Phylogeny of Hepatica (Ranunculaceae) and Origin of Hepatica Maxima Nakai Endemic to Ullung Island, Korea

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Phylogeny of Hepatica (Ranunculaceae) and origin of Hepatica maxima Nakai endemic to Ullung Island, Korea

Article in Biological Journal of the Linnean Society · January 2011

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Phylogeny of Hepatica (Ranunculaceae) and origin of Hepatica maxima NAKAI endemic to Ullung Island, Korea

M. PFOSSER*, B.-Y. SUN, T. F. STUESSY, CH.-G. JANG, Y.-P. GUO, K. TAEJIN, K. C. HWAN, H. KATO, & T. SUGAWARA

Abstract: Nuclear ITS, chloroplast atpB-rbcL spacer and trnL-F intron and spacer sequences, as well as [<C[QP"<<< Hepatica¤YCRP endemic H. maxima[T[<<<Hepatica 'Anemone C`P"<'HepaticaX'"< CPtrnL-F #"6+<H. asiatica, H. henryi, H. insularis, and H. maxima#H. asiatica and H. insularis <'[< H. maximaH. asiaticaR' !R[Q[H. maxima'H. asiatica R'<^CH. nobilisP 'H. asiatica and H. insularis'<[Q' <<<< <

Zusammenfassung7Y!"atpB-rbcL-IGS und trnL-F&YY:!&!" '[<C[QPT'< <:Hepatica[H. maxima, einem Endemiten Y¤Y"'T[&!&= =<:Hepatica<'[ :Anemone'HepaticaC`P" T:Hepatica['trnL-F !<Y=&@[T '6+<H. asiatica, H. henryi, H. insularis und H. maxima H. asiatica und H. insularis X<" [TT[ H. maximaH. asiatica- R'![Q&T 'H. maxima-H. asiatica&Q '' CH. nobilisP RT'H. asiatica und H. insularis- [Q&T'[' T&X<'@ &!

Key words7 [Q <" Hepatica" " Y! " T[ " plastid microsatellites, speciation.

16 STAPFIA: reports © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at PFOSSER 'L!Hepatica maxima STAPFIA 95 (2011): 16–27

H. nobilis H. asiatica H. transsilvanica H. insularis H. maxima H. americana H. falconeri H. acutiloba H. henryi H. nobilis var. japonica var. pubescens

Manchuria

A11-12 East Sea

Korea A10 A9 Ullung Island A8 M1-7 A7 Japan A4-6 A1-2 A3 P1-3 J3 J1-2 I11 I12 I15 P4-5 I14 I13

I1-I10

200 km

Fig. 1]UK!Hepatica'QU^p!N" H. asiatica (A1-12), H. insularis$::D;UH. maxima (M1-7), H. nobilisI'japonica (J1-3), H. nobilis I'pubescens$A:D;'

Introduction @< < <" X'- Y < < YCBALDWIN & SANDERSON6**BARRIER6***P" 7 < - ^QYCSTUESSY & ONO6**P"<Y '["'< CFRANCISCO-ORTEGA 4556 RAY 6**P" = ""< CFRANCISCO-ORTEGA6***VARGAS6***P << <'- CEMERSON4554GRANT6**STUESSY & ONO6** WHITTAKER6**P! R¤&CPFOSSER4554P< <<CCRAWFORD & - STUESSY6**`P- <' " ' & < CSUN & STUESSY6**P

STAPFIA: reports 17 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at PFOSSER 'L!Hepatica maxima STAPFIA 95 (2011): 16–27

