Delivered by Publishing Technology to: Yale University IP: 130.132.27.226 on: Tue, 25 Sep 2012 14:55:00 Copyright (c) American Society for Plant Taxonomists. All rights reserved. pnteNwadOdWorlds. Old and New the of exception span the With 1988). (i.e. Wang and species few (i.e. contain Hemisphere that Northern the in ciose in- the of among relationships each resolved analyses them. of confidently previous monophyly or the genera, the of cluded Dono- tested none in adequately However, (reviewed 2003). have sequences al. DNA et and morphology ghue 1994) on al. based et analyses (Judd recent all and in well-supported species), 15 (ca. hamel. contains Pyck clade and This Backlund 1998). sensu al. APG str. s. 1998; et Caprifoliaceae Donoghue = (sensu 2003; and 2001, Caprifoliaceae Bell of within being 2003; Caprifolieae exception tribe al. important most et the 2005), (Donoghue Donoghue Dipsacales within ships oyih 08b h mrcnSceyo ln Taxonomists Plant of Society American the by 2008 Copyright © Botany Systematic yuke mr( including Amur invasives, eysuckles costly ( become honeysuckle have Japanese and several States including United worldwide, ornamentals Lonicera have as species bred of number been A relief. inflammation for species medicines from extracts while of tribes, American Native several within of Species species carpos properties. Many Philip- medicinal 1946). the have Steenis and Caprifolieae van Malaysia, 1903; India, their (Rehder of extending pines areas species tropical several into with range Central areas, and subtropical North and to native America. being North remainder representatives the Asian China, and in while American Japan) and North (China both has cies, species), arfleeicue eeata r mn h otspe- most the among are that genera includes Caprifolieae relation- many resolved have studies phylogenetic Recent Lonicera h ao pi sbtensubgenus between is split major the Nintooa and h otseisrc ftetaiinlgnr,adissbiiin,ne ob vlae.I hssuyw sdsqecso h T region ITS the of sequences used we study this In evaluated. ( be regions to non-coding need chloroplast subdivisions, five its and and DNA genera, ribosomal nuclear traditional of the of species-rich most the eann eeawti h rb,and tribe, the within genera remaining ewe h l n e ols(..subsections (e.g. Worlds New and subgenus Old in stems the hollow between (e.g. characters morphological subgenus several in in homoplasy of instances highlights Triosteum, hlgntc fteCpioia and Caprifolieae the of Phylogenetics hools o-oigregion, non-coding Keywords—chloroplast eetpyoeei nlsso h isclssrnl upr arfleecaewti arflaeeincluding Caprifoliaceae within clade Caprifolieae a support strongly Dipsacales the of analyses phylogenetic Abstract—Recent oti laod n aoisue eiial by medicinally used saponins and alkaloids contain psbM–trnD pce hc aeecpdfo utvto nthe in cultivation from escaped have which species Lonicera aeln enue nsm rdtoa Chinese traditional some in used been long have r etdwti h aahltcsection paraphyletic the within nested are 20) 34:p.776–783 pp. 33(4): (2008), 1 rodAbrtmo avr nvriy 2Dvnt v,Cmrde ascuet 23 U.S.A. 02138 Massachusetts Cambridge, Ave, Divinity 22 University, Harvard of Arboretum Arnold and Symphoricarpos, .Frhroe u aaidct osbeisacso yrdzto nscinNnoaadboegahcdisjuctions biogeographic and Nintooa section in hybridization of instances possible indicate data our Furthermore, Lonicera). Lonicera 3 urn drs:PatSi,adIsc cecs nvriyo ascuet tAmherst, at Massachusetts of University Sciences, Insect and Soil, Plant Address: Current .maackii L. pcr oadesteepolm.Orrslsidct that indicate results Our problems. these address to spacer) .(a 0 species), 200 (ca. L. 2 eateto clg n vltoayBooy aeUiest,PO o 208106, Box P.O. University, Yale Biology, Evolutionary and Ecology of Department .japonica L. 4 Symphoricarpos smil itiue ntemperate in distributed mainly is uhr o orsodne(tesnmuaseu [email protected]) ([email protected]; correspondence for Authors ur) atra ( Tartarian Rupr.), Triosteum Triosteum eainhp ihnCpioia,hwvr eanqieucran n h oohl of monophyly the and uncertain, quite remain however, Caprifolieae, within Relationships Lonicera. iaTheis, Nina Triosteum, hn. n h uhhon- bush the and Thunb.) Caprifolium Leycesteria Leycesteria Leycesteria, .( pce) n is and species), (6 L. Lonicera and Symphoricarpos a utoespecies one just has e ae,Cneiu 62-16U.S.A. 06520-8106 Conneticut Haven, New hools N Sequences DNA Chloroplast honeysuckle, Heptacodium, ihjs i spe- six just with mes,Msahsts003U.S.A. 01003 Massachusetts Amherst, 1,3,4 n subgenus and Leycesteria omnctn dtr eaStruwe Lena Editor: Communicating and .tatarica L. Isika. n genera and ) itga n Alpigenae). and Distegiae ihe .Donoghue, J. Michael alc.(6 Wallich. Symphoricarpos h genera the Symphori- Section Hsu , Du- ihnsubgenus Within Lonicera. L.), Nintooa Lonicera rpoB–trnC 776 omacaeta ssse oamonophyletic a to sister is that clade a form and Isika, losteum including several subsections, rejected Rehder’s but of system, classification Rehder’s adopted Chinese eetees hlgntcrltosisaogadwithin problem. and the among considered agreed relationships have generally phylogenetic who Nevertheless, been botanists most have by therefore, upon sections, four der’s inwti section within tion nei oJpn( Japan to endemic oee,teegop aerrl enacpe ihthe with accepted section been of rarely exception have groups these However, sections 5 pce,cnitdo orscin ( sections four Isoxylosteum of consisted species, 150 ( honeysuckle Morrow’s and thae, 13)pooe ayscin n uscin,including subsections, and sections sections many proposed (1938) inflorescences. the subtending subgenus leaves whereas ate leaves, free Caprifolium and cymes Subgenus two-flowered subsections. four of subgenus, second Caprifolium The 1913). 1909, (Rehder within subsections combined which, of first in the subgenera two recognized (1903) Rehder changes. of treatment published Lonicera taxonomic first the comprehensive was most This and system. classification of a proposed synopsis and detail a published (1903) Lonicera Rehder evaluation. nomic ( honeysuckle Bell’s brid a lob o-oohltc u nlsso h genus the of analysis Our non-monophyletic. be also may okiza egl,Zabelia. Weigela, Kolkwitzia, nhsteteto h aaeeseisof species Japanese the of treatment his In Historically, Heptacodium Praeflorentes spacer, Dpaae)Bsdo ula and Nuclear on Based (Dipsacales) , hri ervee h aooi ieauein literature taxonomic the reviewed he wherein , Tataricae n a ic enaotdwdl ihol minor only with widely adopted been since has and eaia,Euchranthae Cerasinae, 2 Bracteatae Lonicera Breviflorae n inu Li Jianhua and ,wt prxmtl 2seis a composed was species, 22 approximately with ), atpB–rbcL a he-lwrdcmsi hrsadperfoli- and whorls in cymes three-flowered has and , sections Lonicera, Lonicera ssse oCaprifolieae, to sister is , nsection in Nintooa Ramosissimae Chamaecerasus , pce,HuadWn 18)largely (1988) Wang and Hsu species, .gracilipes L. Isika spacer, rgatsia,Monanthae Fragrantissimae, and Monanthae a eevdtems xesv taxo- extensive most the received has yHr 18) ntertetetof treatment their In (1983). Hara by Caprifolium .