Ecosystem - Wikipedia

Definition. An ecosystem (or ecological system) consists of all the organisms and the abiotic pools (or physical environment) with which they interact. Ecosystem FromWikipedia,thefreeencyclopedia Jumptonavigation Jumptosearch Communityoflivingorganismstogetherwiththenonlivingcomponentsoftheirenvironment Thisarticleisaboutnaturalecosystems.Forthetermusedinman-madesystems,seeDigitalecosystem.Forthesystemofclassifyingecologicallyhomogeneouslandunits,seeBiome. "Biosystem"redirectshere.Forthejournal,seeBioSystems. Left:Coralreefecosystemsarehighlyproductivemarinesystems.[1]Right:Temperaterainforest,aterrestrialecosystem. PartofaseriesonBiologyScienceoflife Index Outline Glossary History(timeline) Keycomponents Celltheory Ecosystem Evolution Phylogeny Propertiesoflife Adaptation Energyprocessing Growth Order Regulation Reproduction Responsetoenvironment DomainsandKingdomsoflife Archaea Bacteria Eukarya(Animals,Fungi,Plants,Protists) Branches Anatomy Biotechnology Botany Cellbiology Ecology Evolutionarybiology Genetics Marinebiology Microbiology Molecularbiology Mycology Neuroscience Paleontology Phycology Physiology Protistology Virology Zoology Research Biologist(list) Listofbiologyawards Listofjournals Listofresearchmethods Listofunsolvedproblems Applications Agriculturalscience Biomedicalsciences Healthtechnology Pharming  Biologyportal  Categoryvte Anecosystem(orecologicalsystem)consistsofalltheorganismsandthephysicalenvironmentwithwhichtheyinteract.[2]: 458 Thesebioticandabioticcomponentsarelinkedtogetherthroughnutrientcyclesandenergyflows.Energyentersthesystemthroughphotosynthesisandisincorporatedintoplanttissue.Byfeedingonplantsandononeanother,animalsplayanimportantroleinthemovementofmatterandenergythroughthesystem.Theyalsoinfluencethequantityofplantandmicrobialbiomasspresent.Bybreakingdowndeadorganicmatter,decomposersreleasecarbonbacktotheatmosphereandfacilitatenutrientcyclingbyconvertingnutrientsstoredindeadbiomassbacktoaformthatcanbereadilyusedbyplantsandmicrobes. Ecosystemsarecontrolledbyexternalandinternalfactors.Externalfactorssuchasclimate,parentmaterialwhichformsthesoilandtopography,controltheoverallstructureofanecosystembutarenotthemselvesinfluencedbytheecosystem.Internalfactorsarecontrolled,forexample,bydecomposition,rootcompetition,shading,disturbance,succession,andthetypesofspeciespresent.Whiletheresourceinputsaregenerallycontrolledbyexternalprocesses,theavailabilityoftheseresourceswithintheecosystemiscontrolledbyinternalfactors.Therefore,internalfactorsnotonlycontrolecosystemprocessesbutarealsocontrolledbythem. Ecosystemsaredynamicentities—theyaresubjecttoperiodicdisturbancesandarealwaysintheprocessofrecoveringfromsomepastdisturbance.Thetendencyofanecosystemtoremainclosetoitsequilibriumstate,despitethatdisturbance,istermeditsresistance.Thecapacityofasystemtoabsorbdisturbanceandreorganizewhileundergoingchangesoastoretainessentiallythesamefunction,structure,identity,andfeedbacksistermeditsecologicalresilience.Ecosystemscanbestudiedthroughavarietyofapproaches—theoreticalstudies,studiesmonitoringspecificecosystemsoverlongperiodsoftime,thosethatlookatdifferencesbetweenecosystemstoelucidatehowtheyworkanddirectmanipulativeexperimentation.Biomesaregeneralclassesorcategoriesofecosystems.However,thereisnocleardistinctionbetweenbiomesandecosystems.Ecosystemclassificationsarespecifickindsofecologicalclassificationsthatconsiderallfourelementsofthedefinitionofecosystems:abioticcomponent,anabioticcomplex,theinteractionsbetweenandwithinthem,andthephysicalspacetheyoccupy. Ecosystemsprovideavarietyofgoodsandservicesuponwhichpeopledepend.Ecosystemgoodsincludethe"tangible,materialproducts"ofecosystemprocessessuchaswater,food,fuel,constructionmaterial,andmedicinalplants.Ecosystemservices,ontheotherhand,aregenerally"improvementsintheconditionorlocationofthingsofvalue".Theseincludethingslikethemaintenanceofhydrologicalcycles,cleaningairandwater,themaintenanceofoxygenintheatmosphere,croppollinationandeventhingslikebeauty,inspirationandopportunitiesforresearch.Manyecosystemsbecomedegradedthroughhumanimpacts,suchassoilloss,airandwaterpollution,habitatfragmentation,waterdiversion,firesuppression,andintroducedspeciesandinvasivespecies.Thesethreatscanleadtoabrupttransformationoftheecosystemortogradualdisruptionofbioticprocessesanddegradationofabioticconditionsoftheecosystem.Oncetheoriginalecosystemhaslostitsdefiningfeatures,itisconsidered"collapsed".Ecosystemrestorationisthoughttocontributetoall17SustainableDevelopmentGoals. Contents 