¤&6` < R55'^CQ6P"- AnemoneCEHRENDORFER455*EHREN- ` 4`)` )65 +`)65 _) DORFER & SAMUEL4556HOOT6**+MEYER4565 `2 SCHUETTPELZ4554P<<- *+ strate that Hepatica<Anemone that x`"< CKIM 6*" P & [¤ [ 4` - Island endemic H. maxima. <"' <Y!" <' plastid trnL-F intron and spacer, and atpB-rbcL spacer sequences K 6<C IM6*P[<- <<Hepatica worldwide. To "'"<[ <" <¤&"- <[[ R'^ <C[QP YANG <®<C 6**_P< <'¤Y - ^R [`55¤Y Materials and Methods `"4+ CLEE & YANG6*6SUN & STUESSY6**P- Taxa Sampled<Hepatica '¤Y R"¤&CRP" Y RC*+P"^C_P= CRP""["¤!["^6***&4556 C* CKIM 6*PP Q [CH. transsilvanica, H. henryiP- < #:P 'Acer, A. okamotoa- " " num and A. takesimense"<'CPFOSSER et =#[- 4554PA. mono and A. ¤' pseudosieboldianum"<"R [¤Y"H. maxima" DNA ExtractionT[' &' DOYLE DOYLE Hepatica MILLERC@P & C6*`P'['- CMELCHIOR6*_+P"CTAMURA6**P"CWEISS '`553[#C4 4554P""- [#"655="6+="45=T["54- " [ "X5P5_5 553 [CQ6P- <C4+6P'' [['' 6+ ["T[ ¤H. maxima. NAKAICNA- ''553'' KAI6*`NAKAI6*4PR' '`56553 "H. asiatica and H. insularis' DNA Sequencing & - ¤Y"H. maxima,'' 'trnL(UAA) intron and the H. nobilis"japonicapu- CY:!P ' trnL(UAA)-) bescens^< the trnF(GAA)'[@ CHIROE6*`P# H. henryiH. CPFOSSER & SPETA 6***P atpB-rbcL transsilvanica CP [ '[4MANEN et al. [(H. acutiloba and H. americana; C6**+PY!'[' STEYERMARK & STEYERMARK6*_5P) and the threatened species Y!6&6 C)&M)P Y!+ C)&- H. falconeri'R M)P[[&T[' ''Hepatica, "' <[#Y``C 'CJALAS & SUOMINEN "¤RP'T<- 6**P[ H. maxima' C["¤![P#T< "<"- <!@<@RC[#<" "H. asiatica- ^P"CPFOSSER & SPETA RCSUN & STUESSY6**P'"R 6***P [ ' - endemic, H. insularis Y Y!4 C)&&)P Y! R" < < C)&&)P' related to H. asiatica CSUN & STUESSY6**WOO4554P Y!"6 #- ' H. asiaticaCLEE6*`_PARK6*`+P < Phylogenetic Analysis!' Hepatica['- T[6555[+55<

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57 Hepatica asiatica A13 Hepatica henryi 1 Hepatica insularis I16 H. nobilis var. japonica J4 H. nobilis var. pubescens P1 Hepatica acutiloba 3 Hepatica americana Hepatica nobilis 8 98 Hepatica transsilvanica 4 99 Anemone narcissiflora 2 100 Anemone narcissiflora 3 Anemone obtusiloba 100 Anemone flaccida 70 Anemone keiskeana 93 100 Anemone antucensis Anemone tenuicaulis 1 Anemone canadensis 100 Clematis afoliata 74 Clematis hexapetala Knowltonia vesicatoria 63 Pulsatilla patens 100 Pulsatilla montana 97 67 Pulsatilla koreana Pulsatilla occidentalis Anemone raddeana 100 66 Anemone nemorosa 1 100 Anemone sylvestris 1 74 Anemone blanda 1 Anemone drummondii Anemone rivularis 1 Ranunculus ficaria 98 Trautvetteria carolinensis 100 Trautvetteria grandis Trautvetteria japonica 5 changes

Fig. 2Y"He- patica'!Anemone, Clematis, PulsatillaUKnowltonia' !TrautvetteriaRanunculus'QI$:/U///;!D/ I'!'I! x = %x = _!]- !!'p!H. maxima'

T¤+5TT['& +5+CSWOFFORD4555P=' ¤::'C:& < 6"555 - :6**+P- &C#@P'Q- QT[' CFITCH6*`6P!'' < '' < < [=# '& C[6P<<- CFELSENSTEIN6*P'[ <C=P& C^P Y CP ''[¤ the trnL-F'"

STAPFIA: reports 19 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at PFOSSER 'L!Hepatica maxima STAPFIA 95 (2011): 16–27

66 Hepatica asiatica A1a Hepatica henryi 1 Hepatica insularis I2 Hepatica americana 100 Hepatica acutiloba 2 Hepatica nobilis 3 Hepatica transsilvanica 1 64 H. nobilis var. japonica J1 H. nobilis var. pubescens P1a 74 96 Anemone narcissiflora 2 99 Anemone narcissiflora 1 55 Anemone obtusiloba 100 Anemone keiskeana

