COMPARISONOFTHEEARTHWORMSLumbricusterrestris,

Lumbricusrubellus,ANDAporrectodeatrapezoidsPREFERENCE
FORDIFFERENTSOILTYPES,LEAFLITTER,ANDTREE
COMPOSITION

J.T.Field,

D.Ager,M.Berhrmann,J.Dufour,K.GabelmanM.Lehr,L.Palm,

UniversityofMichigan,MI


AbstractThepurposeofthisstudywastodetermineifthereisacorrelationbetweentheabundance
ofworms(Lumbricusterrestris,Lumbricusrubellus,andAporrecteatrapezoidscombined)andavariety
offactorssuchasleaflitterweight,soilpH,soiltype(sandvs.clay),andforesttype(conifervs.
hardwood).WithinColonialPoint(locatednearBurtLakeattheUniversityofMichigansbiological
station),plotswererandomlytakenalongatransectlinewithinthefollowinglocationtypes:
conifer/sandysites,conifer/clay,hardwood/sand,andhardwood/clay.Amountofworms,leaflitter
weight,soilpH,soiltype,andforesttypewereallclassifiedandmeasured.Theresultssuggesteda
significantcorrelationbetweensoiltypeandwormabundance.Thereisalsoapossibletrendbetween
wormabundanceandpH,howeverthedataonforesttypesandleaflittersuggestedthatthese
variablesdidnothaveasignificanteffectonwormabundance.Wormspreferredtheclaysoilswith
higherpH,yettheydidnotseemtohaveasmuchofapreferenceforforesttypesorleaflitterweight.

KeywordsLumbricusterrestris,Lumbricusrubellus,Aporrecteatrapezoids,Soiltype,Foresttype,and
Leaflitter.

INTRODUCTION

Earthwormsareextremelyimportanttothedecompositionofsoilandleaflitter(Bohlen

et.al,1997),playakeyroleinnutrientcycles,andevencreateassimiblecarbohydratesfor
plants(Berlin,1988).Theyareimportanttotreeregenerationandmaintenanceofbiological
diversity(Belote,2008).Foragriculture,wormshaveapositiveeffectuponsoilfertility,and
theirburrowingopenupdeeperlyinglayersintherain(Beddard,1922).Howeverinforests,
earthwormsoftenchangethechemicalandphysicalcompositionofthesoil,andtherefore
changewhatvegetationthesoilcanhold(Satoko,2003).TheyoftendepletetheOhorizon,
whichinturndecreasesnutrient,moisture,androotmediumavailabilityfortrees.Becauseof
this,invasiveearthwormspeciescanhaveadetrimentaleffectonforests(Haleet.al.,2005).
Therefore,thisstudy,whichfocusesonwormabundancesunderdifferentconditions,would
potentiallyhelpbetterpredictandpreventwhereinvasiveearthwormsmayinhabitnext.It
mayalsohelpdetermineratesofdecompositionandnutrientcyclingwithincertainareas.

ColonialPoint,locatedintheUniversityofMichigansBiologicalStation,isaforest

hometoanabundanceofearthwormssuchasAporrecteatrapezoids,Lumbricusterrestris,and
Lumbricusrubellus.ColonialPointhasbeensurveyed,andconifer,northernhardwoodand
otherstandtypeshavebeenidentified(Albert,1987).WithindifferentsectionsofColonial
Point,thereareamultitudeofdifferentsoiltexturetypes.Theseincludesandyandclay
texturesinwhichtheearthwormslive.Wormsare7590%madeofwater,yettheylackthe
abilitytomaintaintheirownwaterbodycontent.Therefore,whenwormsareindriersoil,they
actuallydecreaseinsize.Whilesomewormsareabletotoleratedryconditions,worms
generallylikemoremesicsoilconditions,andtypicallyclaysoilsaremoremesicthansandy

(Bohlenet.al.1996).AnothersoilconditionfavorableforearthwormsisacircumneutralpH,
whileacidicsoilslowerthanapHof6arenotverytolerable(Wherry1924).Alongthosesame
lines,wormbehaviorisgreatlydictatednotonlybythesoilmoisturecontent,andpH,butfood
content,leafcoverandevengasconcentrations(Laverack,1963).L.terrestrishavespecificleaf
eatingpatterns,whileAporrectodeaandL.rubellusaregeneralists(Curryet.al.,2007).Withthe
basicunderstandingofwhatconditionsearthwormsthrivedin,thepurposeofourstudywasto
determineifthereisadifferenceinwormabundanceincorrelationwithlandscapefeatures
suchassoilcomposition(sandyvs.clay),leafliter,andtreespecies(primarilyhardwoodor
coniferdominatedforeststands).Conifertreeshaveacidicneedleswhichcreatehighersoil
acidity.Itispredictedthatclaysoils,withhardwooddominatedleaflitterandforeststands,will
haveasignificantlyhigherabundanceofwormsthanthesandysoilsandorconiferdominated
leaflitterandforeststand.