1Definition 1.1Originanddevelopmentoftheterm 2Ecosystemprocesses 2.1Externalandinternalfactors 2.2Primaryproduction 2.3Energyflow 2.4Decomposition 2.4.1Decompositionrates 2.5Dynamicsandresilience 2.6Nutrientcycling 2.7Functionandbiodiversity 3Studyapproaches 3.1Ecosystemecology 3.2Classifications 3.3Examples 4Humaninteractionswithecosystems 4.1Ecosystemgoodsandservices 4.2Ecosystemdegradationanddecline 4.3Ecosystemmanagement 4.4Ecosystemrestorationandsustainabledevelopment 5Seealso 6References 6.1Notes Definition Anecosystem(orecologicalsystem)consistsofalltheorganismsandtheabioticpools(orphysicalenvironment)withwhichtheyinteract.[3][4]: 5 [2]: 458 Thebioticandabioticcomponentsarelinkedtogetherthroughnutrientcyclesandenergyflows.[5] "Ecosystemprocesses"arethetransfersofenergyandmaterialsfromonepooltoanother.[2]: 458 Ecosystemprocessesareknownto"takeplaceatawiderangeofscales".Therefore,thecorrectscaleofstudydependsonthequestionasked.[4]: 5  Originanddevelopmentoftheterm Theterm"ecosystem"wasfirstusedin1935inapublicationbyBritishecologistArthurTansley.ThetermwascoinedbyArthurRoyClapham,whocameupwiththewordatTansley'srequest.[6]Tansleydevisedtheconcepttodrawattentiontotheimportanceoftransfersofmaterialsbetweenorganismsandtheirenvironment.[4]: 9 Helaterrefinedtheterm,describingitas"Thewholesystem,...includingnotonlytheorganism-complex,butalsothewholecomplexofphysicalfactorsformingwhatwecalltheenvironment".[3]Tansleyregardedecosystemsnotsimplyasnaturalunits,butas"mentalisolates".[3]Tansleylaterdefinedthespatialextentofecosystemsusingtheterm"ecotope".[7] G.EvelynHutchinson,alimnologistwhowasacontemporaryofTansley's,combinedCharlesElton'sideasabouttrophicecologywiththoseofRussiangeochemistVladimirVernadsky.Asaresult,hesuggestedthatmineralnutrientavailabilityinalakelimitedalgalproduction.Thiswould,inturn,limittheabundanceofanimalsthatfeedonalgae.RaymondLindemantooktheseideasfurthertosuggestthattheflowofenergythroughalakewastheprimarydriveroftheecosystem.Hutchinson'sstudents,brothersHowardT.OdumandEugeneP.Odum,furtherdevelopeda"systemsapproach"tothestudyofecosystems.Thisallowedthemtostudytheflowofenergyandmaterialthroughecologicalsystems.[4]: 9  Ecosystemprocesses Rainforestecosystemsarerichinbiodiversity.ThisistheGambiaRiverinSenegal'sNiokolo-KobaNationalPark. FloraofBajaCaliforniaDesert,Cataviñaregion,Mexico Externalandinternalfactors Ecosystemsarecontrolledbybothexternalandinternalfactors.Externalfactors,alsocalledstatefactors,controltheoverallstructureofanecosystemandthewaythingsworkwithinit,butarenotthemselvesinfluencedbytheecosystem.Onbroadgeographicscales,climateisthefactorthat"moststronglydeterminesecosystemprocessesandstructure".[4]: 14 Climatedeterminesthebiomeinwhichtheecosystemisembedded.Rainfallpatternsandseasonaltemperaturesinfluencephotosynthesisandtherebydeterminetheamountofenergyavailabletotheecosystem.[8]: 145  Parentmaterialdeterminesthenatureofthesoilinanecosystem,andinfluencesthesupplyofmineralnutrients.Topographyalsocontrolsecosystemprocessesbyaffectingthingslikemicroclimate,soildevelopmentandthemovementofwaterthroughasystem.Forexample,ecosystemscanbequitedifferentifsituatedinasmalldepressiononthelandscape,versusonepresentonanadjacentsteephillside.[9]: 39 [10]: 66  Otherexternalfactorsthatplayanimportantroleinecosystemfunctioningincludetimeandpotentialbiota,theorganismsthatarepresentinaregionandcouldpotentiallyoccupyaparticularsite.Ecosystemsinsimilarenvironmentsthatarelocatedindifferentpartsoftheworldcanendupdoingthingsverydifferentlysimplybecausetheyhavedifferentpoolsofspeciespresent.[11]: 321 Theintroductionofnon-nativespeciescancausesubstantialshiftsinecosystemfunction.