90 82 Anemone flaccida 92 Anemone tenuicaulis 2 97 Anemone antucensis 87 Anemone tenuicaulis 1 Anemone canadensis 1 Clematis hexapetala 62 Anemone x intermedia 78 Anemone sylvestris 2 73 Anemone coronaria Anemone ranunculoides Anemone nemorosa 2

61 98 Anemone pavonina Anemone hortensis 97 Anemone blanda 2 97 Anemone blanda 1 Anemone palmata 61 Anemone drummondii 92 Anemone rivularis 2 54 94 Anemone rivularis 1 Anemone hupehensis 78 98 Pulsatilla occidentalis Pulsatilla grandis Knowltonia vesicatoria Ranunculus ficaria 5 changes

Fig. 3YatpB-rbcL- Hepatica'!Anemone, Clematis, PulsatillaUKnowltonia' 'QI$:/U///;!D/ I'!'I! x = %x = _!]!- !'p!H. maxima'

C' AFLP Analysis.[Q<' P<'- ' [Q < C " Q "¤RP'[455T[' C<&<&[P'trnL-F ' < Eco@Y 66*&64*" _+&_*4" `_5& MseY3[C3P' ``6" 654_&65+ ' ' ' ' ` [53T['& ' [T'<CX#[YTP45 =5_CMADDISON & MADDISON6**4P <45*+ "+5_ "5`4

-/ STAPFIA: reports © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at PFOSSER 'L!Hepatica maxima STAPFIA 95 (2011): 16–27

H. acutiloba 1-2

H. asiatica A3, A10-12

H. insularis I1-4

H. transsilvanica 1-3 H. insularis I5-I15 67 H. asiatica A1-2, A4-6, A8

H. henryi 1-4 93 H. asiatica A7, A9

H. nobilis var. japonica J1-3

63 H. nobilis var. pubescens P1-5 H. nobilis 1-7

100 Anemone narcissiflora 1

Anemone canadensis 2 Clematis afoliata

95 Anemone palmata Anemone hortensis

Anemone coronaria

Fig. 4 " Y trnL-F - 'Anemone'x = _'Q I$:/U///;!D/I'p!H. maxima '

@'65& Results [ ' Eco@Y MseI Q&Eco@Y'- Nuclear ITS sequences [Q6_ " @ ' <C"- <"P Y! ' +6+ CY!67 4_5 Y!47 6+P 7 Eco&[[Mse&[" Eco&[:Mse&:" <&'< Eco&[[:Mse&!['- (Anemone, Clematis, Hepatica, Knowlto- '<['45*+ " nia, Pulsatilla, Ranunculus, and Trautvetteria)=- 45_ "5`4 '<64<'- <<<< <5` <T _`*C<Y5_6`"@Y 4<'_ 5`_"<@5+`+P!<- !['_- C 4`44+" Y 5`+" @Y 5_`" @ <<'C:- 5_5P"''Q4"'- 55@"[#<P[#Y`` 'Y' 5&55 Ranunculus or Trautvetteria as '<<O5P- =<<Hepatica<C* O6P P[<" 56'< Hepatica and Anemone spp. with [@®¤Y66C!6**`P& - x`<-

<'FST'- < Clematis and Pulsatilla < '' C655*`"

STAPFIA: reports 21 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at PFOSSER 'L!Hepatica maxima STAPFIA 95 (2011): 16–27

I1 51 I2 I3 I4 85 A10a A10b A11 76 78 A12 A3a A3b 2 98 3 4 1 A2 A1a A1b A6 A8 A5a A5b A4 KOREA I11 I9 I14 I10 I15 I12 75 84 A9 A7 84 I13 79 I6 100 I5 I7 I8 P1b P1c P2 P3 P4 P5a P5b JAPAN 98 P1a 89 J2 86 J1 J3a J3b 58 98 1 2 N AMERICA 86 1 2 3 4 3 EUROPE 5 6 7 2 1

0.1 changes

Fig. 5A!\]$UU;!!- Hepatica'QI$:/U///;!D/Y 'K!!!!'

