Delivered by Publishing Technology to: K. Marhold IP: 78.99.182.222 on: Sun, 11 Mar 2012 10:45:36 Copyright (c) American Society for Plant Taxonomists. All rights reserved. arcocrec ftomrhlgclysmlradoften 1979). and 1974, similar (Lack morphologically taxa, sym- two the are confused by of complicated however, be occurrence to Asia, seem patric they in as known, variation poorly distributionstill morphological exact and Its 1974, 2002). (Lack range McMullen Australia 1994; and Holzapfel Africa, 1979; to southern introduced been America, has North and Asia, highly widespread to is extending and It Europe, Europe. critical across in genus most this taxonomically of species the polymorphic of one certainly O 10.1600/036364412X616828 DOI © Botany Systematic ruso lsl eae pce,a ela oyopi spe- polymorphic variation. infraspecific as complex well with cies as Greuter species, related 2000; closely of Chaudhary groups 1994; (Holzapfel Holzapfel Minor 2006–2009). 1993; Asia and Lack Basin and Mediterranean the cen- in evolutionary Africa, ters putative with Eurasia, Zealand, New in and distributed Australia, species 40–50 approximately 2008; Waterway and Smith 2008; 2006; Scho al. Sahuquillo et Jørgensen and com- (e.g. Pimentel unravel histories species to and the help patterns and facilitate plex taxa related systems, closely mating of delimitation patterns, and distribution data, on requirements These knowledge ecological detailed data). the size with genome along or several karyological, morphometric, employing molecular, (e.g. simultaneously approaches techniques of and comparative sampling methods large-scale species- and the in to material due made traditionally mainly been systematics, are decades, have level taxa recent advances significant In these controversies. however, and taxonomic establish, by surrounded to concepts difficult taxonomic sound are data), morphological ambi- and species genetic the such within between of discrepancies observed variation, morphological continual Because patterns (e.g. variation research. the of taxonomic guity for challenge undoubtedly are a histories evolutionary complicated and tion oyih 02b h mrcnSceyo ln Taxonomists Plant of Society American the by 2012 Copyright oyopi ln pce ihlreitapcfcvaria- intraspecific large with species plant Polymorphic h genus The ¨ hi lca uvvladpsgailclnzto ots ogdsac ipras nhooei nrdcin n eetsra are spread recent and introductions endemic anthropogenic Balkan A to dispersals, close well. genetically as Long-distance be structure to routes. genetic appears their colonization defined, shaped postglacial have to and suggested survival glacial their rntrlhbtt thge liue.Ti nrseii ramn togycnrsswt h rdtoa aooi ocps hc recogniz which concepts, taxonomic traditional the with lectotypes. contrasts of strongly subspecies treatment 10 infraspecific to This up altitudes. higher at habitats natural or and altitudes, low at habitats man-made opoye,wihaecnretwt h w angntcgopnsrvae yAL akr.I scnlddta w subspecies two altitude’ that concluded ‘lower is the It and markers. altitude’ AFLP by ‘higher revealed the groupings genetic delimited, genetic and species, main are morphometrics this two multivariate morphotypes within the using recognized Two be with examined should data. congruent and are (AFLP) sampled which were polymorphism morphotypes, populations length 104 fragment Altogether, amplified Europe. in species controversial setre l 09 ad ta.21,2011). 2010, al. et Bardy 2009; al. et nswetter Keywords— Abstract— 2 eateto oay aut fSine hre nvriy Bena University, Charles Science, of Faculty Botany, of Department 1 irshieracioides Picris nttt fBtn,Soa cdm fSine,Du Sciences, of Academy Slovak Botany, of Institute 21) 71:p.258–278 pp. 37(1): (2012), Picris irshieracioides Picris Picris h opooia n eei aito ntePlmrhcSpecies Polymorphic the in Variation Genetic and Morphological The .japonica P. h rsn ae rvdsalresaetxnmcrvso of revision taxonomic large-scale a provides paper present The ae Slova Marek FP,Atrca,ifapcfcvrain utvraemrhmtis taxonomy. morphometrics, multivariate variation, infraspecific Asteraceae, AFLPs, nldssvrltxnmclycritical taxonomically several includes .(opste atca)comprises Lactuceae) (Compositae, L. .hieracioides P. hn.and Thunb. abr ag eei aito,adtolnae a ercgie ihnec useis otlkl reflecting likely most subspecies, each within recognized be can lineages two and variation, genetic large harbors 3 ´ uhrfrcrepnec:MrkSlova Marek correspondence: for Author k, 1,3 nErp.A dniiainkyadanmnltrlacutaepeetd nldn h einto of designation the including presented, are account nomenclatural a and key identification An Europe. in Cmoia,Lcuee nErp togyCnrsswith Contrasts Strongly Europe in Lactuceae) (Compositae, Jaromı .hieracioides P. rdtoa aooia Concepts Taxonomical Traditional .hieracioides P. .nuristanica P. .hieracioides P. irshieracioides Picris ´ Kuc r omnctn dtr hisnGemmill Chrissen Editor: Communicating ˇ subsp. era, . subsp. Bornm. 1 ao Marhold, Karol umbellata .is L. hieracioides ´ 258 bravska en otyasotlvdprnilta rw nmsc semi-natural mesic, in grows that perennial short-lived a mostly being , nmsc eintrlo aua aias(uha alherb tall as (such (Slova habitats altitudes higher natural at meadows) or semi-natural mesic, in predominantly occurring perennial, short-lived in to a latter often the habitats annual is whereas mountains), (HA) man-made usually in altitudes low often is at (or (LA) sunny, lowlands former dry, the occupies forms: biennial, life prefer- and ecological their ences in differ apparently morphotypes and two (Slova (LA) types altitude’ (HA) ‘lower altitude’ the ‘higher as named informally recog- and were Carpathians types nized morphological western two morphometric the Pannonia, recent adjacent from and the populations In of missing. investigation still is taxonomic however, advanced using tools, and area distinguish variation geographic species large to the a across of used evaluation are critical commonly indument A subspecies. involucral most these the characters size and the the involucre length, among the peduncle of The color subspe- 1). ten and Table of (see total recognized a with cies checklists, in in found and several Floras be but European can species, treatments single infraspecific a controversial as highly Presently, treated described. usually is been complex have the levels, intraspecific and described cies originally (1753) infraspecific Linnaeus hieracioides sound date. a hampering to factors concept main of the communities) been mesic have alpine low- to xerothermous, habitats Mediterranean land (from amplitude eco- and logical area distribution broad a plasticity), phenotypic large N mut a elc h aooi eeoeet and heterogeneity taxonomic the nuclear reflect in may variation amounts material intraspecific DNA was high the It the in patterns. that found geographic hypothesized some was Approximately also values suggesting size. 2C examined, genome in sig- variation the revealed 1.37-fold in populations heterogeneity 54 nificant of screening flow-cytometric 2 with diploid strictly is that showed Europe across sampled populations opiigana obena lnsocpigdy un,often sunny, dry, occupying plants biennial to annual comprising , xesv opooia aito poal including (probably variation morphological Extensive eetyudrae aylgclsre fnumerous of survey karyological undertaken recently A .hispidissima P. ´ ´ ([email protected]) k et ,S-4 3 rtsaa lvkRepublic. Slovak Bratislava, 23, SK-845 9, cesta irshieracioides Picris 1,2 ´ ic httm,anme ftx,bt ttespe- the at both taxa, of number a time, that since ; tska n uiaZozomova Judita and ´ lhuhmrhlgclyadeooial well ecologically and morphologically although , ,C-2 1Paa2 zc Republic. Czech 2, Praha 01 CZ-128 2, ihyplmrhcadtaxonomically and polymorphic highly a , n =10(Slova ´ ´ ´ n ahl 07.The 2007). Marhold and k -Lihova n ahl 2007). Marhold and k ´ ta.20) However, 2008). al. et k ´ 1 .hieracioides P. .hieracioides P. e Picris Delivered by Publishing Technology to: K. Marhold IP: 78.99.182.222 on: Sun, 11 Mar 2012 10:45:36 Copyright (c) American Society for Plant Taxonomists. All rights reserved. ad ta.2010). al. et Mered Bardy Lihova 2006; (e.g. al. taxa et typical diverged patterns, intraspecific recently complex for of cases in other efficient several proven has here employed molecular (AFLPs) with fingerprinting approach morphometric multivariate combina- a The of species. tion confused, often and of related, these delimitation from genetic and morphological the hieracioides from P. well-differentiated proven morphologically recently and endemic, Balkan ecologically be west to (a Koch J. D. its W. (Bartl.) of sampling restricted relatives, a close With relationships. and patterns (Slova selfing nuristanica P. aooi ramn of suggest treatment taxonomic to a (4) and groups; morphological relate observed the and patterns to environmental them variation genetic similar explore under to (3) cultivation conditions; after stability field- samples,retain field-collected to in for diagnostic (1) potentially variation differences, follows: morphological morphological whether investigate as to of (2) populations; are sampled patterns here the addressed examine objectives data. main morphometric and The (AFLP) polymorphism length ment hieracioides P. shown allogamous. was strictly it is and species rejected, the recently that was The 1935) variation. (Bergman intraspecific in play the formation seed shaping apomictic also reported previously in modes role significant Reproduction with investigations a approach. further multi-method for called a also study (Slova The species this 2009a). of history evolutionary complex the 1982 Pignatti 1979 Hayek SLOVA 1976 1975, Sell 2012] i parentheses). (in aulradMe 2000 Muer and Haeupler Dosta Kergue Bolo 1989 Geltman useisaeas niae:A–Ap,A pniePnnua P–Bla eisl,C eta uoe E–esenErp,I Iberian – IP Europe, eastern – western – EE WE Carpathians, Europe, Western Central – WC – species, CE the of Peninsula, area Balkan whole Europe. the – southern – BP WA of Europe, Peninsula, regions southern Apennine – mountain SE – – Pyrenees, SEm – AP P Europe, Alps, Basin, Mediterranean – – A M Peninsula, indicated: also are subspecies ohae 2002 Rothmaler S eciane l 2004 al. et Aeschimann rue 2006–2009 Greuter ˇ te ˇ hssuypeet ag-cl aooi eiinof revision taxonomic large-scale a presents study This Table pa ` (France) (Catalonia) (Italy) Europe) (Central (Europe) (Germany) fre Czechoslovakia) (former (Russia) (Germany) (Alps) Republic) (Czech (Europe) n io1995 Vigo and s ´ ´ e 2004 nek landC ´ e 1999 len .Ifapcfctxnmccnet of concepts taxonomic Infraspecific 1. oreIfapcfcTx eonzdWithin Recognized Taxa Infraspecific Source * ˇ rek 1992 ervenka ´ ;Slova nteohrhn,dslyacranlvlof level certain a display hand, other the on , ketal.2008). nErp yepoiggntcapiidfrag- amplified genetic employing by Europe in * .jpnc,P nuristanica, P. japonica, P. nyteErpa ato usai osdrdhr.I pcfe ntepriua eeec,aesrpre o h individual the for reported areas reference, particular the in specified If here. considered is Russia of part European the only ˇ ta.20;SihadWtra 2008; Waterway and Smith 2008; al. et a ´ ta.20b,w loamdt describe to aimed also we 2009b), al. et k .hieracioides P. subsp. WA subsp. subsp. A,WE subsp. AP BP, CE, subsp. subsp. subsp. subsp. E E WE EE, CE, subsp. SEm subsp. subsp. P P E WE CE, BP, AP, subsp. hieracioides hieracioides auriculata auriculata grandiflora auriculata hieracioides grandiflora grandiflora grandiflora hieracioides grandiflora elcigteeinferred these reflecting ´ ´ TA. AOOYO IRSHEAIIE 259 HIERACIOIDES PICRIS OF TAXONOMY AL.: ET K ta.20;Kuc 2003; al. et irsjaponica Picris and irshieracioides Picris subsp. IP subsp. subsp. WA subsp. WA subsp. subsp. WC subsp. WA subsp. WA subsp. WA subsp. .hispidissima P. .hieracioides P. .hieracioides P. spinulosa longifolia crepoides hieracioide hieracioides hieracioides crepoides hieracioides hieracioides hieracioides ´ ketal. ˇ and nrcn uoenltrtr.Rfrne r copne yteae covered area the by accompanied are References literature. European recent in era s subsp. P subsp. subsp. A subsp. IP subsp. subsp. WA subsp. E M SE, subsp. M subsp. IP subsp. ln oln odtos nacmo adneprmn tthe at experiment garden (48 common Bratislava Sciences, of a (resem- Academy Slovak conditions in Botany, similar of conditions) Institute under 1) cultivated lowland Table were Supplementary bling plants 1, the Appendix sown; populations, and (32 area of further geographic from seeds conditions mature environmental and analyses, by scoring influenced largely character characters detailed sampled. a plants fewer allow were exception preserve to on to per plants;only and 15–20 paper of capitulum typically consisted shape to sample population well-developed Each tape measurements. and translucent single size adhesive a by their bracts, from specimens attached involucral taken were inner the plant three ligules Before bracts, 1A). three involucral conserved Fig. and outer and three 1, plants) dried, field-collected total) Table were as in Supplementary to individuals 1, referred (1,861 (Appendix (hereafter specimens field of herbarium the as in comparison collected morphological were detailed A (Slova to elsewhere caution. relationships hispidissima with and P. variation taken their available of be plants analyses few hieracioides included to the also P. were due relatives, taxa Nevertheless, these two study. of first morphological were The the populations) analyses. two in genetic from the sampled in plants included Montenegro, and (Croatia P eehretd rsre shraimseiesadue for used and specimens herbarium as preserved harvested, were i ,SplmnayTbe1.I diin e niiul fthe of individuals few a addition, taxa, In (Appen- 1). related localities Table type closely respective Supplementary possible, also the 1, whenever from sampling dix and, collected wide The also subspecies, a habitats. were recognized comprised samples alpine currently they to all and Spain lowland included Romania, in from Mts. in range, Nevada Carpathians Sierra altitudinal southern the populations from the The area, in 1). to geographic distribution Table large its Supplementary of a 1, part spanned Appendix significant 1A: a covering (Fig. localities, Europe 104 at field the 17 . Material— Plant oeaieteetn ftepeoyi lsiiyadt eliminate to and plasticity phenotypic the of extent the examine To of samples population 101 analyses, morphometric For villarsii rielii paleacea paleacea longifolia villarsii hieracioide spinulosa spinulosa longifolia nuristanica 04 0 15 00 ;10m o he er 20–06.Foeigindividuals Flowering (2003–2006). years three for m) 180 E; s Krii,pat rw rmses,and seeds), from grown plants (Kirgizia, hudb eadda lutaieol,adne to need and only, illustrative as regarded be should with ´ ta.2009b). al. et k M subsp. subsp. A subsp. SE CE, subsp. WC subsp. E E WE EE, CE, subsp. SEm subsp. IP+France subsp. ln aeilof material Plant aeil n Methods and Materials .hieracioides P. irshieracioides Picris .japonica P. spinulosa spinulosa sonchoides spinulosa paleacea villarsii villarsii rielii .hieracioides P. Jpn lnsgonfo seeds), from grown plants (Japan, a enpromdadpublished and performed been has M subsp. WE CE, subsp. M subsp. E ,WE M, CE, subsp. irshieracioides Picris eecletdars wide a across collected were spinulosa villarsii spinulosa spinulosa a olce in collected was .hispidissima P. E P WE IP, CE, BP, AP, subsp. .hieracioides P. 10 villarsii 00 15 00 N, Delivered by Publishing Technology to: K. Marhold IP: 78.99.182.222 on: Sun, 11 Mar 2012 10:45:36 Copyright (c) American Society for Plant Taxonomists. All rights reserved. eoie ntehraimSV(omrne l 1990). al. et (Holmgren SAV herbarium the in deposited Europe. in species the useis(codn otetp oaiis–l . rrgosseiidi h rtlge)aeas indicated. also are – t protologues) according ME the proportionally Italy, in colored specified – reco bar, regions traditionally IT vertical or to Hungary, attributable a c., Populations – l. by 1. represented – HU Table localities Supplementary is Croatia, type and individual – the 1 Each HR to Appendix here. (according follow