History of Earth

6/19/2017 HistoryofEarthWikipedia
HistoryofEarth
FromWikipedia,thefreeencyclopedia
ThehistoryofEarthconcerns
thedevelopmentofplanetEarth
fromitsformationtothepresent
day.[1][2]Nearlyallbranchesof
naturalsciencehavecontributed
totheunderstandingofthemain
eventsofEarth'spast.Theageof
Earthisapproximatelyonethird
oftheageoftheuniverse.An
immenseamountofgeological
changehasoccurredinthat
timespan,accompaniedbythe
emergenceoflifeandits
subsequentevolution.
Earthformedaround4.54billion
yearsagobyaccretionfromthe
solarnebula.[3][4][5]Volcanic
outgassingprobablycreatedthe
primordialatmosphereandthen
theoceanbuttheatmosphere
containedalmostnooxygenand
sowouldhavebeentoxictomost
modernlifeincludinghumans.
MuchoftheEarthwasmolten
becauseoffrequentcollisions
withotherbodieswhichledto
extremevolcanism.AgiantimpactcollisionwithaplanetsizedbodynamedTheiawhileEarthwasinitsearliest
stage,alsoknownasEarlyEarth,isthoughttohavebeenresponsibleforformingtheMoon.Overtime,theEarth
cooled,causingtheformationofasolidcrust,andallowingliquidwatertoexistonthesurface.
Thegeologicaltimescale(GTS)clock(seegraphic)depictsthelargerspansoftimefromthebeginningofthe
EarthaswellasachronologyofsomedefinitiveeventsofEarthhistory.TheHadeanEonrepresentstimebefore
thereliable(fossil)recordoflifebeginningonEarthitbeganwiththeformationoftheplanetandendedat4.0
billionyearsagoasdefinedbyinternationalconvention.[6]TheArcheanandProterozoiceonsfollowthey
producedtheabiogenesisoflifeonEarthandthentheevolutionofearlylife.Thesucceedingeonisthe
Phanerozoic,whichisrepresentedbyitsthreecomponenteras:thePalaeozoictheMesozoic,whichspannedthe
rise,reign,andclimacticextinctionofthehugedinosaursandtheCenozoic,whichpresentedthesubsequent
developmentofdominantmammalsonEarth.
Hominins,theearliestdirectancestorsofthehumanclade,rosesometimeduringthelatterpartoftheMiocene
epochtheprecisetimemarkingthefirsthomininsisbroadlydebatedoveracurrentrangeof13to4mya.The
succeedingQuaternaryperiodisthetimeofrecognizablehumans,i.e.,thegenusHomo,butthatperiod'stwo
millionyearplustermoftherecenttimesistoosmalltobevisibleatthescaleoftheGTSgraphic.(Notesrethe
graphic:Gameans"billionyears"Ma,"millionyears".)
https://en.wikipedia.org/wiki/History_of_Earth 1/32
TheearliestundisputedevidenceoflifeonEarthdatesatleastfrom3.5billionyearsago,[7][8][9]duringthe
EoarcheanEraafterageologicalcruststartedtosolidifyfollowingtheearliermoltenHadeanEon.Thereare
microbialmatfossilssuchasstromatolitesfoundin3.48billionyearoldsandstonediscoveredinWestern
Australia.[10][11][12]Otherearlyphysicalevidenceofabiogenicsubstanceisgraphitein3.7billionyearold
metasedimentaryrocksdiscoveredinsouthwesternGreenland[13]aswellas"remainsofbioticlife"foundin4.1
billionyearoldrocksinWesternAustralia.[14][15]Accordingtooneoftheresearchers,"Iflifearoserelatively
quicklyonEarththenitcouldbecommonintheuniverse."[14]
Livingformsderivedfromphotosynthesisappearedbetween3.2and2.4billionyearsagoandbeganenrichingthe
atmospherewithoxygen.Liferemainedmostlysmallandmicroscopicuntilabout580millionyearsago,when
complexmulticellularlifearose,developedovertime,andculminatedintheCambrianExplosionabout541
millionyearsago.ThiseventdrovearapiddiversificationoflifeformsonEarththatproducedmostofthemajor
phylaknowntodayanditmarkedtheendoftheProterozoicEonandthebeginningoftheCambrianPeriodofthe
PaleozoicEra.Morethan99percentofallspecies,amountingtooverfivebillionspecies,[16]thateverlivedon
Earthareestimatedtobeextinct.[17][18]EstimatesonthenumberofEarth'scurrentspeciesrangefrom10million
to14million,[19]ofwhichabout1.2millionhavebeendocumentedandover86percenthavenotyetbeen
described.[20]Morerecently,inMay2016,scientistsreportedthat1trillionspeciesareestimatedtobeonEarth
currentlywithonlyonethousandthofonepercentdescribed.[21]
GeologicalchangehasbeenaconstantofEarth'scrustsincethetimeofitsformation,andbiologicalchangesince
thefirstappearanceoflife.Speciescontinuetoevolve,takingonnewforms,splittingintodaughterspeciesor
goingextinctintheprocessofadaptingordyinginresponsetoeverchangingphysicalenvironments.Theprocess
ofplatetectonicscontinuestoplayadominantroleintheshapingofEarth'soceansandcontinentsandtheliving
speciestheyharbor.Changesinthebiospherenowdominatedbyhumanactivitycontinue,inturn,toproduce
significanteffectsontheatmosphereandothersystemsoftheEarth'ssurface,suchastheintegrityoftheozone
layer,theproliferationofgreenhousegases,theconditionsofproductivesoilsandcleanairandwater,andothers.
Contents
1 Eons
2 Geologictimescale
3 SolarSystemformation
4 HadeanandArcheanEons
4.1 FormationoftheMoon
4.2 Firstcontinents
4.3 Oceansandatmosphere
4.4 Originoflife
5 ProterozoicEon
5.1 Oxygenrevolution
5.2 SnowballEarth
5.3 Emergenceofeukaryotes
5.4 SupercontinentsintheProterozoic
5.5 LateProterozoicclimateandlife
6 PhanerozoicEon
6.1 Tectonics,paleogeographyandclimate
6.2 Cambrianexplosion
6.3 Colonizationofland
6.4 Evolutionoftetrapods
6.5 Extinctions
6.6 Diversificationofmammals
6.7 Humanevolution
7 Seealso
8 Notes
9 References
10 Furtherreading
11 Externallinks
Eons
Ingeochronology,timeisgenerallymeasuredinmya(megayearsormillionyearsago),eachunitrepresentingthe
periodofapproximately1,000,000yearsinthepast.ThehistoryofEarthisdividedintofourgreateons,starting
4,540myawiththeformationoftheplanet.EacheonsawthemostsignificantchangesinEarth'scomposition,
climateandlife.Eacheonissubsequentlydividedintoeras,whichinturnaredividedintoperiods,whichare
furtherdividedintoepochs.
Time
Eon Description
(mya)
TheEarthisformedoutofdebrisaroundthesolarprotoplanetarydisk.Thereisnolife.
4,540 Temperaturesareextremelyhot,withfrequentvolcanicactivityandhellishenvironments.
Hadean
4,000 Theatmosphereisnebular.Possibleearlyoceansorbodiesofliquidwater.Themoonis
formedaroundthistime,probablyduetoaprotoplanet'scollisionintoEarth.
Prokaryotelife,thefirstformoflife,emergesattheverybeginningofthiseon,inaprocess
4,000
Archean knownasabiogenesis.ThecontinentsofUr,VaalbaraandKenorlandmayhavebeen
2,500
formedaroundthistime.Theatmosphereiscomposedofvolcanicandgreenhousegases.
Eukaryotes,amorecomplexformoflife,emerge,includingsomeformsofmulticellular
organisms.Bacteriabeginproducingoxygen,shapingthethirdandcurrentofEarth's
2,500 atmospheres.Plants,lateranimalsandpossiblyearlierformsoffungiformaroundthis
Proterozoic
541 time.Theearlyandlatephasesofthiseonmayhaveundergone"SnowballEarth"periods,
inwhichalloftheplanetsufferedbelowzerotemperatures.Theearlycontinentsof
Columbia,RodiniaandPannotiamayhaveformedaroundthistime,inthatorder.
Complexlife,includingvertebrates,begintodominatetheEarth'soceaninaprocessknown
astheCambrianexplosion.PangaeaformsandlaterdissolvesintoLaurasiaandGondwana.
541 Gradually,lifeexpandstolandandallfamiliarformsofplants,animalsandfungibegin
Phanerozoic
present appearing,includingannelids,insectsandreptiles.Severalmassextinctionsoccur,among
whichbirds,thedescendantsofdinosaurs,andmorerecentlymammalsemerge.Therise
andevolutionofthehumanspeciesoccurattheveryrecentphasesofthiseon.
Geologictimescale
ThehistoryoftheEarthcanbeorganizedchronologicallyaccordingtothegeologictimescale,whichissplitinto
intervalsbasedonstratigraphicanalysis.[2][22]Thefollowingfourtimelinesshowthegeologictimescale.Thefirst
showstheentiretimefromtheformationoftheEarthtothepresent,butthiscompressesthemostrecenteon.
Therefore,thesecondscaleshowsthemostrecenteonwithanexpandedscale.Thesecondscalecompressesthe
mostrecentera,sothemostrecenteraisexpandedinthethirdscale.Thethirdscalecompressesthemostrecent
period,sothemostrecentperiodisexpandedinthefourthscale.
