Spurring Plant Diversification Are Floral Nectar Spurs a Key Innovation

Scott A Hodges Michael L Arnold

Proceedings Biological Sciences Vol 262 No 1365 (Dec 22 1995) pp 343-348

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Spurring plant diversification are floral nectar spurs a key innovation

SCOTT A E J O D G E S 1 A N D 1 4 I C H A E L I 1ZRNOLD2 Oeparlnrenl cfLcoiogy Erolr~ion and LlIarine ISioloqy Inicersity oJCali)mia Santa Rarhara (A 9306 lS1 - )epartnienl qf Genetic 1~niversity q Geotyia 1~Aen (il30602 lS A

SUMMARY

f-Iigh lcvcls of spccies d i~~crs i ty a ltcy innovatioil in taxonomic grorlps have often I)ccn esplairlcd by Hovccr thr difliculty in establishing a causal rolc l~etwecn a proposed key inrlovatior~ anci iricrcased s1)ccies tliversity as well as in substantiating that tlicrsity pattcrns a r r diflicrcnt Crorrl rrull motlels has lcd t o major criticisms of ltey irlrlovatiorl hypotheses Hcre we show that pattcrns ordiversification within and among clades that havc cvolvcci floral nectar spurs strongly srlpport thc hypothesis that floral nectar spurs rcprcsent a key innovation 130th reprociuctivc success and rcprotluctive isolation can I)(- irlfluerlccd I)y sirnplc charlgrs in rlrctar spur morl)hology arid thc acquisition of nectar spurs in a wide array of plant groups is highly correlated with incrcased spccics diversity

1 INTRODUCTION

11 Strrldarnrr~tal goal in biology is to llndcrstaild the factors that irlflueilcc organismal diversity i U a r ~ i n 1872 PJayr 1942 Simpson 1953) Key inno~ations are novel rnorf)l~ological or I)chavionral traits t l~ought cithcr to open new atlaptivc zones (Simpsorl 1953) or to olrer tlic ability to rapidly spcciate after cnvirorl- rricrltal c l~ange (Tiem 1973) Il~us the evolution or n kry innovatior1 is thorrght to l)c a partic~rlarly important evcrlt affecting suhscqncnt patterns of speciatiorl aridor cxtinctiorr 1)cspite thc commorl tlsc o l kcy innovations as an csplanation Tor cladc-spccific irrcrtascs irr diversity t l ~ c concept has heen criticized as bcing essentially untcstal)lc (Cracract 1990) First demonstrating how a particular trait can cause ratcs of diessification to increase is often prol)lcrnatic ((2racract 1990) Second large diftc~renccs irr spccics nurnbcrs bctwern sister taxa can he explained so1tly through stochastic processes i~ l iaup et al 1973 Slovirrski amp Grlyer 1989 199Sj H r r c ve corrsidcr these points irr tlctermining if floral nectar spurs arc a kcy inrrovation 1)articularly in tlie columbine gcrlus Aq~ile~gia

r r ~uncierlying assumption of most spccics concepts is the necessity Tor reprotluctivc isolation [Uobzhansky 1937 Mayr 1942 Grant 19Gl) Characters that can incrcasc thc likelihood oC rcprotiuctic isolation may therehre ir~crease rates of spcciation Nectar spur morphology is intimately ticd to re1)rotluctiorl arrd car1 ilrflucrrce rcprotluctivc isolation Spurs arc tubular ontgrovths orpetals or other floral parts that increase the distance hctwccn t l ~ c floral reward and tllc relxoductivc parts of the flowcr Relatively sirnplc difFCrtrlccs in the lengtll shape orientation and colonr or nectar spurs arc associated with difterent poll i~~ators

lroc R Soi Lond R (111951262 343 3121 P1ir~1idIn f r m l 111itain

and aflect reproductioll 1or instarlcc in orcl~id species the lerlgtli of a n intlividnal plants spur strongly iilflrrtrrccs rcprodtlctivc srtcccss (Nilssorl 1988) In Aqi~ilegia caerzileti variation among populations in the length ant1 colour of spurs is correlated with ariation in tlie types o l pollinators visiting the plants (Miller 198 1 1 Uifr(rcnccs in spurs corresponciin to diftircnt pollirlators (I-lu~rnningl)irds versus hawkmotlis) h a w also been shown to aid in reproductic isolatiorl hctwecrr two hy lwidizirlg lquilgk spccirs (Grant 1952 Hotlgrs 6i Arnold 1994h Similarly in Diascia ariation among specics in spur lcngth is correlatctl with morphological variation in their bec pollinator (Stcincr amp LVhitcl~cad 1090 Steir~cr amp 1Vhitelicad 1991j logcthrr thesr stl~dics suggest that diftcrcncrs irr ncctar spurs among po~)ulatiorrs of a spccies car1 facilitate rc~)roductivc isolation arrd possi0ly leact to incrrascd spcciation

mong species oC ilyz~ilesqzclthcrc arc lov lcvcls of DNA seqrlencc ciivergcncc despite the vitiespread distribution of the gcnus suggcstirrg a recent radiation rrsulting from L kcy innovation (I-Iodges amp Arrrold 19940) 10 deterrriirre il the evolution or ncctar spurs was a key innovation ti)r this gcuus we sought to temporally link the evolution o l ncctar spurs with an increase ill divcrsification rate ill tlie columbines Using a molecular pllylogrnetic approach we con-structcd phylogenies that irrclrldcd t ~ o spurless spccics Aqzriiegia eialcarata and bSemiciyi~ile~qSia adoxoide rhcse spcies havc bcen suggested to I)( basal to the co11lml)ilrc clacic (RIunz 194Gj and could thus revcal wllcthcr spurs evolved near the time of tile radiation ~dditionally because 11rcttr spurs have cvolvcci indcpc~idelltly in numrrous lineages we rlscd nrlll models to trst i l irrcreaseci diversification is gcncrally linkcd wit11 the evolution or floral ilcctar spurs

2 MATERIALS A N D M E T H O D S

111) logcllctic rrlrlscs vrrc corrduc~tcd r~silig L)N scrl11(11c(sof the rlucle~tr illtcrllrll trrrlscril)cd q)rcx~r r( zio~ltrrcI ir~tcrg(~~ric I ) ~ ~ ( Y ~ I Itll( (lrIoro~~l~st s~)tcc~r tllr ibc l t r ~ t l tr11 gcrlcs Ifodges R rr~olcl I)l~im) r~i~)l if lc t t io~~ l)~oto(ols 11rvc l)r(- t r ~ t I ~c ( luc~~ l ( i r~g v io~~s lyI)cc~r tlcscril)c~tl 1 Hodgcs k lr~olcl I )lio Spccics 11iir1gthctl i ~ l ( ~ l ~ ~ t I r t l i l l o ~ ~ rill rhos(gt IIY(~I pr(iot~s

s11ggcslI tcrnl)o~~l I)ctzcc~l111c eolutior~ o l ~ i c ~ t ~ ~ l i ~ ~ k sp~11str1tI colt~rrlbir~e(11( r ~di~tiorr

10 test if tllc tlicrsiIic~tio~r 01 lgiiriqcz rn~tcllcs I

kc) irlrro~tiorr ll~)otllcsis vr S r~ld(~rso~lusctl R Jgtorlogl~l~c~srn~xim~~~n-likclillootl I1 I rpproaclr 1Sr1rtlc1so118r 11)4) llris rnrtlrotlI ) o ~ ~ o g l r ~ ~ c ( I I I

