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TheeffectsofsoilmoistureonPorcellioscaberssiteselection

Abstract
This study was conducted to see whether soil moisture level would have an effect on the
locationalpreferenceof Porcellioscaber.Asimultaneouschoiceexperimentinwhicha10cmpetri
dish divided into two regions corresponding to dry soil (0% moisture level) and moist soil (40%
moisturelevel)wasusedtodeterminewhetherornotaneffectwouldbepresent.Wemeasuredhow
muchtimeP.scaberspentineachregionandwhicheversidetheP.scaberspentmostitstimewithin
5minuteswastherecordedchoice.Inthefirsttrial,ourresultswerenotsignificantwhere10outof14
P.scaberchosemoistsoil(pvalue=0.109).Inthesecondtrial,againourresultswerenotsignificant,
having13outof18 P.scaber choosingthemoistsite(pvalue=0.059). Considering Porcellio
scabers biological characteristics of lacking a waxy cuticle which makes them
more prone to desiccation (Burton and Burton 2002), as well as prior studies
showing significant values where Porcellio scaber chose the moist environment to
be favourable over dry (Gunn 1987 and Waloff 1941), this implies that our
experiment had design limitations: the number of replicates and time given for
site selection.

Introduction

The Porcellioscaber, mostcommonlyknownasthewoodlice,isasmall,ectothermicinsect


thathasaflat,ellipticalshapedbody.Havingtheabilitytogrowuptoanaveragelengthof17mm,
Porcellioscaberiscomposedofthreeparts:thehead,thorax,andabdomen(Isopoda2013).Itscolours
canrangefrompurelyred,orange,cream,brown,amixtureofredandgrey,oramixtureofyellow
withgrey,butmostcommonlyverydarkgreyincolour(WebbandSillem1906).
The P.scaber originatedfrommainlandEuropeandarecrustaceans.Howeverthroughleaf
matterandhumanhabitation,theyhavetransportedtoplacesallaroundtheworld(WangandSchreiber
1999). Throughmovementofballast,rubble,agriculturalproducts,compostandplants, Porcellio
scaber havebecomeawidelydistributedandanabundantspecies(WangandSchreiber1999).They
canbefoundunderbarks,fallenlogs,leaflitter,dunes,andinsaltmarshesnearthesea,orinforests,
gardensandcellarsinhumanhabitations(HardingandSutton1985).
Thelocationof Porcellioscaber also differsbyseason.Inthesummer, Porcellioscaber are
morelikelytofoundclosertothetopoftreesastheymigratetowardstocanopies(Warburg etal.
1984). This could be accounted forby the thickening oftree canopy in the summer, cutting off
sunlight.Inthewinter,theyarecommonlyfoundnearthetreetrunks,closertothegroundbecausethe
leaflesstreesexposeP.scabertolight(Warburgetal.1984).
Whenexposedtolight, P. Scaber displaysquickmovementinordertofindamoresuitable
environment. Having limiting defence mechanisms against predators, birds and small mammals,
Porcellioscaber willbemostactiveatnight,wherelightintensitiesarelowandriskofpredationis
low.Inthesummer,wheredaysarelonger, Porcellio scaber willbeactivelymovingtodarkhiding
spots,underrocks,leaves,andbarkswhereasinthewinter,whenthenightsarelonger,theywilldepict
verylittlemovement,duetothealreadyfavourabledarkconditions.

