2
No 200 199.60 6.58 - - 100 7 4
Nh 100 113.46 17.25 13.74 0.830 200 5 6
Nh 400 386.89 27.65 27.93 0.920 200 5 2
Nh 400 389.12 29.56 28.02 0.898 400 5 2
Nh 200 226.38 10.77 10.07 0.260 100 7 4
Nb 400 319.25 204.51 - - 372 5 3
Nb 400 485.90 387.42 - - 198 5 2
Nb 200 198.59 11.66 10.47 0.880 100 7 4
Nt 400 407.71 61.81 57.45 0.940 400 5 2
Nt 400 439.96 161.42 148.13 0.940 500 5 3
Nt 200 199.37 6.51 6.85 0.950 100 7 4
TABLE N.2.a. DESCRIPrION

OF THETRIALSOF MODELMt
N (Pl,P2,--,PJ Trial
400 (0.55, 0.55, 0.50, 0.45, 0.45) 1

400 (0.05, 0.10, 0.15, 0.15, 0.05) 2
400 (0.10, 0.10, 0.10, 0.10, 0.01) 3
400 (0.01, 0.01, 0.02, 0.03, 0.03) 4
400 (0.04, 0.0S, 0.03, 0.07, 0.06) 5
400 (0.60,0.40,0.20,0.10,0.10,0.10,0.10,0.10,0.10,0.10) 6
400 (0.50, 0.20, 0.10, 0.10 0.10, 0.10, 0.10) 7
400 (0.60 0.40, 0.20, 0.10, 0.10) 8
400 (0.20, 0.40, 0.30, 0.10, 0.20, 0.30, 0.20) 9
200 (0.30, 0.40, 0.10, 0.40, 0.30) 10
800 (0.02, 0.01, 0.03, 0.03, 0.01) 11
100 (0.05, 0.05, 0.10, 0.15, 0.15) 12
TABLE N.2.b. SIMULATIONRESULTSOF ESTIMATIONPROCEDURESUSED ON DATAGENERATEDACCORD-

ING TO MODEL Mt
Estimator N Ave(N) cr(N) Ave) Coverage R t Trial
Nt 400 399.30 202.39 348.63 0.839 155 5 4

Nt 400 408.57 73.91 74.19 0.949 198 5 3
Nt 100 108.44 37.18 35.77 0.942 989 5 12
Nt 800 1,015.62 630.46 748.23 0.902 194 5 11
Nt 400 400.33 54.50 56.00 0.955 200 5 2
Nt 400 399.28 3.55 3.86 0.960 200 5 1
Nt 400 442.66 187.00 151.93 0.930 200 5 5
Nt 400 397.14 206.97 341.61 0.880 292 5 4
Nt 400 398.59 15.64 16.45 0.940 100 7 7
Nt 400 398.81 8.03 7.57 0.950 100 10 6
Nt 200 198.70 8.98 7.98 0.880 100 5 10
Nt 400 399.41 lQ.16 9.96 0.930 100 7 9
Nt 400 399.45 12.80 11.94 0.940 100 5 8
Nh 400 349.17 27.27 26.54 0.515 200 5 3
Nh 400 100.83 27.37 12.93 0.000 200 5 4
Nh 100 98.70 14.43 13.77 0.924 1,000 5 12
Nh 800 219.60 27.94 20.79 0.000 200 5 11
Nh 400 244.46 24.04 22.17 0.000 200 5 5
Nh 400 98.56 30.27 12.43 0.000 400 5 4
Nh 400 506.72 27.76 22.00 0.000 100 5 8
Nh 400 516.22 31.59 26.91 0.000 100 7 7
Nh 200 456.68 50.28 15.91 0.000 100 10 6
Nh 400 235.94 15.78 12.54 0.170 100 5 10
Nh 200 459.60 18.32 15.30 0.010 100 7 9
Nbhl 400 199.70 21.08 18.08 0.810 100 5 10
Nbhl 400 410.30 54.16 70.10 0.860 100 7 9
Nbh 400 387.07 21.14 27.63 0.540 100 10 6
Nbhl 400 364.06 55.82 65.24 0.510 100 7 7
Nbh 400 342.56 9.48 5.72 0.040 100 5 7
I Note, we use Nbhand NRas equivalent notation.
STATISTICALINFERENCEFRONICAPTUREDATA C)tiS et 1.
127
TABLE N.3.a. DESCRIPrION OF THE TRIALS OF
MODEL Mb
N P C Trial
400 0.30 0.10 1
400 0.20 0.50 11
400 0.20 0.05 2
400 Q.10 0.30 3
400 0.10 0.15 4
200 0.30 0.50 5
200 0.25 0.25 6
200 0.20 0.10 7
100 0.20 0.05 8
100 0.20 0.50 9
100 0.10 0.30 10
100 0.40 0.20 12
TABLEN.3.b.-SIMULATION
RESULTS
OF ESTIMATION
PROCEDURES
USEDONDATAGENERATED
ACCORD-
INGTOMODELMb
Estimator N Ave(N) a(N) Ave) Coverage R t Trial
A
Nb 100 82.00 36.73 64.76 0.708 161 5 10

