Delivered by Publishing Technology to: New Mexico State University IP: 128.123.253.108 on: Mon, 18 Mar 2013 00:39:19 Copyright (c) American Society for Plant Taxonomists. All rights reserved. O 10.1600/036364413X661917 DOI © Botany Systematic eu n xesv iapiaino ae.Frexample, For names. name of the the misapplication of extensive taxonomy the and of genus understanding poor a by complicated 1990; al. et (Bloom Raguso 2001). ecology al. and history et life Roy plant- Bloom Dobes and 2007); 2000; 2001); 2006; al. al. Roy et and al. et 1997; Schranz (Bishop 2007; et genes al. interactions Windsor of et pathogen Kantama 2000; evolution 2007a; al. 2009b); al. et Mitchell-Olds et 2009a, and (Bishop al. Song 2006; genomes et al. Kiefer et in Song 2007; research 2006; Dobes al. of 2001; et al. foci Knight et Other (McKay phylogeography 2012). and al. et Boechera (Beck hybrids these hybrids of loid These many although 2010). and triploid, apomictic al. are are al. et date et Aliyu to Schranz examined 2007; in 2010; al. interrelated 2009, are et issues 2005, Kantama 2004, 2006; al. 2005, et Sharbel 2007b; the of Much 2011). habitats al. of evolutionarily range et on and research wide Rushworth this large a 2001; across a from (Mitchell-Olds occurs to gained that applied insights genus be complex genetic to the species model apomic- allows is numerous and hybrids, It species diploid tic studies. sexual 70 evolutionary ± with plant to and, related of closely diversity relatively a for system iest faoitchbi iegs(ida and (Windham lineages hybrid apomictic and of diploids diversity sexual distinct, a this morphologically that that assemblage, five polytypic indicates includes pro- artificial, work research an of represents taxonomic variety “species” recent a in However, used jects. samples many to applied Mitchell- and Sharbel Dobes 2003; 2001; al. al. et Koch et 2001; Olds Naumova 1995; Roy (Bo polyploidy oyih 03b h mrcnSceyo ln Taxonomists Plant of Society American the by 2013 Copyright h nesv eerhitrs in interest research intensive The Boechera 5 rga on nvriy eateto ln n idieSine,25WdseBidn,Poo th862U .A. S. U. 84602 Utah Provo, Building, Widtsoe 275 Sciences, Wildlife and Plant of Department University, Young Brigham 2 rvosyasge to assigned previously fwl-upre ldsaeidentified. are clades well-supported of non fott lrf h aooyo h ru,w rsn hlgntcaaye fBehra,icuigalgnr ftetieadmost and tribe the of genera all including Boechereae, that of of indicate analyses species phylogenetic analyses present diploid we sexual group, the the of of taxonomy the clarify to effort ongoing Borodinia to missouriensis eann pce ssrnl upre n ean h name the retains and supported strongly is species remaining etr ot mrc.I stelretgnsi rb oceee hc loicue ee te ml genera. small other seven includes also which Boechereae, tribe in genus largest the is It America. North western eaa srcgie ntenwgenus new the in recognized is Nevada, eateto ilg,19Booia cecs(o 03) ueUiest,Dra,NrtCrln 70 .S A. S. U. 27708 Carolina Norht Durham, University, Duke 90338). (Box Sciences Biological 139 Biology, of Department oeua hlgntc n aooyo h Genus the of Taxonomy and Phylogenetics Molecular Keywords— Abstract— 4 Arabidopsis eateto oorpi tde,Msor oaia adn ..Bx29 t oi,Msor 36 .S A. S. U. 63166 Missouri Louis, St. 299, Box P.O. Garden, Botanical Missouri Studies, Monographic of Department nld nrseii dpain eei diversity, genetic adaptation, infraspecific include .holboellii B. A ´ 21) 81:p.192–209 pp. 38(1): (2013), 1 eutn nsvnnwcmiain ( combinations new seven in resulting , Boechera ilg eatet S A,NwMxc tt nvriy a rcs e eio803U .A. S. U. 88003 Mexico New Cruces, Las University, State Mexico New 3AF, MSC Department, Biology .Lo ¨ arc .Alexander, J. Patrick hr15,15,16;RyadReeeg1989; Rieseberg and Roy 1969; 1954, 1951, cher Boechera ¨ , n sbcmn oe eu o ies tde,fcsn atclryo pmxsadhbi pcain spr fan of part As speciation. hybrid and apomixis on particularly focusing studies, diverse for genus model a becoming is and Borodinia e&D Lo D. & ve ooii perstellata Borodinia oue nhbiiain pmxs and apomixis, hybridization, on focuses Hre. A (Hornem.) Boechera Bascca)i ies eu f±7 euldpodseisadnmru pmci yrd ocnrtdin concentrated hybrids apomictic numerous and species diploid sexual 70 ± of genus diverse a is (Brassicaceae) Boechera youla nogune yrdseito,lwcp ula oi oeua systematics, molecular loci, nuclear low-copy speciation, hybrid incongruence, cytonuclear , Boechera Boechera ¨ ercnl a eoeamodel a become has recently ve rbdpi thaliana Arabidopsis scretycrusrbd splpyei,cmrsn he ancae.EsenNrhAeia species American North Eastern clades. main three comprising polyphyletic, is circumscribed, currently as , Boechera loicue pmci dip- apomictic includes also omadsic ld ihteAintaxon Asian the with clade distinct a form ´ .Lo eas eryalhybrids all nearly because ˇ , ta.20a 04,2006, 2004b, 2004a, al. et 6 and uhrfrcrepnec ([email protected]) correspondence for Author ¨ 1,6 e&D Lo D. & ve e oiaeue,icuigtopatdlc,nDAIS n ee o-oyncerlc.These loci. nuclear low-copy seven and ITS, nrDNA loci, plastid two including used, are loci Ten . Boechera oenAllphin, Loreen Yosemitea ooii serotina Borodinia ihe .Windham, D. Michael ooii burkii Borodinia Bascca:Boechereae) (Brassicaceae: omnctn dtr nraWeeks Andrea Editor: Communicating ˇ ¨ s however, is, ta.2004a; al. et L)Heynh. (L.) ehsbeen has ve (as oeie repanda Yosemitea Boechera ). 5 ocearepanda Boechera , n .DnvnBailey Donovan C. and ooii canadensis Borodinia ˇ 192 lhuhrslto within resolution Although . hoooenme fsvn(ahrta ih,frspe- for eight, than in (rather maintained seven cies of number chromosome ifrne ewe h w eeaadterproposed their and genera two of the segregation between differences Lo of establishment circumscribed broadly n htgnrcsgeaini arne Kc ta.1999, al. et (Koch warranted is related American closely segregation ana- not generic phylogenetic are that taxa North of two and the series that on confirmed a have time, lyses work that Since authoritative Brassicaceae. (1993) Rollins’ 2010), America North of Flora of ori- exploration thorough hybrid A and a gins. boundaries on species based of not understanding are precise which efforts, research previous interpret per htnn ftepeiu tde iigti species this citing studies sampled previous actually the has of none of that exception appears the with and, Greenland Bo from known only loid 2006). Al-Shehbaz eea particularly anything genera, of or separation reflexed, Morphological strictly erect, between. are in and paral- septum), (flattened the to latiseptate dendritic Most lel narrow, to are tarns. forked fruits some alpine The least trichomes. at to with slopes perennials, are rocky precipita- species and m arid 4,400 temperature from ca. in to tion level ranging sea habitats near in from Mem- found elevation, California. are of genus state the the the of in in bers diverse particularly most A., is S. It U. East. western Far Russian the and Greenland, system. model a to as is potential genus full the its if reach required therefore is relationships logenetic A-hha 2003). (Al-Shehbaz 2 ¨ ¨ nrdcint Boechera— to Introduction Historically, ae .Beck, B. James e(96.Hwvr hs uhr i o ics other discuss not did authors these However, (1976). ve hrsrsac nGenad(Bo Greenland in research cher’s .ApiaiywsenNrhAeia ld opiigall comprising clade American North western primarily A ). ooii macrophylla Borodinia opooial bratseisedmct h Sierra the to endemic species aberrant morphologically a , Boechera , ooii dentata Borodinia Boechera nlds10seisfudi ot America, North in found species 110 includes Boechera Boechera Boechera oceaholboellii Boechera 2 Arabis 1 ha .Al-Shehbaz, A. Ihsan Arabis .holboellii B. Boechera n r eentaserdt h genus the to transferred herein are and ramn A-hha n Windham and (Al-Shehbaz treatment Arabis from pce eeicue ihna within included were species a is rpsdb Lo by proposed first was ) oacmdt pce ihabase a with species accomodate to . sntawy straightforward always not is L., n eae Genera Related and es tit slmtd number a limited, is stricto sensu , scrusrbdi h recent the in circumscribed As ooii laevigata Borodinia eg oln 91 93.The 1993). 1941, Rollins (e.g. Arabis Boechera .s hsmksi ifcl to difficult it makes This s. s. ¨ Boechera Yosemitea hr15,15,16) it 1969), 1954, 1951, cher .s sa pmci trip- apomictic an is s. s. a o olwdin followed not was aooyadphy- and taxonomy Boechera scoeyrelated closely is . 4 , Borodinia rmother from ¨ Boechera eand ve Delivered by Publishing Technology to: New Mexico State University IP: 128.123.253.108 on: Mon, 18 Mar 2013 00:39:19 Copyright (c) American Society for Plant Taxonomists. All rights reserved. rb oceeeA-hha,Bisen .A Kellogg, A. E. genera & monospecific the Beilstein, with along Al-Shehbaz, Boechereae tribe )eaiaino h tlt fpatddt nphylogenetic in data plastid of utility of analyses the of examination 3) rbda DC. genus Arabideae between sister its divergence with the tribal (along highlights major further a that Arabis to 2010) led Warwick al. 2006; have al. et et 2008) (Al-Shehbaz family (2006, the (2006) of al. al. reorganization et et Bailey Beilstein Broad by and 2010). Brassicaceae of Windham analyses and phylogenetic Al-Shehbaz treatments 2012; 2006, taxonomic 2007b, Al-Shehbaz recent and 2007a, Windham most 2005; in Holmgren 2001; genus (Dorn distinct a as ter Payson, drs he sus )rslto ftebudre fthe of boundaries the of resolution genus 1) to intended issues: are three analyses These address subspecies. treat- as that treated in or unrecognized ment either taxa additional several and America as North included, of are diploid 71 Boechereae the in of 57 genera are recognized all of ( bers loci plastid two and ITS, somal 03 LXNE TA. OCEAPYOEEISADTXNM 193 TAXONOMY AND PHYLOGENETICS BOECHERA AL.: ET ALEXANDER This 2008). 2006, American al. North most et of Beilstein transfer 2006; the prompted al. et Bailey 2001; 2000, 2013] to n/rISwr oiae ysmlsof samples by using dominated that were studies ITS found phylogenetic and/or and additional loci Several nuclear yses. and resulting (2009b) topologies plastid al. Boechera between of et conflict analyses Kiefer was from limited. there separate (2009b) that was presented noted al. Boechereae and within et Boechereae tion Kiefer plastid of and region. ITS taxa nuclear ITS 27 using ribosomal Brassicaceae included of nuclear analysis phylogenetic as the global Boechereae of a taxa 30 of included part from (2006) taxa al. et 10 Bailey than tribe. more the included that Boechereae of analyses Boechera— genera ditypic Phoenicaulis i nlsso oceeeuigsvnlwcp nuclear synthase, low-copy chalcone seven dehydrogenase, luminidependens using (alcohol Boechereae loci of analyses tic in the reconstruction Discussion). with phylogenetic (see in problems data fundamental plastid of be use may and This there genus. speciation the that that in suggests unlinked indicate network. be may (2009a) unrooted differentiation al. an haplotype et in and Kiefer monophyly groupings assess However, for values to confidence inability of the Unfortunately, lack com- the studies. is by network plicated previous this from conclusions in phylogenetic drawing than resolution in more sampling extensive with 2007b). 2007a, to 2006, both Al-Shehbaz due and interpret of (Windham to nature difficult polytypic are the and resolution genus little the yielded of members other of (Dobes representation little with l., s. divaricarpa B. rvosPyoeei nlsso oceeeand Boechereae of Analyses Phylogenetic Previous ol fteCretStudy— Current the of Goals network haplotype plastid a presented (2009a) al. et Kiefer Boechera ˇ Boechera and ta.20a 04;Shaze l 05.Teestudies These 2005). al. et Schranz 2004b; 2004a, al. et a o oohltci ihrpatdo T anal- ITS or plastid either in monophyletic not was Borodinia hr aebe uttopeiu phylogenetic previous two just been have There A-hha 03,adtercgiino h lat- the of recognition the and 2003), (Al-Shehbaz Boechera Boechera ut,and Nutt., A esn A Nelson) (A. )ietfcto fcae within clades of identification 2) ; Cusickiella Boechera , ramn A-hha n ida 2010) Windham and (Al-Shehbaz treatment Mps35 . ne hsrvsdclassification, revised this Under . .Busch, N. Boechera nteohrhn,i sindt the to assigned is hand, other the on , trnL-F , Polyctenium oln and Rollins Nuo ´ .Lo Draba Boechera ,and ¨ nlss nbt ae,resolu- cases, both In analyses. .divaricarpa B. e&D Lo D. & ve pce eonzdi the in recognized species rps16-trnK Nevada . srtie ntetribe the in retained is L.) epeetphylogene- present We pistillata Anelsonia ree swl sthe as well as Greene, Sandbergia 127acsin)and accessions) (1,287 ¨ .H Holmgren, H. N. e and ve, and and ,ncerribo- nuclear ), oceastricta Boechera .F ab.& Macbr. F. J. trnL-F Arabis Boechera Greene. .holboellii B. .holboellii B. Boechera .Mem- ). species trnL-F Arabis Dpa1 ;and Flora , , oceeeada ayo h iegn ili pce of species of diploid genera divergent all the represent of to other many effort by as an provided and in were Boechereae herbar- chosen samples recent were DNA from Samples extracted and researchers. few authors A the specimens. by ium made collections from obtained sdamdfe ihtruhu TBapoc iia othat to similar within approach CTAB (2003). study al. high-throughput this et for Mace in isolated modified silica presented samples the DNA a from remaining was contaminants The used EtOH washing 13. species-level 70% when step in at EtOH Tris-HCl columns existing 70% 10mm of that except instead procedures and (2007), used the al. following et isolated results Alexander was samples of 82 phylogenetic incon- from DNA of taxonomy. evaluation between limited indi- (study allow multiple TreeBASE to gruence by populations on represented different available are from are viduals species matrices Twenty-two is Data S12814). numbers, 1. number accession Appendix GenBank in including given information, Voucher Taxonomy 2010). outgroups. 