Delivered by Publishing Technology to: Yale University IP: 130.132.173.179 on: Fri, 02 Aug 2013 21:54:39 Copyright (c) American Society for Plant Taxonomists. All rights reserved. reeadAeod(98 nlddi in De- it characters. Ronse floral included while on (1988) life-form Akeroyd and and anatomy craene petiole of basis the is.Hrlsn(98 xlddi rmher from it excluded (1978) Haraldson cies). ry 98 os ereee l 00.Teei difference of a placement Ak- is the There and regarding 2000). opinion al. Decraene of et to Decraene (Ronse rectangular Ronse cells 1988; mostly eroyd epidermis of and tepal presence venation, the elongated tepal whorl, inner trifid the nectaries, in mostly reduced number ligonum gonella tribe Persicarieae Koenigia enrcgie,adt aeauiu aefrec taxon each for name unique a discussion. have have under that to taxa and inclusive) recognized, possible, (least smallest as been the much for as names usage have standard by to to principally conform guided to been desire have the we confusion. choices and least these names the making of cause In set will hope a we adopted that have circumscriptions we group, this in tionships to reference lato by sensu con- issues are num underlying botanists the most sidestep that to at point tent the and to groups difficult same become different Communication has the 1978). quite (Haraldson and to ranks taxonomic years applied different the been classification over have of proposed names variety been wide have A systems challenge. taxonomic great ered caria stamens lophilon 4–8 and Within 1988). venation; Persicaria Akeroyd and trifid Decraene Ronse with 1978; pectinate (Haraldson tepals or capitate ciliate 4–5 entire, or ochrea; usually spicate 1925a); (Stanford many-flowered plants panicles by These 1993). characterized (Brandbyge temper- regions are northern tropical the into around and distributed ate vines or herbs genus nial a recognized Haraldson Along Persicaria workers, objectives. modern these most achieving to with closest the comes (1978) oyih 08b h mrcnSceyo ln Taxonomists Plant of Society American the by 2008 Copyright © Botany Systematic 1 2 eateto clg n vltoayBooy aeUiest,PO Bo P.O. University, Yale Biology, Evolutionary and Ecology of Department mn oensses h lsiiaino Haraldson of classification the systems, modern Among Polygonum uhrfrcrepnec,peetades a lnkIsiuefrDev for Institute Planck Max address: present correspondence, for Author eadn h lcmn of placement the regarding Within Persicarieae. entire to this related of closely group most and monophyletic is eergos pcfcly efcso nern h eainhp of relationships the inferring to on respect focus with we results Specifically, these regions. test gene we Here segregates. its h hools genes chloroplast the Aconogonon c.6 pce) and species), 60 (ca. Polygoneae Keywords— Abstract— , yhvn odltdflmnossaeswt the with stamens filamentous nondilated having by ) Oxygonum c.1 species), 16 (ca. c.9seis ob lsl eae eeai her in genera related closely be to species) 9 (ca. opie fsm 5 pce fana rperen- or annual of species 150 some of comprised , aado 17)rcgie orsections: four recognized (1978) Haraldson . 20) 31:p.77–86 pp. 33(1): (2008), Persicarieae ntebodLnaa es,hspeetda presented has sense, Linnaean broad the in , ilg,Seantas 73,D706 ubne,Gray(sang-tae. Germany Tuebingen, D-72076, 37-39, Spemannstrasse Biology, natmtn osr u hlgntcrela- phylogenetic out sort to attempting In . eainhp ihntePlgnca aebe eetyeaie using examined recently been have Polygonaceae the within Relationships hools DNA, chloroplast c.2 species), 25 (ca. (including , oeua hlgn of phylogeny Molecular Fallopia Persicaria Eupersicaria rbcL sdsigihdfo h te major other the from distinguished is Echinocaulon , Tovara trnL–F , .punctata P. Pteropyrum Polygonum ld contains clade hr per ob epsltbetween split deep a be to appears there Eupersicaria , Bistorta trnK c.3seis.Seconsid- She species). 3 (ca. hs eut e h tg o oedtie hlgntcaayi of analysis phylogenetic detailed more a for stage the set results These . intron– agTeKim Sang-Tae c.2 species), 21 (ca. Fagopyrum , Atraphaxis es stricto, sensu c.5 pce) and species), 50 (ca. ITS, , Bistorta Tovara Persicarieae matK omnctn dtr ae .Smith F. James Editor: Communicating Persicarieae Polygonum and , and n ld including clade a and c.1 spe- 16 (ca. and , Echinocaulon psbA Polygonum Persicaria Polygo- Cepha- 1,2 based epeo test. Templeton , Persi- Poly- – Cal- trnH n ihe .Donoghue J. Michael and on U.S.A. G,adncerrbsmlISsqecs ecnld that conclude We sequences. ITS ribosomal nuclear and IGS, ( Eupersicaria es lato sensu ntr,ti ld sms lsl eae to related closely most is clade this turn, In . 77 ut fasuyo oyoaeeb abFr n Kron and Fyre Lamb by re- Polygonaceae the on of test based and study (2003) upon a build of analyses sults Our region. ITS somal are here genes used data chloroplast The the group. from this within studies lutionary inter- of are relationships and we sicaria monophyly especially the Most assessing in relatives. ested closest its of within fication primary relation- relationships Our on their data. sequence is infer DNA focus to using of another monophyly and one the taxa to test ships aforementioned to the is of aim Our each describe. to easy comes worker). to worker from somewhat varied have tions trnK have authors most what for section it as used to who referred name b), have the (1913a, we of Gross choice paper The this taxon. level of purposes name To the for usage. adopted confusion, standard with any keeping avoid a in to taxon, and genus-level taxon sive, use genus-level will a We to taxon. both section-level name applied the been of has case name the in except atic Tovara inet r reeat n ewl s hs ae nyas as- only names rank them these use these will to purposes we and assigned present irrelevant, are ranks our signments For taxonomic 1978). the been (Haraldson has in there 1856), disparity (1826, great Meisner by treatments major the eintosfrpttv lds hs o xml,w will we example, to for refer Thus, simply clades. putative for designations gonon T ewe 0220 nNrhAeia rt,Cia n oe,but Korea, by and field China, the Crete, in America, collected North were Lamb in samples in 2002–2004 our sample between of the STK Most to (2003). corresponds Kron largely and also of Frye it all relatives; represent close to within sumed chosen clades was sample major This the purposes. rooting for cluded of three .amphibia P. nte2 cesosw sdt ersn 4seis ih accessions eight species: 24 represent to used of we species six accessions represent 26 the on .amphibia P. ao Sampling— Taxon ihteenmsi ad h ups forpprbe- paper our of purpose the hand, in names these With lhuhms fteetx aebe eonzdsince recognized been have taxa these of most Although Aconogonon Priaia,Polygonaceae) (Persicarieae, iha xadddtst nldn diinlseisand species additional including dataset, expanded an with ) nrn and intron, n aho h eann i rus A groups. six remaining the of each and rt “genus to or ” ( sapeuet oedtie hlgntcadevo- and phylogenetic detailed more to prelude a as Cephalophilon Polygonum and n h eann pce,adteei togconflict strong is there and species, remaining the and lpetlBooy eateto Molecular of Department Biology, elopmental and rbcL 016NwHvn onciu 06520-8106 Connecticut Haven, New 208106 x .hydropiper P. Tovara Koenigia, rbcL sect. eune,wt nepai on emphasis an with sequences, psbA pedx1poie oce pcmninformation specimen voucher provides 1 Appendix w ahof each two , M Eupersicaria Eupersicaria TRASAND ATERIALS Persicaria Persicaria eune alone. sequences Bistorta 1 – Persicaria [email protected]) trnH rto or ;fu pce of species four ); hc uprstemnpyyo the of monophyly the supports which G,adfo h ula ribo- nuclear the from and IGS, rbcL .Ti prahi unproblem- is approach This ”. nmn ro ramns,using treatments), prior many in Bistorta Tovara toacsin ahwr nlddof included were each accessions (two es aado 17)adispre- its and (1978) Haraldson sensu o h esicuie section- inclusive, less the for atog xc circumscrip- exact (although Persicaria Persicaria , Eupersicaria M trnL–F and Persicaria ETHODS Cephalophilon ahrta o“section to than rather , Eupersicaria Echinocaulon Bistorta partial , n nteidenti- the on and , Rumex o h oeinclu- more the for . Polygonum n n of one and , tef ic this since itself, Eupersicaria h sister The . pce a in- was species eeincluded, were sbsdon based is matK and Euper- Acono- with Delivered by Publishing Technology to: Yale University IP: 130.132.173.179 on: Fri, 02 Aug 2013 21:54:39 Copyright (c) American Society for Plant Taxonomists. All rights reserved. h rgnlsac.Asre fheacia ieiodrtotss(hLR tests ratio likelihood hierarchical of those series to A identical 1985) search. parsimonious settings (Felsenstein original search most bootstrap heuristic the of the with with replicates islands 1,000 support ad- multiple using node sequence sample of evaluated The to order We characters. attempt random trees. all a an with of zero in times dition weighting of 100 equal