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Earthworm
FromWikipedia,thefreeencyclopedia

Anearthwormisatubeshaped,segmentedwormfoundinthe
phylumAnnelida.Earthwormsarecommonlyfoundlivinginsoil, Earthworm
feedingonliveanddeadorganicmatter.Anearthworm'sdigestive
systemrunsthroughthelengthofitsbody.Itconductsrespiration
throughitsskin.Ithasadoubletransportsystemcomposedof
coelomicfluidthatmoveswithinthefluidfilledcoelomanda
simple,closedbloodcirculatorysystem.Ithasacentralanda
peripheralnervoussystem.Thecentralnervoussystemconsistsof
twogangliaabovethemouth,oneoneitherside,connectedtoa
nervecordrunningbackalongitslengthtomotorneuronsand
sensorycellsineachsegment.Largenumbersofchemoreceptorsare
concentratednearitsmouth.Circumferentialandlongitudinal
musclesontheperipheryofeachsegmentenablethewormtomove.
Similarsetsofmuscleslinethegut,andtheiractionsmovethe
digestingfoodtowardtheworm'sanus.[2]
Aspeciesofearthwormwithawell
Earthwormsarehermaphroditeseachindividualcarriesbothmale
developedclitellum
andfemalesexorgans.Theylackeitheraninternalskeletonor
exoskeleton,butmaintaintheirstructurewithfluidfilledcoelom Scientificclassification
chambersthatfunctionasahydrostaticskeleton.
Kingdom: Animalia
"Earthworm"isthecommonnameforthelargestmembersof Phylum: Annelida
Oligochaeta(whichiseitheraclassorasubclassdependingonthe
author).Inclassicalsystems,theywereplacedintheorder Class: Oligochaeta
Opisthopora,onthebasisofthemaleporesopeningposteriortothe Subclass: Oligochaeta(butdisputed,
femalepores,thoughtheinternalmalesegmentsareanteriortothe
seetaxonomy) or
female.Theoreticalcladisticstudieshaveplacedthem,instead,inthe
suborderLumbricinaoftheorderHaplotaxida,butthismayagain Haplotaxida
soonchange.Folknamesfortheearthworminclude"dewworm",
Order: Megadrilacea
"rainworm","nightcrawler",and"angleworm"(duetoitsuseas
fishingbait). Suborder: Lumbricina+
Moniligastrida
Largerterrestrialearthwormsarealsocalledmegadriles(which
translatesto"bigworms"),asopposedtothemicrodriles("small NODCv.8.0,1996[1]
worms")inthesemiaquaticfamiliesTubificidae,Lumbricidae,and Families
Enchytraeidae,amongothers.Themegadrilesarecharacterizedby
havingadistinctclitellum(whichismoreextensivethanthatof
microdriles)andavascularsystemwithtruecapillaries. Acanthodrilidae
Ailoscolecidae
Earthwormsarefarlessabundantindisturbedenvironmentsandare
Almidae
typicallyactiveonlyifwaterispresent.[3]
Benhamiinae
Criodrilidae
Diplocardiinae
Contents
Eudrilidae
1 Anatomy Exxidae
1.1 Formandfunction Glossoscolecidae
1.2 Nervoussystem
1.2.1 Centralnervoussystem Hormogastridae
1.2.2 Peripheralnervoussystem Kynotidae

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1.2.3 Sympatheticnervoussystem Lumbricidae

1.2.4 Movement
1.3 Senses Lutodrilidae
1.3.1 Photosensitivity Megascolecidae
1.4 Digestivesystem Microchaetidae
1.5 Circulatorysystem
1.6 Excretorysystem Moniligastridae
1.7 Respiration Ocnerodrilidae
1.8 Reproduction Octochaetidae
2 Regeneration
3 Locomotionandimportancetosoil Octochaetinae
4 Benefits Sparganophilidae
5 Asaninvasivespecies Tumakidae
6 Specialhabitats
7 Ecology
8 Economicimpact
9 Taxonomyanddistribution
10 Seealso
11 References
11.1 Referencedworks
12 Furtherreading
13 Externallinks
13.1 General
13.2 Academic
13.3 Agricultureandecology
13.4 Wormfarming
13.5 Forchildren

Anatomy
Formandfunction

Dependingonthespecies,anadult
earthwormcanbefrom10mm
(0.39in)longand1mm(0.039in)
wideto3m(9.8ft)longandover
25mm(0.98in)wide,butthe
typicalLumbricusterrestrisgrows
toabout360mm(14in)long.[4]

Fromfronttoback,thebasicshape
oftheearthwormisacylindrical
tube,dividedintoaseriesof
segments(calledmetamerisms)that
compartmentalizethebody.
Groovescalled"furrows"are
generally[5]externallyvisibleonthe
bodydemarkingthesegments Earthwormhead
dorsalporesandnephridiopores
exudeafluidthatmoistensandprotectstheworm'ssurface,allowingittobreathe.Exceptforthemouthand
analsegments,eachsegmentcarriesbristlelikehairscalledlateralsetae[6]usedtoanchorpartsofthebody

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duringmovement[7]speciesmayhavefourpairsofsetaeoneachsegmentormorethaneightsometimes
formingacompletecircleofsetaepersegment.[6]Specialventralsetaeareusedtoanchormatingearthworms
bytheirpenetrationintothebodiesoftheirmates.

