Chapter24

TheOriginofSpecies
LectureOutline
Overview

DarwinvisitedtheGalpagosIslandsandfoundthemfilledwithplantsandanimalsthat
livednowhereelseintheworld.

Herealizedthathewasobservingnewlyemergedspeciesontheseyoungislands.

Speciationtheoriginofnewspeciesisatthefocalpointofevolutionarytheorybecause
theappearanceofnewspeciesisthesourceofbiologicaldiversity.

Microevolutionisthestudyofadaptivechangeinapopulation.

Macroevolutionaddressesevolutionarychangesabovethespecieslevel.

Itdealswithquestionssuchastheappearanceofevolutionarynovelties(e.g.,feathersand
flightinbirds)thatcanbeusedtodefinehighertaxa.

Speciationaddressesthequestionofhownewspeciesoriginateanddevelopthroughthe
subdivisionandsubsequentdivergenceofgenepools.

Thefossilrecordchroniclestwopatternsofspeciation:anagenesisandcladogenesis.

Anagenesis,phyleticevolution,istheaccumulationofchangesassociatedwiththegradual
transformationofonespeciesintoanother.

IG Lecture Outlines 24-1

Cladogenesis,branchingevolution,isthebuddingofoneormorenewspeciesfromaparent
species.

Onlycladogenesispromotesbiologicaldiversitybyincreasingthenumberofspecies.

A.WhatIsaSpecies?

SpeciesisaLatinwordmeaningkindorappearance.

Traditionally,morphologicaldifferenceshavebeenusedtodistinguishspecies.

Today,differencesinbodyfunction,biochemistry,behavior,andgeneticmakeuparealso
usedtodifferentiatespecies.

Areorganismstrulydividedintothediscreteunitswecalledspecies,oristhisclassification
anarbitraryattempttoimposeorderonthenaturalworld?

1.Thebiologicalspeciesconceptemphasizesreproductiveisolation.

In1942,ErnstMayrproposedthebiologicalspeciesconcept.

Aspeciesisdefinedasapopulationorgroupofpopulationswhosemembershavethe
potentialtobreedwitheachotherinnaturetoproduceviable,fertileoffspring,butwho
cannotproduceviable,fertileoffspringwithmembersofotherspecies.

Abiologicalspeciesisthelargestsetofpopulationsinwhichgeneticexchangeis
possibleandthatisgeneticallyisolatedfromotherpopulations.
IG Lecture Outlines 24-2

Speciesarebasedoninterfertility,notphysicalsimilarity.

Forexample,easternandwesternmeadowlarkshavesimilarshapesandcoloration,but
differencesinsonghelppreventinterbreedingbetweenthetwospecies.

Incontrast,humanshaveconsiderablediversity,butweallbelongtothesamespecies
becauseofourcapacitytointerbreed.

2.Prezygoticandpostzygoticbarriersisolatethegenepoolsofbiologicalspecies.

Becausethedistinctionbetweenbiologicalspeciesdependsonreproductiveincompatibility,
theconcepthingesonreproductiveisolation,

theexistenceofbiologicalbarriersthat
preventmembersoftwospeciesfromproducingviable,fertilehybrids.

Asinglebarriermaynotblockallgeneticexchangebetweenspecies,butacombinationof
severalbarrierscaneffectivelyisolateaspeciesgenepool.

Typically,thesebarriersareintrinsictotheorganisms,notduetosimplegeographic
separation.

Reproductiveisolationpreventspopulationsbelongingtodifferentspeciesfrom
interbreeding,eveniftheirrangesoverlap.

Reproductivebarrierscanbecategorizedasprezygoticorpostzygotic,dependingonwhether
theyfunctionbeforeoraftertheformationofzygotes.

Prezygoticbarriersimpedematingbetweenspeciesorhinderfertilizationofovaif
membersofdifferentspeciesattempttomate.
IG Lecture Outlines 24-3

Thesebarriersincludehabitatisolation,behavioralisolation,temporalisolation,
mechanicalisolation,andgameticisolation.

