Delivered by Publishing Technology to: Michael Simpson IP: 174.101.35.52 on: Wed, 11 Jul 2012 04:29:57 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 90 ontn12;Shazr20;Tile 09 Zuloaga 2009; Teillier South 2007; Schwarzer western 1927; (Reiche in Argentina Johnston to 1910; east occur and Bolivia, Chile, species Peru, in 1971, America, 70 Higgins coast 1927; Approximately Payson Pacific 1925; the 1979). (Johnston distributions Texas from with to and east Mexico, America, and southern North to Alaska western from in occurs spe- cies, 130 approximately diversity, greatest The 2011). Simpson treatments, delimited most currently As in regions. tropical of intervening but regions America, in South desert absent western and or America temperate North western mostly both in taxa with tributed, nlsv fsvrlsgeaegnr;w ee oti genus this to refer we as genera; concept segregate several of inclusive O 10.1600/036364412X648706 DOI © Botany Systematic –oe vr ihagnbscsye n ri hti a is that fruit a 2001). and al. et style, (Gottschling nutlets gynobasic of schizocarp a deeply with a 2003), ovary Hilger and 4–lobed (Buys cyme scorpioid circinate, that a inflorescence is an by characterized group a Boraginoideae, onwards). 2001 divided Ehretioideae, (Stevens be Lennoideae Cordioideae, classification and Hydrophylloideae, Heliotropoideae, may Boraginoideae, of which system subfamilies Boraginaceae, (2009) into III broad APG a subgroups. the narrow recognizing accept of or arrangements we III broad different Here APG recognizing with 2007; concepts authors al. family various et Heywood with 2001; Ferguson 2009), family al. 1897; Prantl et the and Gottschling of (Engler 1999; years genus the changed a over has repeatedly family is this flower,” of circumscription “popcorn The Boraginaceae. or eye” “cat’s oyih 02b h mrcnSceyo ln Taxonomists Plant of Society American the by 2012 Copyright ic h 1920s, the Since Cryptantha 1 orotru aawr eune o w eergos h ula iooa ee T,adthe and ITS, gene, ribosomal nuclear the regions, gene two for sequenced were taxa outgroup four ihnsbrb rpatia.Tegnsehbt napirpcdsrbto,ocrigi eprt n eetrgoso etr North western of regions desert and temperate of in species occurring 45 of distribution, samples amphitropic Fifty an America. exhibits South and genus The Cryptanthinae. subtribe within eoe hs aawr sdt sespyoeei eainhp sn asmn,mxmmlklho,adBysa neec methods. genera inference related Bayesian The genera and the likelihood, of maximum resurrection parsimony, propose using relationships phylogenetic assess to Cryptantha used were data These genome. eeal uprstehptei frpae,uiietoa ipra vns rmNrht ot mrc.Gnr eurce include: resurrected Genera America. South to North atwoodii from Oreocarya events, in reduction dispersal study similis, this perenniality, unidirectional incomplete, of repeated, is evolution sampling of Although derived Eremocarya hypothesis cleistogamy. multiple, and the the heterostyly, supports sculpturing, support nutlet generally analyses smooth trait heteromorphism, nutlet Character number, monophyletic. nutlet as supported is Cryptanthinae Plagiobothrys ontnlahlpea ontnlamcoee,Jhsoel parviflora, Johnstonella micromeres, Johnstonella holoptera, Johnstonella fulvocanescens cioeaa ontnlafsiit,Jhsoel grayi Johnstonella fastigiata, Johnstonella echinosepala, welshii Oreocarya suffruticosa schoolcraftii Oreocarya urn drs:Rnh at n oai adnadCaeotGaut nvriy 50NrhCleeAvenue, College North 1500 University, Graduate Claremont and Garden Botanic Ana Santa Rancho address: Current Abstract— Keywords— Cryptantha 21) 73:p.738–757 pp. 37(3): (2012), , a’ ysadPponFoes hlgntcSseaiso h Genus the of Systematics Phylogenetic Flowers: Popcorn and Eyes Cat’s .l a on ob oyhltc ihismmespae mn eea elspotdcae.Bsdo hs nlss we analyses, these on Based clades. well-supported several among placed members its with polyphyletic, be to found was l. s. emn xG o,cmol nw as known commonly Don, G. ex Lehmann Greeneocharis var. a eovda oyhltci he lds hs ldscrepnigt rvosynmdscin rgop fscin.The sections. of groups or sections named previously to corresponding clades these clades, three in polyphyletic as resolved was Cryptantha Cryptantha Amsinckia var. Cryptantha eateto ilg,SnDeoSaeUiest,SnDeo aiona912 .S A. S. U. 92182, California Diego, San University, State Diego San Biology, of Department rncl,Oecrasuffruticosa Oreocarya arenicola, Cryptantha Amsinckia , iia roay grahamii Oreocarya nitida, .l h eu sapirpclydis- amphitropically is genus The l. s. ontnlaaglc,Jhsoel nutfla ontnlacsaa ontnladporca Johnstonella diplotricha, Johnstonella costata, Johnstonella angustifolia, Johnstonella angelica, Johnstonella Brgnca)i ru fapoiaey20ana n eena pce,rpeetn w-hrso h diversity the of two-thirds representing species, perennial and annual 200 approximately of group a is (Boraginaceae) biogeography, , , , Johnstonella, roay eilba roay hclten,Oecrasblfr,Oecrasbaiaa Oreocarya subcapitata, Oreocarya sobolifera, Oreocarya shackletteana, Oreocarya semiglabra, Oreocarya imyblnswti subfamily within belongs firmly a encrusrbdbroadly, circumscribed been has and rse .Hasenstab-Lehman E. Kristen , .l a bu 0 pce (see species 200 about has l. s. roay anbi roay opca roay rsie,Oecracetflti Oreocarya creutzfeldtii, Oreocarya crassipes, Oreocarya compacta, Oreocarya barnebyi, Oreocarya 3 Pectocarya uhrfrcrepnec ([email protected]) correspondence for Author Eremocarya 2 uhrfrcrepnec ([email protected]) correspondence for Author and etcra Plagiobothrys, Pectocarya, Oreocarya. eersle smnpyei n otcoeyrltdt aiu ldswithin clades various to related closely most and monophyletic as resolved were Cryptantha , Greeneocharis , omnctn dtr et Klitgaard Bente Editor: Communicating lrmn,Clfri 11,U .A. S. U. 91711, California Claremont, roay yspia roay ontni roay ochroleuca Oreocarya johnstonii, Oreocarya hypsophila, Oreocarya Cryptantha var. e obntosinclude: combinations New aa roay suffruticosa Oreocarya laxa, var. , .l,eepaso h eae genera related the of exemplars l., s. Johnstonella rpohea ontnlagrayi Johnstonella cryptochaeta, .l (Boraginaceae) l. s. aooy ri evolution. trait taxonomy, 738 1,2 ,and ta.20) he species, Three maritima C. 2008). al. et uigadclr h tahetsa sgnrlyashallow, a generally is scar attachment sculp- The in color. variable is and strongly wall turing pericarp is smaller, The that gynobase. three nutlet the to larger often adnate one are and nutlets, there size detached in these, easily heteromorphic conti- gynobase in are both sculpturing; that the on nutlets and/or to species with fruits Several attached have dispersal. remain nents at or calyx the from within and detach style shape can in a lanceolate and to which ovate generally narrowly-pyramidal are from Nutlets to arise. gynobase) stigma attached (the are tissue these receptacular four; to one from throat. corolla (invag- the are fornices are surrounding and yellow Corollas regions) often species) folded wide. five, inated, few with cm a salverform, two in to to (yellow rotate mm white one universally in than almost ranges less limb corolla from The size 1983). (Grau structures lenticular cleistoga- subgenera be also of Geocarya may members but in Flowers taxa, mous features. most diagnostic in as chasmogamous used are accrescent, often the are 2008; of shape, calyx orientation The fruiting Mabberley and 2012). vestiture, al. 1993, fusion, et 1968; of Kelley degree Wilken 1959, 2009; leaves Hasenstab and Keck and and Simpson Kelley oblanceolate Munz 1971; or 1927; Higgins Payson lanceolate, 1925; cau- branched, linear, to (Johnston highly basal simple, generally to and simple stems line, erect with to ascending herbs, generally perennial or nial, and n ihe .Simpson G. Michael and h ubro ult htdvlpt auiycnvary can maturity to develop that nutlets of number The bien- annual, hispid, to strigose are genus the of Members Oreocarya ontnlapusilla. Johnstonella lnso h atbaigfut oiidinto modified fruits bearing last the of plants , reecai circumscissa Greeneocharis r itiue nbt ot n ot America. South and North both in distributed are , n eonto fanwydelimited newly a of recognition and , var. utls,Oecrasuffruticosa Oreocarya pustulosa, var. mici,Pectocarya, Amsinckia, trnL ry,Jhsoel grayi Johnstonella grayi, UAA .albida C. nrnrgo ftechloroplast the of region intron 3 var. rpata Krynitzkia Cryptantha, ouaa Greeneocharis rosulata, , roay roosiorum Oreocarya and , .circumscissa C. Plagiobothrys Cryptantha Cryptantha var. var. nesiotica, setosa, ,and .s. s. .l. s. ,and and , , Delivered by Publishing Technology to: Michael Simpson IP: 174.101.35.52 on: Wed, 11 Jul 2012 04:29:57 Copyright (c) American Society for Plant Taxonomists. All rights reserved. .leiocarpa the K. described subsequently (1841) genus Meyer his- nomenclatural and for Fischer 1935 1925, tory). Johnston (see (1837) published Don validly but by (1836) Meyer and Fischer by earlier cited h eu nascainwt w ot mrcnspecies American South two with for association ( diagnosis in formal first genus the the provided who (1837), hetero- Don the with in placed genus subsequently geneous or originally were many genera of species some hrceie yhtrclaeple seHrrv and Hargrove evaluate to of limited relationships too (see was sample taxon pollen Their 2003). Simpson heterocolpate by and Cynoglosseae which tribe characterized expanded an to in corresponding clade a Eritrichieae, treatment their in abandoned; tribe be should the that Cryptantha of concluded species authors one included ( Boraginoideae, the of iswithin cies La t 1997). classified r Takhtajan however, (1925), 1991; iBrand Al-Shehbaz b 1873; scar Hooker attachment eC and nutlet by submedial (Bentham r a characterized and gynobase, y elongate tribe pyrimidal an to p 2003), the Simpson and t (Hargrove treatments, pollen aheterocolpate taxonomic n most tin h e a eo rs u b t r i b eC r y p t a n t h i n a e( B r a n d1 9 3 1 ) . Cryptantha publication. original of place and placed ultmrhlg stems motn aooi criterion 2009). Hasenstab taxonomic and important (Simpson most the to 739 Higgins is used morphology 1927; nutlet are 1925, features (Johnston among floral whole distinguish and a vegetative as Numerous genus 1971). diagnos- the deemed the was for that near groove tic or ventral the to is CRYPTANTHA extends It OF that apex. SYSTEMATICS nutlet groove PHYLOGENETIC ventral SIMPSON: a is AND which with HASENSTAB-LEHMAN elevated), continuous or rimmed not (generally areole triangular 2012] Eritrichium Eremocarya Echinosperum Krynitzkia Johnstonella Hemisphaerocarya Greeneocharis Wheelerella Rochelia Piptocalyx Oreocarya Myosotis Lithospermum .glomerata C. .virgata C. ˚ al rsnstevrosgnrcnmsapidt spe- to applied names generic various the presents 1 Table Cryptantha Table ngstro xG Don G. ex xGaudin ex .A Mey. A. C. ¨ Kryntizkia Myosotis c.Foa7 4.1824. 243. 7: Flora Rch. .Gnr ihnwhich within Genera 1. .SeisPatrm1 3.1753. 131. 1: Plantarum Species L. n hs 20) nasuyeautn h tribes the evaluating study a in (2002), Chase and m reePtoi :5.1887. 57. 1: Pittonia Greene ic.& Fisch. or,ie.Uie ttsEpoigExpedition Exploring States United ined. Torr., Lehm. Schrad. .B rn,ie.Blei fteSuhr California Southern the of Bulletin ined. Grant, B. G. ae rnfre to transferred later , reePtoi :5.1887. 58. 1: Pittonia Greene ,ofsection Cryptantha a sal enpae,i o oohltcand monophyletic not is placed, been usually has rn eetru pceu Novarum Specierum Repertorium Brand Gu w xLh.P.Aprf ui.1 1.1818. 113. 1: Nucif. Asperif. Pl. Lehm. ex Sw. .SeisPatrm1 3.1753. 132. 1: Plantarum Species L. .l a encasfe ntetieEritrichieae tribe the in classified been has l. s. and ¨ k am Natu Harms & rke Cryptantha Eritrichium rn eetru pceu Novarum Specierum Repertorium Brand , Cryptantha Lithospermum .microcarpa C. o igeNrhAeia species, American North single a for Cryptantha .l,a urnl icmcie.Although circumscribed. currently as l., s. Oreocarya oohrgenera. other to h name The . .l eeoiial lcdi the in placed originally were l. s. eea itr fteDichlamydeous the of History General A lr evtc :4 7 1828. 57. 4, 2: Helvetica Flora ne eiu S.Petersburg] [St. Seminum Index Cryptantha lns41:33 1837. 373. 4(1): Plants en eeaii 4 9 1927. 59. 24: Vegetabilis Regni 1899. 460. Regist. Prantl] & [Engler en eeaii 1 4.1925. 249. 21: Vegetabilis Regni cdm fSine :2.1906. 28. 5: Sciences of Academy 72:43 .1.1874. 12. t. 413, 17(2): 1841. 52. 7: ¨ Cryptantha , pce n nrseis but infraspecies, and species .Teeseisnmswere names species These ). lce Pflanzenfamilien rlichen Cryptantha Echinospermum, nteraayi.These analysis. their in ) Cryptantha Cryptantha .l pce aebeen have species l. s. sebelow). (see samme fthe of member a as or Cryptantha originated eog to belongs Rochelia, ela itnusigcaatro h genus the of character (including distinguishing a as keel of representatives American of ooy of homonym genus the recognized into igorpi agso hs etos lo rn (1927) Brand genus the Also, of recognition sections. proposed these of ranges in biogeographic sections four recognized Payson Johnston Cryptantha section with monograph of agreed this series), taxa six the recognized that of he monograph which (in a in (1927), North Cryptantha perennial, the that American concluded Johnston studies, these 18) h lcdNrhAeia pce htwudlater would that species become American North placed who (1885), and recognizing Geocarya, Boraginaceae, Krynitzkia, American South Cryptantha of published treatments (1927) Johnston Subsequently, series. 