2/13/2017 EarthwormWikipedia

Earthworm
FromWikipedia,thefreeencyclopedia

Anearthwormisatubeshaped,segmentedwormfoundinthe
[Link], Earthworm
[Link]'sdigestive
[Link]
[Link]
coelomicfluidthatmoveswithinthefluidfilledcoelomanda
simple,[Link]
[Link]
twogangliaabovethemouth,oneoneitherside,connectedtoa
nervecordrunningbackalongitslengthtomotorneuronsand
[Link]
[Link]
musclesontheperipheryofeachsegmentenablethewormtomove.
Similarsetsofmuscleslinethegut,andtheiractionsmovethe
digestingfoodtowardtheworm'sanus.[2]
Aspeciesofearthwormwithawell
Earthwormsarehermaphroditeseachindividualcarriesbothmale
developedclitellum
[Link]
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
[Link],instead,inthe
suborderLumbricinaoftheorderHaplotaxida,butthismayagain Haplotaxida
[Link]"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,[Link]
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

[Link] 1/14
2/13/2017 EarthwormWikipedia

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,[Link]
analsegments,eachsegmentcarriesbristlelikehairscalledlateralsetae[6]usedtoanchorpartsofthebody

[Link] 2/14
2/13/2017 EarthwormWikipedia

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

Generally,withinaspecies,thenumberofsegmentsfoundisconsistentacrossspecimens,andindividualsare
[Link](segment
number1)featuresboththeearthworm'smouthand,overhangingthemouth,afleshylobecalledthe
prostomium,whichsealstheentrancewhenthewormisatrest,butisalsousedtofeelandchemicallysensethe
worm'[Link]
itemssuchasgrassesandleavesintotheirburrow.

Anadultearthwormdevelopsabeltlikeglandularswelling,calledtheclitellum,whichcoversseveral
[Link].
Theposteriorismostcommonlycylindricalliketherestofthebody,butdependingonthespecies,mayalsobe
quadrangular,octagonal,trapezoidal,[Link]'s
anus,ashortverticalslit,isfoundonthissegment.[6]

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

[Link] 3/14
2/13/2017 EarthwormWikipedia

[Link],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
[Link] earthworm
canalinthethirdsegment,inagroovebetweenthebuccal
cavityandpharynx.

Apairofcircumpharyngealconnectivesfromthebrainencirclethepharynxandthenconnectwithapairof
[Link],
subpharyngealgangliaandthecircumpharyngealconnectivesformanerveringaroundthepharynx.

Theventralnervecord(formedbynervecellsandnervefibres)beginsatthesubpharyngealgangliaand
[Link],
organglion,ineachsegment,[Link],whichoccursfromthefifthtothelastsegmentofthe
[Link],onemedialgiantaxon(MGA)andtwolateralgiantaxons(LGAs)on
themiddorsalsideoftheventralnervecord.TheMGAis0.07mmindiameterandtransmitsinananterior
posteriordirectionatarateof32.2m/s.TheLGAsareslightlywiderat0.05mmindiameterandtransmitina
posterioranteriordirectionat12.6m/[Link]
arethereforeconsideredonegiantaxon.[15][16]

Peripheralnervoussystem

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

Sympatheticnervoussystem

Thesympatheticnervoussystemconsistsofnerveplexusesintheepidermisandalimentarycanal.(Aplexusa
webofnervecellsconnectedtogetherinatwodimensionalgrid.)Thenervesthatrunalongthebodywallpass
[Link]

[Link] 4/14
2/13/2017 EarthwormWikipedia

[Link]
connective.

Movement

Onthesurface,[Link]
takinglonger"strides"[Link]
[Link]
stridefrequencies.[17]

Touchinganearthworm,whichcausesa"pressure"responseaswellas(often)aresponsetothedehydrating
qualityofthesaltonhumanskin(toxictoearthworms),stimulatesthesubepidermalnerveplexuswhich
connectstotheintermuscularplexusandcausesthelongitudinalmusclestocontact,therebythewrithing
[Link]
[Link]
[Link]
nervoussystemsofthesegments.[18]

[Link]
[Link],fromthereartothefrontofthe
[Link],asignalisrapidlysentforwardscausingthelongitudinalmusclesin
[Link]
[Link]
[Link](perhapscontractingintoitsburrow
toescapeabird).