Habitatisolation.Twoorganismsthatusedifferenthabitats(eveninthesamegeographic
area)areunlikelytoencountereachothertoevenattemptmating.

TwospeciesofgartersnakesinthegenusThamnophisoccurinthesameareas.Because
onelivesmainlyinwaterandtheotherisprimarilyterrestrial,theyrarelyencountereach
other.

Behavioralisolation.Manyspeciesuseelaboratecourtshipbehaviorsuniquetothespecies
toattractmates.

Inmanyspecies,elaboratecourtshipdisplaysidentifypotentialmatesofthecorrect
speciesandsynchronizegonadalmaturation.

Inthebluefootedbooby,malesperformahighstepdancethatcallsthefemales
attentiontothemalesbrightbluefeet.

Temporalisolation.Twospeciesthatbreedduringdifferenttimesofday,differentseasons,
ordifferentyearscannotmixgametes.

Thegeographicrangesofthewesternspottedskunkandtheeasternspottedskunk
overlap.However,theydonotinterbreedbecausetheformermatesinlatesummerand
thelatterinlatewinter.

Mechanicalisolation.Closelyrelatedspeciesmayattempttomatebutfailbecausetheyare
anatomicallyincompatibleandtransferofspermisnotpossible.

IG Lecture Outlines 24-4

Forexample,mechanicalbarrierscontributetothereproductiveisolationofflowering
plantsthatarepollinatedbyinsectsorotheranimals.

Withmanyinsects,themaleandfemalecopulatoryorgansofcloselyrelatedspeciesdo
notfittogether,preventingspermtransfer.

Gameticisolation.Thegametesoftwospeciesdonotformazygotebecauseof
incompatibilitiespreventingfertilization.

Inspecieswithinternalfertilization,theenvironmentofthefemalereproductivetract
maynotbeconducivetothesurvivalofspermfromotherspecies.

Forspecieswithexternalfertilization,gameterecognitionmayrelyonthepresenceof
specificmoleculesontheeggscoat,whichadhereonlytospecificmoleculesonsperm
cellsofthesamespecies.

Asimilarmolecularrecognitionmechanismenablesaflowertodiscriminatebetween
pollenofthesamespeciesandpollenofadifferentspecies.

Ifaspermfromonespeciesdoesfertilizetheovumofanother,postzygoticbarriers

may
preventthehybridzygotefromdevelopingintoaviable,fertileadult.

Thesebarriersincludereducedhybridviability,reducedhybridfertility,andhybrid
breakdown.

Reducedhybridviability.Geneticincompatibilitybetweenthetwospeciesmayabortthe
developmentofthehybridatsomeembryonicstageorproducefrailoffspring.

IG Lecture Outlines 24-5

ThisistruefortheoccasionalhybridsbetweenfrogsinthegenusRana.Mostdonot
completedevelopment,andthosethatdoarefrail.

Reducedhybridfertility.Evenifthehybridoffspringarevigorous,thehybridsmaybe
infertile,andthehybridcannotbackbreedwitheitherparentalspecies.

Thisinfertilitymaybeduetoproblemsinmeiosisbecauseofdifferencesinchromosome
numberorstructure.

Forexample,whileamule,thehybridproductofmatingbetweenahorseanddonkey,is
arobustorganism,itcannotmate(exceptveryrarely)witheitherhorsesordonkeys.

Hybridbreakdown.Insomecases,firstgenerationhybridsareviableandfertile.

However,whentheymatewitheitherparentspeciesorwitheachother,thenext
generationisfeebleorsterile.

Strainsofcultivatedricehaveaccumulateddifferentmutantrecessiveallelesattwoloci
inthecourseoftheirdivergencefromacommonancestor.

Hybridsbetweenthemarevigorousandfertile,butplantsinthenextgenerationthat
carrytoomanyoftheserecessiveallelesaresmallandsterile.

Thesestrainsareintheprocessofspeciating.

Reproductivebarrierscanoccurbeforemating,betweenmatingandfertilization,orafter
fertilization.