repeated collapse were and swapping, analyses with lengths, branch methods branch (TBR) search maximum reconnection heuristic using bisection performed par- tree Maximum were 2001). (Swofford searches Ronquist 4.0b10 and simony (Huelsenbeck PAUP* 3.1 using MrBayes conducted and the were 2002) sets, from sets, data data request chloroplast combined the upon nrITS, our are or of here) analyses S1816) Phylogenetic published number author. trees first (study the (and again TreeBASE sets alignment, in data for available al. aligned 3.6) Our et Ver. eye. 2004, (Notredame by (Edgar T-COFFEE adjusted MUSCLE used and also 1.35) we Ver. cases 2000, these In gaps. required erhwr iutnosyetmtdvamxmmlklho o all swapping branch for each TBR likelihood for with used Parameters maximum were methods 1). via search (Table Heuristic estimated datasets. set simultaneously data Infor- were each Akaike using search for the PAUP* using (AIC) models in Criterion alternative out mation and Crandall carried hLRT and by were selected (Posada searches models 3.7 likelihood version fit Modeltest Maximum best program evolution 1998). the sequence using of model data which the determine to performed was trnH for 1991) al. et (Taberlet e and d reverse region, (the nrITS ITS3b the and addi- trnL 1990) for and al. ITS2) primers et of (White amplification sequence ITS2 the follows: as using (Qiagen). primers, out Kit tional PCR Purification carried below. PCR was steps QIAquick the a Sequencing using using purified sequencing were were two for products least colonies selected at and were Eight M13Rev, products and protocol. PCR M13For primers supplied using screened the Carls- and (Invitrogen, picked following kit cloning California) TA TOPO a bad, using products PCR the cloned from obtained fragments islandica smaller above. the described of conditions amplification the the using Because below) (see sequencing for primers islandica Koenigia nrITS, amplifying in 72 94 at used 94 we ation markers at chloroplast the predenaturation For following: min. 7 the for extension final a and cycle, 97 at denaturation of 48 cycles 97 at at 40 predenaturation by region: followed ITS min the 1 for follows as out carried were TGGA-3 and 2003) Kron and Frye (Lamb trnL the for 5 the for transcribed (nrITS); internal region the coding for (5 rRNA 1990) 5.8S al. the et including (White region, ITS4 spacer and 1998) al. et (Baum 1.0 and Massachusetts), Ipswich, ␮ Biolabs, England (New dNTPs mmol/L 8SSEAI OAY[oue33 [Volume BOTANY SYSTEMATIC 1 tion, 25 in poly- standard (PCR) using reaction Am- performed chain California). was merase Valencia, DNA speci- stranded (Qiagen, herbarium double Kit from of Mini plification or Plant DNeasy silica-gel a in using dried mens samples leaf from extracted (NHA). Hampshire the New of of (YU) Herbarium barium; University of Yale Samples History. islandica the Natural in of Museum available Peabody are identified and were specimens STK field-collected All 2006. in of Garden Botanical sample living a 78 lge sn lsa Topo ta.19)adajse yeeto eye by adjusted as and such regions 1997) noncoding al. Aligning et improvements. (Thompson slight achieve X Clustal Sauk sequencer. using Research, DNA automated aligned (MJ 3100 ABI 51 an BaseStation or 377, a ABI Perkin- an using 1995, Wisconsin), visualized City, August was B, (Revision and Kit Elmer) Primer Reaction Dye Ready PRISM ABI Sequencing the Cycle by specified protocol the followed of sequencing sequence reverse (the PS1R . nto a oyeae(ign;2.5 (Qiagen); polymerase taq of unit 1.0 o/ mlfcto rmr.Tefloigpieswr sd ITSLeu used: were primers following The primers. amplification mol/L Ј ° N xrcin mlfcto,adSequencing and Amplification, Extraction, DNA lgmnsadPyoeei Analyses Phylogenetic and Alignments GTCATTCTTGG-3 -GTTCCAATTATGCCTCT o 0sc,adafnletninfr1 i.Teewr difficulties were There min. 10 for extension final a and sec., 90 for C – ° and IGS, F o 0sc,etnina 72 at extension sec., 30 for C and , ␮ psbA fMgCl of L .meisneriana P. Ј Ј sal i o il nuhpoutfrdrc eunig we sequencing, direct for product enough yield not did usually o the for ) eeotie rmseiesi h aeUiest Her- University Yale the in specimens from obtained were trnF trnK trnL ° – o 0sc,anaiga 54 at annealing sec., 30 for C trnH rbcL – n the and nrnadpartial and intron F osalrfamnswr mlfe ihadditional with amplified were fragments smaller so , nrnadIS n h rT einwsdfiutand difficult was region nrITS the and IGS, and intron P1 (5 -PS1F G ( IGS rbcL 2 .amplexicaulis P. omk ia ocnrtoso . mlL 1.0 mmol/L, 2.5 of concentrations final make to rbcL a bandfo h ebru fteUniversity the of herbarium the from obtained was psbA trnF oigrgo ( region coding and , Ј -AGGMCATTACTTGAATGC-3 age l 1997); al. et Sang ; nrn( intron rbcL trnL rbcL matK ° o 0sc nraig4sc iheach with sec. 4 increasing sec. 20 for C ␮ P1)for -PS1F) trnL – Ј 12R(5 -1425R ecin otie 1 contained reactions L and ) ° F a bandfo h e York New the from obtained was o 0sc,3 ylso denatur- of cycles 35 sec., 90 for C ein(p- region rbcL rmtehraimetato of extraction herbarium the from – ␮ F Lof10 – aelte l 91;ad1FS and 1991); al. et Taberlet ; 62 .Plmrs hi reactions chain Polymerase ). trnk — Ј ° “ eune eeprimarily were Sequences -CAAAGTATCCATTGCT- rbcL o 5sc,etninat extension sec., 45 for C c 22 Yuge l 1999) al. et (Young -2621 ” × matK ° and o 0sc,annealing sec., 10 for C y emntrcycle terminator Dye . ufr 5 buffer, .arifolia P. — eoi N was DNA Genomic “ ); f ” psbA o G between IGS for ␮ – Ј and and F fQsolu- Q of L 0n DNA, ng 10 and ) matK Koenigia Koenigia -PA1F trnH ° psbA for C rbcL and for by T) R – - eea nMsnGmradKlog1996). Kellogg and Mason-Gamer in as here oeweetplgclcnlc a bevd( observed was conflict topological where node a rmaohrdt e,ad()svrlmodified several (2) and set, data another from 93 sipeetdi AP oass h ee fcnrbto of of types contribution two to of set, W data level trees. between the conflict a the assess for Templeton compared responsible to be test; might PAUP* that (WSR nodes in test specific with implemented signed-ranks out combined as Wilcoxon the data carried a 1983) with between was differences conducted as significant well we test a suggested as sets ILD tests set ILD The data When set. gene data. data each the for partitions of pairwise repartitions al. searches heuristic 999 et and We swapping, (Farris with test. branch we homogeneity TBR test addition, partition markers taxon the (ILD) simple as gene used difference PAUP* different in length implemented from 1995) incongruence 1994, sets the data th out assess among carried To congruence node. each of for probability level posterior a generate to calculated oi ie.B osbese n opsrcueof structure loop by and indicated stem are Possible B. sequences lines. complementary solid Inverted inversion. dicates nepalensis from is neriana I Type of sequence eoig1000gnrtosa a as After generation. generations 100th 100,000 every prob- collected removing likelihood were Trees and MCMC. evolution using sequence abilities to relating the parameters estimate from the using estimated conducted values were GTR+GorGTR+G+I inferences to Bayesian fixed tree. swapping. were likelihood (NNI) maximum tests interchange bootstrap neighbor for nearest Parameters with replicates times 1,000 100 ing repeated were Analyses the branches. with zero-length of collapse and n G otn o iegn akr r rsne in presented are markers gene five for content %GC and al .Sqecso oigrgossc as such regions coding of Sequences 1. Table aito n twsncsayt nrdc nesi the in indels introduce to necessary length was showed alignment. it regions In and noncoding straightforward. variation of therefore sequences were contrast, alignments and length matK netdrpaso 7o 8b in bp 18 in or 17 attaching of bp 21 repeats of inverted phyloge- inversions Short their unclear. rendering significance netic overlapped, variously include or not identical sets did data We most was as 1). except analyses, however, (Table phylogenetic sites, in marker information variable gene indel to each PI in of similar the proportion was The region sites. in nrITS sites set; The data (PI) aligned parsimony-informative bp. the 30% 1335 with to variable, bp most 577 from ranged folia se- longest in in the bp from 221 were differences quences regions; noncoding most in utda nlsswt hs neso removed. inversion con- these also with We analysis details). an for phy- ducted Discussion our in (see included analyses and logenetic alignment for inverted were ments maackiana F lge N Sequences DNA Aligned itrapleacea Bistorta trnL IG 5 p.Teaindsqec egho ahmarker each of length sequence aligned The bp). 