universidade de aveiro · projeto europeu do fp7: “recare – preventing and remediating...
TRANSCRIPT
Universidade de Aveiro
2016
Departamento de Ambiente e
Ordenamento
Tiago van der Worp da Silva
The short-term effects of mulching on the soil-dwelling arthropod community after a wildfire Os efeitos no curto prazo da aplicação de mulch sobre a comunidade de artrópodes que vivem no solo após um incêndio florestal
Universidade de Aveiro 2016
Departamento de Ambiente e Ordenamento
Tiago van der Worp da Silva
The short-term effects of mulching on the soil-dwelling arthropod community after a wildfire Os efeitos no curto prazo da aplicação de mulch sobre a comunidade de artrópodes que vivem no solo após um incêndio florestal Dissertação apresentada à Universidade de Aveiro para cumprimento dos requisitos necessários à obtenção do grau de Mestre em Engenharia do Ambiente, realizada sob a orientação científica da Doutora Filomena Maria Cardoso Pedrosa Ferreira, Professora Associada do Departamento de Ambiente e Ordenamento da Universidade de Aveiro e coorientação do Doutor Nelson Abrantes, investigador auxiliar do Centro de Estudos do Ambiente e do Mar (CESAM), Departamento de Ambiente e Ordenamento da Universidade de Aveiro, e Doutor Jan Jacob Keizer, Investigador Auxiliar do CESAM e Departamento de Ambiente e Ordenamento da Universidade de Aveiro.
Esta dissertação foi desenvolvida no âmbito do projeto europeu do FP7: “RECARE – Preventing and Remediating degradation of soils in Europe through Land Care” (no. 603498)
o júri
presidente Professora Doutora Maria Isabel Aparício Paulo Fernandes Capela
Professora associada do Departamento de Ambiente e Ordenamento
Professor Doutor Fernando José Mendes Gonçalves
Professor associado com agregação do Departamento de Biologia da Universidade de Aveiro
Professora Doutora Filomena Maria Cardoso Pedrosa Ferreira Martins
Professora associada do Departamento de Ambiente e Ordenamento da Universidade de Aveiro
agradecimentos
À orientação muito próxima e muito enriquecedora facultada pela Prof. Doutora Filomena Martins, pelo Doutor Nelson Abrantes e Doutor Jan Jacob Keizer. Ao Doutor Jan Jacob Keizer pela oportunidade dada em trabalhar e colaborar no seio de um grupo de investigação. Em especial ao João Puga por toda a disponibilidade, crítica, conselhos, constante apoio e partilha de conhecimento. À minha família por me proporcionar sempre o melhor e pelo apoio incondicional neste percurso. À Mariana pela presença sempre muito especial. A todas as amizades e aprendizagens que nutri durante este percurso e que levo para a vida.
keywords
mulching effects, wildfires, ground-dwelling arthropods
abstract
Forest fires are common events in the Mediterranean basin, but due to human action and climate change, these have been increasing, especially in Portugal. It is in this context that a larger number of studies have emerged in regard to the impacts of forest fires on ecosystems, with special focus on the effects on soil and mitigation strategies in the post-fire period. In this regard, the application of mulch is one of the mitigation strategies studied since it is considered inexpensive and effective in reducing erosion and loss of soil nutrients in the post-fire. Moreover, the impacts of organic coverages application on soil fauna are still poorly understood. This study aims to assess the effect of the application of various mulch densities in the soil on the invertebrate community in a eucalyptus plantation, immediately after the fire. The ground fauna covered in this study were soil arthropods, a large existing community regarded as reliable bio-indicators. The type of mulch used was chopped eucalyptus bark. The study area, located in North-Central region of Portugal, is characterized by eucalyptus plantations in a schist soil. This study was conducted in order to understand the extent of the effects that mulch could produce in a short period after a fire. In order to achieve adequate results, different areas with and without the application of mulch, at different densities were selected and monitored for 5 months after the fire. Sampling was carried out with pitfall traps and identification of the collected specimens was made at Order level. The results suggest differences between the monitoring months and absence of differences among the densities of mulch application, suggesting seasonality as influence factor on the total community abundance. Small differences in the community richness suggested that a low density of mulch provided a faster ecological succession. The application of mulch also raised the hypothesis that it may have introduced unwanted specimens into the ecosystem.
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palavras-chave
efeitos da aplicação de uma camada orgânica, incêndios florestais, artrópodes que vivem no solo
resumo
Os incêndios florestais são eventos comuns na bacia mediterrânica, mas devido à ação do homem e às alterações climáticas, estes têm vindo a aumentar em Portugal. É neste contexto que um maior número de estudos tem surgido, focando-se nos impactes dos incêndios florestais sobre os ecossistemas, e em especial sobre os seus efeitos no solo e em estratégias de mitigação no período pós-incêndio. Neste sentido, a aplicação de coberturas orgânicas sobre o solo é uma das estratégias de mitigação estudada, sendo considerada barata e eficaz na redução da erosão e perda de nutrientes no pós-incêndio. Por outro lado, o conhecimento dos impactes da aplicação de uma cobertura orgânica sobre a fauna presente no solo é ainda escasso. Este estudo pretende avaliar o efeito da aplicação de várias densidades de cobertura orgânica no solo sobre a comunidade de invertebrados do solo em eucaliptais, imediatamente após o incêndio, utilizando como cobertura orgânica raspas de casca de eucalipto. Os invertebrados do solo, formam uma comunidade muito diversa considerada como um bioindicador fiável. A área de estudo, situada na região Centro-Norte de Portugal, é caracterizada por plantações de eucalipto sobre um solo xistoso. A amostragem foi realizada com armadilhas de solo e a identificação dos espécimenes recolhidos foi feita até ao nível da Ordem. Os resultados obtidos sugerem diferenças entre os meses de monitorização e ausência de diferenças entre as densidades de aplicação e sem cobertura orgânica, sugerindo a sazonalidade como um fator de influência sobre a abundância total da comunidade. Pequenas diferenças na riqueza específica da comunidade sugeriram que uma baixa densidade de cobertura orgânica promoveu uma sucessão ecológica mais célere. A aplicação da cobertura orgânica levanta ainda a hipótese de esta ser uma fonte de introdução de espécies indesejadas no ecossistema.
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Tableofcontents
Acknowledgements
Abstract
ListofFigures
ListofTables
1.Introduction...................................................................................................................................1
1.1WildfiresandthePortuguesecontext..............................................................................1
1.2Wildfireimpactsandstudies............................................................................................2
1.3Ground-dwellingarthropods............................................................................................3
1.4Researchobjectives..........................................................................................................5
2.Materialsandmethods..................................................................................................................7
2.1 StudyareaandSites..........................................................................................................7
2.1.1Samplingmethodologyanddesign..........................................................................8
2.1.1.1Pitfalltraps............................................................................................................8
2.1.1.2Samplingdesign....................................................................................................9
2.2 Ground-dwellingarthropodscollectionandidentification.............................................11
2.3 Ecologicalfunction..........................................................................................................12
2.4 Environmentalvariables.................................................................................................13
2.5 Dataanalysis...................................................................................................................13
3.Results...........................................................................................................................................15
3.1 Environmentalvariablesanalysis....................................................................................15
3.2 Communityanalysis........................................................................................................15
3.2.1RichnessandAbundance........................................................................................15
3.2.1AbundanceandEnvironmentalvariables...............................................................23
3.2.2Diversity(H’)andEvenness(J’)...............................................................................23
ix
3.3 Ecologicalfunctionanalysis............................................................................................24
4.Discussion.....................................................................................................................................29
5.Conclusion....................................................................................................................................37
6.Finalconsiderations.....................................................................................................................39
References........................................................................................................................................41
x
Listoffigures
Figure1.MeanburntareaperyearontheMediterraneanclimatecountries(PORDATA,2016).....1
Figure2.Locationofthestudyfields,Burnt(black)andUnburnt(green)........................................7
Figure3.Pictureofanuncoveredpitfalltrapintheburntsite..........................................................8
Figure4.Samplingdesigndisplaypersamplingmonths.................................................................10
Figure7.Stereoscopicmagnifier......................................................................................................11
Figure5.Secondscreening,identifyingandcountingarthropodsatorderlevelperpitfalltrap....11
Figure6.Firstscreeningprocess,separatingtrappedarthropodsfromlitter.................................11
Figure8.Meanandstandarddeviationfortemperatureandprecipitationbetweenlate
September2015andJanuary2016..................................................................................................15
Figure9.Totalrichnessregardingsamplingmonthsandplots........................................................16
Figure10.Totalabundanceregardingsamplingmonthsandplots.................................................18
Figure11.Meanabundanceandrespectivestandarddeviationpersamplingmonthsandplots..18
Figure12.Relativeabundanceinpercentageperplotduringthesamplingmonths......................20
Figure13.TotalabundanceperorderinSeptember.......................................................................21
Figure14.TotalAbundanceperplotduringsamplingmonths........................................................22
Figure15.TotaldiversityandtotalevennessintheLT,HTandNTplotsduringsamplingmonths23
Figure16.EcologicalfunctiondistributioninSeptember................................................................24
Figure17.Ecologicalfunctionrelativeabundanceperplotduringsamplingmonths.....................25
Figure18.Ecologicalfunctiongroupstotalabundanceperplotduringsamplingmonths..............26
xi
Listoftables
Table1.Two-wayANOVAsummaryregardingmonthandplotwithtotalrichnesspertrap.
Significantstatisticalrelationsinboldtype......................................................................................17
Table2.AllPairwiseTukeymultiplecomparisontestwithinmonthsandtreatments....................17
Table3.Two-wayANOVAsummaryregardingmonthandplotwithtotalabundancepertrap.
Significantstatisticalrelationsinboldtype......................................................................................19
Table4.Pearsoncorrelationbetweenordersoneachsamplingmonth.........................................20
Table5.Pearsoncorrelationbetweentotalabundanceandabioticvariables................................23
Table6.Two-wayANOVAsummaryregardingmonthandtreatmentasfactorsanddiversityand
evennessindexeshasvariables.Significantstatisticalrelationsinboldtype..................................24
Table7.Tukey’smultiplecomparisonofmonthsregardingdiversity.............................................24
Table8.Pearsoncorrelationbetweenecologicalfunctiongroupsoneachsamplingmonth.........25
Table9.PearsoncorrelationbetweenecologicalfunctiongroupsoneachPlot.............................25
Table10.One-WayANOVAonfunctionalgroupsregardingsamplingmonths...............................27
Table11.Tukeymulti-comparisontestonmicrobialfeedersandomnivoresregardingdifferent
months.............................................................................................................................................27
Table12.One-WayANOVAonfunctionalgroupsregardingtreatments........................................27
The short-term effects ofmulching on the soil-dwelling arthropod community after awildfire
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1. Introduction
1.1 WildfiresandthePortuguesecontext
Awildfire isanevent that takesplace in forestsetting influencedbyexisting fuels,climate-
weatherrelations,ignitionagentsandanthropologicactivities(Flannigan,Stocks,Turetsky&
Martínez-Mena, 2009), that ultimately produces effects on soil, hydrological regimes,
vegetation and fauna (Moreira, Catry, Silva & Rego, 2010). Wildfires are seen as major
ecologicaldisturbances,althoughthesehavealsobeenoneimportantenvironmentalchange
factor shaping the landscape in fire-prone ecosystems for millenniums, such has the
Mediterraneanbasin,California,SouthAfricaandSouthwestAustraliaregions(Pausas,Llovet,
Rodrigo&Vallego,2008).
Consideringthispremise,itisbelievedthattheglobalclimatechangescenariowillbeplaying
asignificantroleinincreasingchangesforfireperiodsandintensityregimesinthenearfuture
(Lavorel,Canadell,Rambal&Terradas,1998;Bento-Gonçalves,Vieira,Úbeda&Martin,2012).
Largertemperaturegapsbetweenseasonsaswellaslongerdryperiodsandlessprecipitation
willinfluencedirectlyfireseverityandoccurrence(Flanniganetal.,2013).
PortugalhasnotbeenanexceptionintheMediterraneanBasinareasinceitistheEuropean
countrywith the largest recordofwildfires andburnt area in the lastdecade (IPCC,2013)
(Figure1).Moreover,thisscenarioislikelytocontinueinthenearfuture(Bento-Gonçalveset
al.,2011).