81 three including and the Boechereae, used, follows of are members specimens other 95 11 mens, of total A possible. o Qae,Vlni,Clfri)o y1 iuedgsina 37 at digestion minute 15 by or California) Valencia, (Qiagen, QIAquick Qiagen col products were the PCR using The conditions either samples. cycling cleaned recalcitrant were although with deal the above, to of primer modified those sometimes each follow for generally programs listed cycler authors Thermal well. as included C.Frapiiain of amplifications For HCl. aiona OOT Cloning platysperma This TA samples. TOPO 20 California) possibility, ± this copies. multiple for evaluate amplifying unintelligible were To used was primers the sequence that concern the prompted of majority the Research Cancer Chicago of Facility. University Sequencing the DNA at Center’s out (both carried 25 phosphatase were per alkaline reactions Ohio) shrimp Cleveland, unit one Affymetrix, and from I exonuclease units five eie aulyuigS-l(abu 02.Gpcaatr were characters Gap 2002). and (Rambaut checked under then Se-Al 1994) were using al. alignments manually et resulting refined (Thompson The parameters. ClustalW default using automatically aligned cusickii B. lt N,15units 1.5 DNA, plate and 2007) 1991). al. al. et Taberlet et Addition- 1999). Shaw Doyle sequenced: trnK(UUU)x1, and were loci Bailey plastid pi1254r, 2007), two and and ally, 2007), pi504a Mitchell-Olds Mitchell-Olds (primers and and I Song intron Song Nuo-R, and Mps35-R, Nuo-F (primers and Mps35-F (primers their confirm To to belonging pollen. samples using all detect “ status, to the non-hybrid al. and difficult et assignment (Beck more taxonomic hybrids are diploid that apomictic from 2012) have unknown, excluded analyses previously microsatellite automatically recent identified However, were analyses. specimens morphology phylogenetic flowering All our from pollen 2006). identified this Al-Shehbaz readily and exhibiting be (Windham to specimens them most herbarium of allows typical grains that tricolpate ellipsoid, Brassicaceae) narrowly colpate the irregularly to than (ovoid, rather morphology invested grains pollen in distinctive was lineages a show hybrid effort and Many loid considerable (McDade hybrids. 2004), exclude analyses and Rieseberg phylogenetic identify and on process Linder this 1990; of effects confounding ahdT,6 m 60 dNTP, each 07,tenDAISrgo piesIS n T4 ht ta.1990), al. et White ITS4, and ITS5 (primers region luminidependens ITS nrDNA the 2007), 2000), ( al. synthase et chalcone 2006), al. et Slotte ( dehydrogenase alcohol sequenced: The diploids. non-hybrid species) sexual, sexual as exception, single sole interpretation a their from with derived alleles congruent and heterozygosity unpubl. “ low al. our (i.e. et of (Windham one samples but 2,400 All for data). loci 15 examined that analyses apigadDAIsolation— DNA and Sampling xlso fHybrids— of Exclusion lgmn,Mtie,adPyoeei Analyses— Phylogenetic and Matrices, Alignment, mlfcto n Sequencing— and Amplification ietsqecn of sequencing Direct mlfctoswr odce n25 in conducted were Amplifications Boechera Boechera .Egtt 0coe eune eegnrtdfo ahsample. each from generated were sequences cloned 10 to Eight ). ,023a lr fNrhAmerica North of Flora Dpa1 ld”(e eut)as eeicue nmicrosatellite in included were also Results) (see clade” .tiehmii B. piesL-1 n DX4,Sot ta.2006), al. et Slotte LD-XC4R, and LD-D1F (primers seWnhmadA-hha 07,20b n the and 2007b) 2007a, Al-Shehbaz and Windham (see M piesDa- n p1R ogadMitchell-Olds and Song Dpa1-R, and Dpa1-F (primers .sparsiflora B. ean,20 betaine, aeil n Methods and Materials Taq sdsusdi oedti below. detail more in discussed is , adh Boechera oyeae 0.5 polymerase, osdrn h rvlneo hybridization of prevalence the Considering ile ihyplmrhcras nwhich in reads, polymorphic highly yielded adh chs ,085 Ò m . n 7.5 , M a lnduigIvtoe (Carlsbad, Invitrogen using cloned was o orsc ape (017a samples such four for ld”smlssoe lei patterns allelic showed samples clade” ramn A-hha n Windham and (Al-Shehbaz treatment efmtra o N slto was isolation DNA for material Leaf C,15m 1.5 KCl, adh heialscheiranthoides Phoenicaulis ih ula-noe oiwere loci nuclear-encoded Eight chs rps16-trnK rmr d29 n Adh1043R, and Adh259F primers ; rmr SF n SR,Koch CS-R4, and CS-F1 primers ; trnL-F M m ecin ih1 with reactions L m oiesrmabmnwas albumin serum bovine m M C ecin Sequencing reaction. PCR L M fec rmr . m 0.1 primer, each of Ò piestn n trnF, and trnC (primers gl,ad1 m 10 and MgCl2, piesrs622and rps16x2F2 (primers C uiiainproto- purification PCR Boechera ahlcswas locus Each Boechera ,andB103 Boechera m ftem- of L r trip- are pistillata Boechera  M with C Mps35 speci- Tris- M Nuo as of Delivered by Publishing Technology to: New Mexico State University IP: 128.123.253.108 on: Mon, 18 Mar 2013 00:39:19 Copyright (c) American Society for Plant Taxonomists. All rights reserved. u tde eosrtn itrrltosi ewe ebr of members between by relationship here (represented Halimolobeae jaegeri sister of clade a a previ- and on Camelineae demonstrating based is studies rooting of choice ous this and Camelineae, tribe of members + GTR the uses which 2010), al. (Miller portal CIPRES et online the via 2008) al. et Stamatakis 2006; (Stamatakis 2008), al. and et tree-drifting, 1999). (Goloboff search, (Goloboff techniques sectorial TNT tree-fusing 1999), default in (Nixon using search” ratchet removal technology parsimony conducted using a “new was were the support with replicates for Clade replicates Jack-knife settings trees. 0.36. resampling 10,001 of jack-knife of probability 1,000 fur- maximum by a with followed to estimated replicate, bisection searching per tree trees TBR with 20 ther holding replicates (TBR), sequence (Nixon reconnection addition WINCLADA heuristic and random from 500 involved spawned with 2000) analyses 1999) (Goloboff NONA Parsimony in approaches. searches likelihood imum Discussion). and Results (see species this of origin o 5idvdas ihmsigdt ocnrtdpiaiyin primarily concentrated data missing with individuals, 95 for oyadM nlsswr eeal ogun.Wt few With congruent. generally were analyses ML and mony (i.e. analysis matrix nuclear Borodinia the by identified re from trees of analyses single-locus from and resulting Trees 1. shown Table are locus in each for sampling of completeness char- and acters, gap parsimony-informative of number characters, otide oooiswr o osrie oefrercpoa oohl fthese taxa. remaining of the monophyly and reciprocal outgroups enforce to constrained not were Topologies wallichii Crucihimalaya separately estimated the were locus. as parameters each model for analyses, clear. matrices, not ML combined is characters the bootstrap to gap For to 1,000 evolution prior of from removed models estimated available of were applicability was characters support The Gap and support. bootstrap inferred replicates. computing was when model tree CAT best + GTR the and search The excludes concatenated. were also loci matrix nuclear nuclear seven remaining the and Results) (see lsi arx200265 51.1% 3.8% 2.4% 95 95 95 50 23 155 206 98 803 2,050 5,615 676 matrix plastid trnL-F rps16-trnK matrix nuclear pistillata Nuo Mps35 luminidependens ITS Dpa1 chs adh 38 [Volume BOTANY SYSTEMATIC indepen- and not are individually matrix” they analyzed genome, “plastid not non-recombining and the a dent. were of loci as subsets Simmons plastid as to because, two of (referred The matrix”). matrices method “nuclear combined two coding and indel (Mu SeqState simple in implemented as the (2000) Ochoterena using scored 194 sso h lsi arxadncermti r hw in shown are within matrix groups Distinct nuclear respectively. 4, and and matrix 3 anal- Figs. plastid from the resulting Trees of yses S1–6. Figs. supplementary as able aiu ieiod(L nlsswr odce ihRAxML with conducted were analyses (ML) likelihood Maximum max- and parsimony both with conducted were analyses Phylogenetic asmn s aiu Likelihood— Maximum vs. Parsimony lge egh,nmeso asmn-nomtv nucle- parsimony-informative of numbers lengths, Aligned l nlsswr otdo h rnhcnetn h ld of clade the connecting branch the on rooted were analyses All e ifrn arcswr nlzd ahncerlcsseparately, locus nuclear each analyzed: were matrices different Ten Table Adh pistillata lsBehra Bisene l 06 08 ym ta.2008). al. et Oyama 2008; 2006, al. et (Beilstein Boechereae plus ) .Smayo aito mn h e oiue n h w octntdmtie.Sqecswr o eoee o %o h oa 0loci 10 total the of 3% for recovered not were Sequences matrices. concatenated two the and used loci ten the among variation of Summary 1. a xlddfo h ula arxdet aaoyproblems paralogy to due matrix nuclear the from excluded was ,and adh, r hw nFg.1ad2 epciey while respectively, 2, and 1 Figs. in shown are Yosemitea ITS, and lge eghPrioyifraiecaatr asmn-nomtv a hrcesNme facsin ecn isn data missing Percent accessions of Number characters gap Parsimony-informative characters Parsimony-informative length Aligned uiieedn,Mps35, luminidependens, ,2 2 79 0.6% 95 4.4% 94 27 18 120 1,129 131 1,089 rnbrni bursifolia Transberingia 2 62 51.6% 1.4% 2.4% 1.5% 1.8% 95 1.2% 5.1% 91 92 91 95 95 77 23 64 0 21 16 13 36 86 182 50 133 97 112 921 119 942 704 830 615 759 654 r niae nalfigures. all in indicated are ) oceatiehmii Boechera Results Crucihimalaya G oe hncnutn heuristic a conducting when model u oeiec fahybrid a of evidence to due and otermiigsamples. remaining the to ¨ eut rmparsi- from Results lr2005). ller and Transberingia Nuo Boechera Boechera Halimolobos r avail- are adh r both are Dpa1 .s., s. . .l. s. cuaercntuto frltosisi Boechereae. (100/99), supported in strongly is relationships Boechereae most the of of provide Monophyly to includes reconstruction considered (which is 4) accurate topology Fig. matrix loci; nuclear nuclear seven the data, plastid in paralogy with eaaeyi h lsi topology. plastid the in separately ieiodbosrp,btnti nlsso h plastid the analyses single-locus of in analyses recovered also maximum of in is 97% clade not / This but matrix. jackknife parsimony bootstrap), nuclear 99% the likelihood of 4; analyses in (Fig. support high matrix with recovered is s. s. Notably, topologies. different substantially resolved .perennans B. oooyti ruigi aahltc with paraphyletic, is grouping this topology of placement of exception placements Boechera the with divergent S2–6), Figs. of supplementary 2; and 1 (Figs. discussion. or so uncloned This well, the in as Methods). paralogy accessions unrecognized and is there Materials that suggests (see through out analyses ruled was microsatellite hybrids actually are paralogous individuals amplify these 2006) al. indicate et B103 in Slotte copies and (from 085, primers 023a, the that 017a, accessions for sequences inrcie nfrl orspot(o shown). resolu- (not additional support poor this uniformly received but tion analyses, parsimony than olution res- tree. greater ML provided best generally analyses the likelihood on Maximum occurred also analyses parsimony in sup- jackknife port 50% than greater receiving nodes all exceptions, Anelsonia, l cesosi h ula arxtplg,these topology, matrix lone (the nuclear support strong is the with exception multi- by together represented in grouped is accessions species a accessions if in case, acces- ple one evident one but than clearly all more In also by sion. represented are species of topologies examination plastid and nuclear within aa.Ti si hr otatt h lsi arxtopology, matrix ( plastid species the ten to which unpubl. contrast in sharp al. in et is analyses This Windham data). microsatellite morphology, by of species reexamination and distinct as supported are .fendleri B. eouini oceaadBoechereae— and Boechera in Resolution ula s lsi Resolution— Plastid vs. Nuclear d Clones— Adh chs , Dpa1 Boechera n oceee eo)adsm ntblt nthe in instability some and below) Boechereae, and Boechera , n otmmeso the of members most and .gracilipes B. and , ITS, , oceafernaldiana Boechera oceadavidsonii Boechera .s Fg ) oooia ifrne between differences Topological 3). (Fig. s. s. iegn lcmnso cloned of placements Divergent SplmnayFg ) h osblt that possibility The 1). Fig. (Supplementary .serpenticola B. luminidependens adh oceabreweri Boechera , adh eune n nmlu eut of results anomalous and sequences .laevigata B. snticue nfrhranalysis further in included not is .tiehmii B. h w useiso which of subspecies two the , aetoacsin placed accessions two have ) oee,i h plastid the in However, . , , Mps35 .lyallii B. , lsi n ula loci nuclear and Plastid Borodinia .canadensis B. , seRslto in Resolution (see Nuo , u oconcerns to Due .pendulocarpa B. ld included clade and , , .collinsii B. Boechera pistillata Nevada adh , , , Delivered by Publishing Technology to: New Mexico State University IP: 128.123.253.108 on: Mon, 18 Mar 2013 00:39:19 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 03 LXNE TA. OCEAPYOEEISADTXNM 195 TAXONOMY AND PHYLOGENETICS BOECHERA AL.: ET ALEXANDER 2013] ausblw5 r o hw,ada“”idctsta h oei usinddntocro h ette rmmxmmlklho analysis. likelihood maximum from tree best the on occur not did question in node the that indicates Suppor support. “–” bootstrap a likelihood and maximum shown, indicate branches not below are numbers 50 while below support, values jackknife parsimony indicate branches above Numbers Fig. .Src osnu hlga f1,0 otprioiu re lnt 1,C .3 I=08)fo asmn nlssof analysis parsimony from 0.87) = RI 0.83, = CI 316, = (length trees parsimonious most 10,001 of phylogram consensus Strict 1. DpaI sequences. t Delivered by Publishing Technology to: New Mexico State University IP: 128.123.253.108 on: Mon, 18 Mar 2013 00:39:19 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 9 YTMTCBTN Vlm 38 [Volume BOTANY SYSTEMATIC 196 ausblw5 r o hw,ada“”idctsta h oei usinddntocro h ette rmmxmmlklho analysis. likelihood maximum from tree best the on occur not did question in node the that indicates Suppor support. “–” bootstrap a likelihood and maximum shown, indicate branches not below are numbers 50 while below support, values jackknife parsimony indicate branches above Numbers Fig. .Src osnu hlga f3 otprioiu re lnt 5,C .5 I=08)fo asmn nlssof analysis parsimony from 0.81) = RI 0.75, = CI 758, = (length trees parsimonious most 38 of phylogram consensus Strict 2. pistillata sequences. t Delivered by Publishing Technology to: New Mexico State University IP: 128.123.253.108 on: Mon, 18 Mar 2013 00:39:19 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 03 LXNE TA. OCEAPYOEEISADTXNM 197 TAXONOMY AND PHYLOGENETICS BOECHERA AL.: ET ALEXANDER 2013] upr ausblw5 r o hw,ada“”idctsta h oei usinddntocro h ette rmmxmmlklho analysis. likelihood maximum from tree binomial. best the the on following symbols occur not typographical did matching question by in indicated node are the separately that resolved indicates are that support “–” species bootstrap a likelihood single and maximum a shown, indicate of branches not Accessions below are numbers 50 while below support, values jackknife Support parsimony indicate branches above Numbers matrix. Fig. .Src osnu hlga f1,0 otprioiu re lnt 1,C .3 I=07)fo asmn nlsso h plastid the of analysis parsimony from 0.76) = RI 0.63, = CI 912, = (length trees parsimonious most 10,001 of phylogram consensus Strict 3. . Delivered by Publishing Technology to: New Mexico State University IP: 128.123.253.108 on: Mon, 18 Mar 2013 00:39:19 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 9 YTMTCBTN Vlm 38 [Volume BOTANY SYSTEMATIC 198 arx(loci matrix eo rnhsidct aiu ieiodbosrpspot upr ausblw5 r o hw,ada“”idctsta h oei question in node the that indicates “–” a and analysis. likelihood shown, maximum not from are tree 50 best below the values on Support occur support. not bootstrap did likelihood maximum indicate branches below Fig. .Src osnu hlga f40ms asmnostes(egh=295 I=07,R .0 rmprioyaayi ftenuclear the of analysis parsimony