3.Thebiologicalspeciesconcepthassomemajorlimitations.
IG Lecture Outlines 24-6

Whilethebiologicalspeciesconcepthashadanimportantimpactonevolutionarytheory,it
islimitedwhenappliedtospeciesinnature.

Forexample,onecannottestthereproductiveisolationofmorphologicallysimilar
fossils,whichareseparatedintospeciesbasedonmorphology.

Evenforlivingspecies,weoftenlackinformationoninterbreedingneededtoapplythe
biologicalspeciesconcept.

Inaddition,manyspecies(e.g.,bacteria)reproduceentirelyasexuallyandareassignedto
speciesbasedmainlyonstructuralandbiochemicalcharacteristics.

Manybacteriatransfergenesbyconjugationandotherprocesses,butthistransferis
differentfromsexualrecombination.

4.Evolutionarybiologistshaveproposedseveralalternativeconceptsofspecies.

Severalalternativespeciesconceptsemphasizetheprocessesthatunitethemembersofa
species.

Theecologicalspeciesconcept

definesaspeciesintermsofitsecologicalniche,thesetof
environmentalresourcesthataspeciesusesanditsroleinabiologicalcommunity.

Asanexample,aspeciesthatisaparasitemaybedefinedinpartbyitsadaptationstoa
specificorganism.

Thisconceptaccommodatesasexualandsexualspecies.

IG Lecture Outlines 24-7

Thepaleontologicalspeciesconcept

focusesonmorphologicallydiscretespeciesknown
onlyfromthefossilrecord.

Thereislittleornoinformationaboutthematingcapabilityoffossilspecies,andthe
biologicalspeciesconceptisnotusefulforthem.

Thephylogeneticspeciesconcept

definesaspeciesasasetoforganismswithaunique
genetichistory.

Biologistscomparethephysicalcharacteristicsormolecularsequencesofspeciesto
thoseofotherorganismstodistinguishgroupsofindividualsthataresufficientlydifferent
tobeconsideredseparatespecies.

Siblingspeciesarespeciesthatappearsosimilarthattheycannotbedistinguishedon
morphologicalgrounds.

Scientistsapplythebiologicalspeciesconcepttodetermineifthephylogenetic
distinctionisconfirmedbyreproductiveincompatibility.

Themorphologicalspeciesconcept,theoldestandstillmostpractical,definesaspeciesby
auniquesetofstructuralfeatures.

Themorphologicalspeciesconcepthascertainadvantages.Itcanbeappliedtoasexual
andsexualspecies,anditcanbeusefulevenwithoutinformationabouttheextentofgene
flow.

However,thisdefinitionreliesonsubjectivecriteria,andresearcherssometimesdisagree
aboutwhichstructuralfeaturesidentifyaspecies.

IG Lecture Outlines 24-8

Inpractice,scientistsusethemorphologicalspeciesconcepttodistinguishmostspecies.

Eachspeciesconceptmaybeuseful,dependingonthesituationandthetypesofquestionswe
areasking.

B.ModesofSpeciation

Twogeneralmodesofspeciationaredistinguishedbythewaygeneflowamongpopulations
isinitiallyinterrupted.

Inallopatricspeciation,

geographicseparationofpopulationsrestrictsgeneflow.

Insympatricspeciation,

speciationoccursingeographicallyoverlappingpopulationswhen
biologicalfactors,suchaschromosomalchangesandnonrandommating,reducegeneflow.

1.Allopatricspeciation:geographicbarrierscanleadtotheoriginofspecies.

Severalgeologicalprocessescanfragmentapopulationintotwoormoreisolated
populations.

Mountainranges,glaciers,landbridges,orsplinteringoflakesmaydivideonepopulation
intoisolatedgroups.

Alternatively,someindividualsmaycolonizeanew,geographicallyremoteareaand
becomeisolatedfromtheparentpopulation.

IG Lecture Outlines 24-9

Forexample,mainlandorganismsthatcolonizedtheGalpagosIslandswereisolated
frommainlandpopulations.

Howsignificantabarriermustbetolimitgeneexchangedependsontheabilityoforganisms
tomoveabout.