650 : .A neso eunetpsi the in types sequence Inversion A. 1. . n h .Srgo fnISaecnevdi sequence in conserved are nrITS of region 5.8S the and , yeI a rsn in present was II Type . – rbcL “ , F admaddition random .arifolia P. , ( Rumex “ .meisneriana P. ongaislandica Koenigia ettree, test wt oidl)cnandidl htwr not were that indels contained indels) no (with , , p:93b) n 1 pi rT ( nrITS in bp 117 and bp), 963 sp.: .maackiana P. escracapitata Persicaria ” “ rbcL h titcnesste nerdfo given a from inferred tree consensus strict the ia trees: rival oes iemlingnrtoswr u to run were generations million Five models. ” .sagittata, P. psbA pin otta et eepromdus- performed were tests Bootstrap option. a es aibe otiig9 PI 9% containing variable, least was Tp Ii i.1A.Teefrag- These A). 1 Fig. in II (Type R — and , ESULTS itrapleacea Bistorta hwdtesots sequences shortest the showed ( h ag fsqec length sequence of range The ” “ .pensylvanica P. unin burn 1 h titcnesste inferred tree consensus strict the (1) P.meisneriana. n hto yeI from II Type of that and psbA , .nepalensis P. ” psbA – 0 osnu rewas tree consensus 50% a “ trnH ettrees test ’ test , – .sagittata P. escracapitata Persicaria trnH ’ 9 p,14bp 134 bp), 393 : and hddprinin- portion Shaded G eefound were IGS rbcL G ein The region. IGS ” , ’ .arifolia P. rival osrie at constrained atof part a , .lapathi- P. sequences. ’ r used are .meis- P. , , P. P. e e Delivered by Publishing Technology to: Yale University IP: 130.132.173.179 on: Fri, 02 Aug 2013 21:54:39 Copyright (c) American Society for Plant Taxonomists. All rights reserved. o fM re 8261 2 2481 60 837 1 19 836 64 1386 229 2 30 4420 1626 356 608 60 18 753 17 623 1030 233 58 6 3667 21 317 403 115 61 1107 2 269 7 189 103 59 1335 28 409 174 16 155 58 648 265 27 trees MP of length trees 577 MP of No. (%) sites PIS/variable (%) length seq. PIS/aligned PIS of No. sites variable of No. (bp) length seq. Aligned oe eetdGgiGgGgiGgGgiGgG+g+i 0.776 0.644 G+g 0.734 19634. 19 0.704 4969.158 0.572 4977.958 G+g+i 0.650 0.822 14224.537 14251.055 0.697 G+g 0.785 4862.155 4894.937 0.839 0.741 3840.002 G+g+i 3878.535 0.822 0.792 2406.142 2441.061 0.603 G+g 0.700 2637.860 0.817 2667.459 G+g+i 0.675 0.766 0.844 0.739 BA in 0.822 L -ln Mean ML in L -ln selected Model (RI) Index Retention sites) invariant (excl. CI (CI) Index Consistency ie;M otparsimonious most = MP sites; intron; G otn alsts 23.6 sites) (all content %GC ie;M aiu ieiod A=Bysa nlss rtmtcma o mean Arithmetic * analysis. Bayesian = BA likelihood; maximum = ML sites; Bistorta The analyses. our within in clades uncertain the remained of Relationships sis. of for except 0.99) psbA over PP 77%; over values (bootstrap nocaulon p- in supported weakly in to unresolved but and B) 67/62/0.9) = 2 ML/PP Fig. 1.00; of PP psbA of phyly for trees p- gene resulting the sum- and trnL are 1 set Table data and in gene marized likelihood, chloroplast each maximum of analyses parsimony, Bayesian maximum our with Note: eunelnt b)172 (bp) length Sequence 79 PERSICARIA OF PHYLOGENY MOLECULAR DONOGHUE: & KIM 2008] psbA including clade the Bistorta, whereas A), 2 (Fig. analyses p- the in except Aconogonon including reso- clade the clear the between the and relationship of sister exception a the of with lution sets data other in solved togspot(ottasoe 7;P .0 in 1.00) = PP 97%; over (bootstraps support strong solved. psbA the to sister 0.93 PP (with analysis Bayesian in found oeaespot(ottas63 and (bootstraps support moderate B). A, 2 (Fig. analyses ML the and to sister being clade A.B.K. the for aaSet Data gonella the with linked matK trnL T matK nlsso pN eeDt Sets Data Gene cpDNA of Analyses nlsso obndcDADt e n nnrITS an and Set Data cpDNA Combined a of Analyses .amphibia; P. Persicaria ABLE – – ,p- trnL 1 F F and nlss(PM/P=5/307) h monophyly The 59/53/0.72). = (MP/ML/PP analyses nlss eainhp among Relationships analyses. re u eeal hsrltosi eandunre- remained relationship this generally but tree, 4 and , negncsae einbetween region spacer intergenic r rsne nFg .Vr iia re for trees similar Very 2. Fig. in presented are .Aindsqec nomto n h umr fmxmmprioy max parsimony, maximum of summary the and information sequence Aligned 1. obnddtstwith dataset combined Fg ,B,admdrtl 7/209)s np- in so (78/82/0.99) moderately and B), A, 2 (Fig. ihhg upr M/LP 99/.0 in 99/99/1.00) = (MP/ML/PP support high with Polygonum r icse eo,btaentson h mono- The shown. not are but below, discussed are matK, and , and — – trnL Eupersicaria F nogunelnt ifrnetsswt com- a with tests difference length Incongruence Persicaria- Fallopia and a togyspotdi vr pN analy- cpDNA every in supported strongly was analyses. Koenigia – and i.2A.Tecae representing clades The A). 2 Fig. F Cephalophilon matK Polygonum aast oe 9 otta upr n a and support bootstrap 89% (over sets data Koenigia Aconogonon-Koenigia trnL .sr and str. s. eels la nthe in clear less were rewhere tree ABK ld)rcie esspotin support less received clade) (A.B.K. ...caewt ihspotol in only support high with clade A.B.K. a elspotdi nlsswith analyses in supported well was – Fallopia F Euperscaria rbcL eea iereversible time general = G ; Fg )aaye.Srn support Strong analyses. B) 2 (Fig. eesrnl ikdi ahtree each in linked strongly were psbA .str.- s. Polygonella eeas togysupported strongly also were nlssadwr oryre- poorly were and analyses ,p- psbA Koenigia ssrnl (100/100/1.00) strongly is Polygonella Fagopyrum – – matK – 3832.6 33.8 9 557 393 2;P .)i p- in 0.9) = PP 72%; 1 psbA , Echinocaulon , Polygonum rbcL — ld a ikdwith linked was clade and rbcL – omdacaewith clade a formed Persicaria Persicaria a o nlddin included not was ttsisassociated Statistics .0,btnti MP in not but 1.00), and , Cephalophilon psbA trnH ld in clade a lcda a as placed was nlss(owing analyses p- trnL matK ; – – analyses. 69443.9 36.9 2 1324 626 2 Tovara Aconogonon .str., s. matK and – partial 2 however, , mn iert aito oee oftadsrt am distribution gamma discrete a fit to modeled variation rate site among = g ; F ld was clade ; psbA Tovara rbcL rbcL 5 rbcL obnddtstwith dataset combined Tovara. , (MP/ clade matK matK Poly- Echi- rbcL and and and ,or in , and bLtrnL rbcL – – 38829 1328 5132.6 45.1 trnK rvdpyoeei eouinwt ihrnd confi- among node relationships However, higher C). im- 2 with (Fig. provided dences resolution analyses phylogenetic cpDNA proved combined separate the analyses, the to that Compared gene 2). indicated (Table also combinable Pair- were gene set 0.372). chloroplast data = each (P with partitions signifi- tests gene no wise found the genes among chloroplast differences four cant the of set data bined Echinocaulon, h 0 osnu re rmtreaaye.Relationship analyses. three from trees among consensus 50% the h orcDAmresidctdta p- that indicated of each markers versus cpDNA nrITS the four with tests the null ILD sets Pairwise the data 0.0001). two < rejected between (P difference nrITS) significant versus no of (cpDNA hypothesis partitions two ing Markers Gene All including Similarly, analysis. combined cpDNA the rT n pN nlsscnendseisrelationships species concerned analyses cpDNA the and between differences nrITS Several analysis. within Bayesian the groups in major only four instead the including analyses; Persicaria clade nrITS clade a A.B.K. our the with tree, in linked cpDNA appear the to not contrast did In analy- D). Bayesian 2 the (Fig. in sis except place- supported well the not and were results clade A.B.K. Some of the 0.91). ment concerned combined = set the (PP data and nrITS cpDNA support the moderate between differences with topological clade a two formed the suggested analysis as Bayesian paraphyly, analysis. MP the the within in 66%) of where support analyses, bootstrap (with sequence nrITS in lematic eentsgiiatydfeetfo rT Tbe2.Results 2). (Table nrITS from different significantly not were lophilon of runcinata species other to sister as placed strongly with clade a formed with of taxa other all amphibia to D). sister C, as 2 placed (Fig. clades named the within w usatrbr in. burn after runs two f reIcnrec n nlsso aaStIncluding Set Data a of Analyses and Incongruence Tree 5 Ј .hydropiper P. intron; whereas , Eupersicaria psbA ntecDAcmie nlss u a linked was but analysis, combined cpDNA the in ) Cephalophilon lhuhthe although , omdawal upre ld within clade supported weakly a formed .runcinata P. mmlklho,adBysa nlsso aiu datasets. various of analyses Bayesian and likelihood, imum – – ,p- 3 – 5657.6 35.6 6 533 963 F negncsae einbetween region spacer intergenic and 3 matK .nepalensis P. Tovara — ntenISanalysis. nrITS the in , , rbcL Echinocaulon nogunelnt ifrnetssus- tests difference length Incongruence .sagittata P. comb-cpDNA and ld.I eea,hwvr h nrITS the however, general, In clade. , trnL .amphibia P. Persicaria eeabgosadursle in unresolved and ambiguous were .capitata P. n rT;PS=parsimony-informative = PIS nrITS; and perda itrt h clade the to sister as appeared 4 ntenISaayi,btwas but analysis, nrITS the in and , ld a elsupported well was clade Eupersicaria ouain plus populations ntecDAtree. cpDNA the in Tovara escrapunctata Persicaria escrameisneriana Persicaria rT comb-all nrITS trnL – – .nepalensis P. 67.7 650 Eupersicaria Tovara matK eeas prob- also were Echinocaulon and (excluding invariable = i ; Bistorta Eupersicaria Eupersicaria trnF and appeared