15 into placed 18b rae w e eeafo h pce of species the from genera Greene new Additionally, two created cymes. (1887b) dichotomous and pedicels tent segregating 1), Table Cryptantha below]; see Harms; & icuieof (inclusive include to and genus this of cumscribed species American North authors) of subsequent perennial by the annual as from but perennial, as Brand by perennial the genus the previously been in had placed that species genus American North the numerous expanded calyces, greatly (1887c) persistent Greene Subsequently, 1). (Table inflorescences, styles persistent enlarged, and dichotomous and teate, and leaves, radical Oreocarya ilb icse ute eo ntecneto u phyloge- our of context results. the netic in below further discussed be will Greeneocharis Gu latter. ao a rae yalsbeun uhr sete a either as authors subsequent genus this all However, by taxa. Cryptantha treated these was dis- in the present taxon of nutlets because globular species associated tinct, closely four and 1984] pass ae eonzda a as recognized later circumscissa G. etoso ontn(97 n asn(97 osubgenera. to (1927) Payson and (1927) Johnston of sections of inclusion pce of species xeto h eena aa ulse eie monograph revised a another published of taxa, (1971), perennial the Higgins on unchanged. expert largely remained have imnln ftegenus the of Dismantling rn 12)rcgie h genus the recognized (1925) Brand ic h 90,txnmsshv ecie eea new several described have taxonomists 1920s, the Since Oreocarya, Cryptantha Piptocalyx Krynitzkia Oreocarya. rpatacinerea Cryptantha Oreocarya Oreocarya ¨ Cryptantha k n am 19)dsrbdytaohrgenus, another yet described (1899) Harms and rke ossigo inilo eena eb with herbs perennial or biennial of consisting , eto rsubgenus or section , Cryptantha : Allocarya Oreocarya ae ntepeec ntefre fpersis- of former the in presence the on based ofr n ag,hmgnu eu.Payson genus. homogenous large, one form to Eucryptantha shvn he etosi ot America: South in sections three having as Oreocarya Piptocalyx [ n gedwt ontnadPyo nthe on Payson and Johnston with agreed and .racemosa C. Tbe1.I oorpi ramn fthe of treatment monographic a In 1). (Table [= ree(87)frhrrfndtetreatment the refined further (1887a) Greene . Wheelerella rmcra Greeneocharis Eremocarya, taserdfrom (transferred sebelow). (see hudb obndwith combined be should al rvdsdansi etrsand features diagnostic provides 2 Table Greeneocharis Eremocarya and ree from Greene) Oreocarya and .l,bttescin fPyo (1927) Payson of sections the but l., s. pce hudas eicue in included be also should species Krynitzkia Piptocalyx within lv xBnh [= Benth. ex Oliv. [ Cryptantha .jamesii C. nd] ossigo n species, one of consisting ined.], , Plagiobothrys and [= Eucryptantha Eritrichium Cryptantha Cryptantha ,wt i 7rcgie species recognized 57 his with ], .inequata C. itnuse ydne brac- dense, by distinguished , Cryptantha Eritrichium n h eann members remaining the and Oreocarya setal ibnigthe disbanding essentially , Lithospermum nd;seCoqite al. et Cronquist see ined.; , or a leiiaelater illegitimate (an Torr. Hemisphaerocarya hs aooi issues taxonomic These . Krynitzkia. rmcra Johnstonella, Eremocarya, Johnstonella ic.&C .Meyer A. C. & Fisch. Cryptantha ], ontn(95 cir- (1925) Johnston , a ntae yGray by initiated was [= ramme fthe of member a or , telte described latter (the ,and ecra Krynitzkia Geocarya, lvtn h four the elevating , Greeneocharis oigteventral the noting , Cryptantha Piptocalyx ree(1887b) Greene Cryptantha. .,wihwas which L.), Johnstonella Plagiobothrys osegregate to oinclude to Krynitzkia oencom- to Oreocarya .From ]. Gu from ¨ rke In ), : Delivered by Publishing Technology to: Michael Simpson IP: 174.101.35.52 on: Wed, 11 Jul 2012 04:29:57 Copyright (c) American Society for Plant Taxonomists. All rights reserved. etdwti agrCngosa .l a o e aaof taxa complex. few the too in Far 2007) l. Schwarzer s. Cynoglosseae Cryptantha of larger a Cryptantheae within nested tribe to clade well-supported (corresponding a form together and monophyletic of each species two included Plagiobothrys La al. to et Weigend similar results using obtained Boraginoideae the in ships 2007). (Schwarzer species one only by represented is it section and polytomy, unresolved Section clade. supported section section of species four other the to 1 pce) Section species). (10 monophyle mostly Plagiobothrys largely a are species), interrelationships incl but whose unresolved, taxa, remaining the Cryptantha unresolved larger, the within nested and monophyletic a shows tree This Pectocarya species. outgroup Cryptantheae tribe two of species and 34 of cladogram a presents using data, analysis own his from 1925). (2007), (Brand Schwarzer Cryptantheae tribe as to referred group, phyletic Pectocarya, eena rbena eb,pouigol hsoaosflowers. chasmogamous only producing herbs, biennial or Perennial Oreocarya North in Distributed flowers. chasmogamous only having plants Annual Krynitzkia with flowers cleistogamous Basal flowers. cleistogamous with Plants Geocarya to similar flowers Cleistogamous flowers. cleistogamous with Plants Eucryptantha range. biogeographic and features diagnostic their and (1927), Payson ra es ( sense broad uulytetda eto of section a Cryptantha as treated (usually whic in data, cpDNA intron 37 [Volume circumscribing in Higgins Johnston, and of treatments Payson, the follow classifications recent Most BOTANY SYSTEMATIC 740 rm(ie sH .Hilger H. H. as (cited gram and Meisn., ex DC. genera the of members analyses. phylogenetic on based not were suggestions the of lineage speculated diverging of both species (1971) perennial the Higgins that and (1927) Payson complex. in rsbeea oee,torltvl eetfloras recent from relatively recognized separate 1987) two Weber However, 1951; (Abrams subgenera. or tions ial,Wiede l 21) nasuyo rblrelation- tribal of study a in (2010), al. et Weigend Finally, Table h israino cwre 20)fcsdo h Peruvian the on focused (2007) Schwarzer of dissertation The this on published been have studies phylogenetic Few etitdt ot America. North to Restricted America. South and to Restricted America. structures. South lenticular into specialized highly fruits the America. throughout South often to and Restricted leaves inflorescence. of axils in Cleistogamous located corolla. flowers closed for except flowers, chasmogamic .Scin of Sections 2. Geocarya (including fv pce,with species, (five and .s., s. eeue oeaut hlgntcrelationships phylogenetic evaluate to used were .l,section l., s. [ = n n of one and , Cryptantha 1 pce) n nunresolved an and species), (15 Cryptantha Cryptantha. Plagiobothrys Eremocarya toseis,tetofrigawell- a forming two the species), (two Eremocarya Plagiobothrys Allocarya Cryptantha dn monophyletic a uding Amsinckia ] Cryptantha .l) secmasn l orsec- four all encompassing as l.), s. Cryptantha Amsinckia Krynitzkia , ebr ftegenera the of members h .s omawl-upre mono- well-supported a form s. s. nulse)bsdon based unpublished) , Oreocarya Harpagonella .setosa P. , of .l eonzdb ontn(97 and (1927) Johnston by recognized l. s. Greeneocharis ˚ .Thisworkincludesaclado- ngstro Plagiobothrys Lehm., i u atyunresolved partly but tic Oreocarya trnL-F lgoohy) Amsinckia, Plagiobothrys), toseis ssse to sister is species) (two hs he eeaare genera three These . .l ope,btthese but complex, l. s. fv pce)frsan forms species) (five fsection of ¨ osiue h earliest the constituted n hs (2002). Chase and m Pectocarya Cryptantha Oreocarya Plagiobothrys .Gray, A. pN sequences, cpDNA ,and Cryptantha, Cryptantha sursle,as unresolved, is Amsinckia smonophyletic is Johnstonella trnS-G Gruvelia )sisterto , sagenus a as Cryptantha Oreocarya, Pectocarya Allocarya .Within cpDNA trnL w of two nthe in sister (four ) UAA ei eainhp fmmeso h genus the of members of relationships netic of comprised genera clade the a and this to whether corresponds test We group 1931). of (Brand expanded Cryptanthinae Cryptantheae subtribe an to tribe recognize recognize we the 2007), (2010) of Schwarzer (inclusive al. Cynoglosseae et tribe Weigend and (2002) fCytnhne(sdfndhr)wr sampled: were here) cies), defined (as At Cryptanthinae collected. section, of recently recognized been currently each not Krinitzkia of had representative that one her- species supplement least to for used specimens were collections barium field Vouchered 1). (Appendix oro iercgie etos.W eeual osqec h mono- the sequence to unable were We specific sections). recognized five of four Museo (SDSU), University Natural State Diego Diego San (RSA), San Garden (SD), Bota Botanic Museum Ana Museum History Colorado Santa of Rancho University (MO), the at Concepcio de herbaria Universidad (COLO), from loan on material oo DyeadDye18;Fir20) mlfctoswr oein done were Amplifications 2005). Friar 1987; Doyle pro- 25 bromide) and ammonium (Doyle trimethyl (cetyl tocol CTAB the of version modified 1). (Appendix of species of each Myosotis species one tion, aafo the from data Plagiobothrys genera similar morphologically oass igorpi itr ihrsett h amphi- the to respect of and with distribution that studies history tropic features taxonomic biogeographic past of assess in evolution to used assess been to traditionally is used have phylogeny is Additionally, of phylogeny complex. this estimate this of best classification the The revise to evaluated. used these also encompassing are are clades taxa complex the the among in Relationships sections tested. and subgenera, genera, ognized of monophyly of The Cryptanthinae, regions. the ITS ribosomal nuclear and locus, ottete.Alsqecsaeoiia oti td,ecp o the for except study, this to original are sequences of All sequences tree. the root r osdrdabgosyaind nlssicuigadexcluding and including thus Analyses and aligned. homology, was ambiguously positional alignment Some assign considered to 2005). final are difficult Maddison A were and alignments 15. ITS (Maddison within and two 4.08 regions 2 MacClade 1997); of in al. manually costs chromatograms Sequences done et gap-opening sequences. using with (Thompson done ITS ClustalW 2004) were the using Groothuis aligned especially were polymorphisms, and for (Griekspoor checked 1.7 version primers same an the on using or Garden Facility Botanic amplification. for Ana Core as Santa Microchemical Rancho 3100 University at was ABI 3130xl State an pellet ABI Diego on The done San rpm. was the Sequencing 14,000 at ethanol. at 80% minutes ice-cold with 15 washed for centrifugation by pelleted lcl80 PG n25MNC,uigeulvlm fPGt PCR to PEG of volume 37 equal at using polyethylene incubated NaCl, 20% was M mixture with 2.5 the precipitated in product, were (PEG) 8000 amplicons thermal 2720 glycol PCR Biosystems Applied The an on cycler. out carried were Amplifications iduigtefloigpies trnL5 primers: following the using fied T5 n T4(or ta.20) The 2006). al. et (Moore ITS4 and ITS5a 1–10 of total 1.5 n ees rmr,1.25 primers, reverse and ln Samples— Plant m Sequencing— and Isolation DNA euneEiigadAlignment— and Editing Sequence h rmr betv fti td st sesphyloge- assess to is study this of objective primary The ie httercn tde fLa of studies recent the that Given m ´ oue otiig 2.5 containing: volumes l io(I,adteUiest fClfri-ekly(UC-JEPS) California-Berkeley of University the and (SI), nico gl,0.125 MgCl2, M Pectocarya Harpagonella , eeicue ots oohl fteCytnhne Two Cryptanthinae. the of monophyly test to included were Cryptantha Phacelia m Myosotis /lgnmcDA h T einwsapiidusing amplified was region ITS The DNA. genomic g/ml fteCytnhneuigmlclrsequence molecular using Cryptanthinae the of treseis,and species), (three trnL elotiesbaiyBrgniee eeue to used were Boraginoideae, subfamily outside well , mici,Pectocarya, Amsinckia, pcmn eeotie rmdidherbarium dried from obtained were Specimens hc suulyicue nti ope.I addi- In complex. this in included usually is which , aeil n Methods and Materials m and ,0 U/ml 5,000 l and UAA Cryptantha Cynoglossum Geocarya m Cryptantha Phacelia, 200 l hools nrnpr fthe of part intron chloroplast m l10 m ´ eesmld diinlmembers Additional sampled. were , NPs 16.375 dNTP’s, M CN) isuiBtnclGarden Botanical Missouri (CONC), n Taq N a sltduigathree-day, a using isolated was DNA hc eeotie rmGenBank from obtained were which eune eeeie sn 4peaks using edited were Sequences + Plagiobothrys Cryptantha .l. s. oyeae 1.2 polymerase, trnL tnadM-rebfe,1.25 buffer, Mg-free standard 0 and .l,ado rvosyrec- previously of and l., s. mici,Pectocarya, Amsinckia, n trnF and UAA o 5mnts N was DNA minutes. 15 for C Hackelia and ˚ nrnrgo a ampli- was region intron ngstro GAA Plagiobothrys. svnseisttl in total, species (seven .l sbelonging as l. s. Amsinckia n w pce of species two and m Sa ta.2005). al. et (Shaw ¨ Cryptantha Cryptantha lH m n Chase and m l10 2 ,ad1.0 and O, m forward M fu spe- (four Oreocarya trnL-F and and .l. s. m m l l , Delivered by Publishing Technology to: Michael Simpson IP: 174.101.35.52 on: Wed, 11 Jul 2012 04:29:57 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 02 AESA-EMNADSMSN HLGNTCSSEAISO RPATA741 CRYPTANTHA OF SYSTEMATICS PHYLOGENETIC SIMPSON: AND HASENSTAB-LEHMAN 2012] oa f145ncetds ih34prmn informative parimony 354 with nucleotides, 1,475 of total HesnekadRnus 01,wt h T and ITS the with 2001), Ronquist and (Huelsenbeck set. data was combined analysis a likelihood on final done the so recovered, were incongruences ported ob togyspotd ogunebtenISand considered ITS were between greater maximum Congruence or a supported. 70 with strongly of clades be value All to (MPBS) pseudo-replicates replicate. bootstrap per bootstrap parsimony random-additions non-parametric 10 1,000 and with using analyses ITS parsimony the assessed the (Swofford in on was support PAUP* done Branch in independently. initially data test were sequence partition-homogeneity (ILD) analyses conflicts difference a Parsimony test length using 2003). incongruence To done data, data. was and plastid missing unordered test and as as ITS treated treated the were were between Gaps Characters trees optimal weighted. replicate. bisection-reconnection All step. each equally each tree at at trees held with best 10,000 the were addition all parameters: on swapping taxon following branch (TBR) random the Parsi- with of criterion. conducted replicates optimality were parsimony analyses a to mony utilizing submitted 2003) (Swofford were 4.0b6 data aligned were final, they 11794). The number so (study analyses. topology, TreeBASE in final change in no included revealed regions ambiguous these a u sasprt rcs atto rmteISdata. locus ITS plastid the from the partition process analysis, separate a combined as a the run for was combined In were analysis. data the concatenated so final, analysis, either or in 0.95 recovered of (PP) were (PP) probabilities probability greater posterior posterior of a with with deviation clades 0.01, conflicting standard of No calculated. the value after a reached sampled frequencies trees split Consensus from generations. produced 100 were for every ran trees analyses sampling Bayesian All with analyses. generations, Bayesian + 10,000,000 all I for + used TMV was was which locus G, each 2004). for (Nylander selected substitution Mr.Modeltest nucleotide of in model The (AIC) using criterion determined information were Akaike dataset the each for Models independently. analyzed 0 elctsudrteGR+I+Gmdlo vlto.TeISregion ITS The evolution. of model from G estimated + I values + RAxML support and GTR the bootstrap in under with implemented replicates 2008), 100 3.7 analysis al. version (MLBS) et max- Modeltest a (Stamatakis bootstrap with (AIC; from assessed likelihood was nucleotide criterion 2001) support imum Statistical of information Crandall 1998). Crandall model and Akaike and (Posada Posada The the 1974; using 2003). Akaike selected (Swofford imple- was 4.0b10 was evolution 1981) PAUP* Felsenstein in (ML; mented criterion optimality likelihood the mum on illustrated are greater or the 70 tree. consensus Maxi- of of strict above. values analysis parameters bootstrap the bootstrap parsimony using mum calculated, A was from generated. set tree data was consensus concatenated strict analysis A they concatenated analysis. so concatenated sets, this data a two ILD for the combined the between were from incongruent as were support well strong as with data, plastid and The bootstrap ITS test. mapped the with from trees generated consensus values strict the comparing by assessed odifrnewsue o eosrcino ri evolution. trait likeli- of maximum reconstruction the for from used support topology was the statistical inference thus, in hood significant tree; conflict with consensus no MP clades the showed more than tree have both likelihood and maximum Bayes- topology, the and Parsimony of tree topology consensus The America). assumption. ian states Maddison unordered and South the (Maddison using Mesquite 2010), versus in bio- implemented (North were for optimizations and distribution presence/absence), (both geographic cleistogamy charac- additional and two heterostyly for optimization ridges.ters, parsimony and/or the examined turbercles, also no papillae, with We having smooth, sculpturing: rough, nutlet nutlet elements; 4) one and sculpturing heteromorphic, shape; and/or sculpturing; size and in differing size nutlets in all with equal homomorphic, one, heteromorphism: approximately fruit: nutlet per 3) number four; nutlet binary, three, 2) two, perennial; were annual, analyzed duration: Characters characters plant states. four 1) All had were: which cir- fruit, groups. to per number infrageneric recon- used nutlet except been or Characters previously species have observation. that cumscribe personal features diagnostic and were structed literature taxonomic the aeinInference— Bayesian asmn Analysis— Parsimony ri Evolution— Trait aiu ieiodAnalysis— Likelihood Maximum euneMatrices— Sequence trnL UAA p au o h L etws094 ocae htwr recovered were that clades No 0.974. was test ILD the for value nrnwr ntal nlzdsprtl.N togysup- strongly No separately. analyzed initially were intron hrce ttsfrec ao eeotie from obtained were taxon each for states Character nlsswr oeuigMBYSvrin3.1.2 version MRBAYES using done were Analyses hlgntcaaye eedn sn PAUP* using done were analyses Phylogenetic h octntdmti otie a contained matrix concatenated The Results hlgntcifrneuigamaxi- a using inference Phylogenetic trnL UAA eunedata sequence trnL UAA trnL was UAA rpatia srcvrda oohltc(LS=8,P = PP 84, 1.0). = (MLBS monophyletic boot- as recovered likelihood is Cryptanthinae both with maximum here strapvaluesandBayesianposterio presented The is them. tree between incompatibility likelihood no topol- same with the virtually ogy, on converged analyses based model for 1). support (Fig. bootstrap relationship strong this without but species, sampled 23 analysis, Pectocarya cie,aepolyphyletic. are scribed, 0,P 1.0), = PP 100, the to sister is species, three of ples P=10 and 1.0) = PP supported robustly a (MLBS=86,PP=1.0),plustwootherscorrespondingto ednl smn sfv ie.Prnilpatdrto is duration the plant Perennial for is times. apomorphic five duration as plant inde- many annual arose as perenniality an pendently that that and shows Cryptanthinae for 3A seen ancestral Figure is topology 3. ML Fig. the in on characters morphological four of 2). (Fig. 1 s. s. (minus species of support. consisting strong without but other the group, of composed clade minus a to Cryptanthinae, sister are together three all and to sister are together two these hs w oehraesse to sister are four together of two clade these a to sister placed is Plagiobothrys species, two of samples four w aite,i itr(PS=7)to 70) = (MPBS sister is varieties, two support. .l aaaeifre,fu orsodn opeiul named previously to corresponding genera, four inferred, of are clades taxa supported strongly l. Six s. each). for 1.0 100, = = (MLBS PP support strong with monophyletic be to inferred The sampled. species endansi o hsgop eenaiyas evolved also Perenniality group. the this within for diagnostic been s iegn ld ihnCyptanthineae. within clade diverging est as designate we 97), whereas each). for polyphyletic 100 = are (MPBS phyletic circumscribed, Pectocarya currently consensus monophyletic as strict a 78). shows The = 1) 1,435. (MPBS (Fig. Cryptanthinae analysis of parsimony length the tree of tree a with trees parsi- 1,827 220 with nucleotides 616 characters. of informative total mony ITS a the contained and characters, matrix nucleo- informative parsimony 859 134 of with composed tides was matrix cpDNA The characters. .s MB 0) eainhp mn hs i rusare groups six these supported. among strongly Relationships not 100). = (MPBS 2 s. s. genera, 100), = named support, (MPBS bootstrap previously strong four with to all corresponding clades, major six in placed aiu ieiodadBysa Inference— Bayesian and Likelihood Maximum neta tt Reconstructions— State Ancestral asmn Inference— Parsimony Johnstonella Oreocarya Cryptantha Cryptantha Johnstonella Oreocarya Eremocarya steeris iegn ld.A nteparsimony the in As clade. diverging earliest the is and Johnstonella otiigegtsmldseis steearli- the is species, sampled eight containing , pce u ihu togbosrpsupport; bootstrap strong without but species Eremocarya MB 1,pu w eann lds which clades, remaining two plus 91), = (MPBS Greeneocharis reecai,Amsinckia, Greeneocharis, Cryptantha .hispidus P. Amsinckia Cryptantha .l and l. s. .s ssse oalrmiigCryptanthinae, remaining all to sister is 2 s. s. and MB 5 P=1.0), = PP 95, = (MLBS ihtresmlso n pce and species one of samples three with , Oreocarya Cryptantha Pectocarya Cryptantha Eremocarya ld,ocrigin occurring clade, ,adti ld ssse to sister is clade this and ), MB 0,P .) and 1.0), = PP 100, = (MLBS .s ld MB 8 P=1.0). = PP 98, = (MLBS clade 2 s. s. Amsinckia h asmn nlssyielded analysis parsimony The Plagiobothrys r togyspotda mono- as supported strongly are MB 100), = (MPBS .s MB 1 and 71) = (MPBS 1 s. s. Cryptantha lgoohy hispidus Plagiobothrys Johnstonella .jonesii P. ld;ti etr a long has feature this clade; .s ru,wt orsam- four with group, 2 s. s. rbblte hw Fg 2). (Fig. shown probabilities r P .9.Wti h last the Within 0.99). = (PP Cryptantha .s ld MB 96, = (MLBS clade 1 s. s. r itrtx MB 73); = (MLBS taxa sister are Cryptantha and asmn optimization Parsimony scretycircum- currently as , .l and l. s. Oreocarya Greeneocharis n all and lowtotrobust without also , Johnstonella Pectocarya ssse oaclade a to sister is Greeneocharis .racemosa C. .l pce are species l. s. .s ru of group 1 s. s. Plagiobothrys Plagiobothrys iheight with , Eremocarya (PP=0.97), Cryptantha Cryptantha Cryptantha r again are Oreocarya h two The MB = (MPBS MB = (MLBS with , and , Delivered by Publishing Technology to: Michael Simpson IP: 174.101.35.52 on: Wed, 11 Jul 2012 04:29:57 Copyright (c) American Society for Plant Taxonomists. All rights reserved. o the for Allo American. North rpatia Fg B,wt euto nnte number nutlet in reduction with 3B), (Fig. Cryptanthinae orsodn oteSuhAeia sections American clade South each in the and once twice, to most corresponding at evolved duration perennial cainlypeetin present occasionally 37 [Volume BOTANY SYSTEMATIC 742 ri ihfu ult e ri sceryacsrlfor ancestral clearly is fruit per nutlets four with fruit A Fig. = Geocarya Allocarya .Src osnu refo asmn nlsswt otta ausa elspotd( supported well at values bootstrap with analysis parsimony from tree consensus Strict 1. Johnstonella Fg 3A). (Fig. ; Amsi Cryptantha cladeisannual.Within = Amsinckiopsis .holoptera C. eto abbreviations: section ; Plag oee,teacsrlstate ancestral the however, ; = Plagiobothrys Cr Cryptantha = Cryptantha ; Sonn Cryptantha .s 1, s. s. = ; Sonnea Ge = Geocarya . ihsvrlmr oyopi eutosaogtx in taxa Cryptantha among occur, reductions fruit polymorphic sections per more nutlet several one with to reductions independent three curn pt 0tms ihnthe Within times. 10 to up occurring n ot mrc)sosardcinfo ort – nut- 1–2 to section four within from Finally, reduction lets. a shows America) South and ; Kr = rntka Cryptantha, Krynitzkia, Krynitzkia .s clade, 2 s. s. ³ ; 0)internodes. 70%) Or = Oreocarya .maritima C. Oreocarya . and Plagiobothrys * ot mrcntx;alothers all taxa; American South = smlsfo ohNorth both from (samples Geocarya Cryptantha hr r w separate two are there eto abbreviations: section Fg B.I the In 3B). (Fig. .s clade, 1 s. s. Delivered by Publishing Technology to: Michael Simpson IP: 174.101.35.52 on: Wed, 11 Jul 2012 04:29:57 Copyright (c) American Society for Plant Taxonomists. All rights reserved. rmu osvntms eedn nteoptimization the on depending times, seven evolved The model. to heteromorphism up Cryptanthinae; from for nutlets ancestral Homomorphic pericarp calyx. are in single or a within size sculpturing in wall differentially develop nutlets whereby 1– or 1–2 to four 3B). from (Fig. fruit, nutlets 3 per number nutlet of reductions 743 CRYPTANTHA OF SYSTEMATICS PHYLOGENETIC SIMPSON: AND HASENSTAB-LEHMAN 2012] brvain si i.1. Fig. in as Abbreviations iue3 hw eosrcino ultheteromorphism, nutlet of reconstruction shows 3C Figure Fig. .Mxmmlklho hlgn ihM otta aussonaoelnaeadBysa otro rbblte eo lineage. below probabilities posterior Bayesian and lineage above shown values bootstrap ML with phylogeny likelihood Maximum 2. Johnstonella ld nqioal osse com- a possesses unequivocally clade cu nthe in heteromorphism occur of evolution the for optimizations sub- Equivocal 3C). by (Fig. lineages some followed in homomorphism nutlets, to reversals sequent heteromorphic with ancestor mon mohnteshv rsnidpnetyfv ie nthe in times five independently arisen have nutlets Smooth ial,htrmrhs vle neednl noeline- one within age in independently evolved heteromorphism Finally, og ult r neta o rpatia Fg 3D). (Fig. Cryptanthinae for ancestral are nutlets Rough Pectocarya Cryptantha Fg 3C). (Fig. .s and 1 s. s. Cryptantha .s clades. 2 s. s. Delivered by Publishing Technology to: Michael Simpson IP: 174.101.35.52 on: Wed, 11 Jul 2012 04:29:57 Copyright (c) American Society for Plant Taxonomists. All rights reserved. ri.Fu ult/ri ht,oente/ri lc,oetontesprfut=dr ry n-he ult/ri ih ry oeta th that Note gray. light = nutlets/fruit one-three gray, dark = fruit per nutlets by one-two (illustrated black, fruit per = nutlets 4 nutlet/fruit is one Crypanthinae for white, condition ancestral = nutlets/fruit Four fruit. pmrhcfor apomorphic 4 YTMTCBTN Vlm 37 [Volume BOTANY SYSTEMATIC 744 o1 ie.Abeitosa nFg 1. Fig. in as Abbreviations times. 10 to rprnil=ga.Nt htana uain(lutae by (illustrated duration annual that Note gray. = perennial or Fig. AadB hrce vlto,mxmmlklho reson .Patdrto seFg.5,6) nul=wie eena lc,annual black, = perennial white, = Annual 6D). 5A, Figs. (see duration Plant A. shown. tree likelihood maximum evolution, Character B. and 3A Oreocarya n vle neednl within independently evolved and Johnstonella .maritima) C. .intermedia C. n sections and sacsrlfrCytnhne eena uain(lutae by (illustrated duration Perennial Cryptanthinae. for ancestral is .Rdcini ultnme ilsrtdby (illustrated number nutlet in Reduction ). Cryptantha and Geocarya of Cryptantha .s .B ultnme per number Nutlet B. 1. s. s. .ganderi C. curdup occurred ) .humilis C. )is e Delivered by Publishing Technology to: Michael Simpson IP: 174.101.35.52 on: Wed, 11 Jul 2012 04:29:57 Copyright (c) American Society for Plant Taxonomists. All rights reserved. Cryptantha 02 AESA-EMNADSMSN HLGNTCSSEAISO RPATA745 CRYPTANTHA OF SYSTEMATICS PHYLOGENETIC SIMPSON: AND HASENSTAB-LEHMAN 2012] ooopi rhtrmrhc=ga.Nt hthmmrhcntes(lutae by by (illustrated nutlets homomorphic peninsularis Pectocarya that Note by gray. (illustrated = nutlets heteromorphic or homomorphic Fig. .peruviana C. CadD hrce vlto,mxmmlklho reson .Nte eeoopim ooopi ht,htrmrhc=black, = heteromorphic white, = Homomorphic heteromorphism. Nutlet C. shown. tree likelihood maximum evolution, Character D. and 3C .s ,and 2, s. s. sacsrlfrCyatia.A“moh cltrn ilsrtdby (illustrated sculpturing “smooth” A Crypanthinae. for ancestral is ) .Nte ufc cltrn.Ruh=wie moh=bak og rsot ry oeta ruh cltrn (illustrated sculpturing “rough” that Note gray. = smooth or rough black, = smooth white, = Rough sculpturing. surface Nutlet D. . Oreocarya .micromeres C. lds cl as=1m.Abeitosa nFg 1. Fig. in as Abbreviations mm. 1 = bars Scale clades. r pmrhcfor apomorphic are ) Johnstonella n aeeovdidpnetyi the in independently evolved have and .mohavensis C. .scoparia C. a vle neednl nthe in independently evolved has ) r neta o rpatia.Heteromorphic Cryptanthinae. for ancestral are ) Cryptantha .s n lds swl sin as well as clades, 2 and 1 s. s. Cryptantha .s 1, s. s. Delivered by Publishing Technology to: Michael Simpson IP: 174.101.35.52 on: Wed, 11 Jul 2012 04:29:57 Copyright (c) American Society for Plant Taxonomists. All rights reserved. outspotol rmteBysa nlss(P=0.97). = (PP analysis Bayesian the from only support of robust clade the to sister placed rmsection sampled taxon one from the Finally, analyses. all in clade supported Allocarya sections from sampled species and analyses, genus u iie apesz,w ont oepten.Investi- patterns. some note do of we species gated size, of sample limited species our among relationships Plagiobothrys Although clades. four to separate three forming analyses, all in groups. polyphyletic as recovered monophyletic supported, strongly aa h genera outgroup The of taxa. sample our on based supported, is Cryptanthinae Plagiobothrys ot mrc,aeas vdn ntegenera the the in evident in also direction are America, opposite South the in dispersal Cryptantha with single complex the possible in times a four of minimum a occured America has South and to North from unidirectional, as explained best Cryptantha results. our by twice indepen- least evolved at having dently Cryptanthinae, in in only derived clearly found is cleistogamy, Second, section of twice. members having least Cryptanthinae, at within evolved derived clearly is heterostyly ult n lnaegnbs,baigaia tl.Saebr mm. 1 = bars Scale style. C,D. apical branching. bearing aerial gynobase, elongate wide and short, nutlets with Plants B. roots. 