Generally,withinaspecies,thenumberofsegmentsfoundisconsistentacrossspecimens,andindividualsare
bornwiththenumberofsegmentstheywillhavethroughouttheirlives.Thefirstbodysegment(segment
number1)featuresboththeearthworm'smouthand,overhangingthemouth,afleshylobecalledthe
prostomium,whichsealstheentrancewhenthewormisatrest,butisalsousedtofeelandchemicallysensethe
worm'ssurroundings.Somespeciesofearthwormcanevenusetheprehensileprostomiumtograbanddrag
itemssuchasgrassesandleavesintotheirburrow.

Anadultearthwormdevelopsabeltlikeglandularswelling,calledtheclitellum,whichcoversseveral
segmentstowardthefrontpartoftheanimal.Thisispartofthereproductivesystemandproduceseggcapsules.
Theposteriorismostcommonlycylindricalliketherestofthebody,butdependingonthespecies,mayalsobe
quadrangular,octagonal,trapezoidal,orflattened.Thelastsegmentiscalledtheperiprocttheearthworm's
anus,ashortverticalslit,isfoundonthissegment.[6]

Theexteriorofanindividualsegmentisathincuticleoverskin,commonlypigmentedredtobrown,whichhas
specializedcellsthatsecretemucusoverthecuticletokeepthebodymoistandeasemovementthroughsoil.
Undertheskinisalayerofnervetissue,andtwolayersofmusclesathinouterlayerofcircularmuscle,anda
muchthickerinnerlayeroflongitudinalmuscle.[8]Interiortothemusclelayerisafluidfilledchambercalleda
coelom[9]thatbyitspressurizationprovidesstructuretotheworm'sbonelessbody.Thesegmentsareseparated
fromeachotherbytransversedividingwallscalled"septa"(singular"septum")[10]thatareperforated,allowing
thecoelomicfluidtopassbetweensegments.[11]Apairofstructurescallednephrostomesarelocatedatthe
backofeachseptumanephrictubuleleadsfromeachnephrostomethroughtheseptumandintothefollowing

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segment.Thistubulethenleadstothemainbodyfluidfilteringorgan,thenephridiumormetanephridium,
whichremovesmetabolicwastefromthecoelomicfluidandexpelsitthroughporescallednephridioporeson
theworm'ssidesusuallytwonephridia(sometimesmore)arefoundinmostsegments.[12]Atthecenterofa
wormisthedigestivetract,whichrunsstraightthroughfrommouthtoanuswithoutcoiling,andisflanked
aboveandbelowbybloodvessels(thedorsalbloodvesselandtheventralbloodvesselaswellasasubneural
bloodvessel)andtheventralnervecord,andissurroundedineachsegmentbyapairofpallialbloodvessels
thatconnectthedorsaltothesubneuralbloodvessels.

ManyearthwormscanejectcoelomicfluidthroughporesinthebackinresponsetostressAustralian
Didymogastersylvaticus(knownasthe"bluesquirterearthworm")cansquirtfluidashighas30cm(12in).[11]

Nervoussystem

Theearthworm'snervoussystemhasthreeparts:thecentral
nervoussystem(CNS),peripheralnervoussystemandthe
sympatheticnervoussystem.[13][14]

Centralnervoussystem

TheCNSconsistsofabilobedbrain(cerebralganglia,or
suprapharyngealganglia),subpharyngealganglia,circum
pharyngealconnectivesandaventralnervecord.

Earthworms'brainsconsistofapairofpearshapedcerebral Nervoussystemoftheanteriorendofan
ganglia.Thesearelocatedinthedorsalsideofthealimentary earthworm
canalinthethirdsegment,inagroovebetweenthebuccal
cavityandpharynx.

Apairofcircumpharyngealconnectivesfromthebrainencirclethepharynxandthenconnectwithapairof
subpharyngealganglialocatedbelowthepharynxinthefourthsegment.Thisarrangementmeansthebrain,
subpharyngealgangliaandthecircumpharyngealconnectivesformanerveringaroundthepharynx.

Theventralnervecord(formedbynervecellsandnervefibres)beginsatthesubpharyngealgangliaand
extendsbelowthealimentarycanaltothemostposteriorbodysegment.Theventralnervecordhasaswelling,
organglion,ineachsegment,i.e.asegmentalganglion,whichoccursfromthefifthtothelastsegmentofthe
body.Therearealsothreegiantaxons,onemedialgiantaxon(MGA)andtwolateralgiantaxons(LGAs)on
themiddorsalsideoftheventralnervecord.TheMGAis0.07mmindiameterandtransmitsinananterior
posteriordirectionatarateof32.2m/s.TheLGAsareslightlywiderat0.05mmindiameterandtransmitina
posterioranteriordirectionat12.6m/s.ThetwoLGAsareconnectedatregularintervalsalongthebodyand
arethereforeconsideredonegiantaxon.[15][16]

Peripheralnervoussystem

Eighttotennervesarisefromthecerebralgangliatosupplytheprostomium,buccalchamberand
pharynx.
Threepairsofnervesarisefromthesubpharyangealgangliatosupplythe2nd,3rdand4thsegment.
Threepairsofnervesextendfromeachsegmentalgangliatosupplyvariousstructuresofthesegment.