Nb 200 208.09 38.53 40.27 0.930 100 5 7
Nb 200 198.36 13.32 13.13 0.900 100 5 5
Nb 400 401.23 25.43 19.91 0.920 100 5 1
Nh 100 109.83 48.05 - - 199 5 8
Nh 400 400.22 22.38 - - 200 5 1
Nb 400 511.04 408.42 - - 193 5 3
Nb 400 403.56 58.70 - - 200 5 2
Nb 200 lD7.88 13.32 - - 200 5 5
Nb 400 461.48 269.25 - - 196 5 4
Nb 200 206.99 37.11 - - 200 5 7
Nb
A
200 198.59 11.66 10.47 0.880 100 7 6
Nh 100 82.39 6.75 5.09 0.140 100 5 9
Nh 100 176.49 14.40 18.82 0.000 100 5 8
Nh 400 347.67 29.55 26.16 0.495 200 5 4
Nt 100 71.75 4.80 2.43 0.000 100 5 9
Nt 100 299.25 193.37 115.92 0.630 100 5 8
No 100 51.38 7.06 - - 161 5 10
No 100 293.88 109.76 - - 199 5 8
No 400 720.39 57.05 - - 200 5 1
No 400 212.48 13.56 - - 193 5 3
No 400 1,070.94 176.01 - - 200 5 2
No 200 175.18 5.80 - - 200 5 5
No 400 305.73 33.32 - - 196 5 4
No 200 311.67 48.50 - - 200 5 7
TABLE N.4.a.-DESCRIPTION OF THE TRIALSOF MODEL Mh
N Trial
Pl, i = 1,2,. . .,N
400 Pi= O.05, i = 1,200; Pi = 0.15, i = 201,300; Pi = 0.50, i = 301,400. 1
400 Pi= 0.01, i = 1,100; Pi= 0.05, i = 101,200; Pi= 0.10, i = 201,300; 2
Pi= 0-20, i = 301,400.
400 Pi= 0.10, i = 1,100; Pi= 0.20, i = 101,200; Pi = 0.25, i = 201,300; 3
Pi= 0.30, i = 301,400.
400 Pi= 0.01, i = 1,50; Pi= 0.15, i = S1,200; Pi= 0.25, i = 201,300; 4
Pi = 0.30, i = 301,400.
400 Pi= 0-20, i = 1,100; Pi = 0.30, i = 101,200; Pi = 0.40, i = 201,300; 5
Pi= 0.50, i = 301,400.
200 Pi= 0.05, i = 1,50; Pi= 0.15, i = 51,150; Pi= 0.25, i = 151,200. 6
200 Pi= 0-15, i = 1,50; Pi= 0.20, i = 51,100; Pi= 0.25, i = 101,150; 7
Pi= 0.30, i= 151,200.
100 Pi = 0.05, i = 1,40; Pi = 0.10, i = 41,80; Pi = 0.30, i = 81,100. 8
100 Pi :(3,22), i = 1,100. 9
100 Pi (1,22/3), i = 1,100. 10
100 Pi (3/22,1), i = 1,100.1 11
100 Pi = 0.10, i = 1,40; Pi = 0.20, i = 41,80; Pi = 0.30, i = 81,100. 12
400 Pi= 0.05, i = 1,50; Pi= 0.10, i = 51,200; Pi= 0.15, i = 201,300; 13
Pi = 0-25, i = 301,400
200 Pi= 0.05, i = 1,50; Pi= 0.10, i = 51,150; Pi= 0.25, i = 151,200. 14
l Indicates that the probabilities pj were a randomsample of size 100 from a beta probabilitydistributionwith the indicated parameter
TABLE N.4.b.-SIMULATION RESULTSOF ESTIMATIONPROCEDURESUSED ON DATAGENERATEDACCORD-

ING TO MODELMh
Estimator
N Ave(N) a(N) Ave) Coverage R t
Trial
N^h
100 35.41 9.36
_ _
4.38 0.000 200 5 11
N^h
100 79.46 5. 1b 10.91 0.545 200 10
N^h
100 84.66 14.32 11.56 0.675 200 5 8
Nh
400 331.06 23.99 22.69 0.180 200 1
Nh
400 298.06 24.52 24.22 0.000 200 -
2
Nh
100 100.17 21.15 10.54 0.675 200 10 8
Nh
400 460.10 32.31 26.77 0.395 200 f
t,
3
Nh 200 226.78 9.82 11.94 0.440 100 7 7
Nh
400 443.9S 19.71 152*U 0.170 100 5
N^h
400 461.22 32.64 26.92 0.410 100
Nh
400 417.08 30.91 24.92 0.850 100 4
Nh 100 107.57 6.80 12.01 0.810 100 5 12
Nh
200 207.34 6.14 11.59 0.870 100 10
t,
,
NAO 100 26.83 4.79 200 11