( x(l~~tlc ttio11 l~ypotl~csis r~~otl(lI kc) i r ~ r ~ o i r r i t l~cr t l ~ c ~ v i t l r rlo clrrrrgc i r ~~rtc01 divc~silicatior~ rnotl(lor I

ivirll I (-11~11g(~ o11r 01 of 1 1 1 ~in 0111) I I I C 1)irstI li~lcrg(h itr1tI) sis 1 Hotlgrs R r ~ ~ o l t l10040 i a 1 ~ 1it1 icl(litior~ ilrgrorlp is supportrd l hr orlc-I)1rrlnctcr lnotlvl H ( l o s1)ccics tllirt (re urlr rilrI)Ic at t I ~ t ttime -lq~rilqqrtr (11 rcfi1s to tlrcjtl~ rnotl(l ~ i i t l~ i r r tllr cl~ss ol~iiotlc~ls

ic(rco~cta ~ r ~ t ljYitt~c~y~~it~qi(r ~ ~ - c i o ~ ~ s l ) X Ilrllrlctcrs S~r~tl(rsor~ czdoo(ic As tllat 11~s amp I)o~ioglluc I104) tl(s(ril)(tl( o~i11[e~icz111tl ~Ycrt~ll~o~~~zzc~sti~)i(I t i ie~

n r r c used to root the t r c ~ nid lAlJl SolIi)rtl 1)93 virs usctl to a~r~l)sc(he dt t ~ 1Hotlgcs R lr~lolcl 1I94 (I i

Tll( ptttcrl~ of (li(rsifi(itio~ri l l ~v21s~Ic~ii(qr~ i~rlal)sctl 1)) t~sirrg Strrtlc~son rn~xirnu~ir-k l)o~~ogl~uc s lik(lil~ootl ~rlctllod i k l ) ) t Sar~dcrsor~ I)orroglr~~c~ rl~is~rlcrlrotl 11scs a rr~ltlorrl spcciatiorl rnotl(l ro ((st il ol)sc~cd 11t t t( r11~)f i l l tllrrc-t~xorr 1111~- tlicrsity I

logc~ry tzo cl~tIc (hat S I I I I ~ I kc)l)~oposc~I i r~ r~o-i tio or^ I I I C t l~cir sis(c1 gro111)) hl~[)[)ort k(y irrrro~tio~i ir

11) potll(sis itlrill ly~~ic~qi~ rr( tvo there tIisti11ct clirclcs t l i ~ ~ group t11c Erlrol~rarl ~ r l c l~~rrdo~ii irr r(+ isit~l sprcic a1)lrt from the Nortl~ lnlc~icarl sl~ccirs I Hodgcs amp rrroltl is tl~c sister I)lioi Se~i~riciqiii(~gi(i g1-ot11 li)r 24q~ri~qic~ rsrc ligt~rc I ) V( tllr~s corril~lcd t l ~ cclivc~sit) olVt~tnoyuliqii~ ~ v i t l ~io~rc al)ccic~s the Oltl 111d Nc 2orltl col1111l1)illc s~)c(ics($7 1r1(1 23 l)(ci(~ rcsl)c~ctivcl))

111a srlrvcgt 01 ~)ul)lil~c~tl ~ I L ( L v(arrrl ~1111~1l)lislrctl itlrlltifictl c~latlistic arrrlgtscs tllrt tlxa it11 i~lclr~tl(tl 1lcctrr spur Iccar~sc tllr tests ol clic~rsific~irtioli ~)ttt( r~ls olsist(r taxr bze drlxll(l o11 the idc~i t i f ica t io~~ rcstrictc~tl our rr~~l)sis 1)1ol)1bIy(on-to studies ( l r r t trirrctl rellc~sc~~lttlivrs sistc1 groul~s li)r olrll potc~lti~l ( I c I I (Ii(lc ~ l l c ~ c 011cc11rcttr Y I N I ~ S 111vc (~vol(~il hih~(rtaxi1 vcr( i t l~~r~t i l ic t l~ if [ I ] ( tli(rsitgt of v(- ( c s t ( ~ l s~wcics I gt sig~lificantlgt grcrtcr irl the spurrrtl t~xn coml)lrc(l it11 tlrrir rroll-hpl~rrttI sister tasa ~lsillga ~riotlcl~ I I L I I ~ ~ o I I ~sprciittiotl ~ I I I C Icsti~r(tior~111cl l)o(ll I

Vil(oxor~ six11 tcst trr t l Iisl~c~sconrl)ir~ccl ~rol~rl)ility t(st iSlovirlshi R ( ~Igt ( I or1111lltl03) to t ( ~ tli)r 111~ rssocirtiorl I)etccrl the c olr~tio~~ 1lcctrr spurs r r ~ t l01 i~lcrcasctl tlic~sifi(~rtiorl 111( rrurriI)(r 01 s[)r(-i(si l l

cr(ll gro111) ol)trirlctl lkorli ~ ~ a l ) I ) ( ~ r l c ~ 10931 i

3 RESULTS

Tl~c 1111ylogc1rctic recorls(rl~ctio~r of 111dIy~iile~~qio ~-( I r t (~ l ill I trcxctitsa ~-cx~~ltcd il~glcrnor ~ ) ~ r s i m o ~ l i o ~ ~ s vitll ~Yitti~cre~ii~qicz c1(010i(i (loscl ztIig~r(d ~ I I I C III LS I~ to thr c~olr~ml)i~lcs vhc~-r ~slyuii~~qio isi~ra~~mrc~rn1)11((tI vithirl the I)rs~l c1rtlr olthc coltr~r~l)i~rc iscc lig11rc I )

lhe ~)lircclnc~lt of tllr sl~urlcss ~Siinierc~~il~~qcr ir~tlicltcs 11lrt thr3 most prrsilrlollior~s positiorl lor t l ~ c eolr~tiorr01 ri(((tr s11~1ls is closc to tlir c o l ~ ~ r n b i ~ ~ c Irrclirtio~~figure l i l l~is Ijrnrlcli lias ~clatirly li- ir~lirrcd rrrrclc~otiiI~~ (-II III~(s 1 0 ~ I I C I I ~ ~ I I I ( I I ( or~r~c(( i~rg~ l i ( ( o ~ ~ ~ p i ~ r ( x l I ~ ~ ~ i l ~ ~ q i n S ~ ~ ~ i ~ i c z ~ r c i l ~ ~ ~ q i ~ ~ clrtiltclaclr vitl~ the IS~IIIILVI (six vcxsris 19 i~rl irrc~d cliangcs ligrlre 1 ) llirsc tlata