×bella × L. n 0sbetos ae,Rehder Later, subsections. 20 and ) Longiflorae trnS–trnG 1,4 Coelxylosteum .morrowii L. Coeloxylosteum, , n a rae sasubsec- a as treated was and ) , Subsessiliforae Chlamydocarpi hc a igespecies single a has which , Rhodanthae Lonicera (= n section and Zabel). spacer, Within Lonicera. Lonicera Spinosae Triosteum nsection in olxlsem Isika, Coeloxylosteum, ry,adterhy- their and Gray), , petN–psbM scaatrzdby characterized is Isoxylosteum, Distegiae , ,wt oethan more with ), Rotatae ssse othe to sister is Coeloxylosteum and , Periclymenum nsection in Lonicera and Nintooa. Leycesteria, , Pararhodan- Lonicera, Lonicera, Lonicera Tetramerae spacer, n sub- and , nsection in Vesicariae Lonicera and Nakai , Isoxy- Reh- (= , . Delivered by Publishing Technology to: Yale University IP: 130.132.27.226 on: Tue, 25 Sep 2012 14:55:00 Copyright (c) American Society for Plant Taxonomists. All rights reserved. psbM–trnD petN–psbM trnS–trnG - alcoigtcnqe n P and 1. technique, cloning Table tail in T-A se- given are The amplification dimethylsulfoxide. for parameters 10% and primers included quencing reactions ITS amplification. atpB–rbcL ersi erhswt 00rno euneadto elcts with replicates, addition sequence random 1000 and with included in (Huelsenbeck searches analyses (MP) v3.0b4 heuristic conducted parsimony Bayes maximum were for Mr. Options 2001). and analyses Ronquist 2001) Phylogenetic (Swofford 4.0b10 states. PAUP* data character missing as new Gen- treated were in and Gaps unordered. sequence and weighted published equally a were to lonicera ITS-2 them (Diervilla and comparing Bank 5.8S, ITS-1, by Tree- the to determined of limits submitted were Sequence been S2161). have number Matrices (study 1996). BASE (Rambaut v2.0a8 Arbor, Se-Al ing Ann Corporation, edited codes and Gene (Perkin-Elmer/ABI), 4.1, 3730 Michigan). (version or Sequencher 3100 Se- DNA PRISM using automated ABI ABI 377, an using quencer City, analyzed Foster Inc., were out Biosystems Sequences (Applied carried California). kit were V3.1 reactions Terminator hetero- BigDye sequencing sequence a Cycle using for repeats. check ITS to among species geneity representative for sequenced were oiesrmabmnwt ia ocnrto f0.2 of concentration final a with albumin serum bovine 0.4–0.6 albd aiona,200 1 California), components: Carlsbad, 25 reaction in following (10 cyclers the DNA thermal per- with Research was MJ volumes amplification or ␮L Eppendorf All manufactur- either the California). using following Valencia, formed kit (Qiagen, Mini Extract protocol Plant er’s DNeasy a using leaves for obtained newly were study. others this all 2000); out- Zhang Donoghue and as and (2003) (Gould used GenBank al. of sequences were et ITS Donoghue latter (2003). on al. The et based purposes analysis. rooting the for in groups included were (Linnaeae) of cies spe- two both included and we sections, Caprifolieae four of all other cies subsections, From 24 1). the (Appendix of subgenera 21 species, 47 resenting rpoB–trnC aiona olwn manufacturer competent following blue California) XLI and Wisconsin) n h oopcfcgenus monospecific the and ITS non-coding chloroplast five and ( 1995) regions al. ribosomal et nuclear closely (Baldwin of of region DNA ITS relationships the resolving that including genera, in regions related widely DNA used chloroplast been and have nuclear from sequences of monophyly the Lonicera test and Caprifolieae, within relationships phylo- to subjected analysis. been genetic not have subsections of numerous relationships the evolutionary and Further- circumscriptions addressed. the explicitly more, been never have sections these 2008] spacer, hwe l 2005). al. et Shaw o oto h cesos C rdcsfrISwr lnduigthe using cloned were ITS for products PCR accessions, the of most For h dtdsqecswr lge yeeus- eye by aligned were sequences edited Analysis —The Phylogenetic oeua Techniques Molecular T of individuals Material—Fifty-one Plant Regions h rmr betvso h rsn td eet resolve to were study present the of objectives primary The ABLE Weigela Leycesteria o/ ahpies n . units 2.5 and primers, each ␮mol/L .PRprmtr o h T n ieclrpatnnoigregions. noncoding chloroplast five and ITS the for parameters PCR 1. petN–psbM 0 g,1 ng), –100 n t ugnr,scin,adsbetosusing subsections and sections, subgenera, its and rpoB Devlee n n pce ahof each species one and (Diervilleae) nta denaturing Initial ,1mn55 min 1 94°C, 34x min 3 94°C, ,1mn55 min 1 94°C, 34x min 3 94°C, ,3mn3x94 34x min 3 94°C, ,1mn53 min 1 94°C, 34x min 2 94°C, ,1mn55 min 1 96°C, 35x min 5 80°C, ,3mn3x94 34x min 3 94°C, – n he pce ahof each species three and tp time step, trnC –2 M F772 i n i 99.Caatrstates Character 1999). Kim and Kim AF078722, , pcr and spacer, rmr 10 primer, ␮L TRASAND ATERIALS — spacer, o/ ahdT,1.5 dNTP, each ␮mol/L Nswr xrce rmslc-e dried silica-gel from extracted were DNAs Heptacodium. ecsei n Triosteum and Leycesteria gem reps lmdsse Poea Maddison, (Promega, system plamid atpB ’ ntutos w ofv clones five to Two instructions. s psbM × .coli E. M ufr(nirgnCorporation, (Invitrogen buffer – HI TA. HLGNTC FCAPRIFOLIEAE OF PHYLOGENETICS AL.: ET THEIS eprtr,time temperature, rmCpioica,tospe- two Caprifoliaceae, From ETHODS rbcL Symphoricarpos oeratosincluded reactions Some Taq. –trnD Lonicera Denaturing ,4 e 59 sec 45 °C, ,3 e 50 sec 30 °C, el Srtgn,L Jolla, La (Stratagene, cells spacer, Kolkwitzia . mlLMgCl mmol/L –2.0 negncspacer; intergenic eesmld rep- sampled, were eeotie from obtained were oimprove to ␮g/L –5 and template ␮L trnS eprtr,time temperature, rmrannealing Primer and Triosteum ,1mn72 min 1 °C, ,1mn72 min 1 °C, ,1mn72 min 1 °C, ,1mn72 min 1 °C, ,2mn72 min 2 °C, ,1mn72 min 1 °C, – Zabelia trnG –50 2 , ro oti on.Pseirpoaiiis(P rmteaayi are analysis the from tree. (PP) consensus scores rule probabilities majority likelihood Posterior the the on point. of shown million this gen- stabilization 5 100,000 to to for of due prior trees burn-in implemented random A was generations. from erations 100 starting every 3.0b4 run Markov sampling were version of generations, chains Carlo Bayes simultaneous Monte Mr. Four chain 2001). using Ronquist performed and (Huelsenbeck were analyses