Figure1.MeanburntareaperyearontheMediterraneanclimatecountries(PORDATA,2016)
Theserecordsareassociatednotonlywith theclimateregimecharacterisedbywetwinter
seasonsincontrastwithdryandhotsummerseasons,butalsoduetosocio-economicfactors
50 100 150 200 250 300
1980- 1984
1985- 1989
1990- 1994
1995- 1999
2000- 2004
2005- 2009
2010- 2014
MeanburntareainthousandsofhaPortugal Italy Greece France Spain
The short-term effects ofmulching on the soil-dwelling arthropod community after awildfire
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(ICNF, 2013). These factors include: the extensive and intensive plantations of Eucalypt
registeredinPortugalthatstartedinthe80’s;theruralexodusthathasledtotheabandonment
of forest areas leaving them more exposed to this hazard; and the lack of efficient land
managementpoliciesthatwouldpreventit(Shakesby,2011;Shakesby&Doerr,2006;Keeley
etal.,2012).
1.2 Wildfireimpactsandstudies
Awildfireleadstomanyeffectsontheecosystemeitherdirectlyorindirectly,bychangingsoil
structurepropertiesthatlaterleadtochangesinhydrologicalandgeomorphologicalprocesses
andconsequentlychangesinthefaunaandfloraofanecosystem(Certini,2005).
Most of the recent studies undertaken in theMediterranean region have been focused on
physicalandchemicaleffectsonsoilduringthepost-fireseason(Neary,Ryan&DeBano,2005;
Keizeretal.,2008,Shakesby,2011)inordertocomprehendhowbettercopingstrategiescan
beimplementedinthepreandpostfireperiods.
Mostofthemitigationstrategiesarefocusedonreducingsoilerosionbywaterrunoffinthe
burntareas,suchas:seeding,constructionofphysicalbarriersalongtheslope,scarifyingand
ploughing,applicationofhydro-mulchandmulching.Thesetechniquesareappliedaccording
tothephysicalcharacteristicsoftheburntfield,suchastheslopeand/oravailableresources
suchasrockandlogs.Intheoverall,themosteffective,easytoapplyandlowcosttechniqueis
mulching(Ferreiraetal.,2015).
Mulchingconsistsoncoveringthebaresoilsurfacewithabiomasslayerinorderthatinsmall
areas,micro-trapsaregenerated,asthesewillretainashesandstoresomeoftherunoffwater
intheperiodafterthefire.Thistechniqueconductsaverysignificantreductionofwaterrunoff
by50%thatconsequentlyproducesareductionofsoilerosionby90%(Ferreiraetal.,2015).
Thissametechniquehasalsobeenstudiedandappliedintheagriculturalcontext,providing
anincreaseinsoilmicrobialactivity,improvingnutrientbalance,moderatingsoiltemperature
andimprovingwaterinfiltrationandreducingsoilerosion(Gill,McSorley&Branham,2011;
Chalker-Scott,2007;Westerman&Bicudo,2005;Altieri&Nichols,2003).Theseadvantages
havealsobeenobservedinthepost-fireapplicationcontext(Ferreiraetal.,2015).
Mulching also occurs naturally in the ecosystem, especially in settings that had mid-low
severitywildfire events, onwhich leaves remain on trees and later fall in the soil surface,
The short-term effects ofmulching on the soil-dwelling arthropod community after awildfire
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working in the same way has the mulch application effects (Ferreira et al., 2015; Prats,
McDonald,Monteiro,Ferreira&Keizer,2012).Nonetheless,mulchapplicationdependsonthe
localavailabilityandpriceofthematerial(biomass)inordertobeafeasiblesolution.Many
materials have been proven to work effectively like straws, woodchips and hydro-mulch,
althoughforestresidualshavenotbeensubjectedtothesamescrutiny(Ferreiraetal.,2015).
In thatregard,studies inPortugalhave looked into forestresidueshasamulchingmaterial
consideringitsavailability,costandalsothedensityofitsapplication(Pratsetal.,2012).
Followingupthestudiesthatfocusontheeffectivenessofmulchinginthereductionofsoilloss
inburntareas,awarenesshasrisentoothereffectsthatthistechniquemightproduce,specially
to the soil-dwelling arthropod communities (Puga, 2016), which are widely affected by
wildfires(Elia,Lafortezza,Tarasco,Colangelo&Sanesi,2012).
1.3 Ground-dwellingarthropods
ThesoilisoneofthemostcomplexsystemsinEarth,althoughitsbiologicalsystemsareyetto
befullyexplored.It’sthelivingsurfaceforatleastoneormorestagesofalifecycleformany
animals,meaningthatmost,ifnotall,terrestrialorganismsdependdirectlyandindirectlyon
biologicalprocessesinthesoil(Storketal.,1992).
TheArthropodaphylumroughlyaccountsforthreequartersoftheEarth’sorganismshaving
themostdiversetaxaofinvertebratesandbeingpresentinbothwaterandlandecosystems.
Arthropods are characterized by segmented bodies, with bilateral symmetry and chitin
exoskeletons(McGavin,2010).
Theground-dwellingarthropodsaretheoneswhoinhabittheleaflitterinthesoilandrange
inmanydifferentformsandfunctionswhileactinghasdecomposers,playingaveryimportant
role onnutrient cycling and soil structuremaintenance and fertility; as prey andpredator,
regulatingpopulationsofotherorganismsandenhancingthefood-web;andalsoinenabling
floweringplantreproduction(García-Domíngez,Arevalo&Calvo,2010;Buddleetal.,2005;
Danks,1992;Kremenetal.,1993andWiggins,Marshall&Downes,1991).
Inanecologicalcommunity,theground-dwellingarthropodscanbeextremelysusceptibleto
shocksthatmayaltertheirlivingenvironmentsandthereforebeingregardedasbio-indicators
foraccessingecologicalchangesduetotheirhabitatrequirementsandpotentialsensitivityto
The short-term effects ofmulching on the soil-dwelling arthropod community after awildfire
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changes and/or threats. Hence, arthropods have beenwidely studied as bio-indicators for
pollution,habitatdisturbanceandclimatechange(McGeoch,1998).
The threats that affect the integrity and resilience of this group in the Mediterranean
ecosystemsvariedfromhabitatfragmentation,humanexploitation,pestoutbreaks,invasion
ofexoticspecies,overgrazingandforestfires(Eliaetal.,2012;Penman,Beukers,Kavanagh&
Doherty, 2011). Forest fires conduct severe impacts on ecological communities directly,
throughmortalityinducedbyheatandsmoke(Gerson&Kelsey,1997),andindirectbyaltering
vegetation structure in the post-fire period (Whelan, 1995; Pickering, 1997).However, the
extentofseverityfromtheseimpactsisalsodependentonthefireseverity(Certini,2005).
Iffact,fewstudieshavebeenconductedtoassesstheresponseofthisgroupinthepost-fire
period. Nunes et al. (2006) found a decreasing tendency in ground beetle abundance in a
Mediterranean Pine forest, proving that the fire does alter the ecological community
composition.Although theshort-termresponseof insect communities to fire remains tobe
fullyexploredinMediterraneanforestecosystems,especiallywhenpost-fireeffectmitigation
techniquescometoaction.
Another study conducted by Puga (2016) focused on the effects of mulching on ground-
dwellingarthropods,butona laterstageof theecosystemrecovery,approximately5years
afterthefireevent.Inthisstudyitwasobservedahomogenouscommunityineithermulched
anduntreatedplots,suggestingthatatanearlierstageafterawildfire,thepresenceofmulch
can be of bigger influence on the re-establishment of the ground-dwelling arthropods, by
possiblypromotingafasterrecoveryonanearlystage.
Hence,thisstudyfollowsupthestudyundertookbyPuga(2016),byfocusingontheshort-
term impactofmulchingon there-establishmentofground-dwellingarthropods inaburnt
eucalyptplantationarea.
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1.4 Researchobjectives
Themainquestionsaddressedbythisstudywere:
1. How does the ground-dwelling arthropod community responds to fire-inducedecosystemalterationsinthefirstmonthafterthewild-fireevent:1.1. Whicharethemaintaxapresentduringthisperiod?
2. Howdotheabundanceanddiversityoftheground-dwellingarthropodcommunitychangewithtimesincethewildfireinthepresenceofmulchasopposedtowithoutit?2.1. Ismulchpromotingafasterrecovery?2.2. Ismulchpromotingspecifictaxaorecologicalfunctionlevels?2.3. How does a high and low mulch application density impact the re-
establishmentoftheground-dwellingarthropodcommunity?
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2. Materialsandmethods
2.1 StudyareaandSites
ThisstudywasconductedintwositesinthelocationofSemide,MunicipalityofMirandado
Corvo:oneburnt(40°09'57.8"N8°19'29.7"W)andoneunburnt(40°09'49.9"N8°19'44.7"W).
Theareasarelocatedinnorth-centralPortugal.
Figure2:Locationofthestudyfields,Burnt(black)andUnburnt(green).
BetweenAugust9thand10th,thisareawasfustigatedbyawildfirethatburntapproximately
715ha(Figure2).Thefireseveritywasrecordedbetweenmoderateandhigh,accordingtothe
soilseverityindexofVegaetal.(2013).
Regardingwildfirerecords,thestudysiteshadtwowildfiresbefore,in2000and2012.
TheareaischaracterizedbysteephillsthatconnectwiththeCeirariver.Thesoilsderivefrom
pre-OrdovicianschistsoftheHespericMassif(Pereira&FitzPatrick,1995)andareclassified
accordingtotheIUSS(2014)asacidicloamyEpilepticumbrisol.Soilsaretextureequilibrated,
pHrangesbetween4.6and4.8andarerichinsoilorganicmatter,approximately15-18%into
Alayer.Theclimateofthestudyareacanbecharacterizedashumidmeso-thermal(Köppen,
The short-term effects ofmulching on the soil-dwelling arthropod community after awildfire
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Csb),withprolongeddryandwarmsummers.Meanannualtemperatureatthenearestclimate
station(CARAPINHAL(13H/09UG),SNIRH,2012) is12°C.Meanannualprecipitationat the
nearestrainfallstation(12km)is,onaverage,851mm(CARAPINHAL(13H/09UG),SNIRH,
2016).TheareasaredominatedbyEucalyptusglobulusLabillplantations.
Both sites are eucalypt plantations (burnt and unburnt) that present very similar
characteristics,distancing500metersawayfromeachother.Theburntandunburntsitesface
E-NEandEandhaveahillslopedegreeof27°and26°,respectively.Thesitesweresubjected
tothesametypeofsamplingofground-dwellingarthropods:thepitfalltrapmethod.
2.1.1 Samplingmethodologyanddesign
Immediatelyafterthefire,afieldsettingwascarried-outintheselectedburntslopetoassess
theeffectsofforestresiduemulchonrunoffandsoilerosion.Inthisfieldsetting,areastoassess
theeffectsof forestresiduemulchon theground-dwellingarthropodcommunitywerealso
installed.
2.1.1.1 Pitfalltraps
Pitfalltrappingisthemostcommonlyusedcollectionmethod
forsamplingsoilinvertebrates.Itisanefficientandlow-cost
technique to assess ground-dwelling fauna, despite its
disadvantages. It consists, basically, on a cylindrical shaped
container(inthiscasehalfofaplasticbottle)whichisdugin
thegroundwitharimatsurfacelevelthatultimatelywilltrap
mobileanimalsthatfallintoit.Mostofthetimes,itisslightly
coveredbyrockorsomethingsimilarinordertocamouflagethehole,toprotectthetrapfrom
sunlightandfromotherpredators(Figure3).
Thismethodprovides reasonableactivity-densityestimates forgroupssuchbeetles (Baars,
1979),spidersandants(Wang,Strazanac&Butler,2001).