MillionsofYears
SolarSystemformation
ThestandardmodelfortheformationoftheSolar
System(includingtheEarth)isthesolarnebula Naturetimeline
hypothesis.[23]Inthismodel,theSolarSystemformed viewdiscuss
fromalarge,rotatingcloudofinterstellardustandgas 0 Earliesthumans
calledthesolarnebula.Itwascomposedofhydrogen Landlife
Cambrianexplosion
andheliumcreatedshortlyaftertheBigBang13.8Ga 1 Multicellular Earliestsexual
(billionyearsago)andheavierelementsejectedby
life reproduction
supernovae.About4.5Ga,thenebulabegana L
2 i
contractionthatmayhavebeentriggeredbytheshock
f Atmosphericoxygen
waveofanearbysupernova.[24]Ashockwavewould e
3 photosynthesis
havealsomadethenebularotate.Asthecloudbegan
toaccelerate,itsangularmomentum,gravityand Earliestoxygen
inertiaflatteneditintoaprotoplanetarydisk 4 Singlecelledlife
water
Earliestlife
perpendiculartoitsaxisofrotation.Small EarliestEarth(4.54)
SolarSystem
perturbationsduetocollisionsandtheangular 5
momentumofotherlargedebriscreatedthemeansby
whichkilometersizedprotoplanetsbegantoform, 6 cosmicspeedup
orbitingthenebularcenter.[25] AlphaCentauriforms
7
P
r
https://en.wikipedia.org/wiki/History_of_Earth
8 4/32
r
Thecenterofthenebula,nothavingmuchangular 8 i
Darkmatter
momentum,collapsedrapidly,thecompression m Darkenergy
o MilkyWayGalaxy
heatingituntilnuclearfusionofhydrogenintohelium 9 r spiralarmsform
began.Aftermorecontraction,aTTauristarignited
d
andevolvedintotheSun.Meanwhile,intheouterpart 10 i AndromedaGalaxy
a
ofthenebulagravitycausedmattertocondense forms
l
arounddensityperturbationsanddustparticles,and 11 cosmicexpansion
therestoftheprotoplanetarydiskbeganseparating OmegaCentauriforms
intorings.Inaprocessknownasrunawayaccretion,
12
successivelylargerfragmentsofdustanddebris

clumpedtogethertoformplanets.[25]Earthformedin 13
Earliestlight
Earliestquasar
thismannerabout4.54billionyearsago(withan Earliestgalaxy
Earliestgravity
uncertaintyof1%)[26][27][3][28]andwaslargely Earliestuniverse
Axisscale:billionsofyears.
(13.80)
completedwithin1020millionyears.[29]Thesolar
Alsosee:HumantimelineandLifetimeline
windofthenewlyformedTTauristarclearedout
mostofthematerialinthediskthathadnotalready
condensedintolargerbodies.Thesameprocessis
expectedtoproduceaccretiondisksaroundvirtuallyall
newlyformingstarsintheuniverse,someofwhich
yieldplanets.[30]
TheprotoEarthgrewbyaccretionuntilitsinteriorwas
hotenoughtomelttheheavy,siderophilemetals.
Havinghigherdensitiesthanthesilicates,thesemetals
sank.Thissocalledironcatastropheresultedinthe
separationofaprimitivemantleanda(metallic)core
Anartist'srenderingofaprotoplanetarydisk only10millionyearsaftertheEarthbegantoform,
producingthelayeredstructureofEarthandsettingup
[31] [32]
theformationofEarth'smagneticfield. J.A.Jacobs wasthefirsttosuggestthattheinnercoreasolid
centerdistinctfromtheliquidoutercoreisfreezingandgrowingoutoftheliquidoutercoreduetothegradual
coolingofEarth'sinterior(about100degreesCelsiusperbillionyears[33]).
HadeanandArcheanEons
ThefirsteoninEarth'shistory,theHadean,beginswiththeEarth'sformationandisfollowedbytheArcheaneon
at3.8Ga.[2]:145TheoldestrocksfoundonEarthdatetoabout4.0Ga,andtheoldestdetritalzirconcrystalsin
rockstoabout4.4Ga,[34][35][36]soonaftertheformationoftheEarth'scrustandtheEarthitself.Thegiantimpact
hypothesisfortheMoon'sformationstatesthatshortlyafterformationofaninitialcrust,theprotoEarthwas
impactedbyasmallerprotoplanet,whichejectedpartofthemantleandcrustintospaceandcreatedthe
Moon.[37][38][39]
Fromcratercountsonothercelestialbodiesitisinferredthataperiodofintensemeteoriteimpacts,calledtheLate
HeavyBombardment,beganabout4.1Ga,andconcludedaround3.8Ga,attheendoftheHadean.[40]Inaddition,
volcanismwassevereduetothelargeheatflowandgeothermalgradient.[41]Nevertheless,detritalzirconcrystals
datedto4.4Gashowevidenceofhavingundergonecontactwithliquidwater,suggestingthattheEarthalready
hadoceansorseasatthattime.[34]
BythebeginningoftheArchean,theEarthhadcooledsignificantly.
PresentlifeformscouldnothavesurvivedatEarth'ssurface,becausethe
Archeanatmospherelackedoxygenhencehadnoozonelayertoblock
ultravioletlight.Nevertheless,itisbelievedthatprimordiallifebeganto
evolvebytheearlyArchean,withcandidatefossilsdatedtoaround
3.5Ga.[42]Somescientistsevenspeculatethatlifecouldhavebegunduring
theearlyHadean,asfarbackas4.4Ga,survivingthepossibleLateHeavy
BombardmentperiodinhydrothermalventsbelowtheEarth'ssurface.[43]
FormationoftheMoon
Earth'sonlynaturalsatellite,theMoon,islargerrelativetoitsplanetthan
anyothersatelliteinthesolarsystem.[nb1]DuringtheApolloprogram, Artist'sconceptionofHadeanEon
rocksfromtheMoon'ssurfacewerebroughttoEarth.Radiometricdating Earth,whenitwasmuchhotterand
inhospitabletoallformsoflife.
oftheserockshasshownthattheMoonis4.53.01billionyearsold,[46]
formedatleast30millionyearsafterthesolarsystem.[47]Newevidence
suggeststheMoonformedevenlater,4.480.02Ga,or70110million
yearsafterthestartoftheSolarSystem.[48]
TheoriesfortheformationoftheMoonmustexplainitslateformationas
wellasthefollowingfacts.First,theMoonhasalowdensity(3.3times
thatofwater,comparedto5.5fortheearth[49])andasmallmetalliccore.
Second,thereisvirtuallynowaterorothervolatilesonthemoon.Third,
theEarthandMoonhavethesameoxygenisotopicsignature(relative
abundanceoftheoxygenisotopes).Ofthetheoriesthathavebeenproposed
toaccountforthesephenomena,onlyoneiswidelyaccepted:Thegiant Artist'simpressionoftheenormous
impacthypothesisproposesthattheMoonoriginatedafterabodythesize collisionthatprobablyformedthe
ofMars(sometimesnamedTheia[47])strucktheprotoEarthaglancing Moon
blow.[1]:256[50][51]
Thecollisionreleasedabout100milliontimesmoreenergythanthemorerecentChicxulubimpactthatisbelieved
tohavecausedtheextinctionofthedinosaurs.ItwasenoughtovaporizesomeoftheEarth'souterlayersandmelt
bothbodies.[50][1]:256AportionofthemantlematerialwasejectedintoorbitaroundtheEarth.Thegiantimpact
hypothesispredictsthattheMoonwasdepletedofmetallicmaterial,[52]explainingitsabnormalcomposition.[53]
TheejectainorbitaroundtheEarthcouldhavecondensedintoasinglebodywithinacoupleofweeks.Underthe
influenceofitsowngravity,theejectedmaterialbecameamoresphericalbody:theMoon.[54]
Firstcontinents
Mantleconvection,theprocessthatdrivesplatetectonicstoday,isaresultofheatflowfromtheEarth'sinteriorto
theEarth'ssurface.[55]:2Itinvolvesthecreationofrigidtectonicplatesatmidoceanicridges.Theseplatesare
destroyedbysubductionintothemantleatsubductionzones.DuringtheearlyArchean(about3.0Ga)themantle
wasmuchhotterthantoday,probablyaround1,600C(2,910F),[56]:82soconvectioninthemantlewasfaster.
Althoughaprocesssimilartopresentdayplatetectonicsdidoccur,thiswouldhavegonefastertoo.Itislikelythat
duringtheHadeanandArchean,subductionzonesweremorecommon,andthereforetectonicplateswere
smaller.[1]:258[57]
Theinitialcrust,formedwhentheEarth'ssurfacefirstsolidified,totally
disappearedfromacombinationofthisfastHadeanplatetectonicsandthe
intenseimpactsoftheLateHeavyBombardment.However,itisthought
thatitwasbasalticincomposition,liketoday'soceaniccrust,becauselittle
crustaldifferentiationhadyettakenplace.[1]:258Thefirstlargerpiecesof
continentalcrust,whichisaproductofdifferentiationoflighterelements
duringpartialmeltinginthelowercrust,appearedattheendoftheHadean,
about4.0Ga.Whatisleftofthesefirstsmallcontinentsarecalledcratons.