Bayesian eeotie o h rN T n iecDAspacer cpDNA five and ITS ( nrDNA the regions for obtained were rbtdwsslce o l i aast:nrITS, with sets: model data dis- six gamma (GTR) all was rbcL reversible for that selected variation time was site general among tributed and A frequencies base 2004). unequal MrModelt- (Nylander using tests 2.2 ratio est likelihood hierarchical performing by 1999). mined Hasegawa and (Shimodaira PAUP* in of Shimodaira-Hasegawa implemented the (monophyly used as predictions we test down specific broke topology where about char- addition, sections) constant and In data, included. missing as acters treated branch homogeneity gaps TBR off, replicates, partition MulTrees sequence partition swapping, addition heuristic random the 10 thousand using with PAUP* each one in replicates, using implemented was test test ILD homogeneity The 2005). al. et esa nogun fcae ihBP with data clades considered We if incon- 1995). incongruent al. methods: the as et and sets two Farris (ILD; values by test bootstrap difference evaluated length individual gruence was of sets inspection data visual through the of Congruence explored. the find constraint. to each option with search heuristic consistent the trees which using branches shortest PAUP* resolved in single the executed with then trees using Maddison of was and file obtained a (Maddison generated 4.0 were which MacClade individual 2000), (DI) in for command indices index support decay Decay PAUP estimate 1985). to (Felsenstein searches were clades replicates heuristic 1,000 with of using trees (BP) parsimonious analyses conducted most Bootstrap fol- the 15,000. per- of replicates, to all was set 1,000 on MAXTREES per search search exhaustive trees step an 5 by two with keeping lowed swapping, a autoincrease, branch to cases TBR set and other MAXTREES addition all sequence random In with off. formed effect, descent in deepest MULPARS and swapping, branch (TBR) bisection-reconnection tree aylreidl atclryi the in particularly indels large many region ITS 2 in nrDNA among the listed found of was are heterogeneity sequence indices No consistency 2. of Table range the variable the and as of se- distances well as in number pairwise sites informative variation the parsimony on length, and sites, Details alignment 1). lengths, (Appendix quence accessions 64 all spacer, hp mns arfleegnr n within and genera Caprifolieae amongst ships trnC xlddfo h nlsspeetdhr.Mxmmpair- within Maximum here. distances presented wise trees analysis been therefore the identical have from and in not excluded or resulted included were analyses indels whether parsimony the ever ein .%frthe for 2.8% region, pcr .%frthe for 3.7% spacer, h etmdlo uloiesbttto o ahdt e a deter- was set data each for substitution nucleotide of model best The ro ocmiigtedt es ofitbtendt attoswas partitions data between conflict sets, data the combining to Prior eprtr,time temperature, ula n hools aaSets Data Chloroplast and Nuclear hi extension Chain spacer, ,9 e 72 sec 90 °C, ,9 e 72 sec 90 °C, ,2mn72 min 2 °C, ,2mn72 min 2 °C, ,3mn72 min 3 °C, ,1mn72 min 1 °C, pcrrgos ayo hs nesspotdrelation- supported indels these of Many regions. spacer petN–psbM rpoB trnS–trnG – trnC ia extension Final spacer, spacer, lnswti ahseis hr were There species. each within clones –5 ,1 i adi ta.(1995) al. et Baldwin min 10 °C, trnS–trnG tp time step, ,7mnLeadWn(2004) Wen and Lee min 7 °C, ,7mnLeadWn(2004) Wen and Lee min 7 °C, ,7mnHmlo (1999) Hamilton min 7 °C, ,5mnZrwk ta.(94,Canand Crayn (1984), al. et Zurawski min 5 °C, ,5mnSa ta.(2005) al. et Shaw min 5 °C, rpoB–trnC spacer, Lonicera petN–psbM psbM R ESULTS pcr .%frthe for 2.8% spacer, atpB pcr .%frthe for 3.0% spacer, Ն –trnD esrd64 o h ITS the for 6.4% measured 4 eei ofit(..Hansen (e.g. conflict in were 84% un (2000) Quinn – pcr and spacer, rbcL negncsae)from spacer) intergenic opeesequences —Complete trnS–trnG rpoB–trnC spacer, rmrsource Primer psbM–trnD Lonicera, spacer, petN–psbM trnS and atpB–rbcL – spacer. rpoB– how- trnG atpB– 777 Delivered by Publishing Technology to: Yale University IP: 130.132.27.226 on: Tue, 25 Sep 2012 14:55:00 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 4,D )ta a itrt h monophyletic subgenera: two the Lonicera the to Within to 20). sister corresponding = clades was DI 100%; that = 2) (BP = DI 95%; = 84%, (BP Caprifolieae 6). remaining = 0.82). the DI = from split RI that 0.77, and clade is a clade = tree Caprifolieae (CI the consensus 2 to strict sister 1, the Figs. steps; in 2,433 shown of gen- trees analyses MP 2,136 characters, informative. erated 6,536 parsimony were of had which set regions of data 872 DNA combined nuclear The and steps. 504 chloroplast analysis with This trees informative. 1,915 parsimony charac- yielded were 653 with 132 contained which retained set of data were ters DNA nuclear trees The parsi- steps. 200,000 were 1,882 which over of informative, 740 characters, mony 5,883 had DNA 2005). roplast Miller and al. Levin et 2004, (Yoder al. localized et be Hipp to 2001, appear conflicts the data the because combine and sets to Darlu proceeded 2000; we Therefore, al. substi- 2002). et Lecointre (Dolphin and markers 2) of between (Table number rates tution the characters in (PI) differences informative reflect parsimony could with this repeated though clear, were tests incongruent ILD remained partitions (p The the 97%). but excluded, = taxa these (BP tree ITS to the than rather tree chloroplast fragrantissima L. henryi L. 2. Table in genera included other the of between sentatives reported are distances Pairwise asmn nomtv hrces I=cnitnyidx n I=rtninindex. retention = RI and index, consistency = CI characters, informative parsimony eersle ihpseirpoaiiiso 0%(i.1). (Fig. analyses, 100% MP nodes of the Many in probabilities As posterior burn-in. with as resolved all discarded were generations; were 100,000 generations by prior scores likelihood of stabilization subsections section within subgenus within paraphyletic, section did as clade two in evident was henryi incongruence the In the clades. that that indicated suggests gies together ( taken incongruent sets data are cpDNA they 5 all to ITS ing the for 2.5% and spacer, distance pairwise of Range (percent) sites PI (percent) sites Variable length Aligned length raw of Range 778 I RI CI, Isoxylosteum Nintooa Nintooa lhuh otta upr o hstplg a less was topology this for support bootstrap although, lium; hispidula L. T hlgntcRelationships Phylogenetic Sets Data of Congruence aeinaaye ftepriinddt e eutdin resulted set data partitioned the of analyses Bayesian .1.Tecueo h eann eeoeet sun- is heterogeneity remaining the of cause