1 1 ~ s1 1 0 ~rirtiorl rates o l t l i v ( ~ ~ ~ s i l i ( ~ ~ t i o ~ l ~ v oi r ~ v l ~ i l ( t I r ( t o-~jlrtllIctrr lllo(l(~ls 11 111tI 11 I I ~ c it (-lllllg(~i l l

cli~crsificrtiorr rr t( I ) L I I orll i r l ollr I)rallcll of (lrc~ clntl( co~ltrirrillg1 1 1 ~ kc) i11novltion figrrrc 2 ) llrcsc models rrr rcicctctl 1)) the ~llrlysis1 1igr1rc 2 1 3 0)5I 13o(I1 tvo-l)ilr~rlictc~t r~iotlclst11rt arc co~~sistc~~t ~ v i t l rI

kc i~rl~ovt t ior~ 1zcr-c rot ~cjcctcdI ) )Ir)lotlrcsis tl~c irrrrlysis rI~rc~c~-~lrn~rlcrcr 111otl(1s vc~c r~ot tes(ctl I)cctusr tvo-prrlr1Ictcr rnotl(ls arc tos~~ll ic ic~~ltd(-scril)c t l ~ c clrtt ~ S t l ~ d c ~ s o ~ r k L)o~loghuc 10)i]

S ig~~ i l i ( t~~ t ly tlivc~sity t11111 cx l~ (~ r c t l g ~ c ~ ~ r r ~ vs dc~tt3c~trtli r l either lbr~r or lic o11t oltlrc six glo111)s tlri~l lr~cirlclrl)e~ld(rltlycolctl rlcctar spurs ( t r I ) I ( 1 1 Ill( r c l ~ t i o ~ ~ L I I C II rs ((111i ocrI lk)r I)OII~I~~I l(ot~ii~ii~ tlcl)crltlir~g orr (Ire sister grorlp icl(rl(ifictl l s i~ lg tlrc sistc~ g ~ o u ~ ) iqc~o iderrtific~tl 1)) Hoot I) ~sc~lolr rr~orl~l~olog)I kloot 1001 I ~ r ~ dL)N scclucrlccs oIil)( I 11LII ( I 18s rl)N (kioot 1 C1c15 i 111( r(Irtio~l is sigr~ific~~rt OOil) I~J c e ~ i ltqioli)it)= vIrrr(rs ~ii~q 2Yqco I f O(YI IIICI( iiiffirqi irtc~l(ifir~cl I) ) I IJolir~~ss( R ~Jrrlscrl llO3 i l ) ~ ~ t l 1)Ni l ( ~t t i ( t io~~ 1 or1 c l ~ l o ~ ~ o ~ ) l ~ ~ s t sit( rriatio~i the rclatiorl ~s 11ot sigr~ilicarrl I = 00951 lll(gt si11glc cIcrt (x((yjtior~ I1~tlfier~qor~i~tr

( Scvcr spc~cic~s i ~ rthe spr~rrcd group I L L I I C I ) li oltli(~ six groups hat1 ~nol-c spc~ics ill 111(sljt~rrctl clatlc tlialr t l ~ c rlorr-spr~rrcd (little ~rld thus a lilcoxorl six11 st is II (gt I I I ) sigr~ifilttr~(= of t l ~ c sist(r 1 f) 005$1U R(~~tr tI lcss group usctl I)t~~tt~iitr~li)r t r ~ t l I e ~ o ~ u i ~ r Iislicr co~nl)inccl1)rol)rl)ilir) tcst Slozirlski amp ( u)cr ll)i i is lriglrIgt sigrliIi(~~~rt fy2 = 3848 or 4026 (11 == 12 I lt 000 I i

4 DISCUSSION

2 MATERIALS A N D M E T H O D S

111) logcllctic rrlrlscs vrrc corrduc~tcd r~silig L)N scrl11(11c(sof the rlucle~tr illtcrllrll trrrlscril)cd q)rcx~r r( zio~ltrrcI ir~tcrg(~~ric I ) ~ ~ ( Y ~ I Itll( (lrIoro~~l~st s~)tcc~r tllr ibc l t r ~ t l tr11 gcrlcs Ifodges R rr~olcl I)l~im) r~i~)l if lc t t io~~ l)~oto(ols 11rvc l)r(- t r ~ t I ~c ( luc~~ l ( i r~g v io~~s lyI)cc~r tlcscril)c~tl 1 Hodgcs k lr~olcl I )lio Spccics 11iir1gthctl i ~ l ( ~ l ~ ~ t I r t l i l l o ~ ~ rill rhos(gt IIY(~I pr(iot~s

s11ggcslI tcrnl)o~~l I)ctzcc~l111c eolutior~ o l ~ i c ~ t ~ ~ l i ~ ~ k sp~11str1tI colt~rrlbir~e(11( r ~di~tiorr

10 test if tllc tlicrsiIic~tio~r 01 lgiiriqcz rn~tcllcs I

kc) irlrro~tiorr ll~)otllcsis vr S r~ld(~rso~lusctl R Jgtorlogl~l~c~srn~xim~~~n-likclillootl I1 I rpproaclr 1Sr1rtlc1so118r 11)4) llris rnrtlrotlI ) o ~ ~ o g l r ~ ~ c ( I I I

( x(l~~tlc ttio11 l~ypotl~csis r~~otl(lI kc) i r ~ r ~ o i r r i t l~cr t l ~ c ~ v i t l r rlo clrrrrgc i r ~~rtc01 divc~silicatior~ rnotl(lor I

ivirll I (-11~11g(~ o11r 01 of 1 1 1 ~in 0111) I I I C 1)irstI li~lcrg(h itr1tI) sis 1 Hotlgrs R r ~ ~ o l t l10040 i a 1 ~ 1it1 icl(litior~ ilrgrorlp is supportrd l hr orlc-I)1rrlnctcr lnotlvl H ( l o s1)ccics tllirt (re urlr rilrI)Ic at t I ~ t ttime -lq~rilqqrtr (11 rcfi1s to tlrcjtl~ rnotl(l ~ i i t l~ i r r tllr cl~ss ol~iiotlc~ls

ic(rco~cta ~ r ~ t ljYitt~c~y~~it~qi(r ~ ~ - c i o ~ ~ s l ) X Ilrllrlctcrs S~r~tl(rsor~ czdoo(ic As tllat 11~s amp I)o~ioglluc I104) tl(s(ril)(tl( o~i11[e~icz111tl ~Ycrt~ll~o~~~zzc~sti~)i(I t i ie~

n r r c used to root the t r c ~ nid lAlJl SolIi)rtl 1)93 virs usctl to a~r~l)sc(he dt t ~ 1Hotlgcs R lr~lolcl 1I94 (I i

Tll( ptttcrl~ of (li(rsifi(itio~ri l l ~v21s~Ic~ii(qr~ i~rlal)sctl 1)) t~sirrg Strrtlc~son rn~xirnu~ir-k l)o~~ogl~uc s lik(lil~ootl ~rlctllod i k l ) ) t Sar~dcrsor~ I)orroglr~~c~ rl~is~rlcrlrotl 11scs a rr~ltlorrl spcciatiorl rnotl(l ro ((st il ol)sc~cd 11t t t( r11~)f i l l tllrrc-t~xorr 1111~- tlicrsity I

logc~ry tzo cl~tIc (hat S I I I I ~ I kc)l)~oposc~I i r~ r~o-i tio or^ I I I C t l~cir sis(c1 gro111)) hl~[)[)ort k(y irrrro~tio~i ir