[12] Unlikeexternalfactors,internalfactorsinecosystemsnotonlycontrolecosystemprocessesbutarealsocontrolledbythem.[4]: 16 Whiletheresourceinputsaregenerallycontrolledbyexternalprocesseslikeclimateandparentmaterial,theavailabilityoftheseresourceswithintheecosystemiscontrolledbyinternalfactorslikedecomposition,rootcompetitionorshading.[13]Otherfactorslikedisturbance,successionorthetypesofspeciespresentarealsointernalfactors. Primaryproduction Globaloceanicandterrestrialphototrophabundance,fromSeptember1997toAugust2000.Asanestimateofautotrophbiomass,itisonlyaroughindicatorofprimaryproductionpotentialandnotanactualestimateofit. Mainarticle:Primaryproduction Primaryproductionistheproductionoforganicmatterfrominorganiccarbonsources.Thismainlyoccursthroughphotosynthesis.Theenergyincorporatedthroughthisprocesssupportslifeonearth,whilethecarbonmakesupmuchoftheorganicmatterinlivinganddeadbiomass,soilcarbonandfossilfuels.Italsodrivesthecarboncycle,whichinfluencesglobalclimateviathegreenhouseeffect. Throughtheprocessofphotosynthesis,plantscaptureenergyfromlightanduseittocombinecarbondioxideandwatertoproducecarbohydratesandoxygen.Thephotosynthesiscarriedoutbyalltheplantsinanecosystemiscalledthegrossprimaryproduction(GPP).[8]: 124 AbouthalfofthegrossGPPisrespiredbyplantsinordertoprovidetheenergythatsupportstheirgrowthandmaintenance.[14]: 157 Theremainder,thatportionofGPPthatisnotusedupbyrespiration,isknownasthenetprimaryproduction(NPP).[14]: 157 Totalphotosynthesisislimitedbyarangeofenvironmentalfactors.Theseincludetheamountoflightavailable,theamountofleafareaaplanthastocapturelight(shadingbyotherplantsisamajorlimitationofphotosynthesis),therateatwhichcarbondioxidecanbesuppliedtothechloroplaststosupportphotosynthesis,theavailabilityofwater,andtheavailabilityofsuitabletemperaturesforcarryingoutphotosynthesis.[8]: 155  Energyflow Mainarticle:Energyflow(ecology) Seealso:FoodwebandTrophiclevel Energyandcarbonenterecosystemsthroughphotosynthesis,areincorporatedintolivingtissue,transferredtootherorganismsthatfeedonthelivinganddeadplantmatter,andeventuallyreleasedthroughrespiration.[14]: 157 Thecarbonandenergyincorporatedintoplanttissues(netprimaryproduction)iseitherconsumedbyanimalswhiletheplantisalive,oritremainsuneatenwhentheplanttissuediesandbecomesdetritus.Interrestrialecosystems,thevastmajorityofthenetprimaryproductionendsupbeingbrokendownbydecomposers.Theremainderisconsumedbyanimalswhilestillaliveandenterstheplant-basedtrophicsystem.Afterplantsandanimalsdie,theorganicmattercontainedinthementersthedetritus-basedtrophicsystem.[15] Ecosystemrespirationisthesumofrespirationbyalllivingorganisms(plants,animals,anddecomposers)intheecosystem.[16]Netecosystemproductionisthedifferencebetweengrossprimaryproduction(GPP)andecosystemrespiration.[17]Intheabsenceofdisturbance,netecosystemproductionisequivalenttothenetcarbonaccumulationintheecosystem. Energycanalsobereleasedfromanecosystemthroughdisturbancessuchaswildfireortransferredtootherecosystems(e.g.,fromaforesttoastreamtoalake)byerosion. Inaquaticsystems,theproportionofplantbiomassthatgetsconsumedbyherbivoresismuchhigherthaninterrestrialsystems.[15]Introphicsystems,photosyntheticorganismsaretheprimaryproducers.Theorganismsthatconsumetheirtissuesarecalledprimaryconsumersorsecondaryproducers—herbivores.Organismswhichfeedonmicrobes(bacteriaandfungi)aretermedmicrobivores.Animalsthatfeedonprimaryconsumers—carnivores—aresecondaryconsumers.Eachoftheseconstitutesatrophiclevel.[15] Thesequenceofconsumption—fromplanttoherbivore,tocarnivore—formsafoodchain.Realsystemsaremuchmorecomplexthanthis—organismswillgenerallyfeedonmorethanoneformoffood,andmayfeedatmorethanonetrophiclevel.Carnivoresmaycapturesomepreythatispartofaplant-basedtrophicsystemandothersthatarepartofadetritus-basedtrophicsystem(abirdthatfeedsbothonherbivorousgrasshoppersandearthworms,whichconsumedetritus).Realsystems,withallthesecomplexities,formfoodwebsratherthanfoodchains.