Delivered by Publishing Technology to: Michael Simpson IP: 146.244.235.206 on: Mon, 21 Sep 2015 23:37:14 Copyright (c) American Society for Plant Taxonomists. All rights reserved. eei nlssb egn ta.(03.I hi analysis, their In (2013). al. occidentale C. et Weigend by analysis genetic American of North three species other The 2014). 2011, (Cohen includes relatives which Hasenstab- [2012]), of Simpson and (Cryptanthinae Lehman Brand. Amsinckiinae with to sister data, phylogenetic by ported endemic genus Mexican the regated sn opooia vdne Sutory evidence, morphological Using i opooia etrswl econstructed. be will diagnos- features includes morphological that tic system American taxonomic of appropriate members North an other then the from relatives. distinct analysis and remain critical species genus upon the if of Additionally, species American North among of and phylogeny plastid a Cynoglossum use reconstruct to to regions is DNA study ribosomal present nuclear the of objective The includes and Hook. member that a North clade is the western species American of throughout South the relatives and America, its South and G.Don of ex diversification conti- explosive the the the in to colonized prior Indeed, that nent species group genus. American early-diverging North the the an (2013) of represent al. species et Weigend World of Old analyses the of that of species oo,adohr,aense mn h l ol spe- World Old the among nested of cies are others, and Popov, Solenanthus hsi u otofcos n sta utpeseisfrom species multiple that Paracaryum is One factors. including genera, two other to due has is this (2013) Sutory al. and Selvi et (e.g. defined Weigend and 2014) that demonstrated (2011, Cohen phylogenetic Recent by gynobase. work the bear to that attachment character- nutlets is submedial glochidiate Koch, and J. pollen D. heterocolpate W. by Cynoglosseae ized of genus, type The the 1995). is al. which et (Zhu distribution cosmopolitan a with aeo ulcto uy2,2015 23, July publication of Date 10.1600/036364415X688385 DOI © Botany Systematic oyih 05b h mrcnSceyo ln Taxonomists Plant of Society American the by 2015 Copyright orseisof species Four Cynoglossum contrast, Andersonglossum ihfu aetransfers: name four with ncnucinwt igotcmrhlgclfaue,tonwgnr r rpsdfrteespecies, these for proposed are genera new two features, morphological diagnostic with conjunction in inhp mn ebr ftegnsrmi neti.Ti spriual h aefrNrhAeia pce of species American North for case the particularly is This uncertain. remain genus the of members among tionships ebr facaedsic rmteOdWrdseis n hs ot mrcnseisbln otodsic lineages. distinct two to belong species American North these and species, World Old the species from World Old distinct and occidentale New clade the a among relationships of evolutionary members the investigate to reconstructed been of has phylogeny a data, sequence DNA Keywords— Abstract— Adelinia Cynoglossum Oncaglossum Cynoglossum Cynoglossum Boiss., 21) 02:p.611–619 pp. 40(2): (2015), Cynoglossum Ledeb., Adelinia .Ga and Gray A. n eaie nodrt eov relationships resolve to order in relatives and and eetpyoeei vdnesget that suggests evidence phylogenetic Recent .(oaiaee samdu-ie genus medium-sized a is (Boraginaceae) L. Amsinckiinae, .virginianum C. Rindera n te ebr fBrgnca.Tersligpyoeyrsle htNrhAeia pce of species American North that resolved phylogeny resulting The Boraginaceae. of members other and Cynoglossum er esl aln evs eiesrcre nfut n olnwt w hpso oe n otases roe In groove. transverse no and pores of shapes two with pollen and fruit, in recurved pedicels leaves, cauline sessile bears and eeosptoaecuielae,pdcl rc nfut n olnwt nyoepr hp n rnvregroove. transverse a and shape pore one only with pollen and fruit, in erect pedicels leaves, cauline petiolate develops Cynoglossum h eodi httefwNwWorld New few the that is second The . Trachelanthus Sutory etrn nvriy 22 B 70Uiest v. ln,Mcia 80,U .A. S. U. 48504, Michigan Flint, Ave., University 1700 AB, 2212C University, Kettering Cynoglossopsis r ebr fcae eaaefrom separate clades of members are Andersonglossum Pall., .virginianum C. aebe nlddi phylo- a in included been have ´ Omphalodes hssprto a ae sup- later was separation This . nesnlsu boreale Andersonglossum ´ Cryptantha 02,i o oohltc and monophyletic, not is 2012), Pardoglossum r itrseis and species, sister are r aiet ot America. North to native are .pringlei C. si a entraditionally been has it as , Klotzsch, Oncaglossum Brand, oeua systematics, molecular , il and Mill. ´ .wr itrspecies, sister were L. 21)rcnl seg- recently (2010) rem nothe into Greenm. abe Mathez, & Barbier Cryptantha Paracynoglossum Lindelofia Cryptantha pce of Species omnctn dtr hc Bell Chuck Editor: Communicating Cynoglossum Myosotidium , eovdas resolved Brgnca) w e eeafo e World New from Genera New Two (Boraginaceae), .grande C. nesnlsu occidentale Andersonglossum ([email protected]) Cynoglossum Lehm., ae .Cohen I. James Lehm. and Oncaglossum samme fasprt ru.Gvnteeeouinr relationships evolutionary these Given group. separate a of member a is , 611 Cynoglossum Brgnca) ompltngns sntmnpyei,btrela- but monophyletic, not is genus, cosmopolitan a (Boraginaceae), hlgntcaaye rvddeiec that evidence provided their analyses because Cryptanthinae) phylogenetic (as Amsinckiinae of taxonomy Johnst. M. I. including Amsinckiinae, of members virginianum C. h eainhpwsabgos h ld of clade The ambiguous. was relationship the and Greene, ini h rb.Sqec aafo orDArgoswr included were regions regions DNA (cpDNA) DNA four plastid protein-encoding ( from two study: data present Sequence the in tribe. Cynoglosseae. mor the in across both tion from represents S1) sampling Appendix This (supplementary species 55 and ieeiec htohrNwWrdmmeso Cynoglosseae of pro- members (2013) (e.g. World al. New other et that Weigend evidence and vide 2014) (2011, Cohen of results Harms, esfralseisi h rsn td swl svuhrinformation voucher as well as num- study sequences. present GenBank new the for includes in species S1 all Appendix for bers GenBank. from individuals from obtained obtained s were Additional was populations. isolations wild DNA from new inte for material (nrDNA) Leaf DNA ribosomal nuclear ei.T hsed h uhr eie h aooyfor genera: taxonomy nine the includes revised now authors Lehm., which the subtribe, end, the this To letic. swl sohrseista r uaieyrltdto related putatively of are position of members that isolated American the North species these given other North Furthermore, four as Amsinckiinae. of all well members been includes as have study that current species American the among relationships species, phylogenetic elucidate these to exam- adequately order relationships been In not these ined. have but or ambiguous Amsinckiinae, remain to either related closely are . tseslkl htteeNrhAeia pce will species American genera. North other to these transferred be that to likely need seems species type it the L., including genus, the of members matK ao apigadSqec Data— Sequence and Sampling Taxon n21,HsntbLha n ipo evlae the reevaluated Simpson and Hasenstab-Lehman 2012, In taxonomy. , Plagiobothrys .grande C. Cynoglossum and Cryptantha Harpagonella Pectocarya , ndhF nesnlsu virginianum Andersonglossum ,oecDAitrei pcr( spacer intergenic cpDNA one ), oga xLh.wscoeyrltd but related, closely was Lehm. ex Douglas and , aeil n Methods and Materials , Fisch.&C.A.Mey.werenotmonophy- Dasynotus Eremocarya C xMin,and Meisn., ex DC. .grande C. .Gray, A. Andersonglossum .M ont,and Johnst., M. I. Greene, Johnstonella hlgcladgorpi varia- geographic and phological qec aafrmlil species multiple for data equence omdatihtm with trichotomy a formed nltasrbdsae (ITS). spacer transcribed rnal h rsn td includes study present The Cynoglossum ,and Cynoglossum ayou daubenmirei Dasynotus Greeneocharis dlnagrande Adelinia Plagiobothrys Cynoglossum Brand, trnL and Cynoglossum .occidentale C. - Oncaglossum Utilizing . Cynoglossum trnF Adelinia rmother from .officinale C. Cryptantha Amsinckia Oreocarya ,adthe and ), Gu are ¨ k & rke .The . , + ) Delivered by Publishing Technology to: Michael Simpson IP: 146.244.235.206 on: Mon, 21 Sep 2015 23:37:14 Copyright (c) American Society for Plant Taxonomists. All rights reserved. aiencetd ie n 5srcua hrcesaepres- infor- are 343 characters of structural total 35 A and bp). sites (495 nucleotide shortest mative the ITS and [bp]) pairs ,5 uloie,with nucleotides, 5,458 1 YTMTCBTN Vlm 40 [Volume BOTANY SYSTEMATIC 612 eindb h uhr(oe 04.PRmxue,2 Li volume, in mL 25 mixtures, those PCR or 1 2014). primers of (Cohen published included consisted author the either the that of using by amplifications 1990) conducted designed PCR The Doyle were buffer. regions and extraction DNA CTAB (Doyle the in method PVP-40 2% extraction CTAB fied oe hsaon fmsigdt osntapa ocnieal affect considerably there- to phylogeny. appear 2014); resulting not Cohen the the does 2013; in data but include al. missing relationships of missing, that et amount Weigend analyses is this (e.g. fore, previous data sampling with sequence taxon congruent greater the is Ochoterena of and phylogeny (Simmons resulting 33% 2002). coding (Nixon indel Approximately 1.7 simple 2000). ver. using Winclada coded were in Gaps made were adjustments settings. Subsequent default server the MUSCLE at using Institute’s (http://www.ebi.ac.uk/Tools/msa/muscle/) Bioinformatics available European the is by matrix implemented the and S1), 17336). Aligner number (Appendix were (accession CodonCode TreeBASE GenBank Sequences with in Massachusetts). edited deposited Deadham, trace and Sequence Corporation, examined, University. (CodonCode compiled, Cornell were Life at the Center files by Analyzer, Laboratory DNA Core 3730 cycle ABI Sciences an automated with and performed, (ABI), was of sequencing method modification a acetate using ethanol/EDTA/sodium precipitated amplification the were locus-specific products and Sequencing California) primers. [ABI], Biosystems (Applied amplification the with Germany). if purified (Qiagen, agarose determine were kit 1% products purification to PCR PCR a sequencing, QIAquick bromide on to ethidium separated Prior occurred. with were had the Taq products stained listed PCR on and temperatures GXL The annealing gel depending PrimeSTAR (2014). and sample, Cohen of primers DNA the in Eppendorf mL an using mL with 1 Pro, 2.5 performed Mastercycler primers, were to Amplifications of mL concentration. DNA mM 0.1 1 and polymerase, dNTPs, mM 100 etnDs pte otr n hniae eecpue t1 kV 10 at captured were images 6610-LVM. then model and JEOL coater, a sputter with using 5 gold with Desk coated Denton stubs, on a mounted was material plant Dried scope. and evs lwr,ple,adfut of fruits and pollen, flowers, Leaves, trees resulting The iteration iterations. 