Chapter22

DescentwithModification:
ADarwinianViewofLife
LectureOutline
Overview
OnNovember24,1859,CharlesDarwinpublishedOntheOriginofSpeciesbyMeansof
NaturalSelection.
Darwinsbookdrewacohesivepictureoflifebyconnectingwhathadonceseemeda
bewilderingarrayofunrelatedfacts.
DarwinmadetwomajorpointsinTheOriginofSpecies:
1. Todaysorganismsdescendedfromancestralspeciesthatweredifferentfrommodern
species.
2. Naturalselectionprovidedamechanismforthisevolutionarychange.
Thebasicideaofnaturalselectionisthatapopulationcanchangeovertimeifindividuals
thatpossesscertainheritabletraitsleavemoreoffspringthanotherindividuals.
Naturalselectionresultsinevolutionaryadaptation,anaccumulationofinherited
characteristicsthatincreasetheabilityofanorganismtosurviveandreproduceinits
environment.

Eventually,apopulationmayaccumulateenoughchangethatitconstitutesanewspecies.

Inmodernterms,wecandefineevolution

asachangeovertimeinthegenetic
compositionofapopulation.
Evolutionalsoreferstothegradualappearanceofallbiologicaldiversity.
Evolutionissuchafundamentalconceptthatitsstudyisrelevanttobiologyatevery
level,frommoleculestoecosystems.
Evolutionaryperspectivescontinuetotransformmedicine,agriculture,biotechnology,
andconservationbiology.
A.TheHistoricalContextforEvolutionaryTheory
1.Westerncultureresistedevolutionaryviewsoflife.
DarwinsviewoflifecontrastedwiththetraditionalviewofanEarththatwasafew
thousandyearsold,populatedbylifeformsthatwerecreatedatthebeginningandhad
remainedfundamentallyunchanged.
TheOriginofSpecieschallengedaworldviewthathadbeenlongaccepted.
TheGreekphilosopherAristotle(384322B.C.E.)opposedanyconceptofevolutionand
viewedspeciesasfixedandunchanging.

IG Lecture Outline 22-1

Aristotlebelievedthatalllivingformscouldbearrangedonaladderofincreasing
complexity(scalanaturae)withperfect,permanentspeciesoneveryrung.

TheOldTestamentaccountofcreationheldthatspecieswereindividuallydesignedby
Godand,therefore,perfect.
Inthe1700s,naturaltheology

viewedtheadaptationsoforganismsasevidencethatthe
Creatorhaddesignedeachspeciesforapurpose.
CarolusLinnaeus(17071778),aSwedishphysicianandbotanist,foundedtaxonomy,

a
systemfornamingspeciesandclassifyingspeciesintoahierarchyofincreasinglycomplex
categories.
Linnaeusdevelopedthebinomialsystemofnamingorganismsaccordingtogenusand
species.
Incontrasttothelinearhierarchyofthe
scalanaturae,Linnaeusadoptedanested
classificationsystem,groupingsimilarspeciesintoincreasinglygeneralcategories.
ForLinnaeus,similaritybetweenspeciesdidnotimplyevolutionarykinshipbutrather
thepatternoftheircreation.
Darwinsviewswereinfluencedbyfossils,

remainsortracesoforganismsfromthepast
mineralizedinsedimentaryrocks.
Sedimentaryrocksformwhenmudandsandsettletothebottomofseas,lakes,and
marshes.
Newlayersofsedimentcoverolderones,creatinglayersofrockcalledstrata.
Erosionmaylatercarvethroughsedimentaryrocktoexposeolderstrataatthesurface.
Fossilswithinlayersofsedimentaryrockshowthatasuccessionoforganismshave
populatedEarththroughouttime.
Paleontology,thestudyoffossils,waslargelydevelopedbytheFrenchanatomist
GeorgesCuvier(17691832).
InexaminingrockstrataintheParisBasin,Cuviernotedthattheolderthestrata,the
moredissimilarthefossilsfrommodernlife.
Cuvierrecognizedthatextinctionhadbeenacommonoccurrenceinthehistoryoflife.
Insteadofevolution,Cuvieradvocated