11) potll(sis itlrill ly~~ic~qi~ rr( tvo there tIisti11ct clirclcs t l i ~ ~ group t11c Erlrol~rarl ~ r l c l~~rrdo~ii irr r(+ isit~l sprcic a1)lrt from the Nortl~ lnlc~icarl sl~ccirs I Hodgcs amp rrroltl is tl~c sister I)lioi Se~i~riciqiii(~gi(i g1-ot11 li)r 24q~ri~qic~ rsrc ligt~rc I ) V( tllr~s corril~lcd t l ~ cclivc~sit) olVt~tnoyuliqii~ ~ v i t l ~io~rc al)ccic~s the Oltl 111d Nc 2orltl col1111l1)illc s~)c(ics($7 1r1(1 23 l)(ci(~ rcsl)c~ctivcl))

111a srlrvcgt 01 ~)ul)lil~c~tl ~ I L ( L v(arrrl ~1111~1l)lislrctl itlrlltifictl c~latlistic arrrlgtscs tllrt tlxa it11 i~lclr~tl(tl 1lcctrr spur Iccar~sc tllr tests ol clic~rsific~irtioli ~)ttt( r~ls olsist(r taxr bze drlxll(l o11 the idc~i t i f ica t io~~ rcstrictc~tl our rr~~l)sis 1)1ol)1bIy(on-to studies ( l r r t trirrctl rellc~sc~~lttlivrs sistc1 groul~s li)r olrll potc~lti~l ( I c I I (Ii(lc ~ l l c ~ c 011cc11rcttr Y I N I ~ S 111vc (~vol(~il hih~(rtaxi1 vcr( i t l~~r~t i l ic t l~ if [ I ] ( tli(rsitgt of v(- ( c s t ( ~ l s~wcics I gt sig~lificantlgt grcrtcr irl the spurrrtl t~xn coml)lrc(l it11 tlrrir rroll-hpl~rrttI sister tasa ~lsillga ~riotlcl~ I I L I I ~ ~ o I I ~sprciittiotl ~ I I I C Icsti~r(tior~111cl l)o(ll I

Vil(oxor~ six11 tcst trr t l Iisl~c~sconrl)ir~ccl ~rol~rl)ility t(st iSlovirlshi R ( ~Igt ( I or1111lltl03) to t ( ~ tli)r 111~ rssocirtiorl I)etccrl the c olr~tio~~ 1lcctrr spurs r r ~ t l01 i~lcrcasctl tlic~sifi(~rtiorl 111( rrurriI)(r 01 s[)r(-i(si l l

cr(ll gro111) ol)trirlctl lkorli ~ ~ a l ) I ) ( ~ r l c ~ 10931 i

3 RESULTS

Tl~c 1111ylogc1rctic recorls(rl~ctio~r of 111dIy~iile~~qio ~-( I r t (~ l ill I trcxctitsa ~-cx~~ltcd il~glcrnor ~ ) ~ r s i m o ~ l i o ~ ~ s vitll ~Yitti~cre~ii~qicz c1(010i(i (loscl ztIig~r(d ~ I I I C III LS I~ to thr c~olr~ml)i~lcs vhc~-r ~slyuii~~qio isi~ra~~mrc~rn1)11((tI vithirl the I)rs~l c1rtlr olthc coltr~r~l)i~rc iscc lig11rc I )

lhe ~)lircclnc~lt of tllr sl~urlcss ~Siinierc~~il~~qcr ir~tlicltcs 11lrt thr3 most prrsilrlollior~s positiorl lor t l ~ c eolr~tiorr01 ri(((tr s11~1ls is closc to tlir c o l ~ ~ r n b i ~ ~ c Irrclirtio~~figure l i l l~is Ijrnrlcli lias ~clatirly li- ir~lirrcd rrrrclc~otiiI~~ (-II III~(s 1 0 ~ I I C I I ~ ~ I I I ( I I ( or~r~c(( i~rg~ l i ( ( o ~ ~ ~ p i ~ r ( x l I ~ ~ ~ i l ~ ~ q i n S ~ ~ ~ i ~ i c z ~ r c i l ~ ~ ~ q i ~ ~ clrtiltclaclr vitl~ the IS~IIIILVI (six vcxsris 19 i~rl irrc~d cliangcs ligrlre 1 ) llirsc tlata

1 1 ~ s1 1 0 ~rirtiorl rates o l t l i v ( ~ ~ ~ s i l i ( ~ ~ t i o ~ l ~ v oi r ~ v l ~ i l ( t I r ( t o-~jlrtllIctrr lllo(l(~ls 11 111tI 11 I I ~ c it (-lllllg(~i l l

cli~crsificrtiorr rr t( I ) L I I orll i r l ollr I)rallcll of (lrc~ clntl( co~ltrirrillg1 1 1 ~ kc) i11novltion figrrrc 2 ) llrcsc models rrr rcicctctl 1)) the ~llrlysis1 1igr1rc 2 1 3 0)5I 13o(I1 tvo-l)ilr~rlictc~t r~iotlclst11rt arc co~~sistc~~t ~ v i t l rI

kc i~rl~ovt t ior~ 1zcr-c rot ~cjcctcdI ) )Ir)lotlrcsis tl~c irrrrlysis rI~rc~c~-~lrn~rlcrcr 111otl(1s vc~c r~ot tes(ctl I)cctusr tvo-prrlr1Ictcr rnotl(ls arc tos~~ll ic ic~~ltd(-scril)c t l ~ c clrtt ~ S t l ~ d c ~ s o ~ r k L)o~loghuc 10)i]

S ig~~ i l i ( t~~ t ly tlivc~sity t11111 cx l~ (~ r c t l g ~ c ~ ~ r r ~ vs dc~tt3c~trtli r l either lbr~r or lic o11t oltlrc six glo111)s tlri~l lr~cirlclrl)e~ld(rltlycolctl rlcctar spurs ( t r I ) I ( 1 1 Ill( r c l ~ t i o ~ ~ L I I C II rs ((111i ocrI lk)r I)OII~I~~I l(ot~ii~ii~ tlcl)crltlir~g orr (Ire sister grorlp icl(rl(ifictl l s i~ lg tlrc sistc~ g ~ o u ~ ) iqc~o iderrtific~tl 1)) Hoot I) ~sc~lolr rr~orl~l~olog)I kloot 1001 I ~ r ~ dL)N scclucrlccs oIil)( I 11LII ( I 18s rl)N (kioot 1 C1c15 i 111( r(Irtio~l is sigr~ific~~rt OOil) I~J c e ~ i ltqioli)it)= vIrrr(rs ~ii~q 2Yqco I f O(YI IIICI( iiiffirqi irtc~l(ifir~cl I) ) I IJolir~~ss( R ~Jrrlscrl llO3 i l ) ~ ~ t l 1)Ni l ( ~t t i ( t io~~ 1 or1 c l ~ l o ~ ~ o ~ ) l ~ ~ s t sit( rriatio~i the rclatiorl ~s 11ot sigr~ilicarrl I = 00951 lll(gt si11glc cIcrt (x((yjtior~ I1~tlfier~qor~i~tr

( Scvcr spc~cic~s i ~ rthe spr~rrcd group I L L I I C I ) li oltli(~ six groups hat1 ~nol-c spc~ics ill 111(sljt~rrctl clatlc tlialr t l ~ c rlorr-spr~rrcd (little ~rld thus a lilcoxorl six11 st is II (gt I I I ) sigr~ifilttr~(= of t l ~ c sist(r 1 f) 005$1U R(~~tr tI lcss group usctl I)t~~tt~iitr~li)r t r ~ t l I e ~ o ~ u i ~ r Iislicr co~nl)inccl1)rol)rl)ilir) tcst Slozirlski amp ( u)cr ll)i i is lriglrIgt sigrliIi(~~~rt fy2 = 3848 or 4026 (11 == 12 I lt 000 I i

