Patterns and individual-based modeling of spatial competition within two main components of neotropical mangrove ecosystems [Elektronische Ressource] / Cyril Piou

Patternsandindividual-based
modelingofspatialcompetitionwithin
twomaincomponentsofNeotropical
ecosystems.evmangro

DoCyrilctoralPiouthesis

SubmittedtotheFaculty2(Biology&Chemistry),
inpartialfulfillmentoftherequirementsforthedegreeof
DoctorofNaturalSciences(Dr.rer.nat.)

aluators:EvergerBUtaDr.PDProf.Dr.UlrichSaint-Paul

xaminators:eAdditionalDr.IlkaCandyFeller
FilserJulianeDr.Prof.

2007bruaryeF

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Martha

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iv

Preface

I2002havewith“mreally”yfieldstartedtripibnBeingelizeinforterestedmyinmasterEcologythesis.asaThere,scientIfificallintopiclovine
withecologisttheaduringmazingthatentimevironmenbtecauseofImangrostartedvets.ocButonveprtmyarticularlynatural,Ifceltauriositsayn
inintoateractingscienwithtificproandcwessithinofstheiramplingenvandironmendatat.aDuringnalysis,themdealinganyhwithoursspplanentts
inarcBelizeanhitecturesmofangrosingleves,treesIlikoredstructureimaginingoftpheossibleforestproitself.cessesIguessthatitdroisvepartlythe
tofromolsthesetryingttoimesunraasvwellelingthatthecomeshidden.myIinhavterestebforeenmvoerydelinginfluencedandstatisticalduring
thisfirstexperiencebytheownpassionofCandyforthemangroves,aswell
astheinputsofUtainimaginingprocesseswithherKiWimodel...Once
themasterthesiswasfinishedin2003,Iwasluckyandhappytohavethe
supportofUtaandProfessorSaint-PaultogetapositionofPhDstudentin
theMADAMprojecttodevelopamodeloncrabmovements.However,you
willseeinthefirstpartofthepresentthesis,thatIdidnotfocusonlyon
hadcrabstartedecologyin...BelizeMyoPhDnmbangroecameveafconoresttinuationdynamicandaddedimprotovtheemennewtsofewhatxcitingI
topicofUcidescordatus...
ThejumpintothePhDthesismighthavebeenabitfast,andthefirst
experienceoffieldworkinBrazilwasquitehard.Butwiththemoralsupport
ofMartha,myParents,CandyandUta,IthinkIwentthroughsomevery
richpersonalexperiencesandfinallyobtainedinterestingfindings...Atmy
returnfromBrazil,UtaandIproposedtohaveanewexperienceonthefield
tofewinvemontstigatehslater,morethisonttimeheamobitvebmenetttersofcprepared,rabsabasedndonwiththeamoasterbservastionstudenIt
hadoriginmofademostduringofthethedfiatarsteonxptheedition.spatialBeforedistributionthissecondofctriprabs,IthatwlearnedasaCtt++he
anddevelopedthefirstversionsoftheIBUmodel...Iwasnotconvincedat
firstsomehothatwtheIBUonewtouldhatrbringeallybimpelievortaneditnitmadditionalorethanminformation.yself,andVolkfinishederwtaso

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PREFECA

convincemeofthepotential...Theexperienceoftryingtounderstandpat-
ternswithonlyhypothesesonsomehiddenprocesseswasquitechallenging
andfinallysuccessful.ParticularlynowthatIhavedoneasecondstudywith
IBU(Chapter8),Irealizeevenmorethepotentialforfuturetheoretical,but
orks.wappliedalsoFromthedevelopmentandtryingtofitIBUtothefieldpatternswasborn
theneedofdevelopingtheapproachofPOMIC.Thiswasactuallyanother
bigexperienceandpersonaladvancement.Ihadnotmuchknowledgeon
likelihoodmethodsorinformationcriterion.However,fewweeksafterenter-
ingintothistopic,appearedtomethepossibilitythatonecouldestimate
thelikelihoodofanon-linearstochasticmodel,whichmakePOMICinterest-
ingandparalleledtothefamousAkaikeinformationcriterion.Ihavebeen
veryoftendoubtingaboutthemathematicalreliabilityofthe“newmethod”
createdwithit.Afteraskingmanyspecialists,Ibecamequiteconvincedof
itspotential.Andfinallynoneofthesespecialistsfoundthatsomethingwas
wronginthePOMICapproach...SoIjusthavetohopenowthatitwillbe
ofhelptootherindividual-basedmodelers...
InparalleloftheUcidescordatusandPOMICstudies,Ihadalsobeen
workingontheforestdynamicpartandtheKiWimodel.Iwroteandpub-
lishedmyfirstpaper(animportanteventinthelifeofanecologist)withthe
bighelpofCandyandUtaduringmyfirstyearofPhD.Itispresentedhere
becauseitfitsparticularlywellwiththeotherpapersonmangroveforest
dynamic,butthemostimportantpart(thefieldwork)wasdoneduringmy
master.Theoriginsoftheworkonsimulatingtheinfluenceofhurricanes
onmangroveforestdiversitywerealsocomingfrommymaster.However,
inthepaperpresentedhere(Chapter4),mostoftheworkwasdoneduring
thelastthreeyears.Thelastchapterofthefirstpartonmangroveforest
dynamic(Chapter5)wasthenaconnectionbetweenmynewexperienceof
tuningmodels(comingfromtheIBUmodel)andpattern-orientedmodeling
usingthePOMICapproach,sothatitcouldbeseenasparalleltothelast
chapterofthecrabpart.
Finally,theentireworkIhavedoneduring3years(exceptthedevelop-
mentofPOMIC)wasmostlyturningaroundonespecialaspect:theobserva-
tionandunderstandingofpatternshypothesizedtobebecauseof“competi-
tion”...Humansoftenbelievethatcompetitionisoneofthemostimportant
naturalforceatthebaseofanyrelationships,andparticularlyfortheirown
ones.Iwouldlikenottobelieveinthisdirection,andIhavealwaysbeen
quiteattractedtothetopic.Iwouldhavelikedtotrytodemonstratethat
competition-basedrelationshipsisnotthesolutionforhumanpopulations.I
amafraidmyecologicalworkwillneverdemonstratethat...Atleastnotat
allinthisPhDthesis.Butatlast,Icancomfortmyselfwiththeassessment