Sympatheticnervoussystem

Thesympatheticnervoussystemconsistsofnerveplexusesintheepidermisandalimentarycanal.(Aplexusa
webofnervecellsconnectedtogetherinatwodimensionalgrid.)Thenervesthatrunalongthebodywallpass
betweentheoutercircularandinnerlongitudinalmusclelayersofthewall.Theygiveoffbranchesthatform

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theintermuscularplexusandthesubepidermalplexus.Thesenervesconnectwiththecircumpharyngeal
connective.

Movement

Onthesurface,crawlingspeedvariesbothwithinandamongindividuals.Earthwormscrawlfasterprimarilyby
takinglonger"strides"andagreaterfrequencyofstrides.LargerLumbricusterrestriswormscrawlatagreater
absolutespeedthansmallerworms.Theyachievethisbytakingslightlylongerstridesbutwithslightlylower
stridefrequencies.[17]

Touchinganearthworm,whichcausesa"pressure"responseaswellas(often)aresponsetothedehydrating
qualityofthesaltonhumanskin(toxictoearthworms),stimulatesthesubepidermalnerveplexuswhich
connectstotheintermuscularplexusandcausesthelongitudinalmusclestocontact,therebythewrithing
movementswhenwepickupanearthworm.ThisbehaviourisareflexanddoesnotrequiretheCNSitoccurs
evenifthenervecordisremoved.Eachsegmentoftheearthwormhasitsownnerveplexus.Theplexusofone
segmentisnotconnecteddirectlytothatofadjacentsegments.Thenervecordisrequiredtoconnectthe
nervoussystemsofthesegments.[18]

Thegiantaxonscarrythefastestsignalsalongthenervecord.Theseareemergencysignalsthatinitiatereflex
escapebehaviours.Thelargerdorsalgiantaxonconductssignalsthefastest,fromthereartothefrontofthe
animal.Iftherearofthewormistouched,asignalisrapidlysentforwardscausingthelongitudinalmusclesin
eachsegmenttocontract.Thiscausesthewormshortenveryquicklyasanattempttoescapefromapredatoror
otherpotentialthreat.Thetwomedialgiantaxonsconnectwitheachotherandsendsignalsfromthefrontto
therear.Stimulationofthesecausestheearthwormtoveryquicklyretreat(perhapscontractingintoitsburrow
toescapeabird).

Thepresenceofanervoussystemisessentialforananimaltobeabletoexperiencenociceptionorpain.
However,otherphysiologicalcapacitiesarealsorequiredsuchasopioidsensitivityandcentralmodulationof
responsesbyanalgesics.[19]Enkephalinandendorphinlikesubstanceshavebeenfoundinearthworms.
Injectionsofnaloxone(anopioidantagonist)inhibittheescaperesponsesofearthworms.Thisindicatesthat
opioidsubstancesplayaroleinsensorymodulation,similartothatfoundinmanyvertebrates.[20]

Senses

Photosensitivity

Earthwormsdonothaveeyes(althoughsomewormsdo),however,theydohavespecialisedphotosensitive
cellscalled"lightcellsofHess".Thesephotoreceptorcellshaveacentralintracellularcavity(phaosome)filled
withmicrovilli.Aswellasthemicrovilli,thereareseveralsensoryciliainthephaosomewhicharestructurally
independentofthemicrovilli.[21]Thephotoreceptorsaredistributedinmostpartsoftheepidermisbutaremore
concentratedonthebackandsidesoftheworm.Arelativelysmallnumberoccurontheventralsurfaceofthe
1stsegment.Theyaremostnumerousintheprostomiumandreduceindensityinthefirstthreesegmentsthey
areveryfewinnumberpastthethirdsegment.[18]

Digestivesystem

Thegutoftheearthwormisastraighttubewhichextendsfromtheworm'smouthtoitsanus.Itisdifferentiated
intoabuccalcavity(generallyrunningthroughthefirstoneortwosegmentsoftheearthworm),pharynx
(runninggenerallyaboutfoursegmentsinlength),esophagus,crop,gizzard(usually)andintestine.[22]

Foodentersinthemouth.Thepharynxactsasasuctionpumpitsmuscularwallsdrawinfood.Inthepharynx,
thepharyngealglandssecretemucus.Foodmovesintotheesophagus,wherecalcium(fromthebloodand
ingestedfrompreviousmeals)ispumpedintomaintainproperbloodcalciumlevelsinthebloodandfoodpH.
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Fromtherethefoodpassesintothecropandgizzard.Inthegizzard,strongmuscularcontractionsgrindthe
foodwiththehelpofmineralparticlesingestedalongwiththefood.Oncethroughthegizzard,foodcontinues
throughtheintestinefordigestion.TheintestinesecretesPepsintodigestproteins,Amylasetodigest
polysaccharides,Cellulasetodigestcellulose,andlipasetodigestfats.[2]Earthwormsuseinadditiontothe
digestiveproteinsaclassofsurfaceactivecompoundscalleddrilodefensins,whichhelpdigestplant
material.[23]Insteadofbeingcoiledlikeamammalianintestine,anearthworm'sintestineincreasessurfacearea
toincreasenutrientabsorptionbyhavingmanyfoldsrunningalongitslength.Theintestinehasitsownpairof
musclelayerslikethebody,butinreverseorderaninnercircularlayerwithinanouterlongitudinallayer.[24]
Thegizzardisasomewhatofauniqueorgantothewormasitisfoundinveryfewanimals.