France, STRUCTUsubspecies abbreviations shown – population is the The FR common assignment. to Spain, cluster more (according the – the in individuals ES and structure at Germany, admixed Genetic outputs STRUCTURE clustering B. – genetically using Slovakia. different DE clustering of – Austria, Bayesian SK proportion – Slovenia, the the – AT with illustrating SI Andorra, Romania, colors, – – shown different AD is RO populations codes: Montenegro, of some Country in observed sectors admixture B). Genetic NE-LA). into part – blue results, circles dark SW-LA, the – blue dividing light NE-HA, by – red lineage, SW-HA – (yellow colors different aiesml fteosre opooia aito,eooia types, Slova ecological by variation, (reported morphological variants population; observed size per genome represen- the a individuals of be to seven sample chosen were tative to which 1), (two Table Supplementary populations 1, Appendix 61 from nating r for are to circles green and to morphotype’), correspond circles blue Dark/light ae lns ntesm anra h il-olce ones. culti- field-collected as the as to manner same referred the hereafter in plants) individuals; vated (306 analyses morphometric 37 [Volume BOTANY SYSTEMATIC 260 oce pcmn falfedcletdadcliae lnsare plants cultivated and field-collected all of specimens Voucher o FP,w ape 3 niiul of individuals 234 sampled we AFLPs, For Fig. .hieracioides P. .Sml ie n eei tutr of structure genetic and sites Sample 1. subsp. umbellata .hispidissima P. nlddi h opoercaaye ny w eei iegsrsle ihnec useisaeidctdby indicated are subspecies each within resolved lineages genetic Two only. analyses morphometric the in included .hieracioides P. ´ ta.20a n itiuinof distribution and 2009a) al. et k l nlddi h FPaaye.Epycrlsindicate circles Empty analyses. AFLP the in included all , irshieracioides Picris subsp. .hieracioides P. K and 2 = hieracioides K .Rpiaern at runs Replicate 5. = origi- and ie L opoye) e/elwcrlsto circles red/yellow morphotype’), ‘LA (i.e. .hispidissima P. cnae 80Lsanr n h hrceswr esrdwith measured were Microtek characters 2002). a the al. using et (Schols and scanned 2.0 CARNOY were scanner, paper 9800XL the ScanMaker and to ligules attached The 2). . bracts (Table 2 involucral specimen Fig. herbarium in illustrated sampling. each are characters field on morphological Selected scored six the or and semi-quantitative, during measured in two were useful quantitative, binary) taxa as (17 characters inferred infraspecific 25 were Altogether, of that delimitation characters and the for hieracioides P. used traditionally acters irshispidissima Picris opoercAnalyses— Morphometric .Ampdpcigsml ie fteaaye populations. analyzed the of sites sample depicting map A A. . K aesal eut,weestoeat those whereas results, stable gave 2 = nteltrtr,mil hs eotdb ozpe (1994), Holzapfel by reported those mainly literature, the in and .hieracioides P. h opooia hrcesicue char- included characters morphological The .hieracioides P. .hieracioides P. ae nAL akr sinferred as markers AFLP on based subsp. hieracioides subsp. K nerdtwo inferred 5 = umbellata rycircles gray , gnized (‘HA RE o Delivered by Publishing Technology to: K. Marhold IP: 78.99.182.222 on: Sun, 11 Mar 2012 10:45:36 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 0–asn,1–peet seFg 2C). Fig. (see present) – 1 absent, – (0 2B), Fig. (see present) – 1 analyses. (3) (2) (1) peduncle, bracts per branch, involucral capitula stem outer 2A), the per Fig. (see present), capitula of stem branch) – the whole number stem the of of 1 1/2–1/3 along number absent, upper – Maximum in (0 – only branches – (0 stem 1 on peduncle branch, capitula and the of involucre Distribution the on hairs SLOVA 2012] rcso n admycoe aiuu r given; are capitulum chosen randomly one on bracts slse eo.Almrhmti aamtie sdi hssuyare study this in request. used upon matrices author first data the morphometric from available All below. listed as 1976; (Thorpe respect this shown in 1980). deviations been Klecka require against have robust but generally considerably characters, be the space, to analyses of separation three-dimensional distribution Discriminant best normal or the multivariate groups. achieve two- to predefined a largely characters between in all is among weights it variation differentially variation displays overall CDA the the and of individuals when most hierar- the extracts groupings a PCA impose artificial Whereas not do continuous. or analyses, structure cluster few to a chical only contrast in in information and, of amounts dimensions by are large techniques summarize (represented ordination that These populations tools objects. efficient with as character) and each for- of plants discriminant values the mean individual performed, canonical with were and 1990) both Krzanowski 1990) mer 1980; Krzanowski Klecka (CDA; 1973; analysis Sokal (PCA; and computations, analysis the component Sneath Principal distort analyses. may discriminant that char- the correlated 1998) especially highly Legendre of and pairs (Legendre reveal acters to computed first were coefficients ligule fteotrivlca bracts involucral outer the of (cm), ar.I H n H eercre s1(..asneo ar) the hairs), of 0. absence of value (i.e. the NBr given 1 were as color hair recorded the recording were IH DHI PH, DH, and characters DHP If hairs. egho h ligules the of Length admycoe capitula; chosen randomly nour n euce( bet present), – 1 absent, – (0 peduncle and involucre bracts bracts rpl ooe hairs; colored pale or prehis es ar)(e i.2E), Fig. (see hairs) dense – dense 3 – 3 hairs, hairs, sparse sparse – – 2 2 hairs, 2D), no Fig. – (see (1 hairs) peduncle on hairs of Distribution (cm), it fteligules the of Width – nour n euce( bet present), – 1 absent, – (0 peduncle and involucre Table eea aamtie eeasmld n nlsswr performed were analyses and assembled, were matrices data Several correlation (non-parametric) Spearman and (parametric) Pearson ubro tmbranches, stem of Number – tnadsaitclprmtr,sc stemda,te1t,25th, 10th, the median, the as such parameters, (displaying statistical box-plots standard and CDA carefully), interpreted at cultivated be or consideration, should environmental taxonomic least and into by taken influenced be field-collected not most should 1, the (that characters variables identify CDA of to phenotypes and by plants matrix. the tested data ‘HA compare same and was the To two on ‘LA’ Results) based the the was to see which between (attributable respectively; 2 Differentiation PCA morphotype,’ analysis by eliminated. plasticity suggested The was potential groupings the performed. characters where cultivated was conditions, plants of the ecological characters, among similar pattern 25 under variation all overall an correla- of displayed the and matrix (306 experiment plants garden tion cultivated their common all the on of from based individuals) 2, heterogeneity analysis component environmental Principal sam- may by that material characters habitats. the influenced of natural inclusion in the been of patterns first risk have the the variation obtain of to overall irrespective was pled, the aim The into performed. matrix was correlation insights (101 characters, the means 25 and all population plants of field-collected on all based of 1, populations) analysis component Principal 1 NL 1 2 NBP , (mm), (mm), LoB ubro tmleaves, stem of Number – .Ls fmrhlgclcaatr sdi morphometric in used characters morphological of List 2. 1 envle ftremaueet flglso involucral or ligules of measurements three of values Mean egho h ue nourlbracts involucral outer the of Length – aiu ubro rcsprpeduncle, per bracts of number Maximum – RSL WiB e ogtdnlsrpo h ue iue 0–absent, – (0 ligules outer the on strip longitudinal Red – DHI it fteinrivlca bracts involucral inner the of Width – 1 (mm), 1 c 2 (mm), re-rw,lgtbon i rmulti-colored or bi- brown, light green-brown, , itiuino ar nivlce( ohairs, no – (1 involucre on hairs of Distribution – BML NiB a LCT rw-lc,rdbako lc hairs; black or red-black brown-black, 1 (mm), rw ak nteuprpr fligules of part upper the on marks Brown – LTL vrg ubro ne involucral inner of number Average – LBr egho h ool tubes corolla the of Length – egho h ogs et nthe on teeth longest the of Length – LP LiB egho h ogs tmbranch stem longest the of Length – egho h ogs peduncle longest the of Length – NoB egho h ne involucral inner the of Length – vrg ubro the of number Average – ´ DH IH TA. AOOYO IRSHEAIIE 261 HIERACIOIDES PICRIS OF TAXONOMY AL.: ET K PH 1 Color-intermediate – Dark – (mm), 2 Pale – one rmthree from counted NCP a WoB 1 b Maximum – ar nthe on hairs ar nthe on hairs (mm), 1 (mm), Width – DHP NCBr b DC white LL WL – – – – c (5) (4) (6) t enRt emn) U 2 2 Massachusetts), Germany), Leon-Rot, St. Frets,1.5 (Fermentas), T.The CTC. pro- MseI reliable chosen: most the were produced repeatedly files that primers selective of pairs rmr(ple isses,0.04 U/ Biosystems), (5 (Applied primer xeso t72 at extension 19)adtepooo rvddb ple isses(Applied Biosystems Applied using Genomic double-digested by below. described was al. as provided DNA et modifications, some Vos protocol with by 2005) the Biosystems described procedure and general Germany). (1995) the Hilden, followed (Qiagen, kit analyses mini plant AFLP DNeasy the using leaves dried MgCl diinl1 r t16 at hrs 12 added additional 5 was of mix ligation volume a a digestion, in the After ng). (500–1,000 extract DNA r.Terato i f10 of mix reaction The hrs. 3 N rget eedltd11 ihT 1 MTi,01m EDTA) mM 0.1 1 10 Tris, containing in mM umes performed (10 were TE amplifications with Preselective Ligated 1:10 buffer. California). diluted City, were Foster fragments Biosystems, DNA (Applied pair adaptor each hss nttl 7pie ar eesree sn ih individuals polymor- eight using of screened level were appropriate pairs primer an of 27 of total, In profiles phisms. AFLP well-scorable and diluted were amplification buffer. preselective TE the with 1:15 of approximately Products Eppendorf). S, rdc.TePRccepoiews72 was profile cycle PCR The product. 0sc 56 sec, 30 FPFingerprinting— AFLP .hieracioides P. esace o pia eetv rmrpista eutdi clear in resulted that pairs primer selective optimal for searched We -CTC, olce n utvtdpat ie agl nlecdb environ- by influenced largely Results). (i.e. see ment; plants field- cultivated the between three and separation identified the we collected to result, most a dataset contributed As this that 3. characters in CDA invariable from was excluded 2) therefore (Table charac- and DH 24 character on binary based the was ters; plants analysis cultivated predefined, This LA respectively. groups individuals), and on (85 two individuals) based (208 with ‘HA plants was morphotype,’ field-collected the ‘LA LA 3 and analysis the of individuals), discriminant of individuals (178 Canonical individuals characters. plants on cultivated 25 based HA all and was individuals) plants field-collected 2 (396 HA predefined, groups CDA two with Thus, morphotype,’ 1). Supplementary box-plots, 1, (Appendix Table and material available statistics the descriptive of all analyzed the we for field-collected material, we as cultivated both CDA available and in were that Whereas populations and separately. morphotype’ only (‘HA included analyzed 2 were and computed. morphotype’) 1 were steps ‘LA values) in revealed extreme groupings the two and The percentiles 90th and 75th, fte‘Amrhtp, ohbsdo h orlto arxof matrix correlation the on populations) based (48 both means characters. morphotype,’ population 22 ‘LA the on the morphotype,’ based of ‘HA was the 5 of PCA based populations) was while (53 4 means PCA population field- morphotype: the on each on performed for were separately plants, and PCA collected (‘HA’ additional found two dif- types potential morphotypes’), morphological and ‘LA main patterns the variation the within two into ferentiation (the insights predefined deeper groups get two To 3). and PCA by characters, suggested 22 morphotypes of set populations), 101 same from was the individuals 4 (1,861 analysis plants discriminant characters individual Canonical on 22 2). based of Table of DH, matrix populations) WiB, correlation (101 LBr, the means (excl. and population plants on based field-collected but plas- was all CDA 3 environmentally as PCA and identified i.e. PCA characters tic, of to exclusion subjected the again with were plants Field-collected u oterivrac na es n ru.Bxpo graphs Box-plot group. 2000). Institute one (SAS package 8.2 least ana- SAS the All using drawn. at performed also were lyses were in characters selected invariance in variations their depicting character-state-frequency) to the excluded of due were means characters by separately binary analyzed characters); (and 17 (i.e. characters tive japonica relatives: P. close its differ- morphological and the explore to between performed entiation was 5 CDA Finally, 2 m 2 M,0.2 mM), (25 ,ApidBoytm)ad2 and Biosystems) Applied L, o 0scad72 and sec 30 for C EcoRI EcoRI 1 utvtdpat) h nlssicue nyquantita- only included analysis The plants). cultivated (14 o 0mnadcoigt 4 to cooling and min 10 for C rgntn rmfu ifrn egahcrgos Four regions. geographic different four from originating , m m rmr eefursetylblda h 5 the at labeled fluorescently were primers -ACC-(NED)/ 4DAlgs ufr(nldn T)ad1 and ATP) (including buffer ligase DNA T4 L e ape n h ecin eeicbtdfran for incubated were reactions the and sample, per L m m EcoRI L10 m L10 NP 1 Mec) 0.5 each), mM (10 dNTPs L .Telgto i otie 4DAligase DNA T4 U 1 contained mix ligation The C. .hieracioides P. eoi N a xrce rmslc gel- silica from extracted was DNA Genomic -ATC-(6-FAM)/ + + m .nuristanica P. ag ufr(emna)ad5 and (Fermentas) buffer Tango C ufrI ApidBoytm,0.6 Biosystems, (Applied II Buffer PCR o i,wihwr olwdb final a by followed were which min, 2 for C ouecnand5U 5 contained volume L MseI MseI EcoRI CTand -CAT rmErp 36cliae plants) cultivated (306 Europe from NwEgadBoas Ipswich, BioLabs, England (New m m o i,3 ylso 94 of cycles 30 min, 2 for C mlTqDApolymerase DNA AmpliTaq L fdltdrestriction-ligation diluted of L and MseI egtcliae lns and plants) cultivated (eight Msecce pGradient ep (Mastercycler C MseI -CAG, EcoRI m fec preselective each of L nye t37 at enzymes -AGG-(PET)/ EcoRI EcoRI m ecinvol- reaction L -ACG-(VIC)/ (Fermentas, 0 m n.The end. Lofthe m Cfor MseI Cfor Lof m L - Delivered by Publishing Technology to: K. Marhold IP: 78.99.182.222 on: Sun, 11 Mar 2012 10:45:36 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 6 YTMTCBTN Vlm 37 [Volume BOTANY SYSTEMATIC 262 1–n ar,2–sas ar,3–dnehis.E itiuino ar nivlce–DI( ohis prehis es hairs). dense – 3 hairs, sparse – DHP – 2 peduncle hairs, on RSL no hairs – – of ligules (1 Distribution outer DHI D. the – present). on involucre – on strip 1 hairs longitudinal absent, of Red – Distribution B. (0 E. branch). BML hairs). stem – ligules dense of of – 1/2–1/3 part 3 upper upper hairs, in the sparse only on – – marks 2 1 Brown hairs, branch, C. no stem present). – entire – (1 the 1 along absent, – – (0 (0 DC – branches Fig. .Slce opooia hrcesue nmliait opoercaaye of analyses morphometric multivariate in used characters morphological Selected 2. irshieracioides Picris .Dsrbto ftecptl nstem on capitula the of Distribution A. . Delivered by Publishing Technology to: K. Marhold IP: 78.99.182.222 on: Sun, 11 Mar 2012 10:45:36 Copyright (c) American Society for Plant Taxonomists. All rights reserved. inladrgoa ees nSRCUEcmuain,bsdo a on allele based recessive a computations, used popula- we STRUCTURE algorithm, individual, (MCMC) In Carlo the Monte levels. chain at Markov regional patterns genetic and the tional into insights additional riainae xlie 35%ad1.2 ftettlvrain .Cnncldsrmnn nlss(D )wt w r-eie rus ht – white groups: pre-defined two with 1) (CDA analysis discriminant Canonical B. variation. total the of 13.72% hieracioides and P. 23.50% explained axes ordination mlmne nSRCUE22(auhe l 07 a mlydfor method employed was clustering 2007) only al. including Bayesian matrix et data (Falush a the 2.2 analyses STRUCTURE Furthermore, PAUP in bootstrap in 2001). implemented calculated by matrix (Swofford distance using assessed genetic 2006), 