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oftheroleofcompetitiononotherpopulationsandcommunities...Ihave
orientedthecompilationofmyarticlestocoverdifferentaspectsofcompeti-
tionandtheexpressionofitundersomefewpatterns.Youwillrealizethat
Itreatedofinterandintra-specificcompetitioninthetwopartsofthethesis
respectively,andalsodiscussedwiththechapter8ofasymmetricandsym-
metriccompetition.Eveniffarfrombeingexhaustive,Icoveredinteresting
partsofcompetitionsituationinmangroveecosystems.Finally,itleadsme
toapropositionoflookingatcompetitionwithaprobabilisticapproachthat
Ipresentintheconclusionschapter.Letseehowthisideacompeteagainst
other...

nevIerwhanavtetaorrivfinishedttohistheprefacepresentwiththesisanwacithoutknowthehledgemenelpoftmpanartyp.Ieople.wouldI
willstartwithUtaBergerwhosupportedmeinallstepsandadvisedme
alwaysfrankly.IhavealsoabigdebttoCandyFellerwithhertransferof
passionforthemangrovesandconstantsupportonalltheecologicalpartsof
mywork.VolkerGrimmisthenthethird,butnotlessimportant,“mentor”
leathatstfinorthatparticular...IacmonindebtvincedtomethatProfessorIBUUlricwhorks,SainIt-Phavaul,etowhothankmadehimtheseat
forthreeaycceptingearspotossible.evaluateIwomyuldwolikerk,alsoandtoHthankeidemarieWProfessorolterandJulianneJonathanFilser
Montalvotobeonmydefensecommittee.
ThereisalotofpeoplethathelpedmeforsomestudiesandthatIwould
liketothankwarmly:KarenDieleforthefieldworkinBrazilandintroduc-
tiontoUcidescordatus,HannoHildenbrandtforhistalentonmakingC++
programmingsmarter,MarceloProtazioforhisexplanationsongeostatis-
tics,MarcTaJonathanylorandMonGtalvonzalooforOlivhisares-JohnsonmathematicalforfadvisoryruitfulforthediscussionsPOMIConmanpart,y
ecologicaltopics,CoralieD’LimaandUlfMehligforthesharedexperiencein
Brazilianmangroves,SenhorManuel,DomingosdeAraujoandAldodeMelo
forthehelpinfieldworkinBrazil,andFaustinoChiandthepeopleofthe
InstituteStern-PirlotofMalsoarinehelpedStudiesmeforimprothevwingorktheinBEelize.nglishofMarcpartsTaofylortheandthesis.Amanda
andEvIenwoifuldtheylikebetolievtehanktheymyarenparenottsosfmoruchtheirinvoclvedonstanintit,lovIeathinkndsupptheyort.do,
ToalltherestofmyfamillyandfriendsnotyetmentionedIthankthem
alsoforsupportingmeinmanyways.Andfinally,Ihavetoconfessthat
MarthaLilianaFontalvoHerazomademehappyandabletoworkonthis
waPhDyofhduringelp.allShe’llthecontthreeinueytoears.makIeammeveryhappy,gratefulallofwingormeherltoovbeaecomendvafatherrious
insomefewmonths...

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APREFEC

tsentCon

Preface

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FiguresofList

ofListblesaT

ductiontroinGeneral11.1Modelinginecology........................
1.2Competitioninecology......................
1.3Neotropicalmangroves......................
1.3.1Generalities........................
1.3.2The“trees”component..................
1.3.3The“grazers”component................
1.4Objectivesandpresentationofthethesis............
1.4.1Generalandmethodologicalobjectives.........
1.4.2Specificobjectives.....................
1.4.3Administrativepresentationofthethesis........
.............................eferencesR1.5

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125111121315151610242

2Proposinganinformationcriterionforindividualbasedmodel
92selection2.1Abstract..............................30
2.2Introduction............................31
2.3ActualtoolsofPOM.......................33
2.4AninformationcriterionforPOM................36
2.4.1Background........................36
2.4.2Proposition........................38
2.4.3Derivedstatisticsforstronginference..........39
2.4.4Derivedstatisticsforinversemodeling.........41

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CONTENTSx2.5ExampleofApplication......................41
2.5.1DescriptionofSIMOVIM.................42
2.5.2Inversemodelingphase..................44
2.5.3Stronginferencephase..................46
2.6Discussion.............................47
2.7Acknowledgements........................50
2.92.8RAppeferencesendices.........................................................5503
2.9.1AppendixA........................53
2.9.32.9.2AppAppendixendixCB................................................5566
2.9.4AppendixD........................58
IInterspecificcompetitioninCaribbeanmangrove
36forests3ZonationpatternsofBelizeanoffshoremangroveforests41
3y.e1arsAabstfterractacatastrophic..............................hurricane6656
3.2Introduction............................67
3.3Methods..............................