No 100 62.68 2.39 200 5 10
NÔ 100 68.90 2.43 200 8
NO 400 227.83 10.97 200 5 1
N 400 260.29 30.93 200 s
2
No 100 73.72 7.28 200 10
8
NAO 400 365.12 7.15 200 3
Nt
200 193.61 9.70 7.69 0.770 100 7 7
Nt
400 376.18 10.00 7.67 0.240 100
Nt
400 364.31 6.44 17.01 0.460 100 5
N^t
400 333.17 7.92 16.19 0.060 100 tJ
4
Nt
100 87.87 2.07 10.19 0.690 100 5 12
Nt
- 200 173.64 8.68 6.83 0.120 100 10 6
Nt
400 228.02 0.73 7.40 0.000 100 1
200 Pi- 0.25, i = 1,50; Pi 0.50, i - 51,150; P1 = 0.15, P2 = 0.25 p3= 0.05, p4 = Q.1()7 6
FROMCAPTUREDATAtiS
INFERENCE
STATISTICAL et 41. 129
TABLE N.5.a. DESCRIPrION OF THE TRIALS OF MODEL Mth. THE PROBABILITYTHAT THE ith ANIMAL
IS GAUGHTON THE JthTRAPPINGOCCASIONIS EQUALTQ PiPj j = 1, . . ., t AND i-17 ... N
N pX,i = 1,S, . . .,N PJ,i = I,2,. . .,t Trial
4()0 Pi= 0 15, i = 1,10(); Pi - 0.33, i = 1()1,200; Pl = 0-30, P2 = 0.60, p3 = 0.10, p4= ()*30
Pi= 0-67, i = 201n300; Pi= 1.00, p5= 0.60
i = 301,400.
400 Pi - Q.40, i = l,lOQ, Pi= 0.60> i = 101,200; P1 - P2= 0.25, p3= 0.50 p4- 0.15, 2
Pi= 0.80, i = 2017300; PX- 1.00, p5 = 0.25.
i = 301,400.
40() Pi - 0.30, i = 1,150; Pi= 0.40, i = 151,250; Pl - 0.15, P2 = 0.257 p3 - 0.057 p4= (:).10, 3
Pi = 1.007 Pi = 251,400. p5 = 0 3Q
4()0 Pi = (}.03, i = 1,100; Pi = 0.35, i = 101,2()0, P1 = P2 = P3 = 0.35, p4 = o.40, p5 = 0.60. 4
Pi= 0.55, i = 201,300; Pi= 1.00,
i= 301,400.
200 Pi = 0.4Q, i = 1,50; Pi - 0.6Q7i = 51,100; P1 - () 5°7 P2 = P3 = 0-30, P4 - ° 6() 5
Pi = 0.80, i = 101,1$0; Pi = 1.00, p5 = o-2o, P6 = 0 40, P7 = 0.30.
i= 151,200.
Pi = 1.00, i = 151,200. PS = 0 30
400 Pi = 0.45, i = 1i100; Pi - 0.5S, i - 101,200; Pl = 0.65, P2 = 0 75, P3 = 0 55, P4 - 0 35 7
Pi - 0.65, 1 - 2017300; Pi = 0.75, p5 = 0.60> P6 = 0-50, p7 = 0.80.
i = 301,400.
100 Pi - ° 351 i = 1?25; Pi _ 0.45 i = 26750, P1 0.657 I)2 0.75 PS 0-55, P4 = () 35 8
P1= 0-55; i = 51,75; Pi= O.65, Ps= 0.60
i = 7G,100.
TABLE N.5.b. SIMULATIONRE:SULTSOF ESTIMATIONPROCEDURESAPPLIED TO r)ATA GENERATED

ACCORDINGTO MODEL Mth
Estimator N Ave(N) <7,(T) Ave ) Gowerage R t Trial
Nt 400 303.60 14.13 11.74 0.000 100 5 1

Nt 400 369.90 21.69 19.69 0.600 100 5 2
Nt 400 309.04 37.46 35.66 0.340 100 5 3
Nt 400 272.77 12.53 10.32 O.Q00 100 5 4
191; 200 187.93 5.68 5.29 0.360 100 7 5
Nt 2Q0 164.17 34.05 29.11 0.590 10() 5 6
Nh 400 402.88 31.42 23*46 0.850 100 5
Nh 400 478.73 33*55 29 20 0.220 100 S 2
Nh 400 353.9() 25.02 26.62 0.590 100 5 3
Nh 4Q0 352.78 29.38 20.68 0.430 100 5 4
Nh 20Q 217.72 8.42 9.46 ().560 100 7 5
Nh 200 178.21 21.29 18.67 0.760 1()0 5 6
Nbh 400 Failed in all 100 replications 100 5 1
Nbh 400 272.56 11.65 6.27 0.000 100 5 2
Nbh 400 Failed in aI1 100 replicatiolls 100 5 3
Nbh 400 489.06 275.72 319.02 0.840 100 o 4
Nbh 200 179.41 9.27 9.16 0.34() 100 7 5
Nbh 200 23$.91 156.90 337.63 0.793 92 5 6
TABLE N.6.a. DESCRIPrIONOF THE TRIALSOF MODEL Mbh
N P,,C,;i = 1,2, . . ., N TrjAl