5,000,000 (http://www.evolution.reading.ac.uk/ 10,000 V1.3 The BayesTrees.html). a of BayesTrees data. in frequency with Pagel the viewed print were iterations, for and a model Meade 20,000,000 and simplest 2008; burnin, included the 2004, analysis with Meade analyzed BI mix- and were and set Gaps regions, (Pagel length DNA 2008). branch (RJBLS) four reversible-jump the model the as of ture identified and each 2012) categories the al. gamma analyzed for et four model was (Darriba appropriate data jModelTest most which sequence the model, DNA GTR The a characters. with 99,999 resampling. structural jackknife of and entire total the data a of duration and the were replicate, for searches each memory TBR after in 10 held held replicate, trees trees each 10 For with performed. conducted, were 1996) ( al. replicates et jackknife thousand Ten swap- TBR-max, completion. by until trees followed most-parsimonious then the search all and among This times, ping and 1999). 100 fusion (Goloboff repeated tree performed was of strategy rounds were 100 searches afterwards, sectorial drifting; followed tree random downweighting, proba- of of 10% cycles probability ratchet a 1,000 10% parsimony and by a iteration 1,000 and per upweighting held of of trees bility held iterations 20 trees with independent 1999), 1,000,000 10 (Nixon with iterations undertaken, and Meade were memory, analyses 2008; charactersin MP structural 2004, the and Meade gaps), nucleotide Using (e.g. and respectively. 2008), (Pagel Pagel V2 and BayesPhylogenies al. et and (Goloboff TNT 2008) using conducted were analyses phylogenetic (BI) ence hlgntcAnalyses— Phylogenetic irmrhlg n ayooyo ot mrcnCynoglossum— American North of Palynology and Micromorphology euneData— Sequence N xrcinwspromdwt re ln iseuigamodi- a using tissue plant dried with performed was extraction DNA nta lgmnswr efre ihMSL Egr20)as 2004) (Edgar MUSCLE with performed were alignments Initial terminators 3.1 BigDye with performed were reactions Sequencing o Iaaye,tedt a attoe ewe N sequence DNA between partitioned was data the analyses, BI For 2008). al. et (Goloboff TNT with measured was support Clade ayou daubenmirei Dasynotus

+ rmSA X a ufr(aaaBoIc,Japan), Inc., Bio (Takara Buffer Taq GXL PrimeSTAR oehr h orDArgosyielded regions DNA four the Together, eeeaie sn cnigeeto micro- electron scanning a using examined were aiu asmn M)adBysa infer- Bayesian and (MP) parsimony Maximum ndhF Results .grande C. h ogs ein(,0 base (2,103 region longest the 6 eoa rbblt)(Farris probability) removal 36% , .occidentale C. , .virginianum C. , nlne rnhsta r te pce Fg ) hr,in Third, 2). are (Fig. species species these other of are both than analyses, branches longer BI on the of tree to consensus sister is and relatives analysis,and MP the In relationship this although tives, T 9 0 0 20.2% 6.9% 100 378 0 35 100 343 495 5,458 Total ITS trnL-trnF ndhF matk n eaie Fg.1ad2). and 1 (Figs. relatives includesand that clade the of species early-diverging as of species as group monophyletic a of of form species species World not Old World the Old Additionally, ses). the from distinct Cynoglossum clade of members ebr ftecaeta nldsteOdWrdspecies World Old the genus. includes the that of clade the of members nomtv hrces(1) and (115), characters informative ntepeetstudy, included present regions DNA the four in the Of regions. four the among ent 9%J,10 P.Ti ld ssse ooeta includes that one to sister is clade This C.grande,D.daubenmirei PP). 1.00 JK, (99% virginianum C. of jack- >70% received that with analyses (JK). compared MP sup- support the knife were (PP) in analyses probability nodes BI five posterior only the >0.90 in by nodes con- ported 34 strict greater Indeed, MP had the tree. analysis than sensus values BI support the higher struc- The of and the parti- tree resolution 0.75. for consensus one four of The and data. data in (RI) sequence tural DNA resulted index the for retention BayesPhylogenies three consistency tions, a in a and model with 0.53 RJBLS of steps, (CI) 928 index of trees most-parsimonious Lindelofia informative of number greatest The the (100). had nucleotides ITS however, (71); srcntutda itrt pce of species to sister as reconstructed is .grande C. presented are regions DNA 1. and Table four in the (20), of Statistics characters (4). fewest structural of number ino h osnu readtepaeet of placements the resolu- and the tree intermedia exceptions: consensus three the with of tion identical, are analyses BI PP). (0.92 tives l ol pce ftegns eod nteB analysis BI the of in species Second, genus. intermedia the BI C. of the species of than World species Old American resolved North less the is trees Cynoglossum both tree in however, consensus