ThesepiecesoflateHadeanandearlyArcheancrustformthecoresaround
whichtoday'scontinentsgrew.[58]
TheoldestrocksonEartharefoundintheNorthAmericancratonof GeologicmapofNorthAmerica,
Canada.Theyaretonalitesfromabout4.0Ga.Theyshowtracesof colorcodedbyage.Theredsand
metamorphismbyhightemperature,butalsosedimentarygrainsthathave pinksindicaterockfromtheArchean.
beenroundedbyerosionduringtransportbywater,showingthatriversand
seasexistedthen.[59]Cratonsconsistprimarilyoftwoalternatingtypesof
terranes.Thefirstaresocalledgreenstonebelts,consistingoflowgrademetamorphosedsedimentaryrocks.These
"greenstones"aresimilartothesedimentstodayfoundinoceanictrenches,abovesubductionzones.Forthis
reason,greenstonesaresometimesseenasevidenceforsubductionduringtheArchean.Thesecondtypeisa
complexoffelsicmagmaticrocks.Theserocksaremostlytonalite,trondhjemiteorgranodiorite,typesofrock
similarincompositiontogranite(hencesuchterranesarecalledTTGterranes).TTGcomplexesareseenasthe
relictsofthefirstcontinentalcrust,formedbypartialmeltinginbasalt.[60]:Chapter5
Oceansandatmosphere
Earthisoftendescribedashavinghadthreeatmospheres.The
firstatmosphere,capturedfromthesolarnebula,was
composedoflight(atmophile)elementsfromthesolarnebula,
mostlyhydrogenandhelium.Acombinationofthesolarwind
andEarth'sheatwouldhavedrivenoffthisatmosphere,asa
resultofwhichtheatmosphereisnowdepletedofthese
elementscomparedtocosmicabundances.[62]Aftertheimpact
whichcreatedthemoon,themoltenEarthreleasedvolatile
gasesandlatermoregaseswerereleasedbyvolcanoes, Graphshowingrangeofestimatedpartialpressure
completingasecondatmosphererichingreenhousegasesbut
ofatmosphericoxygenthroughgeologictime [61]
poorinoxygen.[1]:256Finally,thethirdatmosphere,richin
oxygen,emergedwhenbacteriabegantoproduceoxygen
about2.8Ga.[63]:8384,116117
Inearlymodelsfortheformationoftheatmosphereandocean,thesecondatmospherewasformedbyoutgassing
ofvolatilesfromtheEarth'sinterior.Nowitisconsideredlikelythatmanyofthevolatilesweredeliveredduring
accretionbyaprocessknownasimpactdegassinginwhichincomingbodiesvaporizeonimpact.Theoceanand
atmospherewouldthereforehavestartedtoformevenastheEarthformed.[64]Thenewatmosphereprobably
containedwatervapor,carbondioxide,nitrogen,andsmalleramountsofothergases.[65]
Planetesimalsatadistanceof1astronomicalunit(AU),thedistanceoftheEarthfromtheSun,probablydidnot
contributeanywatertotheEarthbecausethesolarnebulawastoohotforicetoformandthehydrationofrocksby
watervaporwouldhavetakentoolong.[64][66]Thewatermusthavebeensuppliedbymeteoritesfromtheouter
asteroidbeltandsomelargeplanetaryembryosfrombeyond2.5AU.[64][67]Cometsmayalsohavecontributed.
ThoughmostcometsaretodayinorbitsfartherawayfromtheSunthanNeptune,computersimulationsshowthat
theywereoriginallyfarmorecommonintheinnerpartsofthesolarsystem.[59]:130132
AstheEarthcooled,cloudsformed.Raincreatedtheoceans.Recentevidencesuggeststheoceansmayhavebegun
formingasearlyas4.4Ga.[34]BythestartoftheArcheaneontheyalreadycoveredtheEarth.Thisearlyformation
hasbeendifficulttoexplainbecauseofaproblemknownasthefaintyoungSunparadox.Starsareknowntoget
brighterastheyage,andatthetimeofitsformationtheSunwouldhavebeenemittingonly70%ofitscurrent
power.Thus,theSunhasbecome30%brighterinthelast4.5billionyears.[68]ManymodelsindicatethattheEarth
wouldhavebeencoveredinice.[69][64]Alikelysolutionisthattherewasenoughcarbondioxideandmethaneto
produceagreenhouseeffect.Thecarbondioxidewouldhavebeenproducedbyvolcanoesandthemethaneby
earlymicrobes.Anothergreenhousegas,ammonia,wouldhavebeenejectedbyvolcanosbutquicklydestroyedby
ultravioletradiation.[63]:83
Originoflife
Oneofthereasonsforinterestintheearlyatmosphere
andoceanisthattheyformtheconditionsunder Lifetimeline
whichlifefirstarose.Therearemanymodels,but viewdiscuss
littleconsensus,onhowlifeemergedfromnonliving Earliesthumans
0
chemicalschemicalsystemsthathavebeencreatedin Quaternary P Flowers
h Mammals
thelaboratorystillfallwellshortoftheminimum a Dinosaurs
Karoo
n Landlife
complexityforalivingorganism.[70][71] Andean e
500 r Cambrianexplosion
Thefirststepintheemergenceoflifemayhavebeen Cryogenian
o
z
Ediacarabiota

o
chemicalreactionsthatproducedmanyofthesimpler i Multicellular
c
organiccompounds,includingnucleobasesandamino 1000 life
acids,thatarethebuildingblocksoflife.An

Earliestsexual
experimentin1953byStanleyMillerandHarold reproduction
P
Ureyshowedthatsuchmoleculescouldforminan r
1500 o
atmosphereofwater,methane,ammoniaand t
hydrogenwiththeaidofsparkstomimictheeffectof e
r Eukaryotes
lightning.[72]Althoughtheatmosphericcomposition o
z
2000
wasprobablydifferentfromthecompositionusedby o
i
MillerandUrey,laterexperimentswithmorerealistic c
Huronian
Oxygencrisis
compositionsalsomanagedtosynthesizeorganic
molecules.[73]Recentcomputersimulationshaveeven 2500 Atmosphericoxygen
shownthatextraterrestrialorganicmoleculescould
haveformedintheprotoplanetarydiskbeforethe Pongola
photosynthesis
formationoftheEarth.[74] 3000 A
r
c
h
Thenextstageofcomplexitycouldhavebeenreached e
fromatleastthreepossiblestartingpoints:self 3500
a
Earliestoxygen
n
replication,anorganism'sabilitytoproduceoffspring
thatareverysimilartoitselfmetabolism,itsabilityto
Singlecelled
feedandrepairitselfandexternalcellmembranes, LHBmeteorites
4000 life
whichallowfoodtoenterandwasteproductstoleave, H
a
butexcludeunwantedsubstances.[75] d Earliestlife
e water
4500 a
Earliestwater
EarliestEarth
4500 a
Earliestwater
EarliestEarth
Replicationfirst:RNAworld n Axisscale:millionsofyears.
(4540)
Orangelabels:knowniceages.
Eventhesimplestmembersofthethreemodern Alsosee:HumantimelineandNaturetimeline
domainsoflifeuseDNAtorecordtheir"recipes"and
acomplexarrayofRNAandproteinmoleculesto"read"theseinstructionsandusethemforgrowth,maintenance
andselfreplication.
ThediscoverythatakindofRNAmoleculecalledaribozymecancatalyzebothitsownreplicationandthe
constructionofproteinsledtothehypothesisthatearlierlifeformswerebasedentirelyonRNA.[76]Theycould
haveformedanRNAworldinwhichtherewereindividualsbutnospecies,asmutationsandhorizontalgene
transferswouldhavemeantthattheoffspringineachgenerationwerequitelikelytohavedifferentgenomesfrom
thosethattheirparentsstartedwith.[77]RNAwouldlaterhavebeenreplacedbyDNA,whichismorestableand
thereforecanbuildlongergenomes,expandingtherangeofcapabilitiesasingleorganismcanhave.[78]Ribozymes
remainasthemaincomponentsofribosomes,the"proteinfactories"ofmoderncells.[79]
Althoughshort,selfreplicatingRNAmoleculeshavebeenartificiallyproducedinlaboratories,[80]doubtshave
beenraisedaboutwhethernaturalnonbiologicalsynthesisofRNAispossible.[81][82][83]Theearliestribozymes
mayhavebeenformedofsimplernucleicacidssuchasPNA,TNAorGNA,whichwouldhavebeenreplacedlater
byRNA.[84][85]OtherpreRNAreplicatorshavebeenposited,includingcrystals[86]:150andevenquantum
systems.[87]
In2003itwasproposedthatporousmetalsulfideprecipitateswouldassistRNAsynthesisatabout100C
(212F)andoceanbottompressuresnearhydrothermalvents.Inthishypothesis,lipidmembraneswouldbethe
lastmajorcellcomponentstoappearanduntiltheydidtheprotocellswouldbeconfinedtothepores.[88]
Metabolismfirst:ironsulfurworld
Anotherlongstandinghypothesisisthatthefirstlifewascomposedofproteinmolecules.Aminoacids,the
buildingblocksofproteins,areeasilysynthesizedinplausibleprebioticconditions,asaresmallpeptides(polymers
ofaminoacids)thatmakegoodcatalysts.[89]:295297Aseriesofexperimentsstartingin1997showedthatamino
acidsandpeptidescouldforminthepresenceofcarbonmonoxideandhydrogensulfidewithironsulfideand
nickelsulfideascatalysts.Mostofthestepsintheirassemblyrequiredtemperaturesofabout100C(212F)and
moderatepressures,althoughonestagerequired250C(482F)andapressureequivalenttothatfoundunder7
kilometers(4.3mi)ofrock.Hence,selfsustainingsynthesisofproteinscouldhaveoccurrednearhydrothermal
vents.[90]
Adifficultywiththemetabolismfirstscenarioisfindingawayfororganismstoevolve.Withouttheabilityto
replicateasindividuals,aggregatesofmoleculeswouldhave"compositionalgenomes"(countsofmolecular
speciesintheaggregate)asthetargetofnaturalselection.However,arecentmodelshowsthatsuchasystemis
unabletoevolveinresponsetonaturalselection.[91]
Membranesfirst:Lipidworld
Ithasbeensuggestedthatdoublewalled"bubbles"oflipidslikethosethatformtheexternalmembranesofcells
mayhavebeenanessentialfirststep.[92]ExperimentsthatsimulatedtheconditionsoftheearlyEarthhavereported
theformationoflipids,andthesecanspontaneouslyformliposomes,doublewalled"bubbles",andthenreproduce
themselves.Althoughtheyarenotintrinsicallyinformationcarriersasnucleic
acidsare,theywouldbesubjecttonaturalselectionforlongevityandreproduction.