from 0.80) = RI 0.74, = CI 2,905, = (length trees parsimonious most 480 of phylogram consensus Strict 4. chs , Dpa1 ,ITS, luminidependens , Mps35 , Nuo ,and pistillata .Nmesaoebace niaeprioyjckiespot hl numbers while support, jackknife parsimony indicate branches above Numbers ). Delivered by Publishing Technology to: New Mexico State University IP: 128.123.253.108 on: Mon, 18 Mar 2013 00:39:19 Copyright (c) American Society for Plant Taxonomists. All rights reserved. opooial oeieett,yttepatdtopology genera plastid distinct morphologically the the yet that suggests entity, species. cohesive single a morphologically is of members This of of placements speciation. paraphyly the and in morphological highlighted relationships, of independent nuclear partially least divergence, 3) at (Fig. be Boechereae to in appear relationships plastid inferred al. within cohesive contrast, clades et morphologically and which (Windham genera in within set Resolution Treatment, data also Taxonomic see microsatellite data; extensive unpubl. analyses an and morphology 4) of both (Fig. with matrix congruent nuclear generally the are of analyses phylogenetic our from tribe. the of members other to relative ou ula nlss h oiinof position the analyses, nuclear locus ( ( analyses mony holmgrenii N. ftoohrloci, other two of 199 relationships intergeneric are support only 50% TAXONOMY > AND The receiving PHYLOGENETICS unresolved. BOECHERA AL.: largely ET ALEXANDER are genera the among relationships but support, maximum receive 2013] r) h ape euldpodseiswithin species diploid sexual sampled gen- The these era). between distinctions morphological for Treatment Nevada, with 87; lemmonii Boechera one than more by represented (i.e. sample genera ditypic or monotypic robust the a of each Boechereae, allow Within monophyly. tribal not of assessment does sampling outgroup limited although ou ula nlss(is n n supplementary and 2 and and ITS loci, 1 Two (Figs. S1–6). Figs. analyses nuclear locus of number A 5. Boechera Fig. within recovered species in eight to shown two of groups are well-supported groups these of tributions Boechera the species); macrophylla American North three forms and analysis groups: matrix nuclear the in monophyletic and olsemr ail hnncerlc u otelower al. the et to (Birky genome due plastid the loci for nuclear size population al. than should effective rapidly loci et Plastid more (Kiefer capture. plastid coalesce incom- genus and this: sorting for the lineage explanations plete plausible in two are widespread There divergence 2009b). is haplotype species with coupled within haplotype species of More among pattern a sharing that circumscriptions. shows sampling plastid species extensive with inconsistent strongly-supported is unpubl. and topology these plastid al. 2007b), the et Nonetheless, 2007a, (Windham single data). 2006, sampling a microsatellite Al-Shehbaz within extensive and evalua- morphological from (Windham 4), (Fig. accessions tion together multiple grouped topol- are nuclear which species the of in combination a ogy, by entities real (e.g. as exceptions ported rare with are, chs noguneBtenNceradPatdLoci— Plastid and Nuclear Between Incongruence oceatiehmii Boechera pistillata ,btwt ekspot( support weak with but ), Nevada eaaholmgrenii Nevada and .s r aee nFg n icse below. discussed and 4 Fig. in labeled are s. s. Boechera .l) and l.); s. nloi,Cscila Phoenicaulis, Cusickiella, Anelsonia, lsthe plus 79) hr locus, third A 97/92). : lsfv atr ot mrcnseisof species American North eastern five plus Sandbergia nete h titcnesste rmparsi- from tree consensus strict the either in Mps35 .s cmrsn l u n ftewestern the of one but all (comprising s. s. ihmdrt osrn upr IS 79/ (ITS: support strong to moderate with , Yosemitea aismrel nispaeeti single- in placement its in markedly varies Borodinia chs ri h ette rmM analyses ML from tree best the in or ) ihsrn upr 9/9.Analyses (97/99). support strong with r etdwti t(e Taxonomic (see it within nested are Discussion and ( Boechera ocearepanda Boechera Anelsonia ld (59/63). clade Mps35 < pistillata .fernaldiana B. 0.I h eann single- remaining the In 50). Boechera Borodinia Dpa1 group , .s r ecie) By described). are s. s. .tiehmii B. plus group , grouped , .s n divergent and s. s. Boechera Cusickiella Boechera .Gorpi dis- Geographic ). Boechera ld ( clade and ,srnl sup- strongly ), .tiehmii B. .tiehmii B. sunresolved is Boechera Polyctenium Anelsonia, .tiehmii B. .l snot is l. s. .s and s. s. Borodinia .s sa is s. s. Results (95/95) with with .s. s. ) nlss(..Bie ta.20;Bisene l 06 2008) studies, 2006, these molecular al. to of Prior et relationships. Beilstein series true 2006; their a al. posited et as have Bailey there (e.g. only transferred have analyses Boechereae been most to single fact, assigned recently In a genus. currently a in genera within smaller accommodated less angustiseptate). much been Brassicaceae, and of rarely tribe has (latiseptate variation both types Such and compression incumbent), both and fruit silicles), (accumbent and fruit (siliques positions major family both cotyledon the include of would characteristic It types distinguish Brassicaceae. the to in used vari- A tribes extremely typically 4). be Fig. characters would (see encompassing node Boechereae able, this to by corresponding circumscribed node genus the is to assigned l. well- species s. all only includes The that 2. node Option supported favor to reasons nomenclatural and mono- well-supported genera. smaller, the into phyletic 2) split or be segregates, must the genus of large all or some supported encompasses appropriately the that an of node to boundaries expanded of be the must either one genus 1) large pursued: cases, be such must options in two classification congruent logenetically Drosophila Senecio ta.20) lwrn lnssc as such plants flowering (Schneider 2004), ferns al. asplenioid et including well- encountered groups, relatively been numerous has smaller, in situation This of genera. satellite assortment defined an to) paraphyletic hdne l 2010), al. et Thaden (in genus poorly-defined delimita- species-rich, case generic a this to involves respect This with tion. pattern familiar a exhibits species Boechereae. of of in members other estimations plastid relationships and misleading on phylogenetic provide reliance and to boundaries that likely clear to is in is needed data occurs it capture is Regardless, plastid but which research Boechereae. by genome mechanism Further the nuclear genome. establish paternal two, plastid the a of maternal with fusion than offspring rather nucleus producing sperm the by ( nucleus reduced and (2 a Hedtke unreduced with an by ing scenario, proposed this In recently allow (2011). was Hillis would capture, that species. plastid hybridization mechanism paternal for the additional via to an capture backcrossing Androgenesis, repeated plastid by for followed opportunities Dobes 2007a; vide Al-Shehbaz and of gametes Windham hybridization members unreduced the Frequent of many production loci. in in the plastid at and result at species, cohesive not between genetically could but are loci that capture likely nuclear species more Plastid of a pattern nuclear Boechereae. observed be and in to plastid Levin between seems and loci incongruence 2011) Chan genes for Hillis 2003; explanation nuclear al. and et than Hedtke Tsitrone 2005; rather see plastid species; biased between of introgression (i.e. movement capture haplotypes Plastid towards 2009b). divergent in al. observed et and is Kiefer that species species haplo- between within not plastid is extensive sharing This of loci. type combination over- nuclear the to in with relative result consistent loci should plastid sorting of lineage resolution incomplete so 1983), ntecs fBehra,teeaecmeln taxonomic compelling are there Boechereae, of case the In eei onaisi Boechereae— in Boundaries Generic .l Ple ta.20)adee nml uhas such animals even and 2007) al. et (Pelser l. s. Boechera .l vndrLnee l 07.T civ phy- a achieve To 2007). al. et Linde der (van l. s. .l)itmtl soitdwt adoften (and with associated intimately l.) s. Euphorbia x g ed orpaeeto h egg the of replacement to leads egg ) Boechera .l Bun ta.20) and 2006), al. et (Bruyns l. s. .s Nuoae l 2001; al. et (Naumova s. s. Boechera h rb Boechereae tribe The Draba ˇ ta.20b,pro- 2007b), al. et .s Fg and 3 (Fig. s. s. x .l (Jordon- l. s. pr unit- sperm ) Boechera Boechera .s. s. Delivered by Publishing Technology to: New Mexico State University IP: 128.123.253.108 on: Mon, 18 Mar 2013 00:39:19 Copyright (c) American Society for Plant Taxonomists. All rights reserved. nhraimseieseaie rmBY A,DK,E H ES O D,NC Y M S,U,U,adW,splmne with supplemented WS, and US, UC, RSA, RM, of distribution NY, the NMC, area, mapped NDG, the MO, of outside JEPS, that Note GH, 2011). (Kartesz E, Program DUKE, America North CAS, of Biota BRY, the from from information examined distribution specimens herbarium on 0 YTMTCBTN Vlm 38 [Volume BOTANY SYSTEMATIC 200 xed hog uho aaaadetnsit h usa a at( East Far Russian the into extends and Canada of much through extends n Que and Fig. .Dsrbto ftetregop within groups three the of Distribution 5. ´ e,with bec, ooii macrophylla Borodinia ijnti h usa a East. Far Russian the in disjunct Boechera .l.: s. Boechera .s., s. oceafalcata Boechera Borodinia ,and .Tedsrbto of distribution The ). Yosemitea ntecnietlU .A.Dsrbtosaebased are Distributions A.. S. U. continental the in Borodinia xed eihrlyit Ontario into peripherally extends Boechera .s. s. Delivered by Publishing Technology to: New Mexico State University IP: 128.123.253.108 on: Mon, 18 Mar 2013 00:39:19 Copyright (c) American Society for Plant Taxonomists. All rights reserved. lrsi ijnto,wihhsbe eotdi tleast al. et at Xiang in 1983; Spongberg 1999). reported Wen and 1998; been (Boufford genera has other 65 which disjunction, floristic American North Asian/eastern eastern the of example another ie,etr osalwydnaelae hrceitcof characteristic mm from leaves excluded (2–12 dentate smaller shallowly much Boechera leaves to the basal entire dentate from wide), coarsely different to strikingly repand are wide), mm (10–50 oial iia pce o nlddi u analyses, our in included not morpho- burkii species Two B. 5). similar (Fig. forest logically deciduous in generally Plains, missouriensis esof bers Boechera like but, (70/67) port to sister as The (88/89) supported genus strongly East is type Far which the Russian is monospecific taxa the (Hopkins American of North species species eastern that of Nested of group 2007b). this variety Al-Shehbaz among and a Windham as 1993; treated Rollins to 1937; often al. enough is similar et it is that Kiefer but analyses bootstrap; with phylogenetic clade parsimony a (90% 2009a). formed it support which area. strong in this data in ITS occur also Al-Shehbaz, perstellata Braun) L. (E. rnfre otegenus the to transferred 03 LXNE TA. OCEAPYOEEISADTXNM 201 TAXONOMY AND PHYLOGENETICS BOECHERA AL.: ET ALEXANDER 2013] Smelowskieae, hpbtenteto ncnieaino hs at,we facts, these of consideration In relation- two. close recognize a the support between analyses ship phylogenetic our below, discussed of most clade none morphologically American North Though eastern genus. the to a similar without species a is eaaadajcn ags suiu mn h western to the assigned among species unique American is Sierra North ranges, the adjacent to endemic and to taxon, Nevada This relationship unresolved. whose is genera group other (100/100) supported strongly Boechera soito with association and Halimolobeae, and the genus the in in included included was Nevada eur h eonto fjs n e eu n oa of are total genera 3) a These follows: level. and as species characterized and genus the new at combinations one discrete, new just eight of geographically recognition the and 4), require (Fig. analysis morphologically matrix nuclear 2) the by supported strongly to srbdt h rb.Tasern l ebr fthe advocate we of facts, new members 118 these ca. of splitting all require consideration thus In Transferring would combinations. genus tribe. one currently species into the 120 Boechereae ca. to the of two ascribed just included has cation, avail- oldest The is tribe. name the able in name move. generic other a every nearly such of consequences Boechera lie nomenclatural Boechereae to negative equivalent the genus hyper-diverse single, a of B Y eodortxnmcrsrain eadn h creation the regarding reservations taxonomic our Beyond ORODINIA OSEMITEA Borodinia Polyctenium oceaburkii Boechera Boechera utlrcnl atof part recently (until .s Fg ) ae nteersls h ee mem- seven the results, these on Based 4). (Fig. s. s. Boechera sarcn oecaua noain rdtdby predated innovation, nomenclatural recent a is Pre)Wnhm&A-hha and Al-Shehbaz & Windham (Porter) nti ein opooial nmlu and anomalous Morphologically region. this in .l.: s. a nlddi rvospyoeei nlssof analyses phylogenetic previous in included was .repanda B. Ti ld nldsfv ape pce of species sampled five includes clade —This Tetoacsin of accessions two —The and , Phoenicaulis ld sawoercie eaieywa sup- weak relatively receives whole a as clade Sandbergia Boechera Boechera .canadensis B. .l etitdt atr ot mrc are America North eastern to restricted l. s. r ogsadn eeawt oprevious no with genera long-standing are .l notregnr htae )moderately 1) are: that genera three into l. s. .serotina B. nanwmnseii genus, monospecific new a in a o enicue naypublished any in included been not has Yosemitea Draba .s nmlclrgrounds, molecular on s. s. or utl,wih nisbods appli- broadest its in which, Nuttall, rcnl eoe from removed (recently Borodinia Arabis l r on ato h Great the of east found are All . nwAaiee.Both Arabideae). (now , tceryde o eogto belong not does clearly it , Smelowskia .dentata B. .l. s. hsapast eyet be to appears This . Boechera .repanda B. a sindto assigned was ) , .laevigata B. .dentata B. Cusickiella .l t large Its l. s. .canadensis B. .perstellata B. Halimolobos .laevigata B. Yosemitea .repanda B. Borodinia, Anelsonia Boechera oma form with , was B. . ) . ehv bandmcoaelt aafrom data Second, microsatellite 4). (Fig. obtained clade have this in we placed robustly are all and 2006) of polyantha consideration. segregates B. American under North holboellii clade four B. that included the have evidence to we of First, lines assignment two its are support there but analyses, logenetic of range as known identified samples include 2005) al. “ et Schranz 2004b; al. et srtie o hsclade. the this for of retained one is that and hybrid within a is is parents triploid apomictic indicate this analyses Preliminary that data). unpubl. al. et (Windham trip- apomictic an As 2006). Al-Shehbaz loid, and from (Windham segregates l. apomictic s. and sexual of variety Dobes (e.g. studies previous many though in history nomic 2006). (Bo Al-Shehbaz known Greenland only taxon triploid in apomictic distinctive parsi- a of (56% on of species typified support topology type low ITS The with the bootstrap). although in mony clade (2009a), this al. in et included Kiefer also is East Far 2010). ( western Greenland in one these ( endemics, Quebec of St. eastern and in two few are Lakes one there Great A and the regions, 5). into River extend Lawrence (Fig. that Plains ranges have Great species the primarily of woodland coniferous west or chaparral, habitats, rocky to assigned previously ain is,teei upr o h oyyi aueof nature infor- polytypic the additional for provides support is that there resolution First, mation. finer is there within of divergence early the from other every of nearly taxon connects uninvolved that hybridization is apparently rampant and the species), in other encoun- two rarely just condition in tered (a lineages. glabrous completely two is branches, the between gap davidsonii Boechera genetic and morphological davidsonii B. te ape ebr of members sampled other sssoni i.4srnl uprs(99)asse rela- sister a (99/97) supports strongly between 4 tionship Fig. in shown ysis Nevada. County, Washoe and California Although County, Mono in an as tiehmii originated Boechera have near between may single-locus hybrid the it in intergeneric that in species suggest classification this analyses of this nuclear placements revisit Differing to future. necessary be may inaattecleto fadtoa irstliedata. microsatellite additional of of collection the origin await the tion of understanding tiehmii thorough B. more plausible. a geographically However, hybridization rendering Nevada, with co-occurs it counties, .holboellii, B. B Although eouinwti oceas s.