[15] Decomposition Seealso:Decomposition Sequenceofadecomposingpigcarcassovertime. Thecarbonandnutrientsindeadorganicmatterarebrokendownbyagroupofprocessesknownasdecomposition.Thisreleasesnutrientsthatcanthenbere-usedforplantandmicrobialproductionandreturnscarbondioxidetotheatmosphere(orwater)whereitcanbeusedforphotosynthesis.Intheabsenceofdecomposition,thedeadorganicmatterwouldaccumulateinanecosystem,andnutrientsandatmosphericcarbondioxidewouldbedepleted.[18]: 183  Decompositionprocessescanbeseparatedintothreecategories—leaching,fragmentationandchemicalalterationofdeadmaterial.Aswatermovesthroughdeadorganicmatter,itdissolvesandcarrieswithitthewater-solublecomponents.Thesearethentakenupbyorganismsinthesoil,reactwithmineralsoil,oraretransportedbeyondtheconfinesoftheecosystem(andareconsideredlosttoit).[19]: 271–280 Newlyshedleavesandnewlydeadanimalshavehighconcentrationsofwater-solublecomponentsandincludesugars,aminoacidsandmineralnutrients.Leachingismoreimportantinwetenvironmentsandlessimportantindryones.[10]: 69–77  Fragmentationprocessesbreakorganicmaterialintosmallerpieces,exposingnewsurfacesforcolonizationbymicrobes.Freshlyshedleaflittermaybeinaccessibleduetoanouterlayerofcuticleorbark,andcellcontentsareprotectedbyacellwall.Newlydeadanimalsmaybecoveredbyanexoskeleton.Fragmentationprocesses,whichbreakthroughtheseprotectivelayers,acceleratetherateofmicrobialdecomposition.[18]: 184 Animalsfragmentdetritusastheyhuntforfood,asdoespassagethroughthegut.Freeze-thawcyclesandcyclesofwettinganddryingalsofragmentdeadmaterial.[18]: 186  Thechemicalalterationofthedeadorganicmatterisprimarilyachievedthroughbacterialandfungalaction.Fungalhyphaeproduceenzymesthatcanbreakthroughthetoughouterstructuressurroundingdeadplantmaterial.Theyalsoproduceenzymesthatbreakdownlignin,whichallowsthemaccesstobothcellcontentsandthenitrogeninthelignin.Fungicantransfercarbonandnitrogenthroughtheirhyphalnetworksandthus,unlikebacteria,arenotdependentsolelyonlocallyavailableresources.[18]: 186  Decompositionrates Decompositionratesvaryamongecosystems.[20]Therateofdecompositionisgovernedbythreesetsoffactors—thephysicalenvironment(temperature,moisture,andsoilproperties),thequantityandqualityofthedeadmaterialavailabletodecomposers,andthenatureofthemicrobialcommunityitself.[18]: 194 Temperaturecontrolstherateofmicrobialrespiration;thehigherthetemperature,thefasterthemicrobialdecompositionoccurs.Temperaturealsoaffectssoilmoisture,whichaffectsdecomposition.Freeze-thawcyclesalsoaffectdecomposition—freezingtemperatureskillsoilmicroorganisms,whichallowsleachingtoplayamoreimportantroleinmovingnutrientsaround.Thiscanbeespeciallyimportantasthesoilthawsinthespring,creatingapulseofnutrientsthatbecomeavailable.[19]: 280  Decompositionratesarelowunderverywetorverydryconditions.Decompositionratesarehighestinwet,moistconditionswithadequatelevelsofoxygen.Wetsoilstendtobecomedeficientinoxygen(thisisespeciallytrueinwetlands),whichslowsmicrobialgrowth.Indrysoils,decompositionslowsaswell,butbacteriacontinuetogrow(albeitataslowerrate)evenaftersoilsbecometoodrytosupportplantgrowth.[18]: 200  Dynamicsandresilience Furtherinformation:Resistance(ecology)andEcologicalresilience Ecosystemsaredynamicentities.Theyaresubjecttoperiodicdisturbancesandarealwaysintheprocessofrecoveringfrompastdisturbances.[21]: 347 Whenaperturbationoccurs,anecosystemrespondsbymovingawayfromitsinitialstate.Thetendencyofanecosystemtoremainclosetoitsequilibriumstate,despitethatdisturbance,istermeditsresistance.Thecapacityofasystemtoabsorbdisturbanceandreorganizewhileundergoingchangesoastoretainessentiallythesamefunction,structure,identity,andfeedbacksistermeditsecologicalresilience.[22][23]Resiliencethinkingalsoincludeshumanityasanintegralpartofthebiospherewherewearedependentonecosystemservicesforoursurvivalandmustbuildandmaintaintheirnaturalcapacitiestowithstandshocksanddisturbances.[24]Timeplaysacentralroleoverawiderange,forexample,intheslowdevelopmentofsoilfrombarerockandthefasterrecoveryofacommunityfromdisturbance.[14]: 67  Disturbancealsoplaysanimportantroleinecologicalprocesses.F.StuartChapinandcoauthorsdefinedisturbanceas"arelativelydiscreteeventintimethatremovesplantbiomass".