catastrophism,

speculatingthatboundaries
betweenstratawereduetolocalfloodsordroughtsthatdestroyedthespeciesthen
present.
Hesuggestedthatthedenudedareaswerelaterrepopulatedbyspeciesimmigratingfrom
unaffectedareas.
2.Theoriesofgeologicgradualismpreparedthepathforevolutionarybiologists.
IncontrasttoCuvierscatastrophism,ScottishgeologistJamesHutton(17261797)
proposedatheoryofgradualismthatheldthatprofoundgeologicalchangestookplace
throughthecumulativeeffectofslowbutcontinuousprocessesidenticaltothosecurrently
operating.
Thus,valleyswereformedbyriversflowingthroughrocksandsedimentaryrockswere
formedfromsoilparticlesthaterodedfromlandandwerecarriedbyriverstothesea.
Later,geologistCharlesLyell(17971875)proposedatheoryofuniformitarianism,

whichheldthatgeologicalprocesseshadnotchangedthroughoutEarthshistory.

HuttonsandLyellsobservationsandtheorieshadastronginfluenceonDarwin.

IG Lecture Outline 22-2

First,ifgeologicchangesresultfromslow,continuousprocessesratherthansudden
events,thentheEarthmustbefarolderthanthe6,000yearsestimatedbytheologians
frombiblicalinference.
Second,slowandsubtleprocessespersistingforlongperiodsoftimecanalsoacton
livingorganisms,producingsubstantialchangeoveralongperiodoftime.

3.Lamarckplacedfossilsinanevolutionarycontext.
In1809,FrenchbiologistJeanBaptistedeLamarck(17441829)publishedatheoryof
evolutionbasedonhisobservationsoffossilinvertebratesinthecollectionsoftheNatural
HistoryMuseumofParis.
Bycomparingfossilsandcurrentspecies,Lamarckfoundwhatappearedtobeseveral
linesofdescent.
Eachwasachronologicalseriesofoldertoyoungerfossils,leadingtoamodernspecies.
Heexplainedhisobservationswithtwoprinciples:useanddisuse

ofpartsandthe
inheritanceofacquiredcharacteristics.
Useanddisusewastheconceptthatbodypartsthatareusedextensivelybecomelarger
andstronger,whilethosethatarenotuseddeteriorate.
Theinheritanceofacquiredcharacteristicsstatedthatmodificationsacquiredduringthe
lifeofanorganismcouldbepassedtooffspring.
Aclassicexampleisthelongneckofthegiraffe.Lamarckreasonedthatthelong,
muscularneckofthemoderngiraffeevolvedovermanygenerationsastheancestorsof
giraffesreachedforleavesonhigherbranchesandpassedthischaracteristictotheir
offspring.
Lamarckthoughtthatevolutionarychangewasdrivenbytheinnatedriveoforganismsto
increasingcomplexity.
Lamarckstheorywasavisionaryattempttoexplainthefossilrecordandthecurrent
diversityoflifewithrecognitionofgradualevolutionarychange.
However,moderngeneticshasprovidednoevidencethatacquiredcharacteristicscanbe
inherited.
Acquiredtraitssuchasabodybuildersbiggerbicepsdonotchangethegenes
transmittedthroughgametestooffspring.
B.TheDarwinianRevolution

CharlesDarwin(18091882)wasborninwesternEngland.
Asaboy,hedevelopedaconsuminginterestinnature.
WhenDarwinwas16,hisfathersenthimtotheUniversityofEdinburghtostudy
medicine.

DarwinleftEdinburghwithoutadegreeandenrolledatCambridgeUniversitywiththe
intentofbecomingaclergyman.
Atthattime,mostnaturalistsandscientistsbelongedtotheclergyandviewedtheworld
inthecontextofnaturaltheology.

DarwinreceivedhisB.A.in1831.

AftergraduationDarwinjoinedthesurveyshipHMSBeagleasshipnaturalistand
conversationcompaniontoCaptainRobertFitzRoy.
FitzRoychoseDarwinbecauseofhiseducation,andbecausehisageandsocialclass
weresimilartothatofthecaptain.
IG Lecture Outline 22-3

1.FieldresearchhelpedDarwinframehisviewoflife.
TheprimarymissionofthefiveyearvoyageoftheBeaglewastochartpoorlyknown
stretchesoftheSouthAmericancoastline.
Darwinhadthefreedomtoexploreextensivelyonshorewhilethecrewsurveyedthe
coast.
HecollectedthousandsofspecimensoftheexoticanddiversefloraandfaunaofSouth
America.
DarwinexploredtheBrazilianjungles,thegrasslandsoftheArgentinepampas,the
desolationofTierradelFuegonearAntarctica,andtheheightsoftheAndes.
DarwinnotedthattheplantsandanimalsofSouthAmericawereverydistinctfromthose
ofEurope.
OrganismsfromtemperateregionsofSouthAmericamorecloselyresembledthosefrom
thetropicsofSouthAmericathanthosefromtemperateregionsofEurope.
Further,SouthAmericanfossils,thoughdifferentfrommodernspecies,moreclosely
resembledmodernspeciesfromSouthAmericathanthosefromEurope.