Therearealsoinevitablybiasesinpitfallsamplingwhilecomparingdifferentgroupsofanimals
and different habitats. An invertebrate’s probability of being captured depends on the
structure of its habitat, such as the density of vegetation; activity of a certain taxa (higher
physiologicalactivityand/or largerareaof influence);animal size,bybeing too large tobe
Figure3.Pictureofanuncoveredpitfalltrapintheburntsite
The short-term effects ofmulching on the soil-dwelling arthropod community after awildfire
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trappedortoosmallthatitisnotcollected;andalsofromfactorssuchhastemperatureand
rain,whichalteranimal’sbehaviour.Comparisonsbetweendifferentgroupsmust,therefore,
take into account variation in habitat structure and complexity, changes in ecological
conditionsovertimeandtheinnatedifferencesinspecies(Woodcock&Leather,2007,chap.
3;Spence&Niemela,1994).
Pitfall traps, while taking into consideration its limitations, are able to provide answers
regardingground-dwellingarthropod richness,which is thenumberofdifferent taxonomic
individuals identified; and abundance, which is the total number of collected individuals
disregarding its taxonomical aspects. Taxa richness and abundance combined can provide
informationaboutthechangesonthecommunitiesduringthesamplingperiodandevaluate
howtheyevolvethroughtime(Abbott&LeMaitre,2010).
Thetrapsusedinthisstudyhad8.5cmindiameterand12cmheightplasticbottombottles.
Thebottomofthebottlewasfilledbywater,alittlebitofethanol70%thatslowsdownthe
degradationofthecapturedanimalsandonedropofdetergentthatbreaksthesurfacetension
oftheliquidinordertopromotequickdrowning.
2.1.1.2 Samplingdesign
Atotalof12pitfalltrapswereinstalledatthetwostudysites,3ofwhichattheunburntsite
(UNB)and9attheburntsitetocomparethreedifferenttreatments:heavy-treatment(HT),
lowtreatment(LT)andnotreatment(NT),wheretreatmentstandsforthemulchapplication
atdifferentdensities.ThefieldinstallationwasperformedfromSeptemberuntillateOctober
(Figure4).Theappliedmulchmaterialwaschoppedeucalyptbark,whichisbelievedtobethe
bestfitasmulchmaterialsareregardedconsideringthesite’scharacteristics,availabilityon
siteandapplicationcost.Theunburntsiteactedasacontrollocationbeingareferenceofan
ecosystemwithverysimilarcharacteristicsthathasnotbeenchangedforatleast4years.
Pitfall sampling was performed for a period of seven days, repeated in the months of
September, October, November and January. The selection of sampling months had the
intentionoffollowingtheendofsummer,includingAutumn,whichisaseasonthatcomprises
manybiologicalevents(Gallinat,Primack&Wagner,2015),andearlyWinter.Thecollection
dayswere the25thof September, the14thofOctober, the25thofNovemberand the27thof
January.TheunburntsiteinstallationtookplaceonlyattheendofOctober,thus,itonlybegun
tobesampledfromNovemberon.
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AfterthefirstsamplingperiodofSeptember,whennotreatmenthadyetbeenapplied,theplots
attheburntsiteweremulched:onewithadensityofmulchmaterialof0.26kg.m-2andanother
with0.80kg.m-2;athirdplotremaineduntreated.Eachplotmeasures2metersinwidthand8
metersonlength,spacing10metersfromeachother,withthreepitfalls linearlyplacedand
equallyspacedalongtheslopeofthehill.Theunburntsitefollowedthesamesetting,buthad
onlyoneplot(Figure4).Thisdesignensuredthatafirstassessmenttotheground-dwelling
arthropodcommunitycouldbeexecutedrightafterthewildfireeventandalsoonlaterstages,
conductingtodatathatcouldclarifythequestionsraisedonthisstudyregardingtheuseof
mulch.
Figure4.Samplingdesigndisplaypersamplingmonths.
The short-term effects ofmulching on the soil-dwelling arthropod community after awildfire
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2.2 Ground-dwellingarthropodscollectionandidentification
Beforethesamplingperiodtookplace,everypitfalltraphadtobere-set,inordertoguarantee
that the collection of arthropodswas only from that 7-day period. Re-setting consisted on
cleaningtheinteriorofthepitfallandintroducingwater,ethanolanddetergent,aslistedbefore
(Chapter2.2.1).Afterthe7-dayperiod,theinteriorofeachtrapwasfilteredbyasmallmesh
(±1mm)andthesamplesweretransferredintoarecipient,respectivelylabelled.Thefiltering
process allowed elimination of accumulated water in the pitfall trap during the sampling
period,beingaveryusefulstepduringwinter.However,micro-arthropods(smallerthanthe
mesh size) could have beenwasted in this process. Due to this limitation, theAcariorder
collectedisconsiderednottobefullyrepresentative.
Thesamplescollected inthe fieldwere lateranalysed inthe laboratory.Firstly,ascreening
fromdirtandanimals tookplace (Figure5).Secondly,a screeningatorder level fromeach
pitfall was made (Figure 6). The screening process was done with the assistance of a
stereoscopic magnifier (Figure 7), in order to easily identify the selected taxa. The
identificationprocessfollowedtaxonomickeysfromChinery(1993)andwasalsosupported
byBarrientos(1988).
Figure6.Secondscreening,identifyingandcountingarthropodsatorderlevelperpitfall
trap.
Figure7.Firstscreeningprocess,separatingtrappedarthropodsfromlitter.
Figure5.Stereoscopicmagnifier
The short-term effects ofmulching on the soil-dwelling arthropod community after awildfire
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Identificationwasonlydoneat theOrder level, considering the timeeffort and the levelof
expertiseneededtoidentifylowertaxonomiclevels.Nonetheless,theuseofhighertaxonomic
levelscanprovidegoodoutcomeswhen itcomes to thestudyofarthropodcommunities in
severalecosystems(Simão,Carretero,Amaral,Soares&Mateos,2015).Arthropodlarvaewere
notconsideredinthecommunityanalysis.
2.3 Ecologicalfunction
Fourecologicalfunctionswererecognisedforthetaxathathadbeenidentifiedinthevarious
samples:microbialfeeders,plantfeeders,predatorsandomnivores.Theselectionoftaxa,at
theorderlevel,foreachfunctionalgroupwasperformedconsideringthemorphologicfeatures
foundintheindividualsfromeachsampleandalsobasedonBarrientos(1988).
Forthegroupofpredators,orderssuchasAranaea,Chilopoda,OpilionesandPseudoscorpiones,
were automatically assumed considering the main morphology found in these orders.
Collembolawereconsideredasmicrobialfeeders(feedingfromfungianddeadorganicmatter).
Thysanura,IsopodaandHymenopterawereconsideredasomnivores.Hymenopteraindividuals
wereonlyantsandtherefore,thesewerealsofittedintotheomnivoregroup.
Coleoptera individualshadasub-samplingproceduretocharacterisetheircommunity,since
thisorderisoneofthemostdiversewhenitcomestoecologicalfunctions(Rainio&Niemela,
2003). Considering the large number of individuals in some samples, a sub-sampling was
adopted to classify the collected specimensperecological functionbasedonmorphological
characteristics.Onsampleswithaverylargernumberofindividuals,only10specimenswere
classified.
Acari individuals were not considered in the ecological function since it is not possible to
determinetheirfunctionwithoutidentificationatfamilylevel.Dipterawasalsoexcludedfrom
theseanalysessincethesamplingmethod(pitfalltrap)isnotadequateforthisorder.
The short-term effects ofmulching on the soil-dwelling arthropod community after awildfire
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2.4 Environmentalvariables
Theenvironmentalvariablesconsideredinthisstudywererainfalllevelsandtemperature.
Theselectedenvironmentalvariableswerecollectedbyameteorologicalstationalsoinstalled
bytheESPTeam,neartothestudysites,onthe24thofSeptember.Inordertointerpretdata
provided by this station, which collected data on a 15-minute period, an average of
temperaturewascalculatedandthetotalprecipitationwassummedina10-dayperiod,having
atleast3valuesforeachmonth.The10-dayperiodwasselectedinordertoobtainvaluesthat
wouldmatchthenumberofsamplesinthefoursamplingmonths(3plotsx4months)andalso
considering the level of detail regarding environmental variables intended, which is
seasonality.
2.5 Dataanalysis
The results obtained by the collected samples, after being screened and sorted,were later
analysedstatisticallywithSPSS13.0andMicrosoftExcel2015.
Alltaxawereanalysedatorderlevel,asmentionedbefore,andthedataretrievedwasusedto
calculate ground-dwelling arthropod community’s structural parameters: total abundance,
total richness, Shannon-Weiner diversity index (H’) andPielou’s evenness index (J’). These
parameterswereorganizedbysamplingmonthsandperplots.Theindexeswerecalculated
fromtotalabundanceresultsataplotlevel,inwhichthetotalabundancewasthesumofthe3
pitfall-trapsinstalledoneachplot.
TheShannon-Weinerdiversityindex(H’)andthePielou’sevennessindex(J’)werecalculated
accordinglytothefollowingformulas.
𝐻´ = − 𝑝'𝑙𝑛 𝑝' *
'+,
• pi=fractionoftheentirepopulationfromorderi;
• T=numberofordersfound;
• ∑=sumfromorder1toorderT;
The short-term effects ofmulching on the soil-dwelling arthropod community after awildfire
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𝐽. = 𝐻′
𝐻′012
where,𝐻′012 = − ,*𝑙𝑛*
'+,,*= ln 𝑇
• T=numberofordersfound;
These indexeswere calculated accordingly with the total abundance at plot level for each
samplingmonth.
Pearson’scorrelationswereperformedinordertocomprehendrelationsbetweenordersand
between ecological function groups through the sampling period and also between total
abundanceandtheenvironmentalvariables.Correlationswereconsideredsignificantona2-
tailedtest.
A two-wayANOVAwasused to test for significantdifferencesamong themonths (October,
NovemberandJanuary)aswellasbetweentreatments(NT,LTandHT)regardingthedistinct
community parameters analysed (total abundance, diversity and evenness). Specific
differencesbetweenfactorswereanalysedusingtheTukeymultiplecomparisontest.Inthe
caseoftheecologicalfunctionanalysis,aOne-wayANOVAtestwasusedperfactor(months
andtreatments).Normalityandhomogeneityofvariancesofthedatawereassessedusingthe
Shapiro–WilktestandtheLevenemediantest,respectively.Ifdataisnotstatisticallynormal,
aNaperianlogarithm(ln)wasappliedtoensureanormaldistribution.Allstatisticaltestswere
carriedoutusingaαof0.05.
Inthestatisticalanalysisprocedure,onlytotalabundancewasused.Totalrichnessatorder
levelinthisstudycontextdisplayedverysmalldifferencesthatleadtoinconclusivestatistical
results.TheUNBplotandthesamplingperformedinSeptemberwerenotconsideredonthe
statisticalanalysisregardingdifferentsamplingperiods,becauseinSeptembernomulchwas
presentonanyplotandtheUNBsitewasonlysampledinthelasttwomonthsofthisstudy
(NovemberandJanuary).Duetothesereasons,statisticalcomparisonregardingrichnesswas
notapplicable.AcariandDipterawerealsoexcludedfromthestatisticalanalyses,considering
thepitfallsamplinglimitationsconcerningthesetwoorders.
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3. Results
3.1 Environmentalvariablesanalysis
In Figure 8, it is seen that regarding precipitation, the first big events took place only at late
DecemberandearlyJanuary.Beforethatperiod,smallprecipitationeventswereregistered,always
below 50mm in a 10-day period. Betweenmid-November andmid-December, there were few
precipitationevents.Meanwhile,averagetemperatureina10-dayperiodhadasteadydeclinefrom
lateSeptembertoearlyNovember.ThetemperaturehadamajordeclineduringNovember,from
15°Ctoa7°Caverage.Afterwards,temperatureroseuntilmid-December,reachingapeakat13°C.
Inthesameperiodthattemperaturereacheditslowest,precipitationwasalsoveryscarce.January
displaysameantemperaturearound9°C.