The 0.01). < ABLE ssse to sister is .Cmaio ftevraini h ula iooa ein h ieclrpatnncdn ein,adtecmie aast I= PI set. data combined the and regions, non-coding chloroplast five the region, ribosomal nuclear the in variation the of Comparison 2. Within . eent oee nteM analyses, MP the in However not. were eei w eaaecae n section and clades separate two in were Symphoricarpos Statistic ssse to sister is eentsse pce ihnsubgenus within species sister not were Pileatae eemnpyei,weessections whereas monophyletic, were Nintooa Isika .japonica L. ssse to sister is Lonicera, and , Isoxylosteum .giraldii L. omdmnpyei ldsecp for except clades monophyletic formed ld fteclrpattree, chloroplast the of clade p and Lonicera psbM .1.Cmaio fte topolo- tree of Comparison 0.01). < Vesicariae eut fteIDts compar- test ILD the of —Results B 0%,wiei h T tree, ITS the in while 100%), = (BP section Leycesteria .standishii L. –trnD B 4) rmsection From 84%). = (BP .7 0.82 0.77, 60(25) 1630 h aastfo h chlo- the from set data —The Lonicera 7 (13) 872 .hemsleyana L. 0–0.11 6536 oee,seiso section of species However, . All sections Lonicera . Coeloxylosteum Triosteum negncsae region. spacer intergenic hr eetomajor two were there omdacae(P= (BP clade a formed Lonicera Coeloxylosteum B 8)i the in 98%) = (BP l subsections All . Heptacodium si h aein case the is as .,0.71 0.6, 546 3 (20) 132 (34) 220 Caprifolium 0–0.13 pce formed species 653 ITS .etrusca L. –621 n repre- and YTMTCBOTANY SYSTEMATIC omda formed Isika Isika Lonicera Lonicera Caprifo- Isika, was was and and and and 1074–1296 .3 0.87 0.83, rpoB–trnC 9 (13) 195 (26) 384 0–0.15 1479 u hlgntcrslso isclsteei well- a is containing clade there Lonicera Caprifolieae resolved Dipsacales poorly of but supported results phylogenetic ous o otn upssw included supported. we strongly purposes been rooting not clade For have this l. of s. 2003; relationships Caprifoliaceae al. other precise et to The Zhang 2003). 1999; al. al. et et Donoghue Pyck 1996; Donoghue and lund etmlclraaye aesgetdta h monospe- the that suggested have cific analyses molecular of cent species two and Linnaeeae n a encniee ob nisontie(s 93.A 1983). (Hsu tribe own its in between connection be direct to considered been has and Lonicera of is placement the Dispacales to of support tacodium remainder add analyses the 2003). our al. of et limited, Donoghue sampling 2001; our al. et Although Bell 2000; Smets and (Pyck ld otiigtermiiggnr,floe yaclade a by followed of genera, (BP=88%) remaining the containing 1), clade (Fig. trees our In <65%. phoricarpos ld.Wieti oooyi ossetwt h Bayesian the this of with placement for the consistent for synapomorphies support is 1), clear (Fig. topology result no this of While know clade. morphological we inflores- of terminal but have analysis genera cences, Both phenetic (1983). Hsu early by characters an in suggested lead- branch the long in to exceptionally reflected ing an is with also, uniqueness data This molecular cymes. dense and leaves dium aa aat arflaa P 75). = (PP Caprifoliaeae to of taxa was placement basal there the However, for tree. support the weak in support relatively nodes was deeper the there of notably many Most tree for the analysis. of Bayesian topology the the be- for from support disagreement any higher of was placement strong there the but for not taxa, analyses MP was and Bayesian there the tween Overall 50%. than te iscls r eesr oass h lcmn of placement the assess of to sample necessary Triosteum broader are a Dipsacales, previous including stronger other than studies, provide Caprifolieae additional analyses within studies, our relationships Although for 75%). results = (PP high ( relationships: following the obtained (2003) phoricarpos found (2001) containing al. Dono- Caprifolieae. et within ghue relationships the of analyses ized negnrcRltosisWti Caprifolieae Within Relationships Intergeneric ofitn oooisadpo eouinhv character- have resolution poor and topologies Conflicting Heptacodium smrhlgclyuiu ihissrnl 3-nerved strongly its with unique morphologically is Heptacodium , B 100%). = (BP Leycesteria ihCpioia Fg.1 ) Nevertheless, 2). 1, (Figs. Caprifolieae with .6 0.91 0.86, 590–730 5 (20) 156 atpB 0–0.074 n h eainhpbetween relationship the and 2(12) 92 u ihbosrpspot<0.Zage al. et Zhang <50%. support bootstrap with but , ecsei Lonicera (Leycesteria Triosteum 798 –rbcL Leycesteria a etesse ru fCaprifolieae of group sister the be may , Fg 2). (Fig. Symphoricarpos Triosteum n h itrtaxa sister the and and .9 0.87 0.79, 578–1253 petN 9 (21) 291 9 (14) 197 0–0.082 D 1449 Leycesteria Leycesteria ISCUSSION Triosteum –psbM Symphoricarpos ),alwt otta support bootstrap with all ))), Weigela steol ebcosgroup herbaceous only the is Zabelia and , B 6)i itrt a to sister is 96%) = (BP and ob itrt clade a to sister be to .,0.94 0.9, 557–668 trnS 0 (24) 202 1 (14) 119 rmDevlee Re- Diervilleae. from 0–0.12 Triosteum Leycesteria 878 and Symphoricarpos –trnG Lonicera hc ssse to sister is which , Triosteum Triosteum Triosteum Kolkwitzia Vlm 33 [Volume nprevi- —In eg Back- (e.g. and and Heptaco- sntas not is .9 0.88 0.89, 602–1178 psbM 7 (30) 377 3 (11) 137 0–0.23 sthe as 1279 (Sym- Sym- Sym- from Hep- –trnD was Delivered by Publishing Technology to: Yale University IP: 130.132.27.226 on: Tue, 25 Sep 2012 14:55:00 Copyright (c) American Society for Plant Taxonomists. All rights reserved. hddbxsalohr ntesection the Cupulae in others all boxes shaded iete(i) uprsets(u) yeace(y) hdnhe(h) pnse(p) aaia tr,Vscra (ves). Vesicariae (tar), Tataricae (spi), Spinosae (rho), Rhodanthae (pyr), Pyrenaicae (pur), Purpurascentes (pil), Pileatae yteprioyaayi lf onr.Arnm ntergtidct aooi rusfloigRhe 10,10,11) aa(93 n s and Hsu and (1983) Hara 1913), 1909, (1903, Rehder following groups taxonomic indicate right the on Acronyms corner). (left analysis parsimony the by n eo h rnhsaebosrppretgsdcyidcs h seikidctstedfeettplg fthe of Genus: topology (1988). different the Wang indicates asterisk The indices. percentages/decay bootstrap are branches the below and eune fnDA(T) n pN ( cpDNA and (ITS), nrDNA of sequences 2008] ersn ofitbtenteclrpatadncerdt B 4)i h asmn nlss ubr bv h rnhsaepseirprobabilities posterior are branches the above Numbers analysis. parsimony the in 84%) > (BS data nuclear and chloroplast the between conflict represent F IG .Tecmie aoiyrl osnu refo h aeinaayi n h titcnesste rmteprioyaayi ae on based analysis parsimony the from tree consensus strict the and analysis Bayesian the from tree consensus rule majority combined The 1. . (cup), ypoa ch,Dseie(i) uarflu ec,Farnisme(r) ogfoa ln,Monanthae (lon), Longiflorae (fra), Fragrantissimae (euc), Eucaprifolium (dis), Distegiae (cyh), Cyhpeolae Subgenera: Lonicera. Subsections: Isika. rpoB–trnC HI TA. HLGNTC FCAPRIFOLIEAE OF PHYLOGENETICS AL.: ET THEIS Caprifolium spacer, and liea ap,Bataa ba,Beilre(r) acrte(a) hayoap cl,Ceuee(cae), Coeruleae (chl), Chlamydocarpi (cal), Calcaratae (bre), Breviflorae (bra), Bracteatae (alp), Alpigenae atpB–rbcL Sections: Lonicera. spacer, trnS–trnG Nintooa (Nin), spacer, Isoxylosteum petN–psbM pcr and spacer, ,and (Iso), (mon), Caprifolium Coeloxylosteum psbM–trnD crnhe(c) hnati(phe), Phenianthi (ocr), Ochranthae ugnsa determined as subgenus pcr.Dse lines Dashed spacer). niae in indicated (Coe) 779 Delivered by Publishing Technology to: Yale University IP: 130.132.27.226 on: Tue, 25 Sep 2012 14:55:00 Copyright (c) American Society for Plant Taxonomists. All rights reserved. phoricarpos 780 rrpeettvso ohhv enmpe notete o etosof sections for tree the onto mapped been circle) Lonicera. have is (filled both solid branches im- of (circle), representatives those to branches or above Hollow lines length. branches; branch shortened remaining numerical the indicate the regions, DNA of lines nuclear visualization Dashed and prove lengths. chloroplast of branch data showing combined the of analysis notosubgenera: two both into in clade supported well analyses. a Bayesian form and MP study our in sampled omrhvn w-lwrdcms hl h atrhas 1), (Fig. latter trees Bayesian the and the MP while both with In cymes, cymes. inflorescence, three-flowered two-flowered the having by dis- former significantly are subgenera most the tinctive, Morphologically, Afghanistan, China. America, Central North and and Europe throughout range Lonicera, species, 20 F ugnsDelimitations Subgenus IG .Pyormo h titcnesste ae naparsimony a on based tree consensus strict the of Phylogram 2. . huhisgorpia itiuini ra iha with broad is distribution geographical its though Caprifolium ugse ee l fteseisof species the of All here. suggested Lonicera stesalrsbeu ntegenus the in subgenus smaller the is edr(93 divided (1903) —Rehder and Caprifolium ihjs over just With . YTMTCBOTANY SYSTEMATIC Lonicera Lonicera spinosa o.HuadWn 18) oee,cmie h w sub- or two tube the treated combined corolla and however, the sections (1988), from Wang exerted and Hsu are not. styles and stamens and the 1988) Wang and Hsu s n Wang and Hsu Spinosae 10)rcgie w subsections: Rehder two base. the recognized acti- at nearly (1903) non-gibbous with are flowers that and corollas with leaves, nomorphic shrubs small compact pith, and white are Himalayas solid, plants eastern These in China. distributed southwestern are that species eight infcnl ogrta h nosrie hlgn as phylogeny unconstrained the than longer significantly tooa section of species representative other japonica with clade a of form placement the for although Interestingly, support bracteoles. and monotypic ovaries, the connate spur, bract- including even subsections, or three reduced subsection are are leaves There upper like. their and vines are hroe l he fteescin r etdwti section within section nested are Therefore, sections Isika. these Fur- of non-monophyletic. three is all thermore, it that suggesting clades, separate sparaphyletic. is whereas monophyly, 1), their (Fig. ing trees our 1 In buds. with Coeloxylosteum accessory having corollas in and actinomorphic nectaries, nearly and solid zygomorphic have section plants in These as section. branches, diverse most and largest the lyrcgie nsubgenus in recognized ally reighbt,cmie ihcmsfrigterminal Section tubes. forming slender long, cymes with corollas with and panicles, combined within habits, once creeping than Section 2). more are (Fig. evolved characters branches these hollow of Both with branches. Section shared hollow buds. and accessory corolla of absence the Coeloxylosteum and nectaries, unique a five sec- by example, marked For is tion characters. sections morphological these of of combination Each 1988). Wang steum sub- of monophyly the the test of further species sections. to more needed subge- Nevertheless, are the groups. subgenus separate smaller to inappropriate into be nus may it that dicating etrusca ruswt rmr pce ape ( sampled species more or hispidula 2 with groups thae spp.), subgenus of species ymtyadtefso fbates edr(93 recog- (1903) subgenus Rehder in bractlets, subsections of four nized fusion the and symmetry taxa. two these of recognition the for port subgenus L L etoso ugnsLonicera Subgenus of Sections etoso ugnsCaprifolium Subgenus of Sections ONICERA ONICERA ofr ld,sxmr tp eenee.Ti snot is This needed. were steps more six clade, a form to Isoxylosteum 1s. oee,i u hlgntctes(i.1,sub- 1), (Fig. trees phylogenetic our in However, sp.) (1 , Cyhpeolae Nintooa of Longiflorae) sebde ihnsection within embedded is savreyof variety a as , (subsection and Eucaprifolia Lonicera Calcaratae S S Nintooa ECTION ECTION .dioica L. and , and scaatrzdb lwr ihtwo-lipped with flowers by characterized is eg fteetosections. two these of merge ’s 1 spp.), (10 Nintooa scaatrzdb rnhswt oi pith, solid with branches by characterized is hspoie h is hlgntcsup- phylogenetic first the provides This . hnw ocdalfu pce of species four all forced we When . Isoxylosteum Breviflorae) Isika N hc suiu nhvn ognectar long a having in unique is which , .spinosa L. Caprifolium ontfr oohltcgop,in- groups, monophyletic form not do ) I SOXYLOSTEUM n ecnifrfo u re that trees our from infer can we and INTOOA of Isoxylosteum .myrtillus L. staiinlycircumscribed, traditionally as Isika, ifr rmohr nistiigor twining its in others from differs Cypheolae Rhe 93 aa18;Huand Hsu 1983; Hara 1903; (Rehder Eucaprifolia Spinosae l 0seisi hssection this in species 30 —All Lonicera h nyseiso subsection of species only the , .calcarata L. , .giraldii L. Nintooa omacaeta ssse to sister is that clade a form ahfr ld,support- clade, a form each orscin r gener- are sections —Four — and , u ifri aigboth having in differ but , norcmie tree, combined our In . hs ifri whether in differ These . Caprifolium hsscincnit of consists section This : ae ntecorolla the on —Based Microstylae 7sp) and spp.), (7 Microstylae Coeloxylosteum .tragophylla L. sdvddit two into divided is and , slw tde not does it low, is .carnosifolia L. .henryi L. : supporting , Phenianthi Vlm 33 [Volume (= Nintooa Thoracian- , Lonicera Cupulae Isoxylo- and Isika (sub- Nin- , : – (5 L. L. L. L. is 3 Delivered by Publishing Technology to: Yale University IP: 130.132.27.226 on: Tue, 25 Sep 2012 14:55:00 Copyright (c) American Society for Plant Taxonomists. All rights reserved. ugs htnihro h w uscin omdaclade, a formed subsections of placement two nrDNA the the to and owing of chloroplast