4 YTMTCBTN Vlm 37 [Volume BOTANY SYSTEMATIC within within times three Cryptanthinae: 746 rpataconfertiflora section in of Cryptantha found members several is in height, and which here) in in (see inversely herkogamy vary analyses of several style type and phylogenetic a stamens heterostyly, our the First, 2). to 1, respect Figs. with assessed 3D). (Fig. species sampled three for apomorphy an hlgntcAnalyses— Phylogenetic ial,w oeta h mhtoi itiuinof distribution amphitropic the that note we Finally, w te opooia etrs(napd eealso were (unmapped) features morphological other Two Fig. .Dansi n/raoopi etrso eurce genus resurrected of features apomorphic and/or Diagnostic 4. Amsinckia Cryptantha Amsinckia r ahmnpyei n oehrfr strongly a form together and monophyletic each are .s ld.Idpnetdsesl,fo ot to North from dispersals, Independent clade. 1 s. s. .l,i xlie i ogdsac ipra,is dispersal, distance long via explained if l., s. seFg 2). Fig. (see r eodtesoeo hssuybcueof because study this of scope the beyond are ona lgoohy hispidus, Plagiobothrys Sonnea, Plagiobothrys Amsinckia nteBysa n aiu likelihood maximum and Bayesian the in .s ,adoc within once and 2, s. s. pce (including species Cryptantha and Discussion nalaaye,mnpyyo the of monophyly analyses, all In and Eremocarya section .flava C. and Pectocarya Geocarya Amsinckiopsis Cryptantha ape ee;thus, here); sampled , .spectabilis, A. + Oreocarya Oreocarya Oreocarya r anandas maintained are fSuhAmerica, South of Plagiobothrys Plagiobothrys sconsistently is Pectocarya .micrantha C. .s ,once 1, s. s. aresisterto (including u with but , h last the , sampled and and var. is Eremocarya lepida ae eea n ftecae scmoe foespecies micrantha one section C. of in composed placed is taxa clades two with the of One previously genera. to named corresponding four recovered, consistently are ei nalaaye,a ssection is as analyses, all in letic pce eg ulim n adi 00 01 ilmore will 2011) 2010, relationships. Baldwin these resolve and definitively Guilliams of size (e.g. sample species increased an that predict We itdseisbcueo h itnt lblrnutlets that globular show distinct, results Our the taxa. of these in because present species genus the ciated proposed encompass Brand to 1927, in or biennial Also the was groups that in two root character the perennial separate primary to the used been America has North in separate Even account. definitely will Cryptantha that character ical American of South members examining after that, reasoned Payson 1927). ontnsteteto ot American South of genus treatment the Johnston’s reduced (1927) Payson Oreocarya by monograph A lication. American American North between differences from species of distinguished group. subset Greene a monophyletic include supported genus to the well circumscribed originally a (1887b) Greene forms also lyses, ld scnitnl lcdsse othe to sister placed consistently is The 4D). clade than (Fig. height style persistent in a greater with is nutlets that bracteatemature gynobase 4B), a and (Fig. 4C), width (Fig. is flowers and that height system in branching red- equal aerial having approximately an by 4A), morphologically (Fig. diagnosed roots Taxa pigmented are 1). Table clade see this (1887b; Greene in by described and named as as treated n testo others and all in e.g. remaining found, The (as keel that ventral nutlets a and lack calyx, fruiting persistent duration, perennial .Crla.D ri,wt psplu ay n or homomorphic four, and calyx aposepalous with Fruit, D. Corollas. C. . Cryptantha The A,B. . Oreocarya rpatamicrantha Cryptantha .lepida C. Cryptantha, oascinof section a to hnteSuhAeia pce r ae into taken are species American South the when Cryptantha, var. .l,a urnl icmcie,i lopolyphy- also is circumscribed, currently as l., s. Cryptantha .cinerea C. Krynitzkia Oreocarya ld,wt ih ape aai u ana- our in taxa sampled eight with clade, m. .Ti orsod otegenus the to corresponds This ]. and h atrnm aigpirt fpub- of priority having name latter the Oreocarya hr s“ is there steewresswn diagnostic no saw workers these as , .micrantha C. pce eetaserdb Greene by transferred were species [ .jamesii C. n h nuloein one annual the and var. Cryptantha, micrantha Krynitzkia Krynitzkia ... from n orcoeyasso- closely four and ] o igemorpholog- single a not .Rdprl pigmented Red-purple A. . var. Plagiobothrys Krynitzkia Krynitzkia Oreocarya Krynitzkia Cryptantha nareetwith agreement in yJhso (1925): Johnston by i ao clades major Six . lepida Hemisphaerocarya Plagiobothrys Oreocarya .cinerea C. Cryptantha. [sometimes Eremocarya, clade. n South and Eremocarya Tbe1). (Table ae on based (Johnston Oreocarya species). and .l. s. is ” Delivered by Publishing Technology to: Michael Simpson IP: 174.101.35.52 on: Wed, 11 Jul 2012 04:29:57 Copyright (c) American Society for Plant Taxonomists. All rights reserved. ult eta iw hwn u-pclvnrlgov arw.Saebr mm. 1 = bars Scale (arrow). groove ventral sub-apical showing view, ventral D-F. nutlet, size. large H. view. note ventral view; dorsal Nutlet, C. flowers. showy large, vle nyi eti ebr of members certain in among 5E, only addition, (Fig. evolved roughened In to H-J). rugulose or F, 5C,G) is that (Fig. sculpturing smooth a either and 5F,J) with (Fig. showy nutlets 747 groove and ventral large, sub-apical calyces, a relatively fruiting persistent 5A,D), 5B), (Fig. (Fig. flowers leaves of rosette basal Cryptantha CRYPTANTHA OF SYSTEMATICS ments, PHYLOGENETIC SIMPSON: AND HASENSTAB-LEHMAN of the within well nested 2012] et aa evs .Nte,dra iw hwn uuoesupuig .Nte,vnrlve;nt uaia eta roe(ro) G. (arrow). groove ventral subapical note view; ventral Nutlet, F. sculpturing. rugulose showing view, dorsal Nutlet, E. leaves. basal tent, Fig. Hemisphaerocarya .Dansi n/raoopi etrso eurce genus resurrected of features apomorphic and/or Diagnostic 5. Oreocarya .l yprnilpatdrto ihapersistent a with duration plant perennial by l. s. .roosiorum C. swl icmcie rmremaining from circumscribed well is sufudd ept asnscom- Payson’s Despite unfounded. is ult oslve,wt uuoesupuig I. sculpturing. rugulose with view, dorsal nutlet, , Oreocarya Cryptantha ld ota recognition that so clade Oreocarya. .l,htrsyyhas heterostyly l., s. smentioned As .humilis C. Oreocarya .Woepat ebru he,soigprnilhbtadpersis- and habit perennial showing sheet, herbarium plant, Whole D. . h asmn nlssti ld spae itrto sister analyses, of placed Bayesian and likelihood is the the both In clade 1). (Fig. ported this analysis In Cryptantha varies. parsimony Cryptanthinae within the placement its but analyses, ihtecasfcto sdi w eaieyrcn floristic recent 1987). relatively Weber two 1951; (Abrams in treatments used classification the with of recognition the earlier, .thyrsiflora C. The ontnla Amsinckia, Johnstonella, Cryptantha A-C. . Cryptantha .confertiflora C. ult oslve,wt uuoesupuig . J sculpturing. rugulose with view, dorsal nutlet, , .s ,btti lcmn sntrbsl sup- robustly not is placement this but 1, s. s. .s ld ssse oalrecaecomposed clade large a to sister is clade 2 s. s. .s ld ssrnl upre nall in supported strongly is clade 2 s. s. .Pat eena.B noecne showing Inforescence, B. perennial. a Plant, A. . Oreocarya w ldsof clades two sadsic eu agrees genus distinct a as Plagiobothrys .cinerea C. species, .weberi C. ,nutlet, , Delivered by Publishing Technology to: Michael Simpson IP: 174.101.35.52 on: Wed, 11 Jul 2012 04:29:57 Copyright (c) American Society for Plant Taxonomists. All rights reserved. Fg D,adbataefoes(i.6) ult r not wide are as nutlets high 6D); of as species (Fig. two basally less The flowers 6C). or is (Fig. bracteate more distinctive and that is 6D), that calyx (Fig. branched system a stem a 6B); (Fig. aerial fruit 6A), in cirscumscissile and (Fig. synsepalous roots genus red-pigmented The 2). 1, (Figs. and subgenus) (or section in called relationships been elucidate has to what necessary features is these sampling of members However, extensive other maturity. many at with shared unit are a as easily detach diagnose the assem- both in Cryptantha polyphyletic found a analyses, our form in America, blage South taxa and The section and North complex. in both feature the classified in derived been once have a than that is more evolved cleistogamy have results that may our fact but the relationships, support phylogenetic their evaluate to Geocarya of section of species a section plus section of of member sampled species one the two analyses, alyssoides C. all In chasmogamous) flowers. (and cleistogamous with species contain and 4 YTMTCBTN Vlm 37 [Volume BOTANY SYSTEMATIC the and 748 Geocarya and sampled), to greater needed clades. be a two these will with of data interrelationships studies molecular the confirm Future additional or 2). size and sample 1 clades encompass genus the within it taxa these that to fact sister the be and actually may position phylogenetic uncertain its Given the for features morphological nostic Geocarya ot.B icmcsieclx ieo eicnea ro.C ult eta iw D. view. ventral Nutlet, C. arrow. mm. at 1 dehiscence = of bars line Scale calyx; Circumscissile B. roots. Fg.1 ) hscaecnan ersnaie fsubgenera of representatives contains clade Cryptantha This 2). 1, (Figs. Cryptantha The wnytresmldtx of taxa sampled Twenty-three Fig. Cryptantha .similis, C. .Dansi n/raoopi etrso eurce genus resurrected of features apomorphic and/or Diagnostic 6. Greeneocharis Cryptantha and sampled, and Krynitzkia .s lds hrcesta aebe sdto used been have that Characters clades. 2 s. s. .s ld ihsrn upr nalanalyses all in support strong with clade 1 s. s. oeseissampled), species (one and Cryptantha Cryptantha a on ob elspotdi l analyses all in well-supported be to found was .s Fg.1 ) ebr fsubgenera of Members 2). 1, (Figs. 1 s. s. Krynitzkia .aprica C. .linearis C. ld ftospecies, two of clade .s ld.W antietf n diag- any identify cannot We clade. 1 s. s. nld nana ai n risthat fruits and habit annual an include Greeneocharis Cryptantha cu xlsvl nSuhAmerica South in exclusively occur utb ape oeextensively more sampled be must omawl upre ld with clade supported well a form , 1 pce ape) Subgenera sampled). species (19 rntka .collina. C. Krynitzkia, Cryptantha snse ihnaohrclade another within nested is , Cryptantha .s ,w hoet retain to choose we 1, s. s. rpata .capitulifora C. Cryptantha, Krynitzkia Geocarya sdansdb having by diagnosed is Cryptantha .s dlmtdhr to here (delimited s. s. rpatacircumscissa Cryptantha Cryptantha Greeneocharis Cryptantha Krynitzkia. itiue in distributed , trespecies (three opieour comprise .s clade. 2 s. s. h other The .s and 1 s. s. .l More l. s. Geocarya, hwa show , Greeneocharis racemosa genus the from transferred species two diplotricha C. American North the analyses and all in although the what, within species of tionships Cryptanthinae separate of containing two rest clade and a the to sister to or basal 1), (Fig. either being varies, clade genus subgenus or holoptera section Cryptantha con- of previously part all be were to these sidered analyses; all in supported well ancestry. common via is similarity this additional with research n w ot mrcnseis(al ) n ftenine the was Angustifoliae of One in 3). species (Table American species North American series South his two in American North and 1927) nine (1925, of total Johnston a contained by which Angustifoliae, placed were recognized hetero- species Brand two are by The that margins. acute nutlets with and with morphic herbs suffrutescent perennial, nutfla (excluding the Angustifoliae than roughened less nutlets.” the slightly surpassing definitely sometimes style others; and larger ial abax- nutlet odd in with heteromorphous decidedly or species homomorphous most pale winged, definitely with or knife-like or dark acute nutlets, sides tuberculate with triangular-oblong, or or triangular-ovate roughenings, muricate genus “4, as the Angustifoliae having series in of members it diagnosed phylogenetic (1925) treat Johnston molecular we and and morphological others data), the both with on belong not (based does clearly species this however, outspot neetnl,teol te nw members known other only the Interestingly, support. robust r oehtdsatyrltdi u nlss(is ,2). 1, (Figs. genus analyses the our in related Greeneocharis distantly somewhat are to resemblance remarkable fCytnhnehvn icmcsieclcs sfound as calyces, circumscissile in having Cryptanthinae of Johnstonella ial,the Finally, hs epooeta h 0seiso Johnston’s of species 10 the that propose we Thus, Greeneocharis .holoptera C. Oreocarya .similis C. A-C. . and Plagiobothrys .circumscissa C. and scnitnl itrt ld ffu pce of species four of clade a to sister consistently is hwn hr,wd eilbacigadfoa bracts. floral and branching aerial wide short, showing , a ecie yBad(95,woincluded who (1925), Brand by described was Johnstonella .inaequata J. cu nseisof species in occur , Gen 87) h oiino the of position The 1887b). (Greene .parviflora C. r itrtx,a r h ot American South the are as taxa, sister are Plagiobothrys fti ld a rgnlypae nthe in placed originally was clade this of nalaaye Fg.1 ) u without but 2), 1, (Figs. analyses all in .micrantha C. var. Plagiobothrys rn igoe h eu as genus the diagnosed Brand . ld fegtsmldseisis species sampled eight of clade Eremocarya circumscissa Fg.1 2). 1, (Figs. lds(i.2.Teinterrela- The 2). (Fig. clades Johnstonella )belonginthe mici,Greeneocharis, Amsinckia, Eremocarya .Rdprl pigmented Red-purple A. . u hs w groups two these but , Cryptantha Plagiobothrys. aamyacranif ascertain may taxa Krynitzkia ld aysome- vary clade .micrantha C. : seabove). (see Johnstonella Johnstonella Johnstonella although , .costata C. Further ; Delivered by Publishing Technology to: Michael Simpson IP: 174.101.35.52 on: Wed, 11 Jul 2012 04:29:57 Copyright (c) American Society for Plant Taxonomists. All rights reserved. opooia ris odtx eune o hssuy ot mrcndistribution. American South = * study. this for sequenced = taxa Bold traits. morphological pce,w r ofdn htteohr fJohnston’s and of addi- an to species adding others three propose tional we the addition, these In morphology. that of nutlet seven confident in only belong are Angustifoliae for 2). data we 1, (Figs. sequence study species, this have of we results the Although given 3), Table (see clade grayi C. fastigiata C. echinosepala C. diplotricha* C. 749 costata C. angustifolia C. angelica C. CRYPTANTHA OF SYSTEMATICS PHYLOGENETIC SIMPSON: AND HASENSTAB-LEHMAN 2012] .grayi C. oe eeal eeosit -eddnte.Hwvr in However, nutlet. 