Figure8.MeanandstandarddeviationfortemperatureandprecipitationbetweenlateSeptember2015andJanuary
2016
3.2 Communityanalysis
3.2.1 RichnessandAbundance
Duringthe4samplingmonths,thatcomprisetwoseasons(AutumnandearlyWinter),2303
ground-dwellingarthropodswereretrievedbythepitfalltraps,representingintotal14orders
(Richness): Collembola, Coleoptera, Diptera, Hemiptera, Hymenoptera, Isopoda, Thysanura,
0
75
150
225
300
375
0
5
10
15
20
25
1-Sep-15 1-Oct-15 1-Nov-15 1-Dec-15 1-Jan-16TotalPrecipitatio
n[m
m]
Tempreature[°C]
TotalPrecipitation
AverageTemperature
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Chilopoda,Araneae,Acari,Diplopoda,Pseudoscorpiones,PsocopteraandOpiliones(Appendix1
and2).
Although 14 orderswere found, only amaximum of 12were identified on one single plot
duringthesamplingperiod.TakingintoaccounttheconditionsofthehabitatinSeptember,a
considerablenumberoftaxawasyetidentified,namely8to10orders.
TotalrichnessdifferencesbetweenSeptemberandJanuarywerenolargerthan5.Thishigher
differenceoccurred in theNTplot fromOctober toNovember.TheLTplot remainedquite
equal during the sampling period, with total richness of around 10, whereas the HT plot
registeredapeakfrom8to11betweenSeptemberandNovember.Afterthemulchapplication
that occurred inOctober, it is seen that only theHTplot had an increase in total richness,
whereastheLTremainedstableandtheNThadadecrease.Ontheotherhand,inNovember,
the NT and LT had a richness growth, more accentuated on the NT plot, whilst the HT
decreased.InJanuary,theLTandUNBhadadecreaseintotalrichness,HTremainedidentical
tothepreviousmonthandtheNTincreasedfrom11to12orders(Figure9).
Figure9.Totalrichnessregardingsamplingmonthsandplots
Nostatisticallysignificantdifferenceswerefoundbetweensamplingmonthsandtreatments
viaaTwo-wayANOVAtestregardingtotalrichness(Table1).Thetestpointsoutasignificant
interaction betweenmonths and plots (treatments) (Table 1). Significant differenceswere
foundfortreatmentinNovember,namelywiththeNTandtheHT,andonmonthsintheNT
4
5
6
7
8
9
10
11
12
13
14
September October November January
Totalnum
bero
fide
ntified
Orders
NT
LT
HT
UNB
Mulch Application
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plot,namelywithinNovemberandOctoberandalsowithinJanuaryandOctoberforthesame
plot(Table2).
Table1.Two-wayANOVAsummaryregardingmonthandplotwithtotalrichnesspertrap.Significantstatisticalrelationsinboldtype.
SourceofVariation DF SumofSquares MS F P
Month 2 7.185 3.593 1.830 0.189Treatment 2 6.741 3.370 1.717 0.208
MonthxTreatment 4 23.259 5.815 2.962 0.048Residual 18 35.333 1.963 Total 26 72.519 2.789 NormalityTest(Shapiro-Wilk):passed(p=0.083)
Table2.AllPairwiseTukeymultiplecomparisontestwithinmonthsandtreatments.
Factor1 Factor2 Comparison DiffofMeans p P<0.05Treatment October HTvs.NT 2.000 0.215 No
HTvs.LT 0.333 0.954 DoNotTest
LTvs.NT 1.667 0.334 DoNotTest
November NTvs.HT 3.333 0.024 Yes NTvs.LT 1.333 0.488 No LTvs.HT 2.000 0.215 No January NTvs.HT 2.000 0.215 No NTvs.LT 0.667 0.831 DoNotTest
LTvs.HT 1.333 0.488 DoNotTest
Month NT Novembervs.October 4.000 0.007 Yes Novembervs.January 1.000 0.663 No Januaryvs.October 3.000 0.044 Yes LT Novembervs.October 1.000 0.663 No Novembervs.January 0.333 0.954 DoNotTest
Januaryvs.October 0.667 0.831 DoNotTest
HT Octobervs.November 1.333 0.488 No Octobervs.January 1.000 0.663 DoNotTest
Januaryvs.November 0.333 0.954 DoNotTest
Lookingtotheabundancefoundduringthesamplingperiodsperplot,allofthesampledplots
follow a similar trend. Earlier, it seems that before the mulch application process, the
abundance in the sampledplotsdecreased.After this firstperiod, inNovember, abundance
increased, especially on the NT andHT plots. BetweenNovember and January, abundance
decreased,buttohigherlevelswhilecomparingwithOctober(Figure10).
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Regardingtotalabundancevariationsonthesamplingperiod,atrendisidentifiedwithinthe
differentplots.FromSeptembertoOctober,allplotsexperiencedadropintotalabundance
whichwaslaterfollowedbyanincreaseinallplotsinNovember.Onthelastmonth,January,
all plots had a decrease in total abundance, to levels a little higher than the registered in
September.
Figure10.Totalabundanceregardingsamplingmonthsandplots.
Inordertobetterinterpretthedataanalysedonthepreviousfigure,relativeabundancewas
calculatedbyobtainingmeanabundanceperplot,consideringabundancelevelsoutofsetsof
threepitfalltraps(Figure11).
Figure11.Meanabundanceandrespectivestandarddeviationpersamplingmonthsandplots.
Regardingthemeanabundanceperplot,thetreatedplots(LTandHT)displayhighervaluesof
relativeabundancewhilecomparingwiththeNTplotsandtheUNB.Adecreaseofproximately
0
50
100
150
200
250
300
350
400
450
500
September October November January
Totalnum
bero
fcollectindividu
als
NT
LT
HT
UNB
MulchApplication
0
50
100
150
200
250
300
350
September October November January
Abun
dance
UNB NT LT HT
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50%lessrelativeabundanceisidentifiedbetweenSeptemberandOctoberoneachplot.The
peakintotalabundanceidentifiedinFigure10isalsoseenonrelativeabundance,wherethe
NTandLTplotdisplaysimilarvalues.Ontheotherhand,relativeabundanceontheHTplotin
Novemberhadalargedisparityoftotalabundancebetweenpitfalltraps,sincethestandard
deviation is larger thanthemeanabundanceonthatplot. In January, there isan increasing
relativeabundancefromtheUNBplot,totheNT,LTandtheHTplot,between25and75.
ATwo-wayANOVAtestwasperformed,consideringthesamplingmonthsandthetreatments
asfactors(Table3)(Appendix2).
Table3.Two-wayANOVAsummaryregardingmonthandplotwithtotalabundancepertrap.Significantstatisticalrelationsinboldtype.
SourceofVariation df SumofSquares MS F P
Month 2 4.817 2.408 2.537 0.107Treatment 2 0.282 0.141 0.149 0.863
MonthxTreatment 4 2.484 0.621 0.654 0.632Residual 18 17.089 0.949 Total 26 24.671 0.949
NormalityTest(Shapiro-Wilk):passed(p=0.545)
Althoughthereisnotastatisticallysignificantdifference(p=0.107)betweenmonths,thereare
considerable differences in abundances. Regarding the abundances between treatments
(plots)therearenodifferences.Theinteractionbetweenmonthsandtreatmentswasalsonot
significant.
Regardingtaxadistributionalongthestudyarea(Figure12),Collembolaisthemostabundant
order,foundoneveryplotandincreasingalongeachsamplinginmostoftheplots,followedby
Hymenoptera andAcari. Contrary toCollembola,Hymenoptera individuals decrease steadily
fromSeptembertoJanuary.Dipteraarepresentinallplots,butinlargernumbersduringthe
monthsofNovemberandJanuary.Acariarealsopresentineveryplot,althoughtheyaremore
abundant in September and January in some plots. Araneae and Coleoptera are also
representedinallplots,althoughapatternisnotyetidentifiable.ItisalsoseenthatColeoptera
abundanceintheHTplotinNovemberisthemaincauseforthemarkedincreaseinabundance
foundearlierinFigure9and10.Regardingthelessrepresentedorders,Thysanurawasonly
foundintheUNBplotandDipluraonlypresentinthetreatedplots.
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Figure12.Relativeabundanceinpercentageperplotduringthesamplingmonths.
StatisticalsignificantcorrelationsbetweenordersaredisplayedonTable4.Atabledisplaying
allresultsregardingthePearsoncorrelationtestisfoundonAppendix3.
Table4.Pearsoncorrelationbetweenordersoneachsamplingmonth.
Month OrderPair N PearsonCorrelation Sig.(2-tailed)
OctoberAraneae&Hymenoptera 3 1 0.011*
Collembola&Pseudoscorpionidea 3 0.997 0.048*November Coleoptera&Hymenoptera 3 0.997 0.05*January Acari&Coleoptera 3 -0.997 0.046*
*Correlationissignificantatthe0.05level(2-tailed).
InOctober,statisticallysignificantresultsarefoundregardingacorrelationbetweenAraneae
andHymenopteraandbetweenCollembolaandPseudoscorpionidea(Table2).Bothcorrelation
coefficientspointtowardsariseintotalabundancebetweenthepairsoforders.TheCollembola
andPseudoscorpionideacorrelationmaybeduetoa trophicrelation.OnNovemberthere is
alsoasignificantcorrelationbetweenColeopteraandHymenoptera,althoughecologicallyitis
adoubtable result considering theColeoptera total abundanceon theHTplot inNovember
(Figure12).
0%
10%
20%
30%
40%
50%
60%
70%
80%
90%
100%
NT LT HT UNB NT LT HT UNB NT LT HT
October November January
Relativ
eAb
undance
Thysanura
Psocoptera
Pseudoscorpiones
Orthoptera
Opilionidea
Isopoda
Hymenoptera
Hemiptera
Diptera
Diplopoda
Collembola
Coleoptera
Chilopoda
Araneae
Acarina
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Figure13.TotalabundanceperorderinSeptember
ThetotalabundanceofordersinSeptembercorrespondstotheindividualstrappedinatotal
ofninepitfalls(distributedover3plots,seeFigure4)(Figure13).Atotalof518individuals
were collected in September. It is seen thatHymenoptera is the largest represented order,
followed by Collembola, Acari andAraneae. Richness, considering all plots together in this
month,is11outof14.
Figure14 shows thedistributionof themainordersperplotoneachsamplingmonth.The
“Others”representthetaxathatislessabundant(<50identifiedindividualsduringthewhole
sampling period), namely Diplopoda, Pseudoscorpiones, Psocoptera, Opiliones, Orthoptera,
Hemiptera,Isopoda,ThysanuraandChilopoda.Althoughtheseareinalowernumber,theyare
stillrelevantregardingecologicalfunctioninthepost-fireperiod.
0 50 100 150 200 250
HymenopteraCollembola
AcarinaAraneae
ColeopteraDipteraIsopoda
PseudoscorpionidaChilopoda,Opiliones,Hemiptera
TotalabundanceinSeptember
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Figure14.TotalAbundanceperplotduringsamplingmonths.
TotalabundancefollowsasdescribedonFigure9,showinghowmucheachtaxacontributes
andhoweachtaxaevolvesduringthesamplingperiod.Octoberwasthemonthwiththelowest
abundancewhereasNovemberdisplaysthemaximumtotalabundance.
CollembolashowsasteadyriseinabundanceontheHTplotofalmostfourtimesasmorefrom
September to January.On the other plots, therewas also amarked increase inCollembola,
notedfromOctobertoNovemberintheNTplot,andalsoontheLTplot,althoughnotasmuch
while comparingwith other plots. On theHT plot, there is a very high total abundance of
ColeopteraduringNovember,onemonthaftertheapplicationofmulch.Thiswasalsodisplayed
inthepreviousfigures.TheUNBplot,despitedisplayingalowertotalabundanceasotherplots,
200 individuals inNovember and 100 in January, it shows amore evenly distributed total
abundanceperOrder.
050
100150200250300350400450
Totalabu
ndance
NTPlot
050100150200250300350400450
LTPlot
050
100150200250300350400450
HTPlot
050100150200250300350400450
November January
Totalabu
ndance
UNBPlot
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3.2.1 AbundanceandEnvironmentalvariables
Table5showsthattemperatureandtotalabundancearenotsignificantlycorrelatedbythe
Pearsoncorrelationtest,yetthep-valueisclosetobesignificant.However,evennotsignificant,
thetotalabundancedecreaseswithdecreasingtemperature.
Table5.Pearsoncorrelationbetweentotalabundanceandabioticvariables.