and Cepha- Tovara and 665.026 clade psbA was was trnL 516 in P. P. 5 , Delivered by Publishing Technology to: Yale University IP: 130.132.173.179 on: Fri, 02 Aug 2013 21:54:39 Copyright (c) American Society for Plant Taxonomists. All rights reserved. niae y-.Smosrpeetn oendsidct osrit us constraints indicate nodes some branches the representing above Symbols presented --. are by analyses indicated likelihood maximum / parsimony trnL 0SSEAI OAY[oue33 [Volume BOTANY SYSTEMATIC 80 F IG – F .5%cnesstesfo aeinifrne o A. for inferences Bayesian from trees consensus 50% 2. . ,adD h ula iooa T eunedt e.Pseirprobabili Posterior set. data sequence ITS ribosomal nuclear the D. and ), rbcL ,B. trnL di icxnsge-ak et seTbe3). Table (see tests signed-ranks Wilcoxon in ed – upr fls hn5%bosrpvleo . otro rbblt is probability posterior 0.5 or value bootstrap 50% than less of Support . F .tecmie aastfrfu pN ee ( genes cpDNA four for set data combined the C. , isaepeetdudrtebace.Bosrpvle o maximum for values Bootstrap branches. the under presented are ties psbA ,p- matK , rbcL , Delivered by Publishing Technology to: Yale University IP: 130.132.173.179 on: Fri, 02 Aug 2013 21:54:39 Copyright (c) American Society for Plant Taxonomists. All rights reserved. rT s pDAcombined DNA cp vs. nrITS h ulhptei fn ifrnebtentetoteswt two-tailed with trees two the between difference no of hypothesis null the obnbei L eti rsne y*. by presented is test ILD in combinable aoedaoa) infcn ifrnebtendtst ugsign suggesting datasets between difference Significant diagonal). (above rT vs. nrITS vs. nrITS 81 versus nrITS with test WSR the of PERSICARIA OF PHYLOGENY trnL MOLECULAR DONOGHUE: & rbcL KIM p- psbA 2008] eue tp httedt e ne osdrto scntandb h i the by constrained is consideration under set data the that steps reduced ld a,b,c n 4i i.2adTbe3,adrelation- and 3), Table and 2 Fig. in d4 and c4 b4, a a (a4, of of clade existence existence the the 3), 3), Table Table and 2 and 2 Fig. in lapathifolia d1 and c1 b1, we of (a1, Specifically, placement 3. the Table concerning in conflicts investigated presented are data cpDNA bined matK trnL T rT pDAcmie aastsrc osnu re3 62 0.0006* 20 16 36 tree consensus strict set data combined DNA cp / nrITS / nrITS pDAcmie rT titcnesste 596 0.0001* < 66 9 75 tree consensus strict nrITS / combined DNA cp rbcL tree/ consensus strict nrITS T ABLE ABLE – (d2) of Clade (d1) of Clade (b5) c)Mnpyyof Monophyly (c6) of Clade (c5) of Clade (c4) of Monophyly (c3) of Clade (c2) (c1) of Monophyly (d7) of Monophyly (d6) of Clade (d5) of Clade (d4) of Clade (b4) of Monophyly (b3) of Clade (b2) of Clade (b1) d)Mnpyyof Monophyly (d7) of Monophyly (d6) of Clade (d5) of Clade (d4) (d2) of Clade (d1) of Monophyly (d7) of Monophyly (d6) of Clade (d5) of Clade (d4) of Monophyly (d3) (d2) of Clade (d1) of Monophyly (a6) of Clade (a5) of Clade (a4) of Monophyly (a3) of Clade (a2) (a1) F - rT titcnesste 542 0.0001* < 21 4 25 tree consensus strict nrITS / F .Tmltn(icxnsge-ak)ts o nogunebtendata between incongruence for test signed-ranks) (Wilcoxon Templeton 3. 2. .punctata P. .punctata P. nISsrc osnu re2 60.0016* 16 5 21 tree consensus strict /nrITS .lapathifolia P. .lapathifolia P. .lapathifolia P. trnL rbcL trnL p plus au bandfo arieIDts ewe ahgene each between test ILD pairwise from obtained value – - psbA F F — titcnesste 483 0.0001* < 36 8 44 tree consensus strict .capitata P. maackiana, P. arifolia, P. lapathifolia P. .capitata P. maackiana, P. arifolia, P. lapathifolia P. .capitata P. .aioi,P maackiana P. arifolia, P. lapathifolia P. hydropiper P. .punctata P. .capitata P. sagittata P. .capitata P. sagittata P. punctata P. capitata meisneriana P. P. and sagittata P. punctata P. .pensylvanica P. stesse oters of rest the to sister the is stesse oters of rest the to sister the is stesse oters of rest the to sister the is stesse oters of rest the to sister the is of rest the to sister the is itra Koenigia, Bistorta, Eupersicaria itra Koenigia, Bistorta, Cephalophilon Eupersicaria itraadPersicaria and Bistorta Tovara Bistorta Tovara Tovara, Echinocaulon, Bistorta Tovara aast/Cntansin Constraints / set Data p- .0 .1 .1 0.076 0.811 0.412 0.301 aKrc trnL rbcL matK — and and and and and and and and and and and rbcL and and and and and and and and and .runcinata P. .runcinata P. .runcinata P. .nepalensis P. .nepalensis P. .nepalensis P. .meisneriana P. .meisneriana P. .hydropiper P. .hydropiper P. hydropiper P. titcnesste 01 4<0.0001* < 34 16 50 tree consensus strict uescraecuigP amphibia P. excluding Eupersicaria Eupersicaria Eupersicaria .cespitosa P. .pensylvanica P. .pensylvanica P. .pensylvanica P. ld a,b,c n 2i Fig. in d2 and c2 b2, (a2, clade Persicaria Persicaria plus .sagittata P. .9 .9 0.073 0.897 0.392 and , rbcL and — and and and Eupersicaria and Eupersicaria , Eupersicaria Aconogonon Aconogonon .meisneriana P. .meisneriana P. .meisneriana P. “ trnL ia tree rival excluding excluding Eupersicaria Eupersicaria Eupersicaria Eupersicaria and – .7 0.001* 0.479 F excluding — ” n h com- the and , .meisneriana P. – F excluding excluding ld 0550.1967 5 5 10 clade .amphibia P. amphibia P. .punctata P. excl. excluding excluding .amphibia P. .amphibia P. 0.007* nrITS .amphibia P. .amphibia P. ot- P. .amphibia P. amphibia P. cnrettplg nthe in topology ncongruent et seikidctstesgiiatdfeec at difference significant the indicates Asterisk test. ) h lcmn of placement the 3), infcn ifrne( .05.I otat hnthe when contrast, In 0.0015). = (P difference significant including clade the to of placement the and 3), the of At persicaria relationship 3). the Table concerning and conflicts 2 Fig. in of d5 species three the among ships rT a osrie ythe by the When constrained nrITS. was from inferred nrITS tree parsimonious most the in 3 directions. Table both in ing in out conflicts carried other example, tests For between the in conflict of significance None the showed nrITS. to and significantly cpDNA contribute of trees to placement consensus The seen strict hypothesis. rival null the contrasting rejected tests Many 3). Table .amphibia P. es(e i.2.Gi G,ls L n e ersn diinlor additional represent net and (L) loss (G), Gain 2). Fig. (see sets excluding Cephalophilon 801 0.0001* < 18 0 0.0001* < 18 18 0 18 741 0.0046* 0.0593 0.0067* 0.0005* 13 0.0001* < 8 9 12 4 20 5 1 0 17 0 13 10 12 20 0.1967 0.0015* 5 0.0001* < 18 5 18 7 10 0 25 18 e P-value Net L G nthe in 0.0027* 0.5637 0.3173 9 1 3 0 1 3 9 2 6 0.5637 0.3173 0.0082* 0.5271 1 3 7 0.0588 2 0.1573 0.0833 1 0.1573 3 0 5 4 2 3 2 0.0082* 2 6 0.5271 7 6 1 0 0 4 0 - 0 2 00 6 2 3 2 0 4 4 6 0.0082* 7 0 7 0.0047* 1.0000 0.0455* 0.3173 0.6547 0.0027* 8 0.0082* 0 4 2 0.5637 1 0.3173 9 0 7 2 0.5271 0 1 1 2 3 8 0 2 0 2 4 3 3 1 9 3 7 4 2 6 6 ’ .amphibia P. ia tree rival Bistorta rbcL Eupersicaria Tovara titcnesste,bti ssister is it but tree, consensus strict stesse to sister the is ’ prxmt rbblt P evaluates (P) probability Approximate . a,c n 7i i.2adTable and 2 Fig. in d7 and c6 (a6, rbcL a n 3i i.2adTable and 2 Fig. in d3 and (a3 b,c n 6i i.2and 2 Fig. in d6 and c3 (b3, Cephalophilon “ , section re h S hwda showed WSR the tree, Echinocaulon Cephalophilon Eupersicaria p ” 0.05. < ee estudied we level a,b,c and c5 b5, (a5, .punctata P. and , exclud- Tovara to rbcL Eu- is Delivered by Publishing Technology to: Yale University IP: 130.132.173.179 on: Fri, 02 Aug 2013 21:54:39 Copyright (c) American Society for Plant Taxonomists. All rights reserved. fivrin aebe eotdi h aergo from region same the in reported types been aligned Paeonia Several have in 1). inversions (Fig. bp structure of 18 stem-loop possible or a 17 gesting of of repeats sequence inverted short between n ny06 Pi aeinanalyses. Bayesian values in bootstrap PP 50% 0.64 than only less and with supported, 3). weakly with (Fig. was values directly support linked set high data was combined with cpDNA maintained the in cpDNA-ITS were found combined clades we the most conflict set, of data significant analyses the In of documented. view have in caution must with results combined treated the be in without placement or its Clearly, with analyses. inferred trees topological in no punctata level found this we datasets at because these differences combined Also we analyses. level, between further this conflicts for at significant nrITS no and found cpDNA we and relationship not was test 0.6547). WSR = the (P tree nrITS significant the by constrained were data sal soitdwt tmlo rhipnsrcue have structures hairpin or inversions stem-loop short with contrast, In associated 2000). usually al. et shown (Tsumura al. been conifers et in has (Cosner polymorphism Campanulaceae intrapopulational Ranunculaceae and and 1994), 2004), 1994), Palmer Palmer and and (Hoot (Downie homoplasy Chenopodiaceae in although Cactaceae inversions 2005), and such al. in et documented been Kim Raube- has 2005; 1987; Palmer Jansen and and (Jansen son levels analyses taxonomic phylogenetic higher for at have characters genome