Inaddition,duetotheirphysicalattributesoflackingcuticlesontheirexoskeleton,P.scaber
havewaterpermeablecharacteristics(Mckenzien.d.)Theirwaterpermeableexoskeletonwillmake
themmorepronetodesiccation,anexcessivelossofwater.Asaresult,airhumidityplaysahugefactor
thataffectstheirbehaviourandtheirsurvival.ThePorcellioscaberexhibitshygrokinesis,displaying
highlevelsofmovementindryairandalmostnomotionindampconditions(Gunn1937). When
exposedtoverydryconditions, Porcellioscaber willquicklytrytofindamorefavourablemoist
locationinordertoreducewaterlossortorestoretheiroriginalweightbyabsorptionofmoisture
(Edney2008).ThisexplainswhyPorcellioscaberaremoreabundantinwethabitatssuchasforests,
dunes,saltmarshesandbytheseashore.
TheairhumidityhasanaffectonsoilmoistureinwhichthePorcellioscabercanbefound.Low
airhumiditycorrespondstodrysoilandhighhumiditycorrespondstomoistsoil.ThereforePorcellio
scaberwillmorelikelybefoundinmoistsoilratherthandry.Ithasbeenfoundthatin10cmofsoil,
thesoilmoisturepercentageiswithintherange25%70%inOregon(Perry etal. 2012).Because
PorcellioscaberarefoundinOregonandduetothesimilarclimateandgeographicfeaturesitshares
withBritishColumbia,inourexperimentwewillusethisdatainordertodeducewhatisconsidered
moist.
InordertofurtherdeterminePorcellioscaberspreferredhabitat,wewillbestudyingtheeffect
ofsoilmoisture. SinceithasbeenestablishedthatPorcellioscaberexhibitshygrokinesisinorderto
preventdesiccation,ahypothesisthatP.scaberwillspendmoretimeinthedampsoilthandrysoilis
made.
Methods
Simultaneouschoicetestingwasconductedinordertodeterminethepreferredsoilmoisture
levelofPorcellioscaber.Individualorganismswerepresentedachoiceoftworegionswithina10cm

diameterpetridish.Duetotimelimitations,weusedtwoidenticalpetridishesatthesametime.Halfof
thepetridishhadcompletelydrysoil(madewith0mLofwaterand50mLsoil)andtheotherhalf
contained40%moistsoil(madewith20mLofdechlorinatedwaterand50mLsoil).Thesoilwas
leveledtobeequaldepthsof1.00cmforbothregions.Thesemoisturelevelswerechosenbecausethey
fitintherangeofthenaturalhabitatofP.scaberaswellasbeingsignificantlydifferentsothatthedata
extrapolatedismeaningful.Soilmoistureleveldidnotexceed40%inourexperimentthusensuring
thatthephysicalpropertyofsoiltextureremainedconsistentenoughtonotinterferewithlocomotionof
theorganisms.Asmalllaminateddividerseparatedthetwotreatmentsbelowthesurfaceinorderto
preventdiffusionofwaterintotheothertreatmentwhilealsonotimpactingtheabilityofP.scaberto
choose/changeitschoiceoftreatment.
Beforeeachreplicate,onbothsidesofthearena,thetemperatureandthelightintensitywere
keptasconstantaspossibleandmeasuredinordertomonitorextraneousabioticfactorsintheroom.
Temperaturerangedfrom23C to24C inTrial1and25C to26C inTrial2whilelightintensity
rangedfrom421to489luxand448to488lux,respectively.Adicewasrolledbeforeeveryreplicate
inordertorandomizetheorientationofthepetridishduringtheexperiment,witheachnumberfrom1
6correspondingwith6directionsthatwereevenlyspacedoutbetween360degrees.Thesizeofeach
P.scaberwasmeasuredandrecordedtomonitorthepossibilityofbioticfactorsinthechoicebetween
dryandmoistsoil.
Forthefirsttrial,therewere14replicatesrangingfromasizeof0.68cmto1.24cm.Arandom
P.scaberwasselectedfromajar,containingapproximately20,usingaplasticstraw.TheP.scaber
wasplacedatthecentreofthepetridishandassoonasitleftthestraw,thetimerwasstarted.One
person handled the timer for all replicates to minimize experimental bias. We assumed that five
minuteswouldbeenoughtimefortheorganismtoacclimatizeandmakeaselection,andthusafive