WhileontheBeagle,DarwinreadLyellsPrinciplesofGeology.
Heexperiencedgeologicalchangefirsthandwhenaviolentearthquakerockedthecoast
ofChile,causingthecoastlinetorisebyseveralfeet.
HefoundfossilsofoceanorganismshighintheAndesandinferredthattherocks
containingthefossilshadbeenraisedtherebyaseriesofsimilarearthquakes.
TheseobservationsreinforcedDarwinsacceptanceofLyellsideasandledhimtodoubt
thetraditionalviewofayoungandstaticEarth.

Darwinsinterestinthegeographicdistributionofspecieswasfurtherstimulatedbythe
BeaglesvisittotheGalapagos,agroupofyoungvolcanicislands900kmwestoftheSouth
Americancoast.
Darwinwasfascinatedbytheunusualorganismsfoundthere.
AfterhisreturntoEngland,Darwinnotedthatwhilemostoftheanimalspeciesonthe
Galapagoslivednowhereelse,theyresembledspecieslivingontheSouthAmerican
mainland.
Hehypothesizedthattheislandshadbeencolonizedbyplantsandanimalsfromthe
mainlandthathadsubsequentlydiversifiedonthedifferentislands.
AfterhisreturntoGreatBritainin1836,Darwinbegantoperceivethattheoriginofnew
speciesandadaptationofspeciestotheirenvironmentwerecloselyrelatedprocesses.
Forexample,cleardifferencesinthebeaksamongthe13speciesoffinchesthatDarwin
collectedintheGalapagosareadaptationstothespecificfoodsavailableontheirhome
islands.
Bytheearly1840s,Darwinhaddevelopedthemajorfeaturesofhistheoryofnatural
selectionasthemechanismforevolution.
In1844,hewrotealongessayontheoriginofspeciesandnaturalselection,buthewas
reluctanttopublishandcontinuedtocompileevidencetosupporthistheory.
InJune1858,AlfredRusselWallace(18231913),ayoungnaturalistworkingintheEast
Indies,sentDarwinamanuscriptcontainingatheoryofnaturalselectionessentiallyidentical
toDarwins.
Laterthatyear,bothWallacespaperandextractsofDarwinsessaywerepresentedto
theLinnaeanSocietyofLondon.
IG Lecture Outline 22-4

DarwinquicklyfinishedTheOriginofSpeciesandpublisheditthenextyear.

WhilebothDarwinandWallacedevelopedsimilarideasindependently,thetheoryof
evolutionbynaturalselectionisattributedtoDarwinbecausehedevelopedhisideasearlier
andsupportedthetheorymuchmoreextensively.
ThetheoryofevolutionbynaturalselectionwaspresentedinTheOriginofSpecieswith
immaculatelogicandanavalancheofsupportingevidence.
Withinadecade,TheOriginofSpecies

hadconvincedmostbiologiststhatbiological
diversitywastheproductofevolution.
2.TheOriginofSpeciesdevelopedtwomainideas:thatevolutionexplainslifesunityand
diversityandthatnaturalselectionisthemechanismofadaptiveevolution.

DarwinscarcelyusedthewordevolutioninTheOriginofSpecies.
Insteadheusedthephrasedescentwithmodification.
Allorganismsarerelatedthroughdescentfromacommonancestorthatlivedinthe
remotepast.
Overevolutionarytime,thedescendentsofthatcommonancestorhaveaccumulated
diversemodifications,oradaptations,

thatallowthemtosurviveandreproducein
specifichabitats.

Viewedfromtheperspectiveofdescentwithmodification,thehistoryoflifeislikeatree
withmultiplebranchesfromacommontrunk.
Closelyrelatedspecies,thetwigsonacommonbranchofthetree,sharedthesamelineof
descentuntiltheirrecentdivergencefromacommonancestor.
Linnaeusrecognizedthatsomeorganismsresembleeachothermorecloselythanothers,
buthedidnotexplainthesesimilaritiesbyevolution.
However,histaxonomicschemefitwellwithDarwinstheory.
ToDarwin,theLinnaeanhierarchyreflectedthebranchinghistoryofthetreeoflife.
Organismsatvarioustaxonomiclevelsareunitedthroughdescentfromcommon
ancestors.

