ReintroducingWolvesintoaWildlifeRefuge:

APredatorPreyModel

Group2:KarminaCordero,JadeFrizzell,MauroGirotto,AudreyTung

Abstract
Thereintroductionofwolvesintoecosystemsinvolvingdeer,andtheresultingpredatorprey
modelshavebeenofinterestformanyyearsandhavebeenextensivelystudiedinwildlifeparks
aroundtheworld.Thisreportproposesanequationtomodeltheinteractionofwolfanddeer
populationsoveratimespanof100years.Onemodelassumesunlimitedcarryingcapacity,and
theotherintroducescarryingcapacity.Thiswasdonebyfirstcreatingaconceptualmodel,
transitioningittoamathematicalmodelandplottingitundervariousconditions.Overall,lower
carryingcapacityandhigherinitialpopulationsincreasestability,whileanincreaseintimestep
increasesthemodelsamplitude.Asmallamountoflongtermwolfhuntingissustainable,
throughhigherratescouldleadtowolfextirpation.Thismodelishighlylimitedbythe
assumptionsitoperatesunderhowever,generalpatternscanbeseenandsomegeneral
conclusionscanbemade.

Introduction
Thehistoryofwolfreintroductionhasalonghistory,entangledwithpolitical,socialand
biologicalcontroversy(Fritts
etal.
,1997).Anexcellentexampleisthereintroductionofgray
wolvestoYellowstoneNationalParkinWyoming.Wolveswereoncenativetotheland
however,negativeviewsandapredatorcontrolprogrameventuallyledtotheireradication(Fritts
etal.
,1997).70yearslater,wolveswerereintroducedtotheparkinanattempttocontrolthe
growingelkpopulation.Overall,thereintroductionappearstohavehasapositiveeffectonthe
ecosystem,viaatritrophiccascadehowever,constantmonitoringofpopulationsofanimalsand
plantsisbeingdonetoensurebalanceismaintained(RippleandBeschta,2012).

Thisreportproposesasimplifiedmodelforthereintroductionofwolvestoawildlife
refugewheretheirmainsourceoffoodisdeer.Sincethewolvescommonlypreyonthedeera
classicpredatorpreymodelemerges.Twosuchmodelswillbecreated,withthefirstassuming
unlimiteddeerfoodsupply(carryingcapacity),andthesecondtakingintoconsiderationnatural
disastersthataffectthedeerfoodsupply.Thegoalofthesemodelsistoportraytheinteractions
ofthewolfanddeerpopulationsoveratimespanof100years,focusingonthefollowing
researchquestions:
1)Howdowolfanddeerpopulationsinteractandchangeover100yearswithinfiniteandfinite
carryingcapacity?
2)Howdoesthelengthofthetimestepaffectthemodelsandtheirstability?
3)Howsensitivearebothmodelstodifferentinitialconditionsandparameters.Aretheystable?
4)Howwouldallowingrecreationalhuntingofwolvesaffectbothpopulationsandshouldthisbe
allowed?
Inordertobegincreatingthemodels,someassumptionshadtobemade.Inreality,wolves
anddeerareapartofaverycomplexecosysteminvolvingmanyanimalandplantspecies,such

ascoyotes,eagles,bears,varioustrees,andmanymore(Smith
etal.
,2003).Thissimplified
modeloperatesunderthefollowingmainassumptions:
1)Thewolvesonlysourceoffoodisdeer,andtheyaretheonlyspeciespreyingonthedeer.
2)Therearenoanimalspreyingonthewolves.Theyonlydieofnaturalcauses.
3)Bothpopulationsarecomposedofhalfmaleandhalffemale.
4)Thebirthrateofdeerisconstantwhenthereisunlimitedcarryingcapacity

Whiletheseassumptionsdolimittheaccuracyofthemodel,thegeneralpatternsobserved
arestillimportantandcanbeusedtomakesomebasicpredictions.Theresearchquestionswill
beansweredbyfirstcreatingaconceptualmodel,thentransitioningittoamathematicalmodel
andfinallyusingthemodeltoexaminevariousconditionsandreachconclusions.