useful chloroplast considered the been in inversions scale Large 2000). and ikd hra rmtedtsticuigteivre se- inverted the a including obtained we dataset quence the from whereas linked, in autapomorphy one same of the as in with ception (adjusted species identical other were sequences from strand) included inverted those complimentary and the the dataset to that the necessary found in these We others inverted the them. we match instead to and inversion characters; sequences Ohsako the as code 1997; not themselves gene did al. we events same analyses, et our the Sang In 2000). in 1996; Ohnishi sites Wendel nucleotide and other (Kelchner to compared evolutionary events rates heterogeneous inversion possible of and order mutations base the and determining owing of analyses difficulty phylogenetic the in to problematic as viewed been ocuigta infcnl ifrn aast r homoge- of are chance sets low a data be different to significantly appears 2002; that there et 2004), concluding Yoder Lecointre com- al. 2002; et of and Lutzoni Hipp measure 2001; and Darlu al. Barker a 2002; 2000; as Austin test and al. Dowton ILD et the (Dolphin of binability utility the over versy differ- minor values. only confidence with the in inverted, ences were as topology sequences same the the Analyses when yielded B). A, removed 2 inversions (Fig. these markers with cpDNA other the with sistent psbA within ference lophilon 2SSEAI OAY[oue33 [Volume BOTANY SYSTEMATIC 82 hr neso Sequences Inversion Short eas h anfcso u td so eto level section on is study our of focus main the Because aaCmiaiiyadIncongruence and Combinability Data trn rei which in tree C netn n nldn hs eune aeadif- a made sequences these including and Inverting . Sn ta.19)adi te ee (e.g. genes other in and 1997) al. et (Sang dt o hw) eretained we shown), not (data -rpo pcr from spacer) B psbA Cephalophilon ewe uloiests8 n 0,sug- 109, and 80 sites nucleotide between .capitata P. .capitata P. D Eupersicaria ISCUSSION ihu netn eotie a obtained we inverting without : Fagopyrum — and hr r nesossituated inversions are There plus .nepalensis P. .capitata P. .runcinata P. “ u hsrelationship this but , section, Osk n Ohnishi and (Ohsako .punctata P. — ept contro- Despite ” within eedirectly were ihteex- the with trn ld,con- clade, intron K nthese in Cepha- Tovara P. eainhp,sc sterltosisof relationships the as such the in relationships, differences to due of which mainly sets, placement were data showed cpDNA tests our WSR to our nrITS comparing in conflicts within of (with differences placement markers several different of the exception from the congruence obtained overall trees of the an- observation the between one the that from with different indicated consistent significantly other, tests not are ILD markers pairwise and cpDNA (Darlu Our sites 2002). informative data of when Lecointre number rate) sufficient error a II have Type sets (low combine to enough neous ee(i n oohe Si rp) ti osbethat possible is It prep.). in MS Donoghue, and (Kim al. of gene parentage et possible Consaul the punctata exploring 1980; P. been have McDonald we and 1964; 1991), Timson 1925b; ford with in ticulation associated most incongruence The the misleading. punctata be for may explanation position its likely spe- that the caution of but with remainder 3, tree the the shown for have We without topology a cies. same of and the analyses with in out Analyses resulted carried set. we basis, data this combined On direction. one in lophilon rT,sc stoeinvolving those and as cpDNA such between discordances nrITS, minor, more other, some ra es Jrtk 98 L 1928; throughout poly- (Jaretzky and by sense groups supported broad these is in possibility ploidy This introgression. mic and reee l 00,adwsspotdi rvosphylo- previous De- a 2003). in Ronse of supported 1998; was analysis al. and genetic et 2000), al. Hong et 1988; craene Ronse Akeroyd 1978; (Haraldson and similarities affinity Decraene vegetative close and floral a on However, L 1969). 1997; Federov 1946, between 1957; (Hedberg base Sarkar seven a and of and grains, number ar- arctic pollen chromosome and spinulose-spheroidal alpine unique in the distribution eas, on predominant genus its separate a of as basis segregated been group. often has monophyletic species, a results be our not on would Based this 1988). Akeroyd and Decraene Ronse (i.e. genus The 3). Fig. including Persicarieae evolution. concerted rapid by low affected is- nuclear not especially these genes markers, of copy additional Clarification require 1997). the will Cunningham of sues 1995; some Quei- al. least (de error et at sampling roz of that result possibility a is the incongruence observed out rule yet cannot to and related 1999) groups in in as such hybridization of examples oinclude to between relationships lophilon Persicaria genus the as recognized of monophyly the support sicaria strongly sets data combined h oohl fPriai n eainhpwithin Relationship and Persicaria of Monophyly The Bistorta Fg ,3.Ti stegopta aado (1978) Haraldson that group the is This 3). 2, (Fig. and , ieie hr ssrn upr o itrgroup sister a for support strong is there Likewise, . a eta toiiae hog yrdzto.Re- hybridization. through originated it that be may Koenigia (here Aconogonon Fallopia Fagopyrum Aconogonon sn nrnsqecso igecp nuclear copy single a of sequences intron using lorfethbi pcainado cytoplas- and/or speciation hybrid reflect also , — Eupersicaria Tovara Fagopyrum .punctata P. eut rmoraaye ftesprt and separate the of analyses our from Results Eupersicaria and Bie n tc 92 olnsot tal. et Hollingsworth 1992; Stace and (Bailey Persicaria eefwadsoe infcneonly significance showed and few were , rbc , Nsiooe l 03.Hwvr we However, 2003). al. et (Nishimoto Aconogonon Bistorta Persicaria eune Lm readKron and Frye (Lamb sequences L and i.2.I otat efudstrong found we contrast, In 2). Fig. ; a enpooe rvosy(Stan- previously proposed been has Persicaria ofit ocrighigher-level concerning Conflicts . ), ssmtmscrusrbds as so circumscribed sometimes is Bistorta Tovara and , ö eadL and ve n elspotdclade well-supported a and Echinocaulon opsdo h sections the of composed , .punctata P. a ensgetdbased suggested been has Koenigia , u oexclude to but , Echinocaulon ö Koenigia, Polygonum e15) n by and 1956), ve Persicaria teABK clade; A.B.K. (the nlddi Fig. in included Bistorta , Cephalophilon and , .punctata P. Persicaria ihnine with , Koenigia n the and Cepha- Cepha- nthe in Per- ö ve P. , , Delivered by Publishing Technology to: Yale University IP: 130.132.173.179 on: Fri, 02 Aug 2013 21:54:39 Copyright (c) American Society for Plant Taxonomists. All rights reserved. oge l 98 os ereee l 00 and 2000) al. et Decraene Ronse 1998; al. et 1988; Akeroyd Hong and by Decraene supported (Ronse been evidence has morphological this and genera), as recognized monly include 08 I OOHE OEUA HLGN FPRIAI 83 PERSICARIA OF PHYLOGENY MOLECULAR DONOGHUE: & KIM 2008] fls hn5%bosrpvle r05pseirpoaiiyi indicate is probability posterior 0.5 or values bootstrap 50% parsimony maximum than for less values of Bootstrap branches. the under presented are F The IG .5%cnesste rmBysa neecsfrtecmie aasto set data combined the for inferences Bayesian from tree consensus 50% 3. . Persicarieae Persicaria , a salse yHrlsn(98 to (1978) Haraldson by established was Aconogonon , Bistorta and Koenigia rbc (com- se- L dby--. aiu ieiodaaye r rsne bv h rnhs Support branches. the above presented are analyses likelihood maximum / iegs(i.3.Suiso lrlcaatr aesuggested goneae have characters of floral results transfer these of the Our of Studies 3). comprised 2003). (Fig. group Kron lineages monophyletic and a support Frye strongly (Lamb analysis quence orcDAmresadnISsqecs otro probabilities Posterior sequences. nrITS and markers cpDNA four f into ) Persicarieae Fagopyrum RneDcan n kry 1988; Akeroyd and Decraene (Ronse wihHrlsnpti the in put Haraldson (which Poly- Delivered by Publishing Technology to: Yale University IP: 130.132.173.179 on: Fri, 02 Aug 2013 21:54:39 Copyright (c) American Society for Plant Taxonomists. All rights reserved. iecae(including clade sive name the between the relationship and sister a support not do was analyses this but 2000), the al. in et supported Decraene Ronse 1998; al. et Hong rum Koenigia ol eueu oci e aefrtemr inclusive more the for name new a coin clade. to useful be would of time. present position the the at supported If better is clade circumscription this initial because the and was this because both clade clusive ob aahltc wn oawl-upre lcmn of of placement analyses well-supported a sagittata to P. owing paraphyletic, be to rbcL isof cies a otcoeyrltdto related closely most was the where trees, nrITS, combined in and the unresolved, in in and being genes exceptions lin- chloroplast analyses the three our in other of supported the three strongly of of was comprised relationship clade This a eages. of group sister the the caria among relationships for support base ambiguous Bank. of the number in high relatively pairs the reflect or markers. misidentification, may of other it case our a of represent all may discrepancy on This based placement its with incides i,adte nywt eywa upr.Orcombined Our support. of weak monophyly the very support with strongly analyses only then and sis, of of phyly populations two Tovara sequences the nrITS show the noted also be should it However, supported. Tovara that suggested analyses in hand, other the On clade. this to ter o togyrsle nM n Laayi Fg ,3.How- lon 3). 