Howdoesnaturalselectionwork,andhowdoesitexplainadaptation?

EvolutionarybiologistErnstMayrhasdissectedthelogicofDarwinstheoryintothree
inferencesbasedonfiveobservations.
Observation#1:Allspecieshavesuchgreatpotentialfertilitythattheirpopulationsize
wouldincreaseexponentiallyifallindividualsthatarebornreproducedsuccessfully.
Observation#2:Populationstendtoremainstableinsize,exceptforseasonal
fluctuations.
Observation#3:Environmentalresourcesarelimited.
Inference#1:Productionofmoreindividualsthantheenvironmentcansupportleads
toastruggleforexistenceamongtheindividualsofapopulation,withonlyafraction
oftheoffspringsurvivingeachgeneration.
Observation#4:Individualsofapopulationvaryextensivelyintheircharacteristics;no
twoindividualsareexactlyalike.
Observation#5:Muchofthisvariationisheritable.
Inference#2:Survivalinthestruggleforexistenceisnotrandom,butdependsinpart
oninheritedtraits.Thoseindividualswhoseinheritedtraitsarebestsuitedfor
survivalandreproductionintheirenvironmentarelikelytoleavemoreoffspringthan
lessfitindividuals.
IG Lecture Outline 22-5

Inference#3:Thisunequalabilityofindividualstosurviveandreproducewillleadto
agradualchangeinapopulation,withfavorablecharacteristicsaccumulatingover
generations.

A1798essayonhumanpopulationbyThomasMalthusheavilyinfluencedDarwins
viewsonoverreproduction.
Malthuscontendedthatmuchhumansufferingdisease,famine,homelessness,war
wastheinescapableconsequenceofthepotentialforhumanpopulationstoincreasefaster
thanfoodsuppliesandotherresources.

Thecapacitytooverproduceseemstobeacharacteristicofallspecies.

Onlyatinyfractionofoffspringproducedcompletetheirdevelopmentandreproduce
successfullytoleaveoffspringoftheirown.
Ineachgeneration,environmentalfactorsfilterheritablevariations,favoringsomeover
others.
Differentialreproductivesuccesswherebyorganismswithtraitsfavoredbythe
environmentproducemoreoffspringthandoorganismswithoutthosetraitsresultsin
thefavoredtraitsbeingdisproportionatelyrepresentedinthenextgeneration.
Thisincreasingfrequencyofthefavoredtraitsinapopulationisevolutionarychange.
Darwinsviewsontheroleofenvironmentalfactorsinthescreeningofheritable
variationwereheavilyinfluencedbyartificialselection.
Humanshavemodifiedavarietyofdomesticatedplantsandanimalsovermany
generationsbyselectingindividualswiththedesiredtraitsasbreedingstock.
Ifartificialselectioncanachievesomuchchangeinarelativelyshortperiodoftime,
Darwinreasoned,thennaturalselectionshouldbecapableofconsiderablemodification
ofspeciesoverthousandsofgenerations.

Darwinsmainideascanbesummarizedinthreepoints.
Naturalselectionisdifferentialsuccessinreproduction(unequalabilityofindividualsto
surviveandreproduce)thatresultsfromindividualsthatvaryinheritabletraitsandtheir
environment.
Theproductofnaturalselectionistheincreasingadaptationoforganismstotheir
environment.
Ifanenvironmentchangesovertime,orifindividualsofaspeciesmovetoanew
environment,naturalselectionmayresultinadaptationtothenewconditions,sometimes
givingrisetoanewspeciesintheprocess.