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

Senses

Photosensitivity

Earthwormsdonothaveeyes(althoughsomewormsdo),however,theydohavespecialisedphotosensitive
cellscalled"lightcellsofHess".Thesephotoreceptorcellshaveacentralintracellularcavity(phaosome)filled
[Link],thereareseveralsensoryciliainthephaosomewhicharestructurally
independentofthemicrovilli.[21]Thephotoreceptorsaredistributedinmostpartsoftheepidermisbutaremore
[Link]
[Link]
areveryfewinnumberpastthethirdsegment.[18]

Digestivesystem

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

[Link],
[Link],wherecalcium(fromthebloodand
ingestedfrompreviousmeals)ispumpedintomaintainproperbloodcalciumlevelsinthebloodandfoodpH.
[Link] 5/14
2/13/2017 EarthwormWikipedia

[Link],strongmuscularcontractionsgrindthe
[Link],foodcontinues
[Link],Amylasetodigest
polysaccharides,Cellulasetodigestcellulose,andlipasetodigestfats.[2]Earthwormsuseinadditiontothe
digestiveproteinsaclassofsurfaceactivecompoundscalleddrilodefensins,whichhelpdigestplant
material.[23]Insteadofbeingcoiledlikeamammalianintestine,anearthworm'sintestineincreasessurfacearea
[Link]
musclelayerslikethebody,butinreverseorderaninnercircularlayerwithinanouterlongitudinallayer.[24]
Thegizzardisasomewhatofauniqueorgantothewormasitisfoundinveryfewanimals.

Circulatorysystem

Theearthwormhasadualcirculatorysysteminwhichboththecoelomicfluidandaclosedcirculatorysystem
carrythefood,waste,[Link]:the
dorsal(top)vessel,whichrunsabovethedigestivetracttheventral(bottom)vessel,whichrunsbelowthe
digestivetractthesubneuralvessel,whichrunsbelowtheventralnervecordandtwolateroneuralvesselson
eithersideofthenervecord.[25]Thedorsalvesselmovesthebloodforward,whiletheotherfourlongitudinal
[Link],apairofaorticarchesringsthecoelomand
actsashearts,[Link]
[Link]
[Link],theyreleasenonlivingcellsoffatintothe
fluidfilledcoelom,wheretheyfloatfreelybutcanpassthroughthewallsseparatingeachsegment,moving
foodtootherpartsandassistinwoundhealing.[26]

Excretorysystem

Theexcretorysystemcontainsapairofnephridiaineverysegment,exceptforthefirstthreeandthelast
ones.[27]Thethreetypesofnephridiaare:integumentary,septal,[Link]
[Link]
[Link],
fifthandsixthsegments.[27]Thewasteinthecoelomfluidfromaforwardsegmentisdrawninbythebeating
[Link](wall)viaatubewhichformsaseriesof
[Link]
wastesarethenfinallydischargedthroughaporeontheworm'sside.[28]

Respiration

[Link],
wheretheoxygenispickedupbythehemoglobindissolvedinthebloodplasmaandcarbondioxideisreleased.
Water,aswellassalts,canalsobemovedthroughtheskinbyactivetransport.

Reproduction

Matingoccursonthesurface,[Link],theyhavebothmale
[Link]
[Link],storeandreleasethesperm
viathemalepores.Ovariesandoviductsinsegment13releaseeggsviafemaleporesonsegment14,while
spermisexpelledfromsegment15.Oneormorepairsofspermathecaearepresentinsegments9and10
(dependingonthespecies)whichareinternalsacsthatreceiveandstorespermfromtheotherwormduring
[Link],segment15ofonewormexudesspermintosegments9and10withitsstoragevesicles
[Link].