Methods

Inordertocreateourmodeltheinitialparameterswereestablishedandtheequationswere
derived.Theinitialparametersusedwere:

DeerPopulation
N
D

WolfPopulation
NW

Deerbirth/deathratepercapita
bD

/d
D

Wolfbirth/deathratepercapita
b
/d
W
W

Predatorcoefficient

Preycoefficient

Deercarryingcapacity
K
D

Timestep
t
Firstofallwebeganbydevisingaconceptualmodel(
Figure1
).Therearetwomainstocks,deer
andwolves,bothareaffectedbybirthsanddeaths.Additionally,deerandwolfbirths/deathsare
affectedbythebirthrateanddeathrateconverters,andbythetotalnumberofindividualsineach
population.Also,deeraffectsthebirthrateofwolves,andwolvesaffectthedeathrateofdeer.
Forthesecondscenario,deerbirthsarealsoaffectedbythecarryingcapacity.Wethenproceeded
totranslatetheconceptualmodelintomathematicalequations.
Bothdeerandwolfinitial
populationswereestimatedtobe500
and25respectively,basedon
researchfromYellowstoneNational
Park(Garrot,
etal.
,2005Phillips
andSmith,1997).Deerandwolf
birthrateswerecalculatedusingthe
averagelitterforeachspecies(i.e.2
deerand5wolvespereachfemale
individual).Itwasthenassumedthatbothpopulationswereevenlycomposedbymaleand

femaleindividuals,thuseachbirthratewasdividedbytwotoaccountforhalfthefemale
population.Deathratesforbothdeerandwolveswerecalculatedbytakingtheaveragelifespan
ofasingleindividual(i.e.10yearsfordeerand12forwolves).Thepredatorcoefficient,orthe
effectofwolvesonthedeathrateofdeer,wasfoundbyplottingthed
againstthenumberof
D
wolvesN
,byassumingalineartrendtheslopeofthelineorawasfound(
Appendix
).
W
Similarly,inordertoobtainthepreycoefficient(theeffectofdeeronwolfbirthrates)theb
was
W
plottedvs.thenumberofdeerN
.Again,byassumingalineartrendtheslopeofthelineorb
D
wasfound(
Appendix
).

Forthefirstcase,assumingunlimitedresourcesfordeerthefollowingequationswere
derivedforbothpopulations:

N
=N
+[(b
*N
*N
*N
)*t]
D(t+1)
D(t)
D
D(t)
D(t)
W(t)
N
=N
+[*N
*N
d
*N
)*t]
W(t+1)
W(t)
W(t)
W(t)
W
W(t)
Inthedeerequationthedeathrateisdirectlyproportionaltotheacoefficient,thusitdoesnot
appearintheequationasitisalreadyincorporatedinthepredatorcoefficient.Likewise,thebirth
rateofwolvesisdirectlyproportionaltothebcoefficient,assuchitalsodoesnotappearbecause
itisalreadyincludedinthepreycoefficient.Inbothdeerandwolfequationsthebirthdeath
factorwasmultipliedbythetimestep(t=0.1yr).Subsequently,bothequationswereenteredina
excelspreadsheetandthemodelwastestedfor100years(
Figure1
).Forthisscenariosensitivity
testswerealsocarriedout,initiallybyvaryingtheinitialpopulationvalues,thenbyvaryingthe
timestep(
Figure3
).Furthermore,toproveifthemodelwaseffectivelystableornotweplotted
bothpopulationvaluesagainsteachother(
Figure

2
).

Forthesecondscenario,whichconsistedinaddingacarryingcapacityfordeer,the
followingequationswerederived:
N
=N
+{[b
*(1N
/K
)*N
*N
*N
]*t}
D(t+1)
D(t)
D
D(t)
D
D(t)
D(t)
W(t)
N
=N
+[*N
*N
d
*N
)*t]
W(t+1)
W(t)
W(t)
W(t)
W
W(t)
Thecarryingcapacityterm(1N
/K
)hasnowbeenintroducedinthedeerequation.TheK
D(t)
D
D
influencesdeerbirthrate,thusthetermappearsonthebirthsideoftheequation.Thewolf
equationremainsunchangedinthisscenario.Withanassumedcarryingcapacityof1000deerwe
proceededtorunthemodelfor100yrinexcelwiththenewequations,allotherparameters
remainingthesameasinthebasescenario(
Figure4
).Analogouslytocase1,weransensitivity
testsfirstbyvaryingtheinitialdeerandwolfpopulationvalues,nextbyvaryingthecarrying
capacity
(
Table1
).Lastly,asforscenario1,weplottedthepopulationvaluesobtainedfromthe
modelvs.eachother(
Figures5,7,9
).