4.0b10 Bryant (1979) was and Li’s was (Huson and diagram 4 tree Nei SPLITSTREE NeighborNet program the The the replicates. in in produced 5,000 with node 1985) distances each genetic (Felsenstein (1979) of Li’s and port Nei PAUP on using based constructed tree was NJ The 2006). Harris nlss(C ) qae – squares 2): (PCA analysis eei iiaiis a efre nFM .0 ea(Schlu beta 1.108 FAMD pairwise in calculating performed and for was coefficient similarities, including Jaccard’s genetic both ( using network-generating 2006), taxa analysis, a Asian coordinate neighbor-joining Bryant using two a and by the or excluding (Huson on 1987) method based Nei and NeighborNet analyses Saitou cluster (NJ; (PCoA; algorithm and analysis coordinate 1990) principal by Krzanowski explored were populations data and AFLP request. upon complete author were The first the 2004). analyzed) from analyzed al. available then et is types were matrix (Bonin which morphological independently samples, scored replicated and and populations as different taxa used from and all re-extracted originating dataset, representing indi- final 22 data, and the AFLP present the of as of coded reproducibility (9% the and viduals estimate recorded To were (0). bp absent unambiguous or 50–500 (1) range and size well-scorable the in Only markers scored. and http://hordeum.msu.montana. at edu/genographer/) (available imported 1.6.0 subsequently GENOGRAPHER Biosystems), into (Applied software 3.7 GENESCAN 02 SLOVA (10 amplifications 0.2 mM), (25 selective of 1 contained mix reaction 2012] est,Bailv) ieclbainwsahee sn h nenlsize internal the LIZ using –500 achieved GeneScan was standard, calibration Uni- Comenius Size Biology, Bratislava). sequencer Molecular versity, DNA of Avant fluorescent Department 3100 different ABI Consortium, an four (BITCET using analyzed with and pooled labeled were primers dyes with obtained products 1 C yl rfl a 95 was profile cycle PCR 2 and Biosystems), Applied 55.9 f94 of 0mnadcoigt 4 to cooling and min 10 m FPDt Analyses— Data AFLP h vrl eei tutr n eainhp mn h tde taxa studied the among relationships and structure genetic overall The Fig. M, ec yl eraigb 0.7 by decreasing cycle (each C o 0sc 56 sec, 30 for C MseI .Mrhmti nlsso utvtdpat of plants cultivated of analyses Morphometric 3. t5 at m L10 subsp. m NP 1 Mec) 0.5 each), mM (10 dNTPs L m ) 0.08 M), + hieracioides C odbfe ApidBoytm) 1 Biosystems), (Applied buffer Gold PCR o mn 72 1min, for C Msecce pGain ,Epnof.The Eppendorf). S, Gradient ep (Mastercycler C m o 0mn 3cce f94 of cycles 13 min, 10 for C mlTqGl N oyeae( U/ (5 polymerase DNA Gold AmpliTaq L m a FPdt eeaaye using analyzed were data AFLP Raw .hieracioides P. # * fdltdpeeetv C rdc.The product. PCR preselective diluted of L .hieracioides P. eso .b0(wfod20) h sup- The 2001). (Swofford 4.0b10 version ‘Amrhtp’,bak– black morphotype’), (‘LA ApidBiosystems). (Applied .japonica P. )fr1mn 72 min, 1 for C) o i,floe y72 by followed min, 1 for C subsp. and m and fec rmr( primer each of L ´ .nuristanica P. TA. AOOYO IRSHEAIIE 263 HIERACIOIDES PICRIS OF TAXONOMY AL.: ET K .hispidissima P. hieracioides m ecinvolume) reaction L o i,2 cycles 23 min, 1 for C o 0sc 65– sec, 30 for C irshieracioides Picris .hieracioides P. ‘Amrhtp’,crls– circles morphotype’), (‘LA .Principal ). m * oobtain to ¨ MgCl L EcoRI e and ter version Cfor m L, at 2 subsp. 36pat) ae n2 opooia hrces .Picplcomponent Principal A. characters. morphological 25 on based plants), (306 eutn lseigpten.W sdrn f10 of runs used We patterns. clustering resulting into frequencies each assigned allele For were of Individuals independence K populations. and admixture among assuming model, 1 rnia opnn analysis measured component Principal all thus, (1) pair; analyses. character further in any retained were for characters 0.75 exceed not ariedfeecsbtenidvdas(e’ eedvriy D diversity, gene (Nei’s of individuals proportion average between the differences and the (P%) pairwise calculating markers by polymorphic diversity of genetic percentage the estimated we of analyses, clustering significance The permutations). (10,000 2005). population tests permutation al. among using et tested as was Excoffier well differentiation 3.11; as populations, (ARLEQUIN genetic among total groupings of and partitioning within reveal to variance computed were distances, (Jakobsson pairwise software. 1.1.1 2004) (Rosenberg ver. DISTRUCT CLUMPP and calcu- 2006). 2007) using Rosenberg Ehrich were generated AFLPdat; and parameters was of statistics (part output these Structure-sum-2009 R-script Graphical All details). the more in for lated 2005 replicate to al. respect the et with over change data 3) of the and rate value, of K probability each posterior for runs log estimated the mean of 2) runs, replicate the between coefficients similarity clus- of number ( optimal ters the identify The to (http://www.bioportal.uio.no/). computed were Oslo statistics of following University the of Bioportal FPa a W nErc 2006). Ehrich in DW1 (as AFLPdat Scho (DW; values marker uni eido 10 of period burn-in a ouaingopn n rsn nalisidvdas cluae in (calculated individuals) its a all or in population Schlu certain present FAMD; a and individ- to its confined grouping of (those all population markers in fixed present a necessarily private at not uals), and present grouping (those population markers a was or rare divergence of genetic number The frequency the 2006). calculating (Ehrich by AFLPdat assessed R-script the using vle hc saue-eie ubro lses a e rm1–10. from set was clusters, of number user-defined a is which -value, umbellata opoercAnalyses— Morphometric o ahpplto rpplto ruigta eutdfo the from resulted that grouping population or population each For Euclidean on based (AMOVA), variance molecular of Analyses ouain of populations K htbs elc h eei tutr rsn ntedtst 1) dataset: the in present structure genetic the reflect best that ) K < 0rpiaern eepromdt ettesaiiyo the of stability the test to performed were runs replicate 10 , ‘Amorphotype’). (‘HA .hieracioides P. ¨ 0) rvt akr toecnie oacranpopulation certain a to confined (those markers private 10%), e n ars20)adb h frequency-down-weighted the by and 2006) Harris and ter .hieracioides P. 5 ¨ trtos h nlsswscridota the at out carried was analysis The iterations. setradTish20) sipeetdin implemented as 2005), Tribsch and nswetter subsp. △ K Results K umbellata uniybsdo h eodorder second the on based quantity a – ftelklho ucin(e Evanno (see function likelihood the of h orlto ofiinsdid coefficients correlation The n l 5eaie characters, examined 25 all and ,bsdo 0 field-collected 101 on based 1, ‘Amrhtp’.Tetwo The morphotype’). (‘HA 6 CCieain after iterations MCMC L K ( K )–themeans lses the clusters; Nei ) Delivered by Publishing Technology to: K. Marhold IP: 78.99.182.222 on: Sun, 11 Mar 2012 10:45:36 Copyright (c) American Society for Plant Taxonomists. All rights reserved. RSL LTL WL LCT 2 ohPA2adCA1 ae ntesto 0 culti- 306 of set the on based 1, CDA and 2 PCA Both (2) ihtecnnclae ai ,ai ) o eal ntedfeetPAadCA e aeil n ehd.Frcaatrepaain e al .Th bold. 2. in Table see are explanation, text, character character the For morphological Methods. in and the to Materials of see referred correlations CDA, expressing structure, and structure canonical PCA canonical different total total the and – on 5 eigenvectors details CDA For of 4, 2). values axis CDA 1, higher 1, (axis CDA axes 2); canonical axis the 1, with (axis components principal the with H0.358 PH DH 0.325 DHI DHP NBP 0.295 LP NL LBr NBr 37 [Volume BOTANY SYSTEMATIC 264 DC IH o 0.258 NoB NCP NCBr LL WiB LiB WoB LoB NiB BML Char. Table hrceie ycptl htaetpclydistributed typically morphotype’ are also that ‘LA is capitula morphotype’ the by ‘HA characterized The with 2). Table compared (Supplementary stem when fewer longer but have bracts, ligules involucral apparently outer leaves, morphotype’ stem branches, ‘HA the 1 CDA of Populations 3). by (Table suggested character discriminating was another (RSL) as ligules outer the longitudinal on red a strip of presence the addition, morphotype In the distinction. to the most contributing both thus showed of CDA, and axes PCA (NoB)) discriminating the bracts and with correlations (LL), involucral highest ligules outer of (NBr), of length branches (NL), number stem leaves stem of of number quanti- number four (the and characters (DC)) branches tative stem distribu- on the capitula and of IH) tion (PH, peduncle hairs and color-intermediate involucre and the same char- on pale binary the of Three presence to 1. (the PCA corresponded acters in which found as 3), along morphotypes (Fig. two formed axis groupings first two the showed and characters, experiment 25 garden all common natural the 32 in from collected populations) seeds from (obtained plants vated (HA)). morphotype’ referred altitude ‘higher (further populations the short-lived as by to mostly being higher formed at and habitats was natural altitudes to semi-natural other mesic, in found the alti- while ‘lower Slova the following (2007)), as (LA) to morphotype’ referred tude (further plants biennial annual mainly man-madeto comprising and and altitudes sunny, lower variation; at dry, habitats total popu- in included typically the groupings growing the of lations of One 20.34% shown). (extracting not figure axis first distinct the two influ- in environmentally resulted potentially enced, are that those including .Tenmrclotu fmrhmti nlss C ,PA3 C ievcosepesn orltoso h xmndcharacters examined the of correlations expressing eigenvectors – 4 PCA 3, PCA 2, PCA analyses: morphometric of output numerical The 3. 025012023025014005–.4 .3 0270.418 0.439 –0.211 –0.297 –0.021 0.022 0.338 0.243 0.234 –0.349 –0.124 0.067 0.065 –0.001 0.151 0.144 0.364 –0.199 0.245 0.318 0.011 0.213 0.170 0.153 0.172 0.305 0.308 –0.215 –0.131 –0.033 012003----–.2 .7 - - 0.571 0.278 –0.228 –0.341 - 0.453 - –0.355 0.245 - –0.144 - –0.064 0.244 0.063 0.070 –0.112 0.090 –0.148 –0.066 019028----–.6 .2 0130.152 –0.153 0.223 0.219 –0.166 0.033 0.191 - –0.020 - 0.272 - 0.181 - 0.429 0.147 0.278 0.295 0.346 –0.139 –0.273 –0.101 007007000024029–.5 001013-- - 0.183 –0.021 –0.054 0.289 0.214 0.090 0.077 –0.007 xs1ai xs1ai xs1ai xs1ai xs1ai 2 axis 1 axis 1 axis 1 axis 2 axis 1 axis 2 axis 1 axis 2 axis 1 axis .0 .0 023025034–0.043 0.324 0.285 –0.233 0.206 0.204 .2 .3 .2 .2 .6 0.037 - - 0.265 - - 0.026 –0.029 –0.229 0.220 0.436 0.280 - –0.106 –0.080 0.342 - 0.417 0.331 0.414 - 0.010 –0.136 0.263 0.320 - 0.150 0.057 0.138 0.125 0.324 0.271 .4 .2 .6 .6 .9 .8 .2 .7 .2 –0.176 –0.135 0.221 0.363 0.072 –0.160 0.229 0.343 0.385 0.427 0.095 –0.042 –0.016 0.060 –0.086 0.064 0.126 0.127 0.167 0.141 .6 .1 .1 .3 .3 .6 .4 0090400.100 –0.070 0.420 0.019 –0.079 –0.106 0.348 0.032 0.261 0.210 0.030 0.215 0.339 0.175 0.019 –0.110 0.318 0.303 0.115 0.061 0.019 C 2 PCA i.3A Fig. –0.117 009027–.1 .3 –0.044 0.239 –0.016 0.207 –0.079 0.129 0.005 0.069 .1 .4 .0 0090.178 –0.049 0.104 0.246 0.215 0.096 ruig,sprtdalong separated groupings, 0.270 0.329 0.281 0.285 0.335 0.296 0.269 C 3 PCA i.4A Fig. ´ n Marhold and k .0 .0 .8 –0.152 0.285 0.109 0.003 .3 .7 –0.020 0.272 0.130 0.212 0.316 –0.144 0.209 0.070 0.161 .3 0090.101 –0.049 0.034 .6 .6 .5 –0.472 0.157 0.062 0.161 .8 .2 0.280 0.029 0.080 C 4 PCA i.5 Fig. 3 aoia iciiataayi ,promdo the on performed 2, analysis discriminant Canonical (3) ausosre o h utvtdadfield-collected and the cultivated apparent. in the were shifts plants for substantial also observed CDA; S2) of values Fig. results the Supplementary 3). confirmed 2; cultivated LBr and Table Table the WiB characters (Supplementary (Supplementary of the of 12% morphotype analyses data in this Exploratory only of found plants were morphotype,’ hairs ‘HA the such of as plants many field-collected as the in of observed 43% were hairs LBr. dark and were whereas DH, Indeed, characters WiB, the plastic: three environmentally of Thus, as length identified (LBr). the branch and stem (WiB), of longest bracts width involucral the inner and revealed: the cultivated were the individuals between shift field-collected the to the charac- contributing of two ters S1B), individuals Fig. the (Supplementary on In morphotype’ performed ‘LA S1A). was Fig. which invo- 3, (Supplementary the CDA (DH) on hairs peduncle dark and of lucre involucral presence the inner and of (WiB) width cultivated bracts the the most plants: between that field-collected shift and characters the to two rele- contributed identified strongly taxonomically analysis considered The be vant. not variables, should environmental thus by characters predefined and influenced of largely two identification are the as that on focused plants was cul- of field-collected groups, sets the and and morphotype’ tivated ‘HA the of individuals ‘LA the in one frequent 3). (vs. Table (Supplementary morphotype’) bracts involucral outer on strips the red of mor- occurrence scarce a the ‘LA and in morphotype’), hairs ‘LA pale mainly the (vs. peduncle and involucre in the on being hairs branch color-intermediate mainly (vs. of whole branches presence the photype’), stem the of along 1/2–1/3 upper distributed the in only D 1 CDA 0.559 0.923 0.673 0.580 0.658 .4 0.857 0.842 0.661 0.594 i.3B Fig. D 4 CDA 047-- - –0.457 0.506 .8 0520.414 –0.542 0.484 0.795 0.608 0.563 .4 - - 0.343 0.612 .2 0.788 0.620 i.4B Fig. 0100.259 –0.110 057–0.024 –0.537 .7 –0.432 –0.258 0.670 0.606 -- -- D 5 CDA i.6 Fig. 0.353 e s Delivered by Publishing Technology to: K. Marhold IP: 78.99.182.222 on: Sun, 11 Mar 2012 10:45:36 Copyright (c) American Society for Plant Taxonomists. All rights reserved. .s,P hieracioides P. s., P. ‘Amrhtp’,bak– black morphotype’), (‘LA aito.B aoia iciiataayi CA4 ae n181idvda lnsadtopedfndgop:wie– white groups: pre-defined two and plants individual 1,861 on based 4) (CDA analysis discriminant Canonical B. variation. eei ditr r akdwt rysmos(rage H opoye,coss–‘Amrhtp’ n aee ytercds Populations codes. Suppl their 1, by Appendix labeled see protologues, and outlines: the morphotype’) dashed in ‘LA and specified – arrows areas crosses or by grandiflora localities assigned morphotype’, indicated some type populations ‘HA are showing for the Populations 1) – analyses. are to AFLP squares Table (triangles (according the subspecies black symbols in recognized full included traditionally gray clustering; not to populations with STRUCTURE attributable are Bayesian marked symbols the empty are clustering; in STRUCTURE admixture cluster Bayesian genetic ‘HA’ the the in cluster to ‘LA’ assigned the and to analyses AFLP the in included 6 aty D ,bsdo 0 utvtdpat of plants cultivated 306 on based 5, CDA Lastly, (6) populations 53 on based 4, analysis component Principal (5) SLOVA on based 4, CDA and 3 analysis component Principal (4) 2012] ftelnetse rnh–Lr it fteinrivlca rcs–WB rsneo akhiso h nour n euce–D,det their to due DH, – peduncle and involucre the on hairs dark of of populations – presence squares WiB, populations): – (101 means bracts population involucral on inner based 3) the (PCA of subsp. analysis width component Principal LBr, A. – plasticity). branch environmental stem longest the of Fig. .hieracioides P. details). for below (see samples the the and of can groupings origin trends AFLP geographic considering certain when but observed groupings, be distinct shown), show not 48 not (figure on did morphotype’ based ‘LA 5, PCA the and of 5), populations (Fig. morphotype’ ‘HA the of separa- 3). the (Table to morphotypes the less strips of contributed tion longitudinal (RSL) ligules red of outer of length the presence the on the (IH), and peduncle (LL), and ligules involucre the on of color-intermediate length hairs contrast, In the (LP). and peduncle (LiB), longest bracts the involucral the (NBr), inner branches of stem length (NL), involucral leaves outer invo- stem of (NoB), on number bracts hairs the (PH), pale 2) peduncle of and step presence lucre material; on the capitula (DC), cultivated of branches distribution on stem the (i.e. here: confirmed 1 of also CDA in were found and presence as 2 dis- morphotypes, the PCA two Characters the indicating between plants. criminating 