1-seeded a into develops generally lobes elevation. high at occur necessarily not do the which of in at members independently persist some evolved and or also colonize has many to perenniality where Perennial plants elevations, group. these higher allow that may for duration apomorphic and of Cryptanthinae habit perennial demon- the clearly that study strates This ancestors. perennial from derived hypothesized taxonomists 1971) of annual previous habit Higgins duration, perennial 1927; the plant that Payson to 1925; regard (Johnston With within l. tradi- groups those taxonomic s. of delimit subset to a used are tionally study this in examined characters echinosepala Cryptantha .grayi C. eentincluded. not were of representatives resolution and sequenced little of shows taxa Schwarzer the of among cladogram the ever, that ring see 3; (Table winged 7). narrowly Fig. or examples, margin rimmed, a acute, and often is apex, a that non-acuminate color, acute, in to white obtuse tubercles rounded spaced regularly prominent, with dis- that traits Morphological tinguish is Angustifoliae. in species it this placed have that noted both to he similar 3). however, (Table resemble classification; margin series nutlet but rounded fastigiata to Maritimae, Cryptantha angled an series with nutlets his in Johnstonella (1925) Johnston .racemosa C. pusilla C. parviflora* C. micromeres C. inaequata C. holoptera C. nyoeseisof species one only Table ihnsbaiyBrgniee aho h orovary four the of each Boraginoideae, subfamily Within hrce vlto n igorpi History— Biogeographic and Evolution Character infer- in (2007) Schwarzer of those with agree results Our Cryptantha .micromeres C. var. var. var. mici,Cytnh,Pectocarya, Cryptantha, Amsinckia, .Seisadvreisof varieties and Species 3. Johnstonella Amsinckia cryptochaeta grayi neseotica ao eis(ontn12,12)Nte agnNte eeoopimNte shape Nutlet heteromorphism Nutlet margin Nutlet 1927) 1925, (Johnston Series taxon aai aightrmrhc triangular-ovate heteromorphic, having in taxa .holoptera C. h ato hc a ape here. sampled was which of last the , and a ecie usqett Johnston’s to subsequent described was Johnstonella Cryptantha Johnstonella nld fe eeoopi nutlets heteromorphic often include and Johnstonella Pectocarya n C. and Cryptantha Oreocarya .micromeres C. ntcasfe)Nrol igdHtrmrhcTriangular-ovate Heteromorphic winged Narrowly classified) (not aiia one eeoopi Lance-ovate Heteromorphic Rounded Maritimae aiia nldHtrmrhcLneoaet Ovate to Lance-ovate Heteromorphic Angled Maritimae nutfla arwywne ooopi Triangular-ovate Triangular-ovate Triangular-ovate Lance-ovate Homomorphic Homomorphic Homo.) (rarely Heteromorphic Heteromorphic winged Narrowly winged Narrowly angled to Rounded winged Narrowly Angustifoliae Angustifoliae Angustifoliae Angustifoliae nutfla one ooopi Lance-ovate Homomorphic Rounded Angustifoliae nutfla one ooopi Lance-ovate Homomorphic Rounded Angustifoliae nutfla one ooopi Lance-ovate Homomorphic Rounded Angustifoliae nutfla nldHtrmrhcLneoaet Ovate to Lance-ovate Triangular-ovate Triangular-ovate Triangular-ovate Triangular-ovate Heteromorphic Heteromorphic Homomorphic winged Narrowly Heteromorphic winged Narrowly Homomorphic Angled winged Narrowly Angustifoliae winged Narrowly Angustifoliae Angustifoliae Angustifoliae Angustifoliae Cryptantha Eremocarya .l,epcal ie that given especially l., s. : n n of one and Oreocarya Oreocarya inaequata .ehnspl,C fastigiata, C. echinosepala, C. ie hi iiaiyin similarity their given , r oohltc How- monophyletic. are a rmtv,wt the with primitive, was .s lds(i.3A), (Fig. clades 1 s. s. eurce rtaserdt h genus the to transferred or resurrected n ol likely would and , and sdrvdwithin derived is cu.However, occur. and eepae by placed were Oreocarya Plagiobothrys Greeneocharis Johnstonella Cryptantha were The td hw htnte eeoopimhseovdmul- the evolved within has times, heteromorphism within tiple nutlet that groups shows of study classification the in per fruits tested. more been not or has nutlets idea this larger fewer however, either plant; with have taxa fruit those that per could in nutlets but allocation unclear, resource is to related number be nutlet in reduction of nificance ics hs etrsi hlgntcfaeok(not framework phylogenetic a in features these various discuss in of found significance features floral adaptive the tested; 1983), been unknown. (Grau is not nutlets dispersal smooth animal has to this adapted but more nut- be Roughened nutlets may ones.” smooth roughened lets than that primitive speculated “more who are (1927), Payson the against within independently Cryptantha times five six evolved having to feature, derived large a represent the analysis, studied. our been of The not has plants. survivability nutlets other and “consimilar” versus many widely germination “odd” more in be for found to three potential syndrome other a the remain and dispersed, to parent, fruit the differ- per to This propagule nearer often one detached. enables and easily three attachment are smaller ential other sculpturing, usually nutlet, the rougher are whereas “odd” have remains which gynobase, the ornamented, nutlets, the that less “consimilar” to fact and/or attached the larger firmly usually to heteromor- is related of which be significance may adaptive phism The taxa. eromorphic Fg C,ti etr paetyrpeet napomorphy an represents the apparently variable feature for is this character 3C), this (Fig. of optimization Although 3C). (Fig. Cryptantha oho hc aecesoaosfoes h dpiesig- adaptive that The is flowers. cleistogamous number have nutlet which of sections in both from reduction species of by of clade characterized composed one 1 Interestingly, 3B). s. (Fig. s. clades 2 s. s. ocrusrb pce.Ti td eel htrdcinin within Cryptanthinae, reduction in times that numerous occurred past reveals has the number study nutlet in This used species. feature a circumscribe fruit, to mature per nutlets of number Cryptantha ulthtrmrhs satatotnwihe highly weighted often trait a is heteromorphism Nutlet ent hthtrsyyadcesoayaeapomorphic are cleistogamy and heterostyly that note We of outside rare are nutlets Smooth Oreocarya Johnstonella Johnstonella .l,mn aacnitnl xii euto in reduction a exhibit consistently taxa many l., s. .s ,and 2, s. s. .s lds swl si h genus the in as well as clades, 2 s. s. n ohthe both and ihsre lsiiainaddt o he key three for data and classification series with ld,wihcnan oto h het- the of most contains which clade, Oreocarya Johnstonella Cryptantha Cryptantha lds(i.3) hsgoes This 3D). (Fig. clades , Geocarya Cryptantha Cryptantha Cryptantha .s and 1 s. s. Cryptantha .l aa n we and taxa, l. s. and .s ,and 1, s. s. .l,adin and l., s. Cryptantha, Cryptantha Cryptantha Pectocarya .l Our l. s. .s 1, s. s. Delivered by Publishing Technology to: Michael Simpson IP: 174.101.35.52 on: Wed, 11 Jul 2012 04:29:57 Copyright (c) American Society for Plant Taxonomists. All rights reserved. ult n rjcigsye .Nte,dra iw .Nte,sd iw L. C. R-V. view. side Nutlet, P. K. view. view. dorsal Nutlet, micromeres J. C. style. projecting and nutlets H. view. dorsal shown, cl as=1mm. 1 = bars Scale D. view. dorsal nutlets, small three of One C. ml n n ag hw,dra iw F. view. dorsal shown, large one and small 5 YTMTCBTN Vlm 37 [Volume BOTANY SYSTEMATIC 750 Fig. racemosa. .Mmeso eurce genus resurrected of Members 7. .parviflora C. .Htrmrhcnteso ri,dra iw .Lre eaieysot ult oslve.O n ftresal og ult,dorsal nutlets, rough small, three of One O. view. dorsal nutlet, smooth relatively Large, N. view. dorsal fruit, of nutlets Heteromorphic M. . ,.Pati h il.T ri,soightrmrhcntesadpoetn tl.U ornteso ri.V ag ult eta view. ventral nutlet, Large V. fruit. of nutlets Four U. style. projecting and nutlets heteromorphic showing Fruit, T. field. the in Plant R,S. ihhtrmrhcntes n fec ieson oslve.Q. view. dorsal shown, size each of one nutlets, heteromorphic with , .grayi C. var. grayi ihhmmrhcntes oslve hw.I-K. shown. view dorsal nutlets, homomorphic with , Johnstonella .diplotricha C. .costata, C. .A. n ffu ooopi ult,dra iw E. view. dorsal nutlets, homomorphic four of one ihhmmrhcntes oslve hw.G. shown. view dorsal nutlets; homomorphic with , .angelica C. ihhtrmrhcntes B,C. nutlets. heteromorphic with , .inaequata C. ihhtrmrhcnteso ri hw,dra iw M-O. view. dorsal shown, fruit of nutlets heteromorphic with , .pusilla, C. .holoptera C. n ffu ooopi ult hw,dra view. dorsal shown, nutlets homomorphic four of one .echinosepala C. .angustifolia C. ihhmmrhcntes .Futwt four with Fruit I. nutlets. homomorphic with , .fastigiata C. .Fu,htrmrhcnutlets. heteromorphic Four, B. . ihhtrmrhcntes one nutlets, heteromorphic with , eeoopi ult ffruit of nutlets heteromorphic , Delivered by Publishing Technology to: Michael Simpson IP: 174.101.35.52 on: Wed, 11 Jul 2012 04:29:57 Copyright (c) American Society for Plant Taxonomists. All rights reserved. eaaefrom separate lutae) eeotl sfudi eea pce of species several in found is Oreocarya, Heterostyly illustrated). anann eeata r oohltci olm l four and all lump genera, to recognized is previously monophyletic are that of genera maintaining members other to several than that genera ana- evidence our good in supported have Cryptantha always do not we are lyses, clades relation- major “backbone” of Although ships polyphyletic. is treatments, genus the that is needed trend. are this taxa, corroborate greater American to South substantially of However, a particularly 1983). size, with sample Grau analyses 1963; biogeographic (Raven formal relatively event a of dispersal notion pat- the recent current supporting explain and distribution to making of scenario terns support, unlikely strong highly with a vicariance monophyletic accessions These as localities. resolved American are South is and This North 2006). of both sampling al. our et by Moore distrib- evidenced 1983; strongly amphitropically most Grau 1963; American (Raven taxa other agrees uted study of this scenarios in supported with America South to North from habitat desert their in giving advantage above, 1983). an (Grau flowers former the chasmagamous considerably of of are seedlings cleistogenes those these than of in larger damaged; nutlets or eaten the are plant branches addition, the aerial enable if may even level reproduce ground to at found 1983) Grau (“cleistogenes,” sensu pollinators flowers cleistogamous of specialized adaptation absence highly the an the Calvin in be 1983; reproduce (Grau may other to with feature plants As enabling this analyses. our plants, in cleistogamous twice evolved having clade, in sections studied been has and of 2000), (Baker members al. some mechanism in et outcrossing detail It Barrett an 1997). 1975; as al. Yeo known 1966; et Schoen been 1957; long Chisaki has and Ray 1976; (Ornduff 02 AESA-EMNADSMSN HLGNTCSSEAISO RPATA751 CRYPTANTHA OF SYSTEMATICS PHYLOGENETIC SIMPSON: AND HASENSTAB-LEHMAN 2012] eu seeape nFg 8). Fig. in examples (see the genus both retain robust Cryptantha we without are Thus, clades two support. these of Although interrelationships clade. the this within nested our are of genus) members the of (subgenus) type section of genus members the in remain clade ee hs epooet retain to Plagiobothrys propose we especially denoted Thus, and clades major here. floral, the vegetative, characterizes that would in morphology fruit diversity classification the this called reflect However, be not publication. would approxi- of of which genus priority species, large one, 500 into study) mately this in analyzed not Oreocarya orgenera, four sdsusderir n ftemjrrslso hsstudy this of results major the of one earlier, discussed As dispersal long-distance unidirectional, mostly the Finally, American South of members to restricted is Cleistogamy Plagiobothrys Cryptantha h aawti u designated our within taxa The . swl si oemmeso h genus the of members some in as well as .l r oecoeyrltdt pce fother of species to related closely more are l. s. .s ruswti reduced a within groups 2 s. s. (plus Cryptantha rmcra reecai,Johnstonella, Greeneocharis, Eremocarya, Cryptantha Cryptantha ad otlkl,tegenus the likely, most (and, ˜ aooi Treatment Taxonomic n aet.20) nsection In 2003). Galetto. and o Harpagonella and Cryptantha Geocarya .s ntoo h he anlayses, three the of two in 1 s. s. Oreocarya Cryptantha, .l,a icmcie nrecent in circumscribed as l., s. mici,Cytnh,Pectocarya, Cryptantha, Amsinckia, .s ru alit clade a into fall group 2 s. s. .l pce.Oeslto to solution One species. l. s. o o)adt resurrect to and now) for Cryptantha mici,Pectocarya, Amsinckia, four of Cse 1985). (Casper Cryptantha ie htsequenced that given Cryptantha Cryptantha .maritima C. cnann the (containing Cryptantha Cryptantha Harpagonella .s and 1 s. s. Amsinckia Geocarya .s 1 s. s. .s 1 s. s. from .s. s. and and by , O .Genohrssimilis Greeneocharis 2. .Genohrscircumscissa Greeneocharis all 1. species, second a species, of for members and as type named variety combi- previously a new for Two the needed 1840). 370. are Voy. nations only Beechey Bot. Arn., for & Hook. (basionym 1909 genus 677. 36: Club A Bot. the fruit. in circumscissa in calyx Greeneocharis cirscumscissile a exists and and flowers, synsepalous name bracteate is basally wide, that as is system high that as stem less aerial or branched more a root, red-pigmented yteItrainlCd fBtnclNomenclature Botanical of E Code 2006). al. International et (McNeill required the as taxa, certain for by given gen- are resurrected combinations proposed New four era. the of features diagnostic are ree(87) nvldypbihn h eu ae cited exclude name, we sculpturing. genus these, Of the roughened species. publishing nine to validly in rugulose, (1887b), Greene persistent smooth, ventral sub-apical flowers, and a with large groove nutlets persistent large relatively relatively a leaves, and with calyces, of perennials rosette as diagnosed basal plants, of group 1:13 88.Sbeunl,Mcrd changed MacBride lepida Subsequently, 1878). 193. and (1): 1859) 141. [Emory] micranthum Bot. lepida Bound., Mex. Eremocarya type U.S. designated) Rep. Torr., (here the persistent name, micrantha species: genus a the two and publishing nutlets, validly cited is in mature that (1887b), the gynobase Greene than a style. flowers, height bracteate more in wide, is as greater that high system as stem less aerial or branched a root, pigmented rmtegnsadteti sasnnmof synonym a as it treat and holoptera genus the from G no species, two as needed. or are 2012) combinations al. (which new et varieties Kelley two see here; with recognize species we single a as in treated names published whether validly already are there that lepida REMOCARYA REENEOCHARIS REOCARYA h genus The Oreocarya Eremocarya hrceiain n e Combinations— New and Characterizations o,Mhv eet ..Hgwy35 . in mi 8.7 395, UC). isotype: Highway 1958, RSA; (holotype: Mar U.S. 29 Desert, Adelanto, Mohave Bernardino San California: Co., A. S. U. 1962.—TYPE: 168. 16: aesa .G ipo,cm.nv Basionym: nov. comb. Simpson, G. similis Cryptantha M. & Hasenstab Piptocalyx rnl eit 6.19.[ 1899. 460. Regist. Prantl] 91—YE .S .Clfri:Iy o,Cottonwood Co., Inyo 1949, Aug California: 12 A. Lakes, S. U. 1951.—TYPE: var. Johnst. M. I. o.Basionym: nov. rosulata aB. rc mr cd rs5:55 96 Given 1916. 545. 51: Arts Acad. Amer. Proc. MacBr., nrn,to rank, in blw.O h eann ih pce ie by cited species eight remaining the Of (below). Tre)Gen (basionym Greene (Torrey) ree itna1 7 1887b. 57. 1: Pittonia Greene, or var. Torr. ree itna15.1887b. 1:58. Pittonia Greene, icmcie opooial well-defined morphologically a circumscribes J .Hwl)Hsntb&M .Smsn comb. Simpson, G. M. & Hasenstab Howell) T. (J. Greeneocharis lv xBenth.; ex Oliv. Gu scaatrzdb lnswt red- a with plants by characterized is ¨ k am,NtPlnefm Ege & [Engler Nat.Pflanzenfam. Harms, & rke A ry ree(basionym Greene Gray) (A. rmcramicrantha Eremocarya rosulata rpatacircumscissa Cryptantha lepidum .Mte .H ae,Madron Raven, H. P. & Mathew K. oel26227 Howell Ho.&An)Rd,Bl.Torrey Bull. Ryd., Arn.) & (Hook. scaatrzdb lnswt a with plants by characterized is K ahw&P .Raven) H. P. & Mathew (K. J.T.Howell,Leafl.W.Bot.6:104. .LwsadK ahw1113 Mathew K. and Lewis H. Wheelerella .Ga,Sn l .Ae.2 Amer. N. Fl. Syn. Gray, A. Piptocalyx Ho.&An)Rd var. Ryd. Arn.) & (Hook. ihsemmcircumscissum Lithospermum Cryptantha roay holoptera Oreocarya hltp:CAS). (holotype: rtihu micranthum Eritrichium Tr. reevar. Greene (Torr.) .B rn,ined.] Grant, B. G. or,ie. non ined., Torr., . Ho.&Arn.) & (Hook. h following The Eremocarya Johnstonella Eremocarya Eremocarya Eritrichium Greene ˜ o , Delivered by Publishing Technology to: Michael Simpson IP: 174.101.35.52 on: Wed, 11 Jul 2012 04:29:57 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 5 YTMTCBTN Vlm 37 [Volume BOTANY SYSTEMATIC 752 .Ifoecne hwn lrlbat.V ult oslve.Saebr mm. 1 = bars Scale view. dorsal Nutlet, V. bracts. floral showing Inflorescence, U. litgmu lwr(ro) rudlvlo ln.F ulto hsoaosfoe,dra iw -.Section S,T. G-T. view. dorsal flower, chasmogamous of Nutlet F. plant. G,H. of clade). Section ground-level D-F. (arrow), view. dorsal flower flower, cleistogamous chasmogamous of Nutlet C. (arrow). flowers oslve.O,P. view. dorsal K-L. view. Fig. .nevadensis C. .Eepasof Exemplars 8. .leiocarpa C. .clevelandii C. .pterocarya C. var. tp pce of species (type nevadensis var. Cryptantha c. var. .Ifoecneui,acriae crii ye .Nte,vnrlve.I,J. view. ventral Nutlet, H. cyme. scorpioid circinate, a unit, Inflorescence G. .Ifoecne .Nte,vnrlve.UV Section U,V. view. ventral Nutlet, T. Inflorescence. S. . p .Ifoecne .Wne ult oslve.Q,R. view. dorsal nutlet, Winged P. Inflorescence. O. . .s -.Section A-C. s. s. Krynitzia .K hl ln nfed .Nte,dra iw M,N. view. dorsal Nutlet, L. field. in plant Whole K. ). rpata .capituliflora C. Cryptantha. ecra .aprica C. Geocarya. .Woepato ebru pcmn .Fut fcleistogamous of Fruits B. specimen. herbarium of plant Whole A. . .intermedia C. Krynitzkia .Woepatfo ebru pcmn .Ntesof Nutlets E. specimen. herbarium from plant Whole D. . var. .muricata C. (from i .Foes ls-p .Nte,dra view. dorsal Nutlet, R. close-up. Flowers, Q. . .flaccida C. Cryptantha var. Krynitzkia .Ifoecne .Nte,dorsal Nutlet, J. Inflorescence. I. . m .Ifoecne .Nutlet, N. Inflorescence. M. . .s clade). 2 s. s. (from Cryptantha .maritima C. .s 1 s. s. . Delivered by Publishing Technology to: Michael Simpson IP: 174.101.35.52 on: Wed, 11 Jul 2012 04:29:57 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 02 AESA-EMNADSMSN HLGNTCSSEAISO RPATA753 CRYPTANTHA OF SYSTEMATICS PHYLOGENETIC SIMPSON: AND HASENSTAB-LEHMAN 2012] .Oecrafulvocanescens Oreocarya 8. creutzfeldtii Oreocarya 7. barnebyi Oreocarya 4. .Oecraatwoodii Oreocarya us, 3. to known taxa combinations. new Twenty-one require 1987). however, Weber 1971; genus Higgins the in .Oecragrahamii Oreocarya 9. crassipes Oreocarya 6. compacta Oreocarya 5. eto rsubgenus of species or described section 62 approximately TYPE: the of 1870; Most 479. [Hayden] as Montana type virgatum generic Eritrichium the designate virgata here we Greene, 1 roay johnstonii Oreocarya 11. hypsophila Oreocarya 10. ye R;ioye:CS Y S,UC). RSA, NY, 1981, CAS, S May isotypes: BRY; slope 14 along type: on ft, 10, marker, 6,400 hwy historical canal, nw Creek of Utah, mi Muddy 1.5 A.: at ca road S. S22, dirt U. R6E, T21S, 1982.—TYPE: Co., 203. Emery 42: Naturalist Basin Basionym: nov. stps ,M,N,R,U,US). UC, RM, NY, MO, E, isotypes: 1901, Jun 11 Run, Water- Deer Gunnison shed, County, Mesa Colorado, Basionym: A.: S. nov. U. TYPE: stat. nov., comb. nitida Oreocarya Kelley, B. R. (Greene) Fria Agua of east 1939.—TYPE: miles 1936, 6.5 Apr 397. Co., 13 20: Springs, Brewster Texas: Arbor. A. Arnold S. J. U. Johnston, Basionym: M. nov. I. comb. Simpson, 1968, Jun 18 Range, UTC). RSA, Needle the 1613 of pass Higgins of end west m north 100 Experimental at 21, Range Hwy Desert along of Headquarters west Station mi 8 Utah: ca. A. Co., S. Millard U. 1968.—TYPE: 196–197. 28: Naturalist Basin Basionym: nov. 8748 1947, Barneby Jun C. 17 R. Ouray, & of 1948.—TYPE: Ripley east D. mi 240. D. 30 29: Co., H. Uintah Arbor. Utah: A. Arnold S. J. U. Basionym: Johnston, M. nov. I. comb. Simpson, ipo,cm.nv Basionym: nov. comb. Simpson, Sawtooth WTU). of isotype: 1937, Forest, GH; head Aug (holotype: ridge, 6 National ft, high 11,000 of Sawtooth Mountains, crest 1939.—TYPE: Co., Creek, 395. Blaine 20: Boulder Idaho: Arbor. A. Arnold S. J. U. Johnston, M. Basionym: I. nov. comb. Simpson. Co., Uinta 1933, Wash, Sand May Utah: of 28 mouth A. ft, of 4,500 north S. River Green U. of west 1937.—TYPE: bench 23. 18: Arbor. Basionym: 9ad14o w 9 0My1970, May 20 89, BRY). hwy hwy (holotype: of on junction 164 of N and mi 7 89 Co., Coconino Arizona: A. S. U. 1974.—TYPE: 127. Basionym: 19(2): Naturalist Southw. Higgins, nov. C. L. comb. Simpson, Pre)Gen,Ptoi :5.18b(basionym: 1887b 57. 1: Pittonia Greene, (Porter) Oreocarya rpatagrahamii Cryptantha hltp:BY stps H L O NY, MO, LL, GH, isotypes: BRY; (holotype: ree l ae.[ree :2.1901.— 21. 3: [Greene] Baker. Pl. Greene, rpatacreutzfeldtii Cryptantha Oreocarya rpatacompacta Cryptantha otr rlm e.US el Surv. Geol. U.S. Rep. Prelim. Porter, I .Jhs. .B ely ob nov. comb. Kelley, B. R. Johnst.) M. (I. L .Hgis aesa .G. M. & Hasenstab Higgins) C. (L. I .Jhs. aesa .G. M. & Hasenstab Johnst.) M. (I. I .Jhs. aesa .G. M. & Hasenstab Johnst.) M. (I. L .Hgis .B ely comb. Kelley, B. R. Higgins) C. (L. sePyo 97 bas1951; Abrams 1927; Payson (see L .Hgis aesa .G. M. & Hasenstab Higgins) C. (L. .L oy18613 Cory L. V. .H rhm7924 Graham H. E. I .Jhs. aesa .G. M. & Hasenstab Johnst.) M. (I. S .Wlh .B ely comb. Kelley, B. R. Welsh) L. (S. aevldypbihdnames published validly have S as)Gen var. Greene Wats.) (S. .F ae 95 Baker F. C. .M ontn .Arnold J. Johnston, M. I. .L es 20470 Welsh L. S. .C ar 288 Parry C. C. .W hmsn14129 Thompson W. J. rpatahypsophila Cryptantha rpatajohnstonii Cryptantha hltp:GH). (holotype: rpatacrassipes Cryptantha .C ign,Great Higgins, C. L. rpatabarnebyi Cryptantha rpataatwoodii Cryptantha .D tod2624 Atwood D. N. hltp:GH). (holotype: .L es,Great Welsh, L. S. hltp:GH). (holotype: hltp:RSA; (holotype: Cryptantha Oreocarya GH). , nitida (holo- .C. L. 8 roay subcapitata Oreocarya 18. 7 roay sobolifera Oreocarya 17. 4 roay schoolcraftii Oreocarya 14. roosiorum Oreocarya 13. 0 roay suffruticosa Oreocarya 20. suffruticosa Oreocarya 19. 1 roay suffruticosa Oreocarya 21. shackletteana Oreocarya 16. semiglabra Oreocarya 15. 2 roay ochroleuca Oreocarya 12. lce o,uprMra as ,0 t 0Sp1909, Sep Montana, 10 ft, A.: 6,000 n. S. s. Pass, Jones U. E. Marias M. 1927.—TYPE: upper Co., 305. Glacier 14: Gard. Bot. Basionym: Rd. .B ely ob o. tt o.Basionym: nov. stat. nov., comb. pustulosa Kelley, Oreocarya B. R. (Rydb.) A.: south-facing1968, July S. 26 Eagle, of steep northwest mile U. 1 about on Bluff, Eagle 1969.—TYPE: of slope Co., 28. Fairbanks Southeast 29: Alaska, Naturalist Basin Great Basionym: 1953, Aug 13 6015 ft, Badger of 10,600 east mi 3 Flat, Mts., Inyo of crest Co., Basionym: Inyo California: nov. comb. Simpson, roosiorum G. M. and Canyon Red 1968, Jul of 1788 Higgins 21 south C. Co., L. m Garfield southwestern 100 in outcrop Campground on Co., Utah: A.: Garfield S. U. 1968.—TYPE: 197–198. 28: Naturalist Basin 1968, May 25 an,3 u 1911, Jul 31 Moun-tains, Elk Canyon, Hammond Utah. USA: 1913.—TYPE: ait4:25 96—YE S:Ua.Kn County. 1984, Kane Jun 6 Utah. USA: 1986.—TYPE: T41S,R5W,S26,JohnsonCanyon,LonePinePoint,1,830m, 255. 46: ralist arenicola Jun 23 Dam, Boysen 1980, of west Just Co.; Fremont Wyoming; Madron Lichvar, Basionym: nov. comb. ye H stps ,L,MC,M,N,PI,US). PHIL, NY, MO, MICH, LL, E, isotypes: 1886, GH; Sep type: Chihuahua, 20 Norte, MEXICO: del 1916.—TYPE: Paso near 35. 48: Herb. Basionym: Gray nov. comb. Kelley, multicaulis B. R. Macbr.) Basionym: nov. A. S. 1942, Jun U. 5 4829 Fredonia, 1943.—TYPE: Barneby and of Ripley 197. E miles 3: 2 Co., Coconino Bot. Arizona: W. Basionym: Leafl. Barneby, nov. comb. Simpson, 1984, Jun 23 Corall, Bernards 8775 of Tiehm N A. mile Yellow road Co., 1 Washoe area, Nevada, Hills A.: S. U. 1986.—TYPE: 104. o.Basionym: nov. NY). GH, CAS, .S .Ua:EeyC. rmWosd oCsl Dale, Castle to Woodside from Co., Emery Utah: A. S. U. 1968.—TYPE: 195. 28: Naturalist Basin Great Higgins, C. L. L .Hgis&S .Wlh .B ely ob nov. comb. Kelley, B. R. Welsh) L. S. Basionym: & Higgins C. (L. hltp:RA stps A,GH). CAS, isotypes: RSA; (holotype: .L es n .Moe8629 Moore G. and Welsh L. S. .D on3459 Dorn D. R. .C ign .L es,GetBsnNatu- Basin Great Welsh, L. S. & Higgins C. L. uz ls :14 95—YE .S A.: S. U. 1955.—TYPE: 124. 3: Aliso Munz, .C ign 14296 Higgins C. L. rpataschoolcraftii Cryptantha Tr. reevar. Greene (Torr.) rpatasobolifera Cryptantha rpatacinerea Cryptantha .C ign 1310 Higgins C. L. hltp:N;ioye:CS O RSA). MO, CAS, isotypes: NY; (holotype: hltp:RA stp:GH). isotype: RSA; (holotype: rpataochroleuca Cryptantha ˜ rpatashackletteana Cryptantha .A ybr 9320 Rydberg A. P. hltp:BY stps H NY). GH, isotypes: BRY; (holotype: 83:19 91—YE .S A.: S. U. 1981.—TYPE: 159. 28(3): o yb,Bl.Tre o.Cu 0 480. 40: Club Bot. Torrey Bull. Rydb., hltp:R;ioye NY). isotype: RM; (holotype: Pyo)R .Kle,cm.nov. comb. Kelley, B. R. (Payson) L .Hgis .B ely comb. Kelley, B. R. Higgins) C. (L. Mn)R .Kle,Hasenstab, Kelley, B. R. (Munz) hltp:CS stp:GH). isotype: CAS; (holotype: Brey aesa .G. M. & Hasenstab (Barneby) Dr iha)R .Kelley, B. R. Lichvar) & (Dorn Tem .B ely ob nov. comb. Kelley, B. R. (Tiehm) Tre)Gen var. Greene (Torrey) L .Hgis .B ely comb. Kelley, B. R. Higgins) C. (L. Tre)Gen var. Greene (Torrey) Tre)Gen var. Greene (Torrey) rpatasubcapitata Cryptantha hltp:BRY). (holotype: .C os&A .Roos R. A. & Roos C. J. Gen)Coqitvar. Cronquist (Greene) hltp:BY isotypes: BRY; (holotype: laxa asn n.Missouri Ann. Payson, .G rnl 776 Pringle G. C. hltp:BRY). (holotype: ih,Bitna38(2): Brittonia Tiehm, rpatasemiglabra Cryptantha hltp:NY). (holotype: .F ab. Contr. Macbr., F. J. .C ign,Great Higgins, C. L. .C Higgins, C. L. Cryptantha laxa pustulosa Oreocarya arenicola on& Dorn (holo- J F.(J. Delivered by Publishing Technology to: Michael Simpson IP: 174.101.35.52 on: Wed, 11 Jul 2012 04:29:57 Copyright (c) American Society for Plant Taxonomists. All rights reserved. J welshii Oreocarya 23. 5 ontnlaangustifolia Johnstonella 25. angelica Johnstonella discussion). 24. earlier (see here genus this recognized of varieties), needed members three as are with (one combinations species New eleven for treatments.) recent in ferent Cryptanthinae and the in lignosa 1925) rare is Johnston of which outside see duration, in genus; perennial is the for inaequata J. type the ignated 7 ontnladiplotricha Johnstonella 27. costata Johnstonella 26. lhuhi oetx h uece r mle n more and existing smaller have in varieties, are names two tubercles with one the species, tubercles, Two taxa numerous. whitish some spaced, often in sculpturing regularly although nutlet prominent, The of shape. het- consists are and species size most in of narrowly-winged eromorphic nutlets often the margin, and rounded) angled, (rarely sharply have “knife-like,” usually shape, a in lance-ovate) (rarely ovate or triangular .Ntessraig.. . spreading Nutlets 1. 37 [Volume BOTANY SYSTEMATIC suffruticosa Oreocarya 22. 754 e otegnr fteCytnhnei rsne ee oiid(npr)fo ely(02 n elye l 21) enote We (2012). al. et Kelley and (2012) Kelley from part) (in modified here, presented is genus Cryptanthinae the the that of genera the to key A OHNSTONELLA Johnstonella .Clxlbsi rueul pe >ohr,prl ue,ace vr1nte,wt –0sotsie ahwith each spines stout 5–10 with nutlet, 1 over arched fused, partly others, >> 2 upper unequal, fr in lobes Calyx 2. stps Y WTU). 1979, Campground NY, isotypes: Jun Creek Hot 5 down 2.1 Road, 42(2): ca. Sunnyside, County, Naturalist Nye from Basin Nevada, miles A.: Great S. Higgins, U. 1982.—TYPE: C. 196. L. & Basionym: Thorne nov. comb. Kelley, hltp:CS stps A,G,U,US). UC, GH, Guarda CAS, 1921, la isotypes: Jun 1924.— CAS; 30 de (holotype: 1143. Island, Angel Pond 12: opposite California: 4, Island, Baja ser. Sci. MEXICO. Acad. TYPE: Calif. Proc. Johnst., Basionym: nov. comb. Simpson, 90—YE .S . th erFr oe 7Jn1901, Jun n. 27 s. Cove, Jones Fort E. M. near Utah, A.: S. U. 1910.—TYPE: RETN.LsAds ewe noaat and Antofagasta between 1885, Andes: Jan Calalaste, Los 57.—TYPE: 1891: Chile ARGENTINA. de Santiago Nac., Mus. Basionym: Anales Phil., nov. comb. Simpson, A. S. 1859, U. Apr Brandegee 18 1899.—TYPE: S. Springs, J. Borrego 453. Co., Diego 27: San Basionym: California: Gaz. Bot. nov. Brandegee, comb. Simpson, 1853, Yuma, Fort Co., 1857.—TYPE: Yuma NY). 363. Arizona: (holotype: 5: A. Rep. S. Rail. U. Pacif. Greene, (Torrey) Basionym: nov. comb. Simpson, oe)R .Kle,cm.nv Basionym: nov. comb. Kelley, multicaulis B. R. Jones) I .Jhs. rn,hsntbe eonzda dif- as recognized been not has Brand, Johnst.) M. (I. okdbite;ntes.. . nutlets bristles; hooked Johnstonella Oreocarya Wto)Bad(rn 1925). (Brand Brand (Watson) rn,Rpr.Se.Nv en e.2:29 1925. 249. 21: Veg. Regni Nov. Spec. Repert. Brand, ossso lnswt ult htaeusually are that nutlets with plants of consists Plagiobothrys var. hltp:UC). (holotype: hltp:RSA). (holotype: Avreyo hsspecies, this of variety (A . setosa : ...... 2 ...... racemosa J. .Philippi F. K .Ton .C ign)R B. R. Higgins) C. L. & Thorne H. (K. .H hre578 Thorne H. K. I .Jhso)Hsntb&M G. M. & Hasenstab Johnston) M. (I. Badge aesa .G. M. & Hasenstab (Brandegee) .E oe,Cnr .Bt 3 4. 