Ageologicalfeaturethatisonlyaminorhindrancetoonespeciesmaybeanimpassible
barriertoanother.

ThevalleyoftheGrandCanyonisasignificantbarrierforthegroundsquirrelsthathave
speciatedonoppositesides

Forbirdsthatcanflyacrossthecanyon,itisnobarrier.

Oncegeographicseparationisestablished,theseparatedgenepoolsmaybegintodiverge
throughanumberofmechanisms.

Mutationsarise.

Sexualselectionfavorsdifferenttraitsinthetwopopulations.

Differentselectivepressuresindifferingenvironmentsactonthetwopopulations.

Geneticdriftaltersallelefrequencies.

Asmall,isolatedpopulationismorelikelytohaveitsgenepoolchangedsubstantiallyovera
shortperiodoftimebygeneticdriftandnaturalselection.

IG Lecture Outlines 24-10

Forexample,lessthan2millionyearsago,smallpopulationsofstrayplantsandanimals
fromtheSouthAmericanmainlandcolonizedtheGalpagosIslandsandgaverisetothe
speciesthatnowinhabittheislands.

However,veryfewsmall,isolatedpopulationsdevelopintonewspecies;mostsimplypersist
orperishintheirnewenvironment.

Toconfirmthatallopatricspeciationhasoccurred,itisnecessarytodeterminewhetherthe
separatedpopulationshavebecomedifferentenoughthattheycannolongerinterbreedand
producefertileoffspringwhentheycomebackincontact.

Insomecases,researchersbringtogethermembersofseparatedpopulationsina
laboratorysetting.

Biologistscanalsoassessallopatricspeciationinthewild.

Forexample,femalesoftheGalpagosgroundfinchGeospizadifficilisrespondtothe
songsofmalesfromthesameislandbutignorethesongsofmalesofthesame
speciesfromotherislands.

2.Sympatricspeciation:anewspeciescanoriginateinthegeographicmidstoftheparent
species.

Insympatricspeciation,newspeciesarisewithintherangeoftheparentpopulations.

Herereproductivebarriersmustevolvebetweensympatricpopulations.

IG Lecture Outlines 24-11

Inplants,sympatricspeciationcanresultfromaccidentsduringcelldivisionthatresultin
extrasetsofchromosomes,amutantconditionknownaspolyploidy.

Inanimals,itmayresultfromgenebasedshiftsinhabitatormatepreference.

Anindividualcanhavemorethantwosetsofchromosomes.

Anautopolyploid

mutantisanindividualthathasmorethantwochromosomesets,all
derivedfromasinglespecies.

Forexample,afailureofmitosisormeiosiscandoubleacellschromosomenumber
fromdiploid(2n)totetraploid(4n).

Thetetraploidcanreproducewithitself(selfpollination)orwithothertetraploids.

Itcannotmatewithdiploidsfromtheoriginalpopulation,becauseofabnormalmeiosis
bythetriploidhybridoffspring.

Amorecommonmechanismofproducingpolyploidindividualsoccurswhenallopolyploid
offspringareproducedbythematingoftwodifferentspecies.

Whilethehybridsareusuallysterile,theymaybequitevigorousandpropagate
asexually.

Insubsequentgenerations,variousmechanismsmaytransformasterilehybridintoa
fertilepolyploid.

Thesepolyploidhybridsarefertilewitheachotherbutcannotbreedwitheitherparent
species.

IG Lecture Outlines 24-12

Theythusrepresentanewbiologicalspecies.

Theoriginofpolyploidplantspeciesiscommonandrapidenoughthatscientistshave
documentedseveralsuchspeciationsinhistoricaltimes.

Forexample,twonewspeciesofplantscalledgoatsbeard(Tragopodon)appearedin
IdahoandWashingtonintheearly1900s.

TheyaretheresultsofallopolyploidyeventsbetweenpairsofintroducedEuropean
Tragopodonspecies.

Manyplantsimportantforagriculturearepolyploid.

Forexample,wheatisanallohexaploid,withsixsetsofchromosomesfromthree
differentspecies.

Oats,cotton,potatoes,andtobaccoarepolyploid.