4), indi- intermediate of (CDA level morphologically the cer- at plants a observed 3), was (PCA vidual overlap averages was of population extent overlap of tain level no the Neverthe- and at 4B). differentiation seen (Fig. clear 4 CDA whereas as in less, 4A), plants (Fig. individual groupings morphotypes the two two did in the to resolved corresponded were characters. that populations 25 all 3, and PCA 2) (step In plants cultivated field-collected or either 1) for (step obtained characters those to yielded three similar plasticity), results morpholog- above, environmental 22 highest the the and and exhibiting (excluding 4) 3 CDA characters PCA in ical in plants individual populations 1,861 (101 plants field-collected hieracioides .Mrhmti nlsso il-olce lnsof plants field-collected of analyses Morphometric 4. – .g P. ., .hieracioides P. ‘Amrhtp’,crls–ppltosof populations – circles morphotype’), (‘LA subsp. n h w sa aa nldn 7quan- 17 including taxa, Asian two the and ilri .v. P. – villarsii .hieracioides P. subsp. ogfla–P . .hieracioides P. l., P. – longifolia subsp. and .hieracioides P. ´ umbellata TA. AOOYO IRSHEAIIE 265 HIERACIOIDES PICRIS OF TAXONOMY AL.: ET K .hieracioides P. ‘Amorphotype’). (‘HA subsp. irshieracioides Picris .hieracioides P. subsp. melt .u. P. – umbellata subsp. aeca–P p. P. – paleacea subsp. uiuaa–P a. P. – auriculata lt aae,toAintaxa, Asian two dataset, plete among Differentiation— 33–53 of from number varied average individual The (mean populations per monomorphic. markers be AFLP to Choc found populations. (LUC, was adjacent or population same indi- One the AFLP few from a multilocus originating in 245 found viduals were detected phenotypes We monomor- identical were dataset. phenotypes; markers data the 1%) the (2. across Five of Repli- phic 2.4%). reproducibility individuals. of rate high 256 error a in (an indicated scored were samples bp cated 54–493 from ing iegneo h w sa aa hc eeresolved were which taxa, nodes Asian terminal two the strong the a all for confirmed of PCoA positions; of except divergence diagram intermediate ordination BS, The in 7). lacked (Fig. clusters clusters smaller these few a that and was cluster well-supported (i.e. with Another together 7). of clustering (Fig. and BS (BS) 77% support receiving each bootstrap branches, long 100% on clusters distinct two formed ih2 opooia hrces(xldn he hrces h length the characters, three (excluding characters morphological 22 with vrl eei tutr n h pce n Population and Species the and Structure Genetic Overall Fingerprinting— AFLP .hispidissima P. umbellata lutaie n oepat edt eaaye nfuture. in analyzed be as of to regarded need available plants be plants more should and few results illustrative, these to species, due Asian however, these 3). cases, 2, both Tables (see In Supplementary margin S2, variation Fig. its Supplementary at also placed was it and characters, of dis- morphotype’ reliable ‘HA most Somewhat 3). a unexpectedly, (Table be character to quantitative appears invo-criminating outer (NoB) the of bracts number lucral The entity. distinct a resolved as plants) 6), (Fig. characters titative .hieracioides P. h w riainae xlie 34%ad1.7 ftetotal the of 12.77% and 33.47% explained axes ordination two The , .hieracioides P. , ‘Amrhtp’.Fl lc ice niaepopulations indicate circles black Full morphotype’). (‘HA .hieracioides P. nteuroe Jte,bsdo h com- the on based tree, NJ unrooted the In eersle ntolreloeclusters, loose large two in resolved were ) 9%B) hl h eto h accessions the of rest the while BS), (93% .japonica P. subsp. D=42.57 = SD subsp. oa f21AL akr rang- markers AFLP 241 of total A ili–P r. P. – rielii .hieracioides P. a on ob iia othe to similar be to found was ˇ rpie .c. P. – crepoides .japonica P. ske ´ , 4). .hieracioides P. .hieracioides P. rh,WCarpathians) W vrchy, .nuristanica P. ntequantitative the in and , .hieracioides P. subsp. subsp. .nuristanica P. .hieracioides P. hieracioides pnls – spinulosa ementary (eight subsp. , Delivered by Publishing Technology to: K. Marhold IP: 78.99.182.222 on: Sun, 11 Mar 2012 10:45:36 Copyright (c) American Society for Plant Taxonomists. All rights reserved. ihteicesn ubro clusters, at of achieved number was increasing the with vs. of structure genetic the exclusion resolve the to with taxa, dataset Asian hieracioides the the on of performed were below hieracioides P. npstoscerysprtdfrom separated clearly positions in 37 [Volume BOTANY SYSTEMATIC 266 ape n 2mrhlgclcaatr.Tetogntclnae r niae;fl lc ice EH,fl rycrls–S-A w vertical explained axes two ordination SW-HA; two – The indicated. (gray circles also admixture gray genetic are regions some full Geographic exhibiting NE-HA, codes. Populations variation. their analyses. – total by AFLP the circles labeled the of black in are 14.51% lineages included and full SW-HA 18.25% not indicated; populations and are NE-HA are symbols the lineages Empty between triangles) overlap. genetic of two zone The the show characters. lines morphological 22 and samples niae httercgiino w ( two of recognition the that indicated only 1.0 for high of sonably similarity of coefficient for a with results clustering s ih us epciey;temr omnadmore and common more the respectively); runs, eight vs. and Italy; central Abruzzo, France). both NE Alsace, VSR, MUN, REFU, Slovakia; SW Male STU, (pop. admixture genetic indicating ments, assign- uncertain with individuals included populations few A lsesbs elcstegntcsrcuepeeti h data. the in present structure At genetic the reflects best clusters fte‘Amrhtp’tgte with together morphotype’ populations ‘LA the the of of composed was other the and morphotype,’ nteSRCUEaaye,temean the analyses, STRUCTURE the In Fig. At K K K K a iil loat also visible was ,atog iiaiyaogtern a lorea- also was runs the among similarity although 2, = ,oecutricue h ouain fte‘HA the of populations the included cluster one 2, = .Picplcmoetaayi PA4 of 4) (PCA analysis component Principal 5. ,tocutrn ucmswr band(ntwo (in obtained were outcomes clustering two 5, = and fgr o hw) ute nlsspresented analyses Further shown). not (figure .hispidissima P. K K 06) hs h TUTR analyses STRUCTURE the Thus, (0.67). 5 = ,btasalices ntepo of plot the in increase small a but 2, = K .Rpiaern ile stable yielded runs Replicate 5. = nmr detail. more in .hispidissima P. K .hispidissima P. )adfv ( five and 2) = K h highest The . irshieracioides Picris L ( K increased ) ´ Fg 1B). (Fig. Karpaty, K =5) and △ △ P. K K subsp. ieycutrn sitrrtdhr.I lutae h distinc- the illustrates It of here. tion interpreted is clustering likely ouain rmnrhr tl n n rmN France NE between admixture from of level detectable, one still but and low, a Italy showed Three northern Slovakia. from from LPT population populations the also in and individual populations one (AQU, PIRO) in Italian KRA, the central (VRA, some Croatian in in and evident CAM) also the was between and lineages Admixture ‘LA’ BAN). UGI), west-two and (CDL, two MAR in populations (pop. observed Carpathians Alpine was (W) lineages ‘HA’ ern two the between Admixture lineages. these between admixture indicating pop- uals, individ- Certain intermediate Romania. genetically harbored and however, Slovakia ulations, (NE-LA), ‘LA from morphotype’ range the ‘LA NE the the of of populations 5) populations and Croatia; 4) Italy, and France, Spain, (SW-LA), Alps; range SW and the from morphotype’ Carpathians morphotype’ compris- the ‘HA (NE-HA), range the ing (NE) of northeastern populations the from 3) range mainly Alps; (SW) SW Pyrenees, and Nevada, southwestern Sierra Jura, the including the SW-HA), as from (denoted mainly morphotype’ 1) resolved 1): were (Fig. clusters follows as five The of structure. geographic morphotypes’ certain ‘LA and ‘HA’ the umbellata .hispidissima P. ‘Amrhtp’ ae n5 il-olce populations field-collected 53 on based morphotype’) (‘HA .hispidissima; P. n w eei iegswti both within lineages genetic two and )ppltoso h ‘HA the of populations 2) .hieracioides P. indicating Delivered by Publishing Technology to: K. Marhold IP: 78.99.182.222 on: Sun, 11 Mar 2012 10:45:36 Copyright (c) American Society for Plant Taxonomists. All rights reserved. ‘Amorphotype’; (‘HA T S lvka xiie otiuinfo l four all from contribution a of exhibited lineages genetic Slovakia) (SW population STU the Interestingly, lineages. between SW-LA intermediate and completely SW-HA the as resolved were (REFU VSR) Italy central and Abruzzo, In from MDR). populations (pop. and two lineages addition, MUN), NE-HA and (pop. FUG), SW-LA lineages the SGA, between SW-LA also (populations and SW-HA lineages the NE-LA between and NE-HA the SLOVA 2012] esue n orgop r-eie:surs– squares pre-defined: groups four and used ters oae na nemdaepsto,btente‘A and ‘LA’ the of between position, populations intermediate of ‘HA’ cluster an The in delimited. genetically located clearly the to be to can corresponding clusters regards three with the The also populations. well, admixed clustering ian WH eefudi eea mle lses h two The of clusters. populations smaller the several while in lineage, found NE-HA were popu- comprised the SW-HA cluster of large lations one clusters: several formed Fig. h egbre iga Fg )spotdteBayes- the supported 8) (Fig. diagram NeighborNet The .hispidissima, P. .Cnncldsrmnn nlss(D )bsdo utvtdpat of plants cultivated on based 5) (CDA analysis discriminant Canonical 6. n H’ad‘A ouain of populations ‘LA’ and ‘HA’ 9) seik – asterisks 190), = .hieracioides P. .hieracioides P. seFg 1). Fig. (see .japonica P. h H’populations ‘HA’ The . ´ TA. AOOYO IRSHEAIIE 267 HIERACIOIDES PICRIS OF TAXONOMY AL.: ET K .hispidissima P. .hieracioides P. ( n 4,tinls– triangles 14), = .hieracioides P. subsp. was hieracioides .nuristanica P. U,adFG eepae uti h etro the PCoA of Separate center shown). ‘LA’ not the the (figure by in populations occupied just ‘HA’ space ordination placed and the were STU, between and populations FUG) graph, the STU, from and SGA, individuals some MUN, REFU, as MUN, well as MDR, VSR (FUG, admixed cally of photypes itntgop,matching groups, distinct the shif in somewhat detected be to individ- as appeared these uals populations, here; also MUN apparent was and analyses, STRUCTURE MDR two the other admix- in Genetic the ture lineage. and SW-LA the lineage, clusters; of NE-LA populations three comprised the into to divided corresponded were clus-one populations ‘HA’ ‘LA’ STU different The peculiar cluster across ters. the scattered the found from towards Individuals were shifted populations. population was ‘LA’ that the position of a in clustered together REFU) (VSR, populations intermediate remarkably nln ihterslsgvnaoe CArvae three revealed PCoA above, given results the with line In ‘Amorphotype’; (‘LA irshieracioides Picris ( n =8). .hieracioides P. n t ls eaie,wt 7qatttv charac- quantitative 17 with relatives, close its and n =116),circles– ouain eelda geneti- as revealed Populations . .hispidissima P. e oad h H’clusters. ‘HA’ the towards ted .hieracioides P. n h w mor- two the and subsp. umbellata Delivered by Publishing Technology to: K. Marhold IP: 78.99.182.222 on: Sun, 11 Mar 2012 10:45:36 Copyright (c) American Society for Plant Taxonomists. All rights reserved. lseigadodnto eut.I eeldsignificant revealed It results. between ordination differentiation above the and predefined supported the clustering computations to AMOVA assignment These unequivocal groups. MUN) their and MDR, of SGA, because FUG, STU, VSR, (REFU, populations hieracioides P. analyses above the by ( suggested differenti- groupings genetic the of among degree ation the hier- tested three-level we the AMOVAs In archical them. among occurred variation the of ouain,adters (47.9%, rest the and populations, populations shown). ‘LA’ not (figure admixed above, genetically revealed or as populations outlined the involving as ‘HA’ overlap considerable structuring, with the geographic further only showed including analyses 37 [Volume BOTANY SYSTEMATIC 268 oNiadL’ 17)gntcdsacs(e cl bar). scale (see distances genetic (1979) Li’s and Nei to rnhsidct otta upr rae hn5%(u o hw o emnlndswti ouain) w useis eiie sarsl of result a as delimited subspecies, Two populations). within nodes terminal for shown not (but 50% than study, greater this support bootstrap indicate branches ouain of populations .hispidissima P. natolvlAOA 21 ftevrainwswithin was variation the of 52.1% AMOVA, two-level a In Fig. .Uroe egbrjiigte ae nteAL aaof data AFLP the on based tree neighbor-joining Unrooted 7. .hieracioides P. ,btecuiggntclysrnl admixed strongly genetically excluding but ), vs. .hieracioides P. .hieracioides P. subsp. .hispidissima, P. hieracioides hwn ht2.%( 26.0% that showing , L’v.‘A ouain of populations ‘HA’ vs. ‘LA’ ; F ST ‘Amrhtp’ n subsp. and morphotype’) (‘LA .8 .f 60, = f. d. 0.48, = h L’ad‘HA’ and ‘LA’ the F p CT < 0.26, = 0.001) irsheaiie,P ipdsia .nuristanica P. hispidissima, P. hieracioides, Picris umbellata sinet otetogntccutr,a nerdby the inferred with congruent are as clusters genetic two clusters, the that evident genetic is two at analyses the STRUCTURE to assignments p .f 1, = f. d. aito a on mn ouain ihngop and groups within populations ( 44.8% among found was variation ntemrhmti C Fg A ftefield-collected the of of 4A) (Fig. samples 3 population PCA morphometric the In n h L’ad‘A ouain stogop,televel the groups, ( two 24.8% significant; as highly also populations was differentiation ‘HA’ of and only ‘LA’ considering the and When populations. within rusad4.%( 44.4% and groups .f 2, = f. d. rus hra 04 ( 30.4% whereas groups, yteetregop,wees2.%( 29.6% whereas groups, three these by < oprsno h opooia n eei Structure— Genetic and Morphological the of Comparison ‘Amrhtp’ r loidctd rebace r proportional are branches Tree indicated. also are morphotype’) (‘HA .0)o h aito a mn ouain within populations among was variation the of 0.001) F ST p < .5 .f 151, = f. d. 0.55, = p .0)o oa ainewsepandb h two the by explained was variance total of 0.001) < .0)o h oa ainewsexplained was variance total the of 0.001) F ST F .6 .f 159, = f. d. 0.56, = SC .hieracioides P. K .0 .f 50, = f. d. 0.40, = (excluding 2 = p < .0)wti populations. within 0.001) and .japonica P. eidctdtheir indicated we , F SC .hispidissima P. .0 .f 51, = f. d. 0.40, = p p < ubr along Numbers . .hieracioides P. < .0)o the of 0.001) .0)was 0.001) F CT =0.25, .It ). Delivered by Publishing Technology to: K. Marhold IP: 78.99.182.222 on: Sun, 11 Mar 2012 10:45:36 Copyright (c) American Society for Plant Taxonomists. All rights reserved. orppltosatiual oteN-Alnaewere lineage populations, NE-LA ‘LA’ the the to of morpho- shown) attributable the not populations In (figure overlap. four 5 with PCA albeit central delimited, metric (4) be Pyrenees, and can regions), Nevada, adjacent Italy (Sierra and range Alps, the western westernmost the from the the those well; from (2) as populations, (3) Carpathian space Alps, (1) ordination the the of in groups evident analyzed the is of origin populations structure geographic Some exact more not. the were zone, to BAN) according overlap (CDL, that two in other right the found whereas were lineages STU) some and NE-HA showing UGI, and (MAR, populations SW-HA the Three between center. admixture over- considerable genetic the with in but part evident SW- right lap the the of in those placed whereas were diagram, HA the ordination in the mainly of placed part were left lineage NE-HA NE-HA at the and analyses of STRUCTURE populations SW-HA by the inferred to as lineages, assignments their indicated of we majority the where cluster, dominant belong. individuals range respective variation the the within placed of are MUN) these and MDR, between SGA, admixture at either genetic individuals (seen of few clusters levels a some with populations exhibiting The morphotypes. two SLOVA 2012] ouainabeitosflo pedx1 upeetr al .Saebri ecn distance. percent is bar Scale 1. Table Supplementary 1, Appendix follow abbreviations population hieracioides ntemrhmti C Fg )o h H’populations, ‘HA’ the of 5) (Fig. 4 PCA morphometric the In Fig. .NihoNtdarmof diagram NeighborNet 8. ‘Amrhtp’ n subsp. and morphotype’) (‘LA