Circulatorysystem

Theearthwormhasadualcirculatorysysteminwhichboththecoelomicfluidandaclosedcirculatorysystem
carrythefood,waste,andrespiratorygases.Theclosedcirculatorysystemhasfivemainbloodvessels:the
dorsal(top)vessel,whichrunsabovethedigestivetracttheventral(bottom)vessel,whichrunsbelowthe
digestivetractthesubneuralvessel,whichrunsbelowtheventralnervecordandtwolateroneuralvesselson
eithersideofthenervecord.[25]Thedorsalvesselmovesthebloodforward,whiletheotherfourlongitudinal
vesselscarrythebloodrearward.Insegmentsseventhrougheleven,apairofaorticarchesringsthecoelomand
actsashearts,pumpingthebloodtotheventralvesselthatactsastheaorta.Thebloodconsistsofameboidcells
andhemoglobindissolvedintheplasma.Thesecondcirculatorysystemderivesfromthecellsofthedigestive
systemthatlinethecoelom.Asthedigestivecellsbecomefull,theyreleasenonlivingcellsoffatintothe
fluidfilledcoelom,wheretheyfloatfreelybutcanpassthroughthewallsseparatingeachsegment,moving
foodtootherpartsandassistinwoundhealing.[26]

Excretorysystem

Theexcretorysystemcontainsapairofnephridiaineverysegment,exceptforthefirstthreeandthelast
ones.[27]Thethreetypesofnephridiaare:integumentary,septal,andpharyngeal.Theintegumentarynephridia
lieattachedtotheinnersideofthebodywallinallsegmentsexceptthefirsttwo.Theseptalnephridiaare
attachedtobothsidesoftheseptabehindthe15thsegment.Thepharyngealnephridiaareattachedtofourth,
fifthandsixthsegments.[27]Thewasteinthecoelomfluidfromaforwardsegmentisdrawninbythebeating
ofciliaofthenephrostome.Fromthereitiscarriedthroughtheseptum(wall)viaatubewhichformsaseriesof
loopsentwinedbybloodcapillariesthatalsotransferwasteintothetubuleofthenephrostome.Theexcretory
wastesarethenfinallydischargedthroughaporeontheworm'sside.[28]

Respiration

Earthwormshavenospecialrespiratoryorgans.Gasesareexchangedthroughthemoistskinandcapillaries,
wheretheoxygenispickedupbythehemoglobindissolvedinthebloodplasmaandcarbondioxideisreleased.
Water,aswellassalts,canalsobemovedthroughtheskinbyactivetransport.

Reproduction

Matingoccursonthesurface,mostoftenatnight.Earthwormsarehermaphroditesthatis,theyhavebothmale
andfemalesexualorgans.Thesexualorgansarelocatedinsegments9to15.Earthwormshaveoneortwopairs
oftestescontainedwithinsacs.Thetwoorfourpairsofseminalvesiclesproduce,storeandreleasethesperm
viathemalepores.Ovariesandoviductsinsegment13releaseeggsviafemaleporesonsegment14,while
spermisexpelledfromsegment15.Oneormorepairsofspermathecaearepresentinsegments9and10
(dependingonthespecies)whichareinternalsacsthatreceiveandstorespermfromtheotherwormduring
copulation.Asaresult,segment15ofonewormexudesspermintosegments9and10withitsstoragevesicles
ofitsmate.Somespeciesuseexternalspermatophoresforspermtransfer.

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InHormogastersamniticaandHormogasterelisaetranscriptomeDNA
librariesweresequencedandtwosexpheromones,Attractinand
Temptin,weredetectedinalltissuesamplesofbothspecies.[29]Sex
pheromonesareprobablyimportantinearthwormsbecausetheylivein
anenvironmentwherechemicalsignalingmayplayacrucialrolein
attractingapartnerandinfacilitatingoutcrossing.Outcrossingwould
providethebenefitofmaskingtheexpressionofdeleteriousrecessive
mutationsinprogeny.[30](Alsoseecomplemenation.)
Earthwormcopulation
Copulationandreproductionareseparateprocessesinearthworms.The
matingpairoverlapfrontendsventrallyandeachexchangesspermwith
theother.Theclitellumbecomesveryreddishtopinkishincolor.Some
timeaftercopulation,longafterthewormshaveseparated,theclitellum
(behindthespermathecae)secretesmaterialwhichformsaringaround
theworm.Thewormthenbacksoutofthering,andasitdoesso,it
injectsitsowneggsandtheotherworm'sspermintoit.Astheworm
slipsoutofthering,theendsofthecocoonsealtoformavaguely
lemonshapedincubator(cocoon)inwhichtheembryonicworms
develop.Theyemergeassmall,butfullyformedearthworms,butlack
theirsexstructures,whichdevelopinabout60to90days.Theyattain EarthwormcocoonsfromL.terrestris
fullsizeinaboutoneyear.Scientistspredictthattheaveragelifespan
underfieldconditionsisfourtoeightyears,whilemostgardenvarieties
liveonlyonetotwoyears.Severalcommonearthwormspeciesare
mostlyparthenogenetic.