Additionally,wealsomodeledanewscenarioinwhichweintroducedwolfhuntingtothe
scenariowithadeercarryingcapacityof1000,becausethisscenariowasonethatresembled

realisticnaturalconditions.Thiswasdonebysubtracting0.2wolvespereachtimestep,resulting
in2wolveshuntedperyear.Huntingvalueswerealsovariedinordertoestablishthemodel
stabilityandtheappropriatenumberofwolvestobekilledbyhunting
(
Figure
8).Againastate
diagramwasalsomade(
Figure9).

Results
Westartedouranalysisbyusingexceltomodelourmathematicalequationforbothcase
1andcase2.
Figure1
showstheinteractionsbetweenthedeerandthewolfpopulationbysetting
thedeerpopulationto500andintroducing25wolvesinoursystem.Wedecidedtoshowthe
interactionsinatimeperiodof100years.Wecanseethatthetwopopulationshaverepeated
cyclesbetweenincreasingtoamaximumandgraduallydecreasingtoaminimumvalue.
Moreover,thewolfpopulationgraduallyfollowsthesamepatternofthedeerpopulationthrough
time.Accordingto
Figure2
,thepopulationwithunlimitedcarryingcapacityisspiraling
outwardswhichcouldleadtoextinctionofonepopulationatsomepoint.

Case1modelmakes
sensebecausethedeerpopulationhasunlimitedfoodsupplyanditgrowstoanunreasonable
populationsize.Ontheotherhand,themodelisunrealisticbecauseoftheextremeincreasesand
decreasesinpopulationwithin100years.In
Figure1
,thedeerandwolfpopulationsfluctuateto
alargeextent,thusindicatinginstability.

Figure2.
Thisgraphrepresentsa
statediagram,indicatingthatinfinitecarrying
capacityleadstoinstability.

Figure1
.
Thegraphshowsourmathematicalmodelwhereweassumed
deerpopulationis500andwolfpopulationis25.

In
Figure3
,thetimestepparameterischangedfrom0.1to1.Anincreaseintimestepchange
havecauseanincreaseinamplitudeforbothpopulations.

Figure3.
Thisgraphrepresentstheinitial
modelwithadifferenttimestep,from0.1yrto
1yr.

Forcase2,wehavesetupourmodelwithacarryingcapacityof1000deers.Wecanseein
Figure4
,thewolfanddeerpopulationgraduallystabilizesthroughtime.Thetotalpopulationof
thedeerhasloweredinnumberstoapproximately210deersandthewolfpopulationfollowsa
similartrendbysteadilydecreasingtoapproximately50wolves.Theamplitudebeginsto
decreasewhenthecarrycapacitycomponentisincludedinourequationforcase2.In
Figure5
,
theadditionofcarryingcapacitymakesourmodelstable.Thespiralsareheadinginwardswhich
correspondstoastabilizingpopulation.Thecarryingcapacityvalueischangedtoobservethe
changestoourmodel.Wehavefoundthatsmallercarryingcapacityvalueshaveamorerapid
processofstabilization.Wehavedecreasedourcarryingcapacityfrom1000to300andwehave
foundthatthepopulationstabilizesmuchfaster.Accordingto
Figure6
,thewolfanddeer
populationbeginstostabilizestartingfrom30yearswithalowerK.Wethenlookedinto
graphingoutthestabilitieswhenweloweredthecarrycapacity.Aswecansee,withlower
carryingcapacitythepopulationsplateauatsteadylevelsrapidlyin
Figure6
andthespiralin
Figure7
hastheleastamountofcoils,indicatingfurtherstability.Insummary,wehave
organized
Table1
toillustratethedifferencesbetweenthemaximumandminimumpopulation
whenwedecreasethecarryingcapacity.Asaresult,weseethatthereislessfluctuationandthe
twopopulationsstabilizeatlowercarryingcapacities.See
Table1
toseethestabilityofthe
populationwhenthecarryingcapacityis1000.