tree; MP the First, Region h ot mrcnseisof species American North The Table nteM n Iaaye,teNrhAeia species American North the analyses, BI and MP the In hlgntcAnalyses— Phylogenetic h hlgntcrltosisrsle rmteM and MP the from resolved relationships phylogenetic The Cynoglossum .Smayo N ein nlddi analyses. in included regions DNA of Summary 1. itrt h te pce nteclade. the in species other the to sister , egh(bp) length reeand Greene Aligned Cynoglossum ,1 52 1 9.5% 4.3% 4.3% 115 92 71 20 4 11 95 88 60 1,211 2,103 1,649 Paracaryum n eaie Fg.1ad2 .5P nB analy- BI in PP 0.95 2; and 1 (Figs. relatives and r tl ebr facaedsic rmthe from distinct clade a of members still are srsle ssse oteNrhAmerican North the to sister as resolved is , r itr,frigawl-upre clade well-supported a forming sisters, are Dasynotus uloiesites nucleotide .intermedia C. informative ubrof Number trnL nalsu pringlei Oncaglossum trnL , n tesaedsrbtdamong distributed are others and Dasynotus ,and - trnF and , - trnF h Paaye eutdi 420 in resulted analyses MP The .fendleri C. ile h raetnme of number greatest the yielded Cryptantha eevslwspot(.2PP). (0.52 support low receives informative ubrof Number yolsu occidentale Cynoglossum characters structural pcrpoie h greatest the provided spacer Cryptantha snse among nested is matK ,and Cynoglossum ree(i.1.I the In 1). (Fig. Greene Cryptantha Cryptantha rvddtefewest the provided finformative of oa number Total n eaie,with relatives, and characters Gen. Sutory (Greenm.) n eaie are relatives and Cynoglossum ndhF .daubenmirei D. Pardoglossum r resolved are Cryptantha Cryptantha Cryptantha n rela- and n rela- and a the had informative characters Percent and do ´ , . Delivered by Publishing Technology to: Michael Simpson IP: 146.244.235.206 on: Mon, 21 Sep 2015 23:37:14 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 05 OE:AEII N NESNLSU,TONWGNR RMCNGOSM613 CYNOGLOSSUM FROM GENERA NEW TWO ANDERSONGLOSSUM, AND ADELINIA COHEN: 2015] Fig. .Src osnu reo 2 otprioiu re f98ses I05,R=.5 ubr bv rnhsaejckievle >50%. values jackknife are branches above Numbers RI=0.75. CI=0.53, steps, 928 of trees most-parsimonious 420 of tree consensus Strict 1. Delivered by Publishing Technology to: Michael Simpson IP: 146.244.235.206 on: Mon, 21 Sep 2015 23:37:14 Copyright (c) American Society for Plant Taxonomists. All rights reserved. and .pringlei O. of species three includes also analysis MP the 40 [Volume BOTANY SYSTEMATIC 614 rbblte f1. of probabilities Fig. Cryptantha .Cnesste fBysa neec hlgn.Nmesaoebace r otro rbblte 09 n seik * eoeposterior denote (*) asterisks and >0.9, probabilities posterior are branches above Numbers phylogeny. inference Bayesian of tree Consensus 2. ssse oalre ld opsdof composed clade larger a to sister is n eaie.Spotvle J n P are PP) and (JK values Support relatives. and .pringlei O. samme fasalcaethat clade small a of member a is Pectocarya u nteB analysis, BI the in but , Dasynotus scoeyrltdt pce htba lciit nutlets glochidiate between bear differs of glochid that species of species type to the (although related topologies, both closely in but is placements, both for low Cynoglossum ). Pectocarya .pringlei O. and Delivered by Publishing Technology to: Michael Simpson IP: 146.244.235.206 on: Mon, 21 Sep 2015 23:37:14 Copyright (c) American Society for Plant Taxonomists. All rights reserved. of the with of as just species pringlei but, 2), American and North 1 depend- (Figs. analysis other tree of the type of the regions on different ing two in resolved in is 2010), included was that 05 OE:AEII N NESNLSU,TONWGNR RMCNGOSM615 CYNOGLOSSUM FROM GENERA NEW TWO ANDERSONGLOSSUM, AND ADELINIA COHEN: 2015] ing Cynoglossum of species suoop htaetesm hp Fg 4A). (Fig. shape same the the contrast, are that In pseudocolpi C). other 4B, of the (Fig. grains pollen differs for pseudocolpi case and the colpi not virginianum is C. of (2003) this species Simpson but two and (2014), Hargrove Spaeth sensu and The groove width. transverse in mm a 8–10 and length of in range pollen mm grains pollen 11–15 from The size pseudocolpi. in three and colpi three from those than (4–6) varies, hooks of glochid number each the apex Although virginianum the C. 3B). at (Fig. hooks surface each protuber-of nutlet of small the apex and on the hook), ances grappling at a nutlets hooks (resembling The recurved glochid species. with three The the glochidiate, 3A). among are (Fig. similar papillae also with are trichomes nutlets bearing all with lar, of species American North three Cynoglossum— and Amsinckiinae. bersofthesamecladeastheOldWorldspeciesof Cynoglossum eut rvd vdneta hs ot mrcnspe- group. American monophyletic a North of occidentale members these not that are cies evidence provide results Amsinckiinae, of members include not did and relationships study their species the and resolve three not these could among analyses their pres- study, the in ent those to results congruent recovered authors these including Amsinckiinae, of species the to mrcnseisof species American is exception intermedia The clade Amsinckiinae. The of (2013). members al. other PP), et 1.00 Weigend JK, by of (99% identified supported also