Threeimportantpointsneedtobeemphasizedaboutevolutionthroughnaturalselection.
1. Althoughnaturalselectionoccursthroughinteractionsbetweenindividualorganismsand
theirenvironment,individualsdonotevolve.Apopulation(agroupofinterbreeding
individualsofasinglespeciesthatshareacommongeographicarea)isthesmallestgroup
thatcanevolve.Evolutionarychangeismeasuredaschangesinrelativeproportionsof
heritabletraitsinapopulationoversuccessivegenerations.
2. Naturalselectioncanactonlyonheritabletraits,traitsthatarepassedfromorganismsto
theiroffspring.Characteristicsacquiredbyanorganismduringitslifetimemayenhance
itssurvivalandreproductivesuccess,butthereisnoevidencethatsuchcharacteristics
canbeinheritedbyoffspring.
3. Environmentalfactorsvaryfromplacetoplaceandfromtimetotime.Atraitthatis
favorableinoneenvironmentmaybeuselessorevendetrimentalinanotherenvironment.
Darwinenvisionedthediversityoflifeasevolvingbyagradualaccumulationofminute
changesthroughtheactionsofnaturalselectionoperatingovervastspansoftime.
IG Lecture Outline 22-6

3.Darwinstheoryexplainsawiderangeofobservations.
Thepowerofevolutionbynaturalselectionasaunifyingtheoryisitsversatilityasa
naturalexplanationfordiversedatafrommanyfieldsofbiology.
Wewillconsidertwoexamplesofnaturalselectionasamechanismofevolutionin
populations.

Ourfirstexampleconcernsdifferentialpredationandguppypopulations.

Guppies(Poeciliareticulata)liveinthewildinpoolsintheAripoRiversystemin
Trinidad.
JohnEndlerandDavidReznickhavebeenstudyingthesesmallfishformorethana
decade.
Theresearchersobservedsignificantdifferencesbetweenpopulationsofguppiesthatlive
indifferentpoolsintheriversystem.
Populationsvariedintheaverageageandsizeofsexualmaturity.
Thesevariationswerecorrelatedtothetypeofpredatorpresentineachpool.
Insomepools,themainpredatoristhesmallkillifish,whicheatsjuvenileguppies.
Inotherpools,themajorpredatoristhelargepikecichlid,whicheatsadultguppies.
Guppiesinpopulationspreyedonbypikecichlidsbeginreproducingatayoungerage
andaresmalleratmaturitythanguppiesinpopulationspreyedonbykillifish.
Totestwhetherthesedifferencesareduetonaturalselection,ReznickandEndler
introducedguppiesfrompikecichlidlocationstonewpoolsthatcontainedkillifishbutno
guppies.
Afterelevenyears,thetransplantedguppieswere,onaverage,14%heavieratmaturity
thanthenontransplantedpopulations.
Theiraverageageatmaturityhadalsoincreased.
Theseresultssupportthehypothesisthatnaturalselectioncausedthechangesinthe
transplantedpopulation.
Becausepikecichlidspreymainlyonreproductivelymatureadults,thechancethata
guppywillsurvivetoreproduceseveraltimesislow.
Theguppieswiththegreatestreproductivesuccessinpondswithpikecichlidpredators
arethosethatmatureatayoungageandsmallsize,enablingthemtoproduceatleastone
broodbeforegrowingtoasizepreferredbypikecichlids.
Inpondswithkillifishpredators,guppiesthatsurviveearlypredationcangrowslowly
andproducemanybroodsofyoung.
AsecondexampleofongoingnaturalselectionistheevolutionofdrugresistantHIV
(humanimmunodeficiencyvirus).
ResearchershavedevelopednumerousdrugstocombatHIV,butusingthesemedications
selectsforvirusesresistanttothedrugs.
Afewdrugresistantvirusesmaybepresentbychanceatthebeginningoftreatment.
Thedrugresistantpathogensaremorelikelytosurvivetreatmentandpassonthegenes
thatenablethemtoresistthedrugtotheiroffspring.
Asaresult,thefrequencyofdrugresistanceintheviralpopulationrapidlyincreases.
Scientistsdesignedthedrug3TCtointerferewithreversetranscriptase,theenzymethat
HIVusestocopyitsRNAgenomeintotheDNAofthehostcell.

IG Lecture Outline 22-7

Because3TCissimilarinshapetothecytosinenucleotideofDNA,HIVsreverse
transcriptaseincorporates3TCintoitsgrowingDNAchaininsteadofcytosine.Thiserror
terminateselongationofDNAandthuspreventsHIVreproduction.
3TCresistantvarietiesofHIVhaveaformofreversetranscriptasethatcandiscriminate
betweencytosineand3TC.
Theseviruseshavenoadvantageintheabsenceof3TC.Infact,theyreplicatemore
slowlythanviruseswithnormalreversetranscriptase.
Once3TCisaddedtotheirenvironment,itbecomesapowerfulselectiveagent,
favoringreproductionofresistantindividuals.