4 DISCUSSION

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Sc~rnrclrl~~ilc~g~tiA51a N A m c r ~ c ~ ~ Europe

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c ~ l ) i c I ( ~ ~ ~ ~ i i c r o ( ~ I r ~ ~ ~ ~ ~ ( ~ t c r h Sy 11itI IIIO ~ i i ~ ~ ~ r o ~ r r o r ~ gt I i o l o ~ ~ lo 16 741 755

11001 S 13 lCl)r) 111sccI o ~ i llr) log(~i oftli( K ~ I ~ ~ I I ~ ( ~ I I ~ I ( ~ ( ~ ~ I ~ 11il3 IIIII 11i(1 18s ril)o~or111lDN se(l11(11((( I I ~ lt I 11(11 S col I l l till 1gtscs1

Ii(~iiIlt 1 1)73 k ~ ~ o l ~ ~ ~ i o ~ i ~ r gt ~ ~ ~ i o s ~ ) l i o l o ~ i ~ ~ i ~ lst1 tt(gics t t i c I

i ~ i ~ i o ~ ~ l i o ~ i h ci(~Irli0 l)Iiary~ig(~tl jth Sq Zoo 2 2 ~ 421 111

1 l t l ) l ) ( 1 lc L)J 1))3 TI~III~I-~IIIIL t 11i ( rs i~ i ~~ l i l~ r ic lgc 11 (as

h l a gt r 1 1912 S gt ~ s t ( ~ ~ ~ i t ~ i ( ~ i St t ~ i c l tlrc o r ig i~r o l ~ s ~ ~ c c i c i 1ork ( k ~ l r ~ ~ i i l ~ i ~crsit) Irci t ~ i i

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X i I lt o ~ r ~ 111~ c o l ~ ~ t i o ~ i (Ice)I ICl88 of llo(ri i t11

(~orolliltill)( Ylill~ IO 334 147 I 19 OI~irht( ~clk i ~ h l i ( ~ l ~ ~ ( ~ l s ~ Ilt h l k j 11 H J S c o ~ t ~

Irl~iic~r l)Cl3 h l o ~ i o l ) l ~ ) l ~tlic s t (~ricl~rc~ 01 IIIO iOc~iti- (I( 11ioti 01 t l ~ c i r r111jor I i~ici tg(~s i~i l t~rrccl IIo11i 1 ) S j ~ ~ ( I L I ( I I C V ~ l t i t i IltII ((11(1 8 0 ~700 722 or ~11i1 t o ( ~ i

11 I L ~ ~ I O I l)(j5 t o g c ~ i c i ( ~( ht110ich011 tlrv ~ ( I I L I l(ii~qi(~

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Stcitrcr li 1 amp Ylii~c~lrcicl I ) I)IO I)olliti~lor i(Iilgtt~itioti10 l lo ~ th ~ ~ ~ ( I I ~ I I ~ ~I ) I ( I ( I I I oiIL~c(~r(~t i t1g I ~ I ( I c~ol t i to~t44 170 1 1707

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You have printed the following article

Spurring Plant Diversification Are Floral Nectar Spurs a Key InnovationScott A Hodges Michael L ArnoldProceedings Biological Sciences Vol 262 No 1365 (Dec 22 1995) pp 343-348Stable URLhttplinksjstororgsicisici=0962-84522819951222292623A13653C3433ASPDAFN3E20CO3B2-P

This article references the following linked citations If you are trying to access articles from anoff-campus location you may be required to first logon via your library web site to access JSTOR Pleasevisit your librarys website or contact a librarian to learn about options for remote access to JSTOR

References

Phylogenetics of Seed Plants An Analysis of Nucleotide Sequences from the Plastid Gene rbcLMark W Chase Douglas E Soltis Richard G Olmstead David Morgan Donald H Les Brent DMishler Melvin R Duvall Robert A Price Harold G Hills Yin-Long Qiu Kathleen A KronJeffrey H Rettig Elena Conti Jeffrey D Palmer James R Manhart Kenneth J Sytsma Helen JMichaels W John Kress Kenneth G Karol W Dennis Clark Mikael Hedren Brandon S GautRobert K Jansen Ki-Joong Kim Charles F Wimpee James F Smith Glenn R Furnier Steven HStrauss Qui-Yun Xiang Gregory M Plunkett Pamela S Soltis Susan M Swensen Stephen EWilliams Paul A Gadek Christopher J Quinn Luis E Eguiarte Edward Golenberg Gerald HLearn Jr Sean W Graham Spencer C H Barrett Selvadurai Dayanandan Victor A AlbertAnnals of the Missouri Botanical Garden Vol 80 No 3 (1993) pp 528-548+550-580Stable URLhttplinksjstororgsicisici=0026-649328199329803A33C5283APOSPAA3E20CO3B2-W

Columbines A Geographically Widespread Species FlockScott A Hodges Michael L ArnoldProceedings of the National Academy of Sciences of the United States of America Vol 91 No 11(May 24 1994) pp 5129-5132Stable URLhttplinksjstororgsicisici=0027-8424281994052429913A113C51293ACAGWSF3E20CO3B2-5

httpwwwjstororg

LINKED CITATIONS- Page 1 of 3 -

Aqlrlcgia

Sc~rnrclrl~~ilc~g~tiA51a N A m c r ~ c ~ ~ Europe

R E F E R E N C E S

i s o l ~ t i o ~ i lqIfq(~ ~ l i c lI I ~ I I~ ~I I I I I ~ ~ I I gt I I l ) ( ~ c c ~ i IIIIHOII

1)111 J I O I I I ( ill Sri 1TLY49172 193 2410 Hoot S 13 1991 1 11~Ioqc1 i~ I)rs(~l1111oIt11c ~ lt I I I I I I ~ ( ~ ~ I ~ I ( ~ ~ I c

c ~ l ) i c I ( ~ ~ ~ ~ i i c r o ( ~ I r ~ ~ ~ ~ ~ ( ~ t c r h Sy 11itI IIIO ~ i i ~ ~ ~ r o ~ r r o r ~ gt I i o l o ~ ~ lo 16 741 755

11001 S 13 lCl)r) 111sccI o ~ i llr) log(~i oftli( K ~ I ~ ~ I I ~ ( ~ I I ~ I ( ~ ( ~ ~ I ~ 11il3 IIIII 11i(1 18s ril)o~or111lDN se(l11(11((( I I ~ lt I 11(11 S col I l l till 1gtscs1

Ii(~iiIlt 1 1)73 k ~ ~ o l ~ ~ ~ i o ~ i ~ r gt ~ ~ ~ i o s ~ ) l i o l o ~ i ~ ~ i ~ lst1 tt(gics t t i c I

i ~ i ~ i o ~ ~ l i o ~ i h ci(~Irli0 l)Iiary~ig(~tl jth Sq Zoo 2 2 ~ 421 111

1 l t l ) l ) ( 1 lc L)J 1))3 TI~III~I-~IIIIL t 11i ( rs i~ i ~~ l i l~ r ic lgc 11 (as

h l a gt r 1 1912 S gt ~ s t ( ~ ~ ~ i t ~ i ( ~ i St t ~ i c l tlrc o r ig i~r o l ~ s ~ ~ c c i c i 1ork ( k ~ l r ~ ~ i i l ~ i ~crsit) Irci t ~ i i