K =2or K irshieracioides Picris ;RF,VR T,FUG, STU, VSR, REFU, 5; = umbellata ´ TA. AOOYO IRSHEAIIE 269 HIERACIOIDES PICRIS OF TAXONOMY AL.: ET K ‘Amrhtp’,a ela eei iegsrcgie ihnbt useisaeotie.The outlined. are subspecies both within recognized lineages genetic as well as morphotype’), (‘HA and K .hispidissima P. =5.The eie rmAL akr.Tetosubspecies, two The markers. AFLP from derived nteWCrahasadPnoi.Rr FPmarkers, AFLP Rare Mala KLAK, Pannonia. (pop. and 3 between Carpathians central ranging W in and the populations), Austria, Alps admixed in western the genetically several in of (including Slovenia), south Italy Italy, concentrated (N) and were (northern LUC) variation of (pop. genetic populations 0 high The LPT). between (pop. varied 21.99 percentage markers the polymorphic and 1), of Table Supplementary 1, Appendix see Choc LUC, (pop. 0 between varied hieracioides— the regarding regard- either or geography. their populations seen ing admixed but genetically pattern graph, the ordination no of the placement was of corner there left otherwise lower the in located n anna n loi h yees hncomparing When Pyrenees. the in Cro also N and Italy, Pannonia, (N and Alps the di of highest that south showed the values per DW with and markers fragments populations private specific and rare five of tion to Franche-Comte PDR, up (pop. in in population reaching observed recorded populations, were were markers Italy), 22 (specific) NW Private Piemonte, populations. GAT, all (pop. 20 and Liptovska LPT, (pop. 0.0887 and ihnPplto eei iest n iegnei Picris in Divergence and Diversity Genetic Within-Population h ihnpplto e’ eediversity gene Nei’s within-population The egnewr otyfound mostly were vergence ˇ ta,i h Carpathians W the in atia), ´ ske oln,WCarpathians; W kotlina, ´ ´ ´ .hieracioides P. rne.Tedistribu- The France). , rh,WCarpathians) W vrchy, ar,WCarpathians) W Fatra, .hieracioides P. exhibiting subsp. Delivered by Publishing Technology to: K. Marhold IP: 78.99.182.222 on: Sun, 11 Mar 2012 10:45:36 Copyright (c) American Society for Plant Taxonomists. All rights reserved. n EH iegs opie h ouain fu to up of populations the comprised lineages) NE-HA and 7 YTMTCBTN Vlm 37 [Volume BOTANY SYSTEMATIC 270 nta td ste‘oe liue(A opoye and morphotype’ (LA) (Slova altitude denoted morphotype’ ‘lower (HA) and altitude popula- the Pannonia ‘higher as on the and study based Carpathians that study W in the previous from a tions in suggested already nterslso eei nlss(i.1) TUTR ana- STRUCTURE 1B). (Fig. at seen analyses to lyses genetic be of attributable can results picture populations the same in comprised The subspecies. 4A) cluster traditional (Fig. morphological several compact 3 a PCA time, same on the At 4A). Fig. phenetic (e.g. large space a this be occupying to illustrate heterogeneous, found clearly morphologically were 4A) subspecies and traditional Several 1B of discrepancy. Figs. areas the see and/or description, localities their type described their previously on the assignments (based to subspecies in The here 1). analyzed subspecies populations Table the (see of numerous discordant were recognized and taxo- Europe that traditional concepts the with nomic contrasts strongly here, presented them. among flow gene the in facilitated individuals have and proximity differentiated gardeners close genetically by brought introduction likely unintentional most Western the the locali- and Italian in some ties) in habi- (observed and activities mountain human of through Italy disturbance tats The northern 1A). Fig. and (see Carpathians central observed in were but mainly distributed, randomly not These genomes. were admixed individuals genetically assignments of uncertain existence showing the indicate individuals genetic few two of A pattern lineages. same the reveal clustering) network (i.e.generating and employed neighbor-joining, approaches Bayesian, the analysis, of ordination all pres- flow subspecies, gene the the some between tree, indicating neighbor-joining individuals), intermediate the of in ence lack support a (e.g. bootstrap support strong of subspecies lack patterns the genetic the at Although appropriately most as entities, level, two these recognition the of the with confirms congruent it and is differentiation, markers morphological AFLP from inferred variation is to morphotype morphotype ‘HA’ to the here within attributed below), described Treatment origi- taxa the Taxonomic of several (see study of the rec- material on be Based nal should species. AFLP subspecies this two by within that revealed ognized conclude as we groupings, Thus, genetic data. main two the with ain abrd4 rvt ns(upeetr al 1). Table popu- (Supplementary ones ‘HA’ private the 48 whereas harbored markers, lations AFLP similar private charac- 37 yielded were populations by both ‘LA’ terized The they variation. genetic morphotypes,’ of ‘HA levels and ‘LA’ the ag xet lob ifrn eoeszs(Slova sizes genome different to Slova by and, 2007; also (longevity) Marhold extent, traits characterized large history are life a ecologies, they different but by morphotypes geographically, two overlap The pattern. presentlargely main the this in corroborates Europe study across sampling large-scale The 2007). h nlsspeetdhr eosrtdta h two the that demonstrated within here observed morphotypes presented analyses The h w opooial eldfeetae niiswere entities well-differentiated morphologically two The h udvso of subdivision The atrso nrseii aito nPci hieracioides— Picris in Variation Intraspecific of Patterns K .hieracioides P. hwdta oegntccutr eg SW-HA (e.g. clusters genetic some that showed 5 = .hieracioides P. .hieracioides P. .hieracioides P. ´ ta.20a e loblw.Tegenetic The below). also see 2009a; al. et k subsp. .hieracioides P. Discussion subsp. hieracioides subsp. subsp. .hieracioides P. villarsii, hieracioides notosbpce as subspecies two into umbellata n subsp. and subsp. ´ n Marhold and k r congruent are . .hieracioides P. crepoides n h ‘LA’ the and umbellata ´ kand see , . aebe eotda h otipratcaatr o the Bolo for 1976; characters within subspecies important shape the most leaf of the and delimitation hairs, as of reported color been bracts, have involucral the of length bracts involucral inner below). of outer key the of length also number the (see the and ligules, num- of bracts, the length branches, involucral the stem leaves, of stem number of strips the ber longitudinal ligules, red outer of the presence on the branches, capit- of of stem distribution color on the ula the peduncle, and follows: involucre conditions. garden the as on their environmental a hairs are in same to characters plants diagnostic transferred the Important to those under to only cultivated also not and but applies habitats, finding natural this and here, rlIay(buz)btas a enrpre rmteAlps the from reported been has also but (Abruzzo) Italy tral Further- Europe. throughout more, nearly from reported been have to belonging populations l he aasol esmldt seti hi genetic their ascertain (Slova to precisely from sampled more distinctness populations be morphological should additional and taxa drawn, three all is conclusion nomic of subspecies two the homogenous subspecies differentiates ters same and the different Europe. central as and genetically Alps treated the from the populations be with cannot together which 1), (see admixed genetically Fig. largely be to revealed Abruzzo, were in however, sampled populations The Carpathians. W the and aerle nafwseiesol n aeover- of have continuum and the only of part specimens only few are in variation that a traits on emphasized circumscriptions relied subspecies previ- and have Apparently, treatments keys. taxonomic identification consid- ous in been all important never bracts, have as involucral diagnostic, ered outer being as the capitula here of of identified number (distribution the structure the branches), branches, panicle on stem ligules, leaves, stem of of length number the as such acters (subsp. found morphotype’ be ‘HA can the range in color whole (here the morphotype’ should hairs, color-intermediate ‘LA trait, subsp. the diagnostic as while defined potential carefully: a interpreted be being of the color although to the thus, Nevertheless, inner hairs, contribute meaningful. and, the hairs taxonomically populations, be of of sampled can length the color the among the that discrimination and indicate bracts study involucral present the of ieg loicue bsdstoIeinedmcsubspe- endemic Iberian two cies (besides included also sub- lineage certain clusters while genetic (e.g. different two subspecies, between dispersed were traditional species different three oki progress). in work the of differences from morphological taxa clear state Asian to difficult was from ferentiated to hispidissima close P. very appears still it at but lyses), cluster neighbor-joining genetic the separate in a support tree, bootstrap (93% well as taxon and etBla endemic Balkan west hrcessc stelnt ftepdnls oo and color peduncles, the of length the as such Characters obnto fsvrlqatttv n iaycharac- binary and quantitative several of combination A ae nAL akr,bt sa species, Asian both markers, AFLP on Based .nuristanica P. .hieracioides P. .hieracioides P. .hieracioides P. ` n io19;Hepe n ur20) Results 2000). Muer and Haeupler 1995; Vigo and s .hieracioides P. smrhlgclyadeooial eldif- well ecologically and morphologically is .hieracioides P. .hieracioides P. r hw ob eldfeetae.The differentiated. well be to shown are , hieracioides subsp. subsp. subsp. .hispidissima P. . villarsii grandiflora .hieracioides P. longifolia er aeo tms (rarely) most at or pale bears ) hs eoetefnltaxo- final the before Thus, . seSlova (see umbellata .Frisac,teSW-HA the instance, For ). K em ob well-defined a be to seems nSRCUEana- STRUCTURE in 5 = n subsp. and a ecie rmcen- from described was .hieracioides P. .hieracioides P. .hieracioides P. subsp. ´ .I otat char- contrast, In ). ta.20b,it 2009b), al. et k villarsii rielii .japonica P. Sl 1975, (Sell .Whereas ´ defined )some ketal. that Delivered by Publishing Technology to: K. Marhold IP: 78.99.182.222 on: Sun, 11 Mar 2012 10:45:36 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 21,seAOArsls a bevdin observed component was results) variation AMOVA see distinct within-population 52.1%, a (the representing within populations also individual variation considerable each and phenotype, multilocus almost with hieracioides— tion, Picris of History the or 2004), 2008), Waterway 2010), and al. (Smith et Bardy 2004; 02 SLOVA 2012] re ntxnmclyitiaepatgop,a demon- as in groups, studies plant the intricate by when strated taxonomically successful in bound- species proven aries re-assessing and also concepts DNA taxonomic has revising variable highly markers with fingerprinting data morphometric Combining or 2005), (Robart taxo- few bracteosa or Hemizygia or in one as continuous into such lumped a entities, be nomic show should fact and the in variation at clinal levels recognized infraspecific previously or taxa many specific study, our shown like have that, morphometrics multivariate of means by species shown was negligible. the separation but be morphotype analyzed, and to the and length to measured also contribution and were their width pappus the the achene of on the length study 2007), previous Marhold a and In of separation. complex populations species contrib- Carpathian/Pannonian extremely achenes the on is to characters the uted characters only and floral continuous, variation and and morphological the vegetative that in shown have studies 2000) (Smalla morphometric Conversely, on 1994). Lack mor- and Holzapfel of (Holzapfel set here 1993; comparable identified a as characters by phological other each from phylogenetically distinguished and sub- morphologically to three close are with which species polymorphic species), one including species a aehdsrn mat nteseis evolution species’ (Slova the self-incompatibility on 1) and impacts of variation diversification: strong biology the and had and (increase) ecology have affect the may apparently to species related this factors main Three yees U rmteWsenCrahas n MULA and Carpathians, Western the the from from LUC (LLV some populations Pyrenees, in depauperate values fragments) genetically rare high these dis- (DW, of hand, parameters strongly divergence other were the the and of On only anthropogenically. individuals turbed few recent, localities a these a comprised of all or almost events Indeed, spread. human-mediated dispersal Pyrenees, long-distance the recent 1, via by Appendix explained from colonization be see can few Carpathians; however, W 1), a Table or Supplementary Italy Nevada, southern (e.g. Sierra from populations and the some from in con- variation those into genetic and, populations sites, Decreased differentiated new tact. bring of may colonization addition, rapid fields in the abandoned and expansion or ‘anthropogenic the highways, along roads, in spreading (especially as of such Slova efficiency M. corridors,’ high 1993; (Sheldon 3) Andersen pappi and 1973; well-developed Burrows dispersal bearing and long-distance and achenes of within ability through flow the gene 2) and populations; outcrossing among promotes that 2008) h eei tutr n yohsso h Evolutionary the on Hypotheses and Structure Genetic The eea te uhr eetyivsiaigpolymorphic investigating recently authors other Several genus the of representatives Australian Picris .hieracioides P. pce rmteAainPnnuaadSocotra and Peninsula Arabian the from species lsu montanum Alyssum .hieracioides P. eoiachamaedrys Veronica Oin ta.2006), al. et (Otieno Lc 99 aule l 06,cnbe can 2006), al. et Samuel 1979; (Lack ieai deltoidea Cineraria ae roanensis Carex Veronica subsp. admn amara Cardamine ope (S complex ihoealgntcvaria- genetic overall High hieracioides (Martı .hieracioides P. ´ Brye l 2010). al. et (Bardy TA. AOOYO IRSHEAIIE 271 HIERACIOIDES PICRIS OF TAXONOMY AL.: ET K eiuai bracteosa Pedicularis ˇ n lidspecies allied and ´ aile l 2011). al. et paniel e-reae al. et nez-Ortega Co ta.2007), al. et (Cron Picris ,wihallows which ), ´ .hieracioides P. es obs.); pers. k (Lihova 1 native (10 ´ tal. et k (Slova ´ tal. et ´ k . lobe ouetd(..Kofe l 03 ane al. dispersal et major the Paun of lineages been 2003; two apparently the al. for have et barrier Alps Kropf The (e.g. have 2008). populations documented Alpine been eastern and also Carpathian his- the Lihova evolutionary of 2009; common tory al. and relationship et Scho close Dixon several a Similarly, 2008b; (e.g. for al. et also species Ronikier discussed mountain and other revealed Alpine western was and Iberian populations dispersal the patterns. between long-distance variation relatedness observed and Genetic the to spread contributed have main- recent also been may more has that structure A internal suggests severely Gene tained. little been thus, regions. not ranges probably and, larger has restricted, NE the populations across the of and altitudes among periods flow cold lower SW the at the survived Pleistocene have in may populations both the with pool gene see refugia widespread Mra the we isolated 2006; Conversely, 2008a). al. al. and et et Kropf Ronikier distinct (e.g. flow in gene restricted survival scenario vicariance moun- a species each under expected in of be lineages would genetic as and structure. range, distinct tain flow observe genetic We not gene this did m). shaped past We (289–600 have history, Alps processes, (post)glacial and colonization the Carpathians that line- the NE-HA assume the in the while grows to Alps, westernmost extends age Sierra the and the and Pyrenees, Mts. in and Jura m) Mts., (552–2,370 Montseny altitudes Nevada, high mainly occupies subsp. the favors strongly geographically habitats scenario. former disturbed areas of con- restricted observation their the rather and but certain, differentiation, incomplete in and centration initial con- secondary or either tact individuals document intermediate may found few populations accumulation and The the differentiation. and (longevity) genetic traits differentiation,of life-history morphological in