well result a is relationship this and study. earlier the resolve grande to regions C. in cpDNA two employed limited (2013) al. quite et respectively,Weigend was studies sampling two although in ITS also (2013) and recovered al. and regions regions et cpDNA cpDNA 10 Weigend three utilizing and 2014), (2011, 2014) Cohen (2011, recovered. Cohen result a pce hthv enmmesof members been have that species ie hs hlgntcrltosis h ot American North the relationships, phylogenetic these Given irmrhlg n ayooyo ot American North of Palynology and Micromorphology h ot mrcnseisof species American North The of species American North three the of pollen The h rsn td nldsalfu fteNrhAmerican North the of four all includes study present The Cynoglossum .occidentale C. .boreale C. Cryptantha sntcoeyrltdt h l ol species World Old the to related closely not is , .grande C. .occidentale C. Oncaglossum nteB nlss hc ssse oteNorth the to sister is which analysis, BI the in and Cynoglossum instead, ; n of and enl,aseissmtmssgeae from segregated sometimes species a Fernald, edt rdc ult ihagetrnumber greater a with nutlets produce to tend . rdc olni hc h hp fthe of shape the which in pollen produce yolsu grande Cynoglossum n relatives. and and h irmrhlg ftelae fthe of leaves the of micromorphology The .virginianum C. .grande C. Cynoglossum and ayou daubenmirei Dasynotus .virginianum C. .occidentale C. Cynoglosssum ,and ssse ommeso Amsinckiinae, of members to sister as Cynoglossum .daubenmirei D. Discussion hudn ogrb nlddin included be longer no should and .virginianum C. . .pringlei O. .grande C. .daubenmirei D. Oncaglossum yolsu occidentale Cynoglossum r itr Fg.1ad2), and 1 (Figs. sisters are Cynoglossum Cynoglossum and Cynoglossum ssse to sister is n te ebr of members other and ssse to sister is ni eety(Sutory recently until Cynoglossum .virginianum C. Fg A )develops D) 4A, (Fig. 35hooks). (3–5 Dasynotus eia species Mexican a , .occidentale C. . scoerelatives close as selpodwith ellipsoid is aecliand colpi have , .grande C. r o mem- not are sqiesimi- quite is Cynoglossum Cynoglossum Oncaglossum .Although Cryptantha Dasynotus n the and , (includ- and and and ´ , h eie ed akad n eoe euvd180 recurved becomes and backwards bends pedicel In development. the fruit during changes this but deltate. is blade the and petiolate, are that leaves cauline ielae htaessie n h ld agsi shape in ranges oblanceolate. blade to the lanceolate and to sessile, spatulate are from that leaves line mrcnseisfrel in formerly species American development. of fruit pedicel during the but develop, fruits when ooegns( genus one to virginianum C. in.Tefrtrltst h l ol pce of species World Old modifica- the taxonomic to relates require first will The tions. that issues additional two oaiaee)hspooe rae icmcito for circumscription broader a proposed has boraginaceae/) Amsinckia n peh[04)adtesaeo h olnprs The pores. pollen the of shape of the [2003] presence and pollen Simpson the [2014]) and in Hargrove Spaeth differ (sensu and genera groove two transverse the a of of of species pollen but 4), The (Fig. characteristics. fruit Andersonglossum and pollen, flowers vegetative, subtending bracts or leaves al. of et inflorescences. lack and (Weigend the areolae triangular and broadly 2013) shorter, their by N eunedt n rnhn patterns.and branching and data sequence DNA ( o h ru.Tescn su sta micinehas Amsinckiinae that is issue system second taxonomic included The previously appropriate group. most the the for identify of to species World order Old the species, Cynoglossum American North the As members genus. with remain non-monophyletic widespread species geographically World a of Old American North the the however, for species; that scheme taxonomic resolved useful have a (2014) provides Cohen and (2013) Cynoglossum al. et Weigend ifri eeaiefeatures. vegetative in differ Andersonglossum relatives. World New northern its from for species case Mexican the is this and (Sutory 2014), and Spaeth (Hargrove 2003; pseudoappertures Amsinckiinae Simpson polar of of species with nutlets Some pollen groups. develop the two on the between as glochids, ture well not as but daubenmirei pollen, hairs, D. The of Amsinckiinae. presence of the species other the members betweentheNorthAmericanspeciesof many of feature to a the shape nutlet, to Amsinckiinae, due in keeled of that a differ mentioned (2013) of that al. presence et pseudocolpi Weigend and 4D). (Fig. colpi with Idaho. 