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Evolution

Evolution

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Evolution and clinate change
Evolution and clinate change

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06/14/2009

 
http://www.jstor.org
Evolutionary Responses to Changing ClimateAuthor(s): Margaret B. Davis, Ruth G. Shaw, Julie R. EttersonSource:
Ecology,
Vol. 86, No. 7, (Jul., 2005), pp. 1704-1714Published by: Ecological Society of AmericaStable URL:
Accessed: 12/08/2008 02:54
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Ecology,86(7),2005,pp.1704-1714
?2005bytheEcological Societyof America
EVOLUTIONARYRESPONSESTOCHANGINGCLIMATE
MARGARET B.DAVIS,''3RUTH G.SHAW,'ANDJULIER. ETTERSON2
'DepartmentofEcology,EvolutionandBehavior,University ofMinnesota,St.Paul,Minnesota 55108 USA2Department ofBiology,University ofMinnesota,Duluth,Minnesota55812 USA
Abstract. Untilnow,Quaternarypaleoecologistshaveregardedevolution as a slowprocessrelativeto climatechange,predictingthat theprimarybioticresponsetochangingclimate is notadaptation,butinstead(1)persistenceinsituifchangingclimate remainswithinthespecies'tolerancelimits,(2)rangeshifts(migration)toregionswhere climateiscurrentlywithin thespecies'tolerancelimits,or(3)extinction. Weargueherethatallthree ofthese outcomesinvolveevolutionary processes.Genetic differentiationwithinspeciesisubiquitous, commonlyviaadaptationofpopulationstodifferingenvironmentalconditions. Detectableadaptivedivergenceevolveson atimescalecomparabletochangeinclimate,withindecadesfor herbaceousplantspecies,andwithincenturies or millenniaforlonger-livedtrees,implyingthatbiologically significant evolutionary responsecanac-company temporalchangeinclimate.Models andempiricalstudiessuggestthat thespeedwithwhich apopulation adaptstoachangingenvironment affects invasion rate ofnewhabitat and thusmigrationrate,populationgrowthrateandthusprobabilityofextinction,andgrowthandmortalityofindividualplantsandthusproductivityofregional vegetation.Recent models andexperimentsinvestigatethestabilityofspeciestolerancelimits,theinfluence ofenvironmentalgradientsonmarginal populations,and theinterplayofde-mography, geneflow,mutationrate,and othergenetic processeson therateofadaptationtochangedenvironments. Newtechniquesenableecologiststo documentadaptationtochangingconditionsdirectlybyresurrectingancientpopulationsfrompropagulesburiedindecades-old sediment.Improvedtaxonomicresolution frommorphologicalstudiesofmacrofossilsand DNArecovered frompollen grainsandmacroremainsprovidesadditionalinformation onrangeshifts,changesinpopulationsizes,andextinctions.Collaborationbetweenpaleoecologistsandevolutionarybiologistscan refineinterpretationsofpaleo-records,andimprove predictionsofbioticresponsetoanticipatedclimatechange.
Keywords:adaptation;climatechange; evolutionaryconstraints;Quaternarypaleoecology;range shifts;tolerance limits.
INTRODUCTION
Adaptationtochangesin theenvironment isan im-portantandnearlyuniversalaspectof bioticresponsetoclimatechange.Weargueherethatadaptivere-sponsesaffectspecies persistence,migrationrate,andforestproductivity.Thus morecompleteunderstandingofadaptiveresponsesto climatemust beincorporatedintointerpretationsofQuaternary paleorecords.Fur-thermore,fossil recordscanprovidetests ofevolu-tionary hypotheses,enhancingunderstandingoftheevolutionaryconsequencesof climatechange.Itisremarkable thatpaleoecologistsreconstructinglate-Quaternaryenvironments so seldomdiscussthepossibilityofadaptationtochangingclimate.TheyhavetendedtoadopttheperspectiveofDarwin,whoemphasizedthat vastperiodsofgeologictime allowgreatopportunityfor theaccumulationofmanyslightdifferencesamong organismsthat,collectively,can re-
Manuscriptreceived24November2003;accepted16June2004;finalversionreceived4October 2004.CorrespondingEd-itor(adhoc):L.J.Graumlich.Forreprintsof thisSpecialFeature,seefootnote1,p.1667.3E-mail:mbdavis@ecology.umn.edu
suitinmajordifferencesinbiologicalform.Accord-ingly,a fundamentalassumptionofpaleoecologyhasbeen thatthe rate of evolution is far slower than therateofclimatechange(Bennett1997,Webb1997,Jack-son2000),withmeaningfulevolutionary adaptations,suchas theoriginationoftaxa,occurringon thetime-scaleofmillionsofyears.Climatechangesmuchmorerapidly,becausesuperimposedonglacial cycles100000yearsinlengthare variations onmillennial,centennial,and evendecadal time scales.Forexample,duringthe most recentglacial-interglacialtransition,atime ofrapid warming,therewas a brief reversallastingseveral centuries.Atthis timepaleorecordsfromthenorth Atlanticregionrecordatemperaturedropofsev-eraldegreesC within decades(Huntleyetal.1997).Paleoecologistshavearguedthatsuchrapidenviron-mentalchangesoverwhelmevolutionary processes,causingextinctionexceptwhereclimateremainswithinpreexistingtolerance limits for aspecies(Bennett1997).Contributingto this viewhas beenthesparse-ness of the record of newplant species duringtheQua-ternaryand theapparentlimitations offossils,espe-ciallypollen,torecordtraits involved inadaptationtoclimate,such asphenologyandphysiology.Further-1704
 
PALEOPERSPECTIVESNECOLOGYmore,latitudinalrangeshiftsof treetaxa,welldocu-mented inQuaternarysediments,arecompatiblewiththeviewthatspecies persistonlywithin the environ-ments towhichtheyarealready adapted,even asthoseenvironments shift inspace(Prenticeet al.1991,Jack-son2000).Moreover,paleoclimatereconstructionsrelyon thepremisethateachspecieshas auniqueset of limits inits toleranceto variousaspectsof theenvironment,de-finingamultidimensionalecologicalniche(Jackson2000).Paleoecologistshave focused onphysicalnichedimensions,determiningforparticularspeciesthe re-alized limitsofphysiologicaltolerances to factors suchasminimumtemperatureorgrowing degree days.Whenspecies-specifictolerances areplottedonamapofmodernclimate,theyquite accuratelydelimit thepresentgeographicaldistribution of thespeciesinques-tion(Petersonet al.1999).Paleoclimate reconstructionemploysthese environmental attributesofspeciestoinferthedistribution of climateatan ancienttimefromthedistribution and abundance of fossils.Thus,paleo-climatereconstructiondependson theassumptionthatspeciestolerancelimits remainstableintime,thatis,evolutionarilyinert.Inpractice,acertain amount ofchangemightnot be noticed because reconstructionsare often ata coarsegeographicalscale(Prenticeet al.1991),withbroadconfidence intervalssurroundingtemperatureandprecipitationestimates.Fossilrecordsusedforvalidation are oftensparse, especiallyforoldertimehorizons.Nevertheless,recentreviewsregard rap-id evolutionas theexception,affectingonly"somelocalpopulationsof aspecies...whileothersundergoextinctionordispersetonewlysuitablesites"(Jackson2000:294).Giventhiscontext,discussionsofbioticresponsetofutureclimateemphasize predictedrangeshifts,focusingon seeddispersaland establishment aspotentiallylimitingprocesses (e.g.,Clarket al.1998).Hereweconsider evolution in relation toclimate anddiscuss itsrelevance toprocessespaleoecologistscon-siderimportant,includingpopulation persistenceinsitu,rangeshifts,andextinction. Wereview theevi-dence foradaptationofplantpopulationstospatialvar-iation inenvironment anddiscuss thepotentialfor evo-lutionaryresponsetotemporalvariation.Weemphasizethatclimatechangeon varioustime scalesimposesselectionregimesthatmaylead toadaptivechangesinplantsandanimals,whetheror not theirrangesshift(DavisandShaw2001).Thedegreeofadaptationde-pendsontheinterplayofnatural selection withotherevolutionaryprocesses,such asgeneflow,geneticdrift,andmutation,and also withdemography.Alagin ad-aptationimpliesreduction ingrowthandsurvival ofindividualplantsas well asin the overallproductivityofregionalvegetation(Rehfeldtet al.1999,2002).Therateofadaptationinfluences the rate atwhichpopu-lationsinvadenewlyavailablehabitat(Garcia-RamosandRodriguez2002),andalso theprobabilitythatpop-ulationswilldie out withintheirpresent ranges(Peaseet al.1989,BurgerandLynch1995).Thus,adaptiveevolutionaffects all of theprimary responsestochang-ingclimatepredictedby paleoecologists-"tolerance,migration,or extinction"(Jackson2000:294).We re-interpret examplesfrom theQuaternaryrecordthatdemonstratehowevolutionarymodels canexpandun-derstanding.Last,wereview recent contributionsbyQuaternary paleoecologiststhat utilizepaleorecordstodemonstrateadaptationsto environmentalchange.
GENETIC CHANGE IN PLANTSDURING THEQUATERNARY
For animalphyla,particularlymammals,bothspe-ciation events andextinctions are well documenteddur-ingthepasttwo millionyears.Incontrast,theQua-ternaryfossilrecorddocuments fewexamplesofnewspeciesof vascularplants(Comesand Kadereit1998),reinforcingtheimpressionthat,particularlyintheseorganisms,evolutionis aslowprocess.Speciationis,however,onlyoneaspectofevolutionary change.With-inspecies, geneticdifferentiationamongpopulationsatteststo theubiquityandrapidityofevolutionarychange.Spatial substructuringofpopulationsis evidencedbyvariation atputativelyneutralgeneticmarkerloci,re-flectingcurrentmatingpatternsand/orpreviousiso-lation ofpopulations.Phylogeographersuse suchmark-ers as abasisforinferringlocationsofrefugesduringthe lastglaciation,andpathwaysofmigrationsincethen(e.g.,Cruzan andTempleton2000,Hewitt2000).Fossilevidencehasvaluablyinformedsuch studies.Forexample,allozymeandmitochondrialDNA(mtDNA)divergencehas been calibratedagainsttheactual timesince isolation of treepopulationsintheAmerican Southwest.Timeof isolation wasdeterminedfrommigrationhistoriesdocumentedbymacroremainspreservedinpackratmiddens(Hamricket al.1994).Allozymesin a seriesofpopulationsoflodgepole pine(Pinus contorta),theagesof whichweredeterminedfromradiocarbon-datedfossil-pollenrecords,suggestaprogressiveloss of allelesinthe courseof northwardmigrationin westernCanadaduringthe Holocene(Cwynarand MacDonald1987).Reducedallelic di-versityawayfromputative glacialrefugiahas beendocumentedfor severalplantspecies,whilePinuspum-ila shows reductionsindiversityat some loci andin-creaseddiversityat others(Taniet al.1996).Adaptativedifferentiationamongpopulationswithinaspeciesis documentedbyclinal variationinphysi-ological, phenological,and fitnesstraitsin relationtolatitudinal orelevationalgradientsinclimate. Suchvar-iationhas beenshown formany species, beginningwiththeclassicstudies of Turesson(1922)andClausenetal.(1940)andcontinuingwithrecentpaperstoonu-merousto list here.Thedistributionof climate-sensi-tivetraitswithinspeciesis studiedincommon-garden(provenance) experiments,in whichplantsfrom dif-ferentgeographicallocationsaregrowntogetherat aJuly20051705

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