Plantgeneticistsnowusechemicalsthatinducemeioticandmitoticerrorstocreatenew
polyploidplantswithspecialqualities.

Oneexampleisanartificialhybridcombiningthehighyieldofwheatwiththe
hardinessanddiseaseresistanceofrye.

Whilepolyploidspeciationdoesoccurinanimals,othermechanismsalsocontributeto
sympatricspeciationinanimals.

Reproductiveisolationcanresultwhengeneticfactorscauseindividualstoexploit
resourcesnotusedbytheparent.

OneexampleistheNorthAmericanmaggotfly,Rhagoletispomonella.
IG Lecture Outlines 24-13

Theflysoriginalhabitatwasnativehawthorntrees.

About200yearsago,somepopulationscolonizednewlyintroducedappletrees.

Becauseapplesmaturemorequicklythanhawthornfruit,theapplefeedingflieshave
beenselectedformorerapiddevelopmentandnowshowtemporalisolationfromthe
hawthornfeedingmaggotflies.

Speciationisunderway.

Sympatricspeciationisonemechanismthathasbeenproposedfortheexplosiveadaptive
radiationofcichlidfishesinLakeVictoria,Africa.

Thisvast,shallowlakehasfilledanddrieduprepeatedlyduetoclimatechanges.

Thecurrentlakeisonly12,000yearsoldbutishometo600speciesofcichlidfishes.

Thespeciesaresogeneticallysimilarthatmanyhavelikelyarisensincethelakelast
filled.

Whilethesespeciesareclearlyspecializedforexploitingdifferentfoodresourcesand
otherresources,nonrandommatinginwhichfemalesselectmalesbasedonacertain
appearancehasprobablycontributed,too.

Individualsoftwocloselyrelatedsympatriccichlidspecieswillnotmateundernormallight
becausefemaleshavespecificcolorpreferencesandmalesdifferincolor.

However,underlightconditionsthatdeemphasizecolordifferences,femaleswillmate
withmalesoftheotherspeciesandproduceviable,fertileoffspring.

IG Lecture Outlines 24-14

Itseemslikelythattheancestralpopulationwaspolymorphicforcolorandthat
divergencebeganwiththeappearanceoftwoecologicalnichesthatdividedthefishinto
subpopulations.

Geneticdriftresultedinchancedifferencesinthegeneticmakeupofthesubpopulations,
withdifferentmalecolorsandfemalepreferences.

Sexualselectionreinforcedthecolordifferences.

Thelackofpostzygoticbarriersinthiscasesuggeststhatspeciationoccurredrelatively
recently.

AspollutioncloudsthewatersofLakeVictoria,itbecomesmoredifficultforfemale
cichlidstoseedifferencesinmalecolor.

Thegenepoolsofthesetwocloselyrelatedspeciesmayblendagain.

Wewillsummarizethedifferencesbetweensympatricandallopatricspeciation.

Inallopatricspeciation,anewspeciesformswhilegeographicallyisolatedfromitsparent
population.

Astheisolatedpopulationaccumulatesgeneticdifferencesduetonaturalselectionand
geneticdrift,reproductiveisolationfromtheancestralspeciesmayariseasabyproduct
ofthegeneticchange.

Suchreproductivebarrierspreventbreedingwiththeparentevenifthepopulations
reestablishcontact.

IG Lecture Outlines 24-15

Sympatricspeciationrequirestheemergenceofsomereproductivebarrierthatisolatesa
subsetofthepopulationwithoutgeographicseparationfromtheparentpopulation.

Inplants,themostcommonmechanismishybridizationbetweenspeciesorerrorsincell
divisionthatleadtopolyploidindividuals.

Inanimals,sympatricspeciationmayoccurwhenasubsetofthepopulationis
reproductivelyisolatedbyaswitchinfoodsourceorbysexualselectioninapolymorphic
population.

C.AdaptiveRadiation

Theevolutionofmanydiverselyadaptedspeciesfromacommonancestorwhennew
environmentalopportunitiesariseiscalledadaptiveradiation.