NucleicacidssuchasRNAmightthenhaveformedmoreeasilywithinthe
liposomesthantheywouldhaveoutside.[93]
Theclaytheory
Someclays,notablymontmorillonite,havepropertiesthatmakethemplausible
acceleratorsfortheemergenceofanRNAworld:theygrowbyselfreplicationof
theircrystallinepattern,aresubjecttoananalogofnaturalselection(astheclay
"species"thatgrowsfastestinaparticularenvironmentrapidlybecomes
dominant),andcancatalyzetheformationofRNAmolecules.[94]Althoughthis
ideahasnotbecomethescientificconsensus,itstillhasactive
supporters.[95]:150158[86]
Researchin2003reportedthatmontmorillonitecouldalsoacceleratethe
conversionoffattyacidsinto"bubbles",andthatthebubblescouldencapsulate
RNAattachedtotheclay.Bubblescanthengrowbyabsorbingadditionallipids
anddividing.Theformationoftheearliestcellsmayhavebeenaidedbysimilar Thereplicatorinvirtuallyall
processes.[96] knownlifeis
deoxyribonucleicacid.DNA
Asimilarhypothesispresentsselfreplicatingironrichclaysastheprogenitorsof isfarmorecomplexthanthe
nucleotides,lipidsandaminoacids.[97] originalreplicatorandits
replicationsystemsare
Lastuniversalancestor highlyelaborate.
Itisbelievedthatofthismultiplicityofprotocells,onlyonelinesurvived.Current
phylogeneticevidencesuggeststhatthelastuniversalancestor(LUA)livedduring
theearlyArcheaneon,perhaps3.5Gaorearlier.[98][99]ThisLUAcellisthe
ancestorofalllifeonEarthtoday.Itwasprobablyaprokaryote,possessingacell
membraneandprobablyribosomes,butlackinganucleusormembranebound
organellessuchasmitochondriaorchloroplasts.Likeallmoderncells,itused
DNAasitsgeneticcode,RNAforinformationtransferandproteinsynthesis,and
enzymestocatalyzereactions.Somescientistsbelievethatinsteadofasingle
organismbeingthelastuniversalcommonancestor,therewerepopulationsof
organismsexchanginggenesbylateralgenetransfer.[98]
ProterozoicEon Crosssectionthrougha
liposome
TheProterozoiceonlastedfrom2.5Gato542Ma(millionyears)ago.[2]:130Inthis
timespan,cratonsgrewintocontinentswithmodernsizes.Thechangetoan
oxygenrichatmospherewasacrucialdevelopment.Lifedevelopedfromprokaryotesintoeukaryotesand
multicellularforms.TheProterozoicsawacoupleofsevereiceagescalledsnowballEarths.Afterthelast
SnowballEarthabout600Ma,theevolutionoflifeonEarthaccelerated.About580Ma,theEdiacarabiotaformed
thepreludefortheCambrianExplosion.
Oxygenrevolution
Theearliestcellsabsorbedenergyandfoodfromthesurrounding
environment.Theyusedfermentation,thebreakdownofmorecomplex
compoundsintolesscomplexcompoundswithlessenergy,andusedthe
energysoliberatedtogrowandreproduce.Fermentationcanonlyoccurin
ananaerobic(oxygenfree)environment.Theevolutionofphotosynthesis
madeitpossibleforcellstomanufacturetheirownfood.[100]:377
MostofthelifethatcoversthesurfaceoftheEarthdependsdirectlyor
indirectlyonphotosynthesis.Themostcommonform,oxygenic
photosynthesis,turnscarbondioxide,waterandsunlightintofood.It Lithifiedstromatolitesontheshores
capturestheenergyofsunlightinenergyrichmoleculessuchasATP, ofLakeThetis,WesternAustralia.
whichthenprovidetheenergytomakesugars.Tosupplytheelectronsin Archeanstromatolitesarethefirst
thecircuit,hydrogenisstrippedfromwater,leavingoxygenasawaste directfossiltracesoflifeonEarth.
product.[101]Someorganisms,includingpurplebacteriaandgreensulfur
bacteria,useananoxygenicformofphotosynthesisthatusealternativesto
hydrogenstrippedfromwateraselectrondonorsexamplesarehydrogen
sulfide,sulfurandiron.Suchextremophileorganismsarerestrictedto
otherwiseinhospitableenvironmentssuchashotspringsandhydrothermal
vents.[100]:379382[102]
Thesimpleranoxygenicformaroseabout3.8Ga,notlongafterthe
appearanceoflife.Thetimingofoxygenicphotosynthesisismore
controversialithadcertainlyappearedbyabout2.4Ga,butsome Abandedironformationfromthe
researchersputitbackasfaras3.2Ga.[101]Thelatter"probablyincreased 3.15GaMooriesGroup,Barberton
GreenstoneBelt,SouthAfrica.Red
globalproductivitybyatleasttwoorthreeordersofmagnitude".[103][104]
layersrepresentthetimeswhen
Amongtheoldestremnantsofoxygenproducinglifeformsarefossil
oxygenwasavailable,graylayers
stromatolites.[103][104][61] wereformedinanoxiccircumstances.
Atfirst,thereleasedoxygenwasboundupwithlimestone,iron,andother
minerals.Theoxidizedironappearsasredlayersingeologicalstratacalledbandedironformationsthatformedin
abundanceduringtheSiderianperiod(between2500Maand2300Ma).[2]:133Whenmostoftheexposedreadily
reactingmineralswereoxidized,oxygenfinallybegantoaccumulateintheatmosphere.Thougheachcellonly
producedaminuteamountofoxygen,thecombinedmetabolismofmanycellsoveravasttimetransformedEarth's
atmospheretoitscurrentstate.ThiswasEarth'sthirdatmosphere.[105]:5051[63]:8384,116117
Someoxygenwasstimulatedbyincomingultravioletradiationtoformozone,whichcollectedinalayernearthe
upperpartoftheatmosphere.Theozonelayerabsorbed,andstillabsorbs,asignificantamountoftheultraviolet
radiationthatoncehadpassedthroughtheatmosphere.Itallowedcellstocolonizethesurfaceoftheoceanand
eventuallytheland:withouttheozonelayer,ultravioletradiationbombardinglandandseawouldhavecaused
unsustainablelevelsofmutationinexposedcells.[106][59]:219220
Photosynthesishadanothermajorimpact.OxygenwastoxicmuchlifeonEarthprobablydiedoutasitslevels
roseinwhatisknownastheoxygencatastrophe.Resistantformssurvivedandthrived,andsomedevelopedthe
abilitytouseoxygentoincreasetheirmetabolismandobtainmoreenergyfromthesamefood.[106]
SnowballEarth
ThenaturalevolutionoftheSunmadeitprogressivelymoreluminousduringtheArcheanandProterozoiceons
theSun'sluminosityincreases6%everybillionyears.[59]:165Asaresult,theEarthbegantoreceivemoreheatfrom
theSunintheProterozoiceon.However,theEarthdidnotgetwarmer.Instead,thegeologicalrecordseemsto
suggestitcooleddramaticallyduringtheearlyProterozoic.GlacialdepositsfoundinSouthAfricadatebackto
2.2Ga,atwhichtime,basedonpaleomagneticevidence,theymusthavebeenlocatedneartheequator.Thus,this
glaciation,knownastheMakganyeneglaciation,mayhavebeenglobal.Somescientistssuggestthiswassosevere
thattheEarthwastotallyfrozenoverfromthepolestotheequator,ahypothesiscalledSnowballEarth.[107]
Theiceagearound2.3Gacouldhavebeendirectlycausedbytheincreasedoxygenconcentrationinthe
atmosphere,whichcausedthedecreaseofmethane(CH4)intheatmosphere.Methaneisastronggreenhousegas,
butwithoxygenitreactstoformCO2,alesseffectivegreenhousegas.[59]:172Whenfreeoxygenbecameavailable
intheatmosphere,theconcentrationofmethanecouldhavedecreaseddramatically,enoughtocountertheeffectof
theincreasingheatflowfromtheSun.[108]
However,thetermSnowballEarthismorecommonlyusedtodescribelaterextremeiceagesduringthe
Cryogenianperiod.Therewerefourperiods,eachlastingabout10millionyears,between750and580million
yearsago,whentheearthisthoughttohavebeencoveredwithiceapartfromthehighestmountains,andaverage
temperatureswereabout50C(58F).[109]Thesnowballmayhavebeenpartlyduetothelocationofthe
supercontintentRodiniastraddlingtheEquator.Carbondioxidecombineswithraintoweatherrockstoform
carbonicacid,whichisthenwashedouttosea,thusextractingthegreenhousegasfromtheatmosphere.Whenthe
continentsarenearthepoles,theadvanceoficecoverstherocks,slowingthereductionincarbondioxide,butin
theCryogieniantheweatheringofRodiniawasabletocontinueuncheckeduntiltheiceadvancedtothetropics.