13: Bot. W. Contr. Jones, E. M. .l splpyei norsuyadwl ieyrqiesgeainit eaaegnr ntefuture. the in genera separate into segregation require likely will and study our in polyphyletic is l. s. Tre)Gen var. Greene (Torrey) ...... Pi. aesa .G. M. & Hasenstab (Phil.) Tre)Hsntb&M G. M. & Hasenstab (Torrey) hltp:M;ioye:GH). isotypes: MS; (holotype: I .Jhso)Bad(des- Brand Johnston) M. (I. rpatawelshii Cryptantha rtihu angustifolium Eritrichium rpataangelica Cryptantha rtihu diplotrichum Eritrichium e oteGnr fteCryptanthinae the of Genera the to Key A ontnlaracemosa Johnstonella rpatacostata Cryptantha hltp:BRY; (holotype: .racemosa J. ontn4221 Johnston setosa Krynitzkia Thomas M E. (M. .H. K. .M. I. var. 9 ontnlafastigiata Johnstonella 29. echinosepala Johnstonella 28. 0 ontnlagrayi Johnstonella 30. 4 ontnlamicromeres Johnstonella 34. holoptera Johnstonella 33. 1 ontnlagrayi Johnstonella 31. 6 ontnlapusilla Johnstonella 36. 5 ontnlaparviflora Johnstonella 35. 2 ontnlagrayi Johnstonella 32. EIO aaClfri ot Bj California], [Baja 1939.—TYPE: 1921, Norte Mar 388. 3 20: Bay. California Animas Arbor. Las Arnold Baja J. MEXICO. Johnst., M. Basionym: I. nov. comb. Simpson, G. M. 1913, Mar California 1 GH). (holotype: [Baja Island, California Magdalena Basionym: Sur]: Lower nov. MEXICO. —TYPE: comb. Simpson, echinosepala G. M. ye GH). type: Vsy&Rs)J .Mcr var. Macbr. F. J. Rose) Basionym: & nov. (Vasey comb. Simpson, G. M. var. Simpson .S .Aioa aPzC. hebr,1876, Ehrenberg, Co., 1877.—TYPE: Paz 81. La GH). 12: Arizona: (holotype: Arts A. Acad. S. Amer. U. Proc. Gray, Basionym: A. nov. comb. Simpson, 1924.—TYPE: US). 1146. UC, GH, 12: 4, 1921, ser. May 30 Francisco Island, San Sci. California Sur: Baja MEXICO. Acad. Calif. Proc. .G ipo,cm.nv,Basionym: meres nov., comb. Simpson, G. M. var. Simpson aaClfri Sr:SnJs e ao u 1897, Jun Cabo, del 347 Anthony Jose MEXICO. San 46.—TYPE: [Sur]: 48: California ser. Baja n. 1916. Herb. Gray Contr. hl,F.Aaa.3.16.TP:CIE adr,Dec Caldera, CHILE. 1853, 1860.—TYPE: 39. Atacam. Fl. Phil., Basionym: nov. comb. Simpson, 1883.—TYPE: 90. Cruz, 19: Santa 2472 Co., Arts Cruz Santa California: Acad. A. S. Amer. U. Proc. Gray, Basionym: A. nov. comb. Simpson, ipo var. Simpson .S .Txs iln o,LaoEtcd,3 Mar 30 Estacado, Llano 1856.—TYPE: Co., 171. Midland 2: Texas: 1852, Rep. A. Rail. S. Pacif. U. Gray, A. & Basionym: Torr. nov. comb. Simpson, G. EIO aaClfri Sr:Lgo ed –5Mar 6–15 Head, 1889, Lagoon 1888.—TYPE: [Sur]: 596. 11: California Mus. Baja Natl. MEXICO. S. U. Proc. Rose, & Vasey hltp:GH). (holotype: .Ple 801 Palmer E. Philippi Pope A ry reevar. Greene Gray) (A. .M ontn3947 Johnston M. I. hltp:NY). (holotype: .F ab. ot.Ga eb 6 7 1918. 57. 56: Herb. Gray Contr. Macbr., F. J. hltp:GH). (holotype: grayi hltp:MS). (holotype: cryptochaeta nesiotica ob o.Basionym: nov. comb. , hltp:N;ioye H UC). GH, isotype: NY; (holotype: Vsy&Rs)Hsntb&M G. M. & Hasenstab Rose) & (Vasey Vsy&Rs)Hsntb&M G. M. & Hasenstab Rose) & (Vasey Vsy&Rs)Hsntb&M G. M. & Hasenstab Rose) & (Vasey Tre .Ga)Hsntb&M. & Hasenstab Gray) A. & (Torrey I .Jhso)Hsntb& Hasenstab Johnston) M. (I. A ry aesa .G. M. & Hasenstab Gray) (A. I .Jhso)Hsntb& Hasenstab Johnston) M. (I. A ry aesa .G. M. & Hasenstab Gray) (A. Pi. aesa .G. M. & Hasenstab (Phil.) J .Mcr)Hsntb& Hasenstab Macbr.) F. (J. J .Mcr)Hsntb& Hasenstab MacBr.) F. (J. hltp:CS isotypes: CAS; (holotype: cryptochaeta .M ontn3505 Johnston M. I. rtihu parviflorum Eritrichium rtihu micromeres Eritrichium rtihu holopterum Eritrichium nesiotica rtihu pusillum Eritrichium rpatafastigiata Cryptantha rpatamicro- Cryptantha .R rut15 Orcutt R. C. rpatagrayi Cryptantha rntkagrayi Krynitzkia .M Johnst., M. I. .F Macbr., F. J. Cryptantha Harpagonella .E Jones E. M. Palmer (holo- .W. A. Delivered by Publishing Technology to: Michael Simpson IP: 174.101.35.52 on: Wed, 11 Jul 2012 04:29:57 Copyright (c) American Society for Plant Taxonomists. All rights reserved. rn,A 1927. A. Brand, rn,A 95 rinu atne a Cryptantheae. dar Gattungen neue Drei 1925. A. 1873. Hooker. Brand, D. and J. evolution and The G. 2000. Bentham, Baker. M. A. and Jesson, K. L. H., C. S. Barrett, hetero- of breakdown and functioning evolution, The 1966. G. H. Baker, classifica- Group Phylogeny Angiosperm the of update An 2009. southeastern APGIII. the in Boraginaceae of identification. genera model The 1991. statistical A. at I. look Al-Shehbaz, new A 1974. H. Akaike, rn,A 91 orgnca-orgnieeCytnha.P.22 Pp. Borraginaceae-Borraginoideae-Cryptantheae. 1931. A. Brand, 1951. L. Abrams, pro- for Guilliams B. Matt Ronald sequences. C. Dr. outgroup and and some Columbus nomenclature, viding Travis with J. assistance Dr. for draft, helpful Kelley earlier providing for an McDade on Plant Lucinda comments Native Dr. California funding, the generous and for Society Fund Research Student Graduate ciation 02 AESA-EMNADSMSN HLGNTCSSEAISO RPATA755 CRYPTANTHA OF SYSTEMATICS PHYLOGENETIC SIMPSON: AND HASENSTAB-LEHMAN 2012] nlr .adK rnl 1897. Prantl. K. and A. small Engler, for procedure isolation DNA rapid A 1987. Doyle. L. J. and J. J. Doyle, us .H n .H igr 03 oaiaeecmsaeexclusively are cymes Boraginaceae 2003. Hilger. H. H. and H. M. Buys, .Ntes±eet... . . erect ± Nutlets 1. o,G 1837. G. Don, K. P. and Reveal, L. J. Holmgren, H. N. Holmgren, distylous H. A. in A., Cronquist, Self-incompatibility 1985. B. B. Casper, Calvin Acknowledgments. .Crlawie aeyylo;ctldn o sal oe,i oe litgnspeet.. . present cleistogenes lobed if lobed, present usually not not cleistogenes cotyledons 2-lobed; yellow; cotyledons rarely spines (red-orange); white, without yellow Corolla nutlet, or 1 orange over limb arched tube, 3. not Corolla free, others, > 2 3. upper unequal, if or, equal ± fr in lobes Calyx 2. Vegetabilis Vegetabilis Regni Novarum Specierum plants. flowering in polymorphisms Botany stylar of function Plumbaginaceae. The 349–368. 20: I. systems. incompatibility morphic III. Society APG Linnean plants: the flowering of of Journal families and orders the for tion 1–169. 1: States. United Control Automatic on Transactions Press. University Stanford in uniiso rs eftissue. leaf fresh of quantities botany) and gardening Cryptantha of system general eena herb perennial helicoid. not and scorpioid Engelmann. Wilhelm epi:Engelmann. Leipzig: A. S. Asteraceae) U. west, intermountain 1984. Holmgren. (Boraginaceae). Evolution and atics ˜ ,A n .Glto 03 litgm nterr ihAndean high rare the in Cleistogamy 2003. Galetto. L. and A. o, .Nteswt eta aail roe( groove (adaxial) . ventral a with keel Nutlets (adaxial) ventral a with 4. Nutlets 4. a Pflanzenreich Das u ihhoe rsrih rcls ult e 4 gen nutlets prickles; straight or hooked with but .Pat nul inil rprnil e alrta ie aeyruddt uho–ie aro sal o e or red not usually taproot cushion–like; to rounded rarely wide, than taller gen perennial, or biennial, annual, Plants 5. .Pat nul e ie hntl,otnruddt uho-ie aro e rprl hndry when purple or red taproot cushion-like; to rounded often tall, than wider gen annual, Plants 5. 5 253–265. 85: eea itr fteDclmdosplants Dichlamydeous the of history general A .33 odn .K:GletadRivington. and Gilbert K.: U. London, 373. p. , rn:NwYr oaia Garden. Botanical York New Bronx: . 4 59. 24: .Clxo itntspl,itc nfut yoae>nte . nutlet > gynobase fruit; in intact sepals, distinct of Calyx 6. .Pat nulo aeybena rprnil eeaielae eeal aln,rrl aa;ntessot,papillate, smooth, nutlets basal; rarely cauline, generally leaves vegetative perennial; or roughened, biennial variously rarely or or rugulose, annual smooth, Plants nutlets tufted; or basal 7. leaves vegetative perennial; or biennial Plants 7. .Clxbslyfsd uua,tb icmcsiei ri;gnbs nutlet < gynobase fruit; in circumscissile tube tubular, fused, basally Calyx 6. Hemisphaerocarya upewe r;foe rcsasn rrrl rsn . . present rarely or absent bracts flower dry; when purple otnsann ebru ae) lrlbat present bracts floral paper); herbarium staining (often lutae lr ftePcfcStates Pacific the of flora Illustrated ora fteAnl Arboretum Arnold the of Journal .Pat nul lwr hsoaoso litgmu;nteslnelt,lneoae roae sal homomorphic, usually ovate, or lance-ovate, lanceolate, nutlets cleistogamous; or chasmogamous flowers annual; Plants 8. .Pat nul aeybena prnilin (perennial biennial rarely annual, Plants 8. rpatacapituliflora Cryptantha h e Phytologist New The ueclt,o uiae pclynrol ct oauiae eta roeae lott ultae . . . apex nutlet to almost apex groove ventral acuminate, to acute narrowly . apically . muricate, or apex tuberculate, nutlet below well apex groove obtuse,ventral to rounded broadly apically ...... 3 ...... 3:41–50. 237: nemuti lr:vsua lnso the of plants vascular flora: Intermountain etakteJsu reNtoa akAsso- Park National Tree Joshua the thank We agnruddo nld o igd(except winged not angled, or rounded margin moho ailt/ueclt,i ueclt,tbrlsgn o hts . . . whitish not gen. tubercles tuberculate, if papillate/tuberculate, or smooth . . winged, . narrowly often whitish rounded), usually (rarely tubercles angled tuberculate, margin sparsely sculpturing, or and densely size in heteromorphic often triangular-ovate, ieaueCited Literature 97,ed.A.Engler.Leipzig:Verlagvon Taxon 6:105–121. 161: . htceia Bulletin Phytochemical eetru pceu oau Regni Novarum Specierum Repertorium oue4. Volume eeaplantarum Genera C1:716–723. AC-19: i Natu Die 2 719–724. 52: 9 149–154. 99: 1 249–254. 21: (Boraginaceae). oue4(Corolliflorae): 4 Volume . ¨ ucasAtrde(except Asteridae Subclass lce Pflanzenfamilien rlichen ...... upeetr Series Supplementary Cryptantha odn Reeve. London: . . o.III Vol. 9 11–15. 19: Oiia Title: (Original rpataflava Cryptantha ln System- Plant ...... Repertorium Stanford: ...... 5 s.l.) ...... nasof Annals Evolution Botanical .racemosa J. IEEE A ...... 6 ...... oyoa .pterocarya C. oxygona, C...... 7 ...... ;foesawy hsoaos ult vt or ovate nutlets chasmogamous; always flowers ); ...... ign,L .17.Brgnca ftesuhetr ntdStates. United southwestern the of Boraginaceae 2007. 1979. C. Seberg. L. O. Higgins, and of Culham, revision A A. 1971. C. Brummitt, L. Higgins, K. R. H., V. heterocolpate Heywood, of Ultrastructure 2003. Simpson. G. M. and L. Hargrove, Gu ice,F .L n .A ee.1841. Meyer. A. C. and L. E. F. Fischer, ice,F .L n .A ee.1836. Meyer. A. C. in and L. relationships E. and F. analysis Fischer, Phylogenetic 1999. M. D. Ferguson, ry .18.Cnrbtost h oayo ot America. North of botany the to Contributions 1885. A. Gray, ree .L 87.Sm etAeia seioie I. Asperifoliae: American west Some 1887a. L. E. Greene, ulim,C .adB .Blwn 01 eor fafeun flier– frequent a of Memoirs 2011. Baldwin. G. B. and M. C. Guilliams, esnti,J 91 vltoaytesfo N eune:Amaximum A sequences: DNA from trees Evolutionary 1981. J. Felsenstein, otcln,M,H .Hle,adN in.20.Scnaysrcueof structure Secondary 2001. Diane. N. and Hilger, H. H. M., of Gottschling, amounts large with plants from DNA of Isolation 2005. A. E. Friar, ree .L 87.Sm etAeia seioieIII. Asperifoliae American west Some 1887c. L. E. Greene, the of distribution and biology reproductive form, Life 1983. J. Grau, ree .L 87.Sm etAeia seioie II. Asperifoliae: American west Some 1887b. L. E. Greene, ulim,C .adB .Blwn 00 hlgntcrltosisof relationships Phylogenetic 2010. Baldwin. G. B. and M. C. alignment Guilliams, sequence DNA Peaks 4 2004. Groothuis. T. and A. Griekspoor, ¨ k,R .A .adH .T am.1899. Harms. T. A. H. and M. A. L. R. rke, ra ai Naturalist Basin Great Series Biological Bulletin Science University Young world the of families plant Flowering Sciences in pollen Engelmann. W. Leipzig: Prantl. Natu h oaia oit fAeiaadteSceyfrEooi Bot- Economic A. for S. Society U. the Missouri, Louis, and St. America Taxonomists, any. of Plant of of Society meeting Society Botanical joint American the a Society, 2011, Fern Botany American America. the South and America North eiu S.Petersburg] [St. Seminum 3 253–268. 23: on based Hydrophyllaceae fteAeia cdm fAt n Sciences and Arts of Academy American the of Hamburg Pary. in P. Hamburg: Vereins Naturwissenshaftlichen des oeua hlgntceiec o utpeln-itnedis- ( long-distance flowers multiple popcorn for of persals evidence phylogenetic molecular A. Botanical S. U. the Island, and Rhode Providence, Taxonomists, America. Plant of Society of Society American Society, Petersburg] approach. likelihood h T1tasrp n t plcto narcntuto fthe of reconstruction a in application Boraginales. its the and of phylogeny transcript ITS1 the Press. Elsevier/Academic Diego: data in biochemical 3–14 Pp. metabolites. secondary aiona/hla genus Californian/Chilean rN IS n pN ( 3 cpDNA and (ITS) nrDNA flowers popcorn the mekentosj.com. software. 0 trnK ¨ lce Pflanzenfamilien rlichen ...... nrn.Btn 00 on etn fteAeia Fern American the of meeting joint a 2010, Botany intron). 6:137–151. 164: Cryptantha :52. 7: ...... ihgnrlytohdt nuaewings), undulate to toothed generally with ...... 4 ...... atB d.Zme .A n .H olo.San Roalson. H. E. and A. E. Zimmer eds. B, Part , ...... ora fMlclrEvolution Molecular of Journal ...... 9 293–350. 39: (Boraginaceae). Plagiobothrys :35. 2: trnL-trnF Cryptantha Cryptantha eit,e.H .A nlr .A E. A. K. Engler, A. G. H. ed. Regist., ndhF Plagiobothrys ln Biology Plant e:RylBtncGardens. Botanic Royal Kew: . eunedata. sequence oeua vlto:Pouigthe Producing evolution: Molecular Brgnca) nerdusing inferred (Boraginaceae), negncsae,r spacer, intergenic Krynitzkia p 3–4 in 231–240 Pp. . nentoa ora fPlant of Journal International subgenus rpataglomerata Cryptantha oaiaee between Boraginaceae) , 2 257–310. 12: :629–636. 3: Greeneocharis . :1–62. 8: ,e.K Kubitski. K. ed. 7, ...... 8 ne eiu [St. Seminum Index 7 368–376. 17: Oreocarya Pittonia ytmtcBotany Systematic Plagiobothrys Pittonia Pittonia Cryptantha Greeneocharis p 6 in 460 Pp. . Sonderba s6intron, ps16 Johnstonella Proceedings Eremocarya :107–120. 1: Pectocarya Amsinckia Oreocarya . :55–60. 1: Brigham :8–23. 1: . Index ¨ .s. s. The .l. s. nde Delivered by Publishing Technology to: Michael Simpson IP: 174.101.35.52 on: Wed, 11 Jul 2012 04:29:57 Copyright (c) American Society for Plant Taxonomists. All rights reserved. rdf,R 96 h erdciesse of system reproductive The 1976. R. Ornduff, author. the by distributed Program v2. MrModeltest 2004. A. A. J. Nylander, uz .A n .D ek 1968. Keck. D. D. and A. P. Munz, 1959. Keck. D. D. and A. P. Munz, La hw . .B iky .T ek .B amr .Lu .Mle,K C. K. Miller, J. Liu, W. Farmer, B. S. Beck, T. J. Lickey, B. E. J., Shaw, 2007. C. Schwarzer, Evolutionary 1997. L’Heureux. A.