Figure5
.
Thisgraphrepresentsastatediagram,
indicatingthatbothpopulationsstabilizewhen
carryingcapacityisadded.

Figure4.

Thisgraphrepresentstheinitialmodelwiththeadditionofcarryingcapacityfordeer.

Figure7.
Thisstatediagramindicatesthatboth
populationsstabilizerapidlywhenKisreduced
to300

Figure6.

Thisgraphrepresentsthemodelwithanewdeercarryingcapacity,loweredto300.
Bothpopulationsshowfasterstabilizationthroughtime.

Column1

Carrying Capacity

Maximum

Minimum

Maximum - Minimum

Deer

infinity

8849

119

8730

Wolf

N/A

160

17

143

Deer

1000

511

136

375

Wolf

N/A

73

25

48

Deer

300

500

164

336

Wolf

N/A

33

19

13

Table1.

Thissummarytablerepresentsdeerandwolfpopulationvaluesobtainedbyvaryinginitialpopulationvalues
andcarryingcapacity.

Wehaveincorporatedthehuntingcomponentintoourmodelbysettingourwolveshunting
parameterto2wolvesbeinghuntedperyear.Wehavefoundthathunting2wolvescanstabilize
theamplitudeofthewolfanddeerpopulationthroughtime.Anyvaluesabove2resultedin
instability,withthedeerpopulationincreasingexponentiallyasthewolvesweregradually
broughttoextinction.

Figure9.
Thisstatediagramshowsthatboth
populationseventuallystabilizeevenwiththe
introductionofhunting.

Figure8.

Thisgraphrepresentsthecase2modelwiththeadditionofhunting(2wolves/yr)whenK=1000.

DiscussionandConclusion

Ourresultsindicatethatalloftheparameterswetested(carryingcapacity,timestep,
initialpopulations,andhuntingscenarios)inducedchangesinpopulationdynamics.Firstly,we
foundthatcarryingcapacitywasasignificantdeterminantofpopulationstability:accordingto
ourdata,boththemagnitudeoffluctuationsandthetimerequiredforpopulationstabilization
werelowestatsmallercarryingcapacities,whileanunlimitedcarryingcapacitycreatedlarge
fluctuationsandanindefinitelyunstablepopulationtrajectorythatappearsboundforthe
extinctionofeitherpopulation.Thisphenomenonislikelyduetothefactthatunlimitedorhigh
carryingcapacitiesenabledeerpopulationexplosionsasdeerpopulationgrowthisnotimpeded
bylimitedresources,whichthencauserapidincreasesinthewolfpopulation.However,such
growthisunsustainablesincethepredationpressurefromwolveseventuallydepletesdeer
resourcesandaccordinglyresultsinacrashinwolfpopulationaswell.Thesestaggered
synchroniesareevidentin
Figures1,3,and4
.Asillustratedin
Figures1and2
,these
fluctuationsincreaseinamplitudeuntilinevitableextinctionforunstablesystems.Instable
systems,ontheotherhand,theamplitudesdecreaseuntilsteadypopulationsareeventually
reached(
Figures4,5,6,and7
).Becauseofthecorrelationbetweenhighcarryingcapacityand
instability,wesuggestcarefullymanageddeerhuntingasameansforlimitingdeerpopulations
underhighcarryingcapacities,therebypreventingdangerouspopulationexplosions.
Furthermore,overpopulationofdeercouldinduceotherecologicalriskssuchasirreversible

changesinplantcommunitiesduetoovergrazing,anddecreasesinecosystembiodiversity,thus
makingdeerhuntingaparticularlysalientsolution(Levy2006).Next,althoughlowercarrying
capacitiesreducethemaximumpopulationsreached,theyinfactincreaseminimumpopulations
fordeerinallscenariosandforwolveswhenKisreducedfromanunlimitedcarryingcapacityto
1000(
Table1
).Thissuggeststhatthesupplyofanimalsavailableforhuntingismorereliableat
lowercarryingcapacities,andthatthehuntingshouldonlycoincidewithpopulationpeaksin
highcarryingcapacityenvironmentstopreventextirpation.