— s. Boechera within Resolution OECHERA .holboellii B. oceafalcata Boechera Boechera Boechera Boechera n ia eiinrgrigisgnrcdisposi- generic its regarding decision final a and eaaholmgrenii Nevada .retrofracta B. .l noraaye ( analyses our in l. s. S and , .tiehmii B. shgl upre 1010,epaiigthe emphasizing (100/100), supported highly is hs ape eealcletdotiethe outside collected all were samples these ” . S —hscaeicue h aoiyo species of majority the includes clade .—This .s osqety h eei name generic the Consequently, s. s. a arwdsrbto napn habitats alpine in distribution narrow a has .quebecensis B. .s eev iteo ospot However, support. no or little receive s. s. .s sntaporaefricuini phy- in inclusion for appropriate not is s. s. .s WnhmadA-hha 06.Aside 2006). Al-Shehbaz and (Windham s. s. .holboellii B. sdsiciei hti a hcee caudex thickened has it that in distinctive is ¨ .holboellii B. Boechera .retrofracta B. .davidsonii B. hr15,15,16;Wnhmand Windham 1969; 1954, 1951, cher oceaholboellii Boechera stnaieyrtie in retained tentatively is Trz)A-hha fteRussian the of Al-Shehbaz (Turcz.) pce hti outyplaced robustly is that species a , Boechera .lemmonii B. eerh(e nrdcin and, Introduction) (see research Boechera a o encletdi these in collected been not has .s)(lSebzadWindham and (Al-Shehbaz s.) s. .davidsonii B. .s n rbbyrpeeta represent probably and s. s. .holmgrenii N. ida lSebz and Al-Shehbaz) & Windham ida n Al-Shehbaz and Windham ; n ld opiigall comprising clade a and .l,wihocri open in occur which l., s. .clisi .pendulocarpa B. collinsii, B. h ula arxanal- matrix nuclear The .s hssse ld to clade sister This s. s. nnab y County, Nye nearby in a ofsn taxo- confusing a has Boechera ˇ eprelationships deep , ta.20a Dobes 2004a; al. et and .holboellii B. Boechera is .lemmonii B. .holboellii B. .holboellii B. Boechera .s,it s., s. .s. s. ˇ , , . Delivered by Publishing Technology to: New Mexico State University IP: 128.123.253.108 on: Mon, 18 Mar 2013 00:39:19 Copyright (c) American Society for Plant Taxonomists. All rights reserved. eaaadClfri.Tog lsl related,and of closely is species Though California. lavender/purple-flowered Plateau, and primarily Nevada Colorado the the to to sister endemic taxon white-flowered region. Corners Four through the to California Nevada southern from distributed parapatrically retto WnhmadA-hha 2006). Al-Shehbaz and (Windham orientation 0 YTMTCBTN Vlm 38 [Volume BOTANY SYSTEMATIC 202 fses(–2 bevdi h genus. number the smallest petals in observed narrow the its contain (8–12) by seeds that group of fruits the non-undulate of members and distinguished other is the Nevada, Sierra from southern the of endemic rare omrytetda aite of varieties as treated formerly pulchra B. ral igd(.–. m ed e locule. per of seeds comprising row mm) single a pygmaea (0.8–2.5 containing fruits winged (3– erect wide broadly to relatively ascending have which mm), of 7 all species, three includes ru)i h hlgn ugssidpnetoiisfrthis for origins specialization. independent edaphic suggests phylogeny the in dis- group) three ( in species groups serpenticolous tinct of placement The pentine. suffrutescens B. constancei B. in in mm mm differ- 1.2 by 2.5–5 distinguished (styles length easily style (petals turn, in in ences species are, smaller-flowered which two mm), the 4.5–8 containing clade a larger-flowered the analysis, fruits rollei matrix B. nuclear pendent the seeds mm), In mm) locule. (0.3–1.5 per winged (2.5–6 broadly of wide row single relatively a containing have three All respectively. Mountains, Klamath and Nevada Idaho. Sierra northern central to Cascade constancei region the Boechera Intermountain of northern slope east the and from the Range arc through an Nevada in Sierra distributed central species common relatively a is taxa. diploid sexual three includes clade .holboellii B. .holboellii B. opiigteohrfu pce,alo hc have a to which sister ( diploids is of sexual flowers, three brick-red of all to clade yellow unique species, clade with clade, taxon a this four Within to fruits. sister other pubescent narrower, is wide, the mm comprising >5 fruits glabrous distinctive igi h oae ra ai,adClrd Plateau Colorado and Basin, Great Mojave, occur- fruits the regions. reflexed to in pendent ring with species five includes h aecaeadoe( one and clade same the ( two only 4), Fig. togyspotdcaeo non-“ of clade supported strongly swal upre 6/)a ebro h ag,non- large, the of member a as “ (64/–) supported weakly is pendulocarpa re icsin fsm fteewl-upre rusof groups well-supported these follow. species of diploid some of groups. discussions morphological Brief that recognized clades previously several seeds. to identifies biseriate correspond also analysis often matrix nuclear the and the into insight fruits some pendent providing Beyond widely shares it holboellii P P S UFFRUTESCENS LATYSPERMA ULCHRA .pulchra B. ptl –1m)i elspotd(59)a itrto sister as (95/99) supported well is mm) 8–11 (petals oceaglaucovalvula Boechera ld hticue the includes that clade ” ssse oasrnl upre 1010 clade (100/100) supported strongly a to sister is ihwiet upefoes hs he species, three These flowers. purple to white with ) GROUP .suffrutescens B. nlddi u apig(aee “ (labeled sampling our in included h otdsiciesgeaemorphologically, segregate distinctive most the , .howellii B. and .l ftefu euldpodsgeae of segregates diploid sexual four the Of l. s. suffrutescens cuso ait fsbtae,icuigser- including substrates, of variety a on occurs GROUP r aiydsigihdb eie n fruit and pedicel by distinguished easily are .rollei B. Ti oeaeyspotd(88)clade (78/87) supported moderately —This GROUP .collinsii B. and Ti togyspotd(99)clade (99/98) supported strongly —This and Ti togyspotd(100/100) supported strongly —This .rollei B. r biaeysretclu,whereas serpenticolous, obligately are .I em fsbtaepreferences, substrate of terms In ). group, .platysperma B. and .retrofracta B. oaeDsr nei with endemic Desert Mojave a , .pulchra B. .formosa B. .polyantha B. arerarespeciesfoundinthe .fendleri B. breweri oceaformosa Boechera holboellii )isnestedwithina .constancei B. oceahowellii Boechera r loarclyor allopatrically are , oceasuffrutescens Boechera . ru,and group, .holboelli B. , oceapygmaea Boechera .lincolnensis B. r lcdwithin placed are ) ru,wt which with group, ”taxa. .yorkii B. .lincolnensis B. holboellii largely a , complex, n 0.4– and Boechera Boechera puberula rare a , and , ”in and ,a nldstrespecies: three includes nlss h w cesosof accessions two to northward matrix the nuclear Basin the analysis, In Great Idaho. adjacent the and Washington from eastern occurring species uted and Oregon; southwestern and California northwestern Oregon; of species serpenticolous southwestern facultatively and California western eie rcoe r ifrn.Ti pce sstrongly 2006). Al-Shehbaz and is (Windham species of This ognition to different. sister as are supported trichomes pedicel of variety the to affinities morphological clear with of subspecies two nldssvnseis l fwihehbtwdl pendent widely exhibit which of all species, seven includes .serpenticola B. includes: iuto wisfrhrresearch. further awaits situation uaaaadSnJcnoMutiswietelte slim- Mountains. is Bernardino latter San the the to while ited the Mountains Jacinto in occurring San former and Cuyamaca the California, southern in endemics Desert; Mojave dispar Boechera features, these that on petal suspect Based number. we in ovule differences and major length, style exhibit size, they that given surprising h bec frslto ihnthe within (pannose) resolution hairs. of dendritic pubescent absence The (6–14-rayed) divided densely highly with entire, leaves basal and fruits, ascending parishii species: B. three includes ae npbsec,wihi ofndt h aa leaf basal the to in sepals separable confined and are is in and which margins pubescence, Range, on Cascade based southern and Nevada platysperma B. oehrsse oapltm encompassing polytomy a to sister together h otenClfri taxon California Although southern fruits. the arcuate long, predominantly relatively and with trichomes pedicels simple ascending having species by four These united support. are without though it, to sister placed orfee rista r oeo espbset(otygla- (mostly pubescent less or in more brous are that fruits reflexed to ieydsrbtdi h ra ai n otenSnake southern and Basin Great Plains; the River in distributed widely aymcoaelt nlss(ida ta.upb.data) with, unpubl. conspecific al. et indicate which (Windham analyses microsatellite nary ecology evolutionary puberula ongoing the of much in of research focus the and (Windham 2007b) is Al-Shehbaz lineages and hybrid Windham 2007a; of involved Al-Shehbaz array is and genus, diverse the particularly in a species in widespread North most the the and respectively. of one California, Utah of of Ranges Mountains Wasatch Coast the to endemic omacae u eainhp mn pce ihnthe within species among relationships but puberula clade, a form uhot ta.21) h lcmn of placement The 2011). al. et Rushworth D F P B ENDLERI REWERI UBERULA ISPAR ru r tews unresolved. otherwise are group ru saoaoscmae orslso prelimi- of results to compared anomalous is group .retrofracta B. l aelvne/upefoes divaricate- flowers, lavender/purple have All . GROUP .sparsiflora B. ocealasiocarpa Boechera GROUP .howellii B. GROUP Boechera .arcuata B. GROUP ,and .lasiocarpa B. .platysperma B. cussoaial nadaon h western the around and in sporadically occurs r oebodydsrbtdi h Sierra the in distributed broadly more are .retrofracta B. Ti togyspotd(19)clade (91/93) supported strongly —This .johnstonii B. Ti togyspotd(79)clade (97/96) supported strongly —This .johnstonii B. .polyantha B. Ti togyspotd(910 clade (99/100) supported strongly —This Ti elspotd(49)clade (84/93) supported well —This .breweri B. .subpinnatifida B. ). Aiue l 00 hre ta.2010; al. et Sharbel 2010; al. et (Aliyu .rectissima B. n ctee vrteses leaves, stems, the over scattered and oceapuberula Boechera hrsmn fteefaue,the features, these of many shares ) and .breweri B. oceadispar Boechera and . . .sparsiflora B. sgntclysmlrt,o even or to, similar genetically is oceacusickii Boechera and etrudrtnigo the of understanding better A . .arcuata B. , .serpenticola B. .sparsiflora B. and .puberula B. .parishii B. .sparsiflora B. aiaigtercn rec- recent the validating , ihtosbpce in subspecies two with , l aecoeypendent closely have All . dispar .parishii B. frel rae sa as treated (formerly , and oceapuberula Boechera sdsic species distinct as ru ssomewhat is group ocearetrofracta Boechera .retrofracta B. .johnstonii B. ieydistrib- widely a , .koehleri B. r itrspecies. sister are breweri orhspecies fourth a , , r aespecies rare are .subpinnatifida B. .retrofracta B. r grouped are .koehleri B. r narrow are ru,is group, n the and nthe in and , ,a is , , Delivered by Publishing Technology to: New Mexico State University IP: 128.123.253.108 on: Mon, 18 Mar 2013 00:39:19 Copyright (c) American Society for Plant Taxonomists. All rights reserved. .Fut aiett,trt,o udaglr evsudvddo aeypnaii,soft, . pinnatifid, . rarely or spines undivided pectinate leaves as quadrangular; years or subsequent terete, in latiseptate, persisting Fruits rigid, pectinate, leaves angustiseptate; 1. Fruits 1. entire. stigma repanda Yosemitea complete; accumbent. septum cotyledons winged; siliques, uniseriate, pubescent; Seeds latiseptate linear, or dehiscent, ovules Fruits glabrous white; fruit. petals per Racemes divaricate-ascending. pubescent; 34–50 sparsely to auriculate. or erect glabrous not Sepals pedicels petiolate, fruiting leaves ebracteate; rarely or cauline broadly dentate coarsely rosulate, surface entire; to leaves ground repand Basal obovate, near to rosettes. oblanceolate arising basal of branch, center Stems caudex from simple. per rarely 1 2–6-rayed, usually stalked, trichomes woody; 203 Yosemitea ongoing of subject data). the unpubl. al. are et group (Alexander this and research in boundaries hybrids of species TAXONOMY origin of AND western the PHYLOGENETICS Clarification Nevada, BOECHERA Mexico, Arizona. AL.: New southern ET and ALEXANDER Colorado, Texas, through Wyoming, occur- California southern group, species two Utah, southern widespread in from a arranged is seeds ring This biseriate locule. have addi- per seven In rows the trichomes. of (2-rayed) forked five some tion, least at and fruits 2013] ooii canadensis Borodinia burkii Borodinia B ORODINIA eb,sot oln-ie eenas adcsmsl not mostly caudices perennials; long-lived to short- Herbs, .Fut iert acoaeslqe,rrl ( rarely siliques, lanceolate to ovary/fruit linear per Fruits 4 ovules/seeds silicles; ovoid 2. Fruits 2. SS :17 1921.—TYPE: 137. 2: RSFSR 2.1875. 122. repanda Arabis stp:GH). isotype: 1852, Franklin Pennsylvania: Mercerburg, A. S. Co.: U. 2007.—TYPE: 237. 12: Bot. 1933; 1890; burkii 572. 15. U.S. Southeast. 17: Fl. Man. Club Small, (Porter) Bot. Torrey Bull. Porter, nov. Schulz]. macrophylla oo 34:38 03—YE .S .California: A. S. UC). isotype: U. GH; (holotype: 1866, 2003.—TYPE: Valley. Yosemite Co.: 388. Mariposa 13(4): Novon 1875; 122. 