Amonglumbricidearthworms,parthenogenesisarosefromsexual
relativesmanytimes.[31]ParthenogenesisinsomeAporrectodea
trapezoideslineagesarose6.4to1.1millionyearsagofromsexual
ancestors.[32]

Regeneration
AnearthwormcocoonfromL.
Earthwormshavetheabilitytoregeneratelostsegments,butthisability
rubellus
variesbetweenspeciesanddependsontheextentofthedamage.
Stephenson(1930)devotedachapterofhismonographtothistopic,
whileG.E.Gatesspent20yearsstudyingregenerationinavarietyofspecies,butbecauselittleinterestwas
shown,Gates(1972)onlypublishedafewofhisfindingsthat,nevertheless,showitistheoreticallypossibleto
growtwowholewormsfromabisectedspecimenincertainspecies.Gatessreportsincluded:

Eiseniafetida(Savigny,1826)withheadregeneration,inananteriordirection,possibleateach
intersegmentallevelbacktoandincluding23/24,whiletailswereregeneratedatanylevelsbehind20/21,
i.e.,twowormsmaygrowfromone.[33]
LumbricusterrestrisLinnaeus,1758replacinganteriorsegmentsfromasfarbackas13/14and16/17but
tailregenerationwasneverfound.
PerionyxexcavatusPerrier,1872readilyregeneratedlostpartsofthebody,inananteriordirectionfrom
asfarbackas17/18,andinaposteriordirectionasfarforwardas20/21.
LampitomauritiiKinberg,1867withregenerationinanteriordirectionatalllevelsbackto25/26andtail
regenerationfrom30/31headregenerationwassometimesbelievedtobecausedbyinternalamputation
resultingfromSarcophagasp.larvalinfestation.
CriodriluslacuumHoffmeister,1845alsohasprodigiousregenerativecapacitywithheadregeneration
fromasfarbackas40/41.[34]

AnunidentifiedTasmanianearthwormshowngrowingareplacementheadhasbeenreported.[35]

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Locomotionandimportancetosoil
Earthwormstravelundergroundbythemeansofwavesofmuscular
contractionswhichalternatelyshortenandlengthenthebody(peristalsis).The
shortenedpartisanchoredtothesurroundingsoilbytinyclawlikebristles
(setae)setalongitssegmentedlength.Inallthebodysegmentsexceptthefirst,
lastandclitellum,thereisaringofSshapedsetaeembeddedintheepidermal
pitofeachsegment(perichaetine).Thewholeburrowingprocessisaidedbythe
secretionoflubricatingmucus.Wormscanmakegurglingnoisesunderground
whendisturbedasaresultoftheirmovementthroughtheirlubricatedtunnels.
Earthwormsmovethroughsoilbyexpandingcreviceswithforcewhenforces
aremeasuredaccordingtobodyweight,hatchlingscanpush500timestheir
ownbodyweightwhereaslargeadultscanpushonly10timestheirownbody
weight.[36]

Earthwormsworkasbiological"pistons"forcingairthroughthetunnelsasthey
move.Thusearthwormactivityaeratesandmixesthesoil,andisconduciveto Closeupofanearthwormin
mineralizationofnutrientsandtheiruptakebyvegetation.Certainspeciesof gardensoil
earthwormcometothesurfaceandgrazeonthehigherconcentrationsof
organicmatterpresentthere,mixingitwiththemineralsoil.Becauseahigh
leveloforganicmattermixingisassociatedwithsoilfertility,anabundanceofearthwormsisgenerally
consideredbeneficialbytheorganicgardener.Infact,aslongagoas1881CharlesDarwinwrote:"Itmaybe
doubtedwhethertherearemanyotheranimalswhichhaveplayedsoimportantapartinthehistoryofthe
world,ashavetheselowlyorganizedcreatures."[37]

Benefits
Themajorbenefitsofearthwormactivitiestosoilfertilitycanbesummarizedas:

Biological:Inmanysoils,earthwormsplayamajorroleintheconversionoflargepiecesoforganic
matterintorichhumus,thusimprovingsoilfertility.Thisisachievedbytheworm'sactionsofpulling
belowthesurfacedepositedorganicmattersuchasleaffallormanure,eitherforfoodortoplugit's
burrow.Onceintheburrow,thewormwillshredtheleafandpartiallydigestitandmingleitwiththe
earth.Wormcasts(seebottomright)cancontain40percentmorehumusthanthetop9"(23cm)ofsoil
inwhichthewormisliving.[3]