2003; pollen to Simpson endemic and species shape (Hargrove including same groove 2014), Spaeth the transverse pollen develop are and a also Amsinckiinae pores with colpi of species pollen the Other the 4). (Fig. between and of shape groove, pollen verse in the contrast, differ In pores pseudocolpi. pollen the Adelinia nfoe,tepdcl fteseiso ohgnr r erect, are genera both of species the of pedicels the flower, In h eonto ftonwgnr eie rmteNorth the from derived genera new two of recognition The Andersonglossum nadto ople hrceitc htdfe between differ that characteristics pollen to addition In Cryptantha Adelinia ´ 00.Ti olnfauecnhl itnus the distinguish help can feature pollen This 2010). .Teerltosisaespotdb oethan more by supported are relationships These ). t.,btSmsn(http:/ Simpson but etc.), , Andersonglossum ifrfo h l ol pce of species World Old the from differ n ls eaie hudb evlae in reevaluated be should relatives close and sntmnpyei,adtepeetstudy present the and monophyletic, not is u o yteato)sol etransferred be should author) the by not but , per ob nadtoa rniinlfea- transitional additional an be to appears , n eaie Fg.1ad2,i transitional is 2), and 1 (Figs. relatives and Andersonglossum and and .daubenmirei D. and Cryptantha Adelinia Adelinia Adelinia ayou daubenmirei Dasynotus ak rnvregov,and groove, transverse a lacks pce ftetognr also genera two the of species , Andersonglossum .daubenmirei D. n eaie (e.g. relatives and Cynoglossum ssmlri ieadshape and size in similar is hc srsle ssister as resolved is which , )and ifrfo ahohrby other each from differ /www.sci.sdsu.edu/plants/ .grande C. Adelinia Adelinia Cynoglossum hudhighlight should Andersonglossum Andersonglossum lodevelops also eeoscau- develops Adelinia ean erect remains ,aperennial Plagiobothrys Cynoglossum Cynoglossum a trans- a has oanother to .pringlei O. bears and  , , . Delivered by Publishing Technology to: Michael Simpson IP: 146.244.235.206 on: Mon, 21 Sep 2015 23:37:14 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 1 YTMTCBTN Vlm 40 [Volume BOTANY SYSTEMATIC 616 grande Fig. saebr=200 = bar (scale .Sann lcrnmcocp mgso:A efsraeof surface Leaf A. of: images microscope electron Scanning 3. m ,75 m,

+ ). yolsu occidentale Cynoglossum saebr=100 = bar (scale m ,100 m,

+ .B ultsraeof surface Nutlet B. ). Cynoglossum Delivered by Publishing Technology to: Michael Simpson IP: 146.244.235.206 on: Mon, 21 Sep 2015 23:37:14 Copyright (c) American Society for Plant Taxonomists. All rights reserved. pce omryin formerly include species to subtribe the 617 CYNOGLOSSUM FROM GENERA NEW TWO ANDERSONGLOSSUM, AND ADELINIA COHEN: 2015] Adelinia one to glochids with one associated from is type eglochidiate. color is fruit yellow, corolla in that white, in change are shift a this with that Perhaps corollas orange. bear or Amsinckiinae of Cynoglosseae, cies of members as American such North other among are red, and colors to (Simpson these Amsinckiinae purple H corollas, of a or white members s of among white euncommon n exception to s the blue t with a are and, b2 that 0 0 corollas 9develop ) .B l to u Cynoglosseae e in synapomorphies ,p taxa u Boraginaceae. and other rand p from features l Amsinckiinae iden- e diagnostic ,a on distinguish n focus of dr should e studies dctification future o defined, r well obecome l l a saand r ep r e s e n t evsta r esl;tein cauline the lanceolate sessile; or are elliptical that to leaves compared petiolate, and 2 m Fig. ti neetn ont that note to interesting is It iia to Similar ,9,000 m, xLh. .I Cohen I. J. Lehm.) ex Oncaglossum .Sann lcrnmcocp mgso oln A. pollen. of images microscope electron Scanning 4. Mertensia .I oe,gn nov.—Type: gen. Cohen, I. J. + .C. ). Cynoglossum yolssmoccidentale Cynoglosssum .A h rae iiso h subtribe the of limits broader the As ). aooi Treatment Taxonomic Cynoglossum ohand Roth Dasynotus u h aln evsaedeltate are leaves cauline the but , Hackelia Adelinia lrsecsaeebracteate; are florescences (i.e. n h ot American North the and saebr=10 = bar (scale Adelinia dlnagrande Adelinia pz u otspe- most but Opiz, and , Andersonglossum Andersonglossum m ,1,000 m, (Douglas yolsu grande Cynoglossum

+ .D. ). , ayou daubenmirei Dasynotus ae o eeayseiesctd n oeo Douglas’ of none desig- and was cited, type specimens no any but were Douglas, nor by nated manuscript a on based grande Adelinia better. day every makes who 2014), (b. Cohen Etta ovoid, attach- triangular. abortion, broad attachment, by scar submedial fewer ment yellow, or to 4 included, brown Nutlets glochidiate, style terminal. supe- pyramidal, 2, gynoecium gynobase stigmas 4-parted, ellipsoid; ovary anthers rior, included, 5, fauc lanceolate stamens 5-lobed, flared, glabrous, to salverform, elliptical, linear white, with lobes and base blue the corolla lobes; to pentam- 5-lobed cymes, actinomorphic, calyx scorpioid erous, pedicellate, sericeous, flowered Flowers to ebracteate. 15–25 strigose to terminal, glabrous Inflorescences indument entire, margin evsatrae ipe aln n aa,bslleaves basal 5–21 basal, petiolate, and leaves cauline cauline epetiolate, simple, alternate, Leaves erect are pedicels and groove; fruit. transverse in a has pollen the nhsbo,Lhan(80 etosthat mentions (1830) Lehmann book, his In Etymology— rc eena eb o7 m tmgaru.Taproot. glabrous. Stem cm. 75 to herbs perennial Erect hd iewo erFr acue.1825, In Vancouver. Fort Columbia: n. near s. British wood CANADA. pine shady 1830.—TYPE: 25. 2: grande Cynoglossum saebr=5 = bar (scale noye[eintdhr] iae! M[image]!). BM [image]!; K here]: [designated (neotype saebr=1 = bar (scale hsgnsi ae o yduhe,Adeline daughter, my for named is genus This m Dulse em)J .Chn ob nov. comb. Cohen, I. J. Lehm.) ex (Douglas ,5,000 m,

oga xLh.Nv tr.Pug. Stirp. Nov. Lehm. ex Douglas + .B. ). m ,10,000 m, lapnae rsn,white; present, appendages al yolsu virginianum Cynoglossum

+ ).