1liIl(r Ilt 13 1981 H lt I 11lt11trr1rk~i iot l r ilt I I I ~ It11c g c o g r ~ ~ ) l r i ( ~ 01 f1orrI lrittio~r i l l i(if1111~1i 3 5 ~~I II I~~(I I~~II o~ 7(i 774

hIr111~ 1 19 16 (111iI(~gi~1tlic crrIti ~ t c c l 111cl i I (1 c o I r r ~ l i l ) i ~ i c ~ ~ HIgt~I(II I~III I gtO(OIIIgt 7

X i I lt o ~ r ~ 111~ c o l ~ ~ t i o ~ i (Ice)I ICl88 of llo(ri i t11

(~orolliltill)( Ylill~ IO 334 147 I 19 OI~irht( ~clk i ~ h l i ( ~ l ~ ~ ( ~ l s ~ Ilt h l k j 11 H J S c o ~ t ~

Irl~iic~r l)Cl3 h l o ~ i o l ) l ~ ) l ~tlic s t (~ricl~rc~ 01 IIIO iOc~iti- (I( 11ioti 01 t l ~ c i r r111jor I i~ici tg(~s i~i l t~rrccl IIo11i 1 ) S j ~ ~ ( I L I ( I I C V ~ l t i t i IltII ((11(1 8 0 ~700 722 or ~11i1 t o ( ~ i

11 I L ~ ~ I O I l)(j5 t o g c ~ i c i ( ~( ht110ich011 tlrv ~ ( I I L I l(ii~qi(~

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S I I I ( I ( ~gtOII hl J amp h l J ICI)4 SliilihL ) o t i o g l ~ ~ ~ c ~ it1 (liv~tili(1tio11 r ~ t c tlic oriKiti o l ~ t ~ g i o s ~ ) ( ~ t ~ ~ ~ s it11 s S c ~ r ~ i i ~

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t r ~ i t s 111( ( I L I S ( ~ I 1111l)liIicc1(licriIi( ~tiotl I I ~ ili111ro1(1

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Stcitrcr li 1 amp Ylii~c~lrcicl I ) I)IO I)olliti~lor i(Iilgtt~itioti10 l lo ~ th ~ ~ ~ ( I I ~ I I ~ ~I ) I ( I ( I I I oiIL~c(~r(~t i t1g I ~ I ( I c~ol t i to~t44 170 1 1707

S t ~ i t ~ ( r ~K I amp V l ~ i t c l ~ c ~ ~ c l ~ l))l L I I ( I oil 1) Oil l l o ( ~ t ~ l ) ( ~ gt 1111 tli(r c~iclcticc li)r l )ol l i t i ~tor ~ I ~ L ~ I I I ~ ~ I I I ~ f i ~ o ~ i i i ~ i t

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RI itc11 22 l i i q i~ I)) 4 SI~I~II~II Ij 1~~~1~~1011 lO)T

You have printed the following article

Spurring Plant Diversification Are Floral Nectar Spurs a Key InnovationScott A Hodges Michael L ArnoldProceedings Biological Sciences Vol 262 No 1365 (Dec 22 1995) pp 343-348Stable URLhttplinksjstororgsicisici=0962-84522819951222292623A13653C3433ASPDAFN3E20CO3B2-P

This article references the following linked citations If you are trying to access articles from anoff-campus location you may be required to first logon via your library web site to access JSTOR Pleasevisit your librarys website or contact a librarian to learn about options for remote access to JSTOR

References

Phylogenetics of Seed Plants An Analysis of Nucleotide Sequences from the Plastid Gene rbcLMark W Chase Douglas E Soltis Richard G Olmstead David Morgan Donald H Les Brent DMishler Melvin R Duvall Robert A Price Harold G Hills Yin-Long Qiu Kathleen A KronJeffrey H Rettig Elena Conti Jeffrey D Palmer James R Manhart Kenneth J Sytsma Helen JMichaels W John Kress Kenneth G Karol W Dennis Clark Mikael Hedren Brandon S GautRobert K Jansen Ki-Joong Kim Charles F Wimpee James F Smith Glenn R Furnier Steven HStrauss Qui-Yun Xiang Gregory M Plunkett Pamela S Soltis Susan M Swensen Stephen EWilliams Paul A Gadek Christopher J Quinn Luis E Eguiarte Edward Golenberg Gerald HLearn Jr Sean W Graham Spencer C H Barrett Selvadurai Dayanandan Victor A AlbertAnnals of the Missouri Botanical Garden Vol 80 No 3 (1993) pp 528-548+550-580Stable URLhttplinksjstororgsicisici=0026-649328199329803A33C5283APOSPAA3E20CO3B2-W

Columbines A Geographically Widespread Species FlockScott A Hodges Michael L ArnoldProceedings of the National Academy of Sciences of the United States of America Vol 91 No 11(May 24 1994) pp 5129-5132Stable URLhttplinksjstororgsicisici=0027-8424281994052429913A113C51293ACAGWSF3E20CO3B2-5

httpwwwjstororg

LINKED CITATIONS- Page 1 of 3 -

R E F E R E N C E S

i s o l ~ t i o ~ i lqIfq(~ ~ l i c lI I ~ I I~ ~I I I I I ~ ~ I I gt I I l ) ( ~ c c ~ i IIIIHOII

1)111 J I O I I I ( ill Sri 1TLY49172 193 2410 Hoot S 13 1991 1 11~Ioqc1 i~ I)rs(~l1111oIt11c ~ lt I I I I I I ~ ( ~ ~ I ~ I ( ~ ~ I c

c ~ l ) i c I ( ~ ~ ~ ~ i i c r o ( ~ I r ~ ~ ~ ~ ~ ( ~ t c r h Sy 11itI IIIO ~ i i ~ ~ ~ r o ~ r r o r ~ gt I i o l o ~ ~ lo 16 741 755

11001 S 13 lCl)r) 111sccI o ~ i llr) log(~i oftli( K ~ I ~ ~ I I ~ ( ~ I I ~ I ( ~ ( ~ ~ I ~ 11il3 IIIII 11i(1 18s ril)o~or111lDN se(l11(11((( I I ~ lt I 11(11 S col I l l till 1gtscs1

Ii(~iiIlt 1 1)73 k ~ ~ o l ~ ~ ~ i o ~ i ~ r gt ~ ~ ~ i o s ~ ) l i o l o ~ i ~ ~ i ~ lst1 tt(gics t t i c I

i ~ i ~ i o ~ ~ l i o ~ i h ci(~Irli0 l)Iiary~ig(~tl jth Sq Zoo 2 2 ~ 421 111

1 l t l ) l ) ( 1 lc L)J 1))3 TI~III~I-~IIIIL t 11i ( rs i~ i ~~ l i l~ r ic lgc 11 (as

h l a gt r 1 1912 S gt ~ s t ( ~ ~ ~ i t ~ i ( ~ i St t ~ i c l tlrc o r ig i~r o l ~ s ~ ~ c c i c i 1ork ( k ~ l r ~ ~ i i l ~ i ~crsit) Irci t ~ i i