METHODSANDMATERIALS

WithinColonialPoint,EastandWestplotswererandomlyspaced200metersapart(usinga

randomnumbergenerator)alongonenorthtosouthtransect(Fig.1).Thesepointscontained:primarily
coniferoustreesonsandysoils,coniferoustreesonclaysoil,ornorthernhardwoodtreesoneither
sandyorclaysoils.Withineachlocationwemeasuredvariablessuchastreecover,leaflitter,worm
density,andsoilcomposition.Treecoverwasmeasuredbycreatingfourquadrantssurroundingthe
locationofeachplotpoint(Fig.2).Withineachofthosefourquadrantstwotreeswereclassifiedby
species,andmeasuredfortheirDBHanddistancefromtheendofthetransectpoint.Forourworm
count,fromeachofourplotpoints,tworandomsubplotswereselectedwithin15metersoftheoriginal

plotpoint.Onthesetworandomsubplots,thewormswereshockedtothesurfaceandcountedwithin
a0.25^2metermarker.Toshocktheworms,a250Velectricalgeneratorwasusedfor25minutesusing
4positive/4negative1melectrodes.Leaflitterwasgatheredfromwithinallthe0.25metersquaresin
whichthewormswereshockedandweighed.Theleaveswereseparatedbyspecies,andpercentof
eachtreespeciespresentintheleaflitterwasrecorded.Todeterminesoilcomposition,soilpitswere
dugateachoftheplotpoints.Soilshorizons,andlengthofeachhorizonweremeasured.LaterthepH
ofthesoilswererecorded.KruskalWallis,MannWhitneyU,ChiSquare,andLinearRegressionswere
usedtoanalyzethedatawhereappropriate.

RESULTS

BylookingatFigure3,itisclearthatthereisasignificantdifferencesomewherebetweenthe
fourdifferentsetsofplottypes(hardwoodclay,hardwoodsand,coniferclay,andconifersand).Data
analysisrevealedachisquare=13.874,df=3,andaPvalue=0.003.Next,testsindicatethatthereisa
significantcorrelationbetweenwormdensityandsoiltype(p<0.001)asseeninFigure4.Thistest
indicatedthattherearesignificantlymorewormsinclaysoilsthansandysoils.Thefindingsonforest
typevs.averagewormabundance(Figure5),revealedthattherewasnotasignificantcorrelation
betweenforestcovertypeandwormabundance(p=0.166).
FromFigure6,onecanseethattherewasnotasignificantcorrelationbetweennumberof
wormsandsoilpH(R^2=0.365).However,thereseemstobeatrendinthedataindicatingapositive
relationshipbetweennumberofwormsandpH.Thenexttestrunwastodetermineiftherewasa
relationshipbetweensoilpHandsoiltype.Asignificantcorrelationbetweenthetwowasfound,asclay
tendedtohaveahigherpHthansand(Figure7).Theleaflitterweightvs.averagenumberofworm

correlationwastestedforsignificance(Figure8),howevernosignificantdifferencewasfoundbetween
thetwo(R^2=0.069).

FromFigure9,thepvaluesindicatewhichplottypeshadsignificantdifferencesinworm

abundances.Thereisasignificantdifferenceinwormabundancebetweenhardwoodclayandhardwood
sand(pvalue=0.01),coniferclayandhardwoodsand(pvalue=0.021),conifersandandhardwoodclay
(pvalue=0.009),andconiferclayandconifersand(pvalue=0.018).Thereisnosignificantdifference
betweenconiferclayandhardwoodclay(pvalue=0.163),orhardwoodsandversusconifersand(p
value=0.122).

DISCUSSION

Resultsfromthestudysuggestthatthereisasignificantwormpreferenceforclaysoil,oversandysoil.
WhilepHwasnotseentohaveasignificantcorrelationtowormabundance,apositivetrendwasseen
betweenthetwo(Figure6).Also,claysoilshavebeenfoundtotypicallyhaveahigherpHthanthe
sandysoils,whichmaysuggestthatpHhadaneffectonwormsoilchoice.Thismakesitdifficultto
differentiatewhethersoilpH,ortypewerethereasonforaveragewormabundance.Thereisnota
significantdifferenceinwormabundancebetweenthedifferentforesttypes,suggestingthatthetree
typedoesnothaveasmuchofanimpactontheamountofwormsatagivenpointasthesoildoes.It
maybeconcludedthenthattheacidicneedlesoftheconifertreesdonotimpactwormsasmuchas
acidicsoil.Acidicneedlesofconifertreestypicallymakesoilsmoreacidic,leadingtothequestionof
howmuchofimpactconifertreeneedleshaveontheacidityofthesoil.Also,wormslikesoilsthathold
moremoisture(Bohlenet.al.,1996),andclayretainsmorewaterthansanddoes.Claysoilisgenerally
morecalcareousthansandysoilaswell.CalcareousconditionsindicateahighersoilpH(neutralto
basic),whichwormstendtopreferoveracidicconditions.Finally,theamountofleafliterwasnot
showntoimpactwormabundance.Allinall,ourresultsshowthatsoiltypeandpHhaveaneffecton