[21]: 346 Thiscanrangefromherbivoreoutbreaks,treefalls,fires,hurricanes,floods,glacialadvances,tovolcaniceruptions.Suchdisturbancescancauselargechangesinplant,animalandmicrobepopulations,aswellassoilorganicmattercontent.Disturbanceisfollowedbysuccession,a"directionalchangeinecosystemstructureandfunctioningresultingfrombioticallydrivenchangesinresourcesupply."[2]: 470  Thefrequencyandseverityofdisturbancedeterminethewayitaffectsecosystemfunction.Amajordisturbancelikeavolcaniceruptionorglacialadvanceandretreatleavebehindsoilsthatlackplants,animalsororganicmatter.Ecosystemsthatexperiencesuchdisturbancesundergoprimarysuccession.Alessseveredisturbancelikeforestfires,hurricanesorcultivationresultinsecondarysuccessionandafasterrecovery.[21]: 348 Moresevereandmorefrequentdisturbanceresultinlongerrecoverytimes. Fromoneyeartoanother,ecosystemsexperiencevariationintheirbioticandabioticenvironments.Adrought,acolderthanusualwinter,andapestoutbreakallareshort-termvariabilityinenvironmentalconditions.Animalpopulationsvaryfromyeartoyear,buildingupduringresource-richperiodsandcrashingastheyovershoottheirfoodsupply.Longer-termchangesalsoshapeecosystemprocesses.Forexample,theforestsofeasternNorthAmericastillshowlegaciesofcultivationwhichceasedin1850whenlargeareaswererevertedtoforests.[21]: 340 AnotherexampleisthemethaneproductionineasternSiberianlakesthatiscontrolledbyorganicmatterwhichaccumulatedduringthePleistocene.[25] AfreshwaterlakeinGranCanaria,anislandoftheCanaryIslands.Clearboundariesmakelakesconvenienttostudyusinganecosystemapproach. Nutrientcycling Seealso:Nutrientcycle,Biogeochemicalcycle,andNitrogencycle Biologicalnitrogencycling Ecosystemscontinuallyexchangeenergyandcarbonwiththewiderenvironment.Mineralnutrients,ontheotherhand,aremostlycycledbackandforthbetweenplants,animals,microbesandthesoil.Mostnitrogenentersecosystemsthroughbiologicalnitrogenfixation,isdepositedthroughprecipitation,dust,gasesorisappliedasfertilizer.[19]: 266 Mostterrestrialecosystemsarenitrogen-limitedintheshorttermmakingnitrogencyclinganimportantcontrolonecosystemproduction.[19]: 289 Overthelongterm,phosphorusavailabilitycanalsobecritical.[26] Macronutrientswhicharerequiredbyallplantsinlargequantitiesincludetheprimarynutrients(whicharemostlimitingastheyareusedinlargestamounts):Nitrogen,phosphorus,potassium.[27]: 231 Secondarymajornutrients(lessoftenlimiting)include:Calcium,magnesium,sulfur.Micronutrientsrequiredbyallplantsinsmallquantitiesincludeboron,chloride,copper,iron,manganese,molybdenum,zinc.Finally,therearealsobeneficialnutrientswhichmayberequiredbycertainplantsorbyplantsunderspecificenvironmentalconditions:aluminum,cobalt,iodine,nickel,selenium,silicon,sodium,vanadium.[27]: 231  Untilmoderntimes,nitrogenfixationwasthemajorsourceofnitrogenforecosystems.Nitrogen-fixingbacteriaeitherlivesymbioticallywithplantsorlivefreelyinthesoil.Theenergeticcostishighforplantsthatsupportnitrogen-fixingsymbionts—asmuchas25%ofgrossprimaryproductionwhenmeasuredincontrolledconditions.Manymembersofthelegumeplantfamilysupportnitrogen-fixingsymbionts.Somecyanobacteriaarealsocapableofnitrogenfixation.Thesearephototrophs,whichcarryoutphotosynthesis.Likeothernitrogen-fixingbacteria,theycaneitherbefree-livingorhavesymbioticrelationshipswithplants.[21]: 360 Othersourcesofnitrogenincludeaciddepositionproducedthroughthecombustionoffossilfuels,ammoniagaswhichevaporatesfromagriculturalfieldswhichhavehadfertilizersappliedtothem,anddust.[19]: 270 Anthropogenicnitrogeninputsaccountforabout80%ofallnitrogenfluxesinecosystems.[19]: 270  Whenplanttissuesareshedorareeaten,thenitrogeninthosetissuesbecomesavailabletoanimalsandmicrobes.Microbialdecompositionreleasesnitrogencompoundsfromdeadorganicmatterinthesoil,whereplants,fungi,andbacteriacompeteforit.Somesoilbacteriauseorganicnitrogen-containingcompoundsasasourceofcarbon,andreleaseammoniumionsintothesoil.Thisprocessisknownasnitrogenmineralization.Othersconvertammoniumtonitriteandnitrateions,aprocessknownasnitrification.Nitricoxideandnitrousoxidearealsoproducedduringnitrification.[19]: 277 Undernitrogen-richandoxygen-poorconditions,nitratesandnitritesareconvertedtonitrogengas,aprocessknownasdenitrification.