1liIl(r Ilt 13 1981 H lt I 11lt11trr1rk~i iot l r ilt I I I ~ It11c g c o g r ~ ~ ) l r i ( ~ 01 f1orrI lrittio~r i l l i(if1111~1i 3 5 ~~I II I~~(I I~~II o~ 7(i 774

hIr111~ 1 19 16 (111iI(~gi~1tlic crrIti ~ t c c l 111cl i I (1 c o I r r ~ l i l ) i ~ i c ~ ~ HIgt~I(II I~III I gtO(OIIIgt 7

X i I lt o ~ r ~ 111~ c o l ~ ~ t i o ~ i (Ice)I ICl88 of llo(ri i t11

(~orolliltill)( Ylill~ IO 334 147 I 19 OI~irht( ~clk i ~ h l i ( ~ l ~ ~ ( ~ l s ~ Ilt h l k j 11 H J S c o ~ t ~

Irl~iic~r l)Cl3 h l o ~ i o l ) l ~ ) l ~tlic s t (~ricl~rc~ 01 IIIO iOc~iti- (I( 11ioti 01 t l ~ c i r r111jor I i~ici tg(~s i~i l t~rrccl IIo11i 1 ) S j ~ ~ ( I L I ( I I C V ~ l t i t i IltII ((11(1 8 0 ~700 722 or ~11i1 t o ( ~ i

11 I L ~ ~ I O I l)(j5 t o g c ~ i c i ( ~( ht110ich011 tlrv ~ ( I I L I l(ii~qi(~

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You have printed the following article

Spurring Plant Diversification Are Floral Nectar Spurs a Key InnovationScott A Hodges Michael L ArnoldProceedings Biological Sciences Vol 262 No 1365 (Dec 22 1995) pp 343-348Stable URLhttplinksjstororgsicisici=0962-84522819951222292623A13653C3433ASPDAFN3E20CO3B2-P

This article references the following linked citations If you are trying to access articles from anoff-campus location you may be required to first logon via your library web site to access JSTOR Pleasevisit your librarys website or contact a librarian to learn about options for remote access to JSTOR

References

Phylogenetics of Seed Plants An Analysis of Nucleotide Sequences from the Plastid Gene rbcLMark W Chase Douglas E Soltis Richard G Olmstead David Morgan Donald H Les Brent DMishler Melvin R Duvall Robert A Price Harold G Hills Yin-Long Qiu Kathleen A KronJeffrey H Rettig Elena Conti Jeffrey D Palmer James R Manhart Kenneth J Sytsma Helen JMichaels W John Kress Kenneth G Karol W Dennis Clark Mikael Hedren Brandon S GautRobert K Jansen Ki-Joong Kim Charles F Wimpee James F Smith Glenn R Furnier Steven HStrauss Qui-Yun Xiang Gregory M Plunkett Pamela S Soltis Susan M Swensen Stephen EWilliams Paul A Gadek Christopher J Quinn Luis E Eguiarte Edward Golenberg Gerald HLearn Jr Sean W Graham Spencer C H Barrett Selvadurai Dayanandan Victor A AlbertAnnals of the Missouri Botanical Garden Vol 80 No 3 (1993) pp 528-548+550-580Stable URLhttplinksjstororgsicisici=0026-649328199329803A33C5283APOSPAA3E20CO3B2-W

Columbines A Geographically Widespread Species FlockScott A Hodges Michael L ArnoldProceedings of the National Academy of Sciences of the United States of America Vol 91 No 11(May 24 1994) pp 5129-5132Stable URLhttplinksjstororgsicisici=0027-8424281994052429913A113C51293ACAGWSF3E20CO3B2-5

httpwwwjstororg

LINKED CITATIONS- Page 1 of 3 -

$IIII i ~ ~ c l i ( ~ ( c 11101io1gtli)l~0 1 I I I L I ( I I ~ oil I I I I I I~S I I I I I ~ I O I I I ( I ( ( I ) 8 0 ~(SHi ( ltI9

S I I I ( I ( ~gtOII hl J amp h l J ICI)4 SliilihL ) o t i o g l ~ ~ ~ c ~ it1 (liv~tili(1tio11 r ~ t c tlic oriKiti o l ~ t ~ g i o s ~ ) ( ~ t ~ ~ ~ s it11 s S c ~ r ~ i i ~

I I I 2 6 4 ~I I0 I Ilt Si11111soli( ( I Igtlt T I t t i i l i i i Y~ I I I 01 I ~ o I I o I ~ SC

York ( ~ I I ~ I I I ~ I ~ I I t l i crsit IIIS S l o i ~ i k i ~ J amp (u(r7 ( 198lt1 l(stitly t11c gtto( l i ~s t i~ i ( 1)

01 ~ gt ~ t ( c r t i s org~trisrnal tlictsit i ~ ~ ~ p t o ( c l01 i l l 11~111

111oc1cl I I I I V I 1 3 4 ~I07 )21 SIo i t~ski I 13 ampk ( ~ t gt ( r ~ llt1lt13 1~c(i11g cc r t ~ i t l(1 lt l ~ c ~ I i ~ r

t r ~ i t s 111( ( I L I S ( ~ I 1111l)liIicc1(licriIi( ~tiotl I I ~ ili111ro1(1

1111~t1io(lI)is(cl 011 lt I 1i1oclcl 01 L I I ( Ir1110o111 s p c ( i ~ t i o ~ i c x t i t i ~ ~ t i o t ~ I t t i (I 142 101lt1 102 I

Stcitrcr li 1 amp Ylii~c~lrcicl I ) I)IO I)olliti~lor i(Iilgtt~itioti10 l lo ~ th ~ ~ ~ ( I I ~ I I ~ ~I ) I ( I ( I I I oiIL~c(~r(~t i t1g I ~ I ( I c~ol t i to~t44 170 1 1707

S t ~ i t ~ ( r ~K I amp V l ~ i t c l ~ c ~ ~ c l ~ l))l L I I ( I oil 1) Oil l l o ( ~ t ~ l ) ( ~ gt 1111 tli(r c~iclcticc li)r l )ol l i t i ~tor ~ I ~ L ~ I I I ~ ~ I I I ~ f i ~ o ~ i i i ~ i t

45gt I ampIlt I gtoI S oI1i)rO 1) 1 1)lt13 IgtLr I ~ I I ( I ~ t~iriyI i t ~ o q ~ ~ ~ t ~ ~ t i ~ I

I ( L I I ~ ~ I O ~ I I l l l i t~ois l l l i t io i~ S I IL I~ I~( ~ I i ~ i r ~ ~ l gt i i i g t i ~ H i t o ~ gt c )

RI itc11 22 l i i q i~ I)) 4 SI~I~II~II Ij 1~~~1~~1011 lO)T

You have printed the following article

Spurring Plant Diversification Are Floral Nectar Spurs a Key InnovationScott A Hodges Michael L ArnoldProceedings Biological Sciences Vol 262 No 1365 (Dec 22 1995) pp 343-348Stable URLhttplinksjstororgsicisici=0962-84522819951222292623A13653C3433ASPDAFN3E20CO3B2-P

This article references the following linked citations If you are trying to access articles from anoff-campus location you may be required to first logon via your library web site to access JSTOR Pleasevisit your librarys website or contact a librarian to learn about options for remote access to JSTOR