400 Pi= 0.05, i = 1,200; Pi= 0.15, i = 201,300; Pi = 0.50, i = 301,400.
400 Pi = 0.01, i = 1,50; Pi = 0.15, i = 51,200; Pi = 0.25, i = 201,300; Pi = 030' 2
i = 301,400.
400 Pi = 0.10, i = 1,100; Pi = 0.20, i = 101,200; Pi = 0.25, i = 201,300; Pi = 030' 3
i = 301,400.
4°° Pi = 0 20, i = 1,100; Pi = 0.30, i = 101,200; Pi = 0.40, i = 201,300; Pj - 0 50' 4
i = 301,400.
200 pj= 0.05, i = 1,50; Pi= 0.15, i = 51,150; Pi= 0.25, i = 151,200. 5
200 Pi = 0.15, i = 1,50; Pi = 0.20, i = 51,100; Pi = 0.25, i = 101,150; Pi = 030' 6
i= 151,200.
100 Pi = 0.10, i = 1,40; Pi = 0.20, i = 41,80; Pi= 0.30, i = 81,100. 7
400 Pi = 0.10, i = 1,50; Pi = 0.25, i = 51,200; Pi = 0.35, i = 201,300; Pi = 045' 82
i = 301,400; ci= max[(pi- ran(i)/4),0].
400 Pi= 0.05, i = 1,50; Pi= 0.10, i = 51,200; Pi = 0.15, i = 201,300; Pi= 0.25, 92
i = 301,400; ci = min[(pi + ran(i)/4),1].
400 Pi= 0.05, i = 1,50; Pi= 0.10, i = 51,200; Pi= 0.15, i = 201,300; Pi= 0.25, lO2
i = 301,400; ci= max[(pi- ran(i)/4),0].
100 Pi p(1,22/3), i = 1,1003; ci = min[(pi + ran(i)/2,1)] ll2
100 Pi p(1,22/3), i = 1,1003; ci = max[(pi - ran(i)/2),0] l22
100 pj :(1,22/3), i = 1,1003; ci = min{max[pi + (ran(i) - 0.5)/2,0],1} l32
400 Pi = 0.15, i = 1,100, pl = 0.25, i = 101,300; Pi = 0.35, i = 301,400 ci = 0.5 Pi 14
100 Pi = 0.15, i = 1,33; Pi = 0.25, i = 34,66; Pi = 0.35, i = 67,100. ci = 0.5 Pi 15
' Indicates the trial was used for estimation purposes only. Because the performanceof the estimator associated with Model Mbh
depends only upon N and the probabilitiesof firstcapturepi, recaptureprobabilities;cl need not be specified.
2 The function ran(i)produces a randomvalue of a variable distributeduniformlyover the interval [0,1].
3 Pl (1,22/3) indicates that the randomvariablepi has a beta distributionwith parameters1 and 22/3.
TABLE N.6.b. SIMULATIONRESULTSOF ESTIMATIONPROCEDURESUSED ON DATAGENERATEDACCORD-

ING TO MODEL Mbh
Estimator N Ave(N) a(N) Ave) Coverage R t Trial
Nbhl 400 246.59 38.04 32.26 0.120 100 5 1

Nbh1 400 340.83 57.57 47.90 0.360 100 5 2
Nbh1 400 366.43 41.87 35.63 0.600 100 5 3
Nbh 400 383.00 21.30 15.89 0.560 100 5 4
Nbh1 200 175.51 16.26 13.52 0.380 100 10 5
Nbh1 200 193.72 13.90 13.72 0.780 100 7 6
Nbhl 100 94.14 43.26 41.48 0.620 100 5 7
I Note, we use Nbhand NRas equivalent notations.
TABLE N.7. SIMULATIONRESULTS CONCERNING

THE SIZE AND POWEROF THE TEST T1 OF MODEL
MOVS. MODEL Mh
0.01 0.05 0.10 Data model R t
0.0100 0.0300 0.0550 MovTrial 2 200 5

0.0100 0.0150 0.0450 MO,Trial 6 200 5
0.9900 0.9900 0.9900 Mh, Trial 1 200 5
0.0900 0.1150 0.1400 Mh, Trial 2 200 5
0.0850 0.1350 0.1900 Mh, Trial 3 200 5
0.1350 0.1650 0.1950 Mh, Trial 8 200 5
0.3950 0.5650 0.6250 Mh, Trial 8 200 10
0.1700 0.1750 0.2500 Mh, Trial 10 200 5
0.7450 0.7750 0.8400 Mh, Trial 11 200 5
TABLE N.8. SIMULATEDAND THEORETICALRESULTSCONCERNINGTHE SIZE AND POWEROF THE TEST