[19]: 281  Mycorrhizalfungiwhicharesymbioticwithplantroots,usecarbohydratessuppliedbytheplantsandinreturntransferphosphorusandnitrogencompoundsbacktotheplantroots.[28][29]Thisisanimportantpathwayoforganicnitrogentransferfromdeadorganicmattertoplants.Thismechanismmaycontributetomorethan70Tgofannuallyassimilatedplantnitrogen,therebyplayingacriticalroleinglobalnutrientcyclingandecosystemfunction.[29] Phosphorusentersecosystemsthroughweathering.Asecosystemsagethissupplydiminishes,makingphosphorus-limitationmorecommoninolderlandscapes(especiallyinthetropics).[19]: 287–290 Calciumandsulfurarealsoproducedbyweathering,butaciddepositionisanimportantsourceofsulfurinmanyecosystems.Althoughmagnesiumandmanganeseareproducedbyweathering,exchangesbetweensoilorganicmatterandlivingcellsaccountforasignificantportionofecosystemfluxes.Potassiumisprimarilycycledbetweenlivingcellsandsoilorganicmatter.[19]: 291  Functionandbiodiversity Mainarticle:BiodiversitySeealso:Ecosystemdiversity LochLomondinScotlandformsarelativelyisolatedecosystem.Thefishcommunityofthislakehasremainedstableoveralongperioduntilanumberofintroductionsinthe1970srestructureditsfoodweb.[30] SpinyforestatIfaty,Madagascar,featuringvariousAdansonia(baobab)species,Alluaudiaprocera(Madagascarocotillo)andothervegetation. Biodiversityplaysanimportantroleinecosystemfunctioning.[31]: 449–453 Ecosystemprocessesaredrivenbythespeciesinanecosystem,thenatureoftheindividualspecies,andtherelativeabundanceoforganismsamongthesespecies.Ecosystemprocessesaretheneteffectoftheactionsofindividualorganismsastheyinteractwiththeirenvironment.Ecologicaltheorysuggeststhatinordertocoexist,speciesmusthavesomeleveloflimitingsimilarity—theymustbedifferentfromoneanotherinsomefundamentalway,otherwise,onespecieswouldcompetitivelyexcludetheother.[32]Despitethis,thecumulativeeffectofadditionalspeciesinanecosystemisnotlinear:additionalspeciesmayenhancenitrogenretention,forexample.However,beyondsomelevelofspeciesrichness,[11]: 331 additionalspeciesmayhavelittleadditiveeffectunlesstheydiffersubstantiallyfromspeciesalreadypresent.[11]: 324 Thisisthecaseforexampleforexoticspecies.[11]: 321  Theaddition(orloss)ofspeciesthatareecologicallysimilartothosealreadypresentinanecosystemtendstoonlyhaveasmalleffectonecosystemfunction.Ecologicallydistinctspecies,ontheotherhand,haveamuchlargereffect.Similarly,dominantspecieshavealargeeffectonecosystemfunction,whilerarespeciestendtohaveasmalleffect.Keystonespeciestendtohaveaneffectonecosystemfunctionthatisdisproportionatetotheirabundanceinanecosystem.[11]: 324  Anecosystemengineerisanyorganismthatcreates,significantlymodifies,maintainsordestroysahabitat.[33] Studyapproaches Ecosystemecology Mainarticle:Ecosystemecology Seealso:Ecosystemmodel Ahydrothermalventisanecosystemontheoceanfloor.(Thescalebaris1m.) Ecosystemecologyisthe"studyoftheinteractionsbetweenorganismsandtheirenvironmentasanintegratedsystem".[2]: 458 Thesizeofecosystemscanrangeuptotenordersofmagnitude,fromthesurfacelayersofrockstothesurfaceoftheplanet.[4]: 6  TheHubbardBrookEcosystemStudystartedin1963tostudytheWhiteMountainsinNewHampshire.Itwasthefirstsuccessfulattempttostudyanentirewatershedasanecosystem.Thestudyusedstreamchemistryasameansofmonitoringecosystemproperties,anddevelopedadetailedbiogeochemicalmodeloftheecosystem.[34]Long-termresearchatthesiteledtothediscoveryofacidraininNorthAmericain1972.Researchersdocumentedthedepletionofsoilcations(especiallycalcium)overthenextseveraldecades.[35] Ecosystemscanbestudiedthroughavarietyofapproaches—theoreticalstudies,studiesmonitoringspecificecosystemsoverlongperiodsoftime,thosethatlookatdifferencesbetweenecosystemstoelucidatehowtheyworkanddirectmanipulativeexperimentation.[36]Studiescanbecarriedoutatavarietyofscales,rangingfromwhole-ecosystemstudiestostudyingmicrocosmsormesocosms(simplifiedrepresentationsofecosystems).[37]AmericanecologistStephenR.Carpenterhasarguedthatmicrocosmexperimentscanbe"irrelevantanddiversionary"iftheyarenotcarriedoutinconjunctionwithfieldstudiesdoneattheecosystemscale.Insuchcases,microcosmexperimentsmayfailtoaccuratelypredictecosystem-leveldynamics.