neither both that from (2 suggest diploid Sequences mem- are 1966). their species Green between all hybrids though numerous bers, In yellow. are to yel- fading there or and white addition, red, with are tinged flowers rarely their white, and lowish glabrous, rarely yellow. to are to pink trast, fading are not flowers and their and white, of throughout, Members glabrous more differences. are morphological minor by aaofrsrn upr o h itrrltosi of relationship sister the ryi for support strong offer data r henryi era of relationship closer the nize otdcaeapast emr lsl eae osections to related closely more be to appears clade ported pN eecnodn.Hwvr oeacsin are result. and this accessions nrDNA evaluate more on critically to However, based needed trees concordant. since were unlikely topol- cpDNA is be the found even for we might explanation ogy an and as related, 1); Hybridization, closely (Fig. con-specific. are species each species of these recovered accession clearly we one Instead, containing tree. clades nuclear two or chloroplast acces- the that noteworthy either also is It of 1988). sions Wang and (Hsu tions within two The fruits. red and habit, upright a subsections, an have is to ovaries, tendency It evanescent the distinct and Africa. by corollas, two-lipped North characterized distinctly and pith, and group, north Europe, homogeneous Asia, Himalayas, rather Central the and of eastern west from species 14 about ebr n re 96.Tu,hy- Thus, 1966). Green and (Rüdenberg species diploid are test to needed are data of more monophyly Thus, Shimodaira- 0.5). the > nonparametric (P one-tailed test Hasegawa the by judged 2008] Fg ) o xml,subsection sections lin- example, other the of For with grade 1). allied a (Fig. closely form being section some with this eages, to of paraphyly assigned the groups support species other trees Our to 1903). affinity (Rehder clear cies no exhibited that groups he When species. tion section two to related or were one section corolla. only the with of these proposed shape of the half within in subsections Isika, 12 and recognized buds, (1903) outer Rehder the winter Accordingly, in habit, bracteoles, the and in bracts of diverse the most scales of also shape and is size group the This in the genus. throughout the distributed is of it range and species, 75 about with era, xmn h vltoayhsoyo h eto iha ex- an with section further sampling. the to taxon of desirable tensive history is evolutionary it incongru- the Therefore, phylogenetic examine species. the the of among cause ence the be may bridization rnhsapa ohv rsnmr hnoc in once than more arisen have to appear branches subgenus Rhe 93.Hwvr norpyoeei re section trees phylogenetic our in Nintooa subgenus fused However, not in are 1903). inflorescence found (Rehder the climbing subtending also and leaves are the branches though that hollow traits have habit, members its because L thsbe otltdthat postulated been has It L and ONICERA ONICERA Coeloxylosteum Tataricae .japonica L. per ohv rsnfo ihnsection within from arisen have to appears .xylosteum L. sahxpod(2 hexaploid a is edrpitdotta oeseisi this in species some that out pointed Rehder Isika, Lonicera S S Ochranthae ECTION ECTION uprigtemre ftetosubsec- two the of merger the supporting , oee,tencerrN aarecog- data rDNA nuclear the however, ; hl tl tesfre lal defined clearly formed others still while , hrfr,tiighbtadhollow and habit twining Therefore, . Nintooa I SIKA C n of and OELOXYLOSTEUM and hsi h ags eto in section largest the is —This n .morrowii L. Within . 4,wieteohrtospecies two other the while 54), = .koeheana L. Tataricae Nintooa .henryi L. Isoxylosteum Ochranthae Rhodanthae HI TA. HLGNTC FCAPRIFOLIEAE OF PHYLOGENETICS AL.: ET THEIS Nintooa hsscincontains section —This r itnuse only distinguished are , ik h w subgenera two the links (subsection ontfr ldsin clades form not do n with 8 edr1903; Rehder 18; = n test sec- to others and .giraldii L. u chloroplast our pce,i con- in species, sawl sup- well a as Lonicera Caprifolium, Ochranthae Lonicera Tataricae Isika . .hen- L. Lonic- Lonic- Isika– spe- of . ) hrce vlto,hbiiain n h biogeographic the into of and insights history hybridization, further many evolution, provide character surely iden- have will we which that phy- tified, clades detailed major more the for within stage analyses section the logenetic the set for these sections require results other of will with Our along status species. biogeography additional the their of of as inclusion appraisal well critical as A subsections Worlds. and an New Old indicate the the data between our disjunction Therefore, biogeographic Asia. interesting ( and Europe species to 14 stricted the of sis- subsection two is of which members America, to North ter western and northern to native ry,D .adC .Qin 00 h vlto fthe of evolution The 2000. Quinn. J. C. and M. D. Crayn, Dipsacales 2001. Donoghue. J. M. and Kim, T. S. Edwards, J. E. D., C. Bell, comparing Dipsacales: the Dating 2005. Donoghue. J. M. and D. C. Bell, S. C. Wojciechowski, F. M. Porter, M. J. Sanderson, J. M. G., B. new two Baldwin, Linnaeaceae, and Diervillaceae 1998. Pyck. N. and A. Backlund, for for Beijing Gar- Botanical at Quarryhill Sinica of for Chen Academia Higson den Howard Dr. of and thank Botany McNamara we Bill of material spinosa, Institute leaf the For of generosity advice. their Zhi-duan helpful for Boufford and David resources and with Normark, Ben Levin Rachel sen, w pce,rpeetdi u nlssby analysis our in represented species, two to Subsections Rehder porting speculation. the of this analysis test thorough and a to do to region necessary DNA is it but here, role genae placing hemsleyana in morphology L. with consistent is data chloroplast our akud .adM .Dngu.19.Mrhlg n hlgn of phylogeny and Morphology 1996. Donoghue. J. M. and A. Backlund, for classification ordinal An 1998. (APG). Group Phylogeny Angioperm ahw for Mathews rnhstriae ytoailr itrbd (Rehder however, buds tree, Alpigenae winter ITS the axillary have In and two 1903). hispid by setosely often terminated are branches subsections the of plants pce of species Nintooa eto rcetefrsarbs ld ihseiso sub- of species with clade robust section a result subsection. forms would Bracteatae supported and section well data another our by yet upheld in is 1913) 1909, of (Rehder merge proposed The clades. subsection. the of species other to related distantly Pileatae A te uscin ihnsection within subsections Other Alpigenae .gracilipes. L. CKNOWLEDGMENTS ei pcri h pcis(rcls n t ytmtcadevolu- and systematic its and (Ericales) epacrids the in spacer genic University. Uppsala Uppsala: Backlund. hlgn ae nclrpatDAsequences. DNA Botany chloroplast on based phylogeny implications. evolutionary Botany and genes, nuclear models, of region phy- angiosperm ITS on evidence The of source 1995. logeny. valuable a Donoghue. - J. DNA ribosomal M. and Campbell, caprifolioids. of families h re iscls p 1 Pp. Dipsacales. order the den plants. flowering of families the Alpigenae essetta eunehtrgniymgtpa a play might heterogeneity sequence that suspect We . .dfeiay,L giraldii L. deflexicalyx, L. 5 531–553. 