[Link] 6/14
2/13/2017 EarthwormWikipedia

InHormogastersamniticaandHormogasterelisaetranscriptomeDNA
librariesweresequencedandtwosexpheromones,Attractinand
Temptin,weredetectedinalltissuesamplesofbothspecies.[29]Sex
pheromonesareprobablyimportantinearthwormsbecausetheylivein
anenvironmentwherechemicalsignalingmayplayacrucialrolein
[Link]
providethebenefitofmaskingtheexpressionofdeleteriousrecessive
mutationsinprogeny.[30](Alsoseecomplemenation.)
Earthwormcopulation
[Link]
matingpairoverlapfrontendsventrallyandeachexchangesspermwith
[Link]
timeaftercopulation,longafterthewormshaveseparated,theclitellum
(behindthespermathecae)secretesmaterialwhichformsaringaround
[Link],andasitdoesso,it
injectsitsowneggsandtheotherworm'[Link]
slipsoutofthering,theendsofthecocoonsealtoformavaguely
lemonshapedincubator(cocoon)inwhichtheembryonicworms
[Link],butfullyformedearthworms,butlack
theirsexstructures,[Link] [Link]
[Link]
underfieldconditionsisfourtoeightyears,whilemostgardenvarieties
[Link]
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
[Link]:

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
[Link].
CriodriluslacuumHoffmeister,1845alsohasprodigiousregenerativecapacitywithheadregeneration
fromasfarbackas40/41.[34]

AnunidentifiedTasmanianearthwormshowngrowingareplacementheadhasbeenreported.[35]

[Link] 7/14
2/13/2017 EarthwormWikipedia

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

Earthwormsworkasbiological"pistons"forcingairthroughthetunnelsasthey
[Link],andisconduciveto Closeupofanearthwormin
[Link] gardensoil
earthwormcometothesurfaceandgrazeonthehigherconcentrationsof
organicmatterpresentthere,[Link]
leveloforganicmattermixingisassociatedwithsoilfertility,anabundanceofearthwormsisgenerally
[Link],aslongagoas1881CharlesDarwinwrote:"Itmaybe
doubtedwhethertherearemanyotheranimalswhichhaveplayedsoimportantapartinthehistoryofthe
world,ashavetheselowlyorganizedcreatures."[37]

Benefits
Themajorbenefitsofearthwormactivitiestosoilfertilitycanbesummarizedas:

Biological:Inmanysoils,earthwormsplayamajorroleintheconversionoflargepiecesoforganic
matterintorichhumus,[Link]'sactionsofpulling
belowthesurfacedepositedorganicmattersuchasleaffallormanure,eitherforfoodortoplugit's
[Link],thewormwillshredtheleafandpartiallydigestitandmingleitwiththe
[Link](seebottomright)cancontain40percentmorehumusthanthetop9"(23cm)ofsoil
inwhichthewormisliving.[3]

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

[Link] 8/14
2/13/2017 EarthwormWikipedia

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

Earthwormsacceleratenutrientcyclinginthesoilplantsystemthroughfragmentation&mixingofplantdebris
physicalgrinding&chemicaldigestion.[3]Theearthworm'[Link].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
[Link],someopencastmining
[Link]
observedadvancesinthereturnofecosystemservicesthatpreviouslytookmuchlongertoreestablish.[40]
ResearchfromtheStationd'cologieTropicaledeLamtoassertsthattheearthwormspositivelyinfluencethe
rateofmacroaggregateformation,animportantfeatureforsoilstructure.[41]Thestabilityofaggregatesin
responsetowaterwasalsofoundtobeimprovedwhenconstructedbyearthworms.[41]

Asaninvasivespecies
Fromatotalofaround6,000species,[Link]
theperegrineorcosmopolitanearthworms.[42]

Specialhabitats
While,asthenameearthwormsuggests,themainhabitatofearthwormsisin
soil,[Link]
[Link]
fromVancouverIslandandtheOlympicPeninsulaisgenerallyfoundin
[Link],Sparganophilusspp.,andseveral
[Link],someaquatic
andsomeeuryhaline(saltwatertolerant)andlittoral(livingontheseashore,
[Link]).Eveninthesoilspecies,specialhabitats,suchas
soilsderivedfromserpentine,haveanearthwormfaunaoftheirown.