22 STAPFIA: reports © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at PFOSSER 'L!Hepatica maxima STAPFIA 95 (2011): 16–27

I6 (4) I7 (8) I15 (4) I12 (4) I10 (10) I14 (4) I9 (7) I13 (4) I5 (4) I8 (7) I11 (5) A1 (10) A6 (5) A2 (10) A8 (4) A5 (9) KOREA A7 (6) A9 (5) A10 (5) I3 (9) A3 (8) I4 (10) A4 (4) I1 (5) I2 (5) P2 (5) P5 (6) P3 (10) P1 (8) P4 (7) JAPAN J2 (10) J3 (8) J1 (5) 2 (8) N AMERICA 4 (10) 3 (9) EUROPE 1 (10) 2 (7)

2.0

Fig. 6!!H. asiatica, H. insularis, H. maxima, H. nobilis

I'pubescens, H. nobilisI'japonica, H. acutilobaUH. nobilisF"I ]^A'[! !':Q'I#'

Chloroplast atpB-rbcL spacer sequences C655P"<'- CQP= [atpB-rbcL<65`*"6`` [ " H. maxima, H. asiatica, H. ''<'<&<- insularis, and H. henryi,C__P!Hepatica ''Q** Anemone spp. with x = `C`+ CY 5`" @Y 55" @ 5**P ! ' P!ClematisC*5P" <='`_`CY 'Anemone<- 5*64"@Y5**"@5_PHepatica<<- 'Pulsatilla and KnowltoniaC*P

STAPFIA: reports 23 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at PFOSSER 'L!Hepatica maxima STAPFIA 95 (2011): 16–27

Chloroplast trnL-F intron and spacer sequences asiatica and H. insularis[^ <[Q' trnL-FT[66+*"'66 '[H. acutiloba. `<&& <46'- <CY5*+"@Y55"@544P!- '< Discussion 4''6*+5CY5*`_"@Y 5*"@5*4PCQ+P < ' @ [- <<Hepatica clade < C*P[[[ < CEHRENDORFER 6** STARODUBTSEV 6**6 as well as H. transsilvanica ULBRICH 6*5_P @<" < C_PH. nobilis CEHRENDORFER & SAMUEL4556HOOT6**HOOT et Sister to Hepatica are and A. canadensis 6**+ SCHUETTPELZ 4554P ' 'x = `! T [ x = `ClematisC655P"'< Anemone HOOT C6**+P HOOT C6**P Anemone'x = " Hepatica, the trnL intron and the trnL-F"<- Knowltonia, and Pulsatilla with Anemone, EHRENDORFERC6**P CYTP<CYT6"YT444"YT_ ' < "YT++"YT6+"YT_4P"' - <&C66*&64*"`_5& Anemone ``6"654_&65+P<&[C x = `x = Y _+&_*4P & < - <"HOOTC6**PSCHUETTPELZ et al. < C4554P[[ within Hepatica CQ P[ T[ " Anemone H. maxima ' << <C+PH. asiatica popu- 'Clematis was used as an [`[*R' Trautvetteria !RC'Q6#PH. @ asiatica and H. insularis ' than Clematis[<" T[Y" Anemonex <'#^- = `Trautvetteria or Ranunculus was used pan, the diploid H. nobilisjaponicaH. Clematis, nobilispubescens'- ' C*P['^- ''atpB-rbcL [H. acutiloba and the tetraploid H. <"'Clematis occurred as sister to Hepatica transsilvanica'<'<- and x = `AnemoneC*5P"'x CPH. nobilis appeared = Anemone'<<x <Hepatica. = `Anemonex = <&<CKURITA6*P¤ AFLP analysis Clematis'- '" HOOT Y"6_'+&65" 'x = `C et al. +H. nobilis"6H. acu- 6**+P"" tiloba"H. nobilisjaponica" <<<Hepatica within H. nobilis pubescens" _ H. maxima" 65 [ < @ X'" H. asiatica"6H. insularis '[<Y! ' < ' [Q [ &T[- 4*`<"4*C5P' '" <@<[ <46C5`*P4_+4'" C< 6 C5_P ' ' < ' He- P'H. nobilis C` patica[@ <4*P"H. acutilobaC- CSOLTIS6***P"<- P"^C+6_`P"H. maxima "" C45*P"R- @<"<< H. asiatica and H. insularisC+6_`P& TrolliusC@P''' < X" DESPRES 'FST[ = " [ [ C et al. H. maximaR- 455P[<"Y!T[ ^CQ_P ' 'H. [Q""

24 STAPFIA: reports © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at PFOSSER 'L!Hepatica maxima STAPFIA 95 (2011): 16–27