+ –0c,leaf cm, 2–10 .grande C. saebr= bar (scale Douglas is Delivered by Publishing Technology to: Michael Simpson IP: 146.244.235.206 on: Mon, 21 Sep 2015 23:37:14 Copyright (c) American Society for Plant Taxonomists. All rights reserved. da,R .20.MSL:mlil eune lgmn with alignment sequenced multiple MUSCLE: 2004. C. tissue. fresh R. from DNA plant Edgar, of Isolation 1990. Doyle. L. J. and J. J. Doyle, jModelTest2: 2012. Posada. D. and of Doallo, Recipient R. Taboada, – L. Anderson G. R. D., William Darriba, 2009. Weller. G. S. and F. T. Daniel, molecu- and morphological of analysis phylogenetic A 2014. I. J. Cohen, molecu- and morphological of analysis phylogenetic A 2011. I. J. Cohen, species. order studied in the visit of variation to morphological me the allowing understand better or specimens to the loaning for through WTU International th and Aligner A&M for TEX/LL, Texas funding CodonCode Cor- Program. provided of CodonCode Grant University license species. License Gallegos studied Kellogg, Aligner free M. CodonCode the a D. and of carefully Solis provided C. images for L. poration and Bell SEM manuscript. D. the material, wonderful C. on took plant and commenting providing reviewers, and anonymous reading for three Meyers Anderson, C. C. S. and .Crlawiet let e oprl,ifoecne batae olnwt nyeutra prue n pseudoapertures; and apertures equatorial only with pollen ebracteate, inflorescences purple, to red to blue to white Corolla 1. etsa ra triangular. attach- ovoid, broad attachment, abortion, scar submedial by ment yellow, fewer to or brown included, 4 glochidiate, Nutlets style terminal. pyramidal, 2, gynobase stigmas pur- 4-parted, supe- to gynoecium ovary ellipsoid; red anthers rior, to to included, white 5, red stamens present, to ple; appendages white faucal and flared, to blue 5- corolla glabrous, salverform, lobes; purple, the linear to with 5-lobed base calyx pentamerous, actinomorphic, cellate, cyme scorpioid sericeou flowered 15–25 to strigose indument n aa,eeilt,3–38 epetiolate, cauline basal, simple, alternate, and Leaves Taproot. sericeous, to pilose pedicels and shape; in fruit. differ in grains recurved are pseudo- pollen and the apertures of the apertures ebracteate; are inflorescences the .Crlagent eo elwt rw,ifoecnesmwa rcet,ple iheutra prue and apertures equatorial with pollen bracteate, somewhat inflorescence brown, to yellow lemon to 40 green [Volume Corolla 1. BOTANY SYSTEMATIC is Andersonglossum species, the for the of neotype one describing the be as before to designated saw likely Douglas’ seems Lehmann type. Douglas, which specimens and a Kew, Lehmann at as of specimen one 1825 spirit designating the in annotation Therefore, an bear specimens 618 Acknowledgments. rc eena eb o1m tmhsi osrgs to strigose to hispid Stem m. 1 to herbs perennial Erect iia to Similar .Cuielae esl,pdcl euvdi ri,ple oe fdfeetsae,sespbse . . . pubescen stems shapes, glabrous different stems of shape, pores same pollen the fruit, of in pores recurved pollen pedicels fruit, sessile, in leaves recurved Cauline not pedicels petiolate, leaves 2. Cauline 2. ihacrc n ihthroughput. high 1792–1797. and accuracy high Calif.) Francisco, (San Focus 772. computing. parallel 9: and heuristics new models, More Award. Gray Asa 2008 the evolution. character of patterns taxonomy,and relationships, Evolutionary Boraginaceae: of characters lar evolution. character taxa: related 559–580. and and relationships (Boraginaceae) L. Evolutionary Lithospermum of characters lar virginianum e to Key plantsoftheU.S.AandCanada 2 ...... suoprue n oa suoprue;pat fMxc . . . Mexico of plants pseudoapertures; polar and pseudoapertures .Crlaprl ord tl mi egh lnso aionaadOregon and California of plants length; in mm 4 > style red, to purple Corolla 3. .Crlabu owie style white, to blue Corolla 3. andCanada ...... A Cynoglossum DELINIA L)J .Cohen I. J. (L.) .I oe,gn nov.—Type: gen. Cohen, I. J. ,A h uhrwudlk otakM .Alford H. M. thank to like would author The ieaueCited Literature NDERSONGLOSSUM u h aln evsaesessile; are leaves cauline the but , 2 13–15. 12: ytmtcBotany Systematic + .–1c,la agnentire, margin leaf cm, 0.5–11 .grande A. eproject.IamgratefultoMICH, ,ercet.Foespedi- Flowers ebracteate. s, ,Ifoecne terminal, Inflorescences s, < oe,lbselpia,erect elliptical, lobes lobed, Cladistics . mi egh lnso h otenadesen(o etr)U .A. S. U. western) (not eastern and northern the of plants length; in mm 3.5 uli cd Research Acids Nucleic , . AND 4 1–3. 34: 0 139–169. 