Chemical:Inadditiontodeadorganicmatter,theearthwormalso
ingestsanyothersoilparticlesthataresmallenoughincluding
sandgrainsupto1/20ofaninch(1.25mm)intoitsgizzard,
whereinthoseminutefragmentsofgritgrindeverythingintoa
finepastewhichisthendigestedintheintestine.Whentheworm
excretesthisintheformofcasts,depositedonthesurfaceor
deeperinthesoil,mineralsandplantnutrientsarechangedtoan
accessibleformforplantstouse.InvestigationsintheUnited
Statesshowthatfreshearthwormcastsarefivetimesricherin
availablenitrogen,seventimesricherinavailablephosphates,and Faecesinformofcasts
11timesricherinavailablepotassiumthanthesurroundingupper
6inches(150mm)ofsoil.Inconditionswherehumusisplentiful,
theweightofcastsproducedmaybegreaterthan4.5kg(10lb)perwormperyear.[3]
Physical:Theearthworm'sburrowingcreatesamultitudeofchannelsthroughthesoilandisofgreat
valueinmaintainingthesoilstructure,enablingprocessesofaerationanddrainage.Permacultureco
founderBillMollisonpointsoutthatbyslidingintheirtunnels,earthworms"actasaninnumerablearmy
ofpistonspumpingairinandoutofthesoilsona24hourcycle(morerapidlyatnight)".[38]Thus,the
earthwormnotonlycreatespassagesforairandwatertotraversethesoil,butalsomodifiesthevital

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organiccomponentthatmakesasoilhealthy(seeBioturbation).Earthwormspromotetheformationof
nutrientrichcasts(globulesofsoil,stableinsoil[mucus])thathavehighsoilaggregationandsoil
fertilityandquality.[3]

Earthwormsacceleratenutrientcyclinginthesoilplantsystemthroughfragmentation&mixingofplantdebris
physicalgrinding&chemicaldigestion.[3]Theearthworm'sexistencecannotbetakenforgranted.Dr.W.E.
ShewellCooperobserved"tremendousnumericaldifferencesbetweenadjacentgardens",andworm
populationsareaffectedbyahostofenvironmentalfactors,manyofwhichcanbeinfluencedbygood
managementpracticesonthepartofthegardenerorfarmer.[39]

Darwinestimatedthatarablelandcontainsupto53,000wormsperacre(13/m2),butmorerecentresearchfrom
RothamstedExperimentalStationhasproducedfiguressuggestingthatevenpoorsoilmaysupport
250,000/acre(62/m2),whilstrichfertilefarmlandmayhaveupto1,750,000/acre(432/m2),meaningthatthe
weightofearthwormsbeneathafarmer'ssoilcouldbegreaterthanthatofthelivestockuponitssurface.

Theabilitytobreakdownorganicmaterialsandexcreteconcentratednutrientsmakestheearthworma
functionalcontributorinrestorationprojects.Inresponsetoecosystemdisturbances,someopencastmining
siteshaveutilizedearthwormstopreparesoilforthereturnofnativeflora.Sitesusingthismethodhave
observedadvancesinthereturnofecosystemservicesthatpreviouslytookmuchlongertoreestablish.[40]
ResearchfromtheStationd'cologieTropicaledeLamtoassertsthattheearthwormspositivelyinfluencethe
rateofmacroaggregateformation,animportantfeatureforsoilstructure.[41]Thestabilityofaggregatesin
responsetowaterwasalsofoundtobeimprovedwhenconstructedbyearthworms.[41]

Asaninvasivespecies
Fromatotalofaround6,000species,onlyabout150speciesarewidelydistributedaroundtheworld.Theseare
theperegrineorcosmopolitanearthworms.[42]

Specialhabitats
While,asthenameearthwormsuggests,themainhabitatofearthwormsisin
soil,thesituationismorecomplicatedthanthat.ThebrandlingwormEisenia
fetidalivesindecayingplantmatterandmanure.Arctiostrotusvancouverensis
fromVancouverIslandandtheOlympicPeninsulaisgenerallyfoundin
decayingconiferlogs.Aporrectodealimicola,Sparganophilusspp.,andseveral
othersarefoundinmudinstreams.Somespeciesarearboreal,someaquatic
andsomeeuryhaline(saltwatertolerant)andlittoral(livingontheseashore,
e.g.Pontodriluslitoralis).Eveninthesoilspecies,specialhabitats,suchas
soilsderivedfromserpentine,haveanearthwormfaunaoftheirown.