T2 OF MODELMOVS. MODEL Mb
a
Method 0.01 0.05 0.10 Data model R t
Simulation 0.0100 0.0550 0.086Q MO Trial 2 198 5

Theoretical 0.0100 0.0500 0.1000 M,,,Trial 2 198 5
Simulation 0.0280 0.0830 0.1240 MO,Trial 3 372 5
Theoretical 0.0100 0.0500 0.1000 MO,Trial 3 372 5
Simulation 0.0600 0.1300 0.1400 MO,Trial 4 100 7
Theoretical 0.0100 0.0500 0.1000 Mo Trial 4 100 7
Simulation 1.0000 1.0000 1.0000 Mb,Trial 1 200 5
Theoretical 0.9990 0.9990 0.9940 Mb, Trial 1 200 5
Simulation 0.9150 0.9700 0.9800 Mb, Trial 2 200 5
Simulation 0.9700 0.9900 1.0000 Mb,Trial 5 200 5
132 WILDLIFE
MONOGRAPHS
TABLE N.9.-SIMULATED AND THEORETICALRESULTSCONCERNINGTHE SIZE AND POWEROF THE TEST
T3 OF MODEL MOVS. MODEL Mt
a
Method 0.01 0.05 0.10 Data model R t
Simulation 0.0150 0.0525 0.1100 MoX Trial 2 400 5

Simulation 0.0120 0.0500 0.1060 Mo Trial 3 500 5
Simulation 1.0000 1.0000 1.0000 Mt,Trial 2 200 5
Theoretical 0.9998 0.9999 0.9999 Mt,Trial 2 200 5
Simulation 0.3800 0.6550 0.7350 Mt,Trial 5 200 5
Theoretical 0.4269 0.6635 0.7703 Mt,Trial 5 200 5
Simulation 0.6138 0.8028 0.7984 Mt, Trial 12 989 5
Theoretical 0.5925 0.7984 0.8751 MtnTrial 12 989 5
TABLE N. 10.-SIMULATION RESULTSCONCERNING TABLE N. 12. SIMULATIONRESULTSCONCERNING

THE SIZE AND POWEROF THE GOODNESSOF FIT THE SIZE AND POWEROF THE GOODNESSOF FIT
TEST T4 OF MODEL Mh TEST T6 OF MODEL Mt
0.01 0.05 0.10 Data model R t 0 01 0.05 0.10 Data model R t
0.0200 0.0500 0.0900 Mo Trial 2 100 5 0.0000 0.0200 0.1100 Mt, Trial 13 100 7
0.0000 0.0400 0.0900 Mh, Trial 3 100 5 0.0000 0.0300 0.0600 Mt, Trial 14 100 5
0.0000 0.0500 0.1000 Mh, Trial 8 100 10 0.0000 0.0100 0.0900 Mtb,Trial 1 100 7
0.0000 0.0800 0.1300 Mh, Trial 10 100 5 0.0100 0.0700 0.1200 Mtb, Trial 2 100 5
1.0000 1.0000 1.0000 Mbh,Trial 14 100 7 0.0600 0.2800 0.3600 Mth, Trial 7 100 7
0.0900 0.2300 0.3700 Mbh,Trial 15 100 5 0.0300 0.0700 0.2000 Mth,Trial 8 100 5
1.0000 1.0000 1.0000 Mth,Trial 7 100 7 0.9300 0.9800 0.9900 Mtbh,Trial 1 100 7
0.5400 0.7900 0.8800 Mth, Trial 8 100 5 0.0200 0.0600 0.1000 Mtbh,Trial 2 100 5
1.0000 1.0000 1.0000 Mtbh,Trial 1 100 7
0.1600 0.3300 0.4200 Mtbh, Trial 2 100 5
TABLE N. 13.-SIMULATION RESULTSCONCERNING