Theprocessmayhavefinallybeenreversedbytheemissionofcarbondioxidefromvolcanoesorthe
destabilizationofmethanegashydrates.AccordingtothealternativeSlushballEarththeory,evenattheheightof
theiceagestherewasstillopenwaterattheEquator.[110][111]
Emergenceofeukaryotes
Moderntaxonomyclassifieslifeintothreedomains.Thetimeoftheorigin
ofthesedomainsisuncertain.TheBacteriadomainprobablyfirstsplitoff
fromtheotherformsoflife(sometimescalledNeomura),butthis
suppositioniscontroversial.Soonafterthis,by2Ga,[112]theNeomurasplit
intotheArchaeaandtheEukarya.Eukaryoticcells(Eukarya)arelargerand
morecomplexthanprokaryoticcells(BacteriaandArchaea),andtheorigin
ofthatcomplexityisonlynowbecomingknown.
Aroundthistime,thefirstprotomitochondrionwasformed.Abacterial
cellrelatedtotoday'sRickettsia,[113]whichhadevolvedtometabolize Chloroplastsinthecellsofamoss
oxygen,enteredalargerprokaryoticcell,whichlackedthatcapability.
Perhapsthelargecellattemptedtodigestthesmalleronebutfailed
(possiblyduetotheevolutionofpreydefenses).Thesmallercellmayhavetriedtoparasitizethelargerone.Inany
case,thesmallercellsurvivedinsidethelargercell.Usingoxygen,itmetabolizedthelargercell'swasteproducts
andderivedmoreenergy.Partofthisexcessenergywasreturnedtothehost.Thesmallercellreplicatedinsidethe
largerone.Soon,astablesymbiosisdevelopedbetweenthelargecellandthesmallercellsinsideit.Overtime,the
hostcellacquiredsomegenesfromthesmallercells,andthetwokindsbecamedependentoneachother:thelarger
cellcouldnotsurvivewithouttheenergyproducedbythesmallerones,andtheseinturncouldnotsurvivewithout
therawmaterialsprovidedbythelargercell.Thewholecellisnowconsideredasingleorganism,andthesmaller
cellsareclassifiedasorganellescalledmitochondria.[114]
Asimilareventoccurredwithphotosyntheticcyanobacteria[115]enteringlargeheterotrophiccellsandbecoming
chloroplasts.[105]:6061[116]:536539Probablyasaresultofthesechanges,alineofcellscapableofphotosynthesis
splitofffromtheothereukaryotesmorethan1billionyearsago.Therewereprobablyseveralsuchinclusion
events.Besidesthewellestablishedendosymbiotictheoryofthecellularoriginofmitochondriaandchloroplasts,
therearetheoriesthatcellsledtoperoxisomes,spirochetesledtociliaandflagella,andthatperhapsaDNAvirus
ledtothecellnucleus,[117][118]thoughnoneofthemarewidelyaccepted.[119]
Archaeans,bacteria,andeukaryotescontinuedtodiversifyandtobecomemorecomplexandbetteradaptedto
theirenvironments.Eachdomainrepeatedlysplitintomultiplelineages,althoughlittleisknownaboutthehistory
ofthearchaeaandbacteria.Around1.1Ga,thesupercontinentRodiniawasassembling.[120][121]Theplant,animal,
andfungilineshadsplit,thoughtheystillexistedassolitarycells.Someoftheselivedincolonies,andgraduallya
divisionoflaborbegantotakeplaceforinstance,cellsontheperipherymighthavestartedtoassumedifferent
rolesfromthoseintheinterior.Althoughthedivisionbetweenacolonywithspecializedcellsandamulticellular
organismisnotalwaysclear,around1billionyearsago[122]thefirstmulticellularplantsemerged,probablygreen
algae.[123]Possiblybyaround900Ma[116]:488truemulticellularityhadalsoevolvedinanimals.
Atfirstitprobablyresembledtoday'ssponges,whichhavetotipotentcellsthatallowadisruptedorganismto
reassembleitself.[116]:483487Asthedivisionoflaborwascompletedinalllinesofmulticellularorganisms,cells
becamemorespecializedandmoredependentoneachotherisolatedcellswoulddie.
SupercontinentsintheProterozoic
Reconstructionsoftectonicplatemovementinthepast250millionyears
(theCenozoicandMesozoiceras)canbemadereliablyusingfittingof
continentalmargins,oceanfloormagneticanomaliesandpaleomagnetic
poles.Nooceancrustdatesbackfurtherthanthat,soearlierreconstructions
aremoredifficult.Paleomagneticpolesaresupplementedbygeologic
evidencesuchasorogenicbelts,whichmarktheedgesofancientplates,
andpastdistributionsoffloraandfauna.Thefurtherbackintime,the
scarcerandhardertointerpretthedatagetandthemorediversethe
reconstructions.[124]:370
ThroughoutthehistoryoftheEarth,therehavebeentimeswhencontinents
collidedandformedasupercontinent,whichlaterbrokeupintonew
continents.About1000to830Ma,mostcontinentalmasswasunitedinthe
AreconstructionofPannotia
supercontinentRodinia.[124]:370[125]Rodiniamayhavebeenprecededby (550Ma).
EarlyMiddleProterozoiccontinentscalledNunaand
Columbia.[124]:374[126][127]
AfterthebreakupofRodiniaabout800Ma,thecontinentsmayhaveformedanothershortlivedsupercontinent
around550Ma.ThehypotheticalsupercontinentissometimesreferredtoasPannotiaorVendia.[128]:321322The
evidenceforitisaphaseofcontinentalcollisionknownasthePanAfricanorogeny,whichjoinedthecontinental
massesofcurrentdayAfrica,SouthAmerica,AntarcticaandAustralia.TheexistenceofPannotiadependsonthe
timingoftheriftingbetweenGondwana(whichincludedmostofthelandmassnowintheSouthernHemisphere,
aswellastheArabianPeninsulaandtheIndiansubcontinent)andLaurentia(roughlyequivalenttocurrentday
NorthAmerica).[124]:374ItisatleastcertainthatbytheendoftheProterozoiceon,mostofthecontinentalmasslay
unitedinapositionaroundthesouthpole.[129]
LateProterozoicclimateandlife
TheendoftheProterozoicsawatleasttwoSnowballEarths,soseverethat
thesurfaceoftheoceansmayhavebeencompletelyfrozen.Thishappened
about716.5and635Ma,intheCryogenianperiod.[130]Theintensityand
mechanismofbothglaciationsarestillunderinvestigationandharderto
explainthantheearlyProterozoicSnowballEarth.[131]Most
paleoclimatologiststhinkthecoldepisodeswerelinkedtotheformationof
thesupercontinentRodinia.[132]BecauseRodiniawascenteredonthe
equator,ratesofchemicalweatheringincreasedandcarbondioxide(CO2)
wastakenfromtheatmosphere.BecauseCO2isanimportantgreenhouse
gas,climatescooledglobally.Inthesameway,duringtheSnowballEarths
mostofthecontinentalsurfacewascoveredwithpermafrost,which A580millionyearoldfossilof
decreasedchemicalweatheringagain,leadingtotheendoftheglaciations. Sprigginafloundensi,ananimalfrom
Analternativehypothesisisthatenoughcarbondioxideescapedthrough theEdiacaranperiod.Suchlifeforms
volcanicoutgassingthattheresultinggreenhouseeffectraisedglobal couldhavebeenancestorstothe
manynewformsthatoriginatedin
temperatures.[132]Increasedvolcanicactivityresultedfromthebreakupof
theCambrianExplosion.
Rodiniaataboutthesametime.
TheCryogenianperiodwasfollowedbytheEdiacaranperiod,whichwascharacterizedbyarapiddevelopmentof
newmulticellularlifeforms.[133]Whetherthereisaconnectionbetweentheendofthesevereiceagesandthe
increaseindiversityoflifeisnotclear,butitdoesnotseemcoincidental.Thenewformsoflife,calledEdiacara
biota,werelargerandmorediversethanever.ThoughthetaxonomyofmostEdiacaranlifeformsisunclear,some
wereancestorsofgroupsofmodernlife.[134]Importantdevelopmentsweretheoriginofmuscularandneuralcells.
NoneoftheEdiacaranfossilshadhardbodypartslikeskeletons.Thesefirstappearaftertheboundarybetweenthe
ProterozoicandPhanerozoiceonsorEdiacaranandCambrianperiods.
PhanerozoicEon
ThePhanerozoicisthecurrenteononEarth,whichstartedapproximately542millionyearsago.Itconsistsof
threeeras:ThePaleozoic,Mesozoic,andCenozoic,[22]andisthetimewhenmulticellularlifegreatlydiversified
intoalmostalltheorganismsknowntoday.[135]
ThePaleozoic("oldlife")erawasthefirstandlongesteraofthePhanerozoiceon,lastingfrom542to251Ma.[22]
DuringthePaleozoic,manymoderngroupsoflifecameintoexistence.Lifecolonizedtheland,firstplants,then
animals.Twomajorextinctionsoccurred.ThecontinentsformedatthebreakupofPannotiaandRodiniaatthe
endoftheProterozoicslowlymovedtogetheragain,formingthesupercontinentPangaeainthelatePaleozoic.