-M. and Johnston, O. M. J., D. Schoen, crı Estudios 1910. K. Reiche, asn .B 97 oorp ftesection the of monograph A 1927. B. E. Payson, or,M . .Te n .K asn 06 atrso long-distance of Patterns 2006. Jansen. K. R. and Tye, A. J., M. Moore, 37 [Volume Hawksworth, L. D. Demoulin, V. Burdet, M. H. Barrie, R. system F. modular J., 4.08. a McNeill, Mesquite: Version 2010. Maddison. MacClade R. D. 2005. and Maddison. P. W. P. Maddison, W. and R. D. Maddison, 2008. J. D. Mabberley, BOTANY SYSTEMATIC otherwise or New XI. Boraginaceae the in Studies the 1935. of M. revision I. A Johnston, VI. Boraginaceae the in Studies 1927. M. I. Johnston, American North The IV. Boraginaceae the in Studies 1925. of M. I. inference Johnston, Bayesian MrBayes: 2001. Ronquist. F. and P. J. Huelsenbeck, 756 ely .A n .Wle.1993. Wilken. D. and A. W. 2012. Kelley, Hasenstab. E. K. and Simpson, G. M. B., R. Kelley, 2012. B. R. Kelley, a,P .adH .Ciai 97 tde on Studies 1957. Chisaki. F. H. and M. and P. North Ray, of floras the in relationships Amphitropical 1963. nucle- H. of P. model Raven, best-fit the Selecting 2001. Crandall. A. K. and DNA D. of Posada, model the testing Modeltest: 1998. Crandall. A. K. and D. Posada, ˚ ngstro itlu species. distylous University. Uppsala Centre, Biology Evolutionary Press. California of University Berkeley: fClfri Press. California of Botany of Journal American Press. California of California versity of plants Higher manual: iiu,C .Wne,E .Shlig n .L ml.20.The 2005. Small. chloroplast noncoding L. 142–166. 21 92: R. of analysis. phylogenetic and utility for Relative Schilling, sequences II: DNA E. hare the E. and Winder, tortoise T. C. Siripun, Universita (Boraginaceae). Freie Don G. Lehm., Gattungen der Vertretern in 1090–1099. system 57: mating the of history Cervantes. in Imprenta 217–236 Pp. it. in 529–536. heterostyly 44: of study a with genus, the rgn fapirpcldsucsadGala and disjuncts amphitropical of origins in dispersal H. J. Skog, E. J. http://ibot.sav.sk/icbn/main.htm. Silva, KG. Vegetabile Regnum C. Code). P. (Vienna clature Prado, 2006. J. Turland. J. Nicolson, N. and H. Wiersema, D. Marhold, K. http://mesquiteproject.org. 2.73. Version analysis. evolutionary for Associates. Sinauer Sunderland: Press. University uses and classification their plants, higher data. sequence and of placement and (Boraginaceae) species. noteworthy University Harvard of ium Boraginoideae. American South University Harvard of species trees. phylogenetic dto,es .G adi,D .Glmn .J el .Patterson, R. Press. California Keil, of J. University D. Berkeley: Goldman, Rosatti. H. J. D. T. Baldwin, and G. B. eds. Edition, in 455–468 Pp. University Berkeley: Rosatti. Press. California J. of T. and Patterson, R. Keil, California J. in D. Goldman, of 502–509 Plants 484–485, lar 475, 453–454, Pp. ot America. South substitution. otide substitution. Garden Botanical Missouri the of Annals ¨ ,E n .W hs.20.Tie fBoraginoideae of Tribes 2002. Chase. W. M. and E. m, Sericostoma Plagiobothrys Cryptantha Tiquilia Bioinformatics h esnmna:Vsua lnso California of plants Vascular manual: Jepson The ln ytmtc n Evolution and Systematics Plant hlgntcaayi ae on based analysis phylogenetic A : ¨ h urel eiwo Biology of Review Quarterly The t. ytmtsh nescugna e peruanischen den an Untersuchungen Systematische mici,Hraoel,Pcoay,Plagiobothrys Pectocarya, Harpagonella, Amsinckia, ytmtcBotany Systematic lr eChile de Flora ytmtcBiology Systematic abre’ ln-ok otbedcinr fthe of dictionary portable A plant-book: Mabberley’s 1–114. 74: Bioinformatics subg. ora fteAnl Arboretum Arnold the of Journal ic.&C .My und Mey. A. C. & Fisch. .SecondEdition,eds.B.G.Baldwin,D.H. . otiuin rmteGa ebru of Herbarium Gray the from Contributions ´ 8 1–118. 78: ilmreti ahBooi.Berlin: Biologie. Fach im Diplomarbeit io or afoad Chile: de flora la sobre ticos Tiquilia 3 1163–1177. 93: 4 817–818. 14: Pectocarya aionaFlora California A Cryptantha Amsinckia upeett lr fCalifornia of Flora A to Supplement nihtm cicio,Ogastemma Echiochilon, Antiphytum, otiuin rmteGa Herbar- Gray the from Contributions 7 754–755. 17: Brgnca) mlctosfrthe for implications (Boraginaceae): d .S ika.Bree:Uni- Berkeley: Hickman. S. J. ed. , nentoa oeo oaia nomen- botanical of code International :57–66. 1: o.5 d ece Santiago: Reiche. ed. 5, vol. 0 580–601. 50: r d abig:Cambridge Cambridge: ed. 3rd . 4 211–358. 14: ..e Meisn., ex D.C. 4.A .G ate Verlag Gantner G. R. A. 146. h esnMna:Vascu- Manual: Jepson The p 6–7 in 369–378 Pp. . Amsinckia mrcnJunlo Botany of Journal American mrcnJunlo Botany of Journal American ´ (Boraginaceae). ao sad endemics. Islands pagos Oreocarya mici grandiflora Amsinckia 3:137–153. 234: ekly University Berkeley: . 8 151–177. 38: Cryptantha atpB 6 173–205. 16: .Asnpi of synopsis A I. . of lsi DNA plastid Cryptantha Cryptantha Cryptantha Amsinckia h Jepson The Evolution em ex Lehm. Second . ,a . . . . . JQ582307; confertiflora Reiche, 3108 Simpson circumscissa Cryptantha ont var. Johnst. JQ582303; JQ513401, 137256), var. Mey. A. C. ediko 525 JQ582295; Hendrickson JQ513393, 178190), holoptera elir .20.UaneaepceLen especie nueva Una 2009. S. Teillier, parsimony 1997. using L. analysis A. Takhtajan, phylogenetic PAUP*. 2003. L. D. Swofford, http://www bootstrap website. phylogeny rapid Angiosperm A onwards. 2001 2008. P. Rougemont. Stevens, J. and Hoover, P. A., Stamatakis, 2011. G. M. Simpson, ipo,M .adK .Hsntb 2009. Hasenstab. E. K. and G. M. Simpson, JQ582302; chaetocalyx 991122 Arroyo Kalin JQ582300; JQ513398, 3636210), (MO JQ582299; JQ513397, 13030), JQ582298; alyssoides Greene, Gray) 2791 JQ582293; JQ513391, 575,J531,JQ582321; JQ513419, 457775), JQ582320; JQ513418, JQ582319; Johnst., JQ582317; JQ513415, 124411), JQ582316; ganderi Cryptantha Greene, (Greene) DA9905 Anderson JQ582313; JQ513411, 457666), (COLO ee,W .1987. A. W. Weber, egn,M,M otcln,F ev,adH .Hle.21.Fsi and Fossil 2010. Hilger. H. H. and Selvi, F. Gottschling, M. M., Weigend, 1997. Jeanmougin. F. and Plewniak, F. Higgins, G. G. D. D., J. Thompson, i oeta n fagvntxn,hraimacsin n GenBank and accession, study. herbarium this taxon), in analyzed given and sequenced a specimens for of numbers accession one than more (if Cata 2008. eds. Belgrano, J. M. and Morrone, O. O., F. Zuloaga, heterostyly. of aspects Some 1975. F. P. Yeo, Fshr&CA e. Greene, Mey.) C.A. & (Fischer 16 Hasenstab i., JQ582312; JQ513410, dumetorum Cryptantha diplotricha .M Johnst., M. I. 11397 Rebman JQ582308; JQ513406, Appendix mici eastwoodiae Amsinckia aal lr eChile. de Flora la para Press. University Columbia York: Associates. Sinauer Sunderland: 4. Version methods). other (*and .mobot.org/MOBOT/research/APweb. servers. web RAxML the for algorithm Crossosoma www.sci.sdsu.edu/plants/cryptantha. xatwsenhmshr oaiee n h oyhl of polyphyly the and Boragineae, (Borag Riedl hemisphere “Trigonotideae” western extant Press. University Associated multiple tools. for analysis quality strategies by Research flexible aided interface: alignment sequence windows ClustalX The oorpsi ytmtcBtn rmteMsor oaia Garden Botanical Press. Missouri Garden Botanical the Acanthaceae- Missouri from Louis: St. Botany Dicotyledoneae: Systematic Chile, in 2 Monographs Brasil, volume de ( Sur Uruguay) Fabaceae (Argentina, y Sur Cono Paraguay del vasculares plantas 409–419. 35: SS 77) Q132 JQ582294; JQ513392, 17575), (SDSU egn 2000 Weigend ole 8570 Zollner A ry .F Macbr., F. J. Gray) (A. rpataangustifolia Cryptantha rpatahumilis Cryptantha Pi. Reiche, (Phil.) A C)Reiche, DC.) (A. rpataglobulifera Cryptantha rpatacinerea Cryptantha rpatacostata Cryptantha Pi. .M Johnst., M. I. (Phil.) circumscissa Gen)Payson, (Greene) 4 4876–4882. 24: uo 2745 Munoz SS 82) Q143 JQ582305; JQ513403, 18627), (SDSU .Txn olco/olcinnme,sml number sample number, collector/collection Taxon, 1. S 762,J530,JQ582309; JQ513407, 171652), (SD esrs n. s. Reiser 5 1–59. 35: Abarema-Schizolobium microcarpa SS 84) Q140 JQ582322; JQ513420, 18348), (SDSU S 761,J539,JQ582296; JQ513394, 173641), (SD CL 974,J531,JQ582314; JQ513412, 499734), (COLO emn6803 Rebman .M Johnst., M. I. / rpatainaequata Cryptantha mici intermedia Amsinckia rpatafendleri Cryptantha Cryptantha CN 694,J539,JQ582301; JQ513399, 166914), (CONC ooaoFoa etr slope Western Flora: Colorado rpatadecipiens Cryptantha 642 M 644) Q144 JQ582306; JQ513404, 2614248), (MO iest n lsiiaino lwrn plants flowering of classification and Diversity Ho.&An)I .Jhs.var. Johnst. M. I. Arn.) & (Hook. A ry Greene, Gray) (A. 1, ole .n. s. Zollner .F Macbride, F. J. M 370) Q148 JQ582310; JQ513408, 4317604), (MO SS 22) Q135 JQ582297; JQ513395, 12421), (SDSU M 059) Q146 JQ582318; JQ513416, 6045195), (MO Gen)Jp.&Hoover, & Jeps. (Greene) esrs n. s. Reiser Novon rpataaprica Cryptantha A ry Payson, Gray) (A. rpatacircumscissa Cryptantha rpataintermedia Cryptantha ilir5210 Tiellier rpatagranulosa Cryptantha Gen)Cronquist, (Greene) Brandegee, emap8417 Helmkamp Tre)Greene, (Torrey) nca:Boraginoideae). inaceae: mici tessellata Amsinckia aooyadiae e ie http:// site. web images and taxonomy hite229 Christie emap2588 Helmkamp Co)Reiche, (Clos) uo 2745 Munoz 9 401–403. 19: S 508,J531,JQ582315; JQ513413, 155048), (SD rpatacapituliflora Cryptantha at 183 Barth :Brgnca.P.13–63in 1631–1663 Pp. Boraginaceae. ): M 458) Q149 JQ582311; JQ513409, 2415685), (MO rpataflava Cryptantha ˜ S 343,J530,JQ582304; JQ513402, 134493), (SD s de osa Cryptantha A ry Greene, Gray) (A. h e Phytologist New The .M Johnst., M. I. ic.&C .Mey., A. C. & Fisch. ytmtcBiology Systematic emap6530 Helmkamp mici spectabilis Amsinckia M .Jns .A Heller, A. A. Jones) E. (M. CN 553,JQ513396, 156553), (CONC adr 6173 Sanders adr 7028 Sanders Pi. Reiche, (Phil.) Cryptantha M 370) JQ513400, 4317604), (MO oe 1089 Honer S 481,JQ513405, 140881), (SD RA703) JQ513421, 710334), (RSA S 631,JQ513414, 169361), (SD rpatadiffusa Cryptantha atnz31492 Martinez rpatacollina Cryptantha S 402,JQ513417, 144002), (SD ipo 8III98A Simpson A ry reevar. Greene Gray) (A. fsuhr California. southern of ole,Colorado: Boulder, . rpataaffinis Cryptantha Ri a. .M. I. Pav.) & (Ruiz Ho.&An)I M. I. Arn.) & (Hook. emap8336 Helmkamp rpataleiocarpa Cryptantha emn2994 Rebman .Ga var. Gray A. A es)Payson, Nels.) (A. rpatafoliosa Cryptantha ytmtcBotany Systematic od2221 Boyd (Boraginaceae) circumscissa RA682210), (RSA Co)Reiche, (Clos) S 124729), (SD S 125659), (SD uli Acids Nucleic S 148214), (SD 7 758–771. 57: elir1057 Teillier oa .n. s. Hogan ´ 5 147–153. 75: ood las de logo Cryptantha Cryptantha Cryptantha Cryptantha Cryptantha ic.& Fisch. Simpson (CONC (SDSU New . (Phil.) (Phil.) (RSA (SD (SD (A. t., 2, . Delivered by Publishing Technology to: Michael Simpson IP: 174.101.35.52 on: Wed, 11 Jul 2012 04:29:57 Copyright (c) American Society for Plant Taxonomists. All rights reserved. roosiorum Greene, 1IV95B 2, Greene (Greene) 3043 Simpson JQ582326; JQ513424, 163475), RA229) Q149 JQ582341; JQ513439, 272392), (RSA JQ582340; scoparia Cryptantha 757 CRYPTANTHA OF SYSTEMATICS PHYLOGENETIC SIMPSON: AND HASENSTAB-LEHMAN JQ582323; 2012] 1, micrantha Cryptantha pterocarya nevadensis Greene, 2273 Gross JQ582331; JQ513429, 503599), JQ582330; JQ513428, micromeres Cryptantha var. Johnst. var. Johnst. M. I. (Torr.) Johnst., 20532 Werff JQ582334; JQ513432, 172713), SS 77) Q143 JQ582325; JQ513423, 17572), (SDSU ofne 12 Wolfinger SS 43) Q145 JQ582337; JQ513435, 14135), (SDSU a e ef 20513 Werff der van adr 20412 Sanders emn6305 Rebman rpatalinearis Cryptantha rpatasimilis Cryptantha Tre)Gen var. Greene (Torrey) Munz, .Nl.&P .Kneyvar. Kennedy B. P. & Nels. A. M 181) Q143 JQ582335; JQ513433, 6128118), (MO RA722) Q140 JQ582332; JQ513430, 712528), (RSA . emn11358 Rebman m., SS 84) Q145 JQ582327; JQ513425, 18446), (SDSU S 856,J532,JQ582324; JQ513422, 189566), (SD eekr4846 DeDecker .Nels., A. Tr. .M ont var. Johnst. M. I. (Torr.) rpataminima Cryptantha iru 6185 Biurrun SS 55) Q146 JQ582338; JQ513436, 15452), (SDSU A ry Greene, Gray) (A. RA648) Q141 JQ582333; JQ513431, 614189), (RSA lepida M 181) Q144 JQ582336; JQ513434, 6128115), (MO .Mte .H Raven, H. P. & Mathew K. Cla Greene, (Colla) rpataparviflora Cryptantha eekr5977 DeDecker rpatamaritima Cryptantha rpatamohavensis Cryptantha A ry .M ont 2, Johnst. M. I. Gray) (A. cycloptera RA362) Q147 JQ582339; JQ513437, 346621), (RSA rpatasimilis Cryptantha S 641,J532,JQ582329; JQ513427, 168431), (SD rpatamicrantha Cryptantha S 76) Q146 JQ582328; JQ513426, 87769), (SI Rydberg, rpataracemosa Cryptantha Gen)J .Macbr., F. J. (Greene) nevadensis lepida ai roo993602 Arroyo Kalin ar 401 Lauri RA663) JQ513438, 626035), (RSA rpataperuviana Cryptantha rea 14292 Freeman A ry .M Johnst. M. I. Gray) (A. Pi. Reiche, (Phil.) rpatanemaclada Cryptantha rpatamicrantha Cryptantha rpatamaritima Cryptantha Gen)Gen 1, Greene (Greene) .Mte .H. P. & Mathew K. , Gen)Greene, (Greene) rgr 1305 Gregory ercsn17339 Henrickson SS 16695), (SDSU ipo 2816 Simpson Tr. .M. I. (Torr.) Cryptantha Cryptantha Cryptantha A Gray) (A. (CONC (COLO a der van Burch .M. I. (SD Raven, 181) Q149 JQ582356; JQ513459, 6128112), myosotoides Plagiobothrys Mn)Jeps., (Munz) JQ582350; JQ513451, 169347), JQ582349; JQ513450, penicillata Pectocarya Johnston, M. I. Lehm., color JQ582346; JQ513444, 467583), (COLO Payson, (Osterhout) 11672 A ry .M ont var. Johnst. M. I. Gray) (A. JQ582353; var. Gray JQ513456, A. 709039), (RSA JQ582352; JQ513455, 1731379), congestus Plagiobothrys distans Johnst., M. I. 5108 Howe JQ582345; JQ513443, 171991), thyrsiflora Cryptantha Q144 JQ582351; JQ513454, Q142 JQ582344; JQ513442, Pers., ES175) Q140 JQ582357. JQ513460, 107359), JEPS ( adr 21776 Sanders hr 13408 Ahart Benth., SS 68,J534,JQ582347; JQ513446, 2648), (SDSU hr 9593 Ahart . oe 992 Honer k., eel2390 Reveal ulim 570 Guilliams a e ef 20623 Werff der van ulim 692 Guilliams JP 146,J534,JQ582359; JQ513448, 110466), (JEPS RA604) Q140 JQ582342; JQ513440, 660941), (RSA Ho.&An)A DC., A. Arn.) & (Hook. rhe 977 Orthner etcrapeninsularis Pectocarya rpatatraskiae Cryptantha lgoohy hispidus Plagiobothrys Wd. .M Johnston, M. I. (Wedd.) JP 035,J534,JQ582358; JQ513447, 103365), (JEPS Gen)Payson, (Greene) S 089,J534,JQ582347; JQ513445, 103849), (SD RA609) Q147 0; JQ513457, 680594), (RSA Lh. Brand, (Lehm.) SS 95) Q142 JQ582360; JQ513452, 19853), (SDSU ols hr 9211 Ahart mollis, rpataweberi Cryptantha hclacrenulata Phacelia lgoohy jonesii Plagiobothrys lgoohy shastensis Plagiobothrys SS 95) Q143 JQ582361; JQ513453, 19859), (SDSU CL 746,J534,JQ582343; JQ513441, 475416), (COLO M 181) Q149 JQ582348; JQ513449, 6128114), (MO lgoohy kingii Plagiobothrys yolsu occidentale Cynoglossum .M Johnst., M. I. ot744 Holt .M Johnst., M. I. .Gray, A. a e ef 20645 Werff der van aklafloribunda Hackelia .M Johnst., M. I. .Wto var. Watson S. ar 189 Lauri JP 060,JQ513458; 102650), (JEPS eks n. s. Beck .Gray, A. lgoohy mollis Plagiobothrys alr16824 Taylor etcraanomola Pectocarya ua SN597 Junak CL 521709), (COLO rpatastricta Cryptantha .Gray, A. M 5183035), (MO SS 16855), (SDSU at 135 Barth adr 27585 Sanders ystslaxa Myosotis ystsdis- Myosotis oda .n. s. Rondeau S Watson) (S. minutiflora .Gray, A. (Lehm.) Phacelia Ahart (MO (UC (SD (SD