References

Phylogenetics of Seed Plants An Analysis of Nucleotide Sequences from the Plastid Gene rbcLMark W Chase Douglas E Soltis Richard G Olmstead David Morgan Donald H Les Brent DMishler Melvin R Duvall Robert A Price Harold G Hills Yin-Long Qiu Kathleen A KronJeffrey H Rettig Elena Conti Jeffrey D Palmer James R Manhart Kenneth J Sytsma Helen JMichaels W John Kress Kenneth G Karol W Dennis Clark Mikael Hedren Brandon S GautRobert K Jansen Ki-Joong Kim Charles F Wimpee James F Smith Glenn R Furnier Steven HStrauss Qui-Yun Xiang Gregory M Plunkett Pamela S Soltis Susan M Swensen Stephen EWilliams Paul A Gadek Christopher J Quinn Luis E Eguiarte Edward Golenberg Gerald HLearn Jr Sean W Graham Spencer C H Barrett Selvadurai Dayanandan Victor A AlbertAnnals of the Missouri Botanical Garden Vol 80 No 3 (1993) pp 528-548+550-580Stable URLhttplinksjstororgsicisici=0026-649328199329803A33C5283APOSPAA3E20CO3B2-W

Columbines A Geographically Widespread Species FlockScott A Hodges Michael L ArnoldProceedings of the National Academy of Sciences of the United States of America Vol 91 No 11(May 24 1994) pp 5129-5132Stable URLhttplinksjstororgsicisici=0027-8424281994052429913A113C51293ACAGWSF3E20CO3B2-5

httpwwwjstororg

LINKED CITATIONS- Page 1 of 3 -

You have printed the following article

Spurring Plant Diversification Are Floral Nectar Spurs a Key InnovationScott A Hodges Michael L ArnoldProceedings Biological Sciences Vol 262 No 1365 (Dec 22 1995) pp 343-348Stable URLhttplinksjstororgsicisici=0962-84522819951222292623A13653C3433ASPDAFN3E20CO3B2-P

This article references the following linked citations If you are trying to access articles from anoff-campus location you may be required to first logon via your library web site to access JSTOR Pleasevisit your librarys website or contact a librarian to learn about options for remote access to JSTOR

References

Phylogenetics of Seed Plants An Analysis of Nucleotide Sequences from the Plastid Gene rbcLMark W Chase Douglas E Soltis Richard G Olmstead David Morgan Donald H Les Brent DMishler Melvin R Duvall Robert A Price Harold G Hills Yin-Long Qiu Kathleen A KronJeffrey H Rettig Elena Conti Jeffrey D Palmer James R Manhart Kenneth J Sytsma Helen JMichaels W John Kress Kenneth G Karol W Dennis Clark Mikael Hedren Brandon S GautRobert K Jansen Ki-Joong Kim Charles F Wimpee James F Smith Glenn R Furnier Steven HStrauss Qui-Yun Xiang Gregory M Plunkett Pamela S Soltis Susan M Swensen Stephen EWilliams Paul A Gadek Christopher J Quinn Luis E Eguiarte Edward Golenberg Gerald HLearn Jr Sean W Graham Spencer C H Barrett Selvadurai Dayanandan Victor A AlbertAnnals of the Missouri Botanical Garden Vol 80 No 3 (1993) pp 528-548+550-580Stable URLhttplinksjstororgsicisici=0026-649328199329803A33C5283APOSPAA3E20CO3B2-W

Columbines A Geographically Widespread Species FlockScott A Hodges Michael L ArnoldProceedings of the National Academy of Sciences of the United States of America Vol 91 No 11(May 24 1994) pp 5129-5132Stable URLhttplinksjstororgsicisici=0027-8424281994052429913A113C51293ACAGWSF3E20CO3B2-5

httpwwwjstororg

LINKED CITATIONS- Page 1 of 3 -

Floral and Ecological Isolation Between Aquilegia formosa and Aquilegia pubescensScott A Hodges Michael L ArnoldProceedings of the National Academy of Sciences of the United States of America Vol 91 No 7(Mar 29 1994) pp 2493-2496Stable URLhttplinksjstororgsicisici=0027-8424281994032929913A73C24933AFAEIBA3E20CO3B2-P

Phylogeny of the Ranunculaceae Based on Epidermal Microcharacters and MacromorphologySara B HootSystematic Botany Vol 16 No 4 (Oct - Dec 1991) pp 741-755Stable URLhttplinksjstororgsicisici=0363-6445281991102F1229163A43C7413APOTRBO3E20CO3B2-5

Evolutionary Strategies and Morphological Innovations Cichlid Pharyngeal JawsKarel F LiemSystematic Zoology Vol 22 No 4 (Dec 1973) pp 425-441Stable URLhttplinksjstororgsicisici=0039-79892819731229223A43C4253AESAMIC3E20CO3B2-0

Hawkmoths and the Geographic Patterns of Floral Variation in Aquilegia caeruleaRussell B MillerEvolution Vol 35 No 4 (Jul 1981) pp 763-774Stable URLhttplinksjstororgsicisici=0014-38202819810729353A43C7633AHATGPO3E20CO3B2-23

A Parsimony Analysis of the Asteridae Sensu Lato Based on rbcL SequencesRichard G Olmstead Birgitta Bremer Kathy M Scott Jeffrey D PalmerAnnals of the Missouri Botanical Garden Vol 80 No 3 (1993) pp 700-722Stable URLhttplinksjstororgsicisici=0026-649328199329803A33C7003AAPAOTA3E20CO3B2-L

Phylogenetic Relationships of the Geraniaceae and Geraniales from rbcL SequenceComparisonsRobert A Price Jeffrey D PalmerAnnals of the Missouri Botanical Garden Vol 80 No 3 (1993) pp 661-671Stable URLhttplinksjstororgsicisici=0026-649328199329803A33C6613APROTGA3E20CO3B2-3

httpwwwjstororg

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Nucleotide Sequences of the rbcL Gene Indicate Monophyly of Mustard Oil PlantsJames Rodman Robert A Price Kenneth Karol Elena Conti Kenneth J Systma Jeffrey D PalmerAnnals of the Missouri Botanical Garden Vol 80 No 3 (1993) pp 686-699Stable URLhttplinksjstororgsicisici=0026-649328199329803A33C6863ANSOTRG3E20CO3B2-23

Testing the Stochasticity of Patterns of Organismal Diversity An Improved Null ModelJoseph B Slowinski Craig GuyerThe American Naturalist Vol 134 No 6 (Dec 1989) pp 907-921Stable URLhttplinksjstororgsicisici=0003-014728198912291343A63C9073ATTSOPO3E20CO3B2-8

Testing Whether Certain Traits have Caused Amplified Diversification An Improved MethodBased on a Model of Random Speciation and ExtinctionJoseph B Slowinski Craig GuyerThe American Naturalist Vol 142 No 6 (Dec 1993) pp 1019-1024Stable URLhttplinksjstororgsicisici=0003-014728199312291423A63C10193ATWCTHC3E20CO3B2-R

Oil Flowers and Oil Bees Further Evidence for Pollinator AdaptationKim E Steiner V B WhiteheadEvolution Vol 45 No 6 (Sep 1991) pp 1493-1501Stable URLhttplinksjstororgsicisici=0014-38202819910929453A63C14933AOFAOBF3E20CO3B2-V

httpwwwjstororg

LINKED CITATIONS- Page 3 of 3 -