TABLE N. 1 1. SIMULATIONRESULTSCONCERNING THE SIZE AND POWEROF THE TEST T7 OF MODEL
THE;SIZE AND POWEROF THE GOODNESSOF FIT Mh VS. MODEL Mbh
TEST T5 OF MODEL Mb
cx
0.01 0.05 0.10 Data model R t
0.01 0.05 0.10 Data model R t
0.0100 0.0800 0.1500 Mh,Trial 10 100 S
0.0000 0.0200 0.0200 Mb, Trial 11 100 7 0.0100 0.0300 0.0800 Mh, Trial 13 100 5
0.0000 0.0500 0.1200 Mb, Trial 12 100 5 0.0600 0.0700 0.0800 Mh, Trial 14 100 10
0.0000 ().0600 0.1000 Mbh,Trial 14 100 7 0.4000 0.6700 0.7600 Mbh,Trial 8 100 S
0.0000 0.1000 0.1600 Mbh Trial 15 100 5 0.1500 0.4000 O.S100 Mbh,Trial 9 100 5
1.0000 1.0000 1.0000 Mtb,Trial 1 100 7 0.0900 0.2300 0.3300 Mbh,Trial 10 100 5
0.3300 0.5700 0.7100 Mtb,Trial 2 100 5 0.2800 0.5000 0.6100 Mbh,Trial 11 100 5
1.0000 1.0000 1.0000 Mtbh,Trial 1 100 7 0.0200 0.1400 0.1800 Mbh, Trial 12 100 5
0.0600 0.1800 0.3000 Mtbh,Trial 2 100 5 0.0100 O.OS00 0.1100 Mbh,Trial 13 lOQ 5
STATISTICAL
INFERENCE
TABLE N.14. DESCRIPTIONOF THE TRIALSOF MODELSMtb AND MtBh
Mtbl
N Pj,j = 1,2, . . .,t c Trial
400 P1 = ° °S, P2 = 0.35, p3 = 0.25, p4 = 0.15, 2.5 1

P5 = 0.20, P6= 0.10, p7 = 0.30.
100 P1 = 0 35, P2 = 0.25, p3 = 0.15, p4 = 0.20, 0.5 2
P5= 0.30.
Mtbh
N P,, i = 1,2, . . .,N Pj,j = 1,2, . . .,t c Trial
400 Pj = 0.15, i = 1,100; Pj = 0.25, Pl = 0.65, P2 = 0 75, P3 = 0.55, p4 = 0.45, 2.5 1

i = 101,200; Pi = 0.35, i = 201,300; p5 = 0.60, P6 = 0 50, p7 = 0.70.
Pi= 0.45, i = 301,400.
00 Pi = 0.15, i = 1,25; Pi = 0.25, i = 26,50; P1 = 0.65, P2 = 0 75, P3 = 0.55, p4 = 0.45, 0.75 2
Pi= 0.35, i = 51,75; Pi= 0.45, P5= 0.60.
i = 76,100.
' The probabilityof any animal being capturedon the jth occasion, j = 1,2, . . .,t, is pj if the animal has not previously been caught and
cpj if the animal is being recaptured.
2 The probabilityof the ithanimal being capturedon the jth occasion is PiPiif the animal has not previously been caught and pjpjcif the
animal is being recaptured.
APPENDIXO
Interval Estimation
Use of the CentralLimit Theorem (cf. Mood et al. 1974:195)in setting
so-called"normaltheory"confidenceintervalsforparametersis widespread.
The theorem usually is cited as justificationfor asserting that, for "large
samples,"a 95 percentconfidenceintervalfor the parameterof interest0 is
given by P{0 - 1.96(T(0) S 0 S 0 + 1.96ff(0)}= 0.95, where (0) represents
an estimate of the standarddeviation of 0 (also see Seber 1973:134for an
exampleof an indirectlyconstructedconfidenceintervalbased on asymptotic
normality).Furtherimpetus is given to this argumentwhen ML estimators
are involved by the knowledge that, under certain regularityconditions,
those estimatorsare BAN (Best AsymptoticallyNormal).Furthermore,large
sample (normaltheory) confidence intervals based on ML estimatorsare
knownto have smallerexpected width than intervalsconstructedusing any
otherestimator(cf. Moodet al. 1974:393).
Unfortunately,it is generallytrue thatassertionsconcerningthe operating
characteristicsof large sample intervals cannot be made if sample sizes
are small or regularityconditions are not met or both. Even more un-
fortunateis the fact that small or moderatesample sizes are more often the
rule than the exceptionin capture-recaptureexperiments.Therefore,it was
not unexpectedthat initial simulationof capture-recaptureexperilnentsre-
vealed that "normaltheory"confidence intervalsoften exhibit undesirable
properties.For example, the lower limit of a given intervalfor population
size N may be less than the number of differentanimals capturedin the
experiment.Also, coverage of the confidence interval,i.e., the percentage
of simulated intervals that contain the true value N, is often significantly
less than the nominallevel of 0.95. Because of such problems,2 alternative
interval estimationprocedureswere investigatedin the hope that a more
satisfactoryprocedurecould be suggestedfor practicaluse.
134 WILDLIFE NIONOGE{APHS
The firstprocedureis baseduponthe suppositionthatthe distribution of