TheexamplesoftheguppiesandHIVhighlighttwoimportantpointsaboutnatural
selection.
First,naturalselectionisaneditingmechanism,notacreativeforce.Itcanonlyacton
existingvariationinthepopulation;itcannotcreatefavorabletraits.
Second,naturalselectionfavorstraitsthatincreasefitnessinthecurrent,local
environment.Whatisadaptiveinonesituationisnotadaptiveinanother.
Forexample,guppiesthatmatureatanearlyageandsmallsizeareatanadvantagein
apoolwithpikecichlids,butatadisadvantageinapoolwithkillifish.
Intheabsenceof3TC,HIVwiththemodifiedformofreversetranscriptasegrows
moreslowlythanHIVwithnormalreversetranscriptase.
4.Evidenceofevolutionpervadesbiology.
Inthecasesdescribed,naturalselectionbroughtaboutchangerapidlyenoughthatit
couldbeobserveddirectly.
Darwinstheoryalsoprovidesacohesiveexplanationforobservationsinthefieldsof
anatomy,embryology,molecularbiology,biogeography,andpaleontology.
Descentwithmodificationcanexplainwhycertaintraitsinrelatedspecieshavean
underlyingsimilarityeveniftheyhaveverydifferentfunctions.

Similarityincharacteristictraitsfromcommonancestryisknownashomology.
Forexample,theforelimbsofhuman,cats,whales,andbatssharethesameskeletal
elements,eventhoughtheappendageshaveverydifferentfunctions.
Theseforelimbsarehomologousstructures

thatrepresentvariationsontheancestral
tetrapodforelimb.

Homologiesthatarenotobviousinadultorganismsmaybecomeevidentwhenwelook
atembryonicdevelopment.
Forexample,allvertebrateembryoshavestructurescalledpharyngealpouchesintheir
throatatsomestageintheirdevelopment.
Theseembryonicstructuresdevelopintoverydifferent,butstillhomologous,adult
structures,suchasthegillsoffishortheeustaciantubesthatconnectthemiddleearwith
thethroatinmammals.
Someofthemostinterestinghomologousstructuresarevestigialorgans,

structuresthat
havemarginal,ifany,importancetoalivingorganism,butwhichhadimportantfunctionsin
theorganismsancestors.
Forexample,theskeletonsofsomesnakesandoffossilwhalesretainvestigesofthe
pelvisandlegbonesofwalkingancestors.
Comparativeanatomyconfirmsthatevolutionisaremodelingprocess,analterationof
existingstructures.
IG Lecture Outline 22-8

Becauseevolutioncanonlymodifyexistingstructuresandfunctions,itmayproduce
structuresthatarelessthanperfect.
Forexample,thebackandkneeproblemsofbipedalhumansareanunsurprisingoutcome
ofadaptingstructuresoriginallyevolvedtosupportfourleggedmammals.
Similaritiesamongorganismscanalsobeseenatthemolecularlevel.
Forexample,allspeciesoflifehavethesamebasicgeneticmachineryofRNAandDNA,
andthegeneticcodeisessentiallyuniversal.
Theubiquityofthegeneticcodeprovidesevidenceofasingleoriginoflife.
Itislikelythatthelanguageofthegeneticcodehasbeenpassedalongthroughallthe
branchesofthetreeoflifeeversinceitsinceptioninanearlylifeform.
Homologiesmirrorthetaxonomichierarchyofthetreeoflife.
Somehomologies,suchasthegeneticcode,aresharedbyalllivingthingsbecausethey
aroseinthedeepancestralpast.
Otherhomologiesthatevolvedmorerecentlyaresharedonlybysmallerbranchesofthe
treeoflife.
Forexample,alltetrapods(amphibians,reptiles,birds,andmammals)sharethesame
fivedigitlimbstructure.
Thushomologiesarefoundinanestedpattern,withalllifesharingthedeepestlayerand
eachsmallergroupaddingnewhomologiestothosetheysharewiththelargergroup.
Thishierarchicalpatternofhomologyisexactlywhatwewouldexpectiflifeevolvedand
diversifiedfromacommonancestor.