minuteperiodwasallotted.Whateverregionitspentthemosttimeinwithinthefiveminuteswasthe
recorded choice. For the second trial, the same methods were used, except that we increased the
numberofreplicatesby4fromn=14ton=18,withbodysizerangingfrom0.63cmto1.23cm.By
enlargingthesamplesizeandnumberofreplicates,wewereabletoincreasetheconfidenceofour
results.
Weanalyzedourdatafrombothtrialsseparatelyandgraphedthefrequencyofchoiceof P.
scaber foreachsoilmoisturelevel.WeusedExcel2013toproduceboththegraphs.Twosamplet
tests were calculated via the web page, <Students ttests (http://www.physics.csbsju.edu/stats/t
test.html)>. Chisquare tests were calculated using the web page, <chisquare tests
(http://www.physics.csbsju.edu/stats/chisquare.html)>.
Results
ThepurposeofthisexperimentwastoassessthepreferenceofPorcellioscaberwithrespectto
soilmoisturelevel.InTrial1,mostorganismsfrequentlychose40%moistsoil(n=10)comparedtosoil
containing0%moisture(n=4),suggestingatrendthatfavoursmorechoicesformoistersoil(Figure1).

Figure 1. Trial 1. Choice of soil moisture by Porcellio scaber measured according to time spent in each
treatment during a period of 5 minutes (n=14).

ThemorefrequentchoicemadebyP.scaberinTrial1wasmoistsoilwhichisevidentlyshown
inFigure1.Basedonthechisquaretest,thedataobtainedinTrial1isnotstatisticallysignificant,due
toachisquarevalueof2.57andapvalueof0.109(>0.05,thealpha).Therefore,accordingtothechi
squaretest,wefailtorejectournullhypothesisthatclaimsequalnumbersofP.scaberwouldchoose
thedrysoiltreatmentandthemoistsoiltreatment.Ontheotherhand,wealsoperformedatwosample
ttestgettingavalueof2.91andapvalueof0.007.Accordingtothetwosamplettestourdatais
statisticallysignificant;sincethepvalueof0.007islessthan0.05,wecanconcludethatthemeansof
thetwogroupsaresignificantlydifferent,andrejectthenullhypothesis.
Figure2alsosuggestsatrendinwhichmoistsoilwasmorefrequentlychosenbytheorganisms
underobservation.Thegraphshowsthat,onceagaininTrial2,thechoiceformoistsoilwasmore
frequentthanthechoicefordrysoil;13organismschosethe40%moistsoil,while5chosethedrysoil
with0%moisture.

Figure 2. Trial 2. Choice of soil moisture by Porcellio scaber measured according to time spent in each
treatment during a period of 5 minutes (n=18).

Increasingthesamplesizefrom14to18,comparativelyfromTrial1toTrial2,increasesthe
numberofreplicatesandthereforeincreasestheconfidenceofourresultsgivingusahigherchisquare
valueof3.56comparedto2.57inTrial1.However,thepvalueforTrial2is0.059,whichisgreater

thanalpha(0.05)by0.009,causingustofailtorejectthenullhypothesisaccordingtothechisquare
testsincethedataisnotstatisticallysignificant.Wealsoperformedatwosamplettestforthesecond
trial,gettingavalueof2.98andapvalueof0.005.Accordingtothetwosamplettest,ourdatais
statisticallysignificant;sincethepvalueof0.005islessthan0.05,wecanconcludethatthemeansof
thetwogroupsaresignificantlydifferent,andrejectthenullhypothesis.
Multipleobservationsweremadeinbothtrials,asidefromthechoicebetweenmoistanddry
soil.P.scaberwouldoftenselectaregion(moistordry),crawltowardstheborderofthepetridish,
andremainstationaryinthatareaforapproximatelyaminute.Anothernotableobservanceexhibitedby
theorganismsshowedthatsmallerorganisms(bodysize<0.80cm)weremoreactiveondriersoilfora
longerperiodoftimebeforeeventuallymovingtothemoistsoil.InTrial1,thefeworganisms(n=4)
thatchosedrysoilovermoisthadasmalleraveragebodysizeof0.875cm,comparedtotheorganisms
thatchosemoistsoil(n=10),whichhadanaveragebodysizeof0.992cm.Similarly,inTrial2,the
numberoforganisms(n=5)thatchosedrysoilhadsmalleraveragebodysizeof0.904cmcomparedto
theorganismsthatchosemoistsoil(n=13),whichhadanaveragebodysizeof0.957cm.Attheendof
the5minutes,12outof14organismswereatthemoistregioninTrial1and15outof18wereatthe
moistregioninTrial2.