TheMesozoic("middlelife")eralastedfrom251Mato66Ma.[22]ItissubdividedintotheTriassic,Jurassic,and
Cretaceousperiods.TheerabeganwiththePermianTriassicextinctionevent,themostsevereextinctioneventin
thefossilrecord95%ofthespeciesonEarthdiedout.[136]ItendedwiththeCretaceousPaleogeneextinction
eventthatwipedoutthedinosaurs..
TheCenozoic("newlife")erabeganat66Ma,[22]andissubdividedintothePaleogene,Neogene,andQuaternary
periods.Thesethreeperiodsarefurthersplitintosevensubdivisions,withthePaleogenecomposedofThe
Paleocene,Eocene,andOligocene,theNeocenedividedintotheMiocene,Pliocene,andtheQuaternarycomposed
ofthePleistocene,andHolocene.[137]Mammals,birds,amphibians,crocodilians,turtlesandlepidosaurswereable
tosurvivetheCretaceousPaleogeneextinctioneventthatkilledoffthenonaviandinosaursandmanyotherforms
oflife,andthisistheeraduringwhichtheydiversifiedintotheirmodernforms.
Tectonics,paleogeographyandclimate
AttheendoftheProterozoic,thesupercontinentPannotiahadbrokenapart
intothesmallercontinentsLaurentia,Baltica,SiberiaandGondwana.[138]
Duringperiodswhencontinentsmoveapart,moreoceaniccrustisformed
byvolcanicactivity.Becauseyoungvolcaniccrustisrelativelyhotterand
lessdensethanoldoceaniccrust,theoceanfloorsriseduringsuchperiods.
Thiscausesthesealeveltorise.Therefore,inthefirsthalfofthePaleozoic,
largeareasofthecontinentswerebelowsealevel.
EarlyPaleozoicclimateswerewarmerthantoday,buttheendofthe
Ordoviciansawashorticeageduringwhichglacierscoveredthesouth
pole,wherethehugecontinentGondwanawassituated.Tracesof
glaciationfromthisperiodareonlyfoundonformerGondwana.Duringthe
LateOrdovicianiceage,afewmassextinctionstookplace,inwhichmany
brachiopods,trilobites,Bryozoaandcoralsdisappeared.Thesemarine
speciescouldprobablynotcontendwiththedecreasingtemperatureofthe Pangaeawasasupercontinentthat
seawater.[139]Aftertheextinctionsnewspeciesevolved,morediverseand existedfromabout300to180Ma.
betteradapted.Theywouldfillthenichesleftbytheextinctspecies. Theoutlinesofthemoderncontinents
andotherlandmassesareindicatedon
ThecontinentsLaurentiaandBalticacollidedbetween450and400Ma, thismap.
duringtheCaledonianOrogeny,toformLaurussia(alsoknownas
Euramerica).[140]TracesofthemountainbeltthiscollisioncausedcanbefoundinScandinavia,Scotland,andthe
northernAppalachians.IntheDevonianperiod(416359Ma)[22]GondwanaandSiberiabegantomovetowards
Laurussia.ThecollisionofSiberiawithLaurussiacausedtheUralianOrogeny,thecollisionofGondwanawith
LaurussiaiscalledtheVariscanorHercynianOrogenyinEuropeortheAlleghenianOrogenyinNorthAmerica.
ThelatterphasetookplaceduringtheCarboniferousperiod(359299Ma)[22]andresultedintheformationofthe
lastsupercontinent,Pangaea.[60]
By180Ma,PangaeabrokeupintoLaurasiaandGondwana.
Cambrianexplosion
Therateoftheevolutionoflifeasrecordedbyfossilsacceleratedinthe
Cambrianperiod(542488Ma).[22]Thesuddenemergenceofmanynew
species,phyla,andformsinthisperiodiscalledtheCambrianExplosion.
ThebiologicalfomentingintheCambrianExplosionwasunpreceded
beforeandsincethattime.[59]:229WhereastheEdiacaranlifeformsappear
yetprimitiveandnoteasytoputinanymoderngroup,attheendofthe
Cambrianmostmodernphylawerealreadypresent.Thedevelopmentof
hardbodypartssuchasshells,skeletonsorexoskeletonsinanimalslike
molluscs,echinoderms,crinoidsandarthropods(awellknowngroupof
arthropodsfromthelowerPaleozoicarethetrilobites)madethe
preservationandfossilizationofsuchlifeformseasierthanthoseoftheir Trilobitesfirstappearedduringthe
Proterozoicancestors.Forthisreason,muchmoreisknownaboutlifein Cambrianperiodandwereamongthe
andaftertheCambrianthanaboutthatofolderperiods.Someofthese mostwidespreadanddiversegroups
Cambriangroupsappearcomplexbutarequitedifferentfrommodernlife ofPaleozoicorganisms.
examplesareAnomalocarisandHaikouichthys.
DuringtheCambrian,thefirstvertebrateanimals,amongthemthefirstfishes,hadappeared.[116]:357Acreature
thatcouldhavebeentheancestorofthefishes,orwasprobablycloselyrelatedtoit,wasPikaia.Ithadaprimitive
notochord,astructurethatcouldhavedevelopedintoavertebralcolumnlater.Thefirstfisheswithjaws
(Gnathostomata)appearedduringthenextgeologicalperiod,theOrdovician.Thecolonisationofnewniches
resultedinmassivebodysizes.Inthisway,fisheswithincreasingsizesevolvedduringtheearlyPaleozoic,suchas
thetitanicplacodermDunkleosteus,whichcouldgrow7meters(23ft)long.
Thediversityoflifeformsdidnotincreasegreatlybecauseofaseriesofmassextinctionsthatdefinewidespread
biostratigraphicunitscalledbiomeres.[141]Aftereachextinctionpulse,thecontinentalshelfregionswere
repopulatedbysimilarlifeformsthatmayhavebeenevolvingslowlyelsewhere.[142]BythelateCambrian,the
trilobiteshadreachedtheirgreatestdiversityanddominatednearlyallfossilassemblages.[143]:34
Colonizationofland
Oxygenaccumulationfromphotosynthesisresultedintheformationofan
ozonelayerthatabsorbedmuchoftheSun'sultravioletradiation,meaning
unicellularorganismsthatreachedlandwerelesslikelytodie,and
prokaryotesbegantomultiplyandbecomebetteradaptedtosurvivaloutof
thewater.Prokaryotelineages[144]hadprobablycolonizedthelandasearly
as2.6Ga[145]evenbeforetheoriginoftheeukaryotes.Foralongtime,the
landremainedbarrenofmulticellularorganisms.Thesupercontinent
Pannotiaformedaround600Maandthenbrokeapartashort50million
yearslater.[146]Fish,theearliestvertebrates,evolvedintheoceansaround
Artist'sconceptionofDevonianflora
530Ma.[116]:354Amajorextinctioneventoccurredneartheendofthe
Cambrianperiod,[147]whichended488Ma.[148]
Severalhundredmillionyearsago,plants(probablyresemblingalgae)andfungistartedgrowingattheedgesof
thewater,andthenoutofit.[149]:138140Theoldestfossilsoflandfungiandplantsdateto480460Ma,though
molecularevidencesuggeststhefungimayhavecolonizedthelandasearlyas1000Maandtheplants
700Ma.[150]Initiallyremainingclosetothewater'sedge,mutationsandvariationsresultedinfurthercolonization
ofthisnewenvironment.Thetimingofthefirstanimalstoleavetheoceansisnotpreciselyknown:theoldestclear
evidenceisofarthropodsonlandaround450Ma,[151]perhapsthrivingandbecomingbetteradaptedduetothevast
foodsourceprovidedbytheterrestrialplants.Thereisalsounconfirmedevidencethatarthropodsmayhave
appearedonlandasearlyas530Ma.[152]
Evolutionoftetrapods
AttheendoftheOrdovicianperiod,443Ma,[22]additionalextinction
eventsoccurred,perhapsduetoaconcurrenticeage.[139]Around380to
375Ma,thefirsttetrapodsevolvedfromfish.[153]Finsevolvedtobecome
limbsthatthefirsttetrapodsusedtolifttheirheadsoutofthewaterto
breatheair.Thiswouldletthemliveinoxygenpoorwater,orpursuesmall Tiktaalik,afishwithlimblikefins
andapredecessoroftetrapods.
preyinshallowwater.[153]Theymayhavelaterventuredonlandforbrief
Reconstructionfromfossilsabout
periods.Eventually,someofthembecamesowelladaptedtoterrestriallife
375millionyearsold.
thattheyspenttheiradultlivesonland,althoughtheyhatchedinthewater
andreturnedtolaytheireggs.Thiswastheoriginoftheamphibians.About365Ma,anotherperiodofextinction
occurred,perhapsasaresultofglobalcooling.[154]Plantsevolvedseeds,whichdramaticallyacceleratedtheir
spreadonland,aroundthistime(byapproximately360Ma).[155][156]
About20millionyearslater(340Ma[116]:293296),theamnioticeggevolved,whichcouldbelaidonland,givinga
survivaladvantagetotetrapodembryos.Thisresultedinthedivergenceofamniotesfromamphibians.Another
30millionyears(310Ma[116]:254256)sawthedivergenceofthesynapsids(includingmammals)fromthe
sauropsids(includingbirdsandreptiles).Othergroupsoforganismscontinuedtoevolve,andlinesdivergedin
fish,insects,bacteria,andsoonbutlessisknownofthedetails.