VariablePair N PearsonCorrelation Sig.(2-tailed)Temperature&Totalabundance 3 -.638 0.065Rainfall&Totalabundance 3 -.232 0.548
3.2.2 Diversity(H’)andEvenness(J’)
Regarding diversity (H’) and evenness (J’) indexes there is a decreasing trend in both
throughoutthesamplingperiods(Figure15).Nonetheless,theHTandNTplotsshowsteeper
decreasesinbothdiversityandevennessindifferentperiods:NovemberontheNTplotand
JanuaryontheHTplot.
Figure15.TotaldiversityandtotalevennessintheLT,HTandNTplotsduringsamplingmonths
The Two-way ANOVA test pointed-out significant differences in diversity betweenmonths
(Table 6), namely from October through January (Table 7). No differences were observed
betweendifferenttreatments(Table6).TheTwo-wayANOVAtestdidnotprovidesignificant
differencesregardingevennessindifferentmonthsandplots(Table6).
0,0
0,2
0,4
0,6
0,8
1,0
0,0
0,4
0,8
1,2
1,6
2,0
LTPlot
DiversityIndex(H') Evenness Index(J)
0,0
0,2
0,4
0,6
0,8
1,0
0,0
0,4
0,8
1,2
1,6
2,0
Even
ness
HTPlot
0,0
0,2
0,4
0,6
0,8
1,0
0,0
0,4
0,8
1,2
1,6
2,0
Diversity
NTPlot
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Table6.Two-wayANOVAsummaryregardingmonthandtreatmentasfactorsanddiversityandevennessindexeshasvariables.Significantstatisticalrelationsinboldtype.
Variable SourceofVariation DF SS MS F P
Diversity* Month 2 0.249 0.125 7.779 0.042 Treatment 2 0.0574 0.0287 1.793 0.278 Residual 4 0.0641 0.0160 Total 8 0.371 0.0463
Evenness** Month 2 0.0604 0.0302 3.389 0.138 Treatment 2 0.00777 0.00389 0.437 0.674 Residual 4 0.0356 0.00891 Total 8 0.104 0.0130
*NormalityTest(Shapiro-Wilk):Passed(p=0.427)**NormalityTest(Shapiro-Wilk):Passed(p=0.242)
Table7.Tukey’smultiplecomparisonofmonthsregardingdiversity.
Comparison DiffofMeans p p<0.050
Octobervs.January 0.369 0.050 -Octobervs.November 0.0357 0.938 NoNovembervs.January 0.334 0.067 No
3.3 Ecologicalfunctionanalysis
AssignmentofordersperecologicalfunctiongrouparefoundonAppendix4.
In September, most of the taxa found were omnivore,
followedbymicrobial feeders, predators and, at last, by
very few plant feeders (Figure 16). After the treatment
wasapplied,omnivoresdecreasedandmicrobialfeeders
andplant feeders increased, fromOctobertoNovember.
InNovember,thetotalabundanceofomnivoresstandsout
ontheUNBplot,representingabout50%ofthesampled
individuals.InJanuary,about80%ofthecommunityisa
microbialfeederonallplots(Figure17). Figure16.EcologicalfunctiondistributioninSeptember
MicrobialFeeders33%
PlantFeeders1%
Omnivores51%
Predators15%
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Figure17.Ecologicalfunctionrelativeabundanceperplotduringsamplingmonths
Table8&9displaytheresultsofthePearsoncorrelationsbetweenecologicalfunctiongroups.
AllcorrelationtestresultsarefoundinAppendixes5&6.Whenthemonthsareconsidered,
there is a significant correlation between plant feeders and predators. When plots are
concerned,moresignificantcorrelationswerefound.Thecorrelationshavethepredatorgroup
incommon.OntheNTplotastrongincreasingtrendbetweenmicrobialfeedersandpredators
wasfound.OntheLTplot,therewasadecreaseofmicrobialandplantfeedersrelatedwithan
increaseinthenumberofpredators.
Table8.Pearsoncorrelationbetweenecologicalfunctiongroupsoneachsamplingmonth.
Month GroupPair N PearsonCorrelation Sig.(2-tailed)January PlantFeeders&Predators 3 1.000* 0.007
*Correlationissignificantatthe0.01level(2-tailed)
Table9.PearsoncorrelationbetweenecologicalfunctiongroupsoneachPlot.
Plot GroupPair N PearsonCorrelation Sig.(2-tailed)NT MicrobialFeeders&Predators 3 0.999 0.024
LTMicrobialFeeders&Predators 3 -1.000* 0.007PlantFeeders&Predators 3 -0.997 0.046
*Correlationissignificantatthe0.01level(2-tailed)
0%
10%
20%
30%
40%
50%
60%
70%
80%
90%
100%
NT LT HT UNB NT LT HT UNB NT LT HT
October November January
Predators
Omnivores
PlantFeeders
MicrobialFeeders
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Figure18complementsthestatisticalcorrelationsshownearlier.Onthetreatedplots,LTand
HT, the ecological function groups followed an identical trend. Omnivores and predators
decreased, microbial feeders and plant feeders increased, reaching a peak in November,
although at a different scale. Abundance per ecological function groups present also very
similarnumberof individuals.PlantfeedersontheHTrosesteeplyinNovemberduetothe
highnumberofColeoptera.
Figure18.Ecologicalfunctiongroupstotalabundanceperplotduringsamplingmonths.
ThegreatmajorityofindividualsfoundinSeptemberintheNTplotareomnivores.InOctober,
most of the groups decreased, but the microbial feeders had a slight increase. The slight
increasewas followedbya steep increase inNovember, from15 to160 individuals,whilst
other groups had a very small increase, of about 10 individuals. Ultimately, in January,
omnivores andmicrobial feeders decreased in total abundance, while predators and plant
feedersmaintainedtheirtotalabundancerelativelytoNovember(Figure19).
A One-Way ANOVA was performed to assess significant differences in ecological function
groupsregardingdifferentfactors:monthsandtreatments(Table10and12).
0
30
60
90
120
150
180HT
0
30
60
90
120
150
180LT
0
30
60
90
120
150
180NT
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Table10.One-WayANOVAonfunctionalgroupsregardingsamplingmonths.
EcologicalFunction Sumof
Squares df MeanSquare F P
MicrobialFeedersBetweenGroups 13152.889 2 6576.444 12.333 0.007WithinGroups 3199.333 6 533.222
Total 16352.222 8
PlantFeeders
BetweenGroups 7698.667 2 3849.333 1.609 0.276WithinGroups 14355.333 6 2392.556
Total 22054 8
Omnivores
BetweenGroups 1510.889 2 755.444 24.545 0.001WithinGroups 184.667 6 30.778
Total 1695.556 8
Predators
BetweenGroups 10.889 2 5.444 0.212 0.815WithinGroups 154 6 25.667
Total 164.889 8
Table11.Tukeymulti-comparisontestonmicrobialfeedersandomnivoresregardingdifferentmonths
DependentVariable MonthPairs DiffofMeans p p<0.005
MicrobialFeeders
OctobervsNovember -92.66667 0.006 Yes
OctobervsJanuary -58.00000 0.050 -
NovembervsJanuary 34.66667 0.236 No
Omnivores
OctobervsNovember -17.66667 0.019 Yes
OctobervsJanuary 14.00000 0.049 Yes
NovembervsJanuary 31.66667 0.001 Yes
Table12.One-WayANOVAonfunctionalgroupsregardingtreatments.
EcologicalFunction SumofSquares df Mean
Square F P
MicrobialFeedersBetweenGroups 118.222 2 59.111 0.022 0.978WithinGroups 16234 6 2705.667
Total 16352.222 8
PlantFeedersBetweenGroups 4418 2 2209 0.752 0.511WithinGroups 17636 6 2939.333
Total 22054 8
OmnivoresBetweenGroups 66.889 2 33.444 0.123 0.886WithinGroups 1628.667 6 271.444
Total 1695.556 8
PredatorsBetweenGroups 80.222 2 40.111 2.843 0.135WithinGroups 84.667 6 14.111
Total 164.889 8
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Statisticaldifferenceswere foundonlywhenconsideringthemonthsasa factor(Table10).
Within the different months, microbial feeders and omnivores had statistically significant
differences. These differences were statistically significant between all months in both
ecological groups, except between November and January on the microbial feeders group
(Table11).Asfarasplotsareconcerned,nosignificantdifferenceswerefoundbetweenthe
ecologicalfunctiongroups(Table12).
The short-term effects ofmulching on the soil-dwelling arthropod community after awildfire
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4. Discussion
Wildfiresconductmanydirectandindirectimpactsonecosystems(Bowmanetal.,2013).One
oftheindirectimpactsproducedbywildfiresisthealterationofthegrown-dwellingarthropod
community(Eliaetal.,2012),akeyelementonsoilquality(Stork&Eggleton,1992).Although
some studies report a cause-effect relation between fire and insects (Bowman & Murphy,
2010), few studies are clear on this matter, regarding the ground-dwelling arthropod
community’sresponsetowildfire.Thesestudiesalsodonotprovidehomogeneousoutcomes
regardingthecommunityresponseandresilience(Eliaetal.,2012).Infact,wildfireeventsare
shapedbymanydifferentvariablesthatconsequentlywillimpactontheextentofseverityof
post-fireevents(Certini,2005).
Thewildfireactsdirectlyasamortalityagentona firststage,bycremationand lethalheat
exposuretothelessmobilearborealandotherfoliage-feedingarthropods(New,2014).Soil
microandmacro-faunaaremoresusceptibledependingon the fire severity impact,mostly
relatedwithheat intensity and exposure (Certini, 2005).On a second stage, right after the
wildfireevent,awindowofopportunityopenstothemorefireresistantandmobilegroups,
namelysaproxylicinsectsandthesoil-dwellingzoophagosandsaprophagous(Moretti,Duelli
&Obrist2006).Thesegroups,withhighertolerancetofireandmoremobile,willalsobepart
ofthere-colonizationprocessoftheareaafterafire,sinceaburntsiteprovideswellsuited
habitat fortheiractivitiesandecologicalroles.Generally,predatorsarethemostnegatively
affected by fire and detrivores are more abundant in fire prone sites (Engstrom, 2010).
Indirectly,thewildfiretransformingeffectsonthehabitatsetting,byremovingvegetationand
litter,will alsopromotedifferentoutcomes.Thearthropod community compositionwill be
dependentonecologicalprocesseslikestarvation,predationorimmigration(Engstrom,2010).
Starvationwillberelatedwiththeavailabilityoffoodtosurviveinassociationwiththelossof
vegetation,which isavital source forherbivorearthropods (Haddadetal.,2009).This can
ultimatelylimitotherfood-webrelations,suchhaspredation.Outcomesfromtheseprocesses
willdeterminethepopulationviabilityinthepost-firecontext(Engstrom,2010).Ultimately,
arthropodabundanceandrichnessvariationsareusuallyrelatedwiththeverydistinctclimate
factorsfromtheMediterranean-typeecosystems,suchhastemperature,rainfallanddaylength
(Simãoetal.,2015).
Inthesensethatmostofthebiomass(vegetationandlitterlayer)islostduringawildfire,the
application of mulch, in the form of chopped eucalyptus bark, is an amendment to the
transformedhabitat.Thiswillverylikelyhavesomeecologicalvalue,sinceagreatamountof
The short-term effects ofmulching on the soil-dwelling arthropod community after awildfire
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biomass (2.6ton to 8ton/ha considering each treatment type regarding the mulching
application)isaddedtoanecosystemthatlostagreatamount.Italsohasbeenwitnessedthat
applicationofforestresidualshasamulchingagentalsoincreasestheorganicmattercontent
inthesoil(Pratsetal.,2014),whichecologicallycanhaveapositiveeffect.Ontheotherhand,
erosioncanalsodictatehowfastanecosystemcanrecoverafterawildfire,consideringthat
erosionratesandrunoffinmono-specificplantationsofeucalyptarehigh(Pratsetal.,2015).
Theseeffectspromotethelossofnutrientscontainedinplantbiomassandsoilorganiclayers,
especiallyafterrainfallevents(Machadoetal.,2015),whichinthiscontextcanbevitaltothe
lifecyclesofmanyground-dwellingarthropods.Hence,theapplicationofmulchcontributesin
thereductionoferosion,whichcanalso,potentially,benefit theground-dwellingarthropod
community.