Discussion
In this experiment we determined whether soil moisture affect site selection by Porcellio
scaber.Inbothtrial1and2,weobservedmorePorcellioscaberchoosingmoistsoiloverdryhowever
statistically,usingthechisquaredtest,weobtainedpvaluesgreaterthanthealphathereforefailedto
rejectthenullhypothesis.Thereforefromourtesting,Porcellioscaberssiteselectionisnotaffected

bysoilmoisturelevel.OurpredictionwasnotsupportedbyourresultsinbothTrial1and2. We
predictedthat P.scaber wouldchoosethemoistsitemorefrequentlythanthedrysitebecauseof
lackingawaxycuticle;theabilityforP.scabertochooseasitewheretheycanpreservetheirinternal
moisturelevelinordertopreventdesiccationisessential(BurtonandBurton2002).
D.L.Gunn(1987)testedthehumidityreactionsofPorcellioscaberusingaseparatetreatment
testwhereobservationsweremadein15minuteintervals.P.scaberexhibitedlittletonomovementin
the65%humiditysitewhereasin35%humiditytheyshowedrapidquickmovements.Infavourable
siteswherePorcellioscaberarecomfortable,theydisplayedlittledesiretomovewhereasindrysites
wheredesiccationismorelikely,theydisplayedhygrokinesisandmovedfrequentlyinordertofinda
moresuitableenvironment(Gunn1987).Thisexplainstheratioof5:1of Porcellioscaber foundin
moistregiontodryregionandwhytheproportionofmotionlessanimalsrosewithrespecttothe
humidityincrease.
Anothersimilarstudywasconductedwherethehumidityreactionsof Porcellioscaber was
analyzed using a separate treatment test (Waloff 1941). Similar to Gunns experiment, mentioned
above, P.scaber wereexposedtohumiditylevelsbetween50%to80%anditwasfoundthatthey
tendedtomovelesswhenexposedtohighhumidityasallofthetestedP.scaberstayedmotionlessfor
longerperiodsoftimeinthemoistsite.BothstudiesshowsignificantvaluesofmoreP.scaberstaying
motionlessinmoistsitethusbeingmorecomfortableinthemoistsite.However,thetwostudiesdonot
agreewiththeresultsfromourexperimentwhereweobtainedaratioof2.5:1P.scaberchoosingmoist
sitetodrysite,resultinginvaluesthatarenotsignificant.Thismaybebecause,insteadofmeasuring
airhumidity,wemeasuredsoilmoisturelevel.Whilesoilmoistureandairhumiditybothcontributeto
theamountofwaterinanenvironment,airhumiditymayaffecttheinteractionbetweenP.scaberand

theamountofwatertoagreaterextent.Surroundedbyair, P.scaber directlybreathesinthewater