shift by subspecies a habitats, accompanied sympatric different was largely to which adaptation these of ecological subspecies that through two evolved speculate the can of origin we the lineages. of genetic hieracioides mode P. and and subspecies time the The between small either overlap showing no thus, NE-HA), or in pg 2.94–3.11 SW-HA; populations subsp. the of in of values (2.39 2C the lineage; NE-LA), SW-LA in in pg 2.74–2.75 pg LPT–) (2.30–2.46 pg 2.30–2.75 of range die ouain SU A,BN r on nthe in found are BAN) genetically MAR, other (STU, Three UGI). populations from (CDL, admixed populations Alps two in westernmost between only the there hybridization observed been suggesting has them, admixture, genetic and eoesz of size genome popu- in decrease varia- a genetic and of disruption size. habitat loss lation recent a to indicate due founder rather tion the with and contrast hypothesis in are effect Italy) southern Calabria, from ieigtetosbpce eonzdadterinternal of their populations the and subsp. con- Whereas recognized by lineages. here subspecies genetic explained two be the can variation sidering size genome much Indeed, this species. of this strongly of history evolu- it colonization complex and range; and/or tionary heterogeneity intraspecific taxonomic a an indicated for high extraordinarily h w eei iegsdtce within detected lineages genetic two The osiuu aito a eetyosre nthe in observed recently was variation Conspicuous hieracioides umbellata umbellata antb setie ihtepeetdt,but data, present the with ascertained be cannot .hieracioides P. r agl loarc h WH lineage SW-HA The allopatric. largely are nlzdhr xiie Cvle nthe in values 2C exhibited here analyzed agdfo .331 g(.329 gin pg (2.73–2.94 pg 2.73–3.11 from ranged (Slova .hieracioides P. ´ ta.20a,wihis which 2009a), al. et k ¨ setre l 2002; al. et nswetter subsp. .hieracioides P. ´ ´ .hieracioides P. ta.2007; al. et z ta.2009). al. et umbellata , Delivered by Publishing Technology to: K. Marhold IP: 78.99.182.222 on: Sun, 11 Mar 2012 10:45:36 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 7 YTMTCBTN Vlm 37 [Volume BOTANY SYSTEMATIC 272 usqethbiiain h opooia variation, within continuous morphological largely The is and however, communities) hybridization. three cottage all gardener’s subsequent (as in ornamentals uninten- grow their with by populations by together likely, originated introduction more tional even probably or, events populations long-distance These Carpathians. species ranges. mountain ferent umbellata nlsso w aaes(il-olce n cultivated and (field-collected here. datasets employed of two also plants) data was on exploratory approach and analyses analyses last discriminant This canonical Using 2004). al. et bandb utvto nacmo adnexperiment garden common specimens a using in ( other cultivation the by compris- obtained specimens, one datasets, field-collected two ing with ( values evaluation different factors morphometric to environmental exposure of with experiments cultivation amarum environmental ( characters between morphological qualitative correlations and stud- factors examining taxonomic as in seen such be in approaches can ies, critical plasticity Different highly of 2002). assessment be the Kaplan may to (e.g. it groups though plant even some addressed detail, have in studies recognition. issue taxonomic this taxonomic few a formal only Nevertheless, deserves taxonomi- (and interpreted) as observed so variation cally studies, the taxonomic whether infraspecific recognize in to should al. concern to plasticity major phenotypic et interest of Certainly, be Coleman 1995). great al. (e.g. of et Via biologists thus 1994; evolutionary is and and has ecologists fitness environments, maximizes It that variable plants. response in functional in a phenomenon as viewed display well-known been to a environmental genotype is different to conditions, a response of in phenotypes ability hieracioides— different the Picris plasticity, in Plasticity Phenotypic Phenotypic – high Specimens expected. and been fact, zone in contact colonization has, large admixture their genetic the of considering rate, levels rap- Therefore, outcrossing expand and areas. and ability new fields) to (e.g. abandoned idly corridors’ and ‘anthropogenic motorways, ruderal along these roads, well, spread as coloniza- can refugia distinct glacial populations different indicate Balkans. from may western routes pattern the tion and SW-NE Italy the genetic central Although genetically exhibiting in populations present populations, a with is Carpathian zone admixture form contact most large Spain a do but NE as and group, southern homogeneous Sicily) from their (Calabria, Populations However, weaker. Italy lineages. much genetic is NE differentiation and SW of occurrence h w opoye eemitie nbt aaes As datasets. both in maintained between were differences morphotypes including relative two cases, the all the characters, in WiB mainly but and characters, sizes, LBr organ the other and some plant in to compared. referring also were dark observed values plants were their of field-collected Shifts in presence and shifts – cultivated pronounced (DH when most involucral the character inner exhibited the binary hairs) of one of width taxonomic length and the reliable – the bracts) WiB – are and (LBr they branches, characters stem if quantitative (i.e. Two markers). sta- cultivation retain after samples bility field-collected for diagnostic potentially Cerastium eei aito ntelwad ihyrdrlsub- ruderal highly lowland, the in variation Genetic h oprsnBtenFedCletdadCultivated and Field-Collected Between Comparison The ietladShqil 07,taslnainand transplantation 2007), Sahuquillo and Pimentel ; .hieracioides P. hwn nysih opooia hfsi dif- in shifts morphological slight only showing , rsigadEvn2000; Elven and Brysting , .hieracioides P. subsp. eeaie fcaatr htare that characters if examined we , Potamogeton hieracioides admn acris Cardamine .hieracioides P. aln20) or 2002), Kaplan ; loidctsthe indicates also Anthoxanthum Perny , subsp. ´ ecnie,adtefrtad9t ecnie r given are percentiles 95th 99th and and fifth first of brackets. the the in ranges to and The correspond percentiles, individuals. characters field-collected quantitative to refer ranges hieracioides— characters the on impact diagnostic. small taxonomically a as only here of has identified variation it morphological but species, total this the to cultivation contributes certainly in most maintained and not populations, 2). was Table Supplementary these (see trait of this be to several importantly, observed in were individ- peduncles intermingled but prolonged peduncles, clearly shorter with uals generally have Mediterranean indeed the the in area of altitudes lower features at diagnostic sampled Populations have the of inflorescence, one European congested south be and to considered dense been a of fea- appearance this that confirm we popu- value. taxonomic so mountain no study, has this ture adjacent in seen from were lations populations these continuous of Spanish other a differentiation No two be genetic margins. or undulate to distinction strongly seems morphological to there flat Pyrenean from and other transition well, in observed as been populations tendencies have Nevertheless, margins cultivated. undulate to and seeds from grown nuaela agn n ananti etr when feature this maintain and margins leaf undulate (Bolo sindt h bra endemic populations. Iberian the within traditionally to EST), characters even (LLV, assigned Pyrenees all vary, the from strongly that populations Two leaves already noting has to value, 1993), related taxonomic Lack their and Holzapfel rejected 1974; (Lack studies hs etrshv rvosybe noprtdi identifi- Padalı in (e.g. incorporated keys cation been previously have features Although these here. examined characters the among included been research further requires 2005). and al. understanding et external (Koes its both factors, and regulatory internal, and complex genes enormously an of pro- by network Flavonoid regulated the them. is thus in however, stimulated and synthesis, likely lowlands, is flavonoids the of from duction those of than complemen- Plants radiation a variation. bear- this have and umbellata mind carefully can in interpreted character when ing the value condi- Still, diagnostic lowland tary dropped 12%. value in only this cultivated experiment), to garden were common plants (our tions the when ever, 3 ffedcletdidvdas upeetr al 3) Table Supplementary (in individuals, in commonly field-collected quite and of dark exclusively 43% almost pedun- the and found involucres Regarding were the cles characters. on identified hairs diagnostic dark and (DH), of morphotypes pigmentation set two same same they the the patterns; similar delimited in both resulted cultivated samples, on field-collected either or based analyses, ordination the expected, dniiainKyt h w useiso Picris of Subspecies Two the to Key Identification in plasticity phenotypic that conclude can We the for responsible peduncles, shortened Extremely h ieadsaeo tmadrstelae aenot have leaves rosette and stem of shape and size The .hieracioides P. ` n io19) emt ecaatrzdb strongly by characterized be to seem 1995), Vigo and s ‘Amrhtp’ r xoe ohge UV higher to exposed are morphotype’) (‘HA h ecitosgvnblwadtevalue the and below given descriptions The .hieracioides P. subsp. aooi Treatment Taxonomic ´ kova umbellata ´ 92 Za 1972; subsp. ‘Amrhtp’.How- morphotype’). (‘HA .hieracioides P. ´ gei17) ak nhis in Lack, 1976), ngheri spinulosa .hieracioides P. eg el1976). Sell (e.g. .hieracioides P. subsp. subsp. rielii Delivered by Publishing Technology to: K. Marhold IP: 78.99.182.222 on: Sun, 11 Mar 2012 10:45:36 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 1A irshieracioides Picris aei hp,ots oauea px eilt,salwyto mostly leaves shallowly Rosette petiolate, pinnatifid. apex, or margin, at at acute dentate deeply to lanceo- numerous, obtuse widely leaves shape, to in linear-lanceolate Basal late rosettes, anchor-shaped, hairs. leaf and in spinulose arranged 2-furcate rigid Leaves simple with (1–)2–26(–39). and hairy branches of densely number to usually along stem, moderately usually entire distributed Stem the branches length. less stem entire branched; or the densely more along hairy, numerous hairy densely to to equally green to few grooved, moderately longitudinally bearing reddish-brown, slightly usually Stem taproot, roots. lateral simple a with 02 SLOVA 2012] .Cptl nyi h pe /–/ fse rnhs euce n ue nourlbat ihclritreit bons ogaih rblack or grayish) to (brownish color-intermediate with bracts involucral outer and Peduncles branches. stem of 1/2–1/3 upper the in only Capitula 1. .Cptl itiue ln h hl tmbace.Pdnlsadotrivlca rcswt otypl,rrl rwiht rys btnever (but grayish to brownish rarely pale, mostly with bracts involucral outer and Peduncles branches. stem whole the along distributed Capitula 1. irshieracioides Picris longifolia Picris hispidissima Picris aspera Picris irsangustissima Picris irscorymbosa Picris irssetulosa Picris irsspinulosa Picris ruderalis Picris nult inil ioos 510–0)c ihplants high cm 15–150(–200) vigorous, biennial, to Annual irshieracioides Picris it-u.42 08 99 o.ivl At 24.3). (Art. inval. nom. 1929, 1038. 4/2: Mitt.-Eur. o.Sd :50 83 o.ivl,nmnud. nom inval., nom. 1883, 520. 8: Sada Bot. syn. pro inval., nom. 1875, 735. 3: 32.9). (Art. 74198!). G (lectotype: a vagues [SWITZERLAND] Terrains here, designated 1905.—TYPE: 331. 5: tl:48 82—YE IAY aem (original Fl. Palermo Comp. Arcang., [ITALY] Gibelli) 1882.—TYPE: unknown). & material 418. Pass. Ces., Ital.: ex 1878; (Guss. 469. 2/20: Ital. YE FAC]Evrn ePrinn 1845, n. Perpignan, s. de Environs [FRANCE] TYPE: Gussone!). NAP-herb. ee[TL]Plro 89 [ designated 1839, 1882.—TYPE: Palermo, 418. [ITALY] Ital.: here Fl. Comp. Arcang., 1843; rupibus 14634–010!). a., B-W In type: s. Pragam, REPUBLIC] circa [CZECH Bohemiae 1803.—TYPE: 1558. rsieo uea eohrosotil,43ma.s.l., of m village 453 39 Calabria, outfield, xerothermous Italy, ruderal designated: Frascineto, here 387 113): EPITYPE: (1975: Cliffortianus Lack siccus by Hortus designated 52).—TYPE: (Art. illeg. undulata hieracioides hieracioides .n. s. rrl ae -uct n nhrsae ar;inrivlca rcs(–1–5–6 mln,lgls(.–741.(1.)m og e long red long; (6.9–)7.4–15.8(–19.9) ligules mm long, (9–)10–15(–16) mm bracts involucral inner hairs; anchor-shaped and 2-furcate pale) (rarely tisi h pe ato uems iue yial bet nyrrl rsn . present rarely only absent, typically ligules outermost of part upper the in strips lc)2fraeadaco-hpdhis ne nourlbat 5).–09–16 mln,lgls(.–691.(1.)m og outermos long; mm (6.2–)6.9–12.4(–14.2) ligules long, mm (5–)6.1–10.9(–11.6) bracts involucral inner hairs; anchor-shaped and 2-furcate black) iue otywt e ogtdnlsrp nteruprpr . part upper their in strips longitudinal red with mostly ligules 49.899 hltp:P). (holotype: (SAV!). irshieracioide Picris ii. l i.Ic.1 2.18,nm inval. nom. 1782, 227. 1: Inch. Lit. Fl. Gilib., 0 ua,F.Hautes-Pyre Fl. Dulac, nr.e ruv,TuyIp S.-Peterburgsk. Imp. Trudy Trautv., ex Andrz. us xCs,Ps.&Gbli op Fl. Comp. Gibelli, & Pass. Ces., ex Guss. ,16 N, .W cmd xWld,S.P.e.4 3/3: 4, ed. Pl. Sp. Willd., ex Schmidt W. F. etl xGs. l iu.Sn /:400–401. 2/1: Syn. Sicul. Fl. Guss., ex Bertol. L)Hd. l nl d :32 1798; 342. 3: ed. Angl. Fl. Huds., (L.) L)Lm,F.Franc Fl. Lam., (L.) rn or,F.Fac :34 1852.— 304. 2: France Fl. Godr., & Gren. subsp. eo aot xBis l Orient. Fl. Boiss. ex Lamotte & Lecoq r.Tu. ul eb osir e.2, Ser. Boissier, Herb. Bull. Arv.-Touv., ` 15.305 . p l d ,2 9.1753; 792. 2: 1, ed. Pl. Sp. L., oan,1 et1904, Sept 10 Locarno, .subsp. L. euhieraciodes subsp. s irshieracioides Picris 0 ,7Jl2005, Jul 7 E, hieracioides spinulosa o lcoye BM!), (lectotype: 2 no. , .2 1.1778; 111. 2: ¸. .W cmd .n. s. Schmidt W. F. usn .n. s. Gussone ´ ne ae nHg,Il Fl. Ill. Hegi, in Hayek ´ ´ s 9.16,nom. 1867, 497. es: TA. AOOYO IRSHEAIIE 273 HIERACIOIDES PICRIS OF TAXONOMY AL.: ET K Bro.e Guss.) ex (Bertol. subsp. Slova hnvr .n. s. Chenevard (lectotype: ] ´ Repa & k Hedypnois Bernard setulosa (holo- Crepis Picris ...... 1B 22(2)prcptlm (2.9–)3.1–6.1(–6.7) capitulum, (11–) per Involucral bracts involucral branches. 12–20(–22) Outer brown-black. stem to entire greenish the bracts dis- along peduncle, mostly per few panicles, (1–)1–4(–5) tributed Capitula branch, racemose stem to bracts. per corymbose involucral (1–)2–9(–15) in outer arranged bracts numerous, as peduncle to shape capitulum, similar below of Peduncles bracts black). (0–)1–4(–6) (never grayish with to brownish numerous, or to pale absent bract-like. tomentose, mainly hairs less anchor-shaped reduced, and or 2-furcate more with lanceolate, long, sessile. (0.2–)0.3–5.2(–6.5) leaves and Peduncles smaller stem becoming Stem upwards Uppermost stage. base, petiolate, at fruiting obovate, narrowed to and linear-lanceolate flowering (6–)7–38(–49), leaves the during shriveled yGeirt h ai ebru n1875. in herbarium Paris the to Grenier by h ecito nFec,ietclt h rtlgeand protologue the to an identical “ nomenclature, French, entitled in the description for of the type the stability Regarding (2) the corymbosa, here. of designated Therefore, precise is level. interest epitype the subspecies the the of on purposes in name for the of identified application critically be cannot ogtdnlsrp nuprpr fotrotlglsmostly ligules outermost red acute (6.2–)6.9–12.4(–14.2) Ligules of yellow, present. part ligules linear-lanceolate, upper flowers, in 35–70 strips ca. longitudinal to with Capitula lanceolate apex. at mm, 0.7–1.6(–1.8) ettp ftename the to of lectotype introduced rarely belt, Europe. sub-montane across altitudes) higher the ter- elevations to river lower at lowland (lowlands generally etc., railways, vineyards, roads, orchards, races, along habitats, sites synanthropic abandoned disturbed grasslands, xerothermous long. mm (3.3–)3.4–6.5(–6.9) hairs, deciduous of rows 2 rmr ie.Inrivlca rcs(0)11(1)per (10–)11–15(–16) (5–)6.1–10.9(–11.6) bracts rows, or two involucral in pale Inner arranged capitulum, two lines. mainly in arranged more hairs black)-colored, or numerous, (never grayish to anchor- to and brownish absent 2-furcate rows; hairs with appressed inner pubescent, shaped towards less squarrose, or upright more rows, apex, more usually at becoming irregular acute recurved, to three to obtuse in obovate, to arranged lanceolate mm, (–1.9) esl uiaebtenrb,edn butyit minute a (2.2–)2.3–3.3(–3.6) into beak, abruptly ending ribs, between muricate trans- without versely slightly fusiform, alveolate, black, to brown-black Receptacle Achenes scales. long. mm ligules of (0.3–)0.4–1.3(–1.7) teeth apical long, mm (2.8–)2.8–5.9(–6.7) tube corolla oecaua Notes— Nomenclatural Distribution and Habitat Phenology— irshieracioides Picris atno o.Nt i.Lmada1 0.1844; 303. 1: Lombardia Civ. umbellata Nat. Not. Cattaneo, irshieracioides Picris ...... irscorymbosa Picris nhraimseie nPbastelblwith label the bears P in specimen herbarium an Shak ese lf igeh,Comp. Fingherh., & Bluff ex Nees (Schrank) lwrn rmJn oNovember. to June from Flowering .subsp. L. irshieracioides Picris irshieracioides Picris + morphotype) altitude’ (‘higher subsp. .–.2m.Ppu lms with plumose Pappus mm. 0.6–1.02 o. h pcmnwsdonated was specimen The nob.” 1 h pcmnslce sthe as selected specimen The (1) hieracioides — h useisi on ndry in found is subspecies The umbellata subsp. + morphotype) altitude’ (‘lower 15).–.(33 mm, (1.5–)1.6–3.1(–3.3) .i nopeeand incomplete is L. umbellata Shak e.in Ces. (Schrank) + Shak Ces. (Schrank) (0.7–)0.7–1.6