Onthisstudycontext,itisnecessarytotakeintoaccountoneofthemaincharacteristicsfrom
theburntarea,whichisthesoiluseinthisarea.Theareahasbeenwidelyusedhasaeucalypt
plantation.BremerandFarley(2010)statedthatbiodiversityineucalyptusplantationsislow
and,ifmanagementpracticesandfirehistoryareadded(Oliver,MacNally&York,2000),the
conjugationofbothoftenresultsonalowerbiodiversity.
FocusingontheresultsofSeptember,regardingtotalabundance,oneofthemostrelevantfact
isthatHymenoptera,exclusivelyants,wasthemostabundantordercollected,weighingabout
50%ofthetotalabundance(518individuals,in9pitfalltraps).Matsudaetal.(2011)considers
ant’sresistanttofire,sinceonlyabout2%ofthepopulationofamatureantcolonyisactiveat
surfaceleveland,therefore,exposedtofirehazards.Antsarealsoknowntobeinthelineof
thefirstorganismstocolonisedisturbedareas,forinstancebyfire,duetotheirgeneralistand
opportunistbehaviour,exploitingspaceandresourcesefficiently(Antunes,Curado,Castro&
Gonçalves,2009).ItshouldalsobetakenintoaccountthatinPortugalantsseemtothrivein
eucalyptusplantationecosystems(Zina,Garcia,Valente,Branco&Franco,2015).
Other orders, namely Collembola and Acari, each representing about 20% of the total
abundance and Araneae that represents about 10%, mostly characterize the community
immediatelyafterthewildfireevent.Collembola isaverycommonorderpresentonthesoil
thatfeedsmostlyonfungianddecayingplants(Neher,Weicht&Barbercheck,2012;Anslan,
Bahram&Tederoso,2016)andactingalsoaspreytospidersandbeetles.Itisalsoexpectedto
someextent,dependingontheseverityofthewildfire,thatcollembolansprofitfromtheeffects
producedby awildfire. In fact, there is an increaseofplantmatterdecay, considering that
coarsewoodydebrisleftoversfromthefirethatcontributeinregeneratingmicrobialactivity
The short-term effects ofmulching on the soil-dwelling arthropod community after awildfire
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andinputofnitrogenandphosphorustothesoil(Marañón-Jiménez&Castro,2013;Brais,Sadi,
Bergeron&Grenier,2005).AcarifollowupCollembolahasoneofthemostcommonordersin
forest soils. Acari are highly diverse and regulate litter decomposition and nutrient
mineralizationrates(Camann,Gillette,Lamoncha&Mori,2008). Inthisstudy, it isbelieved
that the population of Acari sampled is mainly predators, due to their overall size and
morphologicfeaturesseenduringthescreeningprocess(Koehler,1999).Thesizeconditionin
Acariwasimposedbythemeshmeasureusedinthesamplingprocess(Chapter2.3),onwhich
Acariindividualssmallerthan1mmcouldhavebeenwashedaway.Addingup,pittfalltrapping
is also not the most adequate sampling method for sampling the full extent of Acari
biodiversity. TheAraneae are predators widely represented in forest ecosystems and also
adapted to forest fires (Buddle,Spence&Langor,2000).Araneae isexpectedonapost-fire
contextatsomelevelsincemostspiderscanrapidlyre-colonizevia long-distancedispersal,
likeballooning,andshort-distancedispersal,mainlyfromunburnedforestpatches(Buddleet
al.,2000).
Regarding ecological function groups in September, the community sampled was 51% of
omnivores,33%ofmicrobialfeedersand15%predators.Theseresultstranslatetheobtained
valuesregardingabundance,whereHymenopterawasthemostrepresentedorder.Only1%of
thesampledindividualswereclassifiedasplantfeeders,whichindicatestosomeextentthe
absenceofvegetationtofeedintherecentlyburntarea.
The second sampling took place in October, after the mulch application. Total abundance
decreasedinallplots.However,theplotwiththelargestdensityofmulchrichnessrosefrom8
to11orders.Althoughtherewasageneraldecreaseoftotalabundanceinthearea,theplot
withahighdensityofmulchmaterialattractedalargerdiversity(orderrichness)ofground-
dwellingarthropods.Infact,theLTplotregisteredasimilarlevelofrichness(10)andtheNT
plothadadecreasefrom9to7.Thisobservationmaybeduetothefactthatthemulchedareas
may provide more opportunities to survival to more groups, such as shelter and food
resources.Nonetheless,assumptionsraisedbyanalysingtotalrichnessinthisstudycaseneed
tobecarefullyconsidered.Inonehand,pitfalltrappingpresentsaflawatthislevel,inthesense
that there is a chance that orders with a very low total abundance were not sampled by
causality.Ontheotherhand,thelevelofdetailfromacommunityprovidedatataxonomiclevel
suchasOrdermaynotbeenoughtopointoutdifferencesbetweentreatments.
The fact that there was a decrease in abundance while comparing with the results from
September,mayberelatedwiththerainfallandtemperatureregisteredduringOctober.Inlate
The short-term effects ofmulching on the soil-dwelling arthropod community after awildfire
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September and early October, very little precipitation occurred, less than 50mm and
temperatureremainedstableatameanofapproximately17°C.Infact,thelackofrainfallona
post-firecontextcanrepresentadualeffect:lowerosionfromrunoff,sincethereisnotenough
rainfallandlowvegetationgrowth,inthesensethatrainfallmostlyimprovessoilmoistureand
nutrientavailabilitythatlaterpromotevegetationgrowth,whichalargenumberofarthropods
relyon(Certini,2005).Ultimately,theseepigeicfactorsshapeseasonality,whichisakeyfactor
influencingarthropoddiversityandabundance(Shakir&Ahmed,2014).
Summing up, October’s unusual meteorological conditions seemed to delay the overall
recoveryprocessoftheecosystemdespitethemulchapplicationissuggestingaslighthigher
totalabundanceandrichnessofground-dwellingarthropodsinthemulchedplots.
InNovember,themostrelevantobservationwastheabundancegrowthnotedinallplots.The
plotsthatregisteredthelargestgrowthweretheHTandNT,from77to429andfrom44to
283individuals,presentingabout6timesmorespecimenswhilecomparingtoOctober.Infact,
consideringthatthetotalabundancegrowthtookplaceineverysampledplot,thissuggests
that seasonality is playing an important role on shaping the ground-dwelling arthropod
community. This observationhas also some support by the statistical analyses, despite the
differences not being entirely significant. Differences were only closer to be significant
regardingdifferentmonthsandonlynotedbetweenOctoberandNovember. Ifweconsider
temperatureandrainfallbeforethesamplingcampaign,itisseenasteepdecreaseofthemean
temperature,apotentialtriggerthatmarkstheendofSummerandthebeginningofAutumn
(Gallinat et al., 2015). When we include functional groups to this assumption, the most
represented group in November are themicrobial feeders. This suggests that possibly the
ecosystem conditions during late October through late November have promoted the
development of fungi and consequently an increase in organic matter decomposition, a
commonecosystemfeatureinAutumnseason(Almagro,López,Querejeta&Martínez-Mena,
2009).Mostof the totalabundanceregistered isdueto themicrobial feeders,which in this
studyonlycompriseCollembola.OtherfunctionalgroupstotalabundanceroseinNovember,
althoughvery little.Differenceswereonly statistically significant formicrobial feeders and
omnivoresbetweendifferentmonths.Consideringtheincreaseofmicrobialfeeders,andthe
total abundance values regarding the population of predators, omnivores and other plant
feeders,itseemsthatthesepopulationsmayhaverespondedtothesechanges,consideringthe
trophicinteractionswithinthesegroups.ThePearsoncorrelationtestunveilsasignificantand
astrongcorrelationbetweenplantfeedersandpredatorsinJanuaryandbetweenmicrobial
feedersandplantfeederswithpredatorsintheUNBandNTplots.Exceptionally,ontheHT
The short-term effects ofmulching on the soil-dwelling arthropod community after awildfire
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plot,theplantfeedersincreasedmuchmoreitsabundanceincomparisonwithothergroups.
ThisobservationisrelatedwiththehighnumberofColeopteracollectedfromapitfallatthe
HTplotandthefurthersub-samplingperformedinthelaboratory.Duringthissub-sampling,
itwaspossibletovisualisethatthelargestnumberoftheseColeopteraindividuals,classified
asplantfeeders,hadmorphologicalfeaturesthatsuggestedthatthesefedonwoodyremains.
Infact,theforestresidualappliedtotheplotsurfacecouldhavelodgedthiskindofarthropods
initslarvaestage.Abioticand/orclimactericconditionssuchastemperature,rainfalloreven
seasonality could have triggered a rapid development of these Coleoptera morphospecies
(Wermelinger&Seifert,1999).ThedevelopmentoffungiintheareaduringNovembercould
also possibly been promoted by the growth of vegetation, by resprouting and by the
developmentofshrubs(Rincón&Pueyo,2010)aswellasbythealreadystatedseasonality.
Atthisstageofsampling,mulchimpactsontheground-dwellingcommunityarenotyetvery
clear, ifweonly take intoaccount totalabundanceatorder levelandecologicalgroups.No
statisticalsignificantresultswerefoundbetweendifferenttreatmentsfortotalabundanceand
ecologicalfunctiongroups.Whencomparingrichnessbetweenplots,itisseenanincreasein
total richness on the NT and LT plots, whereas the HT total richness decreased. This
observation,takingintoaccounttheconsiderationsabovementioned,suggeststhatmulchcan
becontributingtoacertaindegreeofinhibitionintherecoveryofthehabitat,inthesensethat
themulchlayercandismissthegrowthofsomefungiandplants,byeithercreatingaphysical
barrier,blockinglightandevenchemicalcontrolthroughtheleachingofallelopathicchemicals
present inwood (Chalker-Scott, 2007). This assumption can be relatedwith the density of
application,meaningthata largerdensitynormally impliesverylittleexposureoftheother
lowerlayers,whichmayinhibitplantdevelopment,whereasalessdenseapplicationallows
much more heterogeneity of ground cover. This heterogeneity can provide room for the
developmentoffungiandplants,increasingthevarietyofecologicalnichesinthearea(Menta,
2012).Consideringthescaleoftheareawhereground-dwellingarthropodsremainactive,itis
implicitthatamorediversecommunitycanbefoundduetothelargerdiversityofecological
nichesavailable,shapedbyplantrichness(Haddadetal.,2009).Theseassumptionsseemto
beassociatedtoobservedrichnessvariation,consideringagrowthontotalrichnessfrom7to
11taxaontheNTplot,alessaccentuatedincreaseontheLTfrom10to12taxa,thatcontrast
withadecreasefrom11to8taxaontheHTplot.Unfortunately,theUNBwassampledfrom
November on, and therefore, it is not possible to build comparisons taking into account
previousmonths.
The short-term effects ofmulching on the soil-dwelling arthropod community after awildfire
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The last sampled month, January, presents a decrease of abundance when comparing to
November.Winter season conditions only arrived in late December, bringing large rainfall
events followedby lowmean temperatures. Total abundance and season changes between
months, specifically temperature changes were statistical significant, asmentioned before.
Likewise,theUNBplotalsosuggeststhesame,adecreaseofabundancealsobetweenthetwo
sampling periods fromNovember and January.When comparing total abundancewith the
otherplots,abundanceintheUNBplotisrelativelylower.ThissuggeststhattheUNBplot,since
itdidnothaveafirerecently, itdidnotundergothesamephysicalandchemicalconditions
imposedbythewildfireintheburntsite.Additionally,UNBhadarecordofawildfirein2012
and, besides themulch absence, it seems to present amore advanced and stable stage of
ecologicalrecovery,similartoresultsobtainedbyPuga(2016).