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

[Link] 9/14
2/13/2017 EarthwormWikipedia

Earthwormpopulationsdependonbothphysicalandchemicalpropertiesofthe
soil,suchastemperature,moisture,pH,salts,aeration,andtexture,aswellas
availablefood,[Link]
themostimportantenvironmentalfactorsispH,butearthwormsvaryintheir
[Link]
stillpresentinapHof5.4andDendrobaenaoctaedraatapHof4.3andsome
[Link]
[Link],
thesoonerwormsgointodiapauseandthelongertheyremainindiapauseata
pHof6.4.

[Link]
speciesofbirds([Link],thrushes,gulls,crows,Europeanrobinsand Devil'scoachhorsebeetle
Americanrobins),snakes,mammals([Link],foxes,hedgehogs,pigs,moles) preyingonLumbricussp.
andinvertebrates([Link],snails,slugs).
Earthwormshavemanyinternalparasites,includingprotozoa,platyhelminthes,andnematodestheycanbe
foundintheworms'blood,seminalvesicles,coelom,orintestine,orintheircocoons.

Nitrogenousfertilizerstendtocreateacidicconditions,whicharefataltotheworms,anddeadspecimensare
oftenfoundonthesurfacefollowingtheapplicationofsubstancessuchasDDT,limesulphur,andlead
[Link],changesinfarmingpracticessuchastheapplicationofsuperphosphatesonpasturesand
aswitchfrompastoralfarmingtoarablefarminghadadevastatingeffectonpopulationsofthegiantGippsland
earthworm,leadingtotheirclassificationasaprotectedspecies.

Theadditionoforganicmatter,preferablyasasurfacemulch,onaregularbasiswillprovideearthwormswith
theirfoodandnutrientrequirements,andwillcreatetheoptimumconditionsoftemperatureandmoisturethat
willstimulatetheiractivity.

Economicimpact
Variousspeciesofwormsareusedinvermiculture,thepracticeof
[Link]
areusuallyEiseniafetida(oritscloserelativeEiseniaandrei)orthe
Brandlingworm,commonlyknownasthetigerwormorredwiggler.
[Link],the
AfricannightcrawlerEudriluseugeniaeandtheIndianbluePerionyx
excavatusareused.

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

[Link]
Zealand,andreferstoearthwormswhichareconsidereddelicaciesfortheirchiefs.

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

[Link] 10/14
2/13/2017 EarthwormWikipedia

confusion,[Link]
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)

References
1."ITISReportforLumbricina,TaxonomicSerialNo.: [Link](2009).Elementsofthe
69069".ITIS.RetrievedMay14,2012. NatureandPropertiesofSoils(3rdEdition).Prentice
[Link].,[Link],Frances Hall.ISBN9780135014332.
MHickman(1984).IntegratedPrinciplesofZoology 4.Blakemore2012,[Link].
(7thed.).TimesMirror/MosbyCollegePublishing. [Link]&Bohlen1996,p.11.
p.344.ISBN0801621739. [Link]&Gerard1985,pp.36.
[Link]&Bholen1996,p.3.
[Link]&Bohlen1996,p.89.
[Link] 11/14
2/13/2017 EarthwormWikipedia