[38] Classifications Furtherinformation:EcosystemclassificationandBiogeoclimaticecosystemclassification Biomesaregeneralclassesorcategoriesofecosystems.[4]: 14 However,thereisnocleardistinctionbetweenbiomesandecosystems.[39]Biomesarealwaysdefinedataverygenerallevel.Ecosystemscanbedescribedatlevelsthatrangefromverygeneral(inwhichcasethenamesaresometimesthesameasthoseofbiomes)toveryspecific,suchas"wetcoastalneedle-leafedforests". Biomesvaryduetoglobalvariationsinclimate.Biomesareoftendefinedbytheirstructure:atagenerallevel,forexample,tropicalforests,temperategrasslands,andarctictundra.[4]: 14 Therecanbeanydegreeofsubcategoriesamongecosystemtypesthatcompriseabiome,e.g.,needle-leafedborealforestsorwettropicalforests.Althoughecosystemsaremostcommonlycategorizedbytheirstructureandgeography,therearealsootherwaystocategorizeandclassifyecosystemssuchasbytheirlevelofhumanimpact(seeanthropogenicbiome),orbytheirintegrationwithsocialprocessesortechnologicalprocessesortheirnovelty(e.g.novelecosystem).Eachofthesetaxonomiesofecosystemstendstoemphasizedifferentstructuralorfunctionalproperties.[40]Noneoftheseisthe“best”classification. Ecosystemclassificationsarespecifickindsofecologicalclassificationsthatconsiderallfourelementsofthedefinitionofecosystems:abioticcomponent,anabioticcomplex,theinteractionsbetweenandwithinthem,andthephysicalspacetheyoccupy.[40]Differentapproachestoecologicalclassificationshavebeendevelopedinterrestrial,freshwaterandmarinedisciplines. Examples Thefollowingarticlesareexamplesofecosystemsforparticularregions,zonesorconditions: Aquaticecosystem Borealecosystem Freshwaterecosystem Groundwater-dependentecosystems Lakeecosystem(lenticecosystem) Largemarineecosystem Marineecosystem Montaneecosystem Riverecosystem(loticecosystem) Terrestrialecosystem Urbanecosystem Humaninteractionswithecosystems Humanactivitiesareimportantinalmostallecosystems.Althoughhumansexistandoperatewithinecosystems,theircumulativeeffectsarelargeenoughtoinfluenceexternalfactorslikeclimate.[4]: 14  Ecosystemgoodsandservices TheHighPeaksWildernessAreainthe6,000,000-acre(2,400,000 ha)AdirondackParkisanexampleofadiverseecosystem. Mainarticles:EcosystemservicesandEcologicalgoodsandservices Seealso:EcosystemvaluationandEcologicalyield Ecosystemsprovideavarietyofgoodsandservicesuponwhichpeopledepend.[41]Ecosystemgoodsincludethe"tangible,materialproducts"ofecosystemprocessessuchaswater,food,fuel,constructionmaterial,andmedicinalplants.[42][43]Theyalsoincludelesstangibleitemsliketourismandrecreation,andgenesfromwildplantsandanimalsthatcanbeusedtoimprovedomesticspecies.[41] Ecosystemservices,ontheotherhand,aregenerally"improvementsintheconditionorlocationofthingsofvalue".[43]Theseincludethingslikethemaintenanceofhydrologicalcycles,cleaningairandwater,themaintenanceofoxygenintheatmosphere,croppollinationandeventhingslikebeauty,inspirationandopportunitiesforresearch.[41]Whilematerialfromtheecosystemhadtraditionallybeenrecognizedasbeingthebasisforthingsofeconomicvalue,ecosystemservicestendtobetakenforgranted.[43] TheMillenniumEcosystemAssessmentisaninternationalsynthesisbyover1000oftheworld'sleadingbiologicalscientiststhatanalyzesthestateoftheEarth'secosystemsandprovidessummariesandguidelinesfordecision-makers.Thereportidentifiedfourmajorcategoriesofecosystemservices:provisioning,regulating,culturalandsupportingservices.[44]Itconcludesthathumanactivityishavingasignificantandescalatingimpactonthebiodiversityoftheworldecosystems,reducingboththeirresilienceandbiocapacity.Thereportreferstonaturalsystemsashumanity's"life-supportsystem",providingessentialecosystemservices.Theassessmentmeasures24ecosystemservicesandconcludesthatonlyfourhaveshownimprovementoverthelast50years,15areinseriousdecline,andfiveareinaprecariouscondition.[44]: 6–19  TheIntergovernmentalScience-PolicyPlatformonBiodiversityandEcosystemServices(IPBES)isanintergovernmentalorganizationestablishedtoimprovetheinterfacebetweenscienceandpolicyonissuesofbiodiversityandecosystemservices.[45]ItisintendedtoserveasimilarroletotheIntergovernmentalPanelonClimateChange.