abundanceofworms,whileleaflitterweight,andforesttypedoesnot.Fromthesefindingsitmaybe
beneficialinfuturestudiestolookattheeffectsofmoistureandnutrientcontentinsoilonearthworm
abundance(claysoilsareknowntobemoremoistandnutrientrichthansandysoils).Ecologically,by
knowingwhatconditionsearthwormsprefer,nutrientcycleanddecompositionratesmaybemore
easilypredicted.Ourconclusionsmayalsoallownewwaystopredictwhereinvasiveearthwormspecies
choosetoinhabit.Thismayhelppreventforestdamageviaearthwormdestruction.

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Bohlen,P.J.,andEdwards,C.A1996.BiologyandEcologyofEarthworms.ChapmanandHall,Boundary
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HaleCM,FrelichLE,ReichPB(2005)Changesinhardwoodforestunderstoryplantcommunitiesin
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Satoko,M.2003.EffectsofExoticEarthwormsonNorthernHardwoodForestsinNorthAmerica
(UniversityofMinnesota,St.Paul).StudentonlineJournal.

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5:309.


Figure1.NORTHTOSOUTHTRANSECTWITHRANDOMPLOTPOINTS.Reddotscontainplotsamples
usedwithdifferentsoilandtreetypeswithinColonialPoint.

Figure2.TREEQUADRANTSUSEDFORMEASUREMENTS.Centerisplotspoint,withfourquadrants
surroundingthepoint(twotreesperquadrantweremeasuredforDBH,species,anddistancefrom
center).

Averagenumberofworms

40
35
30
25
20
15
10
5
0
HardwoodClay ConiferClay(n=4) HardwoodSand
(n=6)
(n=4)

ConiferSand
(n=4)

ForestandSoiltype

Figure3. AVERAGE NUMBEROFWORMSINTHEHALFMETERPLOTSFORTHE

DIFFERENTFORESTANDSOILCOMBINATIONS.N=6forHardwoodclay.N=4
forConiferClay,HardwoodSand,andConiferSand.KruskalWallIStestused

45

Averagenumberofworms

40
35
30
25
20
15
10
5
0
Clay(n=10)

Sand(n=8)
SoilType

Figure4.THE AVERAGENUMBEROFWORMSFOUNDINHALFMETERPLOTS
WITHCLAYSOIL(N=10)ANDSANDYSOIL(N=8).MannWhitneyUtest
indicatesasignificantdifferencebetweenthetwo(p<0.000).SEMbars

Averagenumberofworms

30
25
20
15
10
5
0
Hardwood(n=10)

Conifer(n=8)

Forestcovertype

Figure5. THEAVERAGENUMBEROFWORMSINCONIFERVERSUSHARDWOOD
FORESTS.theaveragenumberofwormsfoundsinhalfmeterplotsinhardwood
dominatedareas andconiferdominatedareas.MannWhitneyUtestshowno
significantdifferencebetweenthetwo.SEMbarspresent.

Averagenumberofworms

60

R=0.3647

50
40
30
20
10
0
0

SoilpH

Figure6.RELATIONSHIPBETWEENPHOFTHESOILANDTHENUMBEROFWORMSINTHEMEASURED
HALFMETERPLOT.Thecorrelationisnotsignificant,althoughthereappearstobeatrendoccuring.
LinearRegressionusedforanalysis.

7
6
SoilpH

5
4
3
2
1
0
Sand

Clay
SoilType

Figure7.COMPARISONOFSOILTYPEVS.SOILPH.MannWhitneyUtestusedforanalysis.SEMbars
present.

60
Averagenumberofworms

R=0.0686
50
40
30
20
10
0
0

50

100

150

200

250

300

Leaflitterweight(g)

Figure8.LEAFLITTERWEIGHTVS.NUMBEROFWORMS.Therelationshipbetweentheweightoftheleaf
littertothenumberofwormsineachhalfmeterplotisshown.Nosignificantrelationshipisseen(R^2=
0.0686).Linearregressionusedforanalysis.

Hardwood
Hardwood
Conifer

Clay
Sand
Clay
HardwoodSand
0.01

0.021

ConiferClay
0.163
0.122
0.018
ConiferSand
0.009

Figure9.PVALUETABLEWITHDIFFERENTSITETYPES.Tableofpvaluesindicatingwhetherthereisa
significantdifferencebetweenthethreedifferentsitetypes.Theboldedpvaluesindicateasignificant
differentinwormabundance.