85: , Fragrantissimae om ld ihteohrtrescin n is and sections three other the with clade a forms Isoxylosteum :481–499. 6: 284–296. 92: sgopdwt subsection with grouped is Lonicera naso h isuiBtnclGarden Botanical Missouri the of Annals Isika. .hispidula, L. ngnrlhbtadpubescence. and habit general in oehrwt te pce fsubsection of species other with together r rue nawl upre ld,sup- clade, supported well a in grouped are 10)osrain that observations (1903) ’s Similarly, etakDnPitr ra er,KrnHan- Karen Perry, Brian Pfister, Don thank We . . and , and involucrata, L. L hsi upre ymorphology; by supported is This . ITERATURE .alpigena L. Taxon yoeyo h Dipsacales, the of Phylogeny in –55 Coeloxylosteum and .gymnochlamydea L. Alpigenae, 7 657–661. 47: r ahrn Gould Katherine Dr. syringantha, L. naso h isuiBtnclGar- Botanical Missouri the of Annals Chlamydocarpi .hemsleyana L. C ITED and Isika .subspicata, L. Fragrantissimae .hemsleyana L. ahfr hi own their form each ersne eeby here represented Distegiae hni st other to is it than 2 247–277. 82: .hispida L. mrcnJunlof Journal American with avr aesin Papers Harvard .involucrata L. fsubsection of fsubsection of n r Murata Dr. and Distegiae inter- atp␤-rbcL Vesicariae ssimilar is Distegiae fsub- of )isre- while , d A. ed. Isika, Alpi- has 781 L. , Delivered by Publishing Technology to: Yale University IP: 130.132.27.226 on: Tue, 25 Sep 2012 14:55:00 Copyright (c) American Society for Plant Taxonomists. All rights reserved. oohe .J,T rcso,P .Ree,adR .Omta.2001. Olmstead. G. R. and Reeves, A. P. Ericksson, T. J., M. Donoghue, of evolution The 2003. Winkworth. C. R. and Bell, D. C. Noise J., 2000. M. Quicke. Donoghue, J. L. D. and Orme, L. D. C. Belshaw, R. 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Small. chloroplast noncoding L. 142–166. 21 92: R. analysis. of phylogenetic and utility for Relative Schilling, sequences II: DNA E. hare the E. and Winder, tortoise T. C. Siripun, U466 U637 U631 U645 U659 EU265583 EU265519, EU265455, EU265391, EU265327, EU240666, A,AF265277, (A), spacer, nnmu 1549-80G anonymous li n imn947-67-B Zinman and Elsik hn (A), China 88.101, Higson isofadWre 1078-81Mass Warren and Gilsdorf ee,cliae tAnl Arboretum, Arnold at cultivated Regel, Trz)Mkn,cliae tAnl Arboretum, Arnold at cultivated Makino, (Turcz.) ora fteAnl Arboretum Arnold the of Journal C.Y.Wu, sMtn) utvtda rodArboretum, Arnold at cultivated psbM-trnD), (trnS–trnG, Rehd. L., ..oe,cliae tAnl Arboretum, Arnold at cultivated G.N.Jones, sequences. .deflexicalyx L. .perfoliatum T. .formosa L. U467 U633 U647 U641 EU265535, EU265471, EU265407, EU265343, EU240677, .Tx n hi ore oce,adGnakaccession GenBank and voucher, source, their and Taxa 1. 0 408–424. 50: atpB–rbcL atn 1407 Cantino atpB Genetics iXY27 Li – ad-rw,Hj,adHfnr973-81A Heffner and Heja, Hardy-Brown, rbcL hn (A), China , . (A), oeua hlgntc n Evolution and Phylogenetics Molecular Wall, .occidentalis S. iySa,cliae tKwGardens, Kew at cultivated Shaw, Airy oieaalpigena Lonicera pcr and spacer, U638 U632 U646 EU265520, EU265456, EU265392, EU265328, pBtn) utvtda rodArboretum, Arnold at cultivated rpoB-trnC), , aai. utvtda uryilBtnclGar- Botanical Quarryhill at cultivated Batalin., U634 U638 U642 EU265526, EU265462, EU265398, EU265334, 0:735–749. 106: L., hn (A), China , 6 1114–1116. 16: U467 U630 U634 EU265458, EU265394, EU265330, EU240667, ed ISol) utvtda rodArbore- Arnold at cultivated only), (ITS Rehd. hn (A), China , U462 U638 U642 EU265466, EU265402, EU265338, EU240672, BO,AF265292, (BHO), ofode l 29341 al. et Boufford atn 1408 Cantino trnS–trnG nnmu 1104-89B anonymous isofadWre 431 Warren and Gilsdorf . . . U636 EU265390 EU265326, .aurantiacum T. . U644 U658 EU265582. EU265518, EU265454, .calcarata L. .chrysantha L. (A), .etrusca L. petN–psbM 7 442–452. 27: U468 U634 EU265408, EU265344, EU240678, ok utvtda rodArbore- Arnold at cultivated Hook, 7 222–247. 47: U463 U639 EU265403, EU265339, EU240673, U465 U631 EU265405, EU265341, EU240675, . utvtda rodArbore- Arnold at cultivated L., EU240665. psbM–trnD spacer, onemnadDikl 20447B Driskill and Youngerman BO,AF265291, (BHO), . . (A), egl japonica Weigela nnmu 843-84Mass anonymous .orbiculatus S. at,cliae tArnold at cultivated Santi, Hemsl., pcr odaeindicates Boldface spacer; nnmu 1010-86B anonymous mrcnJunlo Botany of Journal American E.P.Bicknell, uc. utvtda Ar- at cultivated Turcz., hn A,AF265276, (A), China , U468 EU265331, EU240668, U633 EU265397, EU265333, hn (A), China , .miconioides H. . i3304 Li okizaamabilis Kolkwitzia . . rotu angus- Triosteum . onemnand Youngerman .praecox W. onemnand Youngerman (A), Symphoricarpos onh culti- Moench, 6 176–189. 26: Vlm 33 [Volume usa(A), Russia , spacer, atr tal., et Walters hn. cul- Thunb., hn (A), China , trnL-F EU240669, EU265335, EU240670, e Tredici Del Systematic Molecular Lonicera. . . . Zabelia .mi- H. .car- L. Rehd. Tien , rpoB– .co- L. L.H- (A), and ’s . Delivered by Publishing Technology to: Yale University IP: 130.132.27.226 on: Tue, 25 Sep 2012 14:55:00 Copyright (c) American Society for Plant Taxonomists. All rights reserved. U469 U635 U649 U643 U657 EU265621. vosa EU265557, EU265493, EU265429, EU265365, EU240699, (PE), EU265622 EU265558, EU265494, EU265430, EU265620. (A), China U655 EU265619. EU265555, Arboretum, Arnold at cultivated Regel, (Rupr.) EU265606 EU265542, nochlamydea EU265478, EU265414, EU265350, EU240684, Arboretum, Arnold at EU265605. 28354 EU265541, EU265477, al. et Boufford os1323-84A mous EU265618. (A), Korea EU265553 178-83D EU265489, Thompson EU265425, and EU265361, Unger Richardson, Brown, EU265607 EU265543, 1776-80A Driskill U638 U642 U646 U650 EU265604 EU265540, EU265476, EU265412, EU265348, U659 EU265603 EU265539, EU265616 (A), China raldii pidula EU265608. EU265544, EU265480, EU265416, EU265352, EU240686, tum, EU265602. Arboretum, Arnold 18360-A at EU265601 cultivated EU265537, Franch, EU265473, EU265409, EU265345, Arboretum, 2008] U468 U634 U648 U642 U656 EU265610 EU265546, EU265482, EU265418, EU265354, EU240688, EU265609 EU265545, 1792-80Mass ryi U463 U639 U643 U647 U651 EU265615 EU265551, EU265487, EU265423, EU265359, EU240693, EU265613 . EU265549, EU265485, EU265421, EU265357, EU240691, EU265612 EU265548, 955-1A anonymous EU265611 (A), EU265547, USA California, 8523, EU265614. 