Ecology
Earthwormsareclassifiedintothreemainecophysiologicalcategories:(1)leaf
litterorcompostdwellingwormsthatarenonburrowing,liveatsoillitter
interfaceandeatdecomposingOM(calledEpigeic)e.g.Eiseniafetida(2) Permanentverticalburrow
topsoilorsubsoildwellingwormsthatfeed(onsoil),burrowandcastwithin
soil,creatinghorizontalburrowsinupper1030cmofsoil(calledEndogeics)and(3)wormsthatconstruct
permanentdeepverticalburrowswhichtheyusetovisitthesurfacetoobtainplantmaterialforfood,suchas
leaves(calledAnecic(meaning"reachingup")),e.g.Lumbricusterrestris.[43]

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Earthwormpopulationsdependonbothphysicalandchemicalpropertiesofthe
soil,suchastemperature,moisture,pH,salts,aeration,andtexture,aswellas
availablefood,andtheabilityofthespeciestoreproduceanddisperse.Oneof
themostimportantenvironmentalfactorsispH,butearthwormsvaryintheir
preferences.Mostfavorneutraltoslightlyacidicsoils.Lumbricusterrestrisis
stillpresentinapHof5.4andDendrobaenaoctaedraatapHof4.3andsome
Megascolecidaearepresentinextremelyacidichumicsoils.SoilpHmayalso
influencethenumbersofwormsthatgointodiapause.Themoreacidicthesoil,
thesoonerwormsgointodiapauseandthelongertheyremainindiapauseata
pHof6.4.

Earthwormsformthebaseofmanyfoodchains.Theyarepreyeduponbymany
speciesofbirds(e.g.starlings,thrushes,gulls,crows,Europeanrobinsand Devil'scoachhorsebeetle
Americanrobins),snakes,mammals(e.g.bears,foxes,hedgehogs,pigs,moles) preyingonLumbricussp.
andinvertebrates(e.g.groundbeetlesandotherbeetles,snails,slugs).
Earthwormshavemanyinternalparasites,includingprotozoa,platyhelminthes,andnematodestheycanbe
foundintheworms'blood,seminalvesicles,coelom,orintestine,orintheircocoons.

Nitrogenousfertilizerstendtocreateacidicconditions,whicharefataltotheworms,anddeadspecimensare
oftenfoundonthesurfacefollowingtheapplicationofsubstancessuchasDDT,limesulphur,andlead
arsenate.InAustralia,changesinfarmingpracticessuchastheapplicationofsuperphosphatesonpasturesand
aswitchfrompastoralfarmingtoarablefarminghadadevastatingeffectonpopulationsofthegiantGippsland
earthworm,leadingtotheirclassificationasaprotectedspecies.

Theadditionoforganicmatter,preferablyasasurfacemulch,onaregularbasiswillprovideearthwormswith
theirfoodandnutrientrequirements,andwillcreatetheoptimumconditionsoftemperatureandmoisturethat
willstimulatetheiractivity.

Economicimpact
Variousspeciesofwormsareusedinvermiculture,thepracticeof
feedingorganicwastetoearthwormstodecomposefoodwaste.These
areusuallyEiseniafetida(oritscloserelativeEiseniaandrei)orthe
Brandlingworm,commonlyknownasthetigerwormorredwiggler.
Theyaredistinctfromsoildwellingearthworms.Inthetropics,the
AfricannightcrawlerEudriluseugeniaeandtheIndianbluePerionyx
excavatusareused.

Earthwormsaresoldallovertheworldthemarketissizable.
AccordingtoDougCollicut,"In1980,370millionwormswere EarthwormsbeingraisedatLa
exportedfromCanada,withaCanadianexportvalueof$13millionand ChonitaHaciendainMexico
anAmericanretailvalueof$54million."

Earthwormsarealsosoldasfoodforhumanconsumption.NokeisaculinarytermusedbytheMoriofNew
Zealand,andreferstoearthwormswhichareconsidereddelicaciesfortheirchiefs.

Taxonomyanddistribution
Withintheworldoftaxonomy,thestable'ClassicalSystem'ofMichaelsen(1900)andStephenson(1930)was
graduallyerodedbythecontroversyoverhowtoclassifyearthworms,suchthatFenderandMcKeyFender
(1990)wentsofarastosay,"Thefamilylevelclassificationofthemegascolecidearthwormsisinchaos."[44]
Overtheyears,manyscientistsdevelopedtheirownclassificationsystemsforearthworms,whichledto

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confusion,andthesesystemshavebeenandstillcontinuetoberevisedandupdated.Theclassificationsystem
usedhere,developedbyBlakemore(2000),isamodernreversiontotheClassicalSystemthatishistorically
provenandwidelyaccepted.[45]

CategorizationofamegadrileearthwormintooneofitstaxonomicfamiliesundersubordersLumbricinaand
Moniligastridaisbasedonsuchfeaturesasthemakeupoftheclitellum,thelocationanddispositionofthesex
features(pores,prostaticglands,etc.),numberofgizzards,andbodyshape.[45]Currently,over6,000speciesof
terrestrialearthwormsarenamed,asprovidedinaspeciesnamedatabase,[46]butthenumberofsynonymsis
unknown.