2, (Fig. among analysis relationships ML ever, and MP in resolved strongly not 97 ijntbtenesenAi ( Asia filiformis eastern between disjunct 1997) within lineages major highlight the briefly of we Persicaria characteristics Here present. distinctive at traits the these interpret of to evolution difficult the and is trees, best our across fashion connect weakly including ses 98 os ereeadAeod18) olni fthe of is Haraldson Pollen 1952; 1988). Akeroyd Li and 1933; Decraene Small Ronse 1930; 1978; (Steward parts anth 1 re- of dispersal having fascicles fruit arranged racemes a motely interrupted as whip-like, (presumably and tip the mechanism), at styles, persistent hooked long, are two which 1962). having in (Hara distinctive Tertiary is middle group the This since existence in been have 4SSEAI OAY[oue33 [Volume BOTANY SYSTEMATIC 84 h lcmn fEpriai ihnPersicaria within Eupersicaria of Placement The vrl,orrslspoiehge eeso eouinand resolution of levels higher provide results our Overall, Tovara, mosaic a in distributed are characteristics Morphological eanpol eovd np- In resolved. poorly remain .W ao h plcto of application the favor We ). Eupersicaria reo abFy n rn(2003) Kron and Frye Lamb of tree hnpeiu tde.Thus, studies. previous than lstermidrof remainder the plus Echinocaulon hni sto is it than rt oeicuiecae(loincluding (also clade inclusive more a to or ) n atr ot mrc ( America North eastern and ) Persicaria n eea rista r hrdbtenthem. between shared are that traits several and , Eupersicaria opsdo he pce Hr 92 u tal. et Suh 1962; (Hara species three of composed Persicarieae Echinocaulon (of rbcL rbcL Echinocaulon Eupersicaria a ekyspotda itrt clade a to sister as supported weakly was eune place sequences rbcL euneof sequence Fagopyrum lsABK ld Fg ) Consequently, 3). (Fig. clade A.B.K. plus elotieof outside well , Echinocaulon a nysgetdi aeinanaly- Bayesian in suggested only was ol eapidete oals inclu- less a to either applied be could td Lm readKo 03.Our 2003). Kron and Frye (Lamb study Persicaria rbcL, Eupersicaria and Eupersicaria ihni.I otat u own our contrast, In it. within ) Tovara with Tovara Eupersicaria hr hs eainhp were relationships these where – suhl nftr nlssit analyses future in upheld is .sagittata P. lwr,ec ihfu peri- four with each flowers, 3 u hswsvr weakly very was this but , , .sagittata P. Cephalophilon Tovara matK and , Aconogonon Persicarieae ula iooa ITS ribosomal Nuclear . smr lsl eae to related closely more is Persicaria Eupersicaria .amphibia P. “ , .filiformis P. psbA sections Tovara, Eupersicaria Echinocaulon analyses hspsil para- possible This . Fg 3). (Fig. .virginiana P. eoie nGen- in deposited and Eupersicaria ihohrspe- other with ld tefwas itself clade and ” , otels in- less the to per obe to appears trnL n hsco- this and , within Bistorta ssse of sister as Eupersicaria and Fagopyrum Echinocau- appeared – — F a sis- was Fagopy- ,may ), .neo- P. analy- nthe In Persi- and and a pct nlrsecsadsml evs sdfiutto difficult is leaves, simple and inflorescences spicate instance, has For 1988). in Park found 1978; those (Haraldson to inflorescences similar have and of species Some 1946). lon (Hedberg reticulum the to several are spiny there have However, also to trichomes. similarities species stellate Most stiff leaves. and/or triangular to hastate Persicaria 1946). too (Hedberg form here to pollen but relationship intermediate close an 1946), a exhibits of (Hedberg hint reticulum a is a there and pores cular of grains egalensis of species in some however, variable num- vidual; quite Perianth 1978). is api- (Haraldson ber trichomes acute similar with and leaves ces, elliptic simple inflorescences, of similar Plants leaves. simple ceolate caria ridge. exine Tovara litct vt evswt utniglbsadwinged and with lobes form nocaulon inflorescence subtending capitate the with share They leaves petioles. ovate to elliptic stem. the on prickles recurved of ence of species from distinguish the u frltvl ihreierde Hdeg14) The 1946). (Hedberg gonon ridges of reticu- exine grains a higher pollen bearing relatively and of furrows lum narrow with tricolpate being grains. pollen their in nocaulon of members However, lophilon 1927). Danser 1893; (Dammer nocaulon aldmrhlgclaayi.Hwvr u re suggest de- trees more first our broader, the However, a that analysis. require morphological will traits tailed these in homoplasy 1946; (Hedberg columellae 1977). fewer Skvarla and and in Nowicke others larger to have similar exines are but grains), nlrsecsapa ohv rgntdin originated have to appear indepen- inflorescences evolved have in may dently inflorescences pollen tricolpate Capitate and parts, grains. perianth five with flowers cences, n ebr 90 n nthe in and 1990) Hedberg and within again possibly and Tovara opie fteefu iegs ntr,i o per that appears of now group it sister turn, In the lineages. four these of in comprised 2000) al. et lon Decraene of Ronse monophyly 1998; the Ak- al. confirming et and Hong Decraene 1988; Ronse eroyd 1978; (Haraldson work phological Echinocaulon in independently evolved have may pollen lnsof Plants of Plants Finally, salsigtedrcino vlto n h aueof nature the and evolution of direction the Establishing nsmay u oeua tde upr rvosmor- previous support studies molecular our summary, In and , aktecaatrsi pn rclso tlaetrichome stellate or prickles spiny characteristic the lack Persicaria ysielk nlrsecswt ayfoesadlan- and flowers many with inflorescences spike-like by tp,wt 2oln oe n eiaertclmof reticulum delicate a and pores oblong 12 with -type, orprat at a aebe eie in derived been have may parts perianth Four . nbigtiopt adto (and tricolpate being in ossetypouefu einhprs h pollen The parts. perianth four produce consistently , aktemliellrsiftihmsfudin found trichomes stiff multicellular the lack n ls linebetween alliance close a and , Cephalophilon Hrlsn17) n hs rusdfe markedly differ groups these and 1978), (Haraldson Eupersicaria a enpooe ae ngosmorphology gross on based proposed been has ytercptt nlrsecsadsgtaeor sagittate and inflorescences capitate their by Cephalophilon Cephalophilon Echinocaulon Eupersicaria tp;plprt n ihardlk structure rod-like a with and polyporate -type; clade. Persicarieae Eupersicaria Cephalophilon Persicaria r fthe of are n h xsec fa of existence the and , Cephalophilon lnsaecaatrzdb broadly by characterized are plants n in and r edl itnuse rmother from distinguished readily are r itnuse rmother from distinguished are a aehdpnclt inflores- paniculate had have may Eupersicaria . Eupersicaria Echinocaulon Eupersicaria sapeiul nae clade unnamed previously a is Eupersicaria r iia otoeof those to similar are Persicaria- Eupersicaria Echinocaulon Eupersicaria Bistorta risaeo hr type, third a of are grains vnwti nindi- an within even , ial,polyapeturate Finally, . Persicaria Eupersicaria Cephalophilon Tovara olngan r of are grains pollen xetfrtepres- the for except ye aig2 cir- 20 having type, , .bungeana P. Tovara ihtricolporate with and n spike-like and , Eupersicaria uhas such , Koenigia Persicaria ,as nta their that in Tovara , Eupersicaria .amphibia P. Echinocau- Echinocau- - Tovara to which , Acono- (Hong Cepha- Tovara .sen- P. Persi- clade Echi- Echi- Echi- have and - Delivered by Publishing Technology to: Yale University IP: 130.132.173.179 on: Fri, 02 Aug 2013 21:54:39 Copyright (c) American Society for Plant Taxonomists. All rights reserved. ars .S,M K M. S., J. Farris, K M. S., J. Farris, the of phylogeny DNA chloroplast A 1994. Palmer. D. J. Noise and R. 2000. S. Quicke. Downie, J. L. D. and Orme, L. D. C. Belshaw, R. com- K., Dolphin, versus Separate 1995. Kim. J. and Donoghue, J. M. A., deQueiroz, Niederlaendisch-Ostindiens. Polygonaceen Die 1927. H. B. Danser, 1 Pp. Polygonaceae. 1893. U. Dammer, otn .adA .Asi.20.Icesdcnrec osntnec- not does congruence Increased 2002. Austin. D. A. and M. Dowton, length incongruence the does When 2002. Lecointre. G. and P. Darlu, data when predict tests incongruence three Can 1997. W. C. DNA Cunningham, Chloroplast 2004. Jansen. K. R. and Raubeson, A. L. E., M. Cosner, da,R .20.MSL:mlil euneainetwt ihac- high with alignment sequence multiple MUSCLE: 2004. C. R. Edgar, upr fNc elns nteYl ebru n m ito h New the of Litt Amy the Garden. and for Herbarium Botanical grateful Yale York the to are in we foundation Cellinese Finally, Niarchos Nico Fel- History. of the Enders support Natural from F. of grant Museum John a Peabody a and the by University provided Yale National was at (Seoul studies lowship these Kim (Kunming STK for Deng Min-Ha facilitating Funding Ming Korea. for and and Korea) China) Ma Seoul, Yunnan, Le University discus- Chang Botany helpful thank of for to lab Institute like Donoghue meth- would the for in We Smith others sion. Stephen to and Moore and Brian advice, to odological problem, this in interest our osu,L . .I awc,adJ cel.19.Alzm aito in variation Allozyme 1991. McNeill. J. and Warwick, I. S. L., L. Consaul, 531 Pp. Polygonaceae. 