+ itudinal (0.7–) Picris Picris t Delivered by Publishing Technology to: K. Marhold IP: 78.99.182.222 on: Sun, 11 Mar 2012 10:45:36 Copyright (c) American Society for Plant Taxonomists. All rights reserved. irscrinita Picris irsvillarsii Picris irscrepoides Picris virens Picris paleacea Picris irslongifolia Picris irsgrandiflora Picris irssonchoides Picris irsauriculata Picris 7 YTMTCBTN Vlm 37 [Volume BOTANY SYSTEMATIC 274 nHg,Il l it-u.42 09 99—YE desig- granitico m. 1929.—TYPE: Vogesorum 1039. in Alsatia, [FRANCE] 4/2: here, Mitt.-Eur. nated Fl. Ill. Hegi, in 1863; ya,Cnp l u. 6.1879; 467. Eur.: Fl. Consp. Nyman, etrs n. s. Reuter descente media veg. herbosis Grigna, in La [ITALY] here, designated 1854–1856.—TYPE: Slova ar.Vl.3p 4 .n h. 148 here, p. 3 designated Vill. Gaert. 1753.—TYPE: L. non Dauphine Pl. Hist. nov.; nom. HG.87257) PTP:dsgae here, designated Provence-Alpes-Co EPITYPE: [FRANCE] [ a. MHNGr.1837.27597!), s. Valgaudemard, / Laper o uLuae,gasadna h lieBotanical Alpine the near 45 grassland Garden, Lautaret, du Col 6.17.TP:dsgae ee [AUSTRIA] here, Kitzbu designated 1879.—TYPE: 467. hieracioides a., s. 108388!). Webb Parisiensis], FI-Herb. (lectotype: [Hortus [FRANCE] here, p. designated H. 1829.—TYPE: 399. Paris.): subsp. sheet). the of side [ right a. the on s. plant Holland, in auch / Veit uCriod .Grnm,4Jl1851, Jul 4 Geronimo, 1281 no. S. Espagne, d’ de desig- Cortijo re 1975.—TYPE: Nevada, 248. au Sierra 71: [SPAIN] Soc. here, Linn. nated J. Bot. Sell, P.D. 1852; 2004, ainlpr rn as,lc it rt iTivo near di slopes Prati rocky 42 the dicto trail, on the loco Camella, Pietra Sasso, above located Grand Abruzzo, park ITALY, here, national designated EPITYPE: sheet), illegible, the 2] [label [ / a., Camelo Pietra s. di e Chiarino, di 1] Selve [label [ITALY] here, designated 1882.—TYPE: 418. Ital.: a., s. Carinthia, ex [AUSTRIA] C Rep. ee lbl1] designated [label 1914.—TYPE: 357. here, 2: 3, ed. Schweiz, Fl. Keller, hieracioides ae.F.2 3.18.TP:dsgae here, designated vom 880 1789.—TYPE: 17 Berge, Tegernseer 334. u Miesbach, Wallberghaus 2: Kreis GERMANY, Fl. 1825; Baier. 273. 2: German. Fl. hieracioides ´ n eas n. s. Repa and k irshieracioides Picris Slova ¨ irshieracioides Picris eoes n. s. Tenore ˇ l[] .a., s. [?], hl noye M!). (neotype: solv:62 98—YE eintdhere, designated 1928.—TYPE: 622. eskoslov.: ef,Tb.E Tabl. Desf., paleacea et,Cmt-ed rv o.Halle Soc. Trav. Compt.-Rend. Reut., et yl l o. 8 1824; 78. Nov.: Pl. Syll. Vest, ´ od,Ct rie ad io:2–0 1848, 29–30. Dijon: Jard. Graines Cat. Jord., os.&Ru. uil l f.Br ipn:69. Hispan.: Bor. Afr. Pl. Pugill. Reut., & Boiss. at,Foa(eesug 3 0.1830; 409. 13: (Regensburg) Flora Saut., k subsp. lcoye 74199!). G (lectotype: ircu pyrenaicum hieracium subsp. subsp. c.Bp,Ccoicohc up. o 124. No. Suppl., Cichoriaceotheca Bip., Sch. (SAV!). 02 29 et lr Rgnbr)3 .1820; 6. 3: (Regensburg) Flora Vest, e. yl l l epl:37 1831; 397, Neapol.: Fl. Pl. Syll. Ten., irshieracioides Picris 0 0 ¨ 092 942 ocu lrcu asper oleraceus Sonchus e e ezeg 1Jl1981, Jul 11 Setzberg, den ber lcoye A!lretfamn on fragment largest NAP! (lectotype: ] Vs)Dmn&Pd. Klı Podp., & Domin (Vest) crepoides atrs n. s. Sauter 00 00 ´ grandiflora sonchoides lcoye G-Boiss!). (lectotype: (SAV!). ,13 N, ´ ,06 N, :18 79 o.ilg At 53), (Art. illeg. nom. 1789, 148. 3: oeBt d Ct l ot Reg. Hort. Pl. (Cat. 3 ed. Bot. cole subsp. subsp. Villars 15D.Sp n Sup. DC. 1145 Su. ya,Cnp l Eur.: Fl. Consp. Nyman, (Saut.) enoo umbellatus Leontodon 33 24 lcoye W!). (lectotype: 0 auriculata ufns n. s. Wulfen 825 0 ˆ Vs)Tel nShn & Schinz in Thell. (Vest) ? Tn)Acn. op Fl. Comp. Arcang., (Ten.) 239 ´ ets n. s. Vest edAu,muti pass mountain d’Azur, te longifolia u odlo u 1854, Aug Mondello, sur .n. s. 00 subsp. 00 ,144m u 2005 Jul 9 m, 1,434 E, .Wl./h en h. / Wild. y. xhr.Desfontaines herb. ex ,207m 5Aug 15 m, 2,067 E, irspyrenaica Picris lcoye GRM (lectotype: ] ´ /[label2]zuSt. insous-alpine, gion Sh i. Hayek Bip.) (Sch. noye TUB!). (neotype: irshieracioides Picris Bis Reut.) & (Boiss. lcoye W!, (lectotype: ] 94 / 2974a irspyrenaica Picris villarsii oreuPl. Bourgeau .Lippert W. ´ Schrank, ´ c ˇ tuberosa ... . 111. r.: (Jord.) Picris Picris Kve Picris infra Vill., ˇ t. irssenecioides Picris rcs(–81(2)prcptlm (2.7–)3.1–6.7(–7.3) capitulum, b per branches. to (8–)8–18(–20) stem bracts of greenish third or bracts half Involucral upper in pedun- mostly per distributed (1–)1–4(–4) cle, branch, stem per corymbose (1–)1–11(–15) in arranged involu-panicles, numerous, outer to few as Capitula shape bracts. cral similar of bracts peduncle below capitulum, bracts (1–)1–6(–9) numerous, with Peduncles to black-colored. absent to pale hairs anchor-shaped and 2-furcate with (0.2–)0.3–7.6(–10. Peduncles bract- like. often reduced, becoming lanceolate, amplexicaul, leaves upwards stem to Uppermost semi-amplexicaul auriculate. base, sessile, at and narrowed smaller and to petiolate, linear-lanceolate flowering margin. obovate, (4–)5–20(–23), the leaves at Stem during stage. undulate persisting fruiting rarely mostly to leaves or entire Rosette petiolate, crenate, leaves apex, dentate, at to Basal acute shallowly to ovate soft. obtuse linear-lanceolate, shape, rather in rosettes, anchor- obovate (1–)2–12(–16). hairs, in and rigid branches arranged 2-furcate numerous, simple of with and number hairy shaped, stem, densely half the usually upper Leaves of in third usually moderately distributed or to branches sparsely and stem Stem size branched; upwards. hair numerous hairy, decreasing to to densely density green to few grooved, moderately longitudinally bearing reddish-brown, slightly usually Stem taproot, roots. lateral simple a with rielii Picris tatrae Picris irsrielii Picris oeo espbset ih2fraeadanchor-shaped and 2-furcate with pubescent, less to appressed or squarrose, apex, more usually rows; rows, at inner towards upright acute irregular more becoming recurved, to three obtuse in obovate, arranged to lanceolate mm, 1.6(–2) irsmonticola Picris irstatrae Picris irsoligocephala Picris inilt hr-ie eena,1–0 mhg plants high cm 15–100 perennial, short-lived to Biennial 2.12.TP:[RNE.Grdsd Lourdens, de Garides Montme [FRANCE]. sur 1926.—TYPE: 325. o.ivl,nm nud. nom. inval., nom. nud. nom. inval., sheet). the of side left the on plant 138382!, MA totype: oeek l.3,500, alt. Hoheneck, edge lvae saa,mre,120130m, 1,220–1,320 marges, 1925, Sept Estavar, 2 Aug, et 30 Llivia [SPAIN] Cerdagne: here, designated 1987.—TYPE: 9. 14: Fontqueria hieracioides Picris 74201!). G (holotype: 1892, an,e lata allaut en valle bains, Mont-Dore, [FRANCE] here, designated TYPE: LW!). type: Kve hieracioides CLF!). type: YE eintdhr,[OAI]I aci subalpinis pascuis Transsilv. In [ROMANIA] here, designated TYPE: n. s. Martin ee SOAI]i liu acri Ta calcareis alpibus in [Tatranska Barlangliget [SLOVAKIA] here, ˇ .Rp C Rep. t. Borba enn ul o.Bt rne7 12) 5.1927, 656. (1926): 73 France Bot. Soc. Bull. Sennen, enn o.Sc br i a.2 1) 7.1929; 170. (11): 28 Nat. Ci. Iber. Soc. Bol. Sennen, ob,Terme Borb., ob,Mg.Bt ao :38 1902; 318. 1: Lapok Bot. Magy. Borb., ... ´ aot,Por l lt et.2 5.1881.— 453. 2: Centr. Plat. Fl. Prodr. Lamotte, ss.n. lcoye P!). (lectotype: subsp. ´ eued ul o.Bt Gene Bot. Soc. Bull. Beauverd, in ore,1 et1926, Sept 17 Bourges, lian, ˇ itaMr,Kl,Jl .a., s. Jul, Kalk, Mare, Piatra cu,Eu.P.Tasiv:31 1866.— 361. Transsilv.: Pl. Enum. Schur, solv:62 98—YE designated 1928.—TYPE: 622. eskoslov.: ` lcoye P202560!). BP (lectotype: subsp. ... tatrae 0 ´ Sennen 8Jl14,[ 1849, Jul 18 , ztu.Ko szettud. 0Ag16 foeigspecimen], [flowering 1862 Aug 10 ´ oln] 3,000 Kotlina], (Borba rielii )ln,mr rls tomentose, less or more long, 1) onbak ue involucral Outer rown-black. , lne ’Epge5353 Espagne d’ Plantes ´ Sne)Bolo (Sennen) )Dmn&Pd. Klı Podp., & Domin s) ¨ l 6 9.19,nom. 1894, 498. 26: zl. aot .n. s. Lamotte 0 –4,000, cu .n. s. Schur ` eueds n. s. Beauverd eSr ,18: 2, Ser. ve ´ Vigo, & s + 0 ´ ,2 Jul 24 m, (0.6–)0.6– read trae (lecto- ] (lecto- Picris ´ (lec- des ´ c ˇ Delivered by Publishing Technology to: K. Marhold IP: 78.99.182.222 on: Sun, 11 Mar 2012 10:45:36 Copyright (c) American Society for Plant Taxonomists. All rights reserved. iie,abtns h eragdVlas ebru in herbarium de Villars’ Artus rearranged by who written botanist however, a above is, to the Miribel, which belong bears specimen not label, second does 1811 mentioned The and in material. Switzerland collected original was in the hand, Pass Villars’ Simplon in the label in a with name them, the of bear that herbarium Villars’ pyrenaicum Hieracium .(ines15) hr saohrltrhmnmof homonym later another is by There occupied 1753). P already was (Linnaeus epithet L. the pyrenaica as of there C. up species taken a be to homonym related earlier is its that wrote (1975) ee 5 eern otename the to Referring designated is (5) epitype appropriate here. an the specimen, of this lectotypification by the from name apart Therefore, interpretation name. be the the cannot of of specimen purpose the the illegible. for damage, is identified the locality unequivocally of one extent sheet, the this of on the Because labels of two the on material localities on original fragments the plant name representing damaged sheet badly herbarium (4) several protologue. in the are Vest in There by mentioned that collected neighboring material region the the from selected was neotype ilcletdb etfo h oaiymnindi the in mentioned locality mate- the or from material of Vest original protologue by dated No collected to referred (3) rial Vest protologue. that Wulfen’s the one to the in apparently belonged some was it of and undated, leaf herbarium is a specimen apparently the is Although one to third belong the them of name the the Two on of Vest. fragments lectotype plant selected three the are with mentioned sheet There locality (2) the protologue. from the neotype name in a this Therefore, for deposited. here specimens be designated is to herbarium known where are M, Schrank by nor comm.) pers. Stoffelen name lyi rw thge liue mnaet liebelt, alpine to (montane moun- European altitudes ranges. across tain altitudes) higher lower to forest at introduced Gener- rarely mountains. grows along in it habitats valleys, secondary ally non-forest other mountain in in and in also roads penetrating habitats shrubs, synanthropic commu- deciduous subalpine of belt; grass subalpine nities and tall slopes supramontane warm belt, the and alpine of dry to relatively on montane communities the species-rich in communities herb long. mm (4.1–)4.2–7.7(–8.2) hairs, deciduous of rows 2 esl uiaebtenrb,edn butyit minute a into trans- abruptly (2.5–)2.5–3.7(–4.1) slightly ending beak, fusiform, ribs, scales. between black-brown, without muricate to alveolate, versely Receptacle red-brown long. Achenes (0.3–)0.4– ligules mm of teeth 1.7(–2.2) apical long, mm (2.5–)2.8–6.7(–9.7) 02 SLOVA rows, two in arranged capitulum, (6.9–)7.4–12.7(–13.7) per to (9–)10–15(–16) grayish t brownish, bracts two pale, in hairs arranged numerous, black-colored, to absent hairs 2012] iue elw e ogtdnlsrpi pe atof part Ligules upper present. rarely in missing, flowers, strip (6.9–)7.4–15.8(–19.9) mostly 30–60 longitudinal ligules ca red with outermost Capitula yellow, apex. ligules at acute lanceolate, . oecaua Notes— Nomenclatural Distribution— and Habitat Phenology— pyrenaica enoo umbellatus Leontodon irsgrandiflora Picris il,namely Vill., L)Getr ae on based Greuter, (L.) irspaleacea Picris lwrn rmMyt October. to May from Flowering

+ + 07).–.(2 m acoaet linear- to lanceolate mm, (0.7–)0.7–1.8(–2) 16).–.(40 m ool tube corolla mm, (1.6–)1.8–3.5(–4.0) .()Teeaetoseiesin specimens two are There (6) L.

e.Teeaeas he different three also are There Ten. + .–.3m.Ppu lms with plumose Pappus mm. 0.6–1.03 .pyrenaica P. 1 ooiia aeilo the of material original No (1) Crepis crn a on nB (Piet BR in found was Schrank eti vial.Teeoe a Therefore, available. is Vest .hieracioides P. h useisi on ntall in found is subspecies The oelns ne involucral Inner lines. more o u oe hti cannot it that noted but , irspyrenaica Picris L)Get. ae on based Gaertn., (L.) ircu pyrenaicum Hieracium P ´ P TA. AOOYO IRSHEAIIE 275 HIERACIOIDES PICRIS OF TAXONOMY AL.: ET K . . pyrenaica pyrenaica subsp. Sonchus irssonchoides Picris il,Lack Vill., h first The . umbellata .(1753) L. species. , sdsgae eea h lectotype. the the to as here herbarium belongs designated Boissier’s collection is from specimen this the and that material indicates original This Reut.!.” in et “ were name which protologue) the locality, under exsiccate type the as the distributed at 1851 in specimens in are Bourgeau to there by collected Nevertheless, referred G. in (as herbarium Boissier’s Reuter no is by There collected (7) as chosen. identified specimen is a relevant as epitype here of appropriate designated application is an precise specimen lectotype, the this as for Therefore, identified name. be the cannot it ambiguous, second demonstrably such, this is as and addition, damaged In badly comm.). is specimen pers. Poncet (Vincent 1827 ac ntefedadt h aet fM .frtergnrlsupport. reviewers. general their anonymous for two Dus S. of M. to of comments ofthesamplesitesandtoZlataKoma go parents valuable thanks the appreciate Special to and also assis- their field We for the (Jaca) Villar in Luis tance and (Bratislava), Repa Peter (Bratislava), ple isses 2005. Biosystems. Applied several in dispersal seed and morphology Diaspore 2004. 1993. Theurillat. C. P. M. J. Andersen, and Moser, M. D. Lauber, K. D., Aeschimann, the by to supported LPP-0239-09 was no. (project study Marhold). Agency Karol This Support Development characters. and morphological Research the of ation Iva (Grenoble), Douzet Rolland (Palermo), Hoda Domina Gianniantonio vno . .Rgat n .Gue.20.Dtcigtenme of number the Detecting 2005. Goudet. J. and Regnaut, S. G., Evanno, hih .20.ALDT olcino ucin o convenient for functions R of collection a AFLPDAT: 2006. D. Ehrich, Bolo u Studien Zytologische 1935. B. Bergman, and Fischer, A. M. Schneeweiss, M. G. Albach, C. D. E., K. Bardy, Dosta analysis Multivariate Scho P. 2007. J., Knox. Ch. Dixon, B. E. 1994. and Ackerly. Balkwill, D. K. D. V., G. and Cron, McConnaughay, M. D. K. S., J. Coleman, 2000. A. S. Chaudhary, The 2000. Elven. R. and K. A. Brysting, and Brochmann, C. Pompanon, F. Eidesen, B. P. Bellemain, E. A., Bonin, ad,K . .Scho P. E., K. Bardy, Acknowledgments. ` ,O n .Vg.1995. Vigo. J. and O. s, ´ iddsesdAsteraceae. wind-dispersed Alpina lseso niiul sn h otaeSRCUE simulation a STRUCTURE: study. software the using individuals of clusters adigo FPdata. AFLP of handling SPN. Bratislava: vnk oaik Foreningen Botaniska Svenska Gattungen Europe. southeastern in Scho P. Biosystems. Applied irtosi uoenhg onanpat( plant mountain high Primulaceae). European a Pleistocene in long-distance migrations supports testing hypothesis Bayesian 2009. in variation Senecioneae). morphological of lution plants. in variation phenotypic Interpreting Water. and Agriculture of istry of revision 189–216. morphological taxonomic errors 49: of a genotyping and analyses variation assess numerical (Caryophyllaceae): and complex track to studies. How genetic population 2004. in Taberlet. P. Barcelona boundaries species blurs flow gene Extensive 2011. among Albach. C. D. eeisadEvolution ( Europe. and southeastern herb genetics in woodland s.l.) a Plantaginaceae of group, taxonomy and evolution ´ lova ,J n .C M. and J. l, ´ Bailv) ioeoPsaaqa(oez) Maria (Cosenza), Passalacqua Nicodemo (Bratislava), :187–191. 