the estimator&-1is moresymmetric(hencecloserto normality)thanthe
distributionof N (cf. Cormack1968).Thus,the followingprocedurewas
proposed.First,calculatethe ML estimatesN and Var(N).Constructa 95
percentconfidenceintervalfor N-1 of the formP{N-t - 1.96N-2v/Var(N) S
N-1< N-1+ 1.96N-2+Var(N)}. Finally,invertthe intervalin the obvious
mannerto arriveat a confidenceintervalforN. Simulation resultsinvolving
ModelMtshowedthat,althoughthe distribution of 19-1was nearlynormal,
the aboveproceduredid not representa significantimprovement overthe
usuallargesampleinterval.Coverageof the 2 procedureswas roughlythe
same,but the (4reciprocal>'procedurehad, on the average,greaterwidth
thanthe usuallargesampleprocedure.Moreover,lowerconfidencelimits
forN-1 sometimeswere less thanzeroandthus uponinversion,the upper
limitfors wasnegative.
The secondalternative methodforintervalestimationinvolvedthe use of
only the likelihcsodfunctionof the sample and is basedlargelyuponthe
likelihoodprinciple(cf.KendallandStuart1973:226). Thatprincipleasserts
that the likelihoodfunctionprovidesall the informationnecessaryfor
makingstatisticalinferencesconcerningthe dataand has been subjected
to serioustheoreticalquestioning(Stein1962,Birnbaum1968>Kendalland
Stuart1973).Neverthelesswe felt thatit wouldbe beneficialto simulate
the operatingcharacteristics of these 4'likelihoodintervals"in capture-
recapture experiments to evaluatetheirpracticalutility.
Briefly,the mechanicsof constructing a likelihoodintervalareas follows.
(The readeris referredto Hudson[19711for a thoroughexplanation.) For
a given dataset X formthe log-likelihood functionInL(8|X).(Assume8 is
a scalarforsimplicityof presentation.)
Underthe assumption that@,the ML
estimatorof @is unique,andthatthe likelihoodfunctionis unimodal,the
likelihoodintervalI(8)is definedas I(0)- {0:lnL(0|X)> InL(0|X)- 2}. I(0)
consists of all those 0 for which InL(@ &X) (the log-likelihoodfunction
evaluatedat #) is no morethan2 unitsawayfromthe maximum valueof the
likelihoodfunctionInL(o|X).The assumptionis thatthese valuesof o are
<'plausible>>
valuesof @>in view of the dataobserved.Thatis, these values
of 8 producevaluesof the likelihoodfunctionthatare not 4far2'fromits
maximumS andthusthey cannotbe discounted.The use of the value2 may
seemarbitrarybutHudson(1971)arguedthatthisvalueleadsto asymptotic
95 percentconfidenceintervals.
Likelihoodintervalsfor populationsize N were constructedfromdata
simulatedaccordingto the 2-sampleremovalexperimenttreatedby Seber
and Whale(1970).Varyingthe valuesof N and p (probability of removal)
did not appearto significantlyaffectcoverageprobabilityof the intervals
althoughit is theoretically
truethatthe coverageprobability variesat least
slightlywith the true valuesof the parameters. Moreoverlowerlimitsof
the intervalsdid not extendbelow the numberof animalsseen. Average
coverageof the likelihoodintervals(95.5%)was approximately the sameas
thatof the "normal theory"confidenceintervals(92.2%) constructedfromthe
samedata.However,averageintervalwidthforthe likelihoodintervalswas
consistentlygreaterthan that of the normalconfidenceintervals;on the
averagetheywereapproximately 10 percentgreater.In view of thesesome-
whatmixedresults,a secondsimulationstudywasperformedcyndatafrom
Moce Mt T lat stucy reveaec* similarresults,in that coverageof the 2
FROMCAPTUREDATAtis et al. 135
procedures wason the averageapproximately the same(96.0%forlikelihood

vs. 93.2Nofor"normal" intervals),andthe averagewidthforthe likelihood
intervalswas alwaysgreaterthanthatof the normalconfidenceintervals.
In one case, the averagewidthof the likelihoodintervalwas morethan3
times the averagewidth of the normalconfidenceinterval.Finally,in
virtuallyevery simulationinvolvinglikelihoodintervals,we noted that
approximately half of those intervalsthatdid not containN were too low
(i.e., the upperlimitof the intervalwasless thanN) andhalfweretoo high
(i.e., the lowerlimit of the intervalwas greaterthanN). This is in sharp
contrastto the resultsof simulatingnormalconfidenceintervals,wherewe
have foundthat the largemajorityof intervalsthat do not containN are
too low.
Onthe basisof the resultsdescribedabove,we recommend continueduse
of the usuallargesampleconfidenceintervalprocedureratherthaneither
of the 2 alternativeproceduresdiscussed.All 3 procedurespossessboth
attractive andunattractiveoperatingcharacteristics,andthe choicetherefore
cannotbe clear-cut.The decisionto continuethe use of largesamplecon-
fidenceintervalswas madefor2 basicreasons.First,biologistsin general
tend to be morefamiliarwith the computation and use of thatprocedure.
Second,andmoreimportant, is thefactthatmoreis knownaboutthetheoreti-
cal propertiesof the largesampleprocedurethanis knownaboutthe 2 al-
ternatives,and,therefore,it maybe possibleto assesstheoretically the small
samplebehaviorof the procedurein certaincapture-recapture situations.
Finally,we wish to reemphasizethat large sample"normal"confidence
intervalsshould be used with greatcautionin many capture-recapture
experiments, andthatmuchmoretheoreticalworkappearsnecessarybefore
moreadequateprocedures areavailable.