1993. J. Brandbyge, floral and Biogeography incongruence 1998. Wendel. the F. of J. and utility Small, L. The R. 2002. A., D. Lutzoni. Baum, M. F. and F. morphology, K. number, Barker, Chromosome 1992. Stace. A. C. and P. J. Bailey, 85 PERSICARIA OF PHYLOGENY MOLECULAR DONOGHUE: & KIM the to corresponds that clade a form these includes that 2008] between Within uncertain. and Cephalophilon with linked is turn in which Fagopyrum authors, other and Haraldson oeo yrdzto n oyliyi h vlto of evolution the the of in studies polyploidy persicaria detailed and more hybridization for of stage role the set analyses These of our placement between the in conflict punctata sequences nrITS significant and is markers cpDNA there Finally, limited. A Within CKNOWLEDGMENTS infcnets o incongruence. for test significance incongruence. of cance restriction repeat inverted and variation. structural site on based Caryophyllales length incongruence the of test. results difference interpreting incongruence: and evidence. Systematics phylogenetic and of analysis bined lien uayadhg throughput. high and curacy analyses. mixed-model in Biology test tematic difference the length of incongruence behavior accuracy-the phylogenetic increased indicate essarily fail? test difference S Botanique, Jardin du letin combined? be should highly of utility genomes. phylogenetic rearranged Campanulaceae: in rearrangements 2261 the plants vascular sets. data ( multiple Baobabs of evolution test. different length genus the in hybrids and (Polygonaceae). species of values DNA and pairing, Eupersicaria oyou lapathifolium Polygonum o.3 d .Ege n .Pat.Lizg nemn Verlag. Engelmann Leipzig: Prantl. K. and Engler A. ed. 3, vol. – hc a elc yrdzto nisancestry. its in hybridization reflect may which , 2270. .amphibia P. n t relatives. its and Persicaria . n ld aeu of up made clade a and ä ä Aconogonon llersj llersj u eainhp mn hs he remain three these among relationships but , ytmtcBotany Systematic o.2 d .Kbtk ta.Bri:Srne Verlag. Springer Berlin: al. et Kubitzki K. ed. 2, vol. 1 19 51: oeua hlgntc n Evolution and Phylogenetics Molecular 6 657 26: eaegaeu oSnaSla o stimulating for Sultan Sonia to grateful are We . ln ytmtc n Evolution and Systematics Plant Eupersicaria ö ö u nlssspota nta pi into split initial an support analyses our ytmtcBiology Systematic .G lg,adC ut 94 etn signifi- Testing 1994. Bult. C. and Kluge, G. A. , n h et huhorsmln sstill is sampling our though rest, the and .G lg,adC ut 95 osrciga Constructing 1995. Bult. C. and Kluge, G. A. , oeua ilg n Evolution and Biology Molecular – ytmtcBiology Systematic L M vltoayBiology Evolutionary BMC 31. oeua ilg n Evolution and Biology Molecular – ITERATURE 681. Cladistics Adansonia é , rie complex. Bistorta uli cd Research Acids Nucleic 8 117 38: hr per ob epsplit deep a be to appears there 9 236 19: – 0 315 10: obcca)a nerdfrom inferred as Bombacaceae) , 6 in 36. ytmtcBiology Systematic C – – ITED 7 181 47: 4 in 544 261. aainJunlo Botany of Journal Canadian and – 1 625 51: – i nat Die 252. ’ il oki hn and China in work field s 319. nulRve fEcology of Review Annual Echinocaulon h aiisadgnr of genera and families The – Koenigia 207. – ü 637. 8:29 180: :1 4: lce Pflanzenfami- rlichen Persicarieae 9 432 19: – 2 1792 32: 7 401 17: 17. 4 570 44: 4 733 14: Together, . – 52. , – 437. – Tovara – – 406. – Fallopia 572. 1797. 740. Eu- Bul- Sys- 69: of P. , esnti,J 95 ofdnelmt npyoeis napoc usin approach an phylogenies: on limits Confidence 1985. J. Felsenstein, 1969. A. Federov, abFy,A .adK .Ko.2003. Kron. A. K. and S. A. Frye, Lamb aa .1962. H. Hara, Polygon les sur Remarques 1913b. H. Gross, Beitr 1913a. H. Gross, oik,J .adJ kal.17.Ple opooyadterelation- the and morphology Pollen 1977. Skvarla. J. and novel W. A J. Nowicke, T-COFFEE: 2000. Heringa. J. and Higgins, D. C., Notredame, incon- Topological 2003. Hasegawa. M. and Ohnishi, O. Y., Nishimoto, the of study biosystematic A 1980. phylogenetic B. for C. McDonald, Testing 1996. Kellogg. A. E. and J. R. Mason-Gamer, L genus The 1952. H.-L. Li, DNA chloroplast Two 2005. Jansen. K. R. and Choi, K.-S. K.-J., Kim, og .P n .Hdeg 90 aalleouino pruenumbers aperture of evolution Parallel 1990. Hedberg. O. and S.-P. Hong, phy- versus Congruence 2004. Sytsma. J. K. and Hall, C. J. L., A. Hipp, genus The 1997. genus O. Hedberg, the in morphology Pollen 1946. O. Hedberg, subfam. Polygonaceae in taxonomy and Anatomy 1978. K. Haraldson, ese,C .15.Plgnca.P.1 Pp. Polygonaceae. 1856. F. C. Meisner, 1826. F. C. Meisner, L inversions minute create Hairpins 1996. Wendel. F. J. and A. S. Polygona- Kelchner, an studien karyologische und Histologische 1928. R. Jaretzky, marks inversion DNA chloroplast A 1987. Palmer. D. J. and K. of R. inference Jansen, Bayesian MrBayes: 2001. Ronquist. F. and P. J. Huelsenbeck, og .P,L .RneDCan,adE mt.19.Sseai sig- Systematic 1998. Smets. E. and DeCraene, Ronse P. L. S.-P., Hong, Ferris. C. and Hollingsworth, M. P. Bailey, P. J. L., M. Hollingsworth, ot .B n .D amr 94 tutrlrarneet,including rearrangements, Structural 1994. Palmer. D. J. and B. S. Hoot, ö ö ve, ve, Nauka. vlto nPolygonaceae. in evolution b 783 bootstrap. the hpo lmaiaee oyoaeeadPiuaeet h order the to Primulaceae Centrospermae. and Polygonaceae Plumbaginaceae, of ship alignments. ogy sequence multiple for method genus the of group gonaceae). hy- urophyllum suggesting the trees in DNA bridization chloroplast and nuclear between gruence iperoides tribe the Biology in Systematic sets data molecular among conflict Notiser American of 1783 evolution (Asteraceae). early family the sunflower the during simultaneously originated inversions 262. n ragmn ntegenera the in arrangement and difference length incongruence the test. revisiting accuracy: logenetic 1 22: emend. Meisn. Polygonoideae Botany l de letin eai auai en Vegetabilis Regni Naturalis tematis 670. DNA. chloroplast of regions non-coding in of States ceen. United the of Sciences of Academy (Asteracceae). America National family sunflower the the of in Proceedings split evolutionary ancient an trees. phylogenetic genera. related iiac ftplsraemrhlg ntribes in morphology surface tepal of nificance (Polygonaceae). ( knotweed inva- giant Polygonaceae). and between knotweed hybridization Japanese of and populations sive variation DNA Chloroplast 1999. Society Linnean the of Journal Botanical significance. taxonomical its and aallivrin,wti h hools eoeof genome chloroplast the within inversions, parallel 91 goneae ü Á Á – hrf cher n .L D. and . n .Sra.15.Ha oeac of tolerance Heat 1957. Sarkar. P. and . – 116. 0:205 302: 791. – – ytmtcBiology Systematic 1792. Jahrb 95. (Polygonaceae). 7 326 37: ü 1:478 110: ’ 4 5818 84: Plgnca)complex. (Polygonaceae) Systematik r cdmeItrainl eGorpi Botanique Geographie de Internationale Academie oyou.Cnda ora fBotany of Journal Canadian Polygonum. ü Sunania ee eei Systems Genetic & Genes hrf cher – ö 217. e 96 hoooeadtxnm fesenNorth eastern of taxonomy and Chromosome 1956. ve. – vlto;ItrainlJunlo rai Evolution Organic of Journal International Evolution; hoooenmeso lwrn plants flowering of numbers Chromosome 332. – ora fMlclrEvolution Molecular of Journal oaia ora fteLnenSociety Linnean the of Journal Botanical oorpiegnrsplgn prodromus polygoni generis Monographiae – Grana mtsna otiuint Botany to Contribution Smithsonian ä 481. ü 5822. ezrKnti e Polygonaceen. der Kenntnis zur ge isncatih Botanik Wissenschaftliche r 5 524 45: ai.versus Rafin. iifrais(xod England) (Oxford, Bioinformatics 9 234 49: Tovara 9 177 29: 3 81 53: oaia ora fteLnenSociety Linnean the of Journal Botanical Koenigia – 545. ytmtcBotany Systematic – (Polygonaceae). – 339. – 89. aezySmoa oaia Upsalienses Botanicae Symbolae Jaretzky 184. vnkBtns Tidskrift Botanisk Svensk onga Persicaria Koenigia, .eed ebr (Polygonaceae). Hedberg emend. L. Tovara o.1,e.A eCnol.Paris. Candolle. de A. ed. 14, vol. mrcnJunlo Botany of Journal American oeua ilg n Evolution and Biology Molecular 8 139 78: – 8,693 186, rbcL 2:295 124: é Adanson. sd l de es ongailnia Botaniska islandica. Koenigia hlgn n character and phylogeny – urn Genetics Current ora fMlclrBiol- Molecular of Journal 153. 9 357 69: Rhodora – 8 274 38: 8 326 28: – 9 in 695 Triticeae Persicarieae 330. ’ seOrientale. Asie Polygonum 4 501 34: oyou hydrop- Polygonum ora fJapanese of Journal Fagopyrum 7 1 37: – 7 754 17: and – 490. – oaiceJahr- Botanische rdou Sys- Prodromus 4 19 54: 281. 332. 