Adaptiveradiationoccurswhenafeworganismsmaketheirwayintonewareasorwhen
extinctionopensupecologicalnichesforthesurvivors.

Amajoradaptiveradiationofmammalsfollowedtheextinctionofthedinosaurs65
millionyearsago.

TheHawaiianarchipelagoisashowcaseofadaptiveradiation.

Located3,500kmfromthenearestcontinent,thevolcanicislandswereformednaked
andgraduallypopulatedbystrayorganismsthatarrivedbywindoroceancurrents.

Theislandsarephysicallydiverse,witharangeofaltitudesandrainfall.

IG Lecture Outlines 24-16

Multipleinvasionsandallopatricandsympatricspeciationeventshaveignitedan
explosionofadaptiveradiationofnovelspecies.

1.Researchersstudythegeneticsofspeciation.

Researchershavemadegreatstridesinunderstandingtheroleofgenesinparticular
speciationevents.

DouglasSchemskeandhiscolleaguesatMichiganStateUniversityexaminedtwospeciesof
Mimulus.

Thetwospeciesarepollinatedbybeesandhummingbirdsrespectively,keepingtheir
genepoolsseparatethroughprezygoticisolation.

Thespeciesshownopostzygoticisolationandcanbematedreadilyinthegreenhouseto
producehybridswithflowersthatvaryincolorandshape.

Researchersobservedwhichpollinatorsvisitwhichflowersandtheninvestigatedthe
geneticdifferencesbetweenplants.

Twogenelocihavebeenidentifiedthatarelargelyresponsibleforpollinatorchoice.

Onelocusinfluencesflowercolor;theotheraffectstheamountofnectarflowersproduce.

Bydeterminingattractivenessoftheflowerstodifferentpollinators,allelicdiversityat
theselocihasledtospeciation.

IG Lecture Outlines 24-17

2.Thetempoofspeciationisimportant.

Inthefossilrecord,manyspeciesappearasnewformsrathersuddenly(ingeologicterms),
persistessentiallyunchanged,andthendisappearfromthefossilrecord.

Darwinnotedthiswhenheremarkedthatspeciesappearedtoundergomodificationsduring
relativelyshortperiodsoftheirtotalexistenceandthenremainedessentiallyunchanged.

PaleontologistsNilesEldredgeandStephenJayGouldcoinedthetermpunctuated

equilibriumtodescribetheseperiodsofapparentstasispunctuatedbysuddenchange.

SomescientistssuggestthatthesepatternsrequireanexplanationoutsidetheDarwinian
modelofdescentwithmodification.

However,thisisnotnecessarilythecase.

Supposethataspeciessurvivedfor5millionyears,butmostofitsmorphologicalalterations
occurredinthefirst50,000yearsofitsexistencejust1%ofitstotallifetime.

Becausetimeperiodsthisshortoftencannotbedistinguishedinfossilstrata,thespecies
wouldseemtohaveappearedsuddenlyandthenlingeredwithlittleornochangebefore
becomingextinct.

Eventhoughtheemergenceofthisspeciesactuallytooktensofthousandsofyears,this
periodofchangeleftnofossilrecord.

Stasiscanalsobeexplained.

Allspeciescontinuetoadaptaftertheyarise,butoftenbychangesthatdonotleavea
fossilrecord,suchassmallbiochemicalmodifications.
IG Lecture Outlines 24-18

Paleontologistsbasehypothesesofdescentalmostentirelyonexternalmorphology.

Duringperiodsofapparentequilibrium,changesinbehavior,internalanatomy,and
physiologymaynotleaveafossilrecord.

Iftheenvironmentchanges,thestasiswillbebrokenbypunctuationsthatleavevisibletraces
inthefossilrecord.

D.FromSpeciationtoMacroevolution

Speciationisattheboundarybetweenmicroevolutionandmacroevolution.

Microevolutionisachangeovergenerationsinapopulationsallelefrequencies,mainly
bygeneticdriftandnaturalselection.

Speciationoccurswhenapopulationsgeneticdivergencefromitsancestralpopulation
resultsinreproductiveisolation.