Otis Etal 1978

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Statistical Inference from Capture Data on Closed Animal Populations

Author(s): David L. Otis, Kenneth P. Burnham, Gary C. White, David R. Anderson

A Publicationof The WildlifeSociety

OCTOBER 1978 No. 62

STATISTICAL INFERENCE FROM

David L. Otis,l KennethP. Burnham,2Gary C. White,3

INTRODUCTION 7 Example --- ---------------- 35

On the Needfor an Objective Selection APPENDIXB 105

INTRODUCTION studies (a removalstudy is, of course,

Onlyclosedpopulationswill be consid- and analysisInethodswe do not cover

We hope this publicationand the asso- Overtonand Davis (1969:404):"Com-

In thatcontext,openandclosedmodels catchable,or nearlyso, is indistinguish-

UnequalCaptureProbabilities casion).This situationmaybe common

Biologistswho have datafromclosed will need to readmostsections,excep-

the manuscriptare also muchappreci- X1 1 X12 , . . Xlt

life Service, providedencouragement _ XN1 XN2 XNt _

(whichwe neverwill forrealdata).It is m. = sumof the mj= E mj.

sumptionandit need not be a sourceof

n. = the totalnumberof capturesdur- estimator is onethatis notsensitivetothe

approximations of N suggestedfor this IntervalEstimation

parturesfrolnthe underlyingassump- theoryforestimatingsuchparalneters as

markingan animalaffectsits probability tiorzclosurewas reasonable,

NUS3ER OF TRAPPING OCCAS

comingtrapshy will causeNoto overes- ESTIMATEDPABILITY OF CAPTWE, P-T - .4672

timateN, and vice versawhen animals

given in TablesN.3.band N.4.bof Ap- APPROXIMATE CONFIDE:NCE

pendixN. Resultsalsorevealthatthe es- FIG. 1. Exampleof populationestimatiollwithcon-

NUSBEROF TRAPPt NG OCCA51ON5 14AS

FIG. 2.Exampleof populationestimationwithcon-

nally,we mentionthatone wouldnotex- rametersis involved(one)if one is con-

riskto captureon the jth trappingocca- algorithmnecessaryto producethe true

TABLE1. RESULTSOF COMPUTER STUDIESOF THE BL&SOF Nt (ALSOSEE APPENDIX

22.0 1,015.6 800 0.01, 0.01, 0.02, 0.03, 0.03 200 5 11

degreeto whichNt is normallydistrib- estimatestend to be prohibitivelylarge

,348.3 0.89 400 0.01 0.01 0.02 0.03 0.03 1,000 5

used in practiceover the last 40 years, OCCAS I ON

matorswith respectto departuresfrom TOTAL AN I HALS CAPTD 45

mals, regardlessof capturehistory,are

casion.Aspreviouslyemphasized, depar- FREQUENCY

turesfromthatassumption maybe dueto EACH* EQUALS 3 POINTS

imal,or the animal'sprobabilityof cap- 18

turebeing alteredafterfirstcapture,or 12 * * * &

both. Althoughboth departuresmay 9

createsignificantbiasin Nt,the natureof _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _

the bias differs.Thatis, departurefrom FIG. 3. Example of populationestimationwith

TABLE 3.-COMPUTER SIMULATIONRESULTSILLUSTRATINGTHE NONROBUSTNESSOF THE DARROCH

Percent Population Number Number of

199.3 299.3 100 100 5 Mb,Trial 8

differencebetween trappingoccasions, Mt+1 = numberof differentanimals

of the jth trappingoccasion, is satisfied.If thatcondition,whichtests

TABLE 4. RESULTS OF COMPUTERSIMULATIONSTUDIES OF THE BIAS OF Nb (ALSO SEE APPENDIX N,

Percent Average of True Probability Number Number of

-18.0 82.0 100 0.10 161 5 10

the asymptoticvarianceestimatorof Nb frequentlyin cases whereN is small,a

TABLE 5.-SIMULATED CONFIDENCEINTERVALWIDTHSAND COVERAGE

Population Probability Number Numberof

253.9 0.71 100 0.10 161 5 10

intervalsfor the parameterN. (Thisre- ulationof size 400 with200anilualsthat

ffRtATI ESTI>TE IS 112 WITH STA<AZ E 31.1

EACH * EOUALS 2 POI NTS FREQ(£M;Y IS 8 6 3 3 3

EACH * EaJ^LS 2 PO I NTS

FIG. 4. Example of populationestimationwith 6 6

el Mb with deer mouse data from E. Larsen

behavioralresponseto firstcaptureand P[{X@} I Ul,U2, * * *,Ut],