9: en ttgr,We:HutVerlag. 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G. nswetter, lr fKndmo ad rba2/3 Arabia Saudi of Kingdom of Flora ˇ ewudlk otakFboCni(Barisciano), Conti Fabio thank to like would We 7 771–786. 57: und nSno(rtsaa o rprn h map the preparing for (Bratislava) Senko an ieaueCited Literature AFLP oeua clg Notes Ecology Molecular 4 2611–2620. 14: Taxon irs vnkBtns isrf tivnaf Utgifven Tidskrift Botanisk Svensk Picris. mrcnJunlo Botany of Journal American 9 155–301. 29: Ò Vel oeua Ecology Molecular lr auldl Paı dels manual Flora 0 108–121. 60: ln apn:Protocol mapping: plant ˇ ky irslongifolia Picris ´ kl ieai deltoidea Cineraria ´ ˇ ¨ rova u and ´ e i otfazn e den bei Fortpflanzung die ber eatu liu–.arcticum alpinum–C. Cerastium c ˇ .arcticum C. aurc na ´ .spicata V. o rprn h illustr- the preparing for irslongifolia Picris rnsi clg Evo- & Ecology in Trends ˇ vnevys ovanie :603–604. 6: 3 3261–3273. 13: eoiachamaedrys Veronica nrsc vitaliana Androsace 3 580–591. 53: 5:497–521. 154: os.&Reut. & Boiss. ag s.l. 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Sell, S Vest, 03 pnst h yelclt rabodrae pcfe ntepooou of corre- protologue the sampled in specified locality names: Super- area broader Slovakia. following the a – the or that Republic, locality SK type Kyrgyz indicate Slovenia, the – to – codes KG sponds SI Japan, populational – Romania, in JP Spain, – Italy, scripts – RO Country – ES Montenegro, IT Germany, population. – Croatia, – – ME DE per HR Austria, France, markers – – AT FR AFLP Andorra, – private/rare AD abbreviations: of number the ape htwr nlddi Slova in included were that samples umbellata P. /,003,0/8; 0.0232, 5/5, Slova nlsstenme fAL eoye,Nisgn iest (D AFLP diversity for gene used Nei’s plants genotypes, the AFLP of of for latitude, number used number altitude, the plants analyses/the locality, analyses, field-collected/cultivated bold), morphometric the (in of the code number populational the presented of are longitude, Data order east. the the to west in the from geographically arranged are Hornes, M. 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Saut., Picris Picris * Nei and 0 9 N, ), – ,1 ,42 m, 1,510 LEI ir,90m 49 m, 900 Diery, 48 m, 224 5 ,47 m, 550 19.434 40.691 12.620 03.789 52.194 54.984 25.081 31.400 45.638 47 rnoio ,6 ,45 m, 1,162 Brentonico, 47 Wu el atcs,156m 42 m, 1,576 Panticosa, la de 38 ar,Fac Fatra, ar,Ja Tatry, 6 ,49 m, 468 45 4 ,49 m, 643 an,170m 45 m, 1,700 Bains, CAN 47 49 le,l iie ,5 ,45 m, 1,556 Rivine, la Alpes, 1 ,38 m, 612 48 0/4; 0/12; oteyade rl,70m 41 m, 740 Brull, el and Montseny BAN ,4 ,42 m, 1,640 T Niedero AT, t cneeg 4 ,47 m, 543 Schneeberg, Mt. le,Uie 5 ,45 m, 552 Ugine, Alpes, 25 18 17 06 01 HOL SCHB Kitzbu Comte 1/8; VAL1 0/6; /,008,0/11; 0.0285, 4/4, FUG BRI 8 /,007,0/9; 0.0477, 4/4, 18, -; -, -, 20/15, 0/10; 0.0572, 3/3, 16, -; -, -, 17, 2/11; 0.0418, 5/5, 29, iTv,144m 42 m, 1,434 Tivo, di Spis Comte ,- -; -, -, ZDI .42 1/14; 0.0412, /,001,1/13; 0.0516, 3/3, 15,-,-,-; T buz,Vled ev oaa ,7 ,42 m, 1,575 Romana, Selva di Valle Abruzzo, IT, /,004,0/9; 0.0545, 4/4, ,- ,-; -, -, 2, LIM 08 /,006,0/11; 0.0365, 5/5, 20/8, 20 20 ¨ irshieracioides Picris ˇ S aaoi,Ta Catalonia, ES, , tebr,Hiebr-clebc,10m 49 m, 100 Heidelberg-Schlierbach, rttemberg, 52.624 38.319 8 32.187 06.938 ske * 34.636 01.921 02.092 37.261 03.231 22.853 14.991 05.416 T imne ioet,160m a44 ca m, 1,600 Limonetto, Piemonte, IT, , 05.181 ORM R Provence-Alpes-Co FR, , LLV T rnioAt dg,PsoPaodleFgze ,5 ,45 m, 1,156 Fugazze, delle Piano Passo Adige, Trentino-Alto IT, , 54.537 2 CDL O ueor,Ba Hunedoara, RO, , S Arago ES, , ,SK,BelianskeTatry,Z ¨ ´ ´ T azug olrbc mPngu 9 ,47 m, 898 Pinzgau, im Hollersbach Salzburg, AT, , 0 8 0 0 0 0 0 0 0 0 * MAR BRE ´ ,2,55 .61 2/20; 0.0631, 5/5, 20, E, otd-od,34m 47 m, 384 Pont-de-Roide, , osdAot ,0 ,46 m, 1,102 Bois-d’Amont, , E,20,-,-,-;E,15,-,-,-; e,90m 47 m, 930 hel, ,15 -; -, N, -, 13, E, 0/9; 0.0498, 3/3, 15, E, ,2/,44 .29 0/6; 0.0259, 4/4, 20/4, E, ,20 N, 11 N, ,FR,Rho ,SK,Slovensky ´ sadln aly 7 ,49 m, 771 valley, dolina nska Hanus VIE ˇ 6 0 0 kovske 8 0 0 S aaoi,Llı Catalonia, ES, , 0 GRA ,19 N, T imne re,70m 44 m, 750 Ormea, Piemonte, IT, , 0 0 0 0 0 0 ,- /,000,0/3; 0.0109, 4/4, -, E, 0 0 ,11 N, ,13 N, 57.112 R Provence-Alpes-Co FR, , ,19 N, T imne ri,79m 45 m, 799 Breia, Piemonte, IT, , ,1,- ,-. -, -, 19, E, ,06 N, E,20,-,-,-; ,4/2 ,- -; -, -, 40/12, E, ,2,54 .42 1/11; 0.0412, 5/4, 20, E, 1/18; 0.0613, 7/7, 13, E, ,2/,55 .41 2/13; 0.0451, 5/5, 20/4, E, 0 ,06 N, 23.341 16.894 ¨ ,08 N, * 53.600 trec,M.O Mt. sterreich, LAU 29.915 39.766 S aaoi,Veh,120m 42 m, 1,230 Vielha, Catalonia, ES, , MUN ,SK,Male 8.514 11.145 ˇ KLAK 23.476 ve 1 ,49 m, 616 ovce, 26.399 E aen rsc,78m 47 m, 768 Graseck, Bayern, DE, , ˆ ´ ´ 0 eAps alie ,0 ,45 m, 1,204 Valloire, ne-Alpes, 12.156 ,Candanchu n, ,13 N, 8 0 0 el adeak ,2 ,48 m, 1,220 saddleback, sedlo 42.499 0 14.543 ,20 N, ,17 N, 0 R Provence-Alpes-Co FR, , 16.501 GAT SOL 24.239 ,11 N, 0 ,19 N, ,FR,Alsace,Munster,853m,47 0 09.700 ,02 N, ,00 N, 38.356 MTC TIM ANN 0 0 00.573 SAG VOG ,1,33 .49 0/13; 0.0479, 3/3, 15, E, ,SK,Vel E,18,-,-,-; 15.525 0 ´ subsp. 29.942 ,2/3 ,- -; -, -, 20/13, E, rg,c 6 ,c 41 ca m, 460 ca rrega, ´ 0 FAL ´ D nor,Sle,190m 42 m, 1,900 Soldeu, Andorra, AD, , E,20,-,-,-; T imne aiaa 8 ,45 m, 289 Gatinara, Piemonte, IT, , 0 43.170 0 apt,Za Karpaty, ESP 0 ,9 /,- 0/3; -, 2/2, 9, E, ,RO,Bras TIT ,2,- ,-; -, -, 20, E, 0 7 E,20,-,-,-; 28.200 0 25.325 ,2/,77 .59 0/13; 0.0579, 7/7, 22/8, E, GER 03.594 a,Sucha Raj, 0 T iiy aem,M.MneCci,61m, 611 Cuccio, Monte Mt. Palermo, Sicily, IT, , ,1,33 .41 0/7; 0.0461, 3/3, 19, E, 7 5 50.600 00.815 ,1,33 .37 0/8; 0.0387, 3/3, 16, E, 06.509 1 ,IT,Sicily,Sagana,637m,38 ˜ T Niedero AT, , 47.963 0 ¨ ,FR,Alsace,Linthal,944m,47 23.055 9 0 ,06 N, ia 4 ,45 m, 540 nita, ´ 45.029 45.503 ,19 N, 0 E aen al 8 ,47 m, 780 Fall, Bayern, DE, , shr a100m a47 ca m, 1,000 ca tscher, i,110m 42 m, 1,170 via, E Baden-Wu DE, , S aaoi,Ept ,7 ,42 m, 1,576 Espot, Catalonia, ES, , ˆ ˇ ,13 N, FIN 04.138 hieracioides SGA edAu,Brianc d’Azur, te ˇ 44.225 irk oiavle,80m 49 m, 890 valley, dolina diarska ´ 0 * ka E,20,4/4,0.0327,0/7; ,1,555m,42 ,SK,Slovensky 0 0 0 DON ´ ACHE ,12 N, ,2/7 ,- -; -, -, 20/17, E, ,1,77 .79 0/17; 0.0769, 7/7, 18, E, EST 0 0 0 8 0 ,1,33 .21 0/7; 0.0221, 3/3, 17, E, ,2/,22 ,1/13; -, 2/2, 20/8, E, ,1,- ,-; -, -, 19, W, 0 ,10 N, R Franche-Comte FR, , ar,M.Kl Mt. Fatra, 0 E,20,5/1,0,0/7; 0 VLK T rnioAt dg,SnGaoo– Giacomo San Adige, Trentino-Alto IT, , ,00 N, 46.702 ,06 N, 19.524 0 ˆ 28.179 22.612 CEK ,06 N, ´ ´ 28.738 edAu,Cld atrt ,6 m, 2,067 Lautaret, du Col d’Azur, te 0 o,Bras ¸ov, 33.825 horska ,15 N, 6 40.234 * R ageo-osiln Estavar, Languedoc-Roussillon, FR, , STU MAD TAC Bela ,SK,Nı T io,Ahnice,90m, 940 Achenkirchen, Tirol, AT, , LUC * * ¨ ,SK,Vel 23.807 MONT 05.480 0 0 trec,Anbr,59m 47 m, 519 Annaberg, sterreich, 14.557 K inn,C Pieniny, SK, , 0 0 ,02 N, ´ ,20 N, ‘oe liue morphotype)— altitude’ (‘lower 54.831 SCHN E,20,-,-,-; ,06 N, 0 FEL * 25.178 0 ´ ,06 N, 28.304 7 ,2,55 .42 1/12; 0.0412, 5/5, 25, E, 0 ,SK,Male ˆ 09.296 ,2/3 /,009,1/15; 0.0491, 5/5, 20/13, E, T iiy Terminy-Caccamo, Sicily, IT, , * * og,92m 48 m, 942 gorge, ytia 2 ,48 m, 220 Bystrica, 44.086 edAu,L Mone Le d’Azur, te ,SK,Za ,SK,Choc o,60m 45 m, 600 ¸ov, ¨ ´ 26.633 47.401 BAL 12 T oaleg Feldkirch, Vorarlberg, AT, , k ar,Dnvl,87m, 867 Donovaly, Tatry, zke tebr,Ttse 5 m, 853 Titisee, rttemberg, 28.387 ˇ o,138m 44 m, 1,308 ¸on, ´ 0 45.897 0 REFU k ,9 ,48 m, 1,290 ak, ,2,54 .45 0/13; 0.0405, 5/4, 20, E, 0 ,19 N, 39 0 ˇ ,2/,44 .64 0/17; 0.0654, 4/4, 20/6, W, S aaoi,between Catalonia, ES, , 0 0 SPH ,1,44 .77 1/19; 0.0747, 4/4, 15, E, 23.780 ka 5 ,2/,55 .64 1/17; 0.0604, 5/5, 20/4, E, ,07 N, a,Grv,80m 48 m, 860 Geravy, Raj, 0 46.015 0 1/9; 0.0572, 3/3, 15, E, T Niedero AT, , 05.181 0 20.937 ´ ,07 N, ,01 N, 41.190 0 S Arago ES, , 10.000 58.660 ,2/2 ,- -; -, -, 20/12, E, 28.217 24.470 ar,Vlkolı Fatra, 07.632 0 54.537 0 ,01 N, 0 ,00 N, ´ 0 3 * PDR ,23 N, PIZ ,08 N, ´ padne T buz,Prati Abruzzo, IT, , VAL2 ´ ,SK,Spis BDA 52 0 HEI 40.505 e is 1 m, 911 Fins, Les , ˇ ,03 /,0.0277, 3/3, 0/3, E, 0 apt,Stupava, Karpaty, UGI 34 0 JAN ske 0 ,1,44 0.0353, 4/4, 15, E, ,2,44 0.0467, 4/4, 20, E, 0 ,2/1 ,- -; -, -, 20/11, E, 0 55.431 7 0 FAC 0 ,00 N, ˇ ,15 N, 0 0 0 KIT ,06 N, ,18 N, ,IT,Sicily,Piano 0 8 ,11 N, 12 08 N, 0 ,14 N, ´ erveny ,2/5 /,-, 2/2, 20/25, E, ,07 N, R Franche- FR, , ´ E Baden- DE, , 8 56.531 31.532 8 rh,Lu vrchy, 8 0 R Franche- FR, , * 04.756 34.000 ,1/,5/5, 19/5, E, 36.520 R Rho FR, , 56.378 ´ ,FR,Rho 0 34.170 ar,Mac Tatry, 11.951 ,SK,Nı 18.300 E,20,-,-,-; * 2 16.48 ,Balneario n, T Tirol, AT, , ´ ,SK,Vel 0 ˇ e,56m, 586 nec, ,2,- ,-; -, -, 20, E, ske ¨ ´ 80 47.241 39.516 55.184 33.547 58.466 14.086 sterreich, 56.081 16.099 25.600 37.063 07.114 44.554 07.381 09.700 0 ˆ ´ ,20/5, E, 0 Kla tier-les- 0 0 0 20/ W, 0 0 0 0 ,0/9, E, ,13 N, ,12 N, ,11 N, 0 0 ,07 N, ,08 N, ,01 N, vrchy, VSR ,20, E, ,17, E, OTS ´ ´ s ˇ ˆ ˆ ´ c ˇ zke 0 tor, ne- ne- ˇ 0 0 0 0 0 ky, 0 0 W, ˇ 0 0 0 0 0 0 N, N, N, N, N, N, N, ka E, E, E, E, E, E, ie 3 ´ , , Delivered by Publishing Technology to: K. Marhold IP: 78.99.182.222 on: Sun, 11 Mar 2012 10:45:36 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 7 YTMTCBTN Vlm 37 [Volume 38.423 083 33 48.329 BOTANY SYSTEMATIC 32.706 20.341 02.172 37 ca m, 1,100 ca Zucchi, 278 z ,6 ,42 m, 1,566 42 6m,42 Lamac pnt ac,122m 39 m, 1,292 Manco, Spineto oa 6 ,46 m, 864 Gora, ’qia 2 ,42 m, 421 L’Aquila, R Dubrovac HR, 48 z VRA 46 0/11; 0.0479, 3/3, t oln,140m 39 m, 1,450 Pollino, Mt. /,002,0/11; 0.0424, 3/3, aara t ua a190m 39 m, 1,900 ca Mula, Mt. Calabria, 2/8; 13 CRO uao 5 ,40 m, 750 Lucano, /,002,0/5; 0.0424, 3/3, T ruiVnzaGui,Mdii,1 ,45 m, 10 Madrisio, Giulia, Friuli-Venezia IT, /,006,0/4; 0.0363, 4/4, R aasaz Zadarska HR, z ˇ ˇ ˇ pnj,Iae,22m 46 m, 232 Ivanec, upanija, pnj,Kk absa ,45 m, 4 Valbiska, Krk, upanija, pnj,na uolv 8 ,45 m, 386 Lupoglav, near upanija, 09.206 09.390 32.877 28.759 FRA T buz,CmoFlc,150m 42 m, 1,570 Felice, Campo Abruzzo, IT, , R rmrk-oasaz Primorsko-goranska HR, , ˇ 0 0 0 0 0 ,202m,48 0 E,20,-,-,-; ,13 N, 0/10; 0.0384, 4/4, 14, E, ,14 N, 0/8; 0.0332, 3/3, 20, E, ,1,- ,-; -, -, 15, E, T aara rsieo 5 ,39 m, 453 Frascineto, Calabria, IT, , 27.489 0 0 0 ,16 N, 0 ,15 N, ,1,- ,-; -, -, 14, E, ,13 N, 10.752 36.341 ˇ 25.265 onrtasaz ko-neretvanska ˇ 0 pnj,Prvc 4m 43 m, 24 Pirovac, upanija, ,14 N, 57.471 KRA CON RIM 52.440 11.951 PIC 28.324 15.573 0 21.718 0 ,14 N, E,20,-,-,-; LAM RET 0 ,1,22 ,1/9; -, 2/2, 15, E, R Krapinsko-zagorska z HR, , T buz,Percml,78m 42 m, 778 Pietracamela, Abruzzo, IT, , T buz,Frae 2 ,42 m, 620 Fornace, Abruzzo, IT, , 12.103 T aara oez,25m 39 m, 225 Cosenza, Calabria, IT, , 0 12.596 E,20,-,-,-; 0 0 53.605 ,2,44 .75 0/19; 0.0705, 4/4, 20, E, 0 STC U Feje HU, , 0 ,17 N, 0 ,15 N, ,13 N, * ,1,33 .78 0/18; 0.0738, 3/3, 11, E, 54 ,SK,Podunajska 06.740 0 0 ,13 N, 21.404 SOL T buz,SnClmo ,0 ,42 m, 1,100 Colombo, San Abruzzo, IT, , 13.230 ,13 N, CAM 0 E,20/7,-,-,-; 0 ˇ ,16 N, pnj,Ploc upanija, 02.672 27.420 BLH 0 47.105 ,2,- ,-; -, -, 20, E, T Niedero AT, , 01.673 ´ ˇ 0 T buz,AsriCmo ,3 m, 1,236 Campo, Assergi Abruzzo, IT, , 0 ,Re r, KRA ,16 N, pnj,Ce,Vaa 5 ,44 m, 158 Vrana, Cres, upanija, ,16 N, 59.3 45.000 T buz,Mjla lc House, Block Majella, Abruzzo, IT, , 24.256 0 04.732 0 E,15,-,-,-; E,13,2/2,-,0/6; 19.305 0 ´ 0 E,19,-,-,-; sia,16m 46 m, 106 tszilas, 0 0/10; 0.0479, 6/6, 0/21, E, ,SI,Gorenjskaz ,14 N, VAB 0 13.718 ,16 N, ´ 07.120 0 0 ,2/1 /,005,1/16; 0.0551, 5/5, 20/11, E, ˇ 49.340 E,20,-,-,-; 0 ,1 ,43 m, 17 e, nı ,14 N, MUR R Primorsko-goranska HR, , PES ´ ¨ 51.694 49.899 z ˇ 13.928 trec,Slea,36m, 346 Sollenau, sterreich, n olns Bratislava- lowlands, ina 29.877 ˇ 0 pnj,Kaia 2 m, 224 Krapina, upanija, 0 ,1,- ,-; -, -, 15, E, ,6 /,- 0/6; -, 2/2, 6, E, 01.000 IVA CUN T buz,Pescara, Abruzzo, IT, , 0 ,15 N, T aiiaa Muro Basilicata, IT, , 0 AQU 0 0 ,12 N, 09.420 ,16 N, ,13 N, LUP 0 ,HR,Varaz ,5 ,- -; -, -, 5, E, * POL ˇ 0 SPI pna Kranjska upanja, K Podunajska SK, , ,- /,0.0305, 5/5, -, E, ,IT,Abruzzo, 40.140 R Istarska HR, , 0 00.937 03.944 10.340 ,IT,Calabria, 48.734 ,1,- ,-; -, -, 14, E, ,IT,Calabria, 15.305 59.000 24.646 MULA 32.323 0 ,20/1, E, ˇ 0 0 0 0 ,17 N, ,13 N, ,16 N, ,18 N, dinska PIRO MDR 0 0 0 ,17, E, ,15, E, PLM ,20, E, ,IT, 0 N, ´ , , , as 4 ,48 m, 544 pass, nı ervu 6 ,48 m, 162 Bebravou, oiavle,67m 49 m, 677 valley, dolina 31.935 56.341 50.071 39.169 09.089 29.422 1 ,45 m, 415 io,Oae n Chis and Oradea Bihor, Lopenı iaai 3 ,40 m, 339 Hirataki, z Podunajska rh,M.Va Mt. vrchy, Cerova okio tuagn tuamr,Mri 6m 43 m, 66 Morai, Atsuta-mura, Atsuta-gun, Hokkaido, 20 18 17 K Podunajska SK, 18 01,- ,-; -, -, 20/18, -; -, -, 20, os) ,0 ,41 m, 2,800 Toosu), LPT K Podunajska SK, .28 0/8; 0.0258, ˇ pnj,Dbonk 4 ,42 m, 145 Dubrovnik, upanija, ´ z ˇ irshispidissima— Picris irsnuristanica— Picris irsjaponica— Picris n olns C lowlands, ina 39.246 20.932 05.556 * 57.973 K Popradska SK, , 0 0 0 ´ 0 0 ´ 0 ,18 N, -; -, -, 20/12, E, E,20,-,-,-; ,23 0/9; N, 0.0465, 5/5, 20/5, E, ,01,31 .08 0/13. 0.0018, 3/1, 0/14, E, c ˇ rhvn,Ha vrchovina, k 3 ,48 m, 337 ek, 0 VRS 0 0 ,01,- ,-; -, -, 0/10, E, ´ 0 ,- /,002,0/18; 0.0628, 4/4, -, E, ,2/,44 .87 1/19; 0.0887, 4/4, 20/8, E, 49.935 ,- /,002,0/22; 0.0420, 4/4, -, E, nı ´ VRT ´ pec DEV z 01.951 ˇ 49.053 ´ ´ * n olns Koma lowlands, ina K il apt,M.Vrs Mt. Karpaty, Biele SK, , nı ˇ nı ,543m,48 ´ 0 ´ * 37.053 ˇ ,25 N, z z ˇ ˇ ,SK,Kova nv,19m 48 m, 119 unovo, O ueor,Dv,20m 45 m, 220 Deva, Hunedoara, RO, , BANB n olns Va lowlands, ina n olns Bratislava-Petrz lowlands, ina ´ 0 0 ´ 15.450 ,1,44 .46 0/12; 0.0446, 4/4, 19, E, ,2,- ,-; -, -, 20, E, JP106 58.660 PIL oln ai,Liptovsky basin, kotlina jnac 22.387 0 NUR CHLA lz 8 ,47 m, 180 ¸laz, ,20 N, ˇ * 01.951 BUK 52.938 a 1 ,48 m, 212 ka, * ,SK,Pohronsky ,SK,Podunajska 02.192 P kt rf,Ktaiagn Tashiro-cho, Kitaakita-gun, pref., Akita JP, , 0 0 ,18 N, ,73 N, ´ 0 c 56.475 G inShn egn FraaKyrka (Fergana Fergana Tian-Schan, KG, , ˇ ,140 N, ovske E a,Bkvi,64m 42 m, 614 Bukovnik, Bar, ME, , * 0 0 ,2,- ,- -, -, 20, E, 37.805 ,SK,Kova ,17 N, 38.559 0 ,19 N, PUI LOP 15.348 ´ ´ 37.280 ´ DEM 0 n,25m 47 m, 275 rno, nk 6 ,48 m, 260 pnik, ,18 N, oc,Vr kopce, DUB 02.388 O ueor,Br,20m 45 m, 260 Baru, Hunedoara, RO, , 0 SOR 26.331 ,2,44 .62 2/17; 0.0622, 4/4, 20, E, 0 * ,18 N, 14.956 K il apt,Nova Karpaty, Biele SK, , 47.592 * 0 ´ ´ 34.611 0 ,1,- ,-; -, -, 19, E, ,SK,Nı ,08 /,004,5/22. 0.0146, 7/5, 0/8, E, R Dubrovac HR, , 16.129 c ˇ nvc Pı Inovec, ´ 18.916 ovske * ˇ 0 ´ K Slovensky SK, , nı tc 3 ,49 m, 730 atec, 0 ,17 N, ,- /,- 0/13; -, 2/1, -, E, ´ roe,58m 49 m, 588 Trnovec, ´ 0 s 07.296 ˇ z 0 ˇ ,19 N, FAG k 2 ,47 m, 221 ok, E,20,-,-,-; 0 n olns Ba lowlands, ina ´ ,2/,- ,-; -, -, 29/8, E, ˇ 0 ,22 N, 0 la 3 ,48 m, 132 alka, oc,Chl kopce, ,1,22 ,1/13; -, 2/2, 19, E, ´ k ar,Dema Tatry, zke ´ ,RO,Bras 10.692 a 7 ,48 m, 470 la, 46.652 0 11.022 VAC ,- /,000,3/23. 0.0503, 6/6, -, E, 57.939 13.658 * 0 ˇ 0 ,1,- ,-; -, -, 19, E, 0 ,18 N, ,SK,Stra ko-neretvanska ´ ,18 N, ˇ 0 21.006 b,12m 47 m, 142 aba, ,2/0 ,- -; -, -, 20/10, E, 50.012 rs Soros Kras, 04.172 o,Fa ¸ov, 49.351 KOM 0 ´ ,1,3/3, 19, E, ORA HAJ oc nad novce JP126 13.260 ´ 32.175 05.578 06.926 0 38.639 08.735 ,141 N, ¨ ´ 0 Bos ,22 N, novska 0 ˘ z 0 ˇ VAP ,18 N, * ,18 N, * ga ovske ,RO, ,SK, ,SK, ˇ ˘ ,JP, a BA ´ ras ˇ 0 0 0 0 ca- 0 0 N, N, N, N, ka E, E, * ¸, ´ ´ , ,