Afteryetanother,themostsevereextinctionoftheperiod(251~250Ma),
around230Ma,dinosaurssplitofffromtheirreptilianancestors.[157]The
TriassicJurassicextinctioneventat200Masparedmanyofthe
dinosaurs,[22][158]andtheysoonbecamedominantamongthevertebrates.
Thoughsomemammalianlinesbegantoseparateduringthisperiod,
existingmammalswereprobablysmallanimalsresemblingshrews.[116]:169
Theboundarybetweenavianandnonaviandinosaursisnotclear,but
Archaeopteryx,traditionallyconsideredoneofthefirstbirds,livedaround
150Ma.[159] Dinosaurswerethedominant
terrestrialvertebratesthroughoutmost
Theearliestevidencefortheangiospermsevolvingflowersisduringthe oftheMesozoic
Cretaceousperiod,some20millionyearslater(132Ma).[160]
Extinctions
ThefirstmassextinctionwastheOrdovicianSilurianextinction.Apossiblecausewastheintenseglaciationof
Gondwana,whicheventuallyledtoasnowballearth.60%ofmarineinvertebratesbecameextinctand25%ofall
families.
ThesecondmassextinctionwastheLateDevonianextinction,probablycausedbytheevolutionoftrees,which
couldleadtothedepletionofgreenhousegases(likeCO2)orleadtoeutrophicationofthewater.70%ofall
speciesbecameextinct.
ThethirdmassextinctionwasthePermianTriassic,ortheGreatDying,eventwaspossiblycausedbysome
combinationoftheSiberianTrapsvolcanicevent,anasteroidimpact,methanehydrategasification,sealevel
fluctuations,andamajoranoxicevent.EithertheproposedWilkesLandcrater[161]inAntarcticaorBedout
structureoffthenorthwestcoastofAustraliamayindicateanimpactconnectionwiththePermianTriassic
extinction.ButitremainsuncertainwhethereithertheseorotherproposedPermianTriassicboundarycratersare
eitherrealimpactcratersorevencontemporaneouswiththePermianTriassicextinctionevent.Thiswasbyfarthe
deadliestextinctionever,withabout57%ofallfamiliesand83%ofallgenerakilled.[162][163]
ThefourthmassextinctionwastheTriassicJurassicextinctioneventinwhichalmostallsynapsidsandarchosaurs
becameextinct,probablyduetonewcompetitionfromdinosaurs.
ThefifthandmostrecentmassextinctionwastheKTextinction.In66Ma,a10kilometer(6.2mi)asteroidstruck
EarthjustofftheYucatnPeninsulasomewhereinthesouthwesterntipofthenLaurasiawheretheChicxulub
crateristoday.Thisejectedvastquantitiesofparticulatematterandvaporintotheairthatoccludedsunlight,
inhibitingphotosynthesis.75%ofalllife,includingthenonaviandinosaurs,becameextinct,[164]markingtheend
oftheCretaceousperiodandMesozoicera.
Diversificationofmammals
Thefirsttruemammalsevolvedintheshadowsofdinosaursandotherlargearchosaursthatfilledtheworldbythe
lateTriassic.Thefirstmammalswereverysmall,andwereprobablynocturnaltoescapepredation.Mammal
diversificationtrulybeganonlyaftertheCretaceousPaleogeneextinctionevent.[165]BytheearlyPaleocenethe
earthrecoveredfromtheextinction,andmammaliandiversityincreased.CreatureslikeAmbulocetustooktothe
oceanstoeventuallyevolveintowhales,[166]whereassomecreatures,likeprimates,tooktothetrees.[167]Thisall
changedduringthemidtolateEocenewhenthecircumAntarcticcurrentformedbetweenAntarcticaandAustralia
whichdisruptedweatherpatternsonaglobalscale.Grasslesssavannasbegantopredominatemuchofthe
landscape,andmammalssuchasAndrewsarchusroseuptobecomethelargestknownterrestrialpredatory
mammalever,[168]andearlywhaleslikeBasilosaurustookcontroloftheseas.
TheevolutionofgrassbroughtaremarkablechangetotheEarth'slandscape,andthenewopenspacescreated
pushedmammalstogetbiggerandbigger.GrassstartedtoexpandintheMiocene,andtheMioceneiswheremany
moderndaymammalsfirstappeared.GiantungulateslikeParaceratheriumandDeinotheriumevolvedtorulethe
grasslands.Theevolutionofgrassalsobroughtprimatesdownfromthetrees,andstartedhumanevolution.The
firstbigcatsevolvedduringthistimeaswell.[169]TheTethysSeawasclosedoffbythecollisionofAfricaand
Europe.[170]
TheformationofPanamawasperhapsthemostimportantgeologicaleventtooccurinthelast60millionyears.
AtlanticandPacificcurrentswereclosedofffromeachother,whichcausedtheformationoftheGulfStream,
whichmadeEuropewarmer.ThelandbridgeallowedtheisolatedcreaturesofSouthAmericatomigrateoverto
NorthAmerica,andviceversa.[171]Variousspeciesmigratedsouth,leadingtothepresenceinSouthAmericaof
llamas,thespectacledbear,kinkajousandjaguars.
ThreemillionyearsagosawthestartofthePleistoceneepoch,whichfeatureddramaticclimacticchangesdueto
theiceages.TheiceagesledtotheevolutionofmodernmaninSaharanAfricaandexpansion.Themegafauna
thatdominatedfedongrasslandsthat,bynow,hadtakenovermuchofthesubtropicalworld.Thelargeamountsof
waterheldintheiceallowedforvariousbodiesofwatertoshrinkandsometimesdisappearsuchastheNorthSea
andtheBeringStrait.ItisbelievedbymanythatahugemigrationtookplacealongBeringiawhichiswhy,today,
therearecamels(whichevolvedandbecameextinctinNorthAmerica),horses(whichevolvedandbecameextinct
inNorthAmerica),andNativeAmericans.Theendingofthelasticeagecoincidedwiththeexpansionofman,
alongwithamassivedieoutoficeagemegafauna.Thisextinction,nicknamed"theSixthExtinction",hasbeen
goingeversince.
Humanevolution
AsmallAfricanapelivingaround6Mawasthelast
animalwhosedescendantswouldincludebothmodern Humantimeline
humansandtheirclosestrelatives,the viewdiscuss
chimpanzees.[116]:100101Onlytwobranchesofits 0 Modernhumans
P Homosapiens
familytreehavesurvivingdescendants.Verysoon
l Neanderthal Earliestclothes
afterthesplit,forreasonsthatarestillunclear,apesin e Earliestcooking
onebranchdevelopedtheabilitytowalk 1 i
upright.[116]:9599Brainsizeincreasedrapidly,andby s Homoerectus
t Earliestfireuse
2Ma,thefirstanimalsclassifiedinthegenusHomo o Earliestexit
hadappeared.[149]:300Ofcourse,thelinebetween 2 c fromAfrica
e Homohabilis
differentspeciesorevengeneraissomewhatarbitrary n
asorganismscontinuouslychangeovergenerations. e
3
https://en.wikipedia.org/wiki/History_of_Earth Earlieststonetools18/32
3
Aroundthesametime,theotherbranchsplitintothe Earlieststonetools
Australopithecus
ancestorsofthecommonchimpanzeeandthe P
ancestorsofthebonoboasevolutioncontinued 4 l Earliestbipedal
i
simultaneouslyinalllifeforms.[116]:100101 Ardipithecus H
o
c
TheabilitytocontrolfireprobablybeganinHomo 5 e o
n Humanlike
erectus(orHomoergaster),probablyatleast m
e apes
790,000yearsago[172]butperhapsasearlyas
Orrorin i
1.5Ma.[116]:67Theuseanddiscoveryofcontrolled 6

firemayevenpredateHomoerectus.Firewas
n

possiblyusedbytheearlyLowerPaleolithic i Possiblybipedal

7 M Sahelanthropus
(Oldowan)hominidHomohabilisorstrong
i d
australopithecinessuchasParanthropus.[173] o
c s
Itismore 8 e
n
difficultto e
establishthe
9 Ouranopithecus
originof
languageitis
unclear Nakalipithecus
whetherHomo 10 Earlierapes
Axisscale:millionsofyears.
erectuscould
Alsosee:LifetimelineandNaturetimeline
speakorifthat
capabilityhad
notbegununtilHomosapiens.[116]:67Asbrainsizeincreased,babieswere
bornearlier,beforetheirheadsgrewtoolargetopassthroughthepelvis.As
aresult,theyexhibitedmoreplasticity,andthuspossessedanincreased
capacitytolearnandrequiredalongerperiodofdependence.Socialskills
Areconstructionofhumanhistory
becamemorecomplex,languagebecamemoresophisticated,andtools
basedonfossildata. [174] becamemoreelaborate.Thiscontributedtofurthercooperationand
intellectualdevelopment.[175]:7Modernhumans(Homosapiens)are
believedtohaveoriginatedaround200,000yearsagoorearlierinAfricatheoldestfossilsdatebacktoaround
160,000yearsago.[176]
ThefirsthumanstoshowsignsofspiritualityaretheNeanderthals(usuallyclassifiedasaseparatespecieswithno
survivingdescendants)theyburiedtheirdead,oftenwithnosignoffoodortools.[177]:17However,evidenceof
moresophisticatedbeliefs,suchastheearlyCroMagnoncavepaintings(probablywithmagicalorreligious
significance)[177]:1719didnotappearuntil32,000yearsago.[178]CroMagnonsalsoleftbehindstonefigurines
suchasVenusofWillendorf,probablyalsosignifyingreligiousbelief.[177]:1719By11,000yearsago,Homo
sapienshadreachedthesoutherntipofSouthAmerica,thelastoftheuninhabitedcontinents(exceptfor
Antarctica,whichremainedundiscovereduntil1820AD).[179]Tooluseandcommunicationcontinuedtoimprove,
andinterpersonalrelationshipsbecamemoreintricate.