2:139 129: Anemone (Gramineae). Leningrad: . Geneva. . 3 7 23: – – 0 371 40: Aconogonon 521. 64. – and – 755. .s lat. s. L. 0 259 30: 7 664 67: Fallopia 25. – 32. – (Poly- – 154. Poly- Bul- 404. 127: and 22: 39: g – – , Delivered by Publishing Technology to: Yale University IP: 130.132.173.179 on: Fri, 02 Aug 2013 21:54:39 Copyright (c) American Society for Plant Taxonomists. All rights reserved. twr,A .13.TePlgnca fEsenAsia. Eastern of Polygonaceae i The variation 1930. of N. A. cause Steward, a as hybridism of Possibilities 1925b. in E. flower-form E. Stanford, and inflorescence The 1925a. E. E. Stanford, 1933. K. J. phy- Small, DNA Chloroplast 1997. Stuessy. F. T. and Crawford, J. D. T., Sang, u,Y,S i,adC-.Pr.19.Apyoeei td of study phylogenetic A 1997. Park. C.-W. and Kim, S. Y., Suh, of Taxonomy 1988. C.-W. Park, isn .16.Asuyo yrdzto in hybridization of study A 1964. G. J. D. Timson, and Jeanmougin, F. Plewniak, F. Gibson, J. T. D., J. endo- Thompson, restriction from Inference Phylogenetic 1983. R. A. Templeton, 33 [Volume BOTANY SYSTEMATIC signifi- Systematic 2000. Smets. E. and Hong, S.-P. P., L. Decraene, in Ronse limits Generic 1988. Akeroyd. R. J. and P. L. Decraene, Ronse plants. of genomes Chloroplast 2005. Jansen. K. R. and A. L. DNA of Raubeson, model the testing Modeltest: 1998. Crandall. A. K. and D. Posada, of phylogeny interspecific and Intra- 2000. Ohnishi. O. and T. Ohsako, 86 ht,T . .Bus .Le n .Tyo.19.Apiiainaddirec and Amplification 1990. Taylor. J. and Lee, S. Bruns, T. J., T. White, in- DNA Chloroplast 2000. Yoshimura. K. and Suyama, Y. Y., Tsumura, primers Universal 1991. Bouvet. J. and Pautou, parsimony G. using Gielly, L. analysis P., Phylogenetic Taberlet, PAUP*. 2002. L. D. Contrasting Swofford, 1998. Brosi. J. B. and Hann, D. S. Wilczek, M. A. E., S. Sultan, oe,A . .A ri,adB .Pyer 01 alr fteIDto ILD the of Failure 2001. Payseur. A. B. and Irwin, A. J. D., A. Yoder, h ryHraimo avr University Harvard of Herbarium Gray the Rhodora Polygonum. subgenus Garden. Botanical Botany of of biogeography Journal and evolution, reticulate logeny, 301 Polygoneae sect. ac ffutmrhlg n ntm ntribes in anatomy and morphology fruit of cance acters. num ing. plants, higher in variation 45 Pp. substitution. naceae). DNA. Botany chloroplast of in regions noncoding of quences wild utpesqec lgmn ie yqaiyaayi tools. analysis Research quality Acids by for aided strategies alignment flexible sequence interface: multiple windows X clustal The 1997. Higgins. evolu- apes. the the to and reference humans particular of with tion maps site cleavage nuclease DNA. somal ilg n Evolution and of Biology populations in polymorphism version (Botany) Society Linnean of Journal DNA. chloroplast of regions Biology non-coding Molecular Plant three of amplification for Associates. Sinauer Sunderland: 4.0b10. v. Methods), other (*and annual co-occuring of breadth ecological fEcology of eemn aacmiaiiyfrso oi phylogeny. loris slow for Biology combinability data determine Press. Academic Diego: San White. Pp. T. phylogenetics. and Innis for genes RNA 315 ribosomal fungal of sequencing – – 337. 2 in 322 Tovara n eae eea(oyoaee ntebsso lrlchar- floral of basis the on (Polygonaceae) genera related and Fagopyrum – oaia ora fteLnenSociety Linnean the of Journal Botanical 0 408 50: 8in 68 eor fteNwYr oaia Garden Botanical York New the of Memoirs 7 573 87: escra Rhodora Persicaria. 6 363 86: (Polygonaceae). C rtcl:Agiet ehd n application and methods to guide A protocols: PCR Plgnca)bsdo T eune fncerribo- nuclear of sequences ITS on based (Polygonaceae) aulo h otesenFlora Southeastern the of Manual iifrais(xod England) (Oxford, Bioinformatics ora fPatBiology Plant of Journal ln iest n vlto:gntpcadphenotypic and genotypic evolution: and diversity Plant – 4 4876 24: 424. – 4 1120 84: – Plgnca)seisbsdo uloiese- mucleotide on based species (Polygonaceae) 582. 383. 7 81 27: 7 1302 17: – 7 1105 17: 4882. d .J er.Ofrsie AIPublish- CABI Oxfordshire: Henry. J. R. ed. – – 1136. oaia ora fteLnenSociety Linnean the of Journal Botanical 89. Polygonum 7 41 27: – Evolution 1312. – 1109. 9 155 59: – 47. 0 47 40: section Polygonum 7 221 37: – :1 5: Abies Polygonum 160. – 52. e ok e York New York: New . – 4 817 14: 8 321 98: 129. Echinocaulon – and 224. 7 1 47: otiuin from Contributions aoi.American Paeonia. section mrcnJournal American sg.Molecular Tsuga. – – Persicarieae species. 818. 371. – 82. Polygonum Polygonum Systematic Persicaria. d.M. eds. , (Polygo- Polygo- Nucleic Journal 134: and n t , EF653786. s.n Lundgren EF653784. EF653787. Crete-39 EF653761, Chae EF653710, EF653735, EF653684, EF653807. EF653806. * EF653780, Gray; EF653728, EF653754, EF653703, (P2); EF653805. EF653779, EF653727, collection. living Garden for Botanical (1998) York al. New speci- et Colorado. from herbarium Sultan from Sample = = done see *** was CO CT men; extraction University DNA Jersey, York, Wesleyan ** New locality; New in specific = more line = NJ Inbred NY given. * Massachusetts, is Notes: = herbarium MA another unless Connecticut, YU at ited EF653737, EF653790. amplexicaulis EF653764, torta EF653712, EF653738, EF653687, China; Yunnan, G M. Schlechtend.) F575 F579 F571 EF653797. EF653771, EF653719, EF653745, EF653796. EF653770, EF653718, ana EF653744, EF653693, USA; CT, Sect. EF653808. EF653809. EF653782, EF653730, Small; EF653756, EF653705, USA; hydropiper P. EF653803. * EF653777, EF653725, EF653751, (P2); 600 Kim F579 EF653785. EF653759, EF653788. EF653762, EF653711, dense EF653736, EF653685, rea; EF653792. EF653766, esculentum EF653714, rum EF653740, Ohashi; EF653689, Hiroyoshi China; & nan, Yonek. f.) Hook. ex EF653791. (Wall. EF653765, EF653713, EF653739, EF653688, USA EF653801. EF653798. EF653772, EF653720, EF653746, EF653695, rsn td.Txn olco()adnme,lclt;Gnakacces- Genbank locality; nrITS, number, for number and sion collector(s) Taxon; study. present F567 F578 F572 EF653774,EF653800. Gaertn.; Sect. EF653722, EF653795. EF653748, EF653769, EF653697, filiformis EF653717, P. EF653743, Tovara EF653692, China; nan, EF653794. EF653768, EF653716, runcinata EF653742, EF653691, China; Yunnan, Ch-Ko-48, EF653793. EF653767, Ma & Kim Sect. EF653799. Gross; on,N . .E tie,adC .dPmhls 99 h evolution The 1999. dePamphilis. W. C. and Steiner, E. K. D., N. Young, WEIR.8 eu Persicaria Genus te genera Other A PPENDIX Nakai; une euea vltoaytasto series. Garden transition Botanical souri evolutionary se- an gene refute plastid quences Scrophulariaceae/Orobanchaceae: in parasitism of i i Ch-Ko-91 Kim & Kim L)Holub; (L.) i 560 Kim i 650 Kim EG.3 J S;E639,E635,E632,E637,EF653802. EF653776, EF653724, EF653750, EF653699, USA; NJ, , T S;E630,E635,E632,E637,EF653804. EF653778, EF653726, EF653752, EF653701, USA; CT, , utns.n. Sultan Bc.Hm xD o)Masam.; Don) D. ex (Buch.-Ham. —— .pensylvanica P. oyou aviculare Polygonum i&KmCh-Ko-89 Kim Kim& .VuhrifrainfrDAetatosue nthe in used extractions DNA for information Voucher 1. T S;E630,E635,E632,EF653781, EF653729, EF653755, EF653704, USA; CT, , ,M,UA F563 F574 F579 EF653760, EF653709, EF653734, EF653683, USA; MA, ., rt,Gec;E638,E633,E630,EF653758, EF653707, EF653732, EF653681, Greece; Crete, , L)Oi;(P1) Opiz; (L.) F573 EF653789. EF653763, , — ehlpio .capitata P. Cephalophilon T S;E630,E635,E633,EF653783, EF653731, EF653757, EF653706, USA; CT, , Moench.; T S;E639,E634,E632,EF653773, EF653721, EF653747, EF653696, USA; CT, , cioalnP arifolia P. Echinocaulon cngnnmolle Aconogonon D o)Greene; Don) (D. T S;E639,E634,E632,EF653775, EF653723, EF653749, EF653698, USA; CT, , i 504 Kim — ó Nakai; ongaislandica Koenigia .nepalensis P. e;** mez; Sect. unn hn;E639,E634,EF653715, EF653741, EF653690, China; Yunnan, psbA L)M G M. (L.) i i Ch-Ko-96 Kim & Kim escraPriai amphibia Persicaria Persicaria 6 876 86: T S;E638,E633,EF653708, EF653733, EF653682, USA; CT, , i i Ch-Ko-102 Kim & Kim ,p- itr rw&Co 4083 Crow & Crow Ritter, ynSnNmo oe;EF653700, Korea; KyungSangNamDo, , i 570 Kim matK ynSnNmo oe;EF653694, Korea; KyungSangNamDo, , Min)H Gross; H. (Meisn.) – L.; ó 893. D o)Hara; Don) (D. mez; i 710 Kim i aCh-Ko-37 Ma & Kim , oyoel articulata Polygonella utn&Hshls.n., Heschel & Sultan rbcL L.; T S;E630,EF653753, EF653702, USA; CT, , L)K aado;** Haraldson; K. (L.) Bc.Hme .Dn .Gros; H. Don) D. ex (Buch.-Ham i,Dngu utn14 Sultan & Donoghue Kim, *uzs.n. **Lutz , trnL i egCh-Ko-62 Deng & Kim .cespitosa P. (*** .meisneriana P. ynSnNmo Ko- KyungSangNamDo, , – i aCh-Ko-8 Ma & Kim F c.Nm 1797/94 Num. Acc. pcmn r depos- are Specimens . i egCh-Ko-54 Deng & Kim yngD,Korea; KyunggiDo, , O S;EF653686, USA; CO, , i aCh-Ko-50 Ma & Kim ue spp Rumex .sagittata P. L)S .Ga;(P1) Gray; F. S. (L.) naso h Mis- the of Annals .lapathifolia P. .virginiana P. oii (NHA); Bolivia , unn China; Yunnan, , Bue Nakai; (Blume) itrapaleacea Bistorta Meisn.; alpascan- Fallopia ae 89-56 Magee .punctata P. Ca.& (Cham. T USA; CT, .maacki- P. ; Fagopy- L)H. (L.) i & Kim i & Kim Yun- , Yun- , ,NY, ), MA, , Bis- (L.) (L.) P. , , ,