11: Arts nov. comb. canadensis nov. comb. o essigi usqetyasa pns... . . spines as years subsequent in persisting not .Pat cps;seslals;sessleypplae aea en ffutvle nsooig. reticulate; minutely anastomosing or valves smooth fruit seeds of leafy; veins stems lateral scapose; papillate; not silvery Plants seeds leafless; stems scapose; 3. Plants 3. vlsses(–1–5 e vr/ri . . . ovary/fruit per (6–)12–250 ovules/seeds rbslaevigata Arabis .Ctldn nubn;cuielae o uiuae ed iges plump wingless, seeds auriculate; not leaves cauline incumbent; Cotyledons 4. .Ctldn cubn;cuielae uiuaeo o;seswne rrrl iges flattened wingless, rarely or winged seeds not; or auriculate leaves cauline accumbent; Cotyledons 4. .Bshi o.Mtr eb lv.Bt Sada Bot. Glavn. Gerb. Mater. Bot. in Busch N. Pre)Wnhm&A-hha,HradPap. Harvard Al-Shehbaz, & Windham (Porter) .J lxne ida,gn nov.—TYPE: gen. Windham, & Alexander J. P. aea en ffut avsntaatmsn (except anastomosing not valves fruits of veins lateral .Tihmsecuieysml;oue/ed –2proayfut. ovary/fruit per 6–12 ovules/seeds simple; exclusively Trichomes 5. .Tihmsbace;oue/ed 23 e ovary/fruit per 12–30 ovules/seeds branched; Trichomes 5. L)A-hha,Nvn1() 8.2003.— 384. 13(4): Novon Al-Shehbaz, (L.) rbscanadensis Arabis rbsrepanda Arabis .E Schulz, E. O. Pre)P .Aeadr&Wnhm comb. Windham, & Alexander J. P. (Porter) aooi Treatment Taxonomic .Wto,Po.Ae.Aa.At 11: Arts Acad. Amer. Proc. Watson, S. S asn .J lxne Windham, & Alexander J. P. Watson) (S. ocearepanda Boechera L)P .Aeadr&Windham, & Alexander J. P. (L.) Mh.e il. or var. Poir. Willd.) ex (Muhl. .Br .n. s. Burk I. .Wto,Po.Ae.Acad. Amer. Proc. Watson, S. .baicalensis B. .macrophylla B. . p l :65 1753; 665. 2: Pl. Sp. L., S asn Al-Shehbaz, Watson) (S. Anelsonia .N oadr4881 Bolander N. H. .Bsh[= Busch N. hltp:PH, (holotype: ...... 2 ...... 3 ...... Trz)O E. O. (Turcz.) e oGnr fBoechereae of Genera to Key rbsburkii Arabis arwyoaesilicles; ovate narrowly ) ...... Boechera Boechera Draba burkii Phoenicaulis ooii serotina Borodinia htcnb itnuse sn h olwn key: following the using distinguished be can Sandbergia,that Polyctenium, Phoenicaulis, ( Nevada, genera nine tains ooii dentata Borodinia ooii laevigata Borodinia ooii missouriensis Borodinia ooii perstellata Borodinia ...... 4 ...... ) ne hsrvsdgnrccrusrpin oceeecon- Boechereae circumscription, generic revised this Under 3 6.1911; 365. 13: nov. comb. GH). (holotype: 1226 tees n. 1907, s. August of Steele 21 vicinity m, Mts., 500 shale, Alleghany disintegrating Co.: Millboro, Bath 2007.—TYPE: Virginia: 249. A. S. 12: U. Bot. Pap. Harvard Al-Shehbaz, TYPE: ae yRva nCfet n avs ao 13:521. 51(3): Taxon Jarvis, BM). and isotype: LINN; Cafferty 2002: in Reveal by nated dentatus nov. ory&A ry l .Ae.1 0 88 o Clairville, var. 1811; non 223. 1838, Suisse 80. 1: herbor. Amer. Man. N. Fl. Gray, A. & Torrey eintdb ulgn hdr 7 4.19:DWC, 1995: 143. 97: GH). Rhodora isolectotype: Mulligan, River, by Ohio designated sandy the Kentucky: A. of S. U. banks 2008.—TYPE: 293. 13: Bot. Pap. superfl; dentata nom. Boechera 389.2003, 13: Novon Al-Shehbaz, 1952; (Fernald) 23. 4: Phytologia Gleason, (Fernald) ob nov. comb. nov. comb. Windham, .S .Msor:Mnir 5My1894, May 15 Montier, Missouri: A. 2003.—TYPE: S. 387. U. 1908; 13(4): 244. Novon Al-Shehbaz, 5: (Greene) Veg. Regni Nov. 2003.—TYPE: n. 386. s. locality, without 13(4): Pennsylvania: A. Novon S. U. Al-Shehbaz, Willd.) ex 1810; 411. 1: Suppl. Encycl. 1801; Poir., 543. 3(2): Pl. Sp. lhr re,woe ilie,6My1936, May 6 Co.: hillsides, Franklin wooded Kentucky: Creek, A. Elkhorn S. U. 2003.—TYPE: 388. 13(4): 1940; nov. comb. NDG). (holotype: ...... hltp:PH). (holotype: shortii hri dentata Shortia oceaperstellata Boechera .Catn400 Clayton J...... 5 ...... Rf)Gen,Ptoi :24 1897; 254. 3: Pittonia Greene, (Raf.) hltp:US). (holotype: rbsserotina Arabis enl,Rooa4:28 1946; 208. 48: Rhodora Fernald, rbsperstellata Arabis rbsmissouriensis Arabis Rf)P .Aeadr&Wnhm comb. Windham, & Alexander J. P. (Raf.) nloi,Behr,Brdna Cusickiella, Borodinia, Boechera, Anelsonia, ...... oceaserotina Boechera Rf)A-hha auci Harvard Zarucchi, & Al-Shehbaz (Raf.) Sel)P .Aeadr&Windham, & Alexander J. P. (Steele) Mh.e il. .J lxne & Alexander J. P. Willd.) ex (Muhl. E ru)P .Aeadr&Windham, & Alexander J. P. Braun) (E. ethn pcmn(ettp,desig- (lectotype, specimen left-hand , a. ui.Bt 7 1840; 17. Bot. Autik. Raf., ...... Gen)P .Aeadr&Windham, & Alexander J. P. (Greene) rbslaevigata Arabis urtslaevigata Turritis E ru)A-hha,Novon Al-Shehbaz, Braun) (E...... 6 ...... tee ot.US al Herb. Natl. U.S. Contr. Steele, .W hr .n. s. Short W. C. .Ban hdr 2 47. 42: Rhodora Braun, E. rbsperstellata Arabis ocealaevigata Boechera oceamissouriensis Boechera Sel)Wnhm& Windham (Steele) ree eet Sp. Repert. Greene, .H .Muhlenberg E. H. G. Mh.e Willd.) ex (Muhl. ul xWilld., ex Muhl. and oceashortii Boechera rbsdentata Arabis .F Bush F. B. .L Braun L. E. (lectotype, Yosemitea Polyctenium .shortii A. Iodanthus .Braun E. Sandbergia Cusickiella Anelsonia (Muhl. Nevada .S. E. 31 ) Delivered by Publishing Technology to: New Mexico State University IP: 128.123.253.108 on: Mon, 18 Mar 2013 00:39:19 Copyright (c) American Society for Plant Taxonomists. All rights reserved. elti,M . .A lSebz .Mtes n .A elg.2008. Kellogg. A. E. and Mathews, S. Al-Shehbaz, A. I. A., M. Brassicaceae Beilstein, 2006. Kellogg. A. E. and Al-Shehbaz, A. I. Rushworth, A., C. M. Al-Shehbaz, Beilstein, A. I. Allphin, L. Alexander, J. P. B., J. Beck, I. S. Jr., O’Kane L. S. Mummenhoff, K. Mayer, S. M. Koch, M. D., C. the Bailey, of utility phylogenetic Potential 1999. Doyle. J. J. and D. varia- C. Quantitative Bailey, 2010. Sharbel. F. T. and 2007. Schranz, Bailey. E. D. M. M., C. O. and Aliyu, Bacon, D. C. Rajanikanth, G. J., P. Alexander, lSebz .A n .D ida.2010. Windham. D. M. Systematics and A. 2006. I. Al-Shehbaz, Kellogg. A. E. and Beilstein, A. M. A., I. of Al-Shehbaz, species American North most of Transfer 2003. A. I. Al-Shehbaz, research This isolation. DEB-0817033. DNA grant NSF for by material supported Robert was and leaf thank samples, to providing DNA like providing for also for McElderry would Schranz We Eric and (UT). Beilstein History (MO), for Mark Natural Garden of material Botanical Museum of Missouri Utah (JEPS), and removal California Survey allowing of History Natural University for Illinois (ILLS), (GH), and University specimens Harvard isolation: of DNA loans for tions Bo lo,T . .M akn n .C akn 01 clgcllf itr of history life Ecological 2001. Baskin. C. C. and Baskin, M. J. C., T. Bloom, of ecology Germination 1990. Baskin. M. J. and Baskin, C. C. T., C. in Bloom, evolution Rapid 2000. to Mitchell-Olds. approach T. and An Dean, M. 1983. A. Fuerst. G., J. P. Bishop, and Maruyama, T. Jr., W. C. Birky, 38 [Volume BOTANY SYSTEMATIC 204 ¨ hr .W 91 yooia n mrooia tde nteamphi- the in studies embryological and Cytological 1951. W. T. cher, Acknowledgments. eunedt:tie n rcoe revisited. and trichomes A and Botany phytochrome tribes data: from sequence inferred phylogeny Brassicaceae evolution. 607–619. trichome and phylogeny the drive hybridization diploid Does in 2012. asexuality Windham. to D. transition M. and Bailey, D. C. a Toward 2006. Al-Shehbaz. 2142–2160. A. 23: Brassicaceae. I. the of and phylogeny Windham, global D. M. Warwick, and ITS Brassicaceae. nrDNA to son gene nuclear low-copy Botany of genus Journal the American in reproduction apomictic for tion recip- a columns. silica-based using and DNA saw plant rocating quality high of recovery inexpensive Rapid Press. University 7 Oxford Oxford: Vol and Mexico York New of Committee. north America North of n hlgn fteBascca Cuiea) noverview. an (Cruciferae): Evolution and Brassicaceae Systematics the of phylogeny and to apomictic 93–108. 128: survivorship. (Brassicaceae): h auttv odadbiennial woodland facultative the Naturalist Midland biennial facultative the plant-pathogen in selection of targets coevolution. Molecular chitinases: plant mitochon- results. in some genes and for chloroplasts, and theory dria genetic evolutionary and population aseVdnkbre Selskab Videnskabernes Danske Boechera .Tihmsasn,sml,o obnto fvrosyfre/tlaeadsml;ltrlsepals lateral simple; and forked/stellate variously of combination a or a simple, with absent, valves Trichomes fruit base; at saccate 6. sepals lateral throughout; dendritic finely Trichomes 6. 5 1307–1327. 95: .Prnil ihwl-eeoe adxo od ot rcoe sal ipeado branched; and/or simple usually trichomes foot; woody or caudex well-developed with Perennials branched) (rarely 7. simple with or glabrous caudex; with perennials rarely biennials, Usually 7. (Brassicaceae). rbsholboellii Arabis o act tbs;futvle ihosueviso aeywt rmnn midvein; prominent few a a with (except rarely ovary/fruit or per veins obscure 20–250 with usually valves seeds/ovules fruit base; at saccate ovary/fruit not per 6–16(–16) seeds/ovules veins; lateral and midvein prominent Evolution oeua hlgntc n Evolution and Phylogenetics Molecular .Bsllae ral bvt obodyolnelt,15c ie oreydnaeo repand; or dentate coarsely wide, cm 1–5 oblanceolate, broadly to obovate broadly leaves Basal 8. .Bsllae iert arwyolnelt rrrl bvt,(.–059–)c ie nieo aeydentate rarely or entire wide, cm (0.2–)0.5–9(–2) obovate, rarely or oblanceolate narrowly to linear leaves Basal 8. etr ot mrc n atr ot mrc otynrho the of (except north border mostly A./Canada America S. North U. eastern and America North western A/Canada S. U. the of (except south border mostly America, North eastern trichomes; 2:214–230. 124: n rnvreRne nClfri n asnRnei eaa.. . Nevada in Range Carson Nevada and Sierra California prominent; in usually Ranges veins Transverse fruit and marginal auriculate; not leaves cauline hogotwsenU .A,Cnd,Aak,adRsinFrEs . . East Far Russian obscure; and usually Alaska, veins Canada, fruit A., marginal S. not; U. rarely western or throughout auriculate leaves cauline pinnatifid; or ewudlk otaktefloiginstitu- following the thank to like would We ieaueCited Literature 7 5322–5327. 97: pistillata rbslaevigata Arabis 5:89–120. 259: trnL 1 1719–1731. 91: Novon h ora fteTre oaia Society Botanical Torrey the of Journal The complex. :1–59. 6: nrnin intron ooii macrophylla Borodinia ndctldnu lns compari- plants: dicotyledonous in 3 381–391. 13: ocea Evolution Boechera? mrcnJunlo Botany of Journal American rbslaevigata Arabis ,es lr fNrhAmerica North of Flora eds. ., oeua ilg n Evolution and Biology Molecular oeua clg Notes Ecology Molecular Boechera ilgseSrfe Kongelige Skrifter Biologiske Sphaerocardamum Genetics variety Boechera p 4–1 in 347–412 Pp. . 0:513–527. 103: mrcnJunlof Journal American oceafalcata Boechera laevigata 3 20–30. 13: 6 985–995. 66: nteRsinFrEs).. . East) Far Russian the in (Brassicaceae). var. . n other and American laevigata :5–9. 7: Arabis Plant ndhF Flora nteRsinFrEs).. . East) Far Russian the in 93: Boechera Bo rys .V,R .Mpy,adT .Hdesn 06 e subgeneric new A 2006. Hedderson. J. T. and Mapaya, J. R. V., P. Bruyns, North Asian-eastern Eastern 1983. Spongberg. A. S. and E. D. Boufford, Bo ooof .A,J .Fri,adK .Nxn 08 N,afe rga for program free a TNT, 2008. Nixon. C. K. and Farris, S. J. A., P. Goloboff, Dobes Dobes Dobes hn .M .adS .Lvn 05 ek rzgtcioainand isolation prezygotic Leaky Dobes 2005. Levin. A. S. and A. M. K. Chan, on .2001. R. Dorn, odnTae,I,I ae .A lSebz n .A oh 2010. Koch. Kantama,L.,T.F.Sharbel,M.E.Schranz,T.Mitchell-Olds,S.deVries,and A. M. and Al-Shehbaz, A. I. Hase, I. I., Jordon-Thaden, 1937. L. M. in Hopkins, 174–418 Pp. Brassicaceae. 2005. H. N. androgenesis of Holmgren, role potential The 2011. Hillis. M. D. and M. S. Hedtke, Tucuma 2. ver. NAME) (NO NONA 2000. solu- A. times: P. reasonable Goloboff, in sets data large Analyzing 1999. A. P. Goloboff, Dobes ¨ ¨ hr .W 94 xeietltxnmcsuisi the in studies taxonomic Experimental 1954. W. T. cher, hr .W 99 ute tde nthe in studies Further 1969. W. T. cher, complex. nISand ITS on for classification century. time Garden twentieth the the to from Linnaeus history of relationships-a phytogeographical American Tidsskrift Botanisk hlgntcanalysis. phylogenetic author. the lxdslylresaecrmsm usiuin n aberrant and 14026–14031. 104: substitutions chromosome chromosomes. large-scale display plex oe frpouto nteNrhAeia genus American research. North Garden of Botanical Missouri decades the the seven of in compilation a reproduction (Brassicaceae): of modes genus evolution. American of aspects North Mechanistic in pseudogenes iaini ot American Boechera North in fication ls altp aito niae litcn yrdzto and American hybridization North Pleistocene of indicates radiation variation haplotype approach. plast integrative markers–an Botany chloroplast of Journal and American nuclear from oosgnms ai nrgeso fmtral neie DNA. inherited maternally of Evolution introgression rapid genomes: porous .d og 07 ili pmcso the of apomicts Diploid 2007. Jong. de H. Evolution and genera. netics related closely most its of identification and genus the of systematics and phylogeny Molecular 155–186. 106–148, 63–98, 39: York New Bronx: Cronquist. Press. West A. Garden Botanical and Holmgren, K. Intermountain P. Holmgren, the H. of N. plants Vascular Flora: 87–109. 60: discordance. phylogenetic cytoplasmic-nuclear in optima. composite for tions Publishing. West and yaiso yrdzto n pmxsi h vlto fthe of evolution the in apomixis and genus hybridization of dynamics ˇ ˇ ˇ ˇ ˇ . .Kc,adT hre.20.Ebylg,krooy and karyology, Embryology, 2006. Sharbel. T. and Koch, M. C., , . .Mthl-ls n .A oh 04.Itapcfcdiversi- Intraspecific 2004a. Koch. A. M. and Mitchell-Olds, T. C., , .H,T icelOd,adM .Kc.20b xesv chloro- Extensive 2004b. Koch. A. M. and Mitchell-Olds, T. H., C. , . .Kee,M ifr n .A oh 07.Plastidic 2007a. Koch. A. M. and Kiefer, M. Kiefer, C. C., , . .F hre,adM oh 07.Twrsudrtnigthe understanding Towards 2007b. Koch. M. and Sharbel, F. T. C., , ...... and .holboellii A. Boechera Borodinia 0 423–439. 70: 9 720–729. 59: + Tidskrift Botanisk Svensk aclrpat fWyoming of plants Vascular psbA-trnH divaricarpa ...... 8 ...... (Brassicaceae). rceig fteNtoa cdm fSine USA Sciences of Academy National the of Proceedings ...... 7 ...... ) (Brassicaceae). Arabis Euphorbia 4 141–161. 64: 55:524–540. ,and eunedata. sequence nesenadcnrlNrhAmerica. North central and eastern in Cladistics 1 2087–2101. 91: Epobaee nsuhr fiabased Africa southern in (Euphorbiaceae) oceaholboellii Boechera Cladistics 3 517–533. 93: oceastricta Boechera ytmtc n Biodiversity and Systematics rbsdumni,A. drummondii, Arabis oeua Ecology Molecular 4 774–786. 24: 8 31–44. 48: ln Biology Plant Taxon 5 415–428. 15: naso h isuiBotanical Missouri the of Annals d .Ceen:Mountain Cheyenne: 3. ed. , rbsholboellii Arabis 5 397–420. 55: Boechera oceaholboellii Boechera Bascca)inferred (Brassicaceae) (= :502–515. 9: ,Vol.2,partB,eds. rbsdrummondii Arabis 3 349–370. 13: Draba ytmtcBiology Systematic oeua Phyloge- Molecular ´ :Pbihdby Published n: rbsholboellii Arabis (Brassicaceae): (Brassicaceae) Intermountain + Phoenicaulis :321–331. 