30: O Andersonglossum NCAGLOSSUM aueMethods Nature Cladistics (T 32: 27: HE ...... N ORTH ars .S,V .Abr,M Ka M. Albert, A. V. S., J. Farris, ooof . .S ars n .C io.20.TT repormfor program free a TNT, 2008. Nixon. C. K. Solu- and times: Farris, S. reasonable J. in P., sets Goloboff, data large Analyzing 1999. P. Goloboff, ae,M n .Mae 04 hlgntcmxuemdlfrdetect- for Published model mixture York: phylogenetic A New 2004. Meade. Ithaca, A. 1.7. and M. ver. Pagel, WinClada 2002. C. K. Nixon, heterocolpate of Ultrastructre 2003. Simpson. G. M. and L. Hargrove, ed,A n .Pgl 08 hlgntcmxuemdlfor model mixture phylogenetic A 2008. Pagel. M. and A. Meade, 1830. C. pop- G. J. and Lehmann, eyes Cat’s 2012. Simpson. G. M. and K. Hasenstab-Lehman, io,K .19.Prioyrtht e ehdfrrpdparsimony rapid for method new a ratchet, Parsimony 1999. C. K. Nixon, ev,F n .Sutory K. and F. Selvi, phylogenetic in heterotachy Modelling 2008. Meade. A. and M. Pagel, from boreale me Andersonglossum inspired who person, systematics. incom- plant an and study to 2009), botanist, Weller professor, and parable Daniel (1942–2013; Anderson nesnlsu virginianum Andersonglossum occidentale made. Andersonglossum been has combination new appropriate the nists, Etymology— lhuhteato osntaccept not does author the Although hlgntcanalysis. phylogenetic optima. composite for tions 99–124. 12: neighbor-joining. outperforms jackknifing Parsimony 1996. n atr-eeoeet ngn euneo hrce-tt data. character-state or sequence Biology Systematic gene in pattern-heterogeneity ing author. by in pollen tion in 29–41 Pp. heterotachy. Academico. Gymansio Hamburgensium Hamburg: genus the of (Boraginaceae). systematics Phylogenetic flowers: corn 137–151. 164: analysis. rnatoso h oa oit fLno.Sre ,Booia Sciences Biological B, Series London. of 3955–3964. Society 363: Royal the of Transactions Carlo. Monte chain Markov reversible-jump by inference Brgnca-yolsee nItaly. in (Boraginaceae-Cynoglosseae) A enl .n. s. Cascapedia Fernald 1905, Little 17, July Quebec: River, CANADA. 1984.—TYPE: ye M[image]!). Virginia, BM type: in Habitat Virginia: U.S.A. virginianum Cynoglossum 2010. var. 3. 13: Notes Bot. Stantec virginianum Cynoglossum nov. rs1:5.17.TP:USA aiona Sierra California: U.S.A. [image]!). 1874, GH 1874.—TYPE: California, and 58. Co. 10: Arts nov. .virginianum A. MERICAN d .Pnaot.Hiebr:Srne Verlag. Springer Heidelberg: Pontarotti. P. ed. , boreale yolsu occidentale Cynoglossum yolsu boreale Cynoglossum ...... Cladistics Cryptantha S PECIES hsgnsi ae o ila Russell William for named is genus This lcoye H[image]!). GH (lectotype: Frad opr.Mcia o.2:166. 23: Bot. Michigan Cooperr. (Fernald) ytmtcBotany Systematic 3 571–581. 53: 5 407–414. 15: ´ ie hti sacpe yohrbota- other by accepted is it that given , (Boraginaceae). 02 yosso h genus the of synopsis A 2012. . oau tMnsCgiau trimPugillus Stirpium Cognitarum Minus et Novarum F ORMERLY Cladistics .F ilas .F oln,&M L. M. & Collins, F. J. Williams, F. E. vltoaybooyfo ocp oapplica- to concept from biology Evolutionary ¨ ...... 3 ...... llersjo Cladistics ...... subsp. Frad .I oe,comb. Cohen, I. J. (Fernald) .S.P.1 3.1753.—TYPE: 134. 1: Pl. Sp. L. enl hdr :20 1905. 250. 7: Rhodora Fernald 4 774–786. 24: I .G emn667 Lemmon G. J. CUE IN NCLUDED ¨ 7 738–757. 37: A ry .I oe,comb. Cohen, I. J. Gray) (A. .Lpcm,adA .Kluge. G. A. and Lipscomb, D. , nentoa ora fPatSciences Plant of Journal International L)J .Chn ob nov. comb. Cohen, I. J. (L.) .Ga rc mr Acad. Amer. Proc. Gray A. boreale ln Biosystems Plant yolsu virginianum Cynoglossum 5 415–428. 15: nesnlsu virginianum Andersonglossum nesnlsu occidentale Andersonglossum .boreale A. Frad .Haines A. (Fernald) lyo 257 Clayton C nalsu pringlei Oncaglossum YNOGLOSSUM dlnagrande Adelinia Cryptantha 4:461–479. 146: Cynoglossum sdistinct as (holotype: Philosophical Cladistics (lecto- ) s.l. . Delivered by Publishing Technology to: Michael Simpson IP: 146.244.235.206 on: Mon, 21 Sep 2015 23:37:14 Copyright (c) American Society for Plant Taxonomists. All rights reserved. Sutory 2009. Hasenstab. 2014. E. R. Spaeth, K. and G. M. Simpson, 05 OE:AEII N NESNLSU,TONWGNR RMCNGOSM619 CYNOGLOSSUM FROM GENERA sequence- NEW in TWO characters ANDERSONGLOSSUM, as Gaps AND 2000. ADELINIA Ochoterena. COHEN: H. and P. M. Simmons, 2015] yolsee rmMexico. from Cynoglosseae, onr ak aiona ooaSaeUniversity. State (Boraginaceae) Sonoma Cryptanthinae California: the Park, Rohnert in variation pollen using California. ae hlgntcanalyses. phylogenetic based ´ .21.Ocgosm e eu fBrgnca,tribe Boraginaceae, of genus new a Oncaglossum, 2010. K. , Crossosoma opooia hrce apn namlclrphylogeny molecular a on mapping character Morphological 5 1–59. 35: ytmtcBiology Systematic Novon 0 463–469. 20: Cryptantha 9 369–381. 49: ..thesis. M.S. . fsouthern of h,G,H id,adR .Kmln 95 oaiaee p 329–427 Pp. Boraginaceae. 1995. Kamelin. V. R. and Riedl, H. G., Zhu, W e i g e n d ,M . ,F .L u e b e r t ,F .S e l v i ,G .B r o k a m p ,a n dH .H .H i l g e r .2 0 1 3 . .Y uadP .Rvn ejn n t oi:SinePesand Press Science Louis: St. and Beijing Raven. Press. Garden H. Botanical P. Missouri and Wu Y. Z. in 604–618. s.str.). (Boraginaceae ed ( ( seeds tongues Hound’s for origins Multiple lr fChina of Flora Omphalodes o.1 Gninca hog oaiaee,eds. Boraginaceae), through (Gentianaceae 16 vol. il)–Tepyoeyo h oaefamily borage the of phylogeny The – Mill.) oeua hlgntc n Evolution and Phylogenetics Molecular Cynoglossum . n Navel and L.) 68: