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Basal(phylogenetics)Wikipedia,thefreeencyclopedia

Basal(phylogenetics)
FromWikipedia,thefreeencyclopedia

Inphylogenetics,basalisthedirectionofthebase(orroot)ofarootedphylogenetictreeorcladogram.CladeCmaybe
describedasbasalwithinalargercladeDifitsrootisdirectlylinked(adjacent)totherootofD.IfCistheonlycladeof
agiventaxonomicrankthatisbasalwithinD,CmaybedescribedasthebasaltaxonofthatrankwithinD.Whilethere
mustalwaysbetwoormoreequallybasalcladessproutingfromtherootofeverycladogram,thosecladesmaydiffer
widelyinrank[n1]and/orspeciesdiversity.Greaterdiversificationmaybeassociatedwithmoreevolutionaryinnovation,
butancestralcharactersshouldnotbeimputedtothemembersofalessspeciesrichbasalcladewithoutadditional
evidence,astherecanbenoassurancesuchanassumptionisvalid.[1][2][3]
Ingeneral,cladeAismorebasalthancladeBifBisasubgroupofthesistergroupofA.Withinlargegroups,"basal"
maybeusedmorelooselytomean'closertotherootthanthegreatmajorityof',andinthiscontextterminologysuchas
"verybasal"mayarise.

Contents
1
2
3
4
5
6

Examples
Criticism
Relevancetobiogeographichistory
Notes
References
Externallinks

Examples
Abasalgroupformsasistergrouptotherestofthelargerclade,suchasinthefollowingexample:
Basalgroup
Nonbasalgroup#1

Nonbasalgroup#2

Nonbasalgroup#3

Itisassumedinthisexamplethattheterminalbranchesofthecladogramdepictalltheextanttaxaofagivenrankwithin
thecladeotherwise,thediagramcouldbehighlydeceptive.
Thetermbasalcanbecorrectlyappliedtocladesoforganisms,butnottolineagesortoindividualtraitspossessedby
theorganismsalthoughitmaybemisusedinthesewaysintechnicalliterature.While"basal"appliestoclades,
charactersortraitsareusuallyconsideredderivediftheyareabsentinabasalgroup,butpresentinothergroups.This
assumptiononlyholdstrueifthebasalgroupisagoodanalogyforthelastcommonancestorofthegroup."Ancestral"
or"plesiomorphic"(butnot"basal",seeCriticism)arepreferredto"primitive",whichmaycarryfalseconnotationsof
inferiorityoralackofcomplexity.
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ThefloweringplantfamilyAmborellaceae,restrictedtoNewCaledoniainthesouthwesternPacific,[n2]isabasalclade
ofextantangiosperms,containingthemostbasalspecies,genus,familyandorderwithinthegroup(outofatotalof
about250,000angiospermspecies).WhilethetraitsofAmborellatrichopodaareregardedasprovidingsignificant
insightintotheevolutionoffloweringplants,theyareamixofplesiomorphic(archaic)andapomorphic(derived)
featuresthathaveonlybeensortedoutviacomparisonwithotherangiospermsandtheirpositionswithinthe
phylogenetictree(thefossilrecordcouldpotentiallyalsobehelpfulinthisrespect,butisabsentinthiscase).[4]
Amborellales
Monocots

Magnoliids
Eudicots

WithintheprimatefamilyHominidae(greatapes),gorillas(easternandwestern)areasistergrouptocommon
chimpanzees,bonobosandhumans.Thesefivespeciesformaclade,thesubfamilyHomininae(Africanapes),ofwhich
Gorillaisthebasalgenus.However,iftheanalysisisnotrestrictedtogenera,theHomoplusPancladeisalsobasal.
ThissemanticambiguityisdiscussedintheCriticismsection.
Humans(Homosapiens)
Bonobos(Panpaniscus)

Commonchimpanzees(Pantroglodytes)

Easterngorillas(Gorillaberingei)

Westerngorillas(Gorillagorilla)

Moreover,orangutansareasistergrouptoHomininaeandarethebasalgenusinthefamilyasawhole.
Orangutans(Pongospp.)
Humans(Homosapiens)

Chimpanzees(Panspp.)

Gorillas(Gorillaspp.)

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SubfamiliesHomininaeandPonginaearebothbasalwithinHominidae,butgiventhattherearenononbasalsubfamilies
inthecladogramitisunlikelythetermwouldbeappliedtoeither.Ingeneral,astatementtotheeffectthatonegroup
(e.g.,orangutans)isbasal,orbranchesofffirst,withinanothergroup(e.g.,Hominidae)maynotmakesenseunlessthe
appropriatetaxonomiclevel(s)(genus,inthiscase)isspecified.Ifthatlevelcannotbespecified(i.e.,ifthecladein
questionisunranked)amoredetaileddescriptionoftherelevantsistergroupsmaybeneeded.
Inthisexample,orangutansdifferfromtheothergeneraintheirEurasianrange.Thisfactplustheirbasalstatusprovides
ahintthatthemostrecentcommonancestorofextantgreatapesmayhavebeenEurasian(seebelow),asuggestionthat
isconsistentwithotherevidence.[5]

Criticism
Despitetheubiquityoftheusageoftheterm,somesystematistsbelieveitsapplicationtoextantgroupsisunnecessary
andmisleading.[6]Thisisbecauseapairofsisterbranchesofaphylogenetictreeareequallyranked(intermsofthe
highesttaxonomicranksoftherespectiveclades)andcanbefreelyrotatedabouttheirlastcommonnode.Theterm
tendstobeappliedwhenonebranch(theonedeemed"basal")islessdiversethananotherbranch(thisbeingthe
situationinwhichonewouldexpecttofindbasaltaxaoflowerminimumrank).
Potentiallymisleadinguse

Descriptionassistergroup

taxonA(nonbasal)

taxonA

taxonB(nonbasal)

taxonC(nonbasal)

taxonD(basalposition)

TaxonDisconsideredbasalwithinthegroupdiagrammed.
However,cladeA+B+Cisalsobasal. [n3]

taxonB

taxonC

taxonD(sistertotheremainingtaxa)

TaxonDcanalsobedescribedasthesistergroupofA,Band
C,i.e.,theremainingtaxawithintheingroup. [6]

Thissituationraisesanequivocalusageforthetermbasalinthatitalsoreferstothedirectionoftherootofthetree.As
therootrepresentsahypotheticalancestor,thisconsequentlymaycarrymisleadingconnotationsthatthesistergroupof
amorespeciesrichcladedisplaysancestralfeatures.[3]

Relevancetobiogeographichistory
Giventhatthedeepestphylogeneticsplitinawidelydispersedgroupislikelytohaveoccurredinitsregionoforigin,
identificationofthemostbasalclade(s)insuchagroupcanprovidevaluableinsightintoitsbiogeographichistory.For
example,extantaustralidelphianmarsupialsconstituteabout238speciesinAustralasiaandonespecies(themonitodel
monte)inSouthAmerica.Thefactthatthemonitodelmonteoccupiesabasalposition(themostbasalspecies,genus,
familyandorder)inthesuperorderisanimportantcluethatitsoriginwasinSouthAmerica.[7][n4]Also,whilethebat
superfamilyNoctilionoideahasover200speciesintheNeotropics,twoinNewZealand,andtwoinMadagascar,the
basalpositionoftheMalagasygenusandfamily[8]suggests(incombinationwiththefossilrecord)thatthesuperfamily
originatedinAfrica.[9]Similarly,whilecoralsnakescomprise16speciesinAsiaandover65speciesintheAmericas,
thefactthatnoneoftheAmericancladesarebasalimpliesthatthegroup'sancestrywasintheOldWorld.[10]

Notes
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1.MeaningthelowesttaxonomicranksoftherespectivecladesseeAmborella
example.
2.NewCaledoniaisviewedasarefugiumi.e.,inthiscasethegeographic
locationofthebasalcladeisnotthoughttoprovideevidenceforthelocalein
whichangiospermsoriginated.
3.Notethatthecladogramaswrittenconveysnoinformationregardingthe
relativepresentorpastspeciesrichnessofthefourtaxa,orwhich,ifanyof
them,maypossessmoreancestraltraits.
4.ThisconclusionisconsistentwiththefactthatDidelphimorphiaisthebasal
orderwithinMarsupialiai.e.,extantmarsupialsasawholealsoappearto
haveoriginatedinSouthAmerica.[7]

References
1.Baum,D.A.(4November2013)."PhylogeneticsandtheHistoryofLife".
ThePrincetonGuidetoEvolution.PrincetonUniversityPress.p.57.
ISBN9781400848065.OCLC861200134.
2.Crisp,M.D.Cook,L.G.(March2005)."Doearlybranchinglineages
Relationshipofbiogeographyand
signifyancestraltraits?".TrendsinEcology&Evolution20(3):122128.
phylogenyofbatsuperfamily
doi:10.1016/j.tree.2004.11.010.
NoctilionoideainferredfromnuclearDNA
3.Jenner,RonaldA(2006)."Unburdeningevodevo:ancestralattractions,model
sequencedata,showingthebasalposition
organisms,andbasalbaloney".DevelopmentGenesandEvolution216:385
oftheMalagasyfamilyMyzopodidae.
394.doi:10.1007/s0042700600845.
4.Essig,F.B.(20140701)."What'ssoprimitiveaboutAmborella?".Botany
Locationswithonlyfossilmembersare
Professor.Retrieved20141004.
indicatedbyredstars.
5.MoyaSola,S.Alba,D.M.Almecija,S.CasanovasVilar,I.Kohler,M.
DeEstebanTrivigno,S.Robles,J.M.Galindo,J.Fortuny,J.(20090616).
"AuniqueMiddleMioceneEuropeanhominoidandtheoriginsofthegreatapeandhumanclade".PNAS106(24):96019606.
doi:10.1073/pnas.0811730106.PMC2701031.PMID19487676..
6.Krell,FrankT.Cranston,PeterS.(2004)."Whichsideofthetreeismorebasal?".SystematicEntomology29:279281.
doi:10.1111/j.03076970.2004.00262.x.
7.Nilsson,M.A.Churakov,G.Sommer,M.VanTran,N.Zemann,A.Brosius,J.Schmitz,J.(20100727).Penny,David,
ed."TrackingMarsupialEvolutionUsingArchaicGenomicRetroposonInsertions".PLoSBiology(PublicLibraryofScience)
8(7):e1000436.doi:10.1371/journal.pbio.1000436.PMC2910653.PMID20668664.
8.Teeling,E.C.Springer,M.Madsen,O.Bates,P.O'Brien,S.Murphy,W.(20050128)."AMolecularPhylogenyforBats
IlluminatesBiogeographyandtheFossilRecord".Science307(5709):580584.doi:10.1126/science.1105113.
PMID15681385.
9.Gunnell,G.F.Simmons,N.B.Seiffert,E.R.(20140204)."NewMyzopodidae(Chiroptera)fromtheLatePaleogeneof
Egypt:EmendedFamilyDiagnosisandBiogeographicOriginsofNoctilionoidea".PLoSONE9(2):e86712.
doi:10.1371/journal.pone.0086712.Retrieved20140205.
10.Slowinski,J.B.Boundy,J.Lawson,R.(June2001)."ThePhylogeneticRelationshipsofAsianCoralSnakes(Elapidae:
CalliophisandMaticora)BasedonMorphologicalandMolecularCharacters".Herpetologica57(2):233245.
JSTOR3893186.

Externallinks
"InterpretingtheTreeDiagramorListofSubgroupsonaTreeofLifePage".TreeofLifewebproject.
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