5: divaricarpa Yosemitea complex. Borodinia nasof Annals Boechera Boechera trnF Rhodora com- UUC ), , Delivered by Publishing Technology to: New Mexico State University IP: 128.123.253.108 on: Mon, 18 Mar 2013 00:39:19 Copyright (c) American Society for Plant Taxonomists. All rights reserved. esr .B,B odntm .W aeet n .E asn 07 An 2007. Watson. E. L. and Kadereit, W. J. Nordenstam, B. B., P. Pelser, Mu 2001. T. Mitchell-Olds, Chan, L. Liebowitz, T. Midford, E. P. Vos, R. Holder, T. M. A., M. Miller, Mitchell- T. and Sala, A. Richards, H. J. Lin, Z. J. Bishop, G. J. K., J. McKay, cae .19.Hbisadpyoeei ytmtc .Pten of Patterns I. systematics phylogenetic and Hybrids 1990. L. McDade, high-throughput A 2003. Crouch. H. J. and Buhariwalla, K. H. S., E. Mace, reticu- of Lo patterns Reconstructing 2004. Rieseberg. H. L. and R. C. Linder, system- Molecular 2001. Mitchell-Olds. T. and Haubold, B. A., M. Koch, evolu- Comparative 2000. Mitchell-Olds. T. and Haubold, B. A., M. 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Reporter breeding Biology molecular Molecular tropical Plant for protocol extraction DNA Notiser Botaniska plants. in evolution late plastidic coding from Evidence nuclear Brassicaceae: the of atics dehydrogenase Evolution and alcohol Biology lar and synthase in chalcone loci of analysis tionary America. Reproduc- author. of the 2001. Boutilier. populations K. apomictic and in (Brassicaceae). Ramulu, development S. tive K. Bergervoet, J. cie pcro ula iooa N IS nNrhAmerican North in tran- (ITS) divaricarpa internal DNA ribosomal the nuclear of of evolution spacer scribed concerted populations: natural in tion tnodUiest Press. University Stanford oeei N datasets. DNA logenetic n vlto of evolution and gradient. stress water of occurring rally adaptation local and holboellii profiling Boechera Expression 2006. Mitchell-Olds. T. 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(Brassicaceae). trnL-F rbsholboellii Arabis nulRve fEcology of Review Annual 3 1714–1724. 93: yoeei n Genome and Cytogenetic oceastricta Boechera oceastricta, Boechera Arabidopsis 9 369–381. 49: oceaholboellii Boechera pcrsequences spacer oeua Ecology Molecular h ln Journal Plant The 0:173–183. 106: 5 758–771. 75: Arabidopsis genetic a : h New The complex Bioinfor- close a , close a 92: a . Delivered by Publishing Technology to: New Mexico State University IP: 128.123.253.108 on: Mon, 18 Mar 2013 00:39:19 Copyright (c) American Society for Plant Taxonomists. All rights reserved. in,Q-. .E ots n .S ots 98 h atr sa and Asian eastern The 1998. Soltis. S. P. and Soltis, E. D. Q.-Y., Xiang, Formanova N. Schranz, E. M. J., A. Windsor, ida,M .adI .A-hha.2012. Al-Shehbaz. A. I. and D. M. Windham, X475 X489 X494 X450 X448 X437 JX146275, JX146367, JX146663. JX146458, JX146086, JX146550, summit JX146181, the JX146934, of E JX146839, km JX146745, 1.57 ft., ca. 6,475 Valley Hill, Long Orleans above A4: of ridge (7,438) on JX146660. JX146655, 341 NV A9: near A3: JX146659, City A7: JX146654, JX146658, cheiranthoides A2: Phoenicaulis A6: JX146657, JX146653, summit A5: JX146273, the A1: JX146656, JX146364, of JX146083, JX146455, E JX146547, km JX146178, JX146931, 3.32 ca. JX146836, Range JX146742, ft., 7,900 Rosa of Peak, SW Granite Santa just of ridge the (9732) On County: in Humboldt Nevada: Summit A.: Hinkey S. U. Gray, A. & Torr. JX146771, JX146207, Schranz, JX146301, E. JX146393, by JX146689. JX146483, provided JX146112, sample JX146937, JX146576, DNA JX146960, Price, JX146842, JX146865, A. R. JX146748, & JX146666. O’Kane, JX146089, Beilstein], JX146184, JX146278, M. 242 JX146370, JX146461, by JX146553, provided sample 0 YTMTCBTN Vlm 38 [Volume BOTANY SYSTEMATIC spe- noteworthy and New 2006. Al-Shehbaz. A. I. and and D. Amplification M. 1990. Windham, Taylor. J. and Lee, S. Bruns, T. J., T. White, 206 ida,M .adI .A-hha.20b e n noteworthy and New 2007b. Al-Shehbaz. A. I. and noteworthy D. and M. New Windham, 2007a. Al-Shehbaz. A. I. and D. M. Windham, ocesecp o ee ape sltdfo rehuematerial are greenhouse from sample isolated a Schranz. herbarium E. samples by for by provided represented seven sequenced and are for successfully samples Supplementary except All not in vouchers sequence.” Loci used “no by “A1”). prefixes indicated the (e.g., by 1 identified Fig. are adh they listed trnL-F, sequenced, are rps16-trnK, accessions order: pistillata, GenBank following sample, Nuo, the each in For loci study. with this in used ples X471 X485 X490 X456 X444 X433 JX146272, South of JX146363, JX146652. (10,895) JX146454, JX146082, summit JX146546, JX146177, the JX146930, head of ft., JX146835, at NW Range 10,050 JX146741, km Toiyabe 1.02 Peak, the ca. of Toiyabe Creek crest Near Chance County: Last Nye of Nevada: A.: S. JX146209, U. JX146303, no JX146394, JX146691. JX146485, JX146114, JX146484, JX146578, JX146962, JX146577, JX146690. JX146867, JX146247, JX146113, JX146961, JX146208, jaegeri sequence, JX146866, JX146302, JX146772, no sequence, Schranz, JX146430, E. Mountain, Beatty JX146521, Dry of JX146718. of JX146152, (7,013) JX146616, south summit peak the JX147000, isolated of SE on km ft., Etna 6,525 3.92 of ca. NNE Basin, County: Creek Elder Box Utah: Station, Barcroft near JX146651. Mountains, JX146081, JX146929, JX146176, JX146834, JX146271, 098 JX146740, JX146362, Beilstein], JX146453, M. JX146545, by sad- provided the near sample ft., Mountain, 13,670 [DNA soil, White rocky of in slope nestled facing dle, southeast the On County: eurycarpa A l eeaecp ocea ooii,adYosemitea. and Borodinia, Boechera, except genera All PPENDIX nloi eurycarpa Anelsonia hlgntcpten nsvndvregenera. diverse Evolution and seven netics in congruent patterns disjunction: floristic phylogenetic American North western and eastern Physiology with Plant conservation promoter the and microsynteny of 2006. extensive sequencing Mitchell-Olds. T. shotgun and Kroymann, Partial J. Schnabelrauch, Rosatti, D. J. Bishop, T. Patterson, R. Press. California Keil, of J. University Berkeley: D. Wilken. Goldman, H. D. H. California and D. of Baldwin, Plants G. Vascular B. Manual: Jepson The Botany of cies Press. Academic Diego: San White. J. T. and Sninsky, J. Gelfand, D. phylogenetics. Innis, for in gene RNA 315–322 ribosomal Pp. fungal of sequencing direct pmci hybrids. apomictic of species Botany in Papers of species rchmly wallichii Crucihimalaya Mn)Rlis N apepoie yE crn,JX146773, Schranz, E. by provided sample DNA Rollins, (Munz) .D ida 3362 Windham D. M. A ry .F ab.&Pyo,U .A:Clfri:Mono California: A.: S. U. Payson, & Macbr. F. J. Gray) (A. 1 61–88. 11: .VuhrifrainadGnakacsin o sam- for accessions GenBank and information Voucher 1. Boechera Boechera Boechera C rtcl:agiet ehd n applications and methods to guide a protocols: PCR 243 370 4:1169–1182. 140: 084 1 257–274. 11: Bascca)I euldiploids. sexual I: (Brassicaceae) avr aesi Botany in Papers Harvard Bascca)II diinlsxa ilisand diploids sexual additional III: (Brassicaceae) 0 178–190. 10: uikel douglasii Cusickiella uikel douglasii Cusickiella eaaholmgrenii Nevada Bascca)I:aoitctriploids. apomictic II: (Brassicaceae) .S . aiona ooCut:White County: Mono California: A.: S. U. , .S . eaa tryCut:No Virginia of N County: Storey Nevada: A.: S. U. , .D ida 2186 Windham D. M. a 05(M) X480 JX146905, JX146810, (NMC), 2005 May 4 , 085 .D ida 3108 Windham D. M. .D ida 99-22 Windham D. M. 087 Ho.f hmsn Al-Shehbaz, Thompson) & f. (Hook. h,Dpa1, chs, .Bisen26299 Beilstein M. .Bisen2001-72 Beilstein M. heialscheiranthoides Phoenicaulis oytnu fremontii Polyctenium hnmultiple When . oceastricta Boechera ITS, N apepoie by provided sample DNA , Rlis .H Holmgren, H. N. (Rollins) A ry oln,U .A.: S. U. Rollins, Gray) (A. ´ Boechera .GburJn,J G. J. Gebauer-Jung, S. , 2 235–257. 12: uiieedn,Mps35, luminidependens, u 00(NMC), 2000 Jun 8 , 9My20 (NMC), 2004 May 29 , 3Jn19 (NMC), 1999 Jun 13 , 01(O [DNA (MO) 2001 , n dto,eds. edition, 2nd , 7Jn20 (MO) 2001 Jun 27 , oeua Phyloge- Molecular 244 avr aesin Papers Harvard p 2–3 in 521–533 Pp. . adh eoereveals genome 083 Halimolobos lnswere clones S Watson) (S. Arabidopsis Anelsonia ,eds.M. ut in Nutt. Harvard 089 . .Shaz X477 X481 X496 X452 X449 JX146398, JX146489, JX146582, JX146695. JX146118, JX146966, JX146213, JX146306, JX146871, JX146777, Schranz, JX146694.E. JX146117, JX146212, bursifolia JX146305, JX146488, JX146581, JX146397, JX146965, JX146870, JX146776, Schranz, E. by provided X489 X494 X450 X447 X436 osqec,JX146211, sequence, JX146693. JX146116, no JX146396, JX146487, JX146580, JX146964, JX146869, JX146662. JX146085, 247 JX146180, sequence, no JX146366, JX146457, JX146549, fTonCekc.52 mN ftesmi 557 fFa o ut,5,535 Butte, Top Flat of (5,567) summit the of ft., tributary NE along km Shoshone 5.27 of ca. Creek NW Thorn County: of Gooding Idaho: A.: S. U. Greene, oes yoy of synonym a Joness, re ln .R 51A . odmlsfo h umre)junction Squaw (unmarked) Fork the North from 40.9319 of miles (at side road 35N18 west R. 2.5 F. 35N18A, the with R. on F. Mountain along Nosoni Creek of E miles Nati 2.6 Shasta Co.: Shasta California: X430 X428 X413 osequence. JX146423, no JX146514, JX146143, JX146607, JX146238, JX146991, JX146896, JX146330, JX146802, of City, locality (NMC), Bear [type Big Springs central Whiskey in of 18 Hwy. SE California miles Cactus of the National1.8 NE at miles Bernardino 18 3.8 Hwy. turnoff, California San Flats of side Mountains, east Canyon, Bernardino Cushenbury Forest, San Co.: ft., JX146661. Bernardino 7,450 JX146084, Peak, JX146179, Granite JX146274, of 355a Rosa JX146365, (9732) Santa JX146456, summit the the JX146548, in of 99-223 Summit Windham ESE D. Hinkey km M. of 3.13 SSW ca. road Range Along Co.: Humboldt JX146671. davidsonsii AC2: JX146670, Boechera AC1: JX146669, A9: JX146668, A2: JX146667, A1: JX146090, JX146185, JX146279, JX146371, JX146462, JX146554, JX146938, 09(UE,J169,J169,J168,J160,JX146509, JX146602, JX146986, JX146891, JX146710. JX146138, JX146797, JX146233, JX146326, JX146418, (DUKE), 2009 S asn lSebzsubsp. Al-Shehbaz Watson) (S. .S ot 7c.70ra ie Eo t ucinwt .S ot 197, Route S. U. with junction its of NE miles road 7.0 ft., along ca. Summit 11,400 Shaniko 97 near Route Shaniko of S. WSW U. Co.: Wasco Oregon: A.: JX146158, S. U. JX146253, JX146344, JX146622. JX146436, JX146063, JX146527, w/CO junction JX146911, the of JX146816, ft., WNW mi at rd Gunnison 7,850 1.6 of ca ESE 114, 50 Co.: IS Gunnison along Colorado: Creek Cabin A.: of S. mouth U. Weber, A. W. Rob.) no JX146712. JX146421, JX146141, JX146512, JX146236, JX146605, sequence, JX146989, JX146894, JX146800, ft., 4,270 (NMC), Quincy, central Campground, of miles Valley Deanes WSW road of miles 1.0 NE 4.3 Creek, miles Rock 1.1 Branch 24N28, East with National junction above the Plumas 24N20 from R. Nevada, F. Sierra of Central side Co.: N Forest, Plumas California: A.: S. U. . ie eto h umt . ie ato eta antCek[type Creek Peak Walnut central North of on east miles Park, 8.8 of summit, State locality the Diablo of west Mount miles Co.: 0.2 Costa Contra JX146713. California: JX146142, JX146237, JX146329, 350a JX146422, JX146513, in University JX146606, Padres Antioch 1001A of of Los NE locality Alexander Saddle, miles type Romero 6.7 the Mountains, of [near Peak, Barbara W Montecito Ynez Santa miles of road ENE Santa 0.5 miles Cielo 2.2 Co.: Camino East Forest, Barbara National Santa California: X471 X485 X490 X456 X444 X433 JX146281, JX146373, JX146464, JX146673. JX146092, JX146556, JX146187, JX146940, JX146845, JX146751, breweri B. h ucinwt .R 7(t40.9120 (at 27 R. F. with junction the fMnel nGsueVle nlwhl erCbeSdn a .6km 3.46 ca. Siding ft., Cobre 5,935 near Well, hill Cobre low of on Valley SSW Goshute in Montello of ft., 3,150 JX146715. JX146147, 120 Dam, JX146242, JX146334, of Arrowrock sequence, side no of N JX146518, JX146611, NW on road miles 3638A along road Creek Windham 0.6 Deer above ca. slope River on Boise Boise of E Co.: Boise JX146626. confluence JX146067, JX146257, its JX146348, JX146162, of sequence, no E JX146531, km JX146915, 1.05 JX146820, ft., JX146726, ca. 4,000 River Creek, Boise Browns the with of Fork Middle along sequence. City JX146420, no JX146511, A JX146140, JX146604, (Fernald) JX146235, JX146988, JX146893, JX146328, JX146799, (NMC), Boechera. .D ida 2055 Windham D. M. oytnu fremontii Polyctenium oceacobrensis Boechera oceadispar Boechera oceabreweri Boechera .D ida 3662 Windham D. M. ´ rbsbreweri Arabis subsp. D. lSebz&OKn,DAsml rvddby provided sample DNA O’Kane, & Al-Shehbaz (DC.) 354a .Lo ¨ .D ida 2102 Windham D. M. e&D Lo D. & ve 5Ar20 NC,J160,J160,JX146995, JX146900, JX146806, (NMC), 2009 Apr 25 , oceaarcuataBoechera 0Ar20 NC,J160,J169,JX146990, JX146895, JX146801, (NMC), 2009 Apr 10 , shastaensis 248 .dispar B. B103 M .Jns)A-hha,U .A:Clfri:San California: A.: S. U. Al-Shehbaz, Joness) E. (M. 3Jn19 NC,J164,J163,JX146932, JX146837, JX146743, (NMC), 1999 Jun 13 , a 00(M) X474 X488 JX146933, JX146838, JX146744, (NMC), 2000 May 9 , M .Jns)Dr,U .A:Nvd:Ek o:SW Co.: Elko Nevada: A.: S. U. Dorn, Joness) E. (M. Gen)N .Hlge,U .A:Nevada: A.: S. U. Holmgren, H. N. (Greene) S asn lSebzsubsp. Al-Shehbaz Watson) (S. adegawhitedii Sandbergia .Wto] ,2 ft., 3,320 Watson], S. N apepoie yE crn,JX146775, Schranz, E. by provided sample DNA , ¨ e .S . dh:Emr o:EEo Idaho of ESE Co.: Elmore Idaho: A.: S. U. ve,  ,270ft., 2,700 ], oceacusickii Boechera ,122.1452 N, 353a ,610ft., 6,150 ], shastaensis .D ida 3094 Windham D. M. .D ida 2375 Windham D. M. 6Ar20 DK) X479 JX146843, JX146749, (DUKE), 2009 Apr 26 , 359a oceaconstancei Boechera Nt. ida lSebz .S A.: S. U. Al-Shehbaz, & Windham (Nutt.) oceacollinsii Boechera nlFrs,nrho aeShasta, Lake of north Forest, onal 7My20 NC,JX146722, (NMC), 2000 May 17 ,  rbsarcuata Arabis .J lxne 993A Alexander J. P.  .J lxne 1004A Alexander J. P. ,122.1763 N, ) hc s52ra ie from miles road 5.2 is which W), ida lSebz .S A.: S. U. Al-Shehbaz, & Windham 003a .J lxne 997A Alexander J. P. Ppr ree N sample DNA Greene, (Piper) .J lxne 969A Alexander J. P. oceacrandallii Boechera S asn Al-Shehbaz, Watson) (S. 009a 306a 7My20 (NMC), 2004 May 27 , rbsjuniperina Arabis 352a 0Ar20 (NMC), 2001 Apr 30 ,  )[yelclt of locality [type W) Rlis Al-Shehbaz, (Rollins) ut] ,5 ft., 3,250 Nutt.], oceacollinsii Boechera 249 oceaevadens Boechera .S . Idaho: A.: S. U. , breweri oceabreweri Boechera Transberingia ,7Apr2009 ,12Apr2009 ,8Apr2009 ,U.S.A.: ,30Mar .D. M. B L. (B. .E. M. .J. P. 086 Delivered by Publishing Technology to: New Mexico State University IP: 128.123.253.108 on: Mon, 18 Mar 2013 00:39:19 Copyright (c) American Society for Plant Taxonomists. All rights reserved. X496 X452 X449 X448 X436 X423 JX146128, JX146223, JX146316, JX146408, JX146703. JX146499, JX146592, JX146976, X461 X458 X447 X435 X422 X417 osequence. no 174 JX146137, JX146232, JX146325, JX146417, JX146508, JX146601, 965A Alexander X446 X435 X423 X419 X404 osequence. no JX146094, koehleri JX146189, Boechera JX146283, ca. JX146375, Station ft., JX146466, 5,265 Kenworthy Peak, above Lion of canyon (6,868) summit 2954 in the of Hemet WSW Lake km 2.82 of SE Co.: Riverside ,0 ft., 7,100 . ie fOdnHl,±85mlsSo sln,Ssio Mountains, Siskiyou Ashland, of of locality S [type miles Forest 8.5 ± National Hill, River Ogden Rogue of W miles 4.5 ± A s h l aWindham&Al-Shehbaz,U.S.A.:Oregon:JacksonCo.:EastsideofMount n d ,5 0y a r d sS Eo ft h et o po ft h es k il i f t ,±0 . 