moleculestotheirpseudotrachea;thus,airhumidity,ratherthansoilmoisturemaybeamorepertinent
factor to consider in site selection. Another experimental difference is the time used to observe
movementfortheselectionofsite.Inourexperimentweallottedonly5minutesinsteadof15(Gunn
1987)or1hour(Waloff1941).5minutesmaynotbeenoughtimeforthePorcellioscabertorecover
fromhandlingaswellasmakeasiteselection.
Inourexperiment,thebiologicalvariationwasmaximizedthusmakingthetestedanimalsmore
similartothoseinthenaturalenvironment.Porcellioscaberusedinthisexperimentvariedinsize.In
both trials, we observed that smaller P. scaber tended to be more active and visited the dry soil
treatmentmorefrequently.AsforthelargeP.scabershowedlittlemovementandstayedononespot
foralongerperiodoftime.P.scaberthatchosethedrysoiltreatmenthadanaveragebodysizethat
wasapproximately0.30cmsmallerthanthosethatchosethemoistsoil.Apossiblereasoningforthis
maybebecause P. scaberissoflatinshape,volumewouldhavelittleaffectandthedifferenceof
surfaceareabetweeneach P.scaber willhaveagreaterimpact.Large Porcellioscaber wouldhave
moresurfaceareaexposedtothedryenvironment;thismaymakethemmorepronetodesiccation
comparedtosmaller P.scaber.Also,fromBergmannsrule,itisknownthatterrestrialecthotherms,
tendtobelargerincolderenvironments(Timofeev2001),thuscanexplainwhylargerP.scaberwere
moredrawntowardsthemoistandcoolerenvironment.
Inaddition,extraneousfactors,liketemperaturemayhavehadanaffectonthesiteselection
becausethetestingapparatusdidnotremainconstantandrangedfrom 23C to24C inTrial1and
25C to26C inTrial2. Porcellioscaber haveahighermortalityastemperatureincreasesbecause
theirbodycanloosewatermorequicklyinhottemperatureleadingtodesiccation(Edney1953).Trial

2temperaturewasslightlyhigherthanTrial1thereforecanexplainwhymoreP.scaberchosemoist
soiloverdryinTrial2.
Ourresultsshowedthatthereisnostatisticaldifferenceinthefrequencyofchoicebetweensites
withdrysoilormoistsoil.ThisimpliesthatPorcelliosacberareabletospendtimeequallyonboth
typesofsoilandthuscanliveinawiderangeofhabitats.Awiderhabitatrangemeansthateach
Porcellioscaberwillhavemoreresources:food,andsheltertousesincethepopulationismorespread
outamongdifferenthabitats.Thismeansthateach P.scaber willspendlessenergycompetingfor
nutrientsandwillinvesttheirenergyongrowing,andreproducing.Thisultimatelyincreasesthefitness
ofP.scaber,makingtheoverallpopulationdensityincrease.
Asmentionedprior,ourstudywaslimitedbytime.Wemeasuredthetime Porcellioscaber
spentineachtreatmentwithinafiveminuteintervalwithoutgivinganytimeforP.scabertogetover
handling.Bylookingatpriorsimilarstudies,fiveminuteswerelikelynotenoughtimeforP.scaberto
adjusttothenewenvironmentaswellashandlethestressfrombeingtransportedandselectasite.By
increasingthetimeintervalandbygivingatleast30secondsfortheP.scabertoacclimatizetothenew
environment, we may obtain significantly different results that may allow us to reject the null
hypothesis.
Ultimately,wefoundthat Porcellioscaber arenotaffectedbysoilmoisturelevel.Justby
observation,wecansaythattheyhaveapreferenceformoistsoilasbothourtrialsshowedmoreP.
scaberchoosingmoistsite.However,statistically,bothtrialsshowednotsignificantpvaluestherefore
wefailedtorejectournullhypothesis.Thismeansourpredictionthatmoistsoilwillbepreferredisnot
supportedbyourresults.
Acknowledgements

Thank you to Andrew Jeong, Daniel Tsai, and Harmeet Gill, my group members, and
cofoundersofthisexperiment.IwouldliketorecognizeLynnNorman,ourlaboratoryinstructor,for
hersupportandinsightduringtheseweeksofresearchandeducation.LastlyIwouldliketothank
XueginHuang,thelabtechnician,fortakingcareoftheorganismsaswellandallowingustousethe
equipmentrequiredforexperimentation.

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