Thedegreeofpopulationfluctuationsalsovariedinresponsetoournextparameter,time
step,withincreasedfluctuationmagnitudesandthus,higherinstabilityoccurringathigher
timestepsettings.Thissuggeststhatourmodelishighlysensitivetotimestep,whichcould
influencetheprecisionofourdata.

Intermsofinitialpopulations,ourdatashowedthatlowinitialpopulationswere
associatedwithanincreasedfluctuationmagnitudeforbothwolfanddeerpopulationswith
highermaximumandlowerminimumvalues,thussuggestingthatsmallerecosystemsaremore
vulnerabletoinstability.Inthisregard,preventinghabitatfragmentationcouldbeanimportant
measureformaintainingecosystemviability.Wealsorecommendthathuntingshouldbe
restrictedtolargerecosystemareas,sincetheincreasedpressurefromhuntingcouldleadto
extirpationifitiscarriedoutduringperiodsofpopulationtroughs.

Finally,afterapplyinghuntingefforttowolvestoourmodelwithadeercarryingcapacity
of1000,wewereabletodetermineasustainablehuntingrateof2wolvesperyear.Althoughthis
huntingratealteredthepopulationstructurebyincreasingtheamountofdeeranddecreasingthe
amountofwolvesinthelongterm,itislikelysafebecausepopulationseventuallystabilizeat
thesenewlevels,ratherthanresultingintheextirpationofthewolvesorcausingdeerpopulation
explosions.However,wedonotrecommendhuntingatlevelsabove2wolvesperyear,since
ourmodelpredictedtheextirpationofwolvesathigherrates.Therefore,issuinghuntingpermits
andimplementingrobustmonitoringandenforcementsystemstopreventpoachingwillbe
essentialforresponsiblyhuntingthesesensitivewolfpopulations.Althoughthehuntingmaybe
sustainableintermsofdeerandwolfpopulations,theexpectedincreaseinthedeerpopulation
couldhavenegativeenvironmentaleffectsontheecosystematlarge(i.e.vegetationand
biodiversityloss).Thus,wesuggestthatapreferablesituationwouldbeacompletebanonwolf
hunting.

Overall,ourresultswerelikelyunrealisticduetothenumerousassumptionsand
limitationsassociatedwiththemodel.Mostnotably,ourassumptionthatourcomponentsofthe
ecosystemarenegligiblemakesourmodeloverlysimplistic.Inreality,bothdeerandwolves
birth/deathratesvaryseasonally,dependingonenvironmentalfactorsandcomplexinteraction

withotherbioticandabioticecosystemcomponents.Wedidnotconsiderthefoodwebina
broaderecologicalcontext,sinceweassumedthatdeeraretheonlyfoodsourceofwolvesand
wolvesaretheonlypredatorsofprey.However,otherpredators,suchascoyotesandmountain
lions,areknowntopreyondeer,andwolvesarealsoknowntopreyonotherspecies(Ballard
et
al.
2001).Wedidnottakeintoaccountcompetitionwithotherspecies(e.g.competitionbetween
deerandotherforagingspecies),whichisanimportantcomponentofpopulationdynamicsin
scenarioswithlimitedresources.Thus,ourmodelshouldbeconsideredasabasictemplatefor
morecomprehensivefurtherresearch,ratherthanarealisticrepresentationofpopulation
dynamics.Despiteourstudyslimitations,thesimplisticnatureofourmodelmadeiteasytouse
sincewedidnothavetosetalargeamountofparameters.

Appendix

References

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ContributionsofGroupMembers

KarminaCordero:
Completedtheresultssectionforboththereportandthepresentation.
Organizedandlabelledallthefiguresinthereportincludingthecaptions.Contributedtofinal
editingofthereport.
JadeFrizzell:
ConceptualModel:Workingtitle,researchquestions
Presentation:Title,background,researchquestionsandassumptionsslides,helpedwithoverall
ideasandorganization
FinalReport:Wroteabstractandintroduction.Helpedwithoveralleditingofwritingandgraphs,
helpedwithoverallideasandorganization
MauroGirotto:
Completedthemethodssectionforboththepresentationandthereport.
Modeledtheequationsandgeneratedthegraphsfromexcel.Helpedwiththeformattingand
editingofthereport.
AudreyTung
:discussionandconclusionforboththepresentationandreport,createdthetable
andstatediagrams,editingofreport