Regardingrichness,theNThaditsrichnessraisedfrom11to12,theLTandUNBdroppedfrom
12to10and10to9respectively,andtheHTremainedat8.Takingintoaccounttheresults
foundintheprevioussamplingmonths,alltheplotspresentlowerrichness,inJanuary,when
comparing to the previous months. Nevertheless, the NT plot presents the highest total
richnessrecordinJanuary,consideringallsamplingmonths.ItseemsthattheNTplot,sinceit
didnothadmulchapplied,tookmoretimetoperforminitialecologicalsuccessionsordidnot
hadthenecessaryavailabilityofresourcestopromoteafasterrecovery,unliketheHTandLT,
whereastherichnesspeak inthetreatedplotswasreachedinOctober(HT)andNovember
(LT).TheHTplotdisplayslessrichnesscomparingtotheUNBplot,pointingoutthatalarger
densityofmulchdoesnotpromoteground-dwellingarthropodsdiversity.Ontheotherhand,
supportingtheabovestatedobservation,theLTplotpresentshigherrichness.Oncemore,the
total richnessrelatedassumptionsmust take intoaccount the fact thatorder level richness
might not provide a reliable answer, as above-mentioned. In addition, the field design, the
distancesbetweentraps,aswellasthenumberoftrapsusedandtheoverallareaofeachplot
mightnothavebeenenoughtotracesignificantdifferencesinthecommunitywithindifferent
treatments.
Diversity(H’)andEvenness(J’)indexessupportmostoftheobservationsdiscussedabove.A
decreasingtrendindiversityandevennessisvisiblefrommonthtomonth,whichsupportsthe
ideathatseasonalityisshapingthecommunity.Thisdecreasingtendencyisalsosupportedby
astatisticallysignificantdifferenceinvariancesbetweenmonths,namelyOctoberandJanuary.
Therearealsosomevisibleyetsmalldifferenceswithindifferenttreatments,wherethemost
importantoneisfoundonJanuaryattheHTplot,whichhasalowerdiversityandevenness
when comparingwith LT andNT. This observation is in linewith the theory that a higher
The short-term effects ofmulching on the soil-dwelling arthropod community after awildfire
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densityofmulchcouldpossiblyreducethenumberanddiversityofecologicalniches.When
comparingthetreated(LTandHT)anduntreatedplots(NT)withtheUNBplot,aslightlylarger
diversityisfoundontheUNBplotregardingdiversityandevenness.Thisobservationisrelated
bythewildfirehistoryandseemstoshowafurtherstageofre-establishmentofthecommunity
in the UNB plot. Once more, these indexes are calculated with total richness values and,
therefore,shouldbeinterpretedconsideringthealreadystatedlimitations.
Taking into account the main orders found during the 4 sampling months, Collembola,
Coleoptera,Diptera,Hymenoptera,AraneaeandAcari,therearesomeobservationsthatarise.
Collembolaistheordercontributingthemostontotalabundanceinallplots,especiallyfrom
Novemberon.Thisfollowsupwithwhatwasalsoobservedregardingtheecologicalfunction
groups.Dipterawereonly found, in a considerablenumber inNovember.Thisorder isnot
suitedtothepitfallsamplingsincethisordercanbeattractedbythesolutionoranimalremains
presentonthetrapstolayeggs.Hymenopterashowaninverseevolutionwhilecomparingwith
Collembola.The population ofHymenoptera decreased from September until January. This
steadydecreaseseemstoberelatedwithseasonality,regardingthetemperaturedropandhow
theactivityofantsisrelatedwithtemperature(Cerda,Retana&Cros,1998).AraneaeandAcari
displayalargerabundanceintheLTplot,whichmightberelatedwiththelargernumberof
nichesthattheLTplotoffers.
The short-term effects ofmulching on the soil-dwelling arthropod community after awildfire
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The short-term effects ofmulching on the soil-dwelling arthropod community after awildfire
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5. Conclusion
Hymenopterawas the most represented order in September, right after the wildfire. This
findingwasexpected,consideringthatantscanusuallysurvivetoawildfireeventduetotheir
burrowing capabilities.Hymenoptera total abundance gradually decreasedduring sampling
months,mostlikelyduetoseasonality.
Althoughdifferenceswithintreatmentandsamplingmonthswereexpected,onlyseasonality
seemedtobethedriverofabundancevariationsinthisstudycontext.Statisticalresultssuggest
seasonalityasbeinga factorshapingthetotalabundanceof theground-dwellingarthropod
community,aswellasthedecreasingtendencyindiversity(H’)andevenness(J’)fromOctober
through January. Slight differences were found on community total richness during the
samplingmonths,althoughnotsignificant.
These non-significant differencesmay be relatedwith the level of taxonomic identification
usedinthisstudy.Familylevelcouldhaveunveiledbetterinterpretationsregardingecological
groups and potential relations between mulch material and certain families, yet, the time
available toproceedwith thisstudywas limitedandtherefore, itwasnotpossible toreach
Familylevel.
A lower density of mulch application seemed to provide faster response regarding the
invertebrate community, and therefore a faster recovery.A largerdensity ofmulch, on the
otherhand,didnotseemtoproduceanidenticaloutput,sincetotalrichnesswaslowerand
totalabundancehigherwhencomparingwithalowdensityofmulchandnomulchedareas.
Inageneralperspective,mulchingseemednottopromoteaspecificorder,sincenoevidence
was found regarding significant differences between orders within different treatments.
Nonetheless,anuncommonobservationofhundredsofindividualsofaColeopteraspecieslead
to the assumption that the application of mulch may have introduced organisms in the
ecosystem.
The short-term effects ofmulching on the soil-dwelling arthropod community after awildfire
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The short-term effects ofmulching on the soil-dwelling arthropod community after awildfire
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6. Finalconsiderations
Literaturesuggeststhatanaccuratemeasureoffireseveritycanprovidemoreevidenceregarding
ground-dwelling arthropod re-establishment after a wildfire (Malmström, 2010). This
interpretationcouldallowabetterunderstandingonhowground-dwellingarthropodcommunity
response isdrivenafterawildfireandhowfardoesmulch impactspositivelydependingon the
wildfireseverity.EnvironmentalfactorssuchaspH,nutrientavailability,qualityoforganicmatter
are related with the fire severity in a post-fire context (Certini, 2005) and are also important
indicators to support ground-dwelling community parameters. Hence, fire severity accurate
assessmentcanbeveryusefulinthisstudycontext.
Forestresidualsasamulchingagentshouldbealsoreviewedinthesensethat itcancarryalien
speciesintotheecosystem,presentingapotentialthreat.
Someoftheresultsobtainedinthisstudyraisedthehypothesisthatmulchcouldhavecontributed
positively immediately on the re-establishment of ground-dwelling arthropods, although
statistically,nosignificantresultswereachieved.Inordertoreachabetterunderstandingabout
the extent of the impacts of mulch on ground-dwelling fauna, future studies should consider
identificationatfamilylevel,largersamplingplotsandtrapswithlargerdistanceintervalswithin
samplingplots.
Addinganunburntplotwith similar characteristics in this typeof studies canbe veryuseful to
comprehendhow theecological successionproceeds after thewildfire, in the sense that itwill
providecontrastingcommunityparameterswhilecomparingwithanecosystemthathadawildfire.
The short-term effects ofmulching on the soil-dwelling arthropod community after awildfire
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Appendixes
Appendix1.Totalabundanceperorderoneachpitfalltrapandsamplingdate.
SamplingDate
Order Total Acari
Araneae
Chilop.
Coleop.
Collem.
Diplop.
Diptera
Hemip.
Hyme.
Isop
Opilio.
Orthop.
Pseudoscor.
Psocop.
Thysan.
Unindentifyed
NTPF1
25/09/15 4 2 3 1 1 16 2 1 1 2 31
14/10/15 1 2 4 5 2 12
25/11/15 9 1 13 117 12 17 2 1 2 1 175
27/01/16 10 7 52 8 1 1 1 1 7 81
NTPF2
25/09/15 9 4 7 4 16 3 40
14/10/15 2 2 10 5 1 8 20
25/11/15 10 4 1 5 23 12 7 1 7 63
27/01/16 10 2 35 1 2 1 4 51
NTPF3
25/09/15 6 4 1 3 25 1 39
14/10/15 1 2 2 1 6 1 12
25/11/15 5 2 1 2 27 5 1 2 1 45
27/01/16 3 1 1 2 8 2 2 1 1 14 21
LTPF1
25/09/15 52 4 5 3 2 1 27 1 1 3 96
14/10/15 8 6 7 1 8 1 4 10 35
25/11/15 27 2 8 44 1 14 26 1 2 44 125
27/01/16 12 2 8 1 3 1 21 27
LTPF2
25/09/15 2 16 1 3 16 1 38
14/10/15 11 2 1 47 9 1 3 1 74
25/11/15 16 3 8 73 4 2 3 1 110
27/01/16 44 1 4 40 2 1 2 2 14 96LTPF3
25/09/15 15 7 67 1 36 1 3 127
14/10/15 2 15 1 6 1 1 25
25/11/15 4 1 5 1 2 2 13
27/01/16 17 7 2 47 1 1 2 22 77
HTPF1
25/09/15 5 6 10 1 24 2 1 48
14/10/15 1 35 1 1 4 4 1 2 47
25/11/15 1 7 1 6 9
27/01/16 3 1 3 35 10 1 6 53
HTPF2
25/09/15 3 2 8 1 25 1 40
14/10/15 3 1 2 3 1 6 1 1 18
25/11/15 2 4 22 1 3 17 49
27/01/16 10 98 3 2 1 4 114
HTPF3
25/09/15 2 9 3 7 1 35 2 4 59
14/10/15 2 1 1 3 5 12
25/11/15 6 4 203 84 46 25 3 25 371
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27/01/16 14 1 6 73 1 1 2 96
UNBPF1
14/10/15 0
25/11/15 3 6 3 4 14 1 4 31
27/01/16 24 1 1 22 3 2 1 2 1 2 57
UNBPF2
14/10/15 0
25/11/15 2 3 13 29 29 22 2 2 16 102
27/01/16 4 4 14 2 1 5 25
UNBPF3
14/10/15 0
25/11/15 3 8 15 5 22 1 3 54
27/01/16 2 3 2 1 5 8
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Appendix2.SampleresultstransformationwiththeNaperianLogarithm.
Mon
th
Plot TrapNumber TotalAbundance ln(TotalAbundance)
Octob
er
NT
Trap1 12 2.4849
Trap2 20 2.9957
Trap3 12 2.4849
LT
Trap1 35 3.5553
Trap2 74 4.3041
Trap3 25 3.2189
HT
Trap1 47 3.8501
Trap2 18 2.8904
Trap3 12 2.4849
Nov
embe
r
NT
Trap1 175 5.1648
Trap2 63 4.1431
Trap3 45 3.8067
LT
Trap1 125 4.8283
Trap2 110 4.7005
Trap3 13 2.5649
HT
Trap1 9 2.1972
Trap2 49 3.8918
Trap3 371 5.9162
Janu
ary
NT
Trap1 81 4.3944
Trap2 51 3.9318
Trap3 21 3.0445
LT
Trap1 27 3.2958
Trap2 96 4.5643
Trap3 77 4.3438
HT
Trap1 53 3.9703
Trap2 114 4.7362
Trap3 96 4.5643
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Appendix3.Pearsoncorrelationtestresultsregardingorders. Araneae
Chilopoda
Coleoptera
Collembola
Diplopoda
Diptera
Hem
iptera
Hym
enopte
ra
Isopoda
Opilionidea
Orthoptera
Pseudoscor
piones
Psocoptera
October
Acarina
PearsonCorrelation
0,971
.a -0,59 0,943
.a -0,08
2
.a 0,975 .a .a .a 0,915
-1,000**
Sig.(2-tailed) 0,153
. 0,598
0,216
. 0,948
. 0,142 . . . 0,265
.
N 3 1 3 3 0 3 1 3 2 1 0 3 2
Araneae
PearsonCorrelation .a -0,38
1
0,836
.a -0,31
7
.a 1,000*
.a .a .a 0,792
-1,000**
Sig.(2-tailed) . 0,751
0,369
. 0,795
. 0,011 . . . 0,418
.
N 1 3 3 0 3 1 3 2 1 0 3 2
Chilopo
da
PearsonCorrelation .a .a .a .a .a .a .a .a .a .a .a
Sig.(2-tailed) . . . . . . . . . . .