2 5m i l e sEo ft h es u m m i t , NC,J169,J168,J168,J160,J160,JX146416, JX146507, JX146600, JX146984, Switzer JX146709. JX146136, JX146231, JX146889, JX146324, above locality JX146795, ridge [type 180 S. on U. (NMC), and 66 Flagstaff S. U. downtown of junction of the of of NE km NE 1.07 ca. Canyon Just Co.: JX146293, Coconino JX146385, JX146475, JX146683. JX146104, JX146568, JX146199, of JX146952, NW ft., km JX146857, 2.68 5,950 ca. JX146763, Mts., Spring, Valley Pine Grove the of Ash slope SE on Creek Leeds Along Rainbow of head the of Saline of NW of locality miles junction 3.2 [type the 190, of Canyon Hwy. California NNE & miles Rd. 2.5 DarwinValley Hills, western Rosa Park, National Santa Valley southern Death Mesa, Co.: Inyo California: A.: S. U. X482 X497 X453 X441 X430 X429 JX146174, JX146269, JX146360, junction hill its JX146451, JX146649. on JX146079, of JX146543, Vernal N miles JX146927, of ft., road N JX146832, 5,550 4.4 Co.: 40., ca. Uintah 44 U.S. Utah Utah: with near A.: Reservoir S. Steinaker JX146359, U. above Al-Shehbaz, JX146648. JX146450, & JX146078, JX146542, Windham JX146173, JX146926, JX146831, JX146268, JX146737, (NMC), o:EEo enlaogtiuayo lf re nBu onanca. Mountain Blue on Creek ft., Cliff 7,650 Spring, of Davis tributary of NNE along km Vernal 2.50 of ESE Co.: JX146066, vivariensis JX146161, JX146256, JX146347, JX146625. JX146439, JX146530, JX146914, eaa y o:Na rs fteTiaeRnea edo Last of head at Range Toiyabe the Peak, Toiyabe of South of (10895) ft., summit crest the 10,000 of NW Near km 1.02 ca. Creek Co.: Chance Nye Nevada: ft., fernaldiana 7,440 Boechera Taos, Garrapata JX146687. JX146110, of JX146205, JX146299, near JX146391, JX146481, N JX146574, 522 miles Mexico 229A New 15 of about Alexander side Forest, W National the Carson On Creek, Co.: Taos Mexico: New JX146686. JX146109, JX146390, JX146204, JX146480, ft., JX146573, JX146298, 7,800 JX146957, JX146862, Fe, JX146768, National Santa (NMC), Fe of Santa 2005 NE of boundary miles the 5 at about 475 Forest, Mexico New Along Co.: JX146700. Fe Santa JX146124, JX146219, JX146312, 226b JX146404, JX146495, 836B JX146588, Alexander J. P. 03 LXNE TA. OCEAPYOEEISADTXNM 207 TAXONOMY AND PHYLOGENETICS BOECHERA AL.: ET ALEXANDER southern Mountain, Bald on of locality lookout a [type the Forest National of of Sequoia W Nevada, Sierra side miles 1.4 East Tr., Co.: Rattlesnake & Tulare Tr. California: A.: S. smallhillontheNsideofShermanPassRd.,nearthejct.withWoodpecker U. Al-Shehbaz, & Windham 2013] em,23mlsN ftesmi fEgtDla onan[ertetype of the W [near Mountain miles Dollar 4.1 Eight of of Creek, locality summit Deer the with of confluence NW miles Snailback the 2.3 of the Selma, NW above miles River Illinois 0.8 the Trail, Beach of side NE Forest, National Siskiyou X450 X449 X437 X424 X419 osequence. no JX146129, JX146224, lignifera the Boechera JX146317, of JX146409, top JX146500, the Crater of of shore WSW 879a ft., N Alexander 8,850 the Forest, miles of National N Umpqua Mountains, miles meters Cascade 12.5 the southern ± Lake, 150 Chemult, of of side WSW ± West miles 15.5 Co.: Thielsen, ± Douglas spire, Oregon: Mount A.: of S. U. JX146349,summit sequence. Weber, summit A. JX146440, W. the no Watson) JX146532, of (S. JX146068, SSE JX146916, JX146163, km JX146821, 5.89 JX146258, ca. JX146727, Peak ft., 9,200 James (NMC), Mountain, of Middle slope of NE (7,078) on Valley Cache lasiocarpa Boechera JX146711. JX146139, JX146234, JX146327, JX146419, JX146510, JX146603, 985A Alexander rbsgracilipes Arabis oceagracilipes Boechera 1Mr20 NC,J165,J164,J164,JX146558, JX146942, JX146847, JX146753, (NMC), 2004 Mar 31 , oceafendleri Boechera .J lxne 875A Alexander J. P. 126b rbsarbuscula Arabis .D ida 2185 Windham D. M. 028 S .Wlh ida lSebz .S . th Uintah Utah: A.: S. U. Al-Shehbaz, & Windham Welsh) L. (S. 6Jl20 NC,J168,J168,J167,JX146593, JX146977, JX146882, JX146788, (NMC), 2008 Jul 26 , oceajohnstoniiBoechera 7Mr20 NC,J169,J169,JX146985, JX146890, JX146796, (NMC), 2009 Mar 27 , p 09(UE,J169,J169,JX146987, JX146892, JX146798, (DUKE), 2009 Apr 3 , 7Jn20 NC,J166,J166,JX146958, JX146863, JX146769, (NMC), 2005 Jun 17 , oceafernaldiana Boechera A esn .A ee,U .A:NwMxc:McKinley Mexico: New A.: S. U. Weber, A. W. Nelson) (A. Hwl)A-hha,U .A:Oeo:JspieCo.: Josephine Oregon: A.: S. U. Al-Shehbaz, (Howell) ree,700ft., 7,000 Greene], 349c Rlis on .S . th ah o:Suhedof end South Co.: Cache Utah: A.: S. U. Dorn, (Rollins) u 08(M) X473 X487 JX146972, JX146877, JX146783, (NMC), 2008 Jul 4 , Rlis .A ee subsp. Weber A. W. (Rollins) S asn .A ee,U .A:NwMexico: New A.: S. U. Weber, A. W. Watson) (S. .D ida 2733 Windham D. M. oceaglaucovalvulaBoechera Gen)Dr,U .A:Ua:Wsigo Co.: Washington Utah: A.: S. U. Dorn, (Greene) rbsglaucovalvula Arabis ree yoy of synonym a Greene, 347a 5Jl20 NC,J168,JX146881, JX146787, (NMC), 2008 Jul 25 , .D ida 2034 Windham D. M. u 00(M) X475 JX146819, JX146725, (NMC), 2000 Jun 8 , Mn)A-hha,U .A:California: A.: S. U. Al-Shehbaz, (Munz) oceagracilipes Boechera .D ida 3593 Windham D. M. .D ida 2195a Windham D. M. 312a (Rollins)W.A.Webersubsp. .D ida 2744 Windham D. M. 229a 041 oceahowellii Boechera u 03(T,JX146738, (UT), 2003 Jun 2 , .E oes,480ft., 4,880 Joness], E. M. oceaformosa Boechera oceafendleri Boechera .J lxne 225B Alexander J. P. rbshowellii Arabis 313a (M.E.Joness)Al-Shehbaz, .koehleri B. fernaldiana ,5May2004(UT), ocealemmonii Boechera .evadens B. ,U.S.A.:Arizona: .D Windham D. M. ,120ft., 1,200 ], ,31May2008 2Jn2000 Jun 12 , u 2003 Jun 3 , .Watson], S. S Watson) (S. ,U.S.A.: ,U.S.A.: ,830ft., 8,300 ], (Greene) ,16Jun 351a 007a 010a .J. P. .J. P. .J. P. .J. P. 213 X485 X490 X456 X453 X442 X430 JX146227, JX146320, JX146412, JX146706. JX146503, JX146132, JX146596, JX146980, JX146885, ie,±14mlsEEo h ihs on nCroCil,±55mlsNW miles 5.5 ± Chiflo, ft., Cerro on 7,270 point Questa, highest Red the and of Grande of Rio Rio ESE the miles of of 1.4 confluence side ± the east River, of the N miles up 6.8 halfway ± Canyon, about Grande Area, Recreation Rivers Wild Trail, JX146065, JX146160, JX146255, JX146346, JX146624. JX146438, JX146529, JX146913, fLk iyna ot fApn uc a .2k S fTesr Falls, Treasure of ESE km 0.32 ca. ft., Gulch Alpine 8,975 of mouth near City Lake of X454 X442 X431 X420 X415 X400 JX146650. JX146080, ft., 5,775 JX146175, end), JX146270, 331a (E JX146361, bridge JX146452, 40 U.S. JX146544, the 3000A of Camp- NNW Gulch Windham km Rabbit 0.93 near Reservoir ca. Starvation ground of shore W Co.: Duchesne X492 X458 X447 X435 X424 X419 JX146064, JX146159, JX146254, JX146345, JX146623. JX146437, JX146528, JX146912, ot lteRvrbig a .7k Wo h umt(04)o e Hill, Red of (10341) summit the near of ft., 285 SW U.S. 9,870 km along 4.37 Fairplay ca. of bridge SE River Just Platte Co.: South Park Colorado: A.: S. U. Weber, X478 X482 X497 X453 X441 X430 JX146259, JX146350, JX146627. km JX146441, JX146069, 1.76 JX146533, JX146164, ca. JX146917, Mountains Spring JX146822, of the NW JX146728, m, in Co.: 3,175 Spring, Trail Clark Mummy Loop of Nevada: WSW North A.: along S. Vegas U. Las JX146410, Al-Shehbaz, JX146704. & JX146501, Windham JX146130, (Tidestr.) JX146594, of JX146225, SE JX146978, miles 30 JX146883, JX146318, ± summit, JX146789, the ft., of (NMC), 5,900 NNW Mountains, miles Cascade 4.5 Randle, ± below Trail, Trail Creek Crest Killen Pacific the on the Adams Mount of side North Co.: Skamania Dee of ESE km 2.63 ca. tower ft., radio 6,215 to road Spring, JX146685. near Gee Range Egan of slope JX146108, W on JX146479, JX146203, JX146572, JX146956, lincolnensis JX146861, JX146297, km JX146767, 2.28 (NMC), JX146389, ca 2005 ft., Wash 7,175 May Mtn, Tohdildonih Zilditloi 7 overlooking of (8574) slope summit the on of Navajo NNW of NE Co.: X470 X484 X499 X455 X442 X432 JX146300, JX146392, JX146482, JX146688. JX146111, JX146575, JX146206, JX146959, JX146864, JX146770, JX146191, JX146285, JX146377, JX146676. JX146468, JX146096, JX146560, JX146944, JX146849, X470 X484 X499 X455 X443 X432 JX146261, JX146352, JX146443, summit the JX146535, of SW JX146919, km 8.75 JX146824, ca. ft., Mts. JX146730, 3,725 Dam Peak, Jarvis Beaver of the (6,529) in of tributary Along Canyon Co.: Washington JX146506, Utah: A.: Bulldog JX146599, S. U. JX146708. Weber, JX146983, A. W. JX146135, JX146888, Watson) (S. JX146230, JX146794, JX146323, (NMC), JX146415, 1999 Jun 5 a .0k Wo h tt ot 8big,805ft., 8,025 bridge, 28 Route Creek State Pine the along of City NW Pass km South 1.70 of JX146372,ca. WSW Co.: Fremont JX146463, Wyoming: A.: JX146672. S. JX146555, U. JX146091, JX146939, JX146186, JX146844, JX146280, JX146750, summit the near (NMC), of Ely WSW of km 3.65 E ca. Co.: Range (3,335)ofCleveCreekBaldy,2,780m, JX146291, Pine Creek Schell White the Nevada: JX146383, in A.: Summit Success S. JX146473, U. Al-Shehbaz, & Windham JX146566, JX146681. JX146950, JX146102, JX146197, JX146855, JX146761, [type Spring Granite of WSW km on of City 5.24 Bear locality ca. Big Lake, of Baldwin ENE overlooking Co.: Bernardino ridge San California: A.: S. JX146707. JX146321, U. Shehbaz, JX146413, JX146133, JX146504, JX146597, JX146228, JX146981, ca. JX146886, Spring JX146792, Moki (UT), at of Canyon South synonym of locality of [type a Spring Cow head Wild of near N Basin km 2.34 Beef of South Co.: iea igo h ot ieo aeelGp 5 ad Wo the of NW Lake, Chief yards ft., White 10,540 150 of ESE Park, ± miles National 1.6 Gap, Sequoia Trail, Farewell Gap Farewell of the side on of point north south highest Nevada, the Sierra on southern King the Mineral of slope West Co.: Tulare California: X498 X454 X442 X431 X420 X415 JX146070, JX146165, JX146260, JX146351, JX146442, bridge, 28 JX146628. Route JX146534, State the South of of JX146918, NNW WSW km Co.: ft., 1.37 Fremont ca. 8,025 Wyoming: Creek A.: Pine near S. City U. sequence. JX146414, Pass Weber, JX146505, A. no W. JX146598, (Greene) JX146134, JX146982, JX146229, JX146887, JX146322, JX146793, (NMC), 2006 rbtr fTalCno nteKia lta erU .8Ac.45road 4.5 ca. ft., 89A 7,300 S. Lake., Jacob U. of near Plateau ENE Kaibab miles the on Canyon Trail of tributary oceapallidifolia Boechera .D ida 2098 Windham D. M. .D ida 2092 Windham D. M. .D ida 99-145 Windham D. M. 168 005 069 rbsparishii Arabis ida lSebz .S . eaa y o:So Lund of S Co.: Nye Nevada: A.: S. U. Al-Shehbaz, & Windham oceaoxylobula Boechera oceapendulina Boechera oceapallidifolia Boechera .pallidifolia B. 8Ar20 NC,J163,J163,JX146928, JX146833, JX146739, (NMC), 2004 Apr 28 , .J lxne 757A Alexander J. P. B335 B242 .D ida 3022 Windham D. M. .Wto] ,8 ft., 6,480 Watson], S. oceapallidifolia Boechera .S . e eio asC. atChiflo East Co.: Taos Mexico: New A.: S. U. , ,720ft., 7,200 ], ocealyallii Boechera 7My20 NC,J162,JX146818, JX146724, (NMC), 2000 May 17 , 6My20 NC,J162,JX146817, JX146723, (NMC), 2000 May 16 , 314a .D ida 2038 Windham D. M. 118a 004 335a u 99(M) X479 JX146823, JX146729, (NMC), 1999 Jun 5 , .S . rzn:Ccnn o:Along Co.: Coconino Arizona: A.: S. U. , .S . ooao isaeC. WSW Co.: Hinsdale Colorado: A.: S. U. , .J lxne 1107 Alexander J. P. .D ida 2159 Windham D. M. .D ida 2349 Windham D. M. ocealyallii Boechera Rlis .A ee,U .A:Utah: A.: S. U. Weber, A. W. (Rollins) oceaoxylobula Boechera oceapendulocarpa Boechera oceaparishii Boechera .D ida 3080 Windham D. M. .D Windham D. M. .J lxne 881A Alexander J. P. 0My20 NC,JX146791, (NMC), 2008 May 20 , a 04(M) JX146755, (NMC), 2004 May 2 , 342a S asn on .S A.: S. U. Dorn, Watson) (S. .D ida 3426 Windham D. M. 013a rbsthompsonii Arabis .S . th a Juan San Utah: A.: S. U. , 015 oceapendulocarpa Boechera 016 .S . Washington: A.: S. U. , .D ida 99-142 Windham D. M. oceanevadensis Boechera a 00(NMC), 2000 May 6 , .D ida 3370 Windham D. M. ,9Jul2009(DUKE), oceapendulina Boechera oceaperennans Boechera u 00(NMC), 2000 Jun 1 , ,24Apr2001(UT), 03 a 2004 May 3 3023, S asn Al- Watson) (S. Gen)W A. W. (Greene) 142 ,17May2004 u 2008 Aug 3 , A Nelson) (A. .L.Welsh, S. Boechera ,26Apr .D. M. , , , Delivered by Publishing Technology to: New Mexico State University IP: 128.123.253.108 on: Mon, 18 Mar 2013 00:39:19 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 0 YTMTCBTN Vlm 38 [Volume BOTANY SYSTEMATIC salong California with Canyon junction Water its of in NNW Valley miles ft., Yucca road 3,600 1.5 of 62, ca. NNW Rd. Co.: Pioneertown Bernardino San JX146629. JX146071, JX146166, 208 a .9mlsN ftesmi 70)o tat ek ,5 ft., 6,450 Peak, Summit Atlanta Wells of near (7805) Hills summit 3015 Limestone the of the Windham NE in miles Atlanta 8.79 of ca. NE Co.: Lincoln Sierra Lake, sequence. no 170 Shaver JX146144, JX146239, JX146331, of JX146424, JX146515, town JX146608, the 1026A of of Alexander locality ESE of [type S of Forest miles NNE miles National miles 9.7 1.7 0.6 Rd., Creek Mountain, Grove McKinley Dinkey & Dinkey of Rd. Creek side Dinkey east of junction the the on sequence. Nevada, Sierra no southern JX146119, JX146214, rectissima Plumas Boechera JX146307, of SE JX146399, km JX146490, m, 14.5 1,285 Almanor, Almanor, to Lake junction 13752 of and NW 89 km 4.5 Hwy. line, sequence.County S. U. no Between Co.: JX146154, JX146249, rectissima Boechera JX146340, JX146432, ft., 9,900 JX146523, Forest, National Inyo Pass, Pass,1060B Mulkey Cottonwood of of N E miles miles Sierra 2.8 southern 1.2 Meadows, the Horseshoe of of slope side East NE Co.: JX146378,Nevada, Inyo San California: A.: JX146469, California: S. U. JX146561, Al-Shehbaz, A.: JX146677. S. JX146945, JX146097, JX146192, U. JX146850, JX146286, Weber, JX146756, ft., 6,035 Mts. A. Spring, Bernardino (NMC), Top San of W. the WNW in km Watson) Flat 1.13 Cactus ca. near S. Lucerne of ex SE Co.: sequence, Bernardino Joness no E. JX146565, JX146680. (M. JX146949, JX146101, JX146854, JX146196, road JX146290, JX146760, 18.5 JX146382, ca. (UT), ft., 5,475 93 Jackpot., 1999 Route at U.S. border May along Idaho Creek the Falls of Salmon S miles of side SE JX146263, Co.: of Elko JX146354, SE JX146636. JX146445, km JX146073, JX146537, 2.58 JX146168, ca. JX146921, Range JX146826, Shoshone ft., JX146732, the 7,200 in Spring, Austin Bradley to sequence. road along no Summit JX146149, puberula JX146244, Boechera JX146336, JX146427, road sequence, ft., 0.25 1,150 ca. Road, Canyon Road Ross Lake 3672a-A with junction Duck its along of Flat NW Robinson miles above bluff on of Omak side ft., E 1,200 JX146716. JX146148, 361a Rock, JX146243, on JX146335, Orchard JX146426, Chasmere sequence, of