[Link]&Bohlen1996,p.1. [Link],HopfFA,MichodRE(1987)."The
[Link]&Gerard1985,p.8. molecularbasisoftheevolutionofsex".[Link].
[Link]&Bohlen1996,p.12. AdvancesinGenetics.24:32370.
[Link]&Bohlen1996,p.6. doi:10.1016/S00652660(08)600127.ISBN9780
13."BrainInvertebrateBrainGanglia,Nervous, 120176243.PMID3324702.
System,andHead".HSBE.RetrievedApril3,2015. [Link],AiraM,BreinholtJW,StojanovicM,
[Link],S."Differentpartsofnervoussystem JamesSW,PrezLosadaM(2015)."Underground
ofearthworm(withdiagram)".BiologyDiscussion. evolution:Newrootsfortheoldtreeoflumbricid
RetrievedApril3,2015. earthworms".[Link].83:719.
15."Experiment:Comparingspeedsoftwonervefiber doi:10.1016/[Link].2014.10.024.PMC4766815 .
sizes".BackyardBrains.RetrievedApril4,2015. PMID25463017.
[Link],C.D.,Landa,[Link],J.L.(1978). [Link],AlmodvarA,NovoM,SimancasB,
"Giantnervefibreactivityinintact,freelymoving DazCosnDJ(2012)."Addingcomplexitytothe
earthworms".TheJournalofExperimentalBiology. complex:newinsightsintothephylogeny,
72:217227.PMID624897. diversificationandoriginofparthenogenesisinthe
[Link],K.J.(1999)."Kinematicscalingof Aporrectodeacaliginosaspeciescomplex
locomotionbyhydrostaticanimals:ontogenyof (Oligochaeta,Lumbricidae)".[Link].
peristalticcrawlingbytheearthwormlumbricus 64(2):[Link].1016/[Link].2012.04.011.
terrestris".JournalofExperimentalBiology.202: PMID22542691.
661674.PMID10021320. [Link]([Link]
18."Earthwormnervoussystem".[Link] [Link])
April3,2015. [Link],G.E.(1January1953)."OnRegenerative
[Link],R.W.(2011)."Painandsufferingin CapacityofEarthwormsoftheFamilyLumbricidae".
invertebrates?".ILARJournal.52(2):17584. TheAmericanMidlandNaturalist.50(2):414419.
PMID21709310. doi:10.2307/2422100.
[Link],J.A."Aquestionofpainininvertebrates". 35."Invertebrata20aitems".
ILARJournal.33(12):2531.RetrievedApril3, [Link],K.J.(2000)."Ontogeneticscalingof
2015. burrowingforcesintheearthwormLumbricus
[Link],P.,Aros,[Link],Sz.(1970)."Fine terrestris"(PDF).JournalofExperimentalBiology.
structureofphotoreceptorcellsintheearthworm, 203(Pt18):[Link]
Lumbricusterrestris".ZeitschriftfrZellforschung April4,2015.
undMikroskopischeAnatomie.104(3):345357. [Link],Charles,Theformationofvegetablemould
doi:10.1007/BF00335687. throughtheactionofworms,withobservationson
[Link]&Bohlen1996,p.13. [Link]
[Link],ManuelStrittmatter,NicoleFearn,Sarah FormationofVegetableMould,byDarwin([Link]
Morgan,[Link],PeterFuchser,JensWallis, [Link]/ebooks/2355)
DavidPalchykov,VitaliiRobertson,Jeremy(2015 [Link],Bill,PermacultureADesigner'sManual,
0804)."Uniquemetabolitesprotectearthworms TagariPress,1988
againstplantpolyphenols".NatureCommunications. [Link],ShewellSoil,HumusAndHealthISBN978
6:[Link].1038/ncomms8869.PMC4532835 . 0583127967
PMID26241769. [Link],StephaneWratten,StephenD.(20100501).
[Link]&Bohlen1996,pp.1315. "Thepotentialofearthwormstorestoreecosystem
[Link]&Gerard1985,p.10. servicesafteropencastminingAreview".Basicand
[Link].,[Link],Frances AppliedEcology.11(3):196203.
MHickman(1984).IntegratedPrinciplesofZoology doi:10.1016/[Link].2009.12.005.
(7thed.).TimesMirror/MosbyCollegePublishing. [Link],Eric(19921201)."Restorationby
pp.344345.ISBN0801621739. earthworms(megascolecidae)ofthemacroaggregate
[Link],H.J."ExcretorySystem".Retrieved29July structureofadestructuredsavannasoilunderfield
2012. conditions".SoilBiologyandBiochemistry.24(12):
[Link].,[Link],Frances [Link].1016/00380717(92)90155Q.
MHickman(1984).IntegratedPrinciplesofZoology [Link]([Link]
(7thed.).TimesMirror/MosbyCollegePublishing. arthworm/[Link])
pp.345346.ISBN0801621739. [Link]:RenewersofAgroecosystems(SAFall,
[Link],RiesgoA,FernndezGuerraA,GiribetG 1990(v3n1))([Link]
(2013)."Pheromoneevolution,reproductivegenes, TR/v3n1/[Link])
andcomparativetranscriptomicsinmediterranean [Link]&McKeyFender(1990).SoilBiologyGuide.
earthworms(annelida,oligochaeta,hormogastridae)". WileyInterscience.ISBN0471045519.
[Link].30(7):161429.
doi:10.1093/molbev/mst074.PMID23596327.