<! R[<<<- < < RHepatica<H. insularis CHODKINSON4555KOOPMAN4556ZHANG4556P <CWOO4554P [T[[Q '<C[P <<Hepat- H. insularis populations ica Y[ H. nobilis"'" <[Q"- '[H. asiatica, H. henryi, '[<"<- H. insularis, and H. maxima"^- '"'<" ies H. nobilisjaponica and H. nobilispubescens. The [Q""< [H. acuti- - loba and the Transcarpatian endemic H. transsilvanica" < <Y ''[CH. acutiloba and H. americanaCSTEYERMARK & STEYERMARK"6*_5PP" <' Acknowledgements species. Since our results show that ITS sequences are identical "<<X'- Q[!QCQQ "<'H. no- 64*_&#Y6+4PR!- bilis['[ Q CR!QP < - ' ' ^ H. nobilis<STEYERMARK & STEYERMARKC6*_5P ' < < = X" < ^"'< = ¤<" <" ^ R ' <H. nobilis"< X"Q<#- "H. !"¤<" "R falconeri" R X< =- "< !¤Y- Literature Cited demic H. maxima- R ^7 C6P atpB-rbcL #ALDWIN#:"=^ SANDERSONC6**P7[[ spacer sequences cluster H. maxima with H. asiatica, H. insu- X''CPN laris, and H. henryiC__PC4P- [<!"¤![957*+54&*+5_ rosatellite data place all H. maxima- BARRIER="#: BALDWIN"@X ROBICHAUX"=T PURUGGANAN <' 'H. asiatica R' C6***P7Y[<<7 !RC+PCP <<X''C[P- N=#- <''H. insu- <167665&666 laris C+ P" ' CHOIX&R" KIM"#& SUN, & H. SHINC6**P7<<Dys- C+P[QH. maxima in a clade taenia<[C<[PY!- with H. asiatica and H. insularis'^- NR^<2876*&6+* [¤ CRAWFORDT"STUESSYC6**`P7 YR 4+*&4_`"@ I RAVEN '¤Y WATSUKI"X N<7! RC6`P DESPRES" GIELLY"# REDOUTET" TABERLETC455P7¤[Q <<- "^XC4`P- [' <''¤Y"Acer oka- <N=< motoanum and A. takesimense,< 2776&6*_ R^CPFOSSER DOYLE^^"^ DOYLEC6*`P7[T[ 4554PX'" N<# ^ 19766&6 '<¤Y EHRENDORFERQC6**P7<[- ^ N!<"!974&4* <R EHRENDORFER Q" @ SAMUEL C4556P7 << < Anemone C@- DystaeniaC[P"'<' &[PN[<!39: CCHOI6**P"¤YCD. takesimanaP 4*&5` [^CD. ibukiensisP EHRENDORFER F., ZIMAN S.N., KÖNIG Ch., KEENER C.S., DUTTON B.E. , [ < ' H. TSARENKO O.N., BULAKH E.V., BOSCAIU M., MÉDAIL F. & A. KÄSTNER (2009): maximaHepatica- " < Anemone section AnemoneC@P N#- R"''"H. asiatica ^!<160764&+ and H. insularis"RQ EMERSON#C4554P77<- H. insularis is restricted to the south- N=- <117*6&*__

STAPFIA: reports -D © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at PFOSSER 'L!Hepatica maxima STAPFIA 95 (2011): 16–27