no of JX146612, N WNW 3668A meters Co.: 50 Windham Chelan ca. Washington: Pinnacles Peshastin A.: S. U. JX146095, JX146190, JX146284, JX146376, JX146467, JX146675. JX146559, JX146943, X457 X444 X434 X422 X418 X413 JX146682. JX146103, JX146198, JX146292, 311a JX146384, JX146474, JX146567, ig ttejnto fF .2N8adF .2,1. ie S of SSW miles 15.4 29, R. F. and 29N58 ft., R. 4,420 F. Forest, National of Trinity junction Hayfork, the Rattlesnake on Mountains at Bolly Yolla Ridge North the of side N Ranges, Klamath no JX146106, sequence. JX146201, JX146295, Moun- JX146387, Choop JX146477, Bully JX146570, JX146954, of summit ft., the 6,800 near tain, Ranges, Coast North Co.: Shasta Rogue Mountains, Siskiyou Valley, Seiad of of locality S miles [type N 0.55 Forest miles ± National Lake, 5.5 Lilypad River ± of side Butte, W Red the of on Trail Crest Pacific the Below X456 X444 X433 X422 X417 X402 A1: JX146072, AC1: JX146634, JX146167, A7: JX146633, A5: JX146262, JX146632, JX146635. A4: JX146631, JX146353, A2: JX146630, m, JX146444, 3,200 Peak, Thor JX146536, of (3,751) Pine 2570 summit Lone the in of Windham Lake SE Mirror km below 0.70 Whitney ca. Mt. Canyon of ESE Co.: Inyo California: 09(M) X485 X489 X494 X460 X457 no JX146517, JX146610, JX146994, JX146899, JX146805, (NMC), 2009 ft., 9,800 Peak, sequence. no 187 Toiyabe JX146098, JX146193, South JX146287, JX146379, of JX146470, (10,895) JX146562, ca. 3104 summit Range Windham Toiyabe the D. the of M. in NNW Canyon Crane km above 3.84 ridge on Summit JX146702. Ophir of JX146127, JX146222, JX146315, 147 JX146407, JX146498, JX146591, 871A Alexander oceaschistacea Boechera ocearetrofracta Boechera oceaserpenticola Boechera ocearollei Boechera oceapolyantha Boechera 3Jn20 NC,J167,J167,J166,JX146583, JX146967, JX146872, JX146778, (NMC), 2006 Jun 23 , u 09(UE,J161,J160,J170,JX146618, JX147002, JX146907, JX146812, (DUKE), 2009 Jul 2 , a 09(M) X488 X492 X497 X463 no JX146613, JX146997, JX146902, JX146808, (NMC), 2009 May 1 , 360a 017a 358a .D ida 2955 Windham D. M. .W alr13690 Taylor W. D. a 04(M) X472 X486 JX146951, JX146856, JX146762, (NMC), 2004 May 2 , 5Jn20 NC,J163,J162,JX146920, JX146825, JX146731, (NMC), 2002 Jun 15 , 4Jl20 NC,J168,J168,JX146975, JX146880, JX146786, (NMC), 2008 Jul 24 , oceapolyantha Boechera oceaserpenticola Boechera oceaplatysperma Boechera 0Ar20 NC,J160,J160,JX146996, JX146901, JX146807, (NMC), 2009 Apr 30 , 2My20 NC,J160,J169,JX146992, JX146897, JX146803, (NMC), 2009 May 22 , Nt. on .S . eaa y o:Eo Ione of E Co.: Nye Nevada: A.: S. U. Dorn, (Nutt.) .S . aiona rsoC. etsoeo the of slope West Co.: Fresno California: A.: S. U. , Rlis lSebz .S . aiona ikyuCo.: Siskiyou California: A.: S. U. Al-Shehbaz, (Rollins) 8My20 NC,J165,J165,JX146946, JX146851, JX146757, (NMC), 2004 May 28 , Gen)A-hha,U .A:Clfri:Plumas California: A.: S. U. Al-Shehbaz, (Greene) Rlis on .S . eaa y o:WNW Co.: Nye Nevada: A.: S. U. Dorn, (Rollins) .S . ahntn kngnC. N of NNW Co.: Okanogan Washington: A.: S. U. , Gaa)A (Graham) .D ida 99-115 Windham D. M. ida lSebz .S . California: A.: S. U. Al-Shehbaz, & Windham 133 167 p 04(M) X474 JX146848, JX146754, (NMC), 2004 Apr 1 , 6Jn19 JP) X475 JX146859, JX146765, (JEPS), 1993 Jun 16 , rbsrectissima Arabis oceaperennans Boechera oceapuberula Boechera ´ .Lo Gen)Wnhm&Al-Shehbaz, & Windham (Greene) .S . aiona rnt Co.: Trinity California: A.: S. U. , rbsrollei Arabis A ry lSebz .S A.: S. U. Al-Shehbaz, Gray) (A. ¨ 401b e&D Lo D. & ve .D ida 2959 Windham D. M. .J lxne 1052A Alexander J. P. oceapygmaea Boechera .D ida 99-101 Windham D. M. 6My19 (NMC), 1999 May 26 , ree,570ft., 5,730 Greene], oln] ,0 ft., 5,800 Rollins], ¨ 144 e .S . Nevada: A.: S. U. ve, ,U.S.A.:California: .S . Nevada: A.: S. U. , oceapulchra Boechera .W Spellenberg W. R. .D Windham D. M. .J Alexander J. P. ,1Apr2004 ,28May (Rollins) .D. M. .D. M. .D. M. 356a 018a .J. P. .J. P. ,25 270 h anCwCek . ie Wo i lno . ie fthe JX146339, of JX146431, N miles JX146522, 0.5 JX146617, ± JX147001, Blanco, Rio JX146906, of ft., with JX146811, SW confluence 7,300 miles its line, 4.5 from County miles ± Garfield 0.3 Creek, Cow ± Creek main Cow the Branch West Along Co.: X489 X494 X450 X448 X437 X426 JX146171, JX146266, JX146357, JX146646. JX146448, JX146076, JX146540, JX146924, JX146829, h eirPaeuc.45 mNEo h umt(,9)o Malmsten on (9,793) summit the of ft., NNE on km 8,075 Creek 4.58 Pole Peak, along ca. Kingston Plateau of NE Sevier Co.: JX146411,the Piute Utah: A.: JX146502, S. U. JX146595, Al-Shehbaz, JX146705. JX146979, JX146131, JX146226, JX146884, JX146319, JX146790, (NMC), . ie N ftejt fU .Hy 4&U .Hy 6i se Park, Estes in 36 Hwy. locality S. type U. [near Forest & National 34 Roosevelt Haven, Hwy. of Glen S. of U. SE of miles jct. 0.4 the ± Trail, of Mountain NNE Crosier miles of side 6.4 E ± Knob, Knapp’s of JX146188, side Southwestern Co.: JX146282, JX146374, JX146674. JX146465, JX146093, JX146557, JX146941, JX146846, a . odmlsWWo h ucinwt O12 aged Cristo de Sangre 142, CO with junction the 159 ft., of CO 7,935 along WSW Creek miles Grant, Culebra overlooking road slope 1.6 on ca. Luis JX146639. San of A11: WSW Co.: A5: JX146645, JX146642, A9: A4: JX146641, spatifolia JX146644, A3: A8: JX146640, JX146170, JX146643, A2: JX146265, JX146638, JX146356, A1: JX146447, Mt. JX146075, of JX146539, (9601) summit JX146923, the of JX146828, N ft., km JX146621. 3.12 6,600 ca. Lewis, Range JX146062, Shoshone the in JX146157, Canyon JX146252, sparsiflora Boechera JX146343, ft., JX146435, 4,000 Creek, Browns River with Boise confluence the its 2374 of of Fork E Middle km along 1.05 City ca Idaho of ESE Co.: Elmore Idaho: X476 X480 X495 X451 X448 X438 JX146296, JX146388, JX146478, JX146684. JX146107, JX146571, JX146202, JX146955, JX146860, JX146766, the of SSE km 1.17 ca. Creek ft., Tamarack 8,400 Peak, of Ralston of head (9235) near summit Kyburz of Lake ENE Co.: Cup Dorado El below California: A.: S. U. Al-Shehbaz, Watson) (S. o:LsCac ag,o h ieo ayn40 fLs Chance Last of S 440m canyon a of of side [isotype N the Mt. on Range, Chance JX146245, Las Co.: JX146337, JX146428, JX146717. JX146519, JX146150, JX146614, JX146998, JX146903, JX146381, JX146472, JX146564, JX146679. JX146100, JX146948, JX146195, ft., JX146853, JX146289, 5,100 JX146759, across lodge, Park, State the (NMC), Mts. Davis from Co.: Davis canyon Jeff JX146358, Texas: the A.: S. JX146449, U. Al-Shehbaz, JX146541, & JX146647. JX146925, JX146077, JX146172, JX146830, JX146267, JX146736, (NMC), JX146380, JX146678. summit JX146471, the JX146099, JX146563, of JX146194, S JX146947, km JX146852, JX146288, 1.12 ft., JX146758, ca. 7,875 Range (NMC), Peak., Rosa Chocolate near Santa of Winnemucca the (8,017) of NNE in Co.: Summit Humboldt Nevada: Hinkey A.: S. U. Al-Shehbaz, X442 X430 X427 X412 JX146698. JX146493, JX146122, JX146586, JX146217, JX146970, JX146310, JX146875, just Stop; of One JX146781, N JX146402, Herman ft., Mt. miles (UARK), 730 from 18 259 Campground, 2005 S. Lake; Creek U. Holly Bow of the E Broken outside miles Co.: 8.8 ca. McCurtain Bow; Oklahoma: Broken A.: S. U. X487 X492 X458 X446 X435 X424 JX146169, of JX146264, (6126) JX146355, JX146637. summit the JX146446, JX146074, of JX146538, ESE JX146922, km JX146827, 9.46 ft., ca. 5,460 Mountains Pass, Red Cricket of sequence. E inhills no JX146146, shockleyi JX146241, JX146333, sequence, re ln tt ot 8c.4. odmlsN fisjnto ihU S. U. with junction its of NE ft., miles 8,125 road Willow 191, 42.1 West Route ca. overlooking 28 slope Route on State along Farson Creek of of NE [holotype Co.: JX146200, Fremont River Wyoming: JX146294, Red JX146386, sequence. no with JX146476, JX146105, JX146569, confluence Rio JX146953, Bottom, of ft., JX146858, Co.: N 6,800 Taos villosa], mi Mexico: B. New 2 A.: Canyon, S. JX146497,Grande U. JX146590, Al-Shehbaz, JX146974, & Windham JX146879, sequence. JX146785, no JX146126, JX146221, (NMC), JX146314, JX146406, 2008 Mt. of Jul NNE ± National 21 miles Toiyabe Peak, 1.6 Reno, central Church ± of of locality Rose, [type of SW Forest Mt. miles side 14 on ± point W Range highest Carson Co.: Houghton, the of Washoe NNW Nevada: miles A.: 0.8 S. U. Al-Shehbaz, sequence. no JX146153, JX146248, Borodinia. rbsspatifolia Arabis 0Ar20 NC,J162,J161,J161,JX146526, JX146910, JX146815, JX146721, (NMC), 2001 Apr 30 , Mn)Dr,U .A:Ua:MladC. Eo lc Rock Black of NE Co.: Millard Utah: A.: S. U. Dorn, (Munz) Rd. ida lSebz .S . ooao Costilla Colorado: A.: S. U. Al-Shehbaz, & Windham (Rydb.) 288 .D ida 2128 Windham D. M. .yorkii B. .D ida 99-105 Windham D. M. ooii canadensis Borodinia .D ida 3057 Windham D. M. .D ida 99-014 Windham D. M. yb] ,0 ft., 7,700 Rydb.], .D ida 3803B Windham D. M. 192 322a 400a ,U.S.A.:Nevada:LanderCo.:SWsideofDean 002a. .L ae 1113 Baker L. W. rbstiehmii Arabis 367a oceayorkii Boechera ,790ft., 7,910 ], oceaspatifolia Boechera oceastricta Boechera oceasparsiflora Boechera oceawilliamsii Boechera .J lxne 811A Alexander J. P. 148 oln] 050ft., 10,500 Rollins], .J lxne 824-1A Alexander J. P. oceasubpinnatifida Boechera .D ida 3127 Windham D. M. .Yr 2611 York D. 8My20 NC,JX146735, (NMC), 2000 May 28 , .J lxne 174D Alexander J. P. 4My20 NC,JX146752, (NMC), 2004 May 14 , 7My17 NC,JX146764, (NMC), 1979 May 17 , 6My19 NC,JX146734, (NMC), 1999 May 26 , L)P .Aeadr&Windham, & Alexander J. P. (L.) .D ida 2579 Windham D. M. .By,U .A:Clfri:Inyo California: A.: S. U. Boyd, S. 150d 025a 1My20 NC,JX146809, (NMC), 2009 May 31 , 310a 1Ar19 NC,JX146733, (NMC), 1999 Apr 21 , .S . ooao i Blanco Rio Colorado: A.: S. U. , 027 .S . ooao Larimer Colorado: A.: S. U. , oceatexana Boechera oceastricta Boechera oceatiehmii Boechera .MEdry544 McElderry R. Nt. on .S A.: S. U. Dorn, (Nutt.) Rlis on .S A.: S. U. Dorn, (Rollins) oceasuffrutescens Boechera 6My20 (MO), 2001 May 16 , 6Jn20 (NMC), 2008 Jun 16 , 180 .J lxne 862A Alexander J. P. oceavillosa Boechera 298 .D Windham D. M. 022a 124a ,31May2004 ,23Apr2005 ,20Jun2008 ,18Jun2002 S Watson) (S. Windham Borodinia (Graham) Boechera Boechera 5Apr 25 , (Rollins) 023a , Delivered by Publishing Technology to: New Mexico State University IP: 128.123.253.108 on: Mon, 18 Mar 2013 00:39:19 Copyright (c) American Society for Plant Taxonomists. All rights reserved. ftejnto fU .Hy7 rassHy8o h ieof side E the on 8 Hwy Arkansas & miles 71 22.5 Hwy ± Oden, S. of U. W ft., miles of 770 2.8 Mena, junction Camp ± the River, Float Ouachita of the Creek E of Shirley bank N Forest, JX146747, the National on Beilstein], Ouachita M. Co.: JX146183, Montgomery JX146277, by JX146665. JX146369, provided JX146460, JX146088, sample Swan JX146552, creek, [DNA of JX146936, the banks JX146841, (MO) crosses the 2001 125 Along route Co.: May Christian state Missouri: where A.: Creek S. U. Windham, eune X494 X499 X465 X450 X449 JX146338, JX146429, JX146520, JX146615, JX146999, JX146904, sequence, hbrvk ertr:An-ak ofuneo h oe n Odola River, and Borea Omnya the with of Rivers confluence Ayno-Maik, JX146121, Territory: Khabarovsky JX146216, JX146309, JX146401, JX146697. JX146492, JX146585, JX146969, X404 X460 X455 X444 X432 X421 JX146156, JX146251, JX146342, JX146434, JX146720. JX146525, JX146620, JX147004, 03 LXNE TA. OCEAPYOEEISADTXNM 209 TAXONOMY AND PHYLOGENETICS BOECHERA AL.: ET ALEXANDER ft., 1,185 CR 48330, on on 8 miles Arkansas 1.2 of E S 48320, miles of 3.5 miles ca. 1.5 Mena, W of 61, SE miles 12 Forest, National canadensis 2013] lios ilC. DTUS ot 1PoetAe;buf fteKaskaskia the of bluffs Area; Project ft., JX146433, 51 500 Route River, U.S. JX146524, IDOT Co.: JX146619, Will JX146719. Illinois: JX146155, JX147003, JX146250, JX146908, JX146341, ft., 560 north JX146813, west, mile 1.5 mile 113, (ILLS), 0.5 Route then along Road, 113) Zlm & on 53 east Route miles of 3 (junction Kankakee Braidwood Preserve; Co.: Nature of Fayette Prairie Illinois: A.: Shrub Wilmington S. Watershed, U. River Windham, & Alexander J. P. (Raf.) JX146494, JX146699. JX146587, JX146123, JX146971, JX146218, JX146876, JX146311, JX146782, JX146403, (UARK), 2005 May 31 095 .S . rass okC. uciaMutis Ouachita Mountains, Ouachita Co.: Polk Arkansas: A.: S. U. , 369 .R hlip 39455 Phillippe R. L. .J lxne 722 Alexander J. P. ooii laevigata Borodinia ooii macrophylla Borodinia 284 ooii laevigata Borodinia .Y aklvs n. s. Barkalov Y. V. 6Mr20 NC,J168,JX146874, JX146780, (NMC), 2008 Mar 26 , 4Ar20 IL) X484 JX146909, JX146814, (ILLS), 2008 Apr 24 , Mh.e il. .J lxne & Alexander J. P. Willd.) ex (Muhl. 404 ...Mrh 2515 Murphy M.J.C. Trz)O .Shl,Russia: Schulz, E. O. (Turcz.) ooii dentata Borodinia .S . Arkansas: A.: S. U. , 5Jl18 M) no (MO), 1989 Jul 25 , .Bisen2001-06 Beilstein M. 403 ooii dentata Borodinia .MEdry1097 McElderry R. ,2May2007 ,U.S.A.: ,21 , X426 X411 osequence. no JX146151, JX146246, .J lxne ida,DAsml rvddb .Schranz, E. by JX146304, provided JX146395, JX146692. sample JX146486, JX146115, DNA JX146579, JX146210, Windham, JX146963, JX146868, & JX146774, Alexander J. P. G) X476 X480 X495 X451 X449 JX146368, JX146459, JX146551, JX146935, barren JX146664. JX146087, JX146840, Lewis, shale JX146182, JX146276, Fort open of JX146746, steep, east of miles Base (GH), 0.8 Co.: Hill, Bath Ratcliff Virginia: at A.: S. the JX146215, U. on Windham, JX146308, 8 Hwy JX146400, JX146696. Arkansas JX146491, & JX146120, 71 JX146584, Oden, Hwy JX146968, of S. ft., W U. JX146873, 770 miles of Mena, 2.8 junction of ± the side Creek River, of E Ouachita Shirley E the miles of Forest, 22.5 bank ± N National the on Ouachita Camp Float Co.: Montgomery Arkansas: X488 X493 X459 X446 X445 X433 JX146220, JX146313, JX146701. JX146405, JX146496, JX146125, JX146589, JX146973, JX146878, repanda B. ftesmi fM.Eesn atsoeo h iraNvd,Inyo Nevada, Sierra the of slope Sabrina SE of east miles Lake 3.3 locality Emerson, ± of [type dam, Mt. shore Forest the of of National East south summit miles Co.: 0.2 the ± Inyo of Trail, Sabrina California: Lake A.: the along S. U. Windham, X488 X493 X469 X456 X445 X432 JX146240, JX146332, at JX146425, JX146714. JX146516, pool JX146145, JX146609, swimming of of JX146993, & locality N JX146898, [type restaurants miles ft., Park the 4,800 0.2 of Watson], National of Trail S. side Yosemite SSW Ledge south Village, former miles Nevada, Curry the 0.3 Sierra of Point, central part Glacier the lower of Valley, slope Yosemite West Co.: Mariposa Yosemitea ,920ft., 9,200 ], . 308b oeie repanda Yosemitea .J lxne 1031A Alexander J. P. 094 .J lxne 845B Alexander J. P. 357a .J lxne 721 Alexander J. P. ooii serotina Borodinia oeie repanda Yosemitea 283a rbsinamoena Arabis 245 ooii missouriensis Borodinia ooii missouriensis Borodinia S asn .J lxne & Alexander J. P. Watson) (S. 4My20 NC,JX146804, (NMC), 2009 May 24 , 6Mr20 NC,JX146779, (NMC), 2008 Mar 26 , u 08(M) JX146784, (NMC), 2008 Jul 6 , .Kee 3710 Keener C. Sel)P .Aeadr& Alexander J. P. (Steele) .S . California: A.: S. U. , ree yoy of synonym a Greene, rbsrepanda Arabis ,14Aug1982 ,U.S.A.: (Greene)