Civilization
Throughoutmorethan90%ofitshistory,Homosapienslivedinsmallbandsasnomadichuntergatherers.[175]:8
Aslanguagebecamemorecomplex,theabilitytorememberandcommunicateinformationresulted,accordingtoa
theoryproposedbyRichardDawkins,inanewreplicator:thememe.[180]Ideascouldbeexchangedquicklyand
passeddownthegenerations.Culturalevolutionquicklyoutpacedbiologicalevolution,andhistoryproperbegan.
Between8500and7000BC,humansintheFertileCrescentintheMiddle
Eastbeganthesystematichusbandryofplantsandanimals:agriculture.[181]
Thisspreadtoneighboringregions,anddevelopedindependently
elsewhere,untilmostHomosapienslivedsedentarylivesinpermanent
settlementsasfarmers.Notallsocietiesabandonednomadism,especially
thoseinisolatedareasoftheglobepoorindomesticableplantspecies,such
asAustralia.[182]However,amongthosecivilizationsthatdidadopt
agriculture,therelativestabilityandincreasedproductivityprovidedby
farmingallowedthepopulationtoexpand.
Agriculturehadamajorimpacthumansbegantoaffecttheenvironmentas
neverbefore.Surplusfoodallowedapriestlyorgoverningclasstoarise,
followedbyincreasingdivisionoflabor.ThisledtoEarth'sfirst
civilizationatSumerintheMiddleEast,between4000and3000
BC.[175]:15AdditionalcivilizationsquicklyaroseinancientEgypt,atthe
IndusRivervalleyandinChina.Theinventionofwritingenabledcomplex
societiestoarise:recordkeepingandlibrariesservedasastorehouseof VitruvianManbyLeonardodaVinci
knowledgeandincreasedtheculturaltransmissionofinformation.Humans epitomizestheadvancesinartand
nolongerhadtospendalltheirtimeworkingforsurvival,enablingthefirst scienceseenduringtheRenaissance.
specializedoccupations(e.g.craftsmen,merchants,priests,etc...).
Curiosityandeducationdrovethepursuitofknowledgeandwisdom,and
variousdisciplines,includingscience(inaprimitiveform),arose.Thisinturnledtotheemergenceofincreasingly
largerandmorecomplexcivilizations,suchasthefirstempires,whichattimestradedwithoneanother,orfought
forterritoryandresources.
Byaround500BC,therewereadvancedcivilizationsintheMiddleEast,Iran,India,China,andGreece,attimes
expanding,attimesenteringintodecline.[175]:3In221BC,Chinabecameasinglepolitythatwouldgrowtospread
itsculturethroughoutEastAsia,andithasremainedthemostpopulousnationintheworld.Thefundamentalsof
WesterncivilizationwerelargelyshapedinAncientGreece,withtheworld'sfirstdemocraticgovernmentand
majoradvancesinphilosophy,science,andmathematics,andinAncientRomeinlaw,government,and
engineering.[183]TheRomanEmpirewasChristianizedbyEmperorConstantineintheearly4thcenturyand
declinedbytheendofthe5th.Beginningwiththe7thcentury,ChristianizationofEuropebegan.In610,Islamwas
foundedandquicklybecamethedominantreligioninWesternAsia.TheHouseofWisdomwasestablishedin
AbbasideraBaghdad,Iraq.[184]ItisconsideredtohavebeenamajorintellectualcenterduringtheIslamicGolden
Age,whereMuslimscholarsinBaghdadandCairoflourishedfromtheninthtothethirteenthcenturiesuntilthe
MongolsackofBaghdadin1258AD.In1054ADtheGreatSchismbetweentheRomanCatholicChurchandthe
EasternOrthodoxChurchledtotheprominentculturaldifferencesbetweenWesternandEasternEurope.
Inthe14thcentury,theRenaissancebeganinItalywithadvancesinreligion,art,andscience.[175]:317319Atthat
timetheChristianChurchasapoliticalentitylostmuchofitspower.In1492,ChristopherColumbusreachedthe
Americas,initiatinggreatchangestothenewworld.Europeancivilizationbegantochangebeginningin1500,
leadingtothescientificandindustrialrevolutions.Thatcontinentbegantoexertpoliticalandculturaldominance
overhumansocietiesaroundtheworld,atimeknownastheColonialera(alsoseeAgeofDiscovery).[175]:295299
Inthe18thcenturyaculturalmovementknownastheAgeofEnlightenmentfurthershapedthementalityof
Europeandcontributedtoitssecularization.From1914to1918and1939to1945,nationsaroundtheworldwere
embroiledinworldwars.EstablishedfollowingWorldWarI,theLeagueofNationswasafirststepinestablishing
internationalinstitutionstosettledisputespeacefully.AfterfailingtopreventWorldWarII,mankind'sbloodiest
conflict,itwasreplacedbytheUnitedNations.Afterthewar,manynewstateswereformed,declaringorbeing
grantedindependenceinaperiodofdecolonization.TheUnitedStatesandSovietUnionbecametheworld's
dominantsuperpowersforatime,andtheyheldanoftenviolentrivalryknownastheColdWaruntilthe
dissolutionofthelatter.In1992,severalEuropeannationsjoinedintheEuropeanUnion.Astransportationand
communicationimproved,theeconomiesandpoliticalaffairsofnationsaroundtheworldhavebecome
increasinglyintertwined.Thisglobalizationhasoftenproducedbothconflictandcooperation.
Recentevents
Changehascontinuedatarapidpacefromthemid1940stotoday.
Technologicaldevelopmentsincludenuclearweapons,computers,genetic
engineering,andnanotechnology.Economicglobalization,spurredby
advancesincommunicationandtransportationtechnology,hasinfluenced
everydaylifeinmanypartsoftheworld.Culturalandinstitutionalforms
suchasdemocracy,capitalism,andenvironmentalismhaveincreased
influence.Majorconcernsandproblemssuchasdisease,war,poverty,
violentradicalism,andrecently,humancausedclimatechangehaverisen
astheworldpopulationincreases.
In1957,theSovietUnionlaunchedthefirstartificialsatelliteintoorbit
and,soonafterward,YuriGagarinbecamethefirsthumaninspace.Neil
Armstrong,anAmerican,wasthefirsttosetfootonanotherastronomical
object,theMoon.Unmannedprobeshavebeensenttoalltheknown AstronautBruceMcCandlessII
planetsinthesolarsystem,withsome(suchasVoyager)havingleftthe outsideofthespaceshuttle
solarsystem.TheSovietUnionandtheUnitedStatesweretheearliest Challengerin1984
leadersinspaceexplorationinthe20thcentury.Fivespaceagencies,
representingoverfifteencountries,[185]haveworkedtogethertobuildtheInternationalSpaceStation.Aboardit,
therehasbeenacontinuoushumanpresenceinspacesince2000.[186]TheWorldWideWebbecameapartof
everydaylifeinthe1990s,andsincethenhasbecomeanindispensablesourceofinformationinthedeveloped
world.
Seealso
Chronologyoftheuniverse Riskstocivilization,humans,andplanetEarth
Detailedlogarithmictimeline Timelineofevolutionaryhistoryoflife
Evolutionaryhistoryoflife Timelineofnaturalhistory
FutureoftheEarth
GeologicalhistoryofEarth
Notes
1.Pluto'ssatelliteCharonisrelativelylarger,[44]butPlutoisdefinedasadwarfplanet.[45]
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Dalrymple,G.B.(1991).TheAgeoftheEarth.California:StanfordUniversityPress.ISBN9780804715690.
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Dawkins,Richard(2004).TheAncestor'sTale:APilgrimagetotheDawnofLife.Boston:HoughtonMifflinCompany.
ISBN9780618005833.
Gradstein,F.M.Ogg,JamesGeorgeSmith,AlanGilbert,eds.(2004).AGeologicalTimeScale2004.Reprintedwith
corrections2006.CambridgeUniversityPress.ISBN9780521786737.
Gradstein,FelixM.Ogg,JamesG.vanKranendonk,Martin(2008).OntheGeologicalTimeScale2008(http://www.ny
sm.nysed.gov/nysgs/resources/images/geologicaltimescale.pdf)(PDF)(Report).InternationalCommissionon
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Levin,H.L.(2009).TheEarththroughtime(9thed.).SaundersCollegePublishing.ISBN9780470387740.
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Melosh,H.J.Vickery,A.M.&Tonks,W.B.(1993).Impactsandtheearlyenvironmentandevolutionoftheterrestrial
planets,inLevy,H.J.&Lunine,J.I.(eds.):ProtostarsandPlanetsIII,UniversityofArizonaPress,Tucson,pp.1339
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Externallinks
Davies,Paul."Quantumleapoflife(https://www.theguardian.com/technology/2005/dec/20/comment.scienc
e)".TheGuardian.2005December20.discussesspeculationontheroleofquantumsystemsintheorigin
oflife
Evolutiontimeline(http://www.johnkyrk.com/evolution.html)(usesShockwave).Animatedstoryoflife
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organismstotheascentofman.
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EvolutionoftheEarth(http://historystack.com/30_Major_Events_in_History_of_the_Earth).Timelineofthe
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