FELSENSTEIN^C6*P7[<7 NAKAIC6*`^HepaticaN^^#<13: N397`&`*6 44`&4+"5&6+ FITCH= C6*`6P7' [ 7 - NAKAIC6*4P7[<RN# [<N!<< !=76&4` 207+5_&+6_ PARK=RC6*`+P7Keys to the herbaceous plants in Korea (Dicotyledo- FRANCISCO-ORTEGA^"^ BARBER"[ SANTOS-GUERRA"@ FEBLES-HER- nae).N!7^ JANSEN (20 NANDEZ"@R 56P7- PFOSSER="^ GUZY-WRÓBELSKA"#& SUN, T. STUESSY, T. SUGAWARA, :C[7[P & N. FUJIIC4554P7AcerC!P- <Y7 ¤Y"R!<#<2776&_` T[ N[ P SPETA (19 ^#<8876_6&6_* FOSSER="Q **P7<X< T[ N[ = # F ORTEGA GOERTZEN SANTOS GUERRA BENABID RANCISCO- ^"@ "[ - "[ , Garden 8674&` JANSEN @ R C6***P7 = !< Asteriscus R C[7 YP Y7 - AY = Q C6**P7 !< Lavatera and Malva C=" T[N!<#< =P&'!<198: 2474+*&4__ 4*& R GENETICS COMPUTER GROUPC6**+P7Program manual for the Wisconsin OHLFQ^C6**`P7NTSYS-PC. Numerical taxonomy and multivariate package, Version 8.N=""¤![ analysis system, Version 2.0.N!"7!' S KUEFFER ROESSLI EXCOFFIER GRANT@C6**P7Evolution on IslandsN7¤- CHNEIDER!"^= "T " C6**`P7AR- < '*"+//14!N¤- <:"!'7:#<< HANG" MCLEWIN"=Q FAYC4556P7=<< S HOOT SAMUEL EHRENDORFER Helleborus C@P ' [& CHUETTPELZ E., S. "@ "Q C4554P7=- = N5076556&656 ! X Anemone C@P H N!< IROE=C6*`P7[<!<Anemone Hepatica 23176+&66 C@P^NBotanical Magazine Tokyo 707+&` SOLTIST"! SOLTIS"=CHASEC6***P7[<- H RENVOICE NI-CHONGHAILE STAPLETON ODKINSON@"![ , G. , = << CHASE ["= C4555P7[Y!T[ NNature 4027+54&+5+ [Q<Phyl- S lostachysC#"PN^@ TARODUBTSEVC6**6P7[7 Systematics and evolutionN 11374*&4_* 7[!!!@ S STEYERMARK HOOT!#C6**P7<AnemoneC@- TEYERMARK ^ [" ! C6*_5P7 Hepatica in North PT[<N [N@62744&44 !<"!974&55 STUESSYQ"=ONO,C6**P7Evolution and speciation of is- HOOT!#"[[ REZNICEK"^T PALMERC6**+P7<- land plantsN7¤ tionships in AnemoneC@P< SUN#&"Q STUESSYC6**P7<- T[N!<#<1976_*&455 ¤Y"R66& STUESSY JALAS^"^SUOMINEN,C6**P7Atlas Florae EuropaeaeNX 202 in Evolution and Speciation of Island PlantsQ = ONON7¤< KIM ^C6*<<¤& S "RN<16764&6 WOFFORDTC4555P7[¤³<<- K <C³P"+N!"=7 IMRC6*P7<¤"R&6 ![ :<N^^[=" T :80764&6 AMURA=C6**P7@_&The Families and Genera of Vascular Plants,R KUBITZKIN#7! KIM RC6*P7<¤"R& U 4<N^ LBRICHC6*5_P7<:- ^[=" :AnemoneN#^- :8074*4&5 !<3776`4&+ KOOPMAN ZEVENBERGEN VAN DEN BERG VARGAS"X[ MCALLISTER" MORTON"! JURY"=^WILKIN- ^="=^ "@: C4556P7 (19 Species relationships in LactucaC"[PN SON **P7<HederaC[P7<- [^#<766&6` Y!N!<21976_&6`* KURITA=C6*P7@Y²- W SUN STUESSY KIM KATO WAKA- '6+&6_& EISS X" #& , T. " X , H. " = species in Anemone and in Ranunculus@#- BAYASHIC4554P7R<<¤Y- Y¤<_76&` CRPR^ N#^!<1387*&65 LEE YANG (19 "Y! 6P7¤YT W Y_6&*! HITTAKER@^C6**P7Island BiogeographyN7¤ and Dogdo Islands, ed. ANONYMOUSN!7R[ WOOX&R"^&X KIM, S.-H. YEAU, & N. S. LEEC4554P7= LEEC6*`_"#N!7!' <RHepaticaC@- < PN!<2367&++ X SOLTIS SOLTIS MADDISON"T@ MADDISONC6**4MacClade: Analysis of phy- IANG®&"T "! C6**P7[ logeny and character evolution. Version 3.0.N!"=- '[ 7 7![ <N=<- 1076`&6*5 MELCHIOR XC6*_+!)%&N#& Y 7:# ANG!C6**_P7A pioneer study on geology of Korean peninsula.N 7R<¤< MEYER K.M., HOOT S.B. & M.T.K. ARROYO (2010): <[- Z COMES KADEREIT (20 ![ Anemone C@P" HANG&#"X "^ 56P7<< "Barneoudia and OreithalesNY- < < Sol- ^!6`674&6 danellaCPY![QN[- ^#<88746&4+

26 STAPFIA: reports © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at PFOSSER 'L!Hepatica maxima STAPFIA 95 (2011): 16–27

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