[Link] 12/14
2/13/2017 EarthwormWikipedia

[Link],RobertJ.(March2006)."RevisedKeyto 46."Earthwormspeciesnamedatabase".
WorldwideEarthwormFamiliesfromBlakemoreplus
ReviewsofCriodrilidae(includingBiwadrilidae)and
Octochaetidae"(PDF).ASeriesofSearchableTextson
EarthwormBiodiversity,EcologyandSystematics
[Link].
RetrievedMay15,2012.

Referencedworks
Blakemore,RobertJ.(2012).CosmopolitanEarthwormsanEcoTaxonomicGuidetothePeregrine
SpeciesoftheWorld.(5thEd).Yokohama,Japan:VermEcologySo(i)lutions.
Sims,ReginaldWilliamGerard,B(1985).Earthworms:KeysandNotesfortheIdentificationandStudy
[Link]:PublishedforTheLinneanSocietyofLondonandtheEstuarineandBrackish
[Link]/[Link].
Edwards,CliveArthurBohlen,PatrickJ.(1996).BiologyandEcologyofEarthworms,[Link].

Furtherreading
Edwards,CliveA.,Bohlen,P.J.(Eds.)[Link],2005.3rdedition.
Edwards,CliveA.(Ed.)[Link]:CRCPress,[Link].
ISBN084931819X
Lee,[Link]:[Link].
Sydney,1985.ISBN0124408605
Stewart,[Link]:[Link],N.C.:
AlgonquinBooks,2004.ISBN1565123379

Externallinks
General WikimediaCommonshas
mediarelatedto
TheEarthwormSocietyofBritain([Link] Earthworms.
uk/)
WormWatchFieldguidetoearthworms([Link]
Earthwormsaspestsandotherwise([Link]
theUNTGovernmentDocumentsDepartment([Link]

Academic
InfographyaboutEarthworms([Link]
Earthwormresources([Link]
OpusculaZoologicaBudapest,onlinepapersonearthwormtaxonomy([Link]
ASeriesofSearchableTextsonEarthwormBiodiversity,EcologyandSystematicsfromVariousRegions
oftheWorld([Link]
earthwormtaxonomyspecialist

Agricultureandecology

EarthwormInformation([Link]
MinnesotaInvasiveEarthworms([Link]
[Link])MinnesotaDNRinformationonthenegativeimpactsofearthworms

Wormfarming

[Link] 13/14
2/13/2017 EarthwormWikipedia

Amultitieredwormfarm([Link]
constructingatieredwormfarmingsystem
HowtoMakeaWormFarm([Link]
CompostingandFishing

Forchildren
BioKIDSSegmentedworms([Link]

Retrievedfrom"[Link]

Categories: Annelids Soilbiology

Thispagewaslastmodifiedon12February2017,at18:04.
TextisavailableundertheCreativeCommonsAttributionShareAlikeLicenseadditionaltermsmay
[Link],[Link]
trademarkoftheWikimediaFoundation,Inc.,anonprofitorganization.

[Link] 14/14