Whilethechangesafteranyspeciationeventmaybesubtle,thecumulativechangeover
millionsofspeciationepisodesmustaccountformacroevolution,thescaleofchanges
seeninthefossilrecord.

1.Mostevolutionarynoveltiesaremodifiedversionsofolderstructures.

TheDarwinianconceptofdescentwithmodificationcanaccountforthemajor
morphologicaltransformationsofmacroevolution.

IG Lecture Outlines 24-19

Itmaybedifficulttobelievethatacomplexorganlikethehumaneyecouldbetheproductof
gradualevolution,ratherthanafinisheddesigncreatedspeciallyforhumans.

However,thekeyistorememberisthataverysimpleeyecanbeveryusefultoananimal.

Thesimplesteyesarejustclustersofphotoreceptors,lightsensitivepigmentedcells.

Thesesimpleeyesappeartohavehadasingleevolutionaryorigin.

Theyarenowfoundinavarietyofanimals,includinglimpets.

Thesesimpleeyeshavenolensesandcannotfocusanimage,buttheydoallowtheanimalto
distinguishlightfromdark.

Limpetsclingtightlytotheirrockswhenashadowfallsonthem,reducingtheirriskof
predation.

Complexeyeshaveevolvedseveraltimesindependentlyintheanimalkingdom.

Examplesofvariouslevelsofcomplexity,fromclustersofphotoreceptorstocameralike
eyes,canbeseeninmolluscs.

Themostcomplextypesdidnotevolveinonequantumleap,butbyincremental
adaptationoforgansthatbenefitedtheirownersateachstage.

Evolutionarynoveltiescanalsoarisebygradualrefinementofexistingstructuresfornew
functions.

Structuresthatevolveinonecontext,butbecomecooptedforanotherfunction,are
exaptations.

IG Lecture Outlines 24-20

Itisimportanttorecognizethatnaturalselectioncanonlyimproveastructureinthecontext
ofitscurrentutility,notinanticipationofthefuture.

Anexampleofanexaptationisthechangingfunctionoflightweight,honeycombedbonesof
birds.

Thefossilrecordindicatesthatlightbonespredatedflight.

Therefore,theymusthavehadsomefunctionontheground,perhapsasalightframefor
agile,bipedaldinosaurs.

Onceflightbecameanadvantage,naturalselectionwouldhaveremodeledtheskeletonto
betterfittheiradditionalfunction.

Thewinglikeforelimbsandfeathersthatincreasedthesurfaceareaoftheseforelimbs
werecooptedforflightafterfunctioninginsomeothercapacity,suchascourtship,
thermoregulation,orcamouflage.

2.Genes

thatcontroldevelopmentplayamajorroleinevolution.

Evodevoisafieldofinterdisciplinaryresearchthatexamineshowslightgenetic
divergencescanbecomemagnifiedintomajormorphologicaldifferencesbetweenspecies.

Aparticularfocusisongenesthatprogramdevelopmentbycontrollingtherate,timing,and
spatialpatternofchangesinformasanorganismdevelopsfromazygotetoanadult.

Heterochrony,anevolutionarychangeintherateortimingofdevelopmentalevents,hasled
tomanystrikingevolutionarytransformations.
IG Lecture Outlines 24-21

Allometricgrowthtrackshowproportionsofstructureschangeduetodifferentgrowthrates
duringdevelopment.

Changerelativeratesofgrowthevenslightly,andyoucanchangetheadultform
substantially.

Differentallometricpatternscontributetothecontrastofadultskullshapesbetween
humansandchimpanzees,whichbothdevelopedfromfairlysimilarfetalskulls.

Heterochronyappearstoberesponsiblefordifferencesinthefeetoftreedwellingversus
grounddwellingsalamanders.

Thefeetofthetreedwellersareadaptedforclimbingvertically,withshorterdigitsand
morewebbing.

Thismodificationmayhaveevolvedduetomutationsintheallelesthatcontrolthe
timingoffootdevelopment.

Stuntedfeetmayhaveresultedifregulatorygenesswitchedofffootgrowthearly.