[46]TheconceptualframeworkoftheIPBESincludessixprimaryinterlinkedelements:nature,nature’sbenefitstopeople,anthropogenicassets,institutionsandgovernancesystemsandotherindirectdriversofchange,directdriversofchange,andgoodqualityoflife.[47] Ecosystemdegradationanddecline TheForestLandscapeIntegrityIndexmeasuresglobalanthropogenicmodificationonremainingforestsannually.0=Mostmodification;10=Least.[48] Seealso:Ecosystemcollapse,Climatechangeandecosystems,andHumanecology Ashumanpopulationandpercapitaconsumptiongrow,sodotheresourcedemandsimposedonecosystemsandtheeffectsofthehumanecologicalfootprint.Naturalresourcesarevulnerableandlimited.Theenvironmentalimpactsofanthropogenicactionsarebecomingmoreapparent.Problemsforallecosystemsinclude:environmentalpollution,climatechangeandbiodiversityloss.Forterrestrialecosystemsfurtherthreatsincludeairpollution,soildegradation,anddeforestation.Foraquaticecosystemsthreatsalsoincludeunsustainableexploitationofmarineresources(forexampleoverfishing),marinepollution,microplasticspollution,theeffectsofclimatechangeonoceans(e.g.warmingandacidification),andbuildingoncoastalareas.[49] Manyecosystemsbecomedegradedthroughhumanimpacts,suchassoilloss,airandwaterpollution,habitatfragmentation,waterdiversion,firesuppression,andintroducedspeciesandinvasivespecies.[50]: 437  Thesethreatscanleadtoabrupttransformationoftheecosystemortogradualdisruptionofbioticprocessesanddegradationofabioticconditionsoftheecosystem.Oncetheoriginalecosystemhaslostitsdefiningfeatures,itisconsideredcollapsed(seealsoIUCNRedListofEcosystems).[51]Ecosystemcollapsecouldbereversibleandinthiswaydiffersfromspeciesextinction.[52]Quantitativeassessmentsoftheriskofcollapseareusedasmeasuresofconservationstatusandtrends. Ecosystemmanagement Mainarticles:Ecosystemmanagement,Ecosystem-basedmanagement,andEcosystemapproach Whennaturalresourcemanagementisappliedtowholeecosystems,ratherthansinglespecies,itistermedecosystemmanagement.[53]Althoughdefinitionsofecosystemmanagementabound,thereisacommonsetofprincipleswhichunderliethesedefinitions:Afundamentalprincipleisthelong-termsustainabilityoftheproductionofgoodsandservicesbytheecosystem;[50]"intergenerationalsustainability[is]apreconditionformanagement,notanafterthought".[41]Whileecosystemmanagementcanbeusedaspartofaplanforwildernessconservation,itcanalsobeusedinintensivelymanagedecosystems[41](see,forexample,agroecosystemandclosetonatureforestry). Ecosystemrestorationandsustainabledevelopment Seealso:RestorationecologyandUNDecadeonEcosystemRestoration Societyisincreasinglybecomingawarethatecosystemservicesarenotonlylimitedbutalsothattheyarethreatenedbyhumanactivities.Tohelpinformdecision-makers,manyecosystemservicesarebeingassignedeconomicvalues,oftenbasedonthecostofreplacementwithanthropogenicalternatives.Theongoingchallengeofprescribingeconomicvaluetonature,forexamplethroughbiodiversitybanking,ispromptingtransdisciplinaryshiftsinhowwerecognizeandmanagetheenvironment,socialresponsibility,businessopportunities,andourfutureasaspecies.[citationneeded] Ecosystemrestorationwillcontributetoall17SustainableDevelopmentGoals,inparticulartoSDG2(ZeroHunger),SDG6(CleanWaterandSanitation),SDG14(Lifebelowwater)andSDG15(LifeonLand).[54][citationneeded]Paragraph27oftheMinisterialDeclarationoftheHigh-LevelPoliticalForumontheSDGsheldinJuly2018setsoutcommitmentsmadetoachievesustainablemanagementofalltypesofforests,haltdeforestation,restoredegradedforests,andsubstantiallyincreaseafforestationandreforestationgloballyby2020.[55] Integratedconservationanddevelopmentprojects(ICDPs)aimtoaddressconservationandhumanlivelihood(sustainabledevelopment)concernsindevelopingcountriestogether,ratherthanseparatelyaswasoftendoneinthepast.[50]: 445  Seealso Earthsciencesportal Ecologyportal Environmentportal Complexsystem Earthscience Ecosystem-basedadaptation Ecosystemsinspecificregionsoftheworld: LeuserEcosystem Longleafpineecosystem TarangireEcosystem Tropicalsaltpondecosystem Ecosystemsgroupedbycondition: Agroecosystem Closedecosystem Depauperateecosystem 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