485-82B Jones volucrata es. utvtda rodArboretum, Arnold at cultivated Hemsl., ai.(1), Maxim. nnmu 696-86C anonymous U471 U637 U641 U645 U659 EU265623 EU265559, EU265495, EU265431, EU265367, EU240701, ed,cliae tQarhl oaia Garden, Botanical Quarryhill at cultivated Rehd., .Ga,cliae tRnh at oaia Garden, Botanical Santo Rancho at cultivated Gray, A. ak,cliae tAnl roeu,Kopf, Arboretum, Arnold at cultivated Banks, (A), . .japonica L. .fragrantissima L. .morrowii L. .maackii L. .nervosa L. es.() utvtda rodArboretum, Arnold at cultivated (2), Hemsl. hn (A), China , U466 U632 U646 U640 EU265554, EU265490, EU265426, EU265362, EU240696, U468 U634 U648 U642 EU265556, EU265492, EU265428, EU265364, EU240698, U464 U630 U644 U648 EU265552, EU265488, EU265424, EU265360, EU240694, esmnadMceze345-94C Mackenzie and Weissman (A), U460 U636 U640 U644 EU265538, EU265474, EU265410, EU265346, EU240680, hn (A), China , ofode l 28660 al. et Boufford (A), U462 U638 U642 U646 EU265550, EU265486, EU265422, EU265358, EU240692, (A), . . . . hn. utvtda rodArboretum, Arnold at cultivated Thunb., . .inconspicua L. .gracilipes L. .myrtillus L. .iberica L. hn (A), China , .hemsleyana L. ur,cliae tAnl Arboretum, Arnold at cultivated Rupr., .hispida L. ry utvtda rodArboretum, Arnold at cultivated Gray, A U467 U633 U647 EU265491, EU265427, EU265363, EU240697, ai.(2), Maxim. onemnadDikl 1723-80E Driskill and Youngerman (A), U460 U636 U640 EU265484, EU265420, EU265356, EU240690, U467 U633 U647 EU265481, EU265417, EU265353, EU240687, id.&Px. utvtda rodArbore- Arnold at cultivated Paxt., & Lindl. U469 U635 U649 EU265483, EU265419, EU265355, EU240689, U465 U631 U645 EU265479, EU265415, EU265351, EU240685, U461 U637 U641 EU265475, EU265411, EU265347, EU240681, .ib,cliae tAnl Arboretum, Arnold at cultivated M.Bieb., .gynochlamydea L. okf Thoms., & Hook.f. Pall, Miq., Batal., Rehd., U463 U639 EU265413, EU265349, EU240683, ofode l 29295 al. et Boufford hn (A), China , ofode l 29096 al. et Boufford uaas.n Murata HI TA. HLGNTC FCAPRIFOLIEAE OF PHYLOGENETICS AL.: ET THEIS ofode l 28458 al. et Boufford . e rdc 77694 Tredici Del , .nigra L. li,EwnadHofstetter and Erwin Elsik, EU265617. onemnadDriskill and Youngerman pi (A), Spain , hn (A), China , nnmu 381-83Mass anonymous ,Jpn(A), Japan ., es.() cultivated (1), Hemsl. U470 EU265366, EU240700, ofode l 28875 al. et Boufford L., . .graebneri L. ad-rw and Hardy-Brown .Ce,990011 Chen, Z. onemnand Youngerman .maximowiczii L. . isn92.327, Higson hn (A), China , hn (A), China , .ferdinandi L. hn (A), China , ihe Wall Michael hn (A), China , hn (A), China , EU240679, EU240682, EU240695, i271 Li . . Hardy- . .hen- L. Rehd., anony- .ner- L. .his- L. .gy- L. .in- L. .gi- L. (A), . L. , , caria U472 U638 U642 U656 U650 EU265634 EU265570, EU265506, EU265442, EU265378, EU240712, U471 U637 U641 U655 U659 EU265633 EU265569, EU265505, EU265441, EU265377, EU240711, EU265632 EU265568, (A), Tajikistan 299-78B, Carey EU265631. (A), China U650 U654 EU265638 EU265574, EU265510, Arboretum, EU265637 EU265573, EU265509, 815-84A Driskill and erman EU265636. (A), tria EU265635 EU265571, losteum EU265507, EU265443, EU265379, EU240713, ica hn (A), China EU265630 861-82A, EU265566, EU265502, Driskill and man EU265629. (PE), 1999 utvtda rodArboretum, Arnold at (A), cultivated EU265644 China EU265580, voucher, EU265516, no EU265452, Garden, EU265388, Botanical Quarryhill at vated EU265643 EU265579, 1652-77E EU265515, EU265451, Lovejoy and Michener EU265642 EU265578, 694-88C Driskill koehneana (A), Arboretum, Arnold at EU265639 cultivated EU265575, EU265511, EU265447, EU265383, Arboretum, nold U650 U654 EU265628 796-74B EU265564, EU265500, Groves and Dumaine EU265627. U473 U639 U643 U657 U651 EU265645. EU265581, EU265517, EU265453, EU265389, EU240723, ruprechtiana (A), EU265626. (A), China 160 EU265625 EU265561, 346-94A EU265624. 18-92F pileata . utvtda mt College, Smith at cultivated L., U478 U634 U648 U652 U656 EU265640 EU265576, EU265512, EU265448, EU265384, EU240718, U479 U635 U649 U653 U657 EU265641 EU265577, EU265513, EU265449, EU265385, EU240719, (TEX), o. utvtda rodArboretum, Arnold at cultivated Kom., lvr utvtda rodArboretum, Arnold at cultivated Oliver, .() utvtda rodArboretum, Arnold at cultivated (1), L. (A), pi (A), Spain , ed 2,cliae tAnl Arboretum, Arnold at cultivated (2), Rehd. U474 U630 U644 U658 EU265572, EU265508, EU265444, EU265380, EU240714, .koehneana L. .sempervirens L. .standishii L. .tatarica L. .sovetkinae L. .pyrenaica L. U477 U633 U647 U651 EU265565, EU265501, EU265437, EU265373, EU240707, U475 U631 U645 U649 EU265563, EU265499, EU265435, EU265371, EU240705, nnmu 1002-86A anonymous U479 U635 U649 U653 EU265567, EU265503, EU265439, EU265375, EU240709, U474 U630 U644 U648 EU265562, EU265498, EU265434, EU265370, EU240704, ee,cliae tAnl Arboretum, Arnold at cultivated Regel, U472 U638 U642 U646 EU265560, EU265496, EU265432, EU265368, EU240702, hn (A), China 469-97A, Driskill and Youngerman (A), . . . . utvtda rodArboretum, Arnold at cultivated L., .rupicola L. .subsessilis L. .tragophylla L. ed 1,cliae tAnl Arboretum, Arnold at cultivated (1), Rehd. ok utvtda rodArboretum, Arnold at cultivated Hock, U473 U639 U643 EU265497, EU265433, EU265369, EU240703, . utvtda rodArboretum, Arnold at cultivated L., ktc. utvtda rodArboretum, Arnold at cultivated Tkatsch., U470 U636 U640 EU265514, EU265450, EU265386, EU240720, . utvtda nv fTxs Austin, Texas, of Univ. at cultivated L., hn (A), China , U470 U636 U640 EU265504, EU265440, EU265376, EU240710, . (A), . . . .tangutica L. okf Thoms., & Hook.f. .spinosa L. efe,HryBon n ea607-87A Heja and Hardy-Brown, Heffner, .nitida L. .modesta L. oe (A), Korea , (A), onemnadDikl 856-84A Driskill and Youngerman ed,cliae rodArboretum, Arnold cultivated Rehd., Hemsl., U476 U632 EU265436, EU265372, EU240706, U478 U634 EU265438, EU265374, EU240708, U476 U632 EU265446, EU265382, EU240716, U475 U631 EU265445, EU265381, EU240715, emn,UA(A), USA Vermont, 001, Theis ..Wlo,cliae tAr- at cultivated Wilson, E.H. . Jacques., Maxim., ed,cliae tArnold at cultivated Rehd., .syringantha L. iadRn4014 Ren and Li nnmu 838-76D anonymous nnmu 239-75D anonymous onemnadDriskill and Youngerman U471 EU265387, EU240721, ofode l 28126 al. et Boufford nnmu 1059-65A anonymous ie,June Tibet, Chen, Z. ofode l 28914 al. et Boufford . . .xylosteum L. .orientalis L. onemnand Youngerman li,Fc and Fich Elsik, ai,culti- Maxim, hn (A), China , anonymous EU240722, EU240717, Younger- . . . Young- .vesi- L. .dio- L. Aus- , .(2), L. Gould Lam., .xy- L. Elsik, 783 (A), (A), . . L. L. , ,