N 1 1 0 1 1 1 1 0 0 1 1
Coleoptera PearsonCorrelation -
0,826
.a -0,75
6
.a -0,397
.a .a .a -0,86
6
.a
Sig.(2-tailed) 0,382
. 0,454
. 0,74 . . . 0,333
.
N 3 0 3 1 3 2 1 0 3 2
Collembola PearsonCorrelation .a 0,25
5.a 0,846 .a .a .a ,997* -
1,000**
Sig.(2-tailed) . 0,836
. 0,358 . . . 0,048
.
N 0 3 1 3 2 1 0 3 2
Diplopo
da
PearsonCorrelation .a .a .a .a .a .a .a .a
Sig.(2-tailed) . . . . . . . .
N 0 0 0 0 0 0 0 0
Diptera
PearsonCorrelation .a -0,3 .a .a .a 0,327
1,000**
Sig.(2-tailed) . 0,806 . . . 0,788
.
N 1 3 2 1 0 3 2
Hem
ipte
ra
PearsonCorrelation .a .a .a .a .a .a
Sig.(2-tailed) . . . . . .
N 1 1 0 0 1 1
Hym
enopte
ra
PearsonCorrelation .a .a .a 0,803
-1,000**
Sig.(2-tailed) . . . 0,407
.
N 2 1 0 3 2
Isopoda
PearsonCorrelation .a .a .a .a
Sig.(2-tailed) . . . .
N 1 0 2 2
Opilioni
dea
PearsonCorrelation .a .a .a
Sig.(2-tailed) . . .
N 0 1 1
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Orthopt
era
PearsonCorrelation .a .a
Sig.(2-tailed) . .
N 0 0Pseudoscor
piones
PearsonCorrelation -1,000**
Sig.(2-tailed) .
N 2
Novem
ber
Acarina
PearsonCorrelation
-0,90
6
-1,000**
-0,82
7
0,07 .a -0,89
9
.a -0,781
-1,000**
.a .a -0,90
6
-1,000**
Sig.(2-tailed) 0,278
. 0,38 0,956
. 0,289
. 0,429 . . . 0,278
.
N 3 2 3 3 2 3 0 3 2 1 1 3 2
Araneae
PearsonCorrelation 1,000**
0,512
0,359
.a 0,629
.a 0,444 1,000**
.a .a 1,000**
1,000**
Sig.(2-tailed) . 0,658
0,766
. 0,567
. 0,707 . . . 0 .
N 2 3 3 2 3 0 3 2 1 1 3 2
Chilopoda PearsonCorrelation 1,00
0**1,000**
.a 1,000**
.a -1,000**
1,000**
.a .a 1,000**
1,000**
Sig.(2-tailed) . . . . . . . . . . .
N 2 2 1 2 0 2 2 1 1 2 2
Coleoptera PearsonCorrelation -
0,618
.a 0,99 .a ,997* 1,000**
.a .a 0,512
1,000**
Sig.(2-tailed) 0,576
. 0,091
. 0,05 . . . 0,658
.
N 3 2 3 0 3 2 1 1 3 2
Collembol
a
PearsonCorrelation .a -0,5 .a -0,677
1,000**
.a .a 0,359
1,000**
Sig.(2-tailed) . 0,667
. 0,526 . . . 0,766
.
N 2 3 0 3 2 1 1 3 2
Diplopo
da
PearsonCorrelation .a .a .a .a .a .a .a .a
Sig.(2-tailed) . . . . . . . .
N 2 0 2 1 1 0 2 1
Diptera
PearsonCorrelation .a 0,976 1,000**
.a .a 0,629
1,000**
Sig.(2-tailed) . 0,14 . . . 0,567
.
N 0 3 2 1 1 3 2
Hem
ipte
ra
PearsonCorrelation .a .a .a .a .a .a
Sig.(2-tailed) . . . . . .
N 0 0 0 0 0 0
Hym
enopte
ra
PearsonCorrelation -1,000**
.a .a 0,444
-1,000**
Sig.(2-tailed) . . . 0,707
.
N 2 1 1 3 2
Isopoda
PearsonCorrelation .a .a 1,000**
1,000**
Sig.(2-tailed) . . . .
N 1 1 2 2
O p i l i o n i d e aPearsonCorrelation .a .a .a
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Sig.(2-tailed) . . .
N 0 1 1Orthopt
era
PearsonCorrelation .a .a
Sig.(2-tailed) . .
N 1 1
Pseudosc
orpionesPearsonCorrelation 1,000
**Sig.(2-tailed) .
N 2
January
Acarina
PearsonCorrelation
0,983
.a -,997*
-0,43
6
.a -0,95
1
1,000**
-0,561
1,000**
.a .a .a .a
Sig.(2-tailed) 0,119
. 0,046
0,713
. 0,2 . 0,621 . . . . .
N 3 1 3 3 1 3 2 3 2 1 0 2 3
Araneae
PearsonCorrelation .a -0,99
3
-0,59
6
.a -0,99
2
1,000**
-0,397
-1,000**
.a .a .a .a
Sig.(2-tailed) . 0,073
0,593
. 0,081
. 0,74 . . . . .
N 1 3 3 1 3 2 3 2 1 0 2 3
Chilopo
da
PearsonCorrelation .a .a .a .a .a .a .a .a .a .a .a
Sig.(2-tailed) . . . . . . . . . . .
N 1 1 0 1 1 1 1 1 0 1 1
Coleoptera PearsonCorrelation 0,5 .a 0,97
1-1,000**
0,5 .a .a .a .a .a
Sig.(2-tailed) 0,667
. 0,154
. 0,667 . . . . .
N 3 1 3 2 3 2 1 0 2 3
Collembol
a
PearsonCorrelation .a 0,693
.a -0,5 1,000**
.a .a .a .a
Sig.(2-tailed) . 0,512
. 0,667 . . . . .
N 1 3 2 3 2 1 0 2 3
Diplopo
da
PearsonCorrelation .a .a .a .a .a .a .a .a
Sig.(2-tailed) . . . . . . . .
N 1 1 1 0 0 0 1 1
Diptera
PearsonCorrelation -1,000**
0,277 1,000**
.a .a .a .a
Sig.(2-tailed) . 0,821 . . . . .
N 2 3 2 1 0 2 3
Hem
iptera
PearsonCorrelation -1,000**
.a .a .a .a .a
Sig.(2-tailed) . . . . . .
N 2 1 1 0 2 2
Hym
enopte
ra
PearsonCorrelation -1,000**
.a .a .a .a
Sig.(2-tailed) . . . . .
N 2 1 0 2 3
Isop odaPearsonCorrelation .a .a .a .a
Sig.(2-tailed) . . . .
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N 1 0 1 2
Opilionidea PearsonCorrelation .a .a .a
Sig.(2-tailed) . . .
N 0 1 1
Orthopt
era
PearsonCorrelation .a .a
Sig.(2-tailed) . .
N 0 0
Pseudoscor
piones
PearsonCorrelation .a
Sig.(2-tailed) .
N 2
**Correlationissignificantatthe0.01level(2-tailed).*Correlationissignificantatthe0.05level(2-tailed).“a”
Cannotbecomputedbecauseatleastoneofthevariablesisconstant.
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Appendix4.Totalabundanceandecologicalfunctionforeachpitfallandsamplingdate.
Order
Collembola
Hemiptera
Psocoptera
Coleoptera
Thysanura
Isopoda
Coleoptera
Hymenoptera
Chilopoda
Pseudoscorpionida
Coleoptera
Aranea
Opiliones
EcologicalFunction
MicrobialFeeders
PlantFeeders
PlantFeeders
PlantFeeders
Omnivores
Omnivores
Omnivores
Omnivores
Predators
Predators
Predators
Predators
Predators
LTPF1
25/09/15 10 24 2 6 14/10/15 35 1 1 4 4 25/11/15 7 1 27/01/16 35 3 1 1
LTPF2
25/09/15 8 25 1 2 14/10/15 3 1 1 1 6 2 1 25/11/15 22 2 2 17 2 27/01/16 98 1 2
LTPF3
25/09/15 7 2 3 35 9 14/10/15 1 1 5 25/11/15 84 162 25 3 41 4 27/01/16 73 1 6 1
NTPF1
25/09/15 1 1 2 2 16 1 2 14/10/15 4 2 5 25/11/15 117 1 9 2 1 17 1 2 27/01/16 52 1 1 1 1
NTPF2
25/09/15 7 16 4 14/10/15 10 5 1 2 25/11/15 23 4 1 1 7 1 4 27/01/16 35 1 2 2
NTPF3
25/09/15 3 1 25 4 14/10/15 2 2 6 25/11/15 27 2 2 5 1 1 2 27/01/16 8 2 2 1 1 1 1
HTPF1
25/09/15 3 1 1 5 27 1 4 14/10/15 7 1 8 4 6 25/11/15 44 8 1 26 2 2 27/01/16 8 3 1 2
HTPF2
25/09/15 3 16 1 16 14/10/15 47 9 3 1 2 125/11/15 73 1 7 2 1 3 327/01/16 40 2 2 1 3 1
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HTPF3
25/09/15 67 1 36 7 114/10/15 15 1 6 25/11/15 5 1 2 27/01/16 47 1 2 1 2 7
UNBPF114/10/15 25/11/15 3 5 1 1 14 27/01/16 22 1 1 2 1 1 2
UNBPF2
14/10/15 25/11/15 29 7 3 22 2 3 227/01/16 14 1 4
UNBPF3
14/10/15 25/11/15 15 1 22 8 27/01/16 3 1
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Appendix5.Pearsoncorrelationresultsregardingfunctionalgroupswithinmonths.
Date PlantFeeders Omnivores PredatorsOctober
MicrobialFeeders
PearsonCorrelation -0,902 0,945 0,99Sig.(2-tailed) 0,285 0,212 0,091
N 3 3 3PlantFeeders PearsonCorrelation -0,993 -0,954
Sig.(2-tailed) 0,073 0,194N 3 3
Omnivores PearsonCorrelation 0,982Sig.(2-tailed) 0,121
N 3
Novem
ber
MicrobialFeeders
PearsonCorrelation -0,624 -0,457 0,984Sig.(2-tailed) 0,571 0,698 0,113
N 3 3 3PlantFeeders PearsonCorrelation 0,98 -0,752
Sig.(2-tailed) 0,127 0,458N 3 3
Omnivores PearsonCorrelation -0,607Sig.(2-tailed) 0,585
N 3
January
MicrobialFeeders
PearsonCorrelation -0,896 0,993 -0,891Sig.(2-tailed) 0,293 0,073 0,3
N 3 3PlantFeeders PearsonCorrelation -0,941 1,000**
Sig.(2-tailed) 0,219 0,007N 3 3
Omnivores PearsonCorrelation -0,937Sig.(2-tailed) 0,227
N 3**Correlationissignificantatthe0.01level(2-tailed).
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Appendix6.Pearsoncorrelationtestresultsregardingfunctionalgroupswithintreatments.
Treatments PlantFeeders Omnivores Predators
NT
MicrobialFeedersPearsonCorrelation 0,914 0,556 ,999*
Sig.(2-tailed) 0,266 0,625 0,024
N 3 3 3
PlantFeeders
PearsonCorrelation 0,846 0,898
Sig.(2-tailed) 0,359 0,29N 3 3
OmnivoresPearsonCorrelation 0,525
Sig.(2-tailed) 0,648
N 3
LT
MicrobialFeeders
PearsonCorrelation 0,997 0,242 -1,000**
Sig.(2-tailed) 0,053 0,844 0,007N 3 3 3
PlantFeedersPearsonCorrelation 0,161 -,997*
Sig.(2-tailed) 0,897 0,046
N 3 3
Omnivores
PearsonCorrelation -0,231
Sig.(2-tailed) 0,851N 3
HT
MicrobialFeedersPearsonCorrelation 0,538 0,384 -0,181
Sig.(2-tailed) 0,638 0,749 0,884
N 3 3 3
PlantFeeders
PearsonCorrelation 0,985 0,732
Sig.(2-tailed) 0,111 0,478N 3 3
OmnivoresPearsonCorrelation 0,839
Sig.(2-tailed) 0,367N 3
*Correlationissignificantatthe0.05level(2-tailed).**Correlationissignificantatthe0.01level(2-tailed).