Thefamilies,withtheirknowndistributionsororigins:[45]

Acanthodrilidae(GondwananorPangaean?)
AiloscolecidaePyreneesandsoutheastUSA
Almidaetropicalequatorial(SouthAmerica,Africa,IndoAsia)
BenhamiinaeEthiopian,Neotropical(apossiblesubfamilyofOctochaetidae)
CriodrilidaesouthwesternPalaearctic:Europe,MiddleEast,RussiaandSiberiatoPacificcoastJapan
(Biwadrilus)mainlyaquatic
Diplocardiinae/idaeGondwananorLaurasian?(asubfamilyofAcanthodrilidae)
Enchytraeidaecosmopolitanbutuncommonintropics(usuallyclassedwithMicrodriles)
EudrilidaeTropicalAfricasouthoftheSahara
ExxidaeNeotropical:CentralAmericaandCaribbean
GlossoscolecidaeNeotropical:CentralandSouthAmerica,Caribbean
Haplotaxidaecosmopolitandistribution(usuallyclassedwithMicrodriles)
HormogastridaeMediterranean
KynotidaeMalagasian:Madagascar
LumbricidaeHolarctic:NorthAmerica,Europe,MiddleEast,CentralAsiatoJapan
LutodrilidaeLouisianasoutheastUSA
Megascolecidae(Pangaean?)
MicrochaetidaeTerrestrialinAfricaespeciallySouthAfricangrasslands
MoniligastridaeOrientalandIndiansubregion
OcnerodrilidaeNeotropics,AfricaIndia
OctochaetidaeAustralasian,Indian,Oriental,Ethiopian,Neotropical
OctochaetinaeAustralasian,Indian,Oriental(subfamilyifBenhamiinaeisaccepted)
SparganophilidaeNearctic,Neotropical:NorthandCentralAmerica
TumakidaeColombia,SouthAmerica

Seealso
Drilosphere,thepartofthesoilinfluencedbyearthwormsecretionsandcastings
TheFormationofVegetableMouldthroughtheActionofWorms,an1881bookbyCharlesDarwin
Soillife
Vermicompost
Wormcharming
fr:Stationd'cologiedeLamto(FrenchWikipedia)

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45.Blakemore,RobertJ.(March2006)."RevisedKeyto 46."Earthwormspeciesnamedatabase".
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RetrievedMay15,2012.

Referencedworks
Blakemore,RobertJ.(2012).CosmopolitanEarthwormsanEcoTaxonomicGuidetothePeregrine
SpeciesoftheWorld.(5thEd).Yokohama,Japan:VermEcologySo(i)lutions.
Sims,ReginaldWilliamGerard,B(1985).Earthworms:KeysandNotesfortheIdentificationandStudy
oftheSpecies.London:PublishedforTheLinneanSocietyofLondonandtheEstuarineandBrackish
WaterSciencesAssociationbyE.J.Brill/Dr.W.Backhuys.
Edwards,CliveArthurBohlen,PatrickJ.(1996).BiologyandEcologyofEarthworms,3rdEd.Springer.

Furtherreading
Edwards,CliveA.,Bohlen,P.J.(Eds.)BiologyandEcologyofEarthworms.Springer,2005.3rdedition.
Edwards,CliveA.(Ed.)EarthwormEcology.BocaRaton:CRCPress,2004.Secondrevisededition.
ISBN084931819X
Lee,KenethE.Earthworms:TheirEcologyandRelationshipswithSoilsandLandUse.AcademicPress.
Sydney,1985.ISBN0124408605
Stewart,Amy.TheEarthMoved:OntheRemarkableAchievementsofEarthworms.ChapelHill,N.C.:
AlgonquinBooks,2004.ISBN1565123379

Externallinks
General WikimediaCommonshas
mediarelatedto
TheEarthwormSocietyofBritain(http://www.earthwormsoc.org. Earthworms.
uk/)
WormWatchFieldguidetoearthworms(http://www.naturewatch.ca/english/wormwatch/)
Earthwormsaspestsandotherwise(http://digital.library.unt.edu/permalink/metadc1514:1)hostedby
theUNTGovernmentDocumentsDepartment(http://digital.library.unt.edu/browse/department/govdocs/)

Academic
InfographyaboutEarthworms(http://www.infography.com/content/370686832988.html)
Earthwormresources(http://pedozoologia.uw.hu/)
OpusculaZoologicaBudapest,onlinepapersonearthwormtaxonomy(http://opuscula.elte.hu/)
ASeriesofSearchableTextsonEarthwormBiodiversity,EcologyandSystematicsfromVariousRegions
oftheWorld(http://www.annelida.net/earthworm/)maintainedbyRobBlakemore,Ph.D.,an
earthwormtaxonomyspecialist

Agricultureandecology

EarthwormInformation(http://www.sarep.ucdavis.edu/worms/)atUniversityofCalifornia,Davis
MinnesotaInvasiveEarthworms(http://www.dnr.state.mn.us/invasives/terrestrialanimals/earthworms/ind
ex.html)MinnesotaDNRinformationonthenegativeimpactsofearthworms

Wormfarming

https://en.wikipedia.org/wiki/Earthworm 13/14
2/13/2017 EarthwormWikipedia

Amultitieredwormfarm(http://workingworms.com/content/view/40/62/)Atechnicalguidefor
constructingatieredwormfarmingsystem
HowtoMakeaWormFarm(http://opentutorial.com/index.php?title=Make_a_worm_farm)Goodfor
CompostingandFishing

Forchildren
BioKIDSSegmentedworms(http://www.biokids.umich.edu/critters/Annelida/)

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