Inthisway,arelativelysmallgeneticchangecanbeamplifiedintosubstantial
morphologicalchange.

Anotherformofheterochronyisconcernedwiththerelativetimingofreproductive
developmentandsomaticdevelopment.

Iftherateofreproductivedevelopmentacceleratescomparedtosomaticdevelopment,thena
sexuallymaturestagecanretainjuvenilestructuresaprocesscalledpaedomorphosis.

IG Lecture Outlines 24-22

Somespeciesofsalamanderhavethetypicalexternalgillsandflattenedtailofanaquatic
juvenilebuthavefunctioninggonads.

Macroevolutioncanalsoresultfromchangesingenesthatcontroltheplacementandspatial
organizationofbodyparts.

Forexample,genescalledhomeotic

genesdeterminesuchbasicfeaturesaswhereapair
ofwingsandapairoflegswilldeveloponabirdorhowaplantsflowerpartsare
arranged.

Theproductsofoneclassofhomeoticgenes,theHoxgenes,providepositionalinformation
inananimalembryo.

Thisinformationpromptscellstodevelopintostructuresappropriateforaparticular
location.

Onemajortransitionintheevolutionofvertebratesisthedevelopmentofthewalkinglegsof
tetrapodsfromthefinsoffishes.

Afishfinthatlacksexternalskeletalsupportevolvedintoatetrapodlimbthatextends
skeletalsupports(digits)tothetipofthelimb.

ThismaybetheresultofchangesinthepositionalinformationprovidedbyHoxgenes
duringlimbdevelopment,determininghowfardigitsandotherbonesshouldextendfrom
thelimb.

3.Evolutionisnotgoaloriented.
IG Lecture Outlines 24-23

Thefossilrecordshowsapparentevolutionarytrends.

Forexample,theevolutionofthemodernhorsecanbeinterpretedtohavebeenasteady
seriesofchangesfromasmall,browsingancestor(Hyracotherium)withfourtoesto
modernhorses(Equus)withonlyonetoeperfootandteethmodifiedforgrazingon
grasses.

ItispossibletoarrangeasuccessionofanimalsintermediatebetweenHyracotheriumand
modernhorsestoshowtrendstowardincreasedsize,reducednumberoftoes,and
modificationsofteethforgrazing.

Ifwelookatallfossilhorses,theillusionofcoherent,progressiveevolutionleadingdirectly
tomodernhorsesvanishes.

Equusistheonlysurvivingtwigofanevolutionarybushthatincludedseveraladaptive
radiationsamongbothgrazersandbrowsers.

Differencesamongspeciesinsurvivalcanalsoproduceamacroevolutionarytrend.

Thespeciesselection

modeldevelopedbyStevenStanleyconsidersspeciesasanalogousto
individuals.

Speciationistheirbirth,extinctionistheirdeath,andnewspeciesaretheiroffspring.

Inthismodel,Stanleysuggestthatjustasindividualorganismsundergonaturalselection,
speciesundergospeciesselection.

Thespeciesthatendurethelongestandgeneratethegreatestnumberofnewspecies
determinethedirectionofmajorevolutionarytrends.
IG Lecture Outlines 24-24

Thespeciesselectionmodelsuggeststhatdifferentialspeciationsuccessplaysarolein
macroevolutionsimilartotheroleofdifferentialreproductivesuccessinmicroevolution.

Totheextentthatspeciationratesandspecieslongevityreflectsuccess,theanalogyto
naturalselectionisevenstronger.

However,qualitiesunrelatedtotheoverallsuccessoforganismsinspecificenvironments
maybeequallyimportantinspeciesselection.

Asanexample,theabilityofaspeciestodispersetonewlocationsmaycontributetoits
givingrisetoalargenumberofdaughterspecies.

Theappearanceofanevolutionarytrenddoesnotimplysomeintrinsicdrivetowarda
preordainedstateofbeing.

Evolutionisaresponsetointeractionsbetweenorganismsandtheircurrentenvironments,
leadingtochangesinevolutionarytrendsasconditionschange.

IG Lecture Outlines 24-25