Anatomicalresemblancesamongspeciesaregenerallyreflectedintheirgenes(DNA)
andgeneproducts(proteins).
Ifhierarchiesofhomologyreflectevolutionaryhistory,thenweshouldexpecttofind
similarpatternswhetherwearecomparingmoleculesorbones.
Differentkindsofhomologieswillcoincidebecausetheyhavefollowedthesame
branchingpatternthroughevolutionaryhistory.
Thegeographicaldistributionofspeciesbiogeography

firstsuggestedevolutionto
Darwin.
Speciestendtobemorecloselyrelatedtootherspeciesfromthesameareathantoother
specieswiththesamewayoflifethatliveindifferentareas.
ConsiderAustralia,hometoauniquegroupofmarsupialmammals,whichcomplete
theirdevelopmentinanexternalpouch.
Somemarsupialmammalssuperficiallyresembleeutherianmammals(which
completetheirdevelopmentintheuterus)fromothercontinents.

Forexample,theAustraliansugargliderandNorthAmericanflying
squirrelareadaptedtothesamemodeoflifeandlooksomewhatsimilar.

However,thesugarglidersharesmorecharacteristicswithother
Australianmarsupialsthanwiththeflyingsquirrel.

Theresemblancebetweenthetwoglidersisanexampleofconvergent
evolution.

Islandsandislandarchipelagoshaveprovidedstrongevidenceofevolution.
Islandsgenerallyhavemanyspeciesofplantsandanimalsthatareendemic,

found
nowhereelseintheworld.

IG Lecture Outline 22-9

 AsDarwinobservedwhenhereassessedhiscollectionsfromtheBeaglesvoyage,these
endemicspeciesaretypicallymorecloselyrelatedtospecieslivingonthenearestmainland
(despitedifferentenvironments)thantospeciesfromotherislandgroups.
Inislandchains,orarchipelagos,individualislandsmayhavedifferent,butrelated,
species.Thefirstmainlandinvadersreachedoneislandandthenevolvedintoseveralnew
speciesastheycolonizedotherislandsinthearchipelago.
Severalwellinvestigatedexamplesofthisphenomenonincludethediversificationof
finchesontheGalapagosIslandsandfruitflies(Drosophila)ontheHawaiian
Archipelago.
Thesuccessionoffossilformsisconsistentwithwhatisknownfromothertypesof
evidenceaboutthemajorbranchesofdescentinthetreeoflife.
Forexample,considerableevidencesuggeststhatprokaryotesaretheancestorsofalllife
andshouldprecedealleukaryotesinthefossilrecord.Infact,theoldestknownfossilsare
prokaryotes.
Fossilfishespredateallothervertebrates,withamphibiansnext,followedbyreptiles,
thenmammalsandbirds.
Thisisconsistentwiththehistoryofvertebratedescentsupportedbymanyothertypesof
evidence.
TheDarwinianviewoflifealsopredictsthatevolutionarytransitionsshouldleavesigns
inthefossilrecord.
Paleontologistshavediscoveredfossilsofmanysuchtransitionalformsthatlinkancient
organismstomodernspecies.
Forexample,fossilevidencedocumentstheoriginofbirdsfromonebranchofdinosaurs.
Recentdiscoveriesincludefossilizedwhalesthatlinktheseaquaticmammalstotheir
terrestrialancestors.
5.WhatistheoreticalabouttheDarwinianviewoflife?

SomepeopledismisstheDarwinianviewasjustatheory.
Aswehaveseen,Darwinsexplanationmakessenseoflargeamountsofdata.
Theeffectsofnaturalselectioncanbeobservedinnature.

Whatistheoreticalaboutevolution?
Thetermtheoryhasaverydifferentmeaninginsciencethanineverydayuse.
Thewordtheoryincolloquialuseisclosertotheconceptofahypothesisinscience.

Inscience,atheoryismorecomprehensivethanahypothesis,accountingformany
observationsanddataandattemptingtoexplainandintegrateagreatvarietyofphenomena.
Aunifyingtheorydoesnotbecomewidelyacceptedunlessitspredictionsstandupto
thoroughandcontinualtestingbyexperimentsandadditionalobservation.
Thathascertainlybeenthecasewiththetheoryofevolutionbynaturalselection.

Scientistscontinuetotestthistheory.
Forexample,manyevolutionarybiologistsnowquestionwhethernaturalselectionisthe
onlymechanismresponsibleforevolutionaryhistory.
Otherfactorsmayhaveplayedanimportantrole,particularlyintheevolutionofgenes
andproteins.

Byattributingthediversityoflifetonaturalcauses,Darwingavebiologyasound
scientificbasis.
IG